Is the range size of migratory birds constrained by their migratory program?

Aim Intuitively, species in which the individuals migrate long distances between summer and winter quarters should be more likely to disperse and colonize new breeding areas than resident species. However, it has repeatedly been noted that many bird species with large ranges are residents. This paradox was tested on land birds breeding in the boreal forest of the Palaearctic, the largest uninterrupted stretch of habitat on earth. Methods The longitudinal distribution of two land bird communities on each side of the Eurasian continent, in Scandinavia and eastern Siberia, were used to test whether migratory birds indeed have a lower colonization success than resident birds. Results The migratory species are significantly less likely than resident species to have a range including both regions. The pattern held true even after controlling for latitudinal effects and local abundance, and was also observed at the level of genus and family. Main conclusions The relatively low colonization success of migratory species into new breeding areas may be because these new areas require novel migratory programs (migratory distance, direction and timing) in order for the birds to reach suitable wintering grounds.     

Habitat selection responses of parents to offspring predation risk: an experimental test

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.     

Cooperative breeding among Chinese passerines: inference from social behavior,diet, and migratory status

Recent estimates suggest that 9% of bird species are cooperative breeders. However, little is known about the breeding behavior of many species, particularly those in the Indomalayan and Neotropical regions. Our objective was to provide an overview of the prevalence of cooperative breeding among Chinese songbirds. Examination of the social behavior, diet, and migratory status of 55 known cooperative‐breeding species of songbirds in China revealed that 90.9% live in small groups, 89.1% are residents in at least one or all their subspecies, 81.8% are insectivores, and 14.5% are omnivores. In contrast, 58.2% of the 55 species are resident insectivores that live in small groups, 10.9% are resident omnivores that live in small groups, and 12.7% include subspecies that are resident insectivores. We used these combinations of traits of known cooperative breeders and phylogenetic relationships to infer that an additional 106 species of songbirds in China are probable cooperative breeders and 22 species are possible cooperative breeders. Our analysis suggests that a maximum of 27.2% (183 of 674 species) of Chinese passerines are cooperative breeders, with more occurring in subtropical southern China than in temperate northern China. Cooperative breeding is the main breeding system of species in the families Corvidae, Pycnonotidae, and, especially, Timaliidae (105 of 183 species, 57%). Based on our analyses, cooperative breeding might be more common than previously assumed, particularly among species in the families Timaliidae, Corvidae, and Sturnidae, and species in southern, subtropical China. Because most cooperative‐breeding species in our study were either inferred cooperative breeders or possible cooperative breeders, additional study of these species is needed to confirm our results. A better understanding of the prevalence of cooperative breeding in birds will improve our insight into the evolutionary and ecological factors that select for cooperative breeding.     

Breeding Phenology of Birds: Mechanisms Underlying Seasonal Declines in the Risk of Nest Predation

Seasonal declines in avian clutch size are well documented, but seasonal variation in other reproductive parameters has received less attention. For example, the probability of complete brood mortality typically explains much of the variation in reproductive success and often varies seasonally, but we know little about the underlying cause of that variation. This oversight is surprising given that nest predation influences many other life-history traits and varies throughout the breeding season in many songbirds. To determine the underlying causes of observed seasonal decreases in risk of nest predation, we modeled nest predation of Dusky Flycatchers (Empidonax oberholseri) in northern California as a function of foliage phenology, energetic demand, developmental stage, conspecific nest density, food availability for nest predators, and nest predator abundance. Seasonal variation in the risk of nest predation was not associated with seasonal changes in energetic demand, conspecific nest density, or predator abundance. Instead, seasonal variation in the risk of nest predation was associated with foliage density (early, but not late, in the breeding season) and seasonal changes in food available to nest predators. Supplemental food provided to nest predators resulted in a numerical response by nest predators, increasing the risk of nest predation at nests that were near supplemental feeders. Our results suggest that seasonal changes in foliage density and factors associated with changes in food availability for nest predators are important drivers of temporal patterns in risk of avian nest predation.     

