Patterns of parasitism in monarch butterflies during the breeding season in eastern North America

1. Migratory behaviour can result in reduced prevalence of pathogens in host populations. Two hypotheses have been proposed to explain this relationship: (i) ‘migratory escape’, where migrants benefit from escaping pathogen accumulation in contaminated environments; and (ii) ‘migratory culling’, where the selective removal of infected individuals occurs during migration. 2. In the host–parasite system between the monarch butterfly (Danaus plexippus Linn.) and its obligate protozoan parasite Ophryocystis elektroscirrha (OE), there is evidence to support both hypotheses, particularly during the monarchs' autumn migration. However, these processes can operate simultaneously and could vary throughout the monarchs' annual migratory cycle. Assessing the relative strength for each hypothesis has not previously been done. 3. To evaluate both hypotheses, parasite infection prevalence was examined in monarchs sampled in eastern North America during April–September, and stable isotopes (δ²H, δ¹³C) were used to estimate natal origin and infer migration distance. There was stronger support for the migratory escape hypothesis, wherein infection prevalence increased over the breeding season and was higher at southern latitudes, where the breeding season tends to be longer compared with northern latitudes. Little support was found for the migratory culling hypothesis, as infection prevalence was similar whether monarchs travelled shorter or longer distances. 4. These results suggest that migration allows individuals to escape parasites not only during the autumn, as shown in previous work, but during the monarchs' spring and summer movements when they recolonise the breeding range. These results imply a potential fitness advantage to monarchs that migrate further north to exploit parasite‐free habitats.     

Associations between host migration and the prevalence of a protozoan parasite in natural populations of adult monarch butterflies

1. Monarch butterflies Danaus plexippus (L.) (Lepidoptera: Nymphalidae) are susceptible to infection by the obligate protozoan parasite Ophryocystis elektroscirrha (McLaughlin and Myers) (Apicomplexa: Neogregarinida). Because monarchs form resident and migratory populations in different parts of the world, this host–parasite system provides the opportunity to examine how variation in parasite prevalence relates to host movement patterns. 2. Parasite prevalence was evaluated using 14 790 adult monarchs captured between 1968 and 1997. Comparison of three populations in North America indicated that parasite prevalence is associated negatively with host dispersal distances. A continuously breeding, nonmigratory population in southern Florida showed high prevalence (over 70% heavily infected). The western population migrates moderate distances to overwintering sites on the Pacific Coast and has intermediate prevalence (30% heavily infected). The eastern migratory population, which travels the longest distance to Mexican overwintering sites, has exhibited less than 8% infection throughout the past 30 years. 3. Variation in parasite loads within North American migratory populations was investigated to determine whether the prevalence of heavy infection and average parasite loads declined during migration or overwintering. Average parasite loads of summer‐breeding adults in western North America decreased with increasing distance from overwintering sites. This suggests that heavily infected monarchs are less likely to remigrate long distances in spring. No differences in the frequency of heavily infected adults were found among eastern or western North American monarchs throughout the overwintering period, however, suggesting that this parasite does not affect overwintering mortality. 4. Changes in the prevalence of monarchs with low parasite loads demonstrate that spore transfer occurs during migration and overwintering, possibly when adult butterflies contact each other as a result of their clustering behaviour. 5. This study of geographical and temporal variation in O. elektroscirrha among populations of D. plexippus demonstrates the potential role of seasonal migration in mediating interactions between hosts and parasites, and suggests several mechanisms through which migratory behaviour may influence parasite prevalence.     

Host immune defence and migration in birds

Migratory birds are exposed to at least two different parasite faunas during their annual cycle, while resident birds only experience a single parasite fauna. Migratory birds should therefore have evolved mechanisms to control or reduce the negative impact of infections from a more diverse parasite fauna. In a comparison of pairs of closely related species of birds that differ with respect to whether they are migratory or residents, the size of two immune defence organs (the bursa of Fabricius and the spleen) was consistently larger in the migratory species. Since the bursa is only found in juvenile, sexually immature birds, we conclude that immune defence adaptations to the impact of a more diverse parasite fauna in migrants already exist before the start of the first migration. Interspecific differences in investment in immune defence between migratory and resident birds have implications for our understanding of complex host–parasite interactions, the acquisition of new hosts by parasites, and the susceptibility of migratory birds to environmental perturbations.     

