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

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

Body size shapes inter‐specific migratory behaviour: evidence from individual tracks of long‐distance migratory shorebirds

Migration is a common phenomenon across many animal taxa. Understanding how migration scales with body size across species is fundamental in the development of migration theory and in making size‐related predictions. Although aerodynamic theory and ecophysiological scaling laws have assisted greatly in generating such predictions, their verifications have been limited by a lack of empirical data across a range of body sizes. The recent development of ultra‐light tracking devices and its rapid application to migration now allows us to put theory to the test. We used tracking data of seven closely related migratory sandpiper species (family Scolopacidae) along the East Asian‐Australasian Flyway to compare their migratory behaviour when migrating towards the breeding grounds as a function of size (50–750 g). We found a marked decline in migration speed (migration distance divided by total migration duration, including time at stopover sites and in flight) with size. Departure date from the non‐breeding (i.e. wintering) ground and arrival date at the breeding ground also scaled negatively with size. Total migration duration, migration distance, total staging duration (the number of days staying at stopover sites plus days preparing, i.e. fuelling, prior to initial migration) and step length (distance covered within one migratory leg) were not significantly related with size. Correction for phylogeny showed consistent results for all variables. Besides improving our fundamental understanding of inter‐specific variation in migration behaviours, the finding of a clear scaling with size in migration speed and migration timing highlight differential size related capabilities and constraints of migrants. Migratory birds, including sandpipers, are declining on a global scale and particularly along the East Asian‐Australasian Flyway. This notion of size‐dependency in migratory traits may have a bearing on their vulnerability to specific environmental disturbances along their flyways.     

Holding steady: Little change in intensity or timing of bird migration over the Gulf of Mexico

Quantifying the timing and intensity of migratory movements is imperative for understanding impacts of changing landscapes and climates on migratory bird populations. Billions of birds migrate in the Western Hemisphere, but accurately estimating the population size of one migratory species, let alone hundreds, presents numerous obstacles. Here, we quantify the timing, intensity, and distribution of bird migration through one of the largest migration corridors in the Western Hemisphere, the Gulf of Mexico (the Gulf). We further assess whether there have been changes in migration timing or intensity through the Gulf. To achieve this, we integrate citizen science (eBird) observations with 21 years of weather surveillance radar data (1995–2015). We predicted no change in migration timing and a decline in migration intensity across the time series. We estimate that an average of 2.1 billion birds pass through this region each spring en route to Nearctic breeding grounds. Annually, half of these individuals pass through the region in just 18 days, between April 19 and May 7. The western region of the Gulf showed a mean rate of passage 5.4 times higher than the central and eastern regions. We did not detect an overall change in the annual numbers of migrants (2007–2015) or the annual timing of peak migration (1995–2015). However, we found that the earliest seasonal movements through the region occurred significantly earlier over time (1.6 days decade^?1). Additionally, body mass and migration distance explained the magnitude of phenological changes, with the most rapid advances occurring with an assemblage of larger‐bodied shorter‐distance migrants. Our results provide baseline information that can be used to advance our understanding of the developing implications of climate change, urbanization, and energy development for migratory bird populations in North America.     

Long-distance travellers stopover for longer: a case study with spoonbills staying in North Iberia

Long-distance migration is widespread among birds, connecting breeding and wintering areas through a set of stopover localities where individuals refuel and/or rest. The extent of the stopover is critical in determining the migratory strategy of a bird. Here, we examined the relationship between minimum length of stay of PVC-ringed birds in a major stopover site and the remaining flight distance to the overwintering area in the Eurasian spoonbill (Platalea l. leucorodia) during four consecutive autumn migrations. We also analysed the potential effect of timing (arrival date), as well as the role of experience in explaining stopover duration of spoonbills. Overall, birds wintering in Africa, and facing long-distance travel from the stopover site (ca. 3,000 km) stay for longer (2.7 ± 0.4 days) than Iberian winterers (1.5 ± 0.2 days) that perform a much shorter migration (ca. 800 km). These differences were consistent between years. Stopover duration was not significantly affected by the age of the bird. However, there was a significant reduction as migration advanced. Our results suggest that spoonbills develop different stopover strategies depending on the expected distance to the wintering grounds. Adults, especially long-distance migratory ones, could reduce the potential negative effects of density-dependence processes by avoiding stopover at the end of the migration period. These findings are of significant relevance for understanding differences in migratory behaviour within single populations, especially for declining waterbirds, as well as stress the relevance of preserving stopover localities for the conservation of intraspecific diversity in migratory species.     

