Animal movement in the absence of predation: environmental drivers of movement strategies in a partial migration system

Animal movement strategies including migration, dispersal, nomadism, and residency are shaped by broad‐scale spatial‐temporal structuring of the environment, including factors such as the degrees of spatial variation, seasonality and inter‐annual predictability. Animal movement strategies, in turn, interact with the characteristics of individuals and the local distribution of resources to determine local patterns of resource selection with complex and poorly understood implications for animal fitness. Here we present a multi‐scale investigation of animal movement strategies and resource selection. We consider the degree to which spatial variation, seasonality, and inter‐annual predictability in resources drive migration patterns among different taxa and how movement strategies in turn shape local resource selection patterns. We focus on adult Galapagos giant tortoises Chelonoidis spp. as a model system since they display many movement strategies and evolved in the absence of predators of adults. Specifically, our analysis is based on 63 individuals among four taxa tracked on three islands over six years and almost 10⁶ tortoise re‐locations. Tortoises displayed a continuum of movement strategies from migration to sedentarism that were linked to the spatio‐temporal scale and predictability of resource distributions. Movement strategies shaped patterns of resource selection. Specifically, migratory individuals displayed stronger selection toward areas where resources were more predictable among years than did non‐migratory individuals, which indicates a selective advantage for migrants in seasonally structured, more predictable environments. Our analytical framework combines large‐scale predictions for movement strategies, based on environmental structuring, with finer‐scale analysis of space‐use. Integrating different organizational levels of analysis provides a deeper understanding of the eco‐evolutionary dynamics at play in the emergence and maintenance of migration and the critical role of resource predictability. Our results highlight that assessing the potential benefits of differential behavioral responses first requires an understanding of the interactions among movement strategies, resource selection and individual characteristics.     

Spatial memory shapes migration and its benefits: evidence from a large herbivore

From fine‐scale foraging to broad‐scale migration, animal movement is shaped by the distribution of resources. There is mounting evidence, however, that learning and memory also guide movement. Although migratory mammals commonly track resource waves, how resource tracking and memory guide long‐distance migration has not been reconciled. We examined these hypotheses using movement data from four populations of migratory mule deer (n = 91). Spatial memory had an extraordinary influence on migration, affecting movement 2–28 times more strongly than tracking spring green‐up or autumn snow depth. Importantly, with only an ability to track resources, simulated deer were unable to recreate empirical migratory routes. In contrast, simulated deer with memory of empirical routes used those routes and obtained higher foraging benefits. For migratory terrestrial mammals, spatial memory provides knowledge of where seasonal ranges and migratory routes exist, whereas resource tracking determines when to beneficially move within those areas.     

How landscape dynamics link individual‐ to population‐level movement patterns: a multispecies comparison of ungulate relocation data

Aim To demonstrate how the interrelations of individual movements form large‐scale population‐level movement patterns and how these patterns are associated with the underlying landscape dynamics by comparing ungulate movements across species. Locations Arctic tundra in Alaska and Canada, temperate forests in Massachusetts, Patagonian Steppes in Argentina, Eastern Steppes in Mongolia. Methods We used relocation data from four ungulate species (barren‐ground caribou, Mongolian gazelle, guanaco and moose) to examine individual movements and the interrelation of movements among individuals. We applied and developed a suite of spatial metrics that measure variation in movement among individuals as population dispersion, movement coordination and realized mobility. Taken together, these metrics allowed us to quantify and distinguish among different large‐scale population‐level movement patterns such as migration, range residency and nomadism. We then related the population‐level movement patterns to the underlying landscape vegetation dynamics via long‐term remote sensing measurements of the temporal variability, spatial variability and unpredictability of vegetation productivity. Results Moose, which remained in sedentary home ranges, and guanacos, which were partially migratory, exhibited relatively short annual movements associated with landscapes having very little broad‐scale variability in vegetation. Caribou and gazelle performed extreme long‐distance movements that were associated with broad‐scale variability in vegetation productivity during the peak of the growing season. Caribou exhibited regular seasonal migration in which individuals were clustered for most of the year and exhibited coordinated movements. In contrast, gazelle were nomadic, as individuals were independently distributed and moved in an uncoordinated manner that relates to the comparatively unpredictable (yet broad‐scale) vegetation dynamics of their landscape. Main conclusions We show how broad‐scale landscape unpredictability may lead to nomadism, an understudied type of long‐distance movement. In contrast to classical migration where landscapes may vary at broad scales but in a predictable manner, long‐distance movements of nomadic individuals are uncoordinated and independent from other such individuals. Landscapes with little broad‐scale variability in vegetation productivity feature smaller‐scale movements and allow for range residency. Nomadism requires distinct integrative conservation strategies that facilitate long‐distance movements across the entire landscape and are not limited to certain migration corridors.     