Density‐independent predation affects migrants and residents equally in a declining partially migratory elk population

Migration is expected to benefit individuals through exposure to higher quality forage and reducing predation rates more than non‐migratory conspecifics. Previous studies of partially migratory ungulates (with migrant and resident individuals) have focused on bottom–up factors regulating resident and migrant segments, yet differential predation between strategies could also be a density‐dependent regulatory mechanism. Our study tested for density‐dependence in mortality, as well as mechanisms of ­bottom–up or top–down regulation in the resident and migrant portions of the partially migratory Ya Ha Tinda elk population. We tested for density dependence in adult female and juvenile survival rates, and then discriminated between predator‐ and food‐regulation hypotheses by testing for density‐dependence amongst mortality causes for adult female elk. Notably, the population declined almost 70% from near previously published estimates of carrying capacity over 10 years, providing ideal conditions to test for density dependence. In contrast to predictions, we found only weak support for density dependence in adult survival and juvenile survival. We also found few differences between migrant and resident elk in adult or juvenile survival, though juvenile survival differences were biologically significant. Predation by humans and grizzly bears was density dependent, but similar between migratory strategies. Predation by wolves was the leading known cause of mortality, yet remained constant with declining elk density equally for both migrant and resident elk, indicating wolf predation was density‐independent. Instead of being strongly regulated by food or predation, we found adult female survival was driven by density‐independent predation and climatic factors. The few differences between migratory strategies suggest equivalent fitness payoffs for migrants and residents. This population is being limited by density‐independent predation leading to declines of both migratory strategies. Our results challenge classical predator–prey theory, and call for better integration between predator–prey and migration theory.     

Coping with Shifting Nest Predation Refuges by European Reed Warblers Acrocephalus scirpaceus

Predation, the most important source of nest mortality in altricial birds, has been a subject of numerous studies during past decades. However, the temporal dynamics between changing predation pressures and parental responses remain poorly understood. We analysed characteristics of 524 nests of European reed warblers monitored during six consecutive breeding seasons in the same area, and found some support for the shifting nest predation refuge hypothesis. Nest site characteristics were correlated with nest fate, but a nest with the same nest-site attributes could be relatively safe in one season and vulnerable to predation in another. Thus nest predation refuges were ephemeral and there was no between-season consistency in nest predation patterns. Reed warblers that lost their first nests in a given season did not disperse farther for the subsequent reproductive attempt, compared to successful individuals, but they introduced more changes to their second nest sites. In subsequent nests, predation risk remained constant for birds that changed nest-site characteristics, but increased for those that did not. At the between-season temporal scale, individual birds did not perform better with age in terms of reducing nest predation risk. We conclude that the experience acquired in previous years may not be useful, given that nest predation refuges are not stable.     

Consequences of information use in breeding habitat selection on the evolution of settlement time

The role of temporal changes and spatial variability in predation risk and prey's means of mitigating such risks is poorly understood in the context of potential threats of global climate change for migratory birds. Yet nest predation, for example, represents a primary source of reproductive mortality in birds. To assess risk birds must spend time prospecting potential breeding sites for cues or signals of predator presence. However, competition for breeding sites with advantage to prior residency poses an evolutionary dilemma as individuals also benefit from early settling. We develop a model to examine adaptive prospecting time for predator cues on breeding grounds characterized by spatial heterogeneity in nest predation risk. We study how populations respond to environmental change represented by variation in habitat specific levels of nest predation, habitat composition, population vital rates, and availability of information (via prospecting) in the form of acoustic predator cues. We identify two mechanisms that regulate and buffer impacts of environmental change on populations. First, the adaptive response to lower population abundance under deteriorating environmental conditions is to increase prospecting time, which in turn increases individuals nest success to counteract population declines. This occurs because reduced competition for sites decreases the benefit of early settlement. Second, per capita success in site choice increases during population declines owing to reduced competition that increases the availability of good sites. We also show that the increased benefit to settling early when competition increases can lead to the paradoxical result that with greater spatial heterogeneity, less effort is placed on discerning good and bad sites. Our analysis thus contributes several novel results by which nest predation, settlement phenology, prospecting time and information gathering can influence species capacity to adapt to changing environments.     