Identifying critical migratory bottlenecks and high‐use areas for an endangered migratory soaring bird across three continents

Migrant birds face a number of threats throughout their annual cycle, including persecution, collision with energy infrastructure, and habitat and climate change. A key challenge for the conservation of migrants is the identification of important habitat, including migratory concentration areas, because species survival rates may be determined by events in geographically very limited areas. Remote‐tracking technology is facilitating the identification of such critical habitat, although the strategic identification of important sites and incorporation of such knowledge in conservation planning remains limited. We tracked 45 individuals of an endangered, soaring migrant (Egyptian vulture Neophron percnopterus), over 75 complete migrations that traversed three continents along the Red Sea Flyway. We summarize and contextualize migration statistics by season and age class, including migration start, midpoint, and end dates, as well as linear and cumulative migration distance, migration duration and speed, and route straightness. Then, using dynamic Brownian bridge movement models, we quantified space use to identify the most important migratory bottlenecks and high‐use areas on the flyway. These areas each accounted for < 5% of the overall movement range of the tracked birds, yet > 20% of all tracks passed through bottlenecks, and > 50% of the overall vulture time spent on migration fell within high‐use areas. The most important sites were located at the southeastern Red Sea coast and Bab‐el‐Mandeb Strait (Saudi Arabia, Yemen, Djibouti), the Suez Canal zone (Egypt), and the Gulf of Iskenderun (Turkey). Discouragingly however, none of the area within the major migratory bottlenecks was protected and < 13% of the high‐use areas were protected. This demonstrates a very concerning gap in the protected area network for migratory soaring birds along the Red Sea Flyway. Because reducing threats at migratory concentrations can be a very efficient approach to protect populations, our work provides clear guidelines where conservation investment is urgently needed to benefit as many as 35 migratory soaring‐bird species that regularly use the Red Sea Flyway.     

Extreme genetic similarity does not predict non‐breeding distribution of two closely related warblers

Detailed knowledge of migratory connectivity can facilitate effective conservation of Neotropical migrants by helping biologists understand where and when populations may be most limited. We studied the migratory behavior and non‐breeding distribution of two closely related species of conservation concern, the Golden‐winged Warbler (Vermivora chrysoptera) and Blue‐winged Warbler (Vermivora cyanoptera). Although both species have undergone dynamic range shifts and population changes attributed to habitat loss and social interactions promoting competition and hybridization, full life‐cycle conservation planning has been limited by a lack of information about their non‐breeding ecology. Because recent work has demonstrated that the two species are nearly identical genetically, we predicted that individuals from a single breeding population would have similar migratory timing and overwintering locations. In 2015, we placed light‐level geolocators on 25 males of both species and hybrids in an area of breeding sympatry at the Fort Drum Military Installation in Jefferson and Lewis counties, New York. Despite extreme genetic similarity, non‐breeding locations and duration of migration differed among genotypes. Golden‐winged Warblers (N = 2) overwintered > 1900 km southeast of the nearest Blue‐winged Warbler (N = 3) and spent nearly twice as many days in migration; hybrids (N = 2) had intermediate wintering distributions and migratory timing. Spring migration departure dates were staggered based on distance from the breeding area, and all birds arrived at the breeding site within 8 days of each other. Our results show that Golden‐winged Warblers and Blue‐winged Warblers in our study area retain species‐specific non‐breeding locations despite extreme genetic similarity, and suggest that non‐breeding locations and migratory timing vary along a genetic gradient. If the migratory period is limiting for these species, our results also suggest that Golden‐winged Warblers in our study population may be more vulnerable to population decline than Blue‐winged Warblers because they spend almost twice as many days migrating.     