Where the wild birds go: explaining the differences in migratory destinations across terrestrial bird species

Given their large movement capacities, migratory birds have in principle a wide range of possible geographical locations for their breeding and non‐breeding destinations, yet each species migrates between consistent breeding and non‐breeding ranges. In this study, we use a macroecological approach to search for the general factors explaining the location of the seasonal ranges of migratory bird species across the globe. We develop a null model to test the hypotheses that access to resources, geographical distance, tracking of temperature, and habitat conditions (separately as well as considered together) have a major influence in the location of species’ migratory destinations, once each species’ geographical constraints are taken into account. Our results provide evidence for a trade‐off between costs associated with distance travelled and gains in terms of better access to resources. We also provide strong support to the hypotheses that all factors tested, with the exception of habitat, have a strong and additive effect on the global geography of bird migration. Indeed, our results indicate that species’ contemporary migratory destinations (i.e. the combination of their breeding and non‐breeding ranges) are such that they allow species to track a temperature regime throughout the year, to escape local competition and reach areas with better access to resources, and to minimise the spatial distance travelled, within the limitations imposed by the geographical location of each species. Our study thus sheds light on the mechanisms underpinning bird migration and provides a strong basis for predicting how migratory species will respond to future change.     

Movement tactics of a mobile predator in a meta‐ecosystem with fluctuating resources: the arctic fox in the High Arctic

Animal movement is a fundamental process shaping ecosystems at multiple levels, from the fate of individuals to global patterns of biodiversity. The spatio‐temporal dynamic of food resources is a major driver of animal movement and generates patterns ranging from range residency to migration and nomadism. Arctic tundra predators face a strongly fluctuating environment marked by cyclic microtine populations, high seasonality, and the potential availability of sea ice, which gives access to marine resources in winter. This type of relatively poor and highly variable environment can promote long‐distance movements and resource tracking in mobile species. Here, we investigated the winter movements of the arctic fox, a major tundra predator often described as a seasonal migrant or nomad. We used six years of Argos satellite telemetry data collected on 66 adults from Bylot Island (Nunavut, Canada) tracked during the sea ice period. We hypothesized that long‐distance movements would be influenced by spatio‐temporal changes in resource availability and individual characteristics. Despite strong annual and seasonal changes in resource abundance and distribution, we found that a majority of individuals remained resident, especially those located in an area characterized by highly predictable pulse resources (goose nesting colony) and abundant cached food items (eggs). Foxes compensated terrestrial food shortage by commuting to the sea ice rather than using long‐distance tracking or moving completely onto the sea ice for winter. Individual characteristics also influenced movement patterns: age positively influenced the propensity to engage in nomadism, suggesting older foxes may be driven out of their territories. Our results show how these mammalian predators can adjust their movement patterns to favor range residency despite strong spatio‐temporal fluctuations in food resources. Understanding the movement responses of predators to prey dynamics helps identifying the scales at which they work, which is a critical aspect of the functioning and connectivity among meta‐ecosystems.     

A general modeling framework for describing spatially structured population dynamics

Variation in movement across time and space fundamentally shapes the abundance and distribution of populations. Although a variety of approaches model structured population dynamics, they are limited to specific types of spatially structured populations and lack a unifying framework. Here, we propose a unified network‐based framework sufficiently novel in its flexibility to capture a wide variety of spatiotemporal processes including metapopulations and a range of migratory patterns. It can accommodate different kinds of age structures, forms of population growth, dispersal, nomadism and migration, and alternative life‐history strategies. Our objective was to link three general elements common to all spatially structured populations (space, time and movement) under a single mathematical framework. To do this, we adopt a network modeling approach. The spatial structure of a population is represented by a weighted and directed network. Each node and each edge has a set of attributes which vary through time. The dynamics of our network‐based population is modeled with discrete time steps. Using both theoretical and real‐world examples, we show how common elements recur across species with disparate movement strategies and how they can be combined under a unified mathematical framework. We illustrate how metapopulations, various migratory patterns, and nomadism can be represented with this modeling approach. We also apply our network‐based framework to four organisms spanning a wide range of life histories, movement patterns, and carrying capacities. General computer code to implement our framework is provided, which can be applied to almost any spatially structured population. This framework contributes to our theoretical understanding of population dynamics and has practical management applications, including understanding the impact of perturbations on population size, distribution, and movement patterns. By working within a common framework, there is less chance that comparative analyses are colored by model details rather than general principles.     