A theoretical investigation of the effect of latitude on avian life histories

Tropical birds lay smaller clutches than birds breeding in temperate regions and care for their young for longer. We develop a model in which birds choose when and how often to breed and their clutch size, depending on their foraging ability and the food availability. The food supply is density dependent. Seasonal environments necessarily have a high food peak in summer; in winter, food levels drop below those characteristic of constant environments. A bird that cannot balance its energy needs during a week dies of starvation. If adult predation is negligible, birds in low seasonal environments are constrained by low food during breeding seasons, whereas birds in high seasonal environments die during the winter. Low food seasonality selects for small clutch sizes, long parental care times, greater age at first breeding, and high juvenile survival. The inclusion of adult predation has no major effect on any life-history variables. However, increased nest predation reduces clutch size. The same trends with seasonality are also found in a version of the model that includes a condition variable. Our results show that seasonal changes in food supply are sufficient to explain the observed trends in clutch size, care times, and age at first breeding.     

High nest failure but better nestling quality for early breeders in an alpine population of Northern Wheatear (Oenanthe oenanthe)

Climate change is leading to the advancement of spring conditions, resulting in an earlier snowmelt and green-up, with highest rates of change in highly seasonal environments, including alpine habitats. Migratory birds breeding at high elevations need to time their arrival and lay dates accurately with this advancement, but also with the annually variable spring conditions at their breeding sites, to maximize nest survival probabilities and reproductive output. Nest survival probability and mean nestling mass were analysed in relation to lay date and habitat conditions in an alpine population of the migratory Northern Wheatear Oenanthe oenanthe collected over six consecutive breeding seasons in the Western Italian Alps. This open grassland species showed the lowest nest survival probability in years with an early onset of spring conditions. Within-season, nest survival was highest when breeding late, at lower elevations, and when grass cover and grass height were higher. Both across- and within-season, severe weather conditions may indirectly lead to higher early season nest failure rates by increasing predation risk. By contrast, mean nestling mass, and thus the quality of the fledglings, was lower when breeding late. This might be driven by a mismatch with the peak in food abundance. Breeding early is thus generally advantageous in terms of chick quality in our high-elevation population, but reproductive success is limited by the risk of nest failure that is higher in early springs and early in the season. This trade-off between breeding early and late may thus allow Northern Wheatears to maximize fitness under highly variable spring conditions. However, climate change may cause disruption to this trade-off, and shifts in phenology could become a threat for migratory alpine birds that might not be able to keep track of advancing spring conditions.     

Avoid nest predation when predation rates are low,and other lessons: testing the tropical–temperate nest predation paradigm

Nest predation is the most important cause of nest failure in most birds and latitudinal differences in nest predation rates and life histories suggest that nest predation has been influential in life history evolution. All else equal, natural selection should favor reduction of nest predation, yet evidence is equivocal. We used Monte Carlo simulations to examine the combined effects of variation in nest predation rates, breeding season length and renesting intervals on the annual number of young fledged. Simulations suggest that selection most strongly favors a reduction in nest predation when breeding seasons are short and predation rates are low (temperate characteristics). Conversely, selection favors shorter renesting intervals when breeding seasons are long and nest predation rates are high (tropical characteristics). Reducing already low rates provides a proportionately greater increase in annual nesting success than does the same reduction when nest predation rates are higher. In some tropical species, individuals increase reproductive success not by avoiding predation in subsequent nesting attempts, which is largely beyond their control, but rather by reducing renesting intervals. We suggest that the emphasis on nest predation avoidance has biased our perspectives for alternative hypotheses of how birds should respond to nest predation and the consequences of those alternatives for life history theory. Similarly to the need to control for phylogenetics in examining life history strategies, future studies must also control for differences in breeding season lengths and renesting intervals to better understand the influence of nest predation on avian life histories.     