Breeding phenology determines evolutionary transitions in migratory behaviour in finches and allies

The breeding season of long‐distance migratory birds often starts later and is shorter than in resident or short‐distance species breeding at the same latitude, but the reason for this is unclear. Here we investigate the association between migration distance and breeding phenology in a group of passerine birds, the finches and their allies, using phylogenetic comparative methods. We confirm that migration distance is related to aspects of the species’ breeding phenology after controlling for the effect of potentially confounding variables. Directional phylogenetic analyses suggest that evolutionary transitions in migration distance are determined by the breeding phenology. A relatively long migration distance is more likely to evolve in birds with a late, short breeding season, whereas transitions to short distance migration are more likely to occur in lineages with an early, long breeding season. These results suggest that migration distance is constrained by breeding phenology and not vice versa. Thus, breeding phenology may be an important ultimate factor shaping the evolution of migratory strategies.     

Migratory birds have higher prevalence and richness of avian haemosporidian parasites than residents

Individuals of migratory species may be more likely to become infected by parasites because they cross different regions along their route, thereby being exposed to a wider range of parasites during their annual cycle. Conversely, migration may have a protective effect since migratory behaviour allows hosts to escape environments presenting a high risk of infection. Haemosporidians are one of the best studied, most prevalent and diverse groups of avian parasites, however the impact of avian host migration on infection by these parasites remains controversial. We tested whether migratory behaviour influenced the prevalence and richness of avian haemosporidian parasites among South American birds. We used a dataset comprising ~ 11,000 bird blood samples representing 260 bird species from 63 localities and Bayesian multi-level models to test the impact of migratory behaviour on prevalence and lineage richness of two avian haemosporidian genera (Plasmodium and Haemoproteus). We found that fully migratory species present higher parasite prevalence and higher richness of haemosporidian lineages. However, we found no difference between migratory and non-migratory species when evaluating prevalence separately for Plasmodium and Haemoproteus, or for the richness of Plasmodium lineages. Nevertheless, our results indicate that migratory behaviour is associated with an infection cost, namely a higher prevalence and greater variety of haemosporidian parasites.     

Molecular Detection of Hematozoa Infections in Tundra Swans Relative to Migration Patterns and Ecological Conditions at Breeding Grounds

Tundra swans (Cygnus columbianus) are broadly distributed in North America, use a wide variety of habitats, and exhibit diverse migration strategies. We investigated patterns of hematozoa infection in three populations of tundra swans that breed in Alaska using satellite tracking to infer host movement and molecular techniques to assess the prevalence and genetic diversity of parasites. We evaluated whether migratory patterns and environmental conditions at breeding areas explain the prevalence of blood parasites in migratory birds by contrasting the fit of competing models formulated in an occupancy modeling framework and calculating the detection probability of the top model using Akaike Information Criterion (AIC). We described genetic diversity of blood parasites in each population of swans by calculating the number of unique parasite haplotypes observed. Blood parasite infection was significantly different between populations of Alaska tundra swans, with the highest estimated prevalence occurring among birds occupying breeding areas with lower mean daily wind speeds and higher daily summer temperatures. Models including covariates of wind speed and temperature during summer months at breeding grounds better predicted hematozoa prevalence than those that included annual migration distance or duration. Genetic diversity of blood parasites in populations of tundra swans appeared to be relative to hematozoa prevalence. Our results suggest ecological conditions at breeding grounds may explain differences of hematozoa infection among populations of tundra swans that breed in Alaska.     

Migration distance does not predict blood parasitism in a migratory songbird

Migration can influence host–parasite dynamics in animals by increasing exposure to parasites, by reducing the energy available for immune defense, or by culling of infected individuals. These mechanisms have been demonstrated in several comparative analyses; however, few studies have investigated whether conspecific variation in migration distance may also be related to infection risk. Here, we ask whether autumn migration distance, inferred from stable hydrogen isotope analysis of summer‐grown feathers (δ²H_f) in Europe, correlates with blood parasite prevalence and intensity of infection for willow warblers (Phylloscopus trochilus) wintering in Zambia. We also investigated whether infection was correlated with individual condition (assessed via corticosterone, scaled mass index, and feather quality). We found that 43% of birds were infected with Haemoproteus palloris (lineage WW1). Using generalized linear models, we found no relationship between migration distance and either Haemoproteus infection prevalence or intensity. There was spatial variation in breeding ground origins of infected versus noninfected birds, with infected birds originating from more northern sites than noninfected birds, but this difference translated into only slightly longer estimated migration distances (~214 km) for infected birds. We found no relationship between body condition indices and Haemoproteus infection prevalence or intensity. Our results do not support any of the proposed mechanisms for migration effects on host–parasite dynamics and cautiously suggest that other factors may be more important for determining individual susceptibility to disease in migratory bird species.     