The distribution and biology of nomadic birds in the Karoo, South Africa

Dryland nomadic bird species, as a proportion of all bird species in a biome in southern Africa, are highest in the arid grassland and arid and semi-arid Karoo in South Africa. Nomadic birds, of which the most widespread species is the greybacked finchlark Eremopterix verticalis (Smith), are most frequently observed in the north-central and north western Nama Karoo. The species richness of nomadic species is inversely correlated with species richness of all bird species in the Karoo. Since the distribution of nomadic birds is in areas where rainfall is patchy, low (<250 mm per year) and aseasonal, this supports the idea that fewer species are able to cope with resources that are patchy in time and space, and that there has been selection for nomadism in the species that are able to use patchy environments. Species richness and abundance of nomadic birds is negatively correlated with rainfall amount but positively correlated with the coefficient of variation of the rainfall and with rainfall in autumn. The frequency of nomadic birds is inversely correlated with altitude range; nomadic species are most often recorded in structurally simple habitats (shrubland and grassland) on open plains. Most nomadic bird species in the Karoo are granivorous. Perennial desert grasses are important components of the habitat and diet of small nomadic granivores, and also provide nest sites and nest material. Nomadic birds can breed throughout the year, without a clearly defined ‘season’ in both the Succulent and Nama Karoo. Average clutch sizes do not differ significantly between resident and all nomadic species in the arid and semi-arid Karoo. Nomadism is an evolutionary stable strategy for individual species only when extremes in environmental conditions are frequent enough, and unpredictable enough, to maintain movements to high resource patches or to maintain dispersal away from low resource patches. If high rainfall years are too regular or infrequent, or peaks in fluctuations of resources in the environment too low, or rainfall patches are randomly distributed, nomadism would not be maintained as part of the individual behaviour pattern.     

Quantifying Migration Behaviour Using Net Squared Displacement Approach: Clarifications and Caveats

Estimating migration parameters of individuals and populations is vital for their conservation and management. Studies on animal movements and migration often depend upon location data from tracked animals and it is important that such data are appropriately analyzed for reliable estimates of migration and effective management of moving animals. The Net Squared Displacement (NSD) approach for modelling animal movement is being increasingly used as it can objectively quantify migration characteristics and separate different types of movements from migration. However, the ability of NSD to properly classify the movement patterns of individuals has been criticized and issues related to study design arise with respect to starting locations of the data/animals, data sampling regime and extent of movement of species. We address the issues raised over NSD using tracking data from 319 moose (Alces alces) in Sweden. Moose is an ideal species to test this approach, as it can be sedentary, nomadic, dispersing or migratory and individuals vary in their extent, timing and duration of migration. We propose a two-step process of using the NSD approach by first classifying movement modes using mean squared displacement (MSD) instead of NSD and then estimating the extent, duration and timing of migration using NSD. We show that the NSD approach is robust to the choice of starting dates except when the start date occurs during the migratory phase. We also show that the starting location of the animal has a marginal influence on the correct quantification of migration characteristics. The number of locations per day (1–48) did not significantly affect the performance of non-linear mixed effects models, which correctly distinguished migration from other movement types, however, high-resolution data had a significant negative influence on estimates for the timing of migrations. The extent of movement, however, had an effect on the classification of movements, and individuals undertaking short- distance migrations can be misclassified as other movements such as sedentary or nomadic. Our study raises important considerations for designing, analysing and interpreting movement ecology studies, and how these should be determined by the biology of the species and the ecological and conservation questions in focus.     