Partial migration in birds: tests of three hypotheses in a tropical lekking frugivore

1. Partially migratory species provide opportunities to understand which ecological factors cause some animals to migrate when others remain resident year round. Partial migration in birds has been explained by the dominance, arrival-time, and body-size hypotheses. 2. Testing these hypotheses has proven difficult due to the similarities of the predictions they make in temperate-breeding long-distance migrants. In tropical altitudinal migrants, however, these hypotheses make different predictions regarding the sex, age, and condition of migrants and residents. 3. Among white-ruffed manakins in Costa Rica, young birds were not more likely to migrate (as predicted by the dominance hypothesis), nor were females more likely to migrate (as predicted by the arrival-time hypothesis). All condition-related variables interacted with sex, together explaining much of the variation in migratory behaviour. 4. I re-articulate the body-size hypothesis in the context of tropical altitudinal bird migration, focusing explicitly on how limited foraging opportunities and differences in individual condition affect fasting ability during torrential rains. Despite ample food, the smallest birds or those stressed by parasites or moult may risk starvation at breeding elevations due to a reduction in foraging time. These results highlight how intrinsic and extrinsic factors may interact to produce observed patterns of within- and among-species variation in migratory behaviour.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

Nest predation is little affected by parental behaviour and nest site in two African Sylvia warblers

Parental behaviour and nest site are supposed to affect nest predation in birds. Few nest visits and high nest attentiveness are assumed to lead to low predation rates. Poorly concealed nests are thought to be more likely to be preyed upon than well concealed nests. Studies on the relationship between parental behaviour, nest site, and nest predation are rare and none have, so far, been conducted in the Afrotropics. We studied the effect of nest site, nest visitation rate, and nest attentiveness on the nest predation rate of the two tropical warblers Sylvia boehmi and Sylvia lugens in Kenya. Parental behaviour and predation on nests of 13 breeding pairs of both species were observed daily in two consecutive breeding seasons. In both species, parental activity at the nest was low [0.9 trips to the nest in 30 min during incubation, maximum 4.6 (S. boehmi) and 5.8 (S. lugens) trips to the nest in the nestling stage]. Predation rates in both species were high (Mayfield nest success 19.4 and 33.2%). Our analysis revealed only weak evidence for an effect of nest site, nest visitation rate, and nest attentiveness on the predation rate. It is suggested that smaller clutches of tropical in comparison to northern temperate birds result from lower feeding rates in tropical ecosystems with high predation rates (Skutchs hypothesis). The underlying mechanism could not be proven in this study.     

Heterospecific attraction and food resources in migrants' breeding patch selection in northern boreal forest

We studied experimentally how heterospecific attraction may affect habitat selection of migrant passerine birds in Finnish Lapland. We manipulated the densities of resident tit species (Parus spp.). In four study plots residents were removed before the arrival of the migrants in the first study year, and in four other plots their densities were increased by releasing caught individuals. In the second year the treatments of the areas were reversed, allowing paired comparisons within each plot. We also investigated the relative abundance of arthropods in the study plots by the sweep-net method. This allowed us to estimate the effect of food resources on the abundance of birds. The heterospecific attraction hypothesis predicts that densities of migrant species (especially habitat generalists) would be higher during increased resident density. Results supported this prediction. Densities and number of the most abundant migrant species were significantly higher when resident density was increased than when they were removed. On the species level the redwing (Turdus iliacus) showed the strongest positive response to the increased abundance of tits. Migrant bird abundances seemed not to vary in parallel with relative arthropod abundance, with the exception of the pied flycatcher (Ficedula hypoleuca) which showed a strongly positive correlation with many arthropod groups. The results of the experiment indicate that migrants can use resident tit species as a cue to a profitable breeding patch. The relationship between the abundance of the birds and arthropods suggests that annual changes in food resources during the breeding season probably do not have a very important effect on bird populations in these areas. The results stress the importance of positive interspecific interactions in structuring northern breeding bird communities. Received: 1 September 1997 / Accepted: 22 January 1998     

