Why fly the extra mile? Using stress biomarkers to assess wintering habitat quality in migratory shorebirds

Migratory birds make decisions about how far to travel based on cost-benefit trade-offs. However, in many cases the net effect of these trade-offs is unclear. We sought to address this question by measuring feather corticosterone (CORT_f), leucocyte profile, avian malaria parasite prevalence and estimating fueling rates in three spatially segregated wintering populations of the migratory shorebird ruddy turnstone Arenaria interpres during their stay in the winter habitat. These birds fly from the high-Arctic breeding ground to Australia, but differ in that some decide to end their migration early (Broome, Western Australia), whereas others travel further to either South Australia or Tasmania. We hypothesized that the extra costs in birds migrating greater distances and overwintering in colder climates would be offset by benefits when reaching their destination. This would be evidenced by lower stress biomarkers in populations that travel further, owing to the expected benefits of greater resources and improved vitality. We show that avian malaria prevalence and physiological stress levels were lower in birds flying to South Australia and Tasmania than those overwintering in Broome. Furthermore, our modeling predicts that birds in the southernmost locations enjoy higher fueling rates. Our data are consistent with the interpretation that birds occupying more costly wintering locations in terms of higher migratory flight and thermoregulatory costs are compensated by better feeding conditions and lower blood parasite infections, which facilitates timely and speedy migration back to the breeding ground. These data contribute to our understanding of cost-benefit trade-offs in the decision making underlying migratory behaviour.     

Evaluating the reproductive cost of migration for females in a partially migrating pond-breeding amphibian

Populations of animals with resident and migratory individuals provide an ideal system for addressing questions concerning the evolution of migration. Partially migratory populations may persist because residents and migrants have equal fitness or because migration is based on conditional asymmetries. Studies measuring the costs and benefits of migration provide empirical data to test hypotheses concerning the maintenance of partial migration within a population. In this study, we measured the reproductive differences between resident and migrant females in a pond-breeding amphibian, the red-spotted newt Notophthalmus viridescens . We used large field enclosures to repeatedly sample egg laying over the prolonged breeding season of this species. Resident females did not lay a greater number of eggs or begin laying eggs earlier, despite beginning the breeding season earlier and having a higher mass than migrant newts. The only difference in reproduction we detected was that the eggs of resident females hatched into larger larvae compared with the larvae of migrant females. We discuss this result in the context of other potential trade-offs between residency and migration. This study illustrates the phenology of egg laying of N. viridescens and our results contribute to understanding the population dynamics of partially migratory species.     

Migratory costs and the evolution of egg size and number in introduced and indigenous salmon populations

The trade-off between reproductive investment and migration should be an important factor shaping the evolution of life-history traits among populations following their radiation into habitats with different migratory costs and benefits. An experimentally induced difference in migratory rigor for families of chinook salmon (Oncorhynchus tshawytscha), of approximately 86 km and 413 m elevation, exacted a cost to somatic energy reserves (approximately 17% reduction in metabolizable mass) and ovarian investment (13.7% reduction in ovarian mass). This cost was associated with a reduction in egg size and paralleled the phenotypic pattern of divergence between two introduced New Zealand populations of common origin, presently breeding at sites with different migration distances. The genetic pattern of divergence of these same populations, detected under common rearing, was consistent with compensation for migratory costs (the population that migrates farther invested more in ovarian mass), but egg number more than egg size was associated with this evolution. These evolutionary patterns are consistent with what is known of the inheritance of these traits and with trade-offs and constraints favoring initial evolution in offspring number over offspring size. Analysis of egg number-size patterns of other Pacific salmon populations in their native range supported the hypothesis that migration strongly influences patterns of reproductive allocation, favoring a higher ratio of egg number to egg size with greater migration distance.     

