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The migratory movements of wild animals can promote abundance and support ecosystem functioning. For large herbivores, mounting evidence suggests that migratory behaviour is an individually variable trait, where individuals can easily switch between migrant and resident tactics. The degree of migratory plasticity, including whether and where to migrate, has important implications for the ecology and conservation of large herbivores in a changing world. Mule deer (Odocoileus hemionus) are an iconic species of western North America, but are notably absent from the body of literature that suggests large herbivore migrations are highly plastic. We evaluated plasticity of migration in female mule deer using longitudinal GPS data collected from 312 individuals across nine populations in the western United States, including 882 animal‐years (801 migrants and 81 residents). We followed both resident and migratory mule deer through time to determine whether individual animals switched migratory behaviours (i.e., whether to migrate) from migratory to residency or vice versa. Additionally, we examined the fidelity of individuals to their migration routes (i.e., where to migrate) to determine whether they used the same routes year after year. We also evaluated whether age and reproductive status affected propensity to migrate or fidelity to migratory routes. Our results indicate that mule deer, unlike other large herbivores, have little or no plasticity in terms of whether or where they migrate. Resident deer remained residents, and migrant deer remained migrants, regardless of age, reproductive status or number of years monitored. Further, migratory individuals showed strong fidelity (>80%) to their migration routes year after year. Our study clearly shows that migration plasticity is not ubiquitous among large herbivores. Because of their rigid migratory behaviour, mule deer may not adapt to changing environmental conditions as readily as large herbivores with more plastic migratory behaviour (e.g., elk). The fixed migratory behaviours of mule deer make clear that conservation efforts aimed at traditional seasonal ranges and migration routes are warranted for sustaining this iconic species that continues to decline across its range. This study reveals that mule deer, unlike many other large herbivores, have little or no plasticity in terms of whether or where they migrate. The authors found resident deer remained residents, and migrant deer remained migrants, regardless of age, reproductive status or number of years monitored.

Partial migration—wherein migratory and non‐migratory individuals exist within the same population—represents a behavioural dimorphism; for it to persist over time, both strategies should yield equal individual fitness. This balance may be maintained through trade‐offs where migrants gain survival benefits by avoiding unfavourable conditions, while residents gain breeding benefits from early access to resources. There has been little overarching quantitative analysis of the evidence for this fitness balance. As migrants—especially long‐distance migrants—may be particularly vulnerable to environmental change, it is possible that recent anthropogenic impacts could drive shifts in fitness balances within these populations. We tested these predictions using a multi‐taxa meta‐analysis. Of 2,939 reviewed studies, 23 contained suitable information for meta‐analysis, yielding 129 effect sizes. Of these, 73% (n = 94) reported higher resident fitness, 22% (n = 28) reported higher migrant fitness, and 5% (n = 7) reported equal fitness. Once weighted for precision, we found balanced fitness benefits across the entire dataset, but a consistently higher fitness of residents over migrants in birds and herpetofauna (the best‐sampled groups). Residency benefits were generally associated with survival, not breeding success, and increased with the number of years of data over which effect sizes were calculated, suggesting deviations from fitness parity are not due to sampling artefacts. A pervasive survival benefit to residency documented in recent literature could indicate that increased exposure to threats associated with anthropogenic change faced by migrating individuals may be shifting the relative fitness balance between strategies. Résumé La migration partielle – dans laquelle les migrants et les résidents existent dans une seul population – représente un dimorphisme comportemental; pour qu'elles persiste au fil du temps, les deux stratégies doivent conférer la même valeur adaptive (fitness) individuelle. Cet équilibre peut être maintenu par un compromis où les migrants obtiennent des bénéfices de survie en évitant des conditions défavorables, tandis que les résidents obtiennent des avantages reproductifs grâce á un accès précoce aux ressources. Peu d'analyses quantitatives ont été entreprise sur les preuves de cet équilibre de fitness. Étant donné que les migrants – en particulier les migrants de longue distance – peuvent être particulièrement vulnérables aux changements environnementaux, il est possible que les impacts anthropiques récents puissent entraîner une modification dans l’équilibre de fitness au sein de ces populations. Nous avons testé ces prédictions en utilisant une méta‐analyse multi‐taxa. Sur 2,939 études examinées, 23 contenaient des informations appropriées pour la méta‐analyse, donnant 129 tailles d'effet. Parmi eux, 73% (n = 94) ont déclaré un fitness supérieur pour les résidents, 22% (n = 28) un fitness supérieur pour les migrants, et 5% (n = 7) une égalité de fitness entre les deux. Après avoir pondéré les données par la précision, nous avons trouvé des avantages équilibrés sur le fitness sur l'ensemble des données, mais parmi les oiseaux et l'herpétofaune (les groupes les mieux échantillonnés), de manière cohérente, les résidents étaient associés à un meilleur fitness comparé à celle des migrants. Les bénéfices d'être résident étaient associés à la survie, et non au succès reproductif, et augmente avec le nombre d'années de données pendant lesquelles la taille d'effet a été calculé; cela suggérerait que les écarts par rapport à la parité de fitness ne sont pas dus à des artefacts d'échantillonnage. Cet avantage de survie pour les résidents, documenté dans la littérature récente, pourrait indiquer qu'une exposition accrue aux menaces associées aux changements anthropiques rencontrés par les individus migrateurs pourrait modifier l'équilibre entre les stratégies. The authors provide the first quantitative analysis of relative fitness in partially migratory populations. Contrary to existing assumptions concerning the evolution and maintenance of migratory behaviours, they find evidence that residency confers greater fitness than migration for certain taxonomic groups and is associated with a benefit to survival, not breeding success.