Photoperiodic response may facilitate adaptation to climatic change in long-distance migratory birds

Recent climatic change is causing spring events in northern temperate regions to occur earlier in the year. As a result, migratory birds returning from tropical wintering sites may arrive too late to take full advantage of the food resources on their breeding grounds. Under these conditions, selection will favour earlier spring arrival that could be achieved by overwintering closer to the breeding grounds. However, it is unknown how daylength conditions at higher latitudes will affect the timing of life cycle stages. Here, we show in three species of Palaearctic-African migratory songbirds that a shortening of migration distance induces an advancement of springtime activities. Birds exposed to daylengths simulating migration to and wintering in southern Europe considerably advanced their spring migratory activity and testicular development. This response to the novel photoperiodic environment will enable birds wintering further north to advance spring arrival and to start breeding earlier. Thus, phenotypic flexibility in response to the photoperiod may reinforce selection for shorter migration distance if spring temperatures continue to rise.     

Local parasite lineage sharing in temperate grassland birds provides clues about potential origins of Galapagos avian Plasmodium

Oceanic archipelagos are vulnerable to natural introduction of parasites via migratory birds. Our aim was to characterize the geographic origins of two Plasmodium parasite lineages detected in the Galapagos Islands and in North American breeding bobolinks (Dolichonyx oryzivorus) that regularly stop in Galapagos during migration to their South American overwintering sites. We used samples from a grassland breeding bird assemblage in Nebraska, United States, and parasite DNA sequences from the Galapagos Islands, Ecuador, to compare to global data in a DNA sequence registry. Homologous DNA sequences from parasites detected in bobolinks and more sedentary birds (e.g., brown‐headed cowbirds Molothrus ater, and other co‐occurring bird species resident on the North American breeding grounds) were compared to those recovered in previous studies from global sites. One parasite lineage that matched between Galapagos birds and the migratory bobolink, Plasmodium lineage B, was the most common lineage detected in the global MalAvi database, matching 49 sequences from unique host/site combinations, 41 of which were of South American origin. We did not detect lineage B in brown‐headed cowbirds. The other Galapagos‐bobolink match, Plasmodium lineage C, was identical to two other sequences from birds sampled in California. We detected a close variant of lineage C in brown‐headed cowbirds. Taken together, this pattern suggests that bobolinks became infected with lineage B on the South American end of their migratory range, and with lineage C on the North American breeding grounds. Overall, we detected more parasite lineages in bobolinks than in cowbirds. Galapagos Plasmodium had similar host breadth compared to the non‐Galapagos haemosporidian lineages detected in bobolinks, brown‐headed cowbirds, and other grassland species. This study highlights the utility of global haemosporidian data in the context of migratory bird–parasite connectivity. It is possible that migratory bobolinks bring parasites to the Galapagos and that these parasites originate from different biogeographic regions representing both their breeding and overwintering sites.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

North‐African house martins endure greater haemosporidian infection than their European counterparts

Afro‐Palearctic migrant species are exposed to parasites at both breeding and over‐wintering grounds. The house martin Delichon urbicum is one such migratory species facing high instances of blood parasite infection. In an attempt to determine whether breeding European house martins harbour similar blood parasite communities to populations breeding in North Africa, birds were sampled at their breeding grounds in Switzerland and Algeria. Moreover, haemosporidian prevalence and parasite communities were compared to published data sets on Spanish and Dutch breeding populations. This study furthermore wanted to establish whether co‐infection with multiple genera or lineages of parasites had negative e?ects on host body condition. Breeding house martins caught in Algeria showed a higher prevalence of avian haemosporidian parasites than did European populations. Swiss house martins showed a prevalence comparable to that of Spanish and Dutch populations. There were slight differences in the haemosporidian community between European and North‐African populations in terms of composition and abundance of each lineage. Similar to the Dutch house martins, but in contrast to the Spanish population, infection status and number of genera of parasites infecting single hosts did not in?uence Swiss house martin body condition.     