Morphological constraints on changing avian migration phenology

Many organisms at northern latitudes have responded to climate warming by advancing their spring phenology. Birds are known to show earlier timing of spring migration and reproduction in response to warmer springs. However, species show heterogeneous phenological responses to climate warming, with those that have not advanced or have delayed migration phenology experiencing population declines. Although some traits (such as migration distance) partly explain heterogeneity in phenological responses, the factors affecting interspecies differences in the responsiveness to climate warming have yet to be fully explored. In this comparative study, we investigate whether variation in wing aspect ratio (reflecting relative wing narrowness), an ecomorphological trait that is strongly associated with flight efficiency and migratory behaviour, affects the ability to advance timing of spring migration during 1960–2006 in a set of 80 European migratory bird species. Species with larger aspect ratio (longer and narrower wings) showed smaller advancement of timing of spring migration compared to species with smaller aspect ratio (shorter and wider wings) while controlling for phylogeny, migration distance and other life‐history traits. In turn, migration distance positively predicted aspect ratio across species. Hence, species that are better adapted to migration appear to be more constrained in responding phenologically to rapid climate warming by advancing timing of spring migration. Our findings corroborate the idea that aspect ratio is a major evolutionary correlate of migration, and suggest that selection for energetically efficient flights, as reflected by high aspect ratio, may hinder phenotypically plastic/microevolutionary adjustments of migration phenology to ongoing climatic changes.     

Migratory connectivity and population-specific migration routes in a long-distance migratory bird

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu''s harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in sub-Saharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by ‘weak (diffuse) connectivity’. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.     

Repeat tracking of individual songbirds reveals consistent migration timing but flexibility in route

Tracking repeat migratory journeys of individual animals is required to assess phenotypic plasticity of individual migration behaviour in space and time. We used light-level geolocators to track the long-distance journeys of migratory songbirds (wood thrush, Hylocichla mustelina), and, for the first time, repeat journeys of individuals. We compare between- and within-individual variation in migration to examine flexibility of timing and route in spring and autumn. Date of departure from wintering sites in Central America, along with sex and age factors, explained most of the variation (71%) in arrival date at North American breeding sites. Spring migration showed high within-individual repeatability in timing, but not in route. In particular, spring departure dates of individuals were highly repeatable, with a mean difference between years of just 3 days. Autumn migration timing and routes were not repeatable. Our results provide novel evidence of low phenotypic plasticity in timing of spring migration, which may limit the ability of individuals to adjust migration schedules in response to climate change.     

Overtaking on migration: does longer distance migration always incur a penalty?

For many migratory bird species, the latitudinal range of the winter distribution spans thousands of kilometres, thus encompassing considerable variation in individual migration distances. Pressure to winter near breeding areas is thought to be a strong driver of the evolution of migration patterns, as individuals undertaking a shorter migration are generally considered to benefit from earlier arrival on the breeding grounds. However, the influence of migration distance on timing of arrival is difficult to quantify because of the large scales over which individuals must be tracked. Using a unique dataset of individually‐marked Icelandic black‐tailed godwits Limosa limosa islandica tracked throughout the migratory range by a network of hundreds of volunteer observers, we quantify the consequences of migrating different distances for the use of stop‐over sites and timing of arrival in Iceland. Modelling of potential flight distances and tracking of individuals from across the winter range shows that individuals wintering further from the breeding grounds must undertake a stop‐over during spring migration. However, despite travelling twice the distance and undertaking a stop‐over, individuals wintering furthest from the breeding grounds are able to overtake their conspecifics on spring migration and arrive earlier in Iceland. Wintering further from the breeding grounds can therefore be advantageous in migratory species, even when this requires the use of stop‐over sites which lengthen the migratory journey. As early arrival on breeding sites confers advantages for breeding success, the capacity of longer distance migrants to overtake conspecifics is likely to influence the fitness consequences of individual migration strategies. Variation in the quality of wintering and stopover sites throughout the range can therefore outweigh the benefits of wintering close to the breeding grounds, and may be a primary driver of the evolution of specific migration routes and patterns.     