EVOLUTION OF AVIAN COOPERATIVE BREEDING: COMPARATIVE TESTS OF THE NEST PREDATION HYPOTHESIS

Comparative analyses carried out on two different phylogenies of cooperatively and noncooperatively breeding Australian passerine birds (parvorder Corvida) were unable to detect a significant difference in nest predation rates after controlling for body mass and risk of predation due to location of the nest (nest safety). Nest predation rates, however, decrease as nest safety and body mass increase. We suggest that cooperative breeding does not bring about a current net change in rates of nest predation among Australian passerines. Species breeding cooperatively may have developed antipredator strategies that produce results similar to those adopted by noncooperatively breeding species. The function of cooperative breeding may lie outside of antipredator strategies.     

Provision of supplementary food for wild birds may increase the risk of local nest predation

In countries such as the UK, USA and Australia, approximately half of all households provide supplementary food for wild birds, making this the public's most common form of active engagement with nature. Year‐round supplementary feeding is currently encouraged by major conservation charities in the UK as it is thought to be of benefit to bird conservation. However, little is understood about how the provision of supplementary food affects the behaviour and ecology of target and non‐target species. Given the scale of supplementary feeding, any negative effects may have important implications for conservation. Potential nest predators are abundant in urban areas and some species frequently visit supplementary feeding stations. We assess whether providing supplementary food affects the likelihood of nest predation in the vicinity of the feeder, by acting as a point attractant for potential nest predators. We provided feeding stations (empty, peanut feeder, peanut feeder with guard to exclude potential nest predators) in an area of suburban parkland in the UK and monitored the predation rate of eggs placed in artificial nests located at distances that replicated the size of typical suburban gardens. Nest predators (Magpies Pica pica, Grey Squirrels Sciurus carolinensis) were frequent visitors to filled feeders, and predation caused by Magpies, European Jays Garrulus glandarius and Grey Squirrels was significantly higher when nests were adjacent to filled feeders. The presence of a feeder guard did not significantly reduce nest predation. As supplementary feeding is becoming increasingly common during the breeding season in suburban habitats, we suggest that providing point attractants to nest predators at this time may have previously unconsidered consequences for the breeding success of urban birds.     

Competition, predation and nest niche shifts among tropical cavity nesters: phylogeny and natural history evolution of parrots (Psittaciformes) and trogons (Trogoniformes)

Nest site selection by birds is a critically important life history trait as competition for suitable sites can be intense, and because birds are at their most vulnerable to predators during nesting. Previous studies show that the clutch size and nestling period evolve in response to competition for nest sites and nest predation, respectively. This provides the opportunity to study the relative contribution of competition and predation to the evolution of nesting niche. Using previously published phylogenies for parrots and trogons, I found evidence for at least 13 independent evolutionary transitions from tree cavities to alternative nesting niches (including termitaria, cliffs, and burrows). I analyzed variations in clutch size, incubation period and nestling period for 16 phylogenetically controlled pairs of species to test the relative roles of competition for tree cavities and nest predation, in favoring evolutionary switches to alternative nest sites. Tree cavity nesting species did not have larger clutch sizes as predicted if competition for tree cavities leads birds to invest heavily in nesting once they obtain a nest site (the limited breeding opportunities hypothesis). Instead I found that shifts to alternative nesting niches were accompanied by an increase in nestling period. As nestling period is a surrogate measure for long-term nest predation rates, this finding suggests that nest predation has been more important than competition in niche diversification among cavity nesting parrots and trogons. The timing of events in South America suggests that the explosive radiation of mammalian nest predators during the Upper-Oligocene, Lower-Miocene (20–30 million years ago) corresponded with the radiation of parrot and trogon taxa that exploit novel nesting niches.     