Migratory diversity predicts population declines in birds

Declines in migratory species are a pressing concern worldwide, but the mechanisms underpinning these declines are not fully understood. We hypothesised that species with greater within‐population variability in migratory movements and destinations, here termed ‘migratory diversity’, might be more resilient to environmental change. To test this, we related map‐based metrics of migratory diversity to recent population trends for 340 European breeding birds. Species that occupy larger non‐breeding ranges relative to breeding, a characteristic we term ‘migratory dispersion’, were less likely to be declining than those with more restricted non‐breeding ranges. Species with partial migration strategies (i.e. overlapping breeding and non‐breeding ranges) were also less likely to be declining than full migrants or full residents, an effect that was independent of migration distance. Recent rates of advancement in Europe‐wide spring arrival date were greater for partial migrants than full migrants, suggesting that migratory diversity may also help facilitate species responses to climate change.     

Parasites hinder monarch butterfly flight: implications for disease spread in migratory hosts

Monarch butterflies (Danaus plexippus) are parasitized by the protozoan Ophryocystis elektroscirrha throughout their geographical range. Monarchs inhabiting seasonally fluctuating environments migrate annually, and parasite prevalence is lower among migratory relative to non‐migratory populations. One explanation for this pattern is that long‐distance migration weeds out infected animals, thus reducing parasite prevalence and transmission between generations. In this study we experimentally infected monarchs from a migratory population and recorded their long‐distance flight performance using a tethered flight mill. Results showed that parasitized butterflies exhibited shorter flight distances, slower flight speeds, and lost proportionately more body mass per km flown. Differences between parasitized and unparasitized monarchs were generally not explained by individual variation in wing size, shape, or wing loading, suggesting that poorer flight performance among parasitized hosts was not directly caused by morphological constraints. Effects of parasite infection on powered flight support a role for long‐distance migration in dramatically reducing parasite prevalence in this and other host–pathogen systems.     

Flying with diverse passengers: greater richness of parasitic nematodes in migratory birds

Environmental changes are simultaneously affecting parasitic diseases and animal migrations, making it important to understand the disease dynamics of migratory species, including their range of infections and investment into defences. There is an urgent need for such knowledge because migratory animals, especially birds, are important for pathogen transmission and also particularly sensitive to environmental changes. Here we compare the nematode species richness and relative immune investment (via relative spleen size) of almost 200 migratory and non‐migratory species within three diverse groups of birds (Anseriformes, Accipitriformes and Turdidae) with worldwide distributions and varied ecology. Our results provide the first large‐scale demonstration that migratory birds face greater challenge from macroparasites as they have significantly dissimilar nematode fauna and higher nematode species richness compared to non‐migratory species. Even though birds with relatively large spleens had more nematode species, there was no difference in relative spleen size between migratory and non‐migratory bird species. The physiological stress of migration can be exacerbated by the potential range of pathologies induced by their richer nematode communities, particularly in combination with environmental perturbations. Altered migration stemming from global changes can also have important consequences for nematode transmission. Synthesis Most studies on parasites of migratory birds versus non‐migratory birds focus upon blood parasites; here we compared the diversity of another important parasite group – nematodes (roundworms) in three orders of birds. We found for any given order, migratory species and species with proportionally larger spleens generally have a wider range of nematodes. It is unclear why migratory species harbour more nematode species. Global climate change is expected to influence both bird migration patterns and infectious diseases, which may increase host susceptibility to parasitism and also introduce diverse nematodes to new areas and potential hosts.     