Winter rainfall predicts phenology in widely separated populations of a migrant songbird

Climate change is affecting behaviour and phenology in many animals. In migratory birds, weather patterns both at breeding and at non-breeding sites can influence the timing of spring migration and breeding. However, variation in responses to weather across a species range has rarely been studied, particularly among populations that may winter in different locations. We used prior knowledge of migratory connectivity to test the influence of weather from predicted non-breeding sites on bird phenology in two breeding populations of a long-distance migratory bird species separated by 3,000 km. We found that winter rainfall showed similar associations with arrival and egg-laying dates in separate breeding populations on an east–west axis: greater rainfall in Jamaica and eastern Mexico was generally associated with advanced American redstart (Setophaga ruticilla) phenology in Ontario and Alberta, respectively. In Ontario, these patterns of response could largely be explained by changes in the behaviour of individual birds, i.e., phenotypic plasticity. By explicitly incorporating migratory connectivity into responses to climate, our data suggest that widely separated breeding populations can show independent and geographically specific associations with changing weather conditions. The tendency of individuals to delay migration and breeding following dry winters could result in population declines due to predicted drying trends in tropical areas and the tight linkage between early arrival/breeding and reproductive success in long-distance migrants.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

Weather conditions promote route flexibility during open ocean crossing in a long-distance migratory raptor

Weather conditions are paramount in shaping birds’ migratory routes, promoting the evolution of behavioural plasticity and allowing for adaptive decisions on when to depart or stop during migration. Here, we describe and analyze the influence of weather conditions in shaping the sea-crossing stage of the pre-breeding journey made by a long-distance migratory bird, the Eleonora’s falcon (Falco eleonorae), tracked by satellite telemetry from the wintering grounds in the Southern Hemisphere to the breeding sites in the Northern Hemisphere. As far as we know, the data presented here are the first report of repeated oceanic journeys of the same individuals in consecutive years. Our results show inter-annual variability in the routes followed by Eleonora’s falcons when crossing the Strait of Mozambique, between Madagascar and eastern continental Africa. Interestingly, our observations illustrate that individuals show high behavioural plasticity and are able to change their migration route from one year to another in response to weather conditions, thus minimising the risk of long ocean crossing by selecting winds blowing towards Africa for departure and changing the routes to avoid low pressure areas en route. Our results suggest that weather conditions can really act as obstacles during migration, and thus, besides ecological barriers, the migratory behaviour of birds could also be shaped by “meteorological barriers”. We briefly discuss orientation mechanisms used for navigation. Since environmental conditions during migration could cause carry-over effects, we consider that forecasting how global changes of weather patterns will shape the behaviour of migratory birds is of the utmost importance.     

Timing of breeding carries over to influence migratory departure in a songbird: an automated radiotracking study

1. Determining how events interact across stages of the annual cycle is critical for understanding the factors that affect individual fitness. However, there is currently little information detailing how breeding events influence migratory behaviour. 2. Using an automated digital telemetry array and an isolated island-breeding population of Savannah sparrows Passerculus sandwichensis, we provide the first direct evidence that the timing of breeding events carries over to influence the timing of migration in a songbird and assess for the first time how weather conditions on the breeding grounds also affect departure dates. 3. Date of migratory departure between September and October was strongly influenced by date of breeding completion in adults and fledging date in juveniles from June to July. 4. With respect to weather, adults departed during the first half of high-pressure systems, while juveniles departed throughout the entirety of high-pressure systems (including rainy evenings on the western edge of systems). 5. By combining both ecological and weather data, we could explain almost all variation in departure date for adults (95%), but weather conditions were not a good predictor of departure date for juveniles. 6. Our results provide strong evidence that the timing of breeding events is an important driver of migration timing and that exact departure dates are fine-tuned according to local weather conditions in adults, but not in juveniles.     

Size and accumulation of fuel reserves at stopover predict nocturnal restlessness in a migratory bird

Early arrival at the breeding site positively affects the breeding success of migratory birds. During migration, birds spend most of their time at stopovers. Therefore, determining which factors shape stopover duration is essential to our understanding of avian migration. Because the main purpose of stopover is to accumulate fat as fuel for the next flight bout, fuel reserves at arrival and the accumulation of fuel are both expected to affect stopover departure decisions. Here, we determined whether fuel reserves and fuel accumulation predict a bird''s motivation to depart, as quantified by nocturnal migratory restlessness (Zugunruhe), using northern wheatears (Oenanthe oenanthe) that were captured and temporarily contained at spring stopover. We found that fuel reserves at capture were positively correlated with Zugunruhe, and negatively correlated with fuel accumulation. This indicates that fat birds were motivated to depart, whereas lean birds were set on staying and accumulating fuel. Moreover, the change in fuel reserves was positively correlated with the concurrent change in Zugunruhe, providing the first empirical evidence for a direct link between fuel accumulation and Zugunruhe during stopover. Our study indicates that, together with innate rhythms and weather, the size and accumulation of fuel reserves shape stopover duration, and hence overall migration time.     