Management of reedbeds: mosaic reed cutting does not affect prey abundance and nest predation rate of reed passerine birds

Reed passerine birds are strict habitat specialists inhabiting reedbed habitats. In Europe, many of these species are threatened due to loss and degradation of natural reedbeds. Another important factor that can negatively affect the abundance of reed passerines is commercial reed harvesting. Previous studies have shown negative impacts of large-scale winter reed cutting on passerine breeding assemblages and arthropod communities. The effect of reed cutting on a small scale, however, has not been studied experimentally to date. The aim of this study was to investigate whether and how small-scale, mosaic reed cutting influences prey abundance and nest predation rate of reed passerines. In June, after the reed had reached maturity, we conducted nest predation experiments with artificial nests and arthropod sampling using pan traps in cut reed patches, adjacent uncut reed patches and unmanaged reedbed. We found no differences in the risk of egg predation between three types of reedbeds. In contrast, the abundance of arthropods in cut and adjacent uncut reed patches was significantly higher than that in unmanaged reedbed. We assume this was caused by habitat heterogeneity, small size of cut patches and their rapid recolonization by arthropods from adjacent uncut patches. Our results suggest that in contrast to large-scale reed cutting, small-scale, mosaic reed cutting has no negative effect on nest survival and food abundance of reed passerine birds. However, given that we performed all experiments in June, i.e., when the reed was mature, our findings cannot be generalized to whole breeding period of all reed passerine birds. Therefore, temporal variation in nest predation rate and arthropod abundance in managed and unmanaged reedbeds during the entire breeding season should be examined in future studies.     

Phenological sensitivity to climate change is higher in resident than in migrant bird populations among European cavity breeders

Many organisms adjust their reproductive phenology in response to climate change, but phenological sensitivity to temperature may vary between species. For example, resident and migratory birds have vastly different annual cycles, which can cause differential temperature sensitivity at the breeding grounds, and may affect competitive dynamics. Currently, however, adjustment to climate change in resident and migratory birds have been studied separately or at relatively small geographical scales with varying time series durations and methodologies. Here, we studied differential effects of temperature on resident and migratory birds using the mean egg laying initiation dates from 10 European nest box schemes between 1991 and 2015 that had data on at least one resident tit species and at least one migratory flycatcher species. We found that both tits and flycatchers advanced laying in response to spring warming, but resident tit populations advanced more strongly in relation to temperature increases than migratory flycatchers. These different temperature responses have already led to a divergence in laying dates between tits and flycatchers of on average 0.94 days per decade over the current study period. Interestingly, this divergence was stronger at lower latitudes where the interval between tit and flycatcher phenology is smaller and winter conditions can be considered more favorable for resident birds. This could indicate that phenological adjustment to climate change by flycatchers is increasingly hampered by competition with resident species. Indeed, we found that tit laying date had an additional effect on flycatcher laying date after controlling for temperature, and this effect was strongest in areas with the shortest interval between both species groups. Combined, our results suggest that the differential effect of climate change on species groups with overlapping breeding ecology affects the phenological interval between them, potentially affecting interspecific interactions.     

A flyway perspective on food resource abundance in a long-distance migrant, the Eurasian teal (Anas crecca)

Two frequent assumptions about the evolution of long-distance migration in birds are that they travel long distances annually to reach food-rich areas for breeding, and that they time their migratory journey to be at staging sites when the latter provide the best feeding conditions. These assumptions have rarely been properly tested, and there is no study in which a species’ major food types have been measured by standardized methods throughout a flyway and over a large part of the year. We here present such data for Eurasian teal (Anas crecca), converted to a common energetic currency, and collected at wintering, spring staging and breeding sites. Teal did not time migration to maximize local food abundance; most birds left wintering and spring staging sites before a sharp increase in invertebrate food abundance occurred. On the other hand, hatching of ducklings coincided with a peak in invertebrate food abundance on boreal breeding lakes. Mean overall food abundance (invertebrates and seeds combined) did not differ between wintering sites in southern France and breeding sites in northern Sweden at the time of breeding. Our results are inconsistent with the hypothesis that long-distance migration in dabbling ducks has evolved because adult birds gain an immediate pay-off in increased food abundance by flying north in spring. However, our data confirm a selective advantage for breeding at higher latitudes, because hatching of ducklings may coincide with a peak in invertebrate emergence and because longer days may increase the duration of efficient foraging.