Cross-species infection of blood parasites between resident and migratory songbirds in Africa

We studied the phylogeny of avian haemosporidian parasites, Haemoproteus and Plasmodium, in a number of African resident and European migratory songbird species sampled during spring and autumn in northern Nigeria. The phylogeny of the parasites was constructed through sequencing part of their mitochondrial cytochrome b gene. We found eight parasite lineages, five Haemoproteus and three Plasmodium, infecting multiple host species. Thus, 44% of the 18 haemospiridian lineages found in this study were detected in more than one host species, indicating that host sharing is a more common feature than previously thought. Furthermore, one of the Plasmodium lineages infected species from different host families, Sylviidae and Ploceidae, expressing exceptionally large host range. We mapped transmission events, e.g. the occurrence of the parasite lineages in resident bird species in Europe or Africa, onto a phylogenetic tree. This yielded three clades, two Plasmodium and one Haemoproteus, in which transmission seems to occur solely in Africa. One Plasmodium clade showed European transmission, whereas the remaining two Haemoproteus clades contained mixes of lineages of African, European or unknown transmission. The mix of areas of transmission in several branches of the phylogenetic tree suggests that transmission of haemosporidian parasites to songbirds has arisen repeatedly in Africa and Europe. Blood parasites could be viewed as a cost of migration, as migratory species in several cases were infected with parasite lineages from African resident species. This cost of migration could have considerable impact on the evolution of migration and patterns of winter distribution in migrating birds.     

Animal migrations and parasitism: reciprocal effects within a unified framework

Migrations, i.e. the recurring, roundtrip movement of animals between distant and distinct habitats, occur among diverse metazoan taxa. Although traditionally linked to avoidance of food shortages, predators or harsh abiotic conditions, there is increasing evidence that parasites may have played a role in the evolution of migration. On the one hand, selective pressures from parasites can favour migratory strategies that allow either avoidance of infections or recovery from them. On the other hand, infected animals incur physiological costs that may limit their migratory abilities, affecting their speed, the timing of their departure or arrival, and/or their condition upon reaching their destination. During migration, reduced immunocompetence as well as exposure to different external conditions and parasite infective stages can influence infection dynamics. Here, we first explore whether parasites represent extra costs for their hosts during migration. We then review how infection dynamics and infection risk are affected by host migration, thereby considering parasites as both causes and consequences of migration. We also evaluate the comparative evidence testing the hypothesis that migratory species harbour a richer parasite fauna than their closest free-living relatives, finding general support for the hypothesis. Then we consider the implications of host migratory behaviour for parasite ecology and evolution, which have received much less attention. Parasites of migratory hosts may achieve much greater spatial dispersal than those of non-migratory hosts, expanding their geographical range, and providing more opportunities for host-switching. Exploiting migratory hosts also exerts pressures on the parasite to adapt its phenology and life-cycle duration, including the timing of major developmental, reproduction and transmission events. Natural selection may even favour parasites that manipulate their host's migratory strategy in ways that can enhance parasite transmission. Finally, we propose a simple integrated framework based on eco-evolutionary feedbacks to consider the reciprocal selection pressures acting on migratory hosts and their parasites. Host migratory strategies and parasite traits evolve in tandem, each acting on the other along two-way causal paths and feedback loops. Their likely adjustments to predicted climate change will be understood best from this coevolutionary perspective.     

The role of migration for spatial turnover of arctic bird species in a circumpolar perspective

Several different factors may determine where species range limits are located within regions of otherwise continuously available habitat and suitable climate. Within the Arctic tundra biome many bird species are migratory and their breeding distributions are affected by migration routes that are in turn limited by factors such as suitable winter habitat, migratory stopover sites, geographical barriers and historical routes of colonization. We identified longitudinal zones in the circumpolar Arctic of pronounced changes in the avian species composition (high species spatial turnover; ‘species divides’). We tested for the association between migratory status and the geographical location and numbers of such species divides for species with non‐breeding habitats mainly within terrestrial, pelagic and coastal ecosystems. Our results demonstrate that migration is of profound importance for both the number and locations of species divides in the Arctic. Long‐distance migration is associated with a large number of divides among terrestrial and coastal arctic birds but with a reduced number of divides among pelagic birds. We suggest that long‐distance migration permits pelagic but not terrestrial and coastal birds to colonize large winter ranges, which in turn causes expansion of breeding ranges, with more homogenous communities and reduction of species divides as consequences, among the long‐distance migrants of pelagic but not of terrestrial and coastal birds. Furthermore, the divides among long‐distance migrants are situated in two main regions, the Beringia and Greenland zones, while divides among short‐distance migrants are more evenly spaced throughout the circumpolar Arctic. The Beringia and Greenland divides result largely from inter‐continental colonization of new breeding ranges but retainment of original winter quarters in a process of evolution through extension of migration routes, leading to aggregated divides in the meeting zones of major global flyways.     