How New Guinea rainforest flower resources vary in time and space: Implications for nectarivorous birds

Abstract Variability in spatial and temporal patterning of flowering by populations of rainforest trees fed upon by honeyeaters and flower-visiting parrots was studied for 2 years in lowland tropical hill forest in Papua New Guinea. All 2200 trees in a 3 ha plot were tagged, identified, mapped and monitored monthly. Of 274 tree species present, 86 flowered during the course of the study; during any given month, approximately 20% of the species flowering that month were visited by nectarivorous birds. Results showed that overall flower resources (total number of species, and number of bird-pollinated species, individuals and flowers) fluctuated during the year, decreased during the dry season and increased during the wet season. In addition, there was a wide range of temporal variation within and among tree species in length and timing of flowering period, percentage of each conspecific population flowering from year to year, and degree of synchrony among flowering conspecifics. Spatial dispersion of tree populations also varied, from clumps to scattered single individuals. Resident bird species were correlated with synchronously flowering trees, whereas nomadic bird species were correlated with asynchronously flowering trees. Resident birds were also associated with smaller blooming displays per tree, whereas nomadic birds were associated with trees that bloomed massively. There was no correlation between avian nomadism and spatial dispersion of tree populations. Thus nomadic birds seem to range in search of rich but unpredictable resources; resident birds may rely more on predictable, but smaller resources.     

Variability between Scales: Predictors of Nomadism in Birds of an Australian Mediterranean-climate Ecosystem

Nomadism in animals is a response to resource distributions that are highly variable in time and space. Using the avian fauna of the Mediterranean-climate region of southcentral Australia, we tested a number of variables to determine if they predicted nomadism. These variables were species body mass, the distance in body mass terms to the edge of a body mass aggregation, and diet (for example, seeds, invertebrates, nectar, or plants). We utilized two different classifications of the avifauna that diverged in their definition of nomadic to build two different predictive models. Using both classifications, distance to the edge of a body mass aggregation was found to be a significant predictor of nomadism. There was also evidence that nomads tend to feed on nectar and tend to be large. The significance of the variables body mass and diet (nectar) may reflect the greater energy requirements of large birds and the inherent variability of nectar as a food source. The significance of the variable distance to the edge of a body mass aggregation provides further evidence of inherent variability in resources between domains of scale. The edges of body mass aggregations are hypothesized to be areas of increased resource variability that reflect the transition from one scale of landscape pattern to another. Received 14 November 2000; accepted 27 June 2001.     

A memory‐based foraging tactic reveals an adaptive mechanism for restricted space use

The restricted area of space used by most mobile animals is thought to result from fitness‐rewarding decisions derived from gaining information about the environment. Yet, assessments of how animals deal with uncertainty using memory have been largely theoretical, and an empirically derived mechanism explaining restricted space use in animals is still lacking. Using a patch‐to‐patch movement analysis, we investigated predictions of how free‐ranging bison (Bison bison) living in a meadow‐forest matrix use memory to reduce uncertainty in energy intake rate. Results indicate that bison remembered pertinent information about location and quality of meadows, and they used this information to selectively move to meadows of higher profitability. Moreover, bison chose profitable meadows they had previously visited, and this choice was stronger after visiting a relatively poor quality meadow. Our work demonstrates a link between memory, energy gains and restricted space use while establishing a fitness‐based integration of movement, cognitive and spatial ecology.     

The influence of habitat fragmentation on genetic diversity of a rare bird species that commonly faces environmental fluctuations

Habitat loss and fragmentation is one of the main causes of biodiversity loss. Rare species are generally thought to be more sensitive to habitat fragmentation than common ones as small populations become even smaller. We did a population genetic study on a rare bird, the Worthen's sparrow Spizella wortheni which is endemic to semi‐arid and arid regions of northeast Mexico. Its population numbers suffer greatly from the transformation of grassland into farmland that leads to a patchy distribution with locally small population sizes. Our data show that its genetic diversity is nevertheless high, few to no differentiation between study localities was found, and gene flow was high. Although we can not exclude that is too early to see an impact on the genetic level, we think that these results might be explained by the species’ biology: like many other birds living in arid areas, the Worthen's sparrow has a nomadic life style; depending on local conditions individuals flexibly move between areas. This behavior could enhance their ability to find suitable habitat patches in a fragmented landscape. Our results imply that nomadic behavior, which is an adaptation to high temporal variability in environmental conditions, may make species more resilient to spatial variability caused by habitat fragmentation. This insight contributes to identifying common factors such as nomadism that predict a species’ sensitivity to habitat fragmentation.     