Migratory animals are predicted to enhance lifetime fitness by obtaining higher quality forage and/or reducing predation risk compared to non‐migratory conspecifics. Despite evidence for behavioural flexibility in other taxa, previous research on large mammals has often assumed that migratory behaviour is a fixed behavioural trait. Migratory behaviour may be plastic for many species, although few studies have tested for individual‐level flexibility using long‐term monitoring of marked individuals, especially in large mammals such as ungulates. We tested variability in individual migratory behaviour using a 10‐year telemetry data set of 223 adult female elk (Cervus elaphus) in the partially migratory Ya Ha Tinda population in Alberta, Canada. We used net squared displacement (NSD) to classify migratory strategy for each individual elk‐year. Individuals switched between migrant and resident strategies at a mean rate of 15% per year, and migrants were more likely to switch than residents. We then tested how extrinsic (climate, elk/wolf abundance) and intrinsic (age) factors affected the probability of migrating, and, secondly, the decision to switch between migratory strategies. Over 630 individual elk‐years, the probability of an individual elk migrating increased following a severe winter, in years of higher wolf abundance, and with increasing age. At an individual elk level, we observed 148 switching events of 430 possible transitions in elk monitored at least 2 years. We found switching was density‐dependent, where migrants switched to a resident strategy at low elk abundance, but residents switched more to a migrant strategy at high elk abundance. Precipitation during the previous summer had a weak carryover effect, with migrants switching slightly more following wetter summers, whereas residents showed the opposite pattern. Older migrant elk rarely switched, whereas resident elk switched more frequently to migrate at older ages. Our results show migratory behaviour in ungulates is an individually variable trait that can respond to intrinsic, environmental and density‐dependent forces. Different strategies had opposing responses to density‐dependent and intrinsic drivers, providing a stabilizing mechanism for the maintenance of partial migration and demographic fitness in this population.

Understanding the causes and consequences of animal movements is of fundamental biological interest because any alteration in movement can have direct and indirect effects on ecosystem structure and function. It is also crucial for assisting spatial wildlife management under variable environmental change scenarios. Recent research has highlighted the need of quantifying individual variability in movement behavior and how it is generated by interactions between individual requirements and environmental conditions, to understand the emergence of population-level patterns. Using a multi-annual movement data set of 213 individual moose ( Alces alces ) across a latitudinal gradient (from 56° to 67° N) that spans over 1100 km of varying environmental conditions, we analyze the differences in individual and population-level movements. We tested the effect of climate, risk, and human presence in the landscape on moose movements. The variation in these factors explained the existence of multiple movements (migration, nomadism, dispersal, sedentary) among individuals and seven populations. Population differences were primarily related to latitudinal variation in snow depth and road density. Individuals showed both fixed and flexible behaviors across years, and were less likely to migrate with age in interaction with snow and roads. For the predominant movement strategy, migration, the distance, timing, and duration at all latitudes varied between years. Males traveled longer distances and began migrating later in spring than females. Our study provides strong quantitative evidence for the dynamics of animal movements in response to changes in environmental conditions along with varying risk from human influence across the landscape. For moose, given its wide distributional range, changes in the distribution and migratory behavior are expected under future warming scenarios.