Weather-related mass-mortality events in migrants

A major perceived cost of migration in birds is the associated mortality. This mortality has proved difficult to measure and separate from mortality during stationary periods of the annual cycle. This paper reviews some major recorded mortality incidents among migratory birds attributed to inclement weather and other factors, including: (1) in‐flight losses, caused by storms and other adverse weather en route, usually over water; (2) unseasonable cold weather soon after arrival in breeding areas; and (3) unseasonable cold weather before departure from breeding areas. Cold weather often kills migrants in their breeding areas, but not the local resident species which can better withstand it at those times. For migrants, cold and snow act to cut off the food supply, and can have a major selective effect on the seasonal timing of migration. Records of in‐flight weather‐induced mortality, involving up to hundreds or thousands of birds at a time, have affected mainly small passerines, but also larger birds, including eagles and swans. Most occurred in conditions of mist, rain or snow storms, and some involved nocturnal collisions with illuminated masts and other tall structures. Records of post‐arrival mortality in breeding areas have involved mainly small insectivores (especially hirundines), but also waders and waterfowl. Such incidents, associated with cold and snow, have reduced local breeding densities from the previous year by 25–90%, depending on species and area, with up to several years required for recovery. Records of pre‐departure mortality on breeding areas have mainly affected hirundines. Two major incidents in central Europe in September 1931 and 1974 killed hundreds of thousands, or even millions, of swallows and martins. After the latter incident, House Martin Delichon urbicum populations in Switzerland the following year were reduced by an estimated 25–30%. Such climatic extremes that occurred in spring or late summer in particular parts of the breeding range have been recorded at approximate mean frequencies of 2–10 per century. Average daily mortality in many bird species can clearly be much greater during migration periods than during stationary periods. Despite the heavy losses of birds on migration, it may be assumed that migration persists in the long term because the fitness costs (in terms of associated mortality) are more than offset by the fitness benefits (in terms of improved overall survival and breeding success) that accrue from migration.     

Investigating connectivity and seasonal differences in wind assistance in the migration of Common Sandpipers

Many migratory bird species have undergone recent population declines, but there is considerable variation in trends between species and between populations employing different migratory routes. Understanding species-specific migratory behaviours is therefore of critical importance for their conservation. The Common Sandpiper Actitis hypoleucos is an Afro-Palaearctic migratory bird species whose European populations are in decline. We fitted geolocators to individuals breeding in England or wintering in Senegal to determine their migration routes and breeding or non-breeding locations. We used these geolocator data in combination with previously published data from Scottish breeding birds to determine the distributions and migratory connectivity of breeding (English and Scottish) and wintering (Senegalese) populations of the Common Sandpiper, and used simulated random migrations to investigate wind assistance during autumn and spring migration. We revealed that the Common Sandpipers tagged in England spent the winter in West Africa, and that at least some birds wintering in Senegal bred in Scandinavia; this provides insights into the links between European breeding populations and their wintering grounds. Furthermore, birds tagged in England, Scotland and Senegal overlapped considerably in their migration routes and wintering locations, meaning that local breeding populations could be buffered against habitat change, but susceptible to large-scale environmental changes. These findings also suggest that contrasting population trends in England and Scotland are unlikely to be the result of population-specific migration routes and wintering regions. Finally, we found that birds used wind to facilitate their migration in autumn, but less so in spring, when the wind costs associated with their migrations were higher than expected at random. This was despite the wind costs of simulated migrations being significantly lower in spring than in autumn. Indeed, theory suggests that individuals are under greater time pressures in spring than in autumn because of the time constraints associated with reproduction.     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.     

Why do earlier‐arriving migratory birds have better breeding success?