Partial migration in roe deer: migratory and resident tactics are end points of a behavioural gradient determined by ecological factors

Ungulate populations exhibiting partial migration present a unique opportunity to explore the causes of the general phenomenon of migration. The European roe deer Capreolus capreolus is particularly suited for such studies due to a wide distribution range and a high level of ecological plasticity. In this study we undertook a comparative analysis of roe deer GPS location data from a representative set of European ecosystems available within the EURODEER collaborative project. We aimed at evaluating the ecological factors affecting migration tactic (i.e. occurrence) and pattern (i.e. timing, residence time, number of migratory trips). Migration occurrence varied between and within populations and depended on winter severity and topographic variability. Spring migrations were highly synchronous, while the timing of autumn migrations varied widely between regions, individuals and sexes. Overall, roe deer were faithful to their summer ranges, especially males. In the absence of extreme and predictable winter conditions, roe deer seemed to migrate opportunistically, in response to a tradeoff between the costs of residence in spatially separated ranges and the costs of migratory movements. Animals performed numerous trips between winter and summer ranges which depended on factors influencing the costs of movement such as between‐range distance, slope and habitat openness. Our results support the idea that migration encompasses a behavioural continuum, with one‐trip migration and residence as its end points, while commuting and multi‐trip migration with short residence times in seasonal ranges are intermediate tactics. We believe that a full understanding of the variation in tactics of temporal separation in habitat use will provide important insights on migration and the factors that influence its prevalence.     

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.     

The effects of hatching date on timing of autumn migration in partial migrants – an individual approach

Timing of spring migration and breeding and their interaction with climate change has been widely studied in recent years, but the possible changes in timing of autumn migration have gained less attention. This work focuses on autumn migration and provides the first multi‐species individual‐based study of how hatching date affects the autumn migration date and migration age by using nestling ring data and re‐trappings of the same individuals during the autumn migration at the Hanko Bird Observatory, Finland. We studied three potentially multibrooded passerines (great tit, blue tit and coal tit) and two single‐brooded birds of prey (goshawk, sparrowhawk), all partially migratory short‐distance migrants. Individuals from late broods migrated at a younger age in all tit species and also in hawks the late hatched individuals tended to migrate at a younger age than the early‐hatched individuals. Late‐hatched individuals migrated later than early‐hatched individuals in blue and coal tits, where the latest hatchers represented second brood individuals. Based on our results, the time from hatching to autumn migration is not constant even among individuals of the same population. Our study indicates that climate warming induced advancement of avian breeding may cause changes in the timing of autumn migration through the frequency of second broods.     

Light‐level geolocation reveals wintering distribution,migration routes,and primary stopover locations of an endangered long‐distance migratory songbird

The importance of understanding the geographic distribution of the full annual cycle of migratory birds has been increasingly highlighted over the past several decades. However, the difficulty of tracking small birds between breeding and wintering areas has hindered progress in this area. To learn more about Kirtland's warbler Setophaga kirtlandii movement patterns throughout the annual cycle, we deployed archival light‐level geolocators across their breeding range in Michigan. We recovered devices from 27 males and analyzed light‐level data within a Bayesian framework. We found that most males wintered in the central Bahamas and exhibited a loop migration pattern. In both fall and spring, departure date was the strongest predictor of arrival date, but in spring, stopover duration and migration distance were also important. Though stopover strategies varied, males spent the majority of their spring migration at stopover sites, several of which were located just before or after large ecological barriers. We argue that loop migration is likely a response to seasonal variation in prevailing winds. By documenting a tight link between spring departure and arrival dates, we provide a plausible mechanism for previously documented carry‐over effects of winter rainfall on reproductive success in this species. The migratory periods remain the least understood periods for all birds, but by describing Kirtland's warbler migration routes and timing, and identifying locations of stopover sites, we have begun the process of better understanding the dynamics of their full annual cycle. Moreover, we have provided managers with valuable information on which to base future conservation and research priorities.     

Estimating Brownian motion dispersal rate, longevity and population density from spatially explicit mark-recapture data on tropical butterflies

1.?We develop a Bayesian method for analysing mark-recapture data in continuous habitat using a model in which individuals movement paths are Brownian motions, life spans are exponentially distributed and capture events occur at given instants in time if individuals are within a certain attractive distance of the traps. 2.?The joint posterior distribution of the dispersal rate, longevity, trap attraction distances and a number of latent variables representing the unobserved movement paths and time of death of all individuals is computed using Gibbs sampling. 3.?An estimate of absolute local population density is obtained simply by dividing the Poisson counts of individuals captured at given points in time by the estimated total attraction area of all traps. Our approach for estimating population density in continuous habitat avoids the need to define an arbitrary effective trapping area that characterized previous mark-recapture methods in continuous habitat. 4.?We applied our method to estimate spatial demography parameters in nine species of neotropical butterflies. Path analysis of interspecific variation in demographic parameters and mean wing length revealed a simple network of strong causation. Larger wing length increases dispersal rate, which in turn increases trap attraction distance. However, higher dispersal rate also decreases longevity, thus explaining the surprising observation of a negative correlation between wing length and longevity.     