A geolocator-tagged fledgling provides first evidence on juvenile movements of Cory’s Shearwater Calonectris borealis

Using geolocator-immersion loggers, we tracked for the first time the migration of one Cory’s Shearwater Calonectris borealis fledgling, from its breeding colony in the Canary Islands, and along its first year of life. The juvenile bird initially followed the same migratory path as the adults but visited different areas of the Central and the South Atlantic Ocean.     

Travel optimization by foraging bumblebees through readjustments of traplines after discovery of new feeding locations

Animals collecting resources that replenish over time often visit patches in predictable sequences called traplines. Despite the widespread nature of this strategy, we still know little about how spatial memory develops and guides individuals toward suitable routes. Here, we investigate whether flower visitation sequences by bumblebees Bombus terrestris simply reflect the order in which flowers were discovered or whether they result from more complex navigational strategies enabling bees to optimize their foraging routes. We analyzed bee flight movements in an array of four artificial flowers maximizing interfloral distances. Starting from a single patch, we sequentially added three new patches so that if bees visited them in the order in which they originally encountered flowers, they would follow a long (suboptimal) route. Bees' tendency to visit patches in their discovery order decreased with experience. Instead, they optimized their flight distances by rearranging flower visitation sequences. This resulted in the development of a primary route (trapline) and two or three less frequently used secondary routes. Bees consistently used these routes after overnight breaks while occasionally exploring novel possibilities. We discuss how maintaining some level of route flexibility could allow traplining animals to cope with dynamic routing problems, analogous to the well-known traveling salesman problem.     

How a simple adaptive foraging strategy can lead to emergent home ranges and increased food intake

Animals often alternate between searching for food locally and moving over larger distances depending on the amount of food they find. This ability to switch between movement modes can have large implications on the fate of individuals and populations, and a mechanism that allows animals to find the optimal balance between alternative movement strategies is therefore selectively advantageous. Recent theory suggests that animals are capable of switching movement mode depending on heterogeneities in the landscape, and that different modes may predominate at different temporal scales. Here we develop a conceptual model that enables animals to use either an area‐concentrated food search behavior or undirected random movements. The model builds on the animals’ ability to remember the profitability and location of previously visited areas. In contrast to classical optimal foraging models, our model does not assume food to be distributed in large, well‐defined patches, and our focus is on animal movement rather than on how animals choose between foraging patches with known locations and value. After parameterizing the fine‐scale movements to resemble those of the harbor porpoise Phocoena phocoena we investigate whether the model is capable of producing emergent home ranges and use pattern‐oriented modeling to evaluate whether it can reproduce the large‐scale movement patterns observed for porpoises in nature. Finally we investigate whether the model enables animals to forage optimally. We found that the model was indeed able to produce either stable home ranges or movement patterns that resembled those of real porpoises. It enabled animals to maximize their food intake when fine‐tuning the memory parameters that controlled the relative contribution of area concentrated and random movements.     

‘Chancing on a spectacle:’ co-occurring animal migrations and interspecific interactions

Migrations of diverse wildlife species often converge in space and time, with their journeys shaped by similar forces (i.e. geographic barriers and seasonal resources and conditions); we term this ‘co-migration’. Recent studies have illuminated multi-speciesmigrations by land and sea including the simultaneous movements of numerous insects, birds, bats and of fish invertebrates marine predators. Beyond their significance as natural wonders, species with overlapping migrations may interact ecologically, with potential effects on population and community dynamics. Direct and indirect ecological interactions (including predation and competition) between migrant species remain poorly understood, in part because migration is the least-studied phase of animals’ annual cycles. To address this gap, we conducted a literature review to examine whether animal migration studies incorporate multiple species and to what extent they investigate interspecific interactions between co-migrants. Following a key word search, we read all migration research papers in 23 relevant peer-reviewed journals during 2008–2017. Thirty percent of animal migration papers reported two or more species with coinciding migrations, suggesting that co-migrations are common, although few studies investigated or discussed these mixed-species migrations further. Synthesizing these, we present examples and describe five types of ecological interactions between migrating species, including predator–prey, host–parasite and commensal relationships. Considering migratory animals as interacting with migrant communities will enhance understanding of the drivers of migration and could improve predictions about wildlife responses to global change. Further research focused on multi-species migrations could also inform conservation efforts for migratory animal populations, many of which are declining or shifting, with unexplored consequences for other co-migratory species.     