In migratory birds, early arrival on breeding sites is typically associated with greater breeding success, but the mechanisms driving these benefits are rarely known. One mechanism through which greater breeding success among early arrivers can potentially be achieved is the increased time available for replacement clutches following nest loss. However, the contribution of replacement clutches to breeding success will depend on seasonal variation in nest survival rates, and the consequences for juvenile recruitment of hatching at different times in the season. In particular, lower recruitment rates of late‐hatched chicks could offset the benefits to early arrivers of being able to lay replacement clutches, which would reduce the likelihood of replacement clutch opportunities influencing selection on migratory timings. Using a simulation model of time‐constrained capacity for replacement clutches, paramaterized with empirically‐derived estimates from avian migratory systems, we show that greater reproductive success among early‐arriving individuals can arise solely through the greater time capacity for replacement clutches among early arrivers, even when later renesting attempts contribute fewer recruits to the population. However, these relationships vary depending on the seasonal pattern of nest survival. The benefits of early arrival are greatest when nest survival rates are constant or decline seasonally, and early arrival is least beneficial when nest success rates increase over the breeding season, although replacement clutches can mitigate this effect. The time benefits of early arrival facilitating replacement clutches following nest loss may therefore be an important but overlooked source of selection on migratory timings. Empirical measures of seasonal variation in nest survival, renesting, and juvenile recruitment rates are therefore needed in order to identify the costs and benefits associated with individual migration phenology, the selection pressures influencing migratory timings, and the implications for ongoing shifts in migration and breeding phenology.     

Flight responses by a migratory soaring raptor to changing meteorological conditions

Soaring birds that undertake long-distance migration should develop strategies to minimize the energetic costs of endurance flight. This is relevant because condition upon completion of migration has direct consequences for fecundity, fitness and thus, demography. Therefore, strong evolutionary pressures are expected for energy minimization tactics linked to weather and topography. Importantly, the minute-by-minute mechanisms birds use to subsidize migration in variable weather are largely unknown, in large part because of the technological limitations in studying detailed long-distance bird flight. Here, we show golden eagle (Aquila chrysaetos) migratory response to changing meteorological conditions as monitored by high-resolution telemetry. In contrast to expectations, responses to meteorological variability were stereotyped across the 10 individuals studied. Eagles reacted to increased wind speed by using more orographic lift and less thermal lift. Concomitantly, as use of thermals decreased, variation in flight speed and altitude also decreased. These results demonstrate how soaring migrant birds can minimize energetic expenditures, they show the context for avian decisions and choices of specific instantaneous flight mechanisms and they have important implications for design of bird-friendly wind energy.     

Patterns and correlates of songbird movements at an ecological barrier during autumn migration assessed using landscape‐ and regional‐scale automated radiotelemetry

Departure decisions of songbirds at ecological barriers they encounter en route can strongly influence time, energy and survival costs of migration. To date, most field studies of departure decisions and their correlates have used indirect methods and followed migrants at a single stopover site, with limited information on what happens to individuals after they depart from the site. We used an automated radiotelemetry array extending 350 km from southwest Nova Scotia to southern Maine to study the migratory and stopover movements of Northern Waterthrushes Parkesia noveboracensis, Red‐eyed Vireos Vireo olivaceus and Yellow‐rumped Warblers Setophaga coronata in relation to fuel load and weather at the northeastern edge of the Gulf of Maine. From the 105 radio‐transmitters we deployed in southwest Nova Scotia, we recorded 42 landscape‐scale stopover flights and 47 migratory flights by 75 individuals. Of the migratory flights, 57% were orientated southwest, a trajectory that, if held, would require individuals to complete a 350–440 km overwater flight. The remaining 43% of migratory flights were orientated northwest, away from the Gulf of Maine, and 15 individuals were confirmed to have detoured around all or a portion of the barrier, as evidenced by their being re‐detected over the Bay of Fundy and/or along the coast of Maine between 4 h and 15 days later. Across all individuals, initial fat score had a positive effect on departure probability, especially for individuals that made stopover flights. Among weather variables, tailwind assistance was the best predictor of migratory departures but did not appear to be the main factor determining whether individuals orientated towards or away from the Gulf of Maine. Weather had little effect on departure decisions of individuals that made stopover flights. These differences in the correlates of migratory departures and stopover flights would probably not have been distinguishable had our study been restricted to a local scale. Therefore, our findings highlight the importance of expanding the scale at which departure decisions and the ecology of stopover in general are studied.     

What triggers facultative winter migration of Great Bustard (Otis tarda) in Central Europe?