Programmed and flexible: long‐term Zugunruhe data highlight the many axes of variation in avian migratory behaviour

Studies of Zugunruhe – the ‘migratory restlessness’ behaviour of captive birds – have been integral to our understanding of animal migration, revealing an inherited propensity to migrate and an endogenous timing and navigation system. However, differences between Zugunruhe in captivity and migration in the wild call for more data, in particular on variation within and among taxa with diverse migration strategies. Here, we characterise Zugunruhe in a long‐term dataset of activity profiles from stonechats (genus Saxicola) with diverse migratory phenotypes (976 migration periods from 414 birds), using a flexible and consistent quantitative approach based on changepoint analysis. For east African, Austrian, Irish, and Siberian stonechats and hybrids, we report key inter‐population differences in the occurrence, timing, and intensity of Zugunruhe. In line with expectations, we found the highest Zugunruhe intensity in the longest‐distance migrants, more variable patterns in short‐distance migrants, and intermediate characteristics of hybrids relative to their parental groups. Inter‐population differences imply high evolutionary lability of Zugunruhe timing within a robustly structured annual cycle. However, counter to theory, Irish partial migrants showed no segregation between migrant and resident individuals, and previously reported nocturnal restlessness was confirmed for resident African stonechats. Further features of nocturnal restlessness that did not align with migratory behaviour of stonechats were juvenile nocturnal restlessness even prior to postjuvenile moult, and protandry in spring, although stonechats winter in heterosexual pairs. Importantly, Zugunruhe of all populations declined with age, and the intensity of an individual bird's Zugunruhe was correlated with activity levels during other parts of the annual cycle. Our results confirm endogenous, population‐specific migration programmes but also reveal apparent discrepancies between Zugunruhe and migration in the wild. We thus highlight both the continued potential of Zugunruhe study and the need for circumspect interpretation when using migratory restlessness to make inferences about migration in the wild.     

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.     

Variable individual consistency in timing and destination of winter migrating fish

Migration is an important event in the life history of many animals, but there is considerable variation within populations in the timing and final destination. Such differential migration at the population level can be strongly determined by individuals showing different consistencies in migratory traits. By tagging individual cyprinid fish with uniquely coded electronic tags, and recording their winter migrations from lakes to streams for 6 consecutive years, we obtained highly detailed long-term information on the differential migration patterns of individuals. We found that individual migrants showed consistent site fidelities for over-wintering streams over multiple migratory seasons and that they were also consistent in their seasonal timing of migration. Our data also suggest that consistency itself can be considered as an individual trait, with migrants that exhibit consistent site fidelity also showing consistency in migratory timing. The finding of a mixture of both consistent and inconsistent individuals within a population furthers our understanding of intrapopulation variability in migration strategies, and we hypothesize that environmental variation can maintain such different strategies.     

Patterns of change in timing of spring migration in North European songbird populations

From 1976 to 1997 passerines were mist-netted and ringed on the island of Christiansø, in the Baltic Sea. Here we present analyses of phenological changes (i.e. time of arrival) for 25 species based on the entire populations of mist-netted songbirds during spring migration. We used two approaches (least square and quantile regression) to test for changes in arrival time of first individuals and three different parts of the songbird populations (i.e. first 5%, 50% and 95% of the total number of trapped individuals corrected for trapping effort). Our results generally confirm earlier spring arrival of migratory passerines with an overall earlier arrival of 0.26 days per year. Changes in the arrival time of first individuals are often the only data available. They are typically analysed on the assumption that they are representative of their respective population. We found a unidirectional, significant change towards earlier arrival for all four measures of arrival timing which seem to support this. However, the four measures of arrival are changing at different rates. First individuals changed arrival time more rapidly than the first 5%, 50% and 95% of the spring total. Such differences are likely to be important for our understanding of population-dynamic changes in relation to climate change. These differences may also have long-term evolutionary consequences. Migration distance seems to affect the degree of change in arrival time, but we found no difference between species wintering in different regions of Africa.     

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.