Altitudinal bird migration involves annual seasonal movements up and down elevational gradients. Despite the fact that species from montane avifaunas worldwide engage in altitudinal migration, the patterns, causes, and prevalence of these movements are poorly understood. This is particularly true in North America where the overwhelming majority of avian migration research has focused on obligate, long-distance, temperate–tropical movements. Elsewhere in the world, most altitudinal migrants are partial migrants, making downhill movements to nonbreeding areas. However, spatial and temporal patterns, the prevalence and predictability of migration at individual and population levels, and the ultimate ecological factors selecting for movement behavior vary considerably among taxa and regions. I conducted a systematic survey of the evidence for altitudinal migration to fill gaps in our understanding of this behavior among the landbirds of North America and Hawaii. Altitudinal migration was as prevalent as in other avifaunas, occurring in >20% of continental North American and nearly 30% of Hawaiian species. Of the species wintering within the USA and Canada, ∼30% engage in altitudinal migrations. Altitudinal migrants are far more common in the West, are taxonomically and ecologically diverse, and North American species exhibit patterns similar to altitudinal migrants elsewhere in the world. Because altitudinal migration systems are relatively tractable, they present excellent opportunities for testing hypotheses regarding migration generally. Altitudinal migration has likely been overlooked in North America due to contingency in the history of ornithological research. Our need to understand the patterns and causes of altitudinal migrations has never been greater due to emerging environmental threats to montane systems.

Knowledge of the relative importance of genetic versus environmental determinants of major developmental transitions is pertinent to understanding phenotypic evolution. In salmonid fishes, a major developmental transition enables a risky seaward migration that provides access to feed resources. In Atlantic salmon, initiation of the migrant phenotype, and thus age of migrants, is presumably controlled via thresholds of a quantitative liability, approximated by body size expressed long before the migration. However, how well size approximates liability, both genetically and environmentally, remains uncertain. We studied 32 Atlantic salmon families in two temperatures and feeding regimes (fully fed, temporarily restricted) to completion of migration status at age 1 year. We detected a lower migrant probability in the cold (0.42) than the warm environment (0.76), but no effects of male maturation status or feed restriction. By contrast, body length in late summer predicted migrant probability and its control reduced migrant probability heritability by 50–70%. Furthermore, migrant probability and length showed high heritabilities and between-environment genetic correlations, and were phenotypically highly correlated with stronger genetic than environmental contributions. Altogether, quantitative estimates for the genetic and environmental effects predicting the migrant phenotype indicate, for a given temperature, a larger importance of genetic than environmental size effects.

Migratory species are widespread in terrestrial, aquatic and aerial environments, and are important both ecologically and economically. Since migration is an adaptive response to particular conditions, environmental changes (climate or otherwise) will potentially alter the selective pressures on movement behavior. Such changes may also interfere with, or disrupt, a species’ ability to migrate. In either case, environmental changes could lead to the reduction or total loss of a migration, yet we have little understanding of when to expect these outcomes to occur. Here, I argue that an understanding of both the proximate and ultimate drivers of migration is needed if we are to predict the fate of migrations under changing environmental conditions. I review what is currently known about the drivers of animal migration, but show that we also need a more complete synthesis of migratory patterns across diverse ecosystems and taxonomic groups. The current understanding of migration indicates that (1) drivers of migration vary across species and ecosystems, and (2) a species’ ability to adapt to environmental change successfully depends in part on its migration drivers. Together, these findings suggest a way forward for studying and generating predictions of how changing environmental conditions will differentially impact species by taxonomic group and geographic region of the world.