The Great Bustard is on the brink of extinction in Central Europe. Its population is known to suffer high mortality during hard winters, particularly when severe weather conditions cause migration. Long-term winter food management in two populations in Germany did not prevent migration events. To identify migration-triggering factors we tested the potential influence of snow, temperature, phase of winter and development of a tradition of migration. Comparing migratory behaviour with long-term local weather records, we found that snow cover is a much stronger trigger for migration than frost and low temperatures. We conclude that snow heavily affects the Great Bustard’s energy balance mediated not only by limited food access but also by the particular properties of its plumage. This could explain migration events despite food availability and is consistent with our results concerning a tendency for females to undertake facultative winter migration more than males. Available data are currently insufficient to confirm or reject the idea that Great Bustard populations develop a tradition of migratory behaviour following a previous winter migration, and we found no evidence for a decrease in the disposition of the Great Bustard to migrate during the course of winter.     

Population dynamics in migratory networks

Migratory animals are comprised of a complex series of interconnected breeding and nonbreeding populations. Because individuals in any given population can arrive from a variety of sites the previous season, predicting how different populations will respond to environmental change can be challenging. In this study, we develop a population model composed of a network of breeding and wintering sites to show how habitat loss affects patterns of connectivity and species abundance. When the costs of migration are evenly distributed, habitat loss at a single site can increase the degree of connectivity (mixing) within the entire network, which then acts to buffer global populations from declines. However, the degree to which populations are buffered depends on where habitat loss occurs within the network: a site that has the potential to receive individuals from multiple populations in the opposite season will lead to smaller declines than a site that is more isolated. In other cases when there are equal costs of migration to two or more sites in the opposite season, habitat loss can result in some populations becoming segregated (disconnected) from the rest of the network. The geographic structure of the network can have a significant influence on relative population sizes of sites in the same season and can also affect the overall degree of mixing in the network, even when sites are of equal intrinsic quality. When a migratory network is widely spaced and migration costs are high, an equivalent habitat loss will lead to a larger decline in global population size than will occur in a network where the overall costs of migration are low. Our model provides an important foundation to test predictions related to habitat loss in real-world migratory networks and demonstrates that migratory networks will likely produce different dynamics from traditional metapopulations. Our results provide strong evidence that estimating population connectivity is a prerequisite for successfully predicting changes in migratory populations.     

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.     

Migratory costs and contemporary evolution of reproductive allocation in male chinook salmon

Energetically demanding migrations may impact the resources available for reproductive trait development and activity, and hence favour evolution of new investment strategies for remaining resources. We conducted a large-scale experiment to evaluate the proximate cost of migration on male reproductive investment in chinook salmon (Oncorhynchus tshawytscha) and contemporary evolution of reproductive allocation. Experimentally induced differences in migratory costs (17 km inland and 17 m elevation vs. 100 km and 430 m) influenced dorsal hump size and upper jaw length, two traits influencing male mating success that are developed during migration. Longer migration also reduced tissue energy reserves available for competition and length of breeding life. Corresponding shifts in the balance between natural and sexual selection appear to have been responsible for heritable population divergence in secondary sexual trait investment, in approximately 26 generations, following colonization of spawning sites with different migratory demands.     

Theoretical insight into three disease-related benefits of migration

Migration (seasonal round-trip movement across relatively large distances) is common within the animal kingdom. This behaviour often incurs extreme costs in terms of time, energy, and/or survival. Climate, food, predation, and breeding are typically suggested as factors favouring the evolution of migration. Although disease regulation has also been considered, few studies consider it as the primary selective pressure for migration. Our aim was to determine, theoretically, under what conditions migration could reduce the long-term disease prevalence within a population, assuming the only benefits of migration are infection-related. We created two mathematical models, one where the population migrates annually and one where the entire population remains on the breeding ground year-round. In each we simulated disease transmission (frequency-dependent and density-dependent) and quantified eventual disease prevalence. In the migration model we varied the time spent migrating, disease-related migration mortality, and the overall migration mortality. When we compared results from the two models, we found that migration generally lowered disease prevalence. We found a population was healthier if it: (1) spent more time migrating (assuming no disease transmission during migration), (2) had higher disease-induced migration mortality, and (3) had an overall higher mortality when migrating (compared to not migrating). These results provide support for two previously proposed mechanisms by which migration can reduce disease prevalence (migratory escape and migratory cull), and also demonstrate that non-selective mortality during migration is a third mechanism. Our findings indicate that migration may be evolutionarily advantageous even if the only migratory benefit is disease control.