Being there and getting there: balancing the costs of migration against the benefits of changing habitat for North Sea plaice

To improve survival and reproductive success, many fish species have evolved migratory life‐histories, showing ontogenetic and/or seasonal changes in habitat use. Individuals move between different areas, each of which is 'best' for a particular activity, such as feeding, growing or spawning. The benefits of moving to a different habitat, however, have to be balanced against the costs of migration, so any behaviour that reduces the cost of movement would be expected to expand the migratory range and thereby increase the range of habitats that can be exploited.
Previous tracking experiments in the southern North Sea have shown that plaice selectively exploit the tidal streams to aid their spawning migration. Here we examine whether this behaviour is primarily (1) an energy‐saving strategy, or (2) a transport mechanism by which fish that are unable to navigate over long distances can be carried reliably between feeding and spawning grounds. Because selective tidal stream transport requires that fish remain stationary on the sea‐bed during the 'non‐transporting' tide, energetic calculations predict that this behaviour is beneficial only when the current speed exceeds a critical, size‐dependent, value. We have used detailed information about migratory behaviour from individual fish to calculate the metabolic costs and the likely reproductive benefits of different migratory strategies. Our results show that plaice use selective tidal stream transport only in areas where the tidal streams are suitably fast, indicating that this behaviour is primarily an energy saving strategy.     

Weak migratory connectivity,loop migration and multiple non-breeding site use in British breeding Whinchats Saxicola rubetra

Determining the links between breeding populations and the pressures, threats and conditions they experience presents a challenge for the conservation of migratory birds which can use multiple sites separated by hundreds to thousands of kilometres. Furthermore, migratory connectivity – the connections made by migrating individuals between networks of breeding and non-breeding sites – has important implications for population dynamics. The Whinchat Saxicola rubetra is declining across its range, and tracking data from a single African non-breeding site implies high migratory spread. We used geolocators to describe the migration routes and non-breeding areas of 20 Whinchats from three British breeding populations. As expected, migratory spread was high, with birds from the three populations overlapping across a wide area of West Africa. On average, in non-breeding areas, British breeding Whinchats were located 652 km apart from one another, with some likely to share non-breeding areas with individuals from breeding populations as far east as Russia. Four males made a direct non-breeding season movement to a second, more westerly, non-breeding location in January. Autumn migration was through Iberia and around the western edge of the Sahara Desert, whereas spring migration was more direct, indicating an anticlockwise loop migration. Weak migratory connectivity implies that Whinchat populations are somewhat buffered against local changes in non-breeding conditions. If non-breeding season processes have played a role in the species’ decline, then large-scale drivers are likely to be the cause, although processes operating on migration, or interactions between breeding and non-breeding processes, cannot be ruled out.     

Remembering the good and the bad: memory-based mediation of the food–safety trade-off in dynamic landscapes

Predator–prey interactions are central to fitness as animals simultaneously avoid death and consume resources to ensure growth and reproduction. Along with direct effects, predators can also exert strong non-consumptive effects. For example, prey shift habitat use in the presence of predators, a potentially learned behavior. The impact of cognition on movement and predator interactions is largely unexplored despite evidence of learned responses to predation threat. We explore how learning and spatial memory influence predator–prey dynamics by introducing predators into a memory-driven movement modeling framework. To model various aspects of risk, we vary predator behavior: their persistence and spatial correlation with the prey’s resources. Memory outperforms simpler movement processes most in patchy environments with more predictable predators that are more easily avoided once learned. In these cases, memory aids foragers in managing the food–safety trade-off. For example, particular parameterizations of the predation memory reduce encounters while maintaining consumption. We found that non-consumptive effects are highest in landscapes of concentrated, patchy resources. These effects are intensified when predators are highly correlated with the forager’s resources. Smooth landscapes provide more opportunities for foragers to simultaneously consume resources and avoid predators. Predators are able to effectively guard all resources in very patchy landscapes. These non-consumptive effects are also seen with the shift away from the best quality habitat compared to foraging in a predator-free environment.