Behavioral and physiological conflicts in migrants: the transition between migration and breeding

Progression of the vernal migratory life history stage to breeding presents a number of apparent behavioral and physiological conflicts. Features that characterize the migratory stage include: high mobility, sociality, repetitive cycles of feeding (hyperphagia and fattening) and migratory flight. Breeding comprises: sedentary, territorial and reproductive behaviors, an initial decline in hyperphagia and reduction of fuel stores. Because morphology, physiology and behavior change, the transition between stages cannot be instantaneous. In many species development of the reproductive system actually occurs during migration, but in others gonadal development may not commence until later. This variation in degree of overlap of migration and reproductive functions is not well understood, but may be related to migratory distance and length of the breeding season, which tends to be shorter at higher latitudes and altitudes. In these habitats, migrants may arrive at their breeding sites to find unpredictable conditions that cannot support breeding. At this juncture, migrants may retreat to refugia and prolong maintenance of facultative migratory functions, termed arrival biology, until conditions improve sufficiently to initiate breeding. In this review, we focus on the Pacific races of the white-crowned sparrow, Zonototrichia leucophyrs, in which the entire spectrum of migratory strategies are represented from resident to long distance migrants and about which much is known. This species presents a unique view of the appearance and variations in arrival biology. Focusing on the juncture between migration and breeding, we discuss the diversity of responses of congeners to a spectrum of environmental conditions that favor survival and reproductive success.     

Variation in haematozoan parasitism at local and landscape levels in the red-billed quelea Quelea quelea

The red-billed quelea Quelea quelea , one of the most abundant birds in the world, presents two fundamental conundrums that we investigate here with a novel approach using blood parasite assemblages at two spatial scales, landscape and individual. The quelea of southern Africa Q. q. lathamii are split by a hypothesized migratory divide, where birds follow rain fronts in one of two directions (NW or SE). This divide is not detectable in the host population using microsatellite data, and here we show that it is also not apparent from our large-scale phylogeographical analyses of the haematozoan parasite. At a finer scale, the colourful and variable breeding plumage of male red-billed quelea has not previously shown a correlation with predictors of quality, as it does in many other bird species. The male's breeding plumage is partially based on carotenoid colouration, the quality of which has been correlated with haematozoan infection in other bird species. However, we found no correlation between intensity of male carotenoid colouration and haematozoan infection. Our results do not contradict the hypothesis that male breeding plumage in this species serves to identify individuals rather than to indicate quality. Finally, we recovered the greatest number of haematozoan lineages from any phylogenetic survey of a single host species to date. Understanding the reasons for the extreme diversity of parasite lineages in this species may assist in explaining the success of the red-billed quelea in anthropogenic landscapes.     

The influence of seasonal migration on range size in temperate North American passerines

Seasonal migration has been alternately proposed to promote geographic range size in some contexts and to constrain it in others, but it remains unclear if migratory behavior has a general effect on range size. Because migration involves movement, most hypotheses about the relationship between migration and range size invoke an influence of migration on the process of dispersal-mediated range expansion. Intuitively, a positive relationship between migratory behavior and dispersal ability could bolster range expansion among migratory species, yet some biogeographic patterns suggest that long-distance migration may instead impede range expansion, especially in the temperate zone. We conducted a comparative analysis of the relationship between migratory behavior and range size by testing the effect of migratory status, migration distance and morphological dispersal ability on breeding range size among all temperate North American passerines. Further, we assessed whether these traits affect range expansion into suitable habitat by analyzing their relationship with range filling (the proportion of climatically-suitable area occupied, or ‘filled’ by a species). Contrary to previous studies, we found migration and dispersal ability to be poor predictors of range size and range filling in North America. Rather, most variation in range size is explained by latitude. Our results suggest that migratory behavior does not affect range size within the scale of a continent, and furthermore, that temperate North American passerines’ breeding ranges are not influenced by their dispersal abilities. To better understand why migratory behavior appears to promote range size in some contexts and constrain it in others, future studies should investigate how migratory behavior affects dispersal at the individual level, as well as the relationship between the evolution of migratory behavior and the breadth of species’ climatic niches.