Conservation of migration requires information on behavior and environmental determinants. The spatial distribution of forage resources, which migration exploits, often are altered and may have subtle, unintended consequences. Supplemental feeding is a common management practice, particularly for ungulates in North America and Europe, and carryover effects on behavior of this anthropogenic manipulation of forage are expected in theory, but have received limited empirical evaluation, particularly regarding effects on migration. We used global positioning system (GPS) data to evaluate the influence of winter feeding on migration behavior of 219 adult female elk ( Cervus elaphus ) from 18 fed ranges and 4 unfed ranges in western Wyoming. Principal component analysis revealed that the migratory behavior of fed and unfed elk differed in distance migrated, and the timing of arrival to, duration on, and departure from summer range. Fed elk migrated 19.2 km less, spent 11 more days on stopover sites, arrived to summer range 5 days later, resided on summer range 26 fewer days, and departed in the autumn 10 days earlier than unfed elk. Time-to-event models indicated that differences in migratory behavior between fed and unfed elk were caused by altered sensitivity to the environmental drivers of migration. In spring, unfed elk migrated following plant green-up closely, whereas fed elk departed the feedground but lingered on transitional range, thereby delaying their arrival to summer range. In autumn, fed elk were more responsive to low temperatures and precipitation events, causing earlier departure from summer range than unfed elk. Overall, supplemental feeding disconnected migration by fed elk from spring green-up and decreased time spent on summer range, thereby reducing access to quality forage. Our findings suggest that ungulate migration can be substantially altered by changes to the spatial distribution of resources, including those of anthropogenic origin, and that management practices applied in one season may have unintended behavioral consequences in subsequent seasons.

1. Population structure, distribution, abundance and dispersal arguably underpin the entire field of animal ecology, with consequences for regional species persistence, and provision of ecosystem services. Divergent migration behaviours among individuals or among populations are an important aspect of the ecology of highly mobile animals, allowing populations to exploit spatially or temporally distributed food and space resources. 2. This study investigated the spatial ecology of lake sturgeon (Acipenser fulvescens) within the barrier free Huron-Erie Corridor (HEC), which connects Lake Huron and Lake Erie of the North American Laurentian Great Lakes. 3. Over 6 years (2011-2016), movements of 268 lake sturgeon in the HEC were continuously monitored across the Great Lakes using acoustic telemetry (10 years battery life acoustic transmitters). Five distinct migration behaviours were identified with hierarchical cluster analysis, based on the phenology and duration of river and lake use. 4. Lake sturgeon in the HEC were found to contain a high level of intraspecific divergent migration, including partial migration with the existence of residents. Specific behaviours included year-round river residency and multiple lake-migrant behaviours that involved movements between lakes and rivers. Over 85% of individuals were assigned to migration behaviours as movements were consistently repeated over the study, which suggested migration behaviours were consistent and persistent in lake sturgeon. Differential use of specific rivers or lakes by acoustic-tagged lake sturgeon further subdivided individuals into 14 \"contingents\" (spatiotemporally segregated subgroups). 5. Contingents associated with one river (Detroit or St. Clair) were rarely detected in the other river, which confirmed that lake sturgeon in the Detroit and St. Clair represent two semi-independent populations that could require separate management consideration for their conservation. The distribution of migration behaviours did not vary between populations, sexes, body size or among release locations, which indicated that intrapopulation variability in migratory behaviour is a general feature of the spatial ecology of lake sturgeon in unfragmented landscapes.

The most common framework under which ungulate migration is studied predicts that it is driven by spatio-temporal variation in plant phenology, yet other hypotheses may explain differences within and between species. To disentangle more complex patterns than those based on single species/ single populations, we quantified migration variability using two sympatric ungulate species differing in their foraging strategy, mating system and physiological constraints due to body size. We related observed variation to a set of hypotheses. We used GPS-collar data from 537 individuals in 10 roe Capreolus capreolus and 12 red deer Cervus elaphus populations spanning environmental gradients across Europe to assess variation in migration propensity, distance and timing. Using time-to-event models, we explored how the probability of migration varied in relation to sex, landscape (e.g. topography, forest cover) and temporally-varying environmental factors (e.g. plant green-up, snow cover). Migration propensity varied acrossstudy areas. Red deer were, on average, three times more migratory than roe deer (56% versus 18%). This relationship was mainly driven by red deer males which were twice as migratory as females (82% versus 38%). The probability of roe deer migration was similar between sexes. Roe deer (both sexes) migrated earliest in spring. While territorial male roe deer migrated last in autumn, male and female red deer migrated around the same time in autumn, likely due to their polygynous mating system. Plant productivity determined the onset of spring migration in both species, but if plant productivity on winter ranges was sufficiently high, roe deer were less likely to leave. In autumn, migration coincided with reduced plant productivity for both species. This relationship was stronger for red deer. Our results confirm that ungulate migration is influenced by plant phenology, but in a novel way, that these effects appear to be modulated by species-specific traits, especially mating strategies.