Natural grazing by horses and cattle promotes bird diversity in a restored European alluvial grassland.

Context: A challenge in grassland conservation is to maintain both the openness and the heterogeneity of the habitat to support the diversity of their animal communities, including birds-a taxon that is known to be sensitive to disturbance. An increasingly used management tool in European grassland conservation, especially in rewilding projects, is grazing by large herbivores such as horses and cattle. These grazers are believed to create and maintain patchy landscapes that promote diversity and richness of other species, but their influence on birds is often debated by conservationists, who raise concerns about the impact of disturbance by the grazers. Objectives: Our aim was to examine the relationship between the abundance and species richness of birds across four foraging guilds and the area utilization patterns of Highland cattle and Konik horses in an alluvial grassland in France. We also aimed to examine the influence of land cover and season on the spatial distribution, including abundance and species richness, of different bird guilds present in the grazed area. Methods: We used GPS-collars on all grazers and recorded their positions on an hourly basis over a study period of 1.5 years, assessing patterns of area usage. We counted birds weekly along three transects to describe their distribution within the grazed area and carried out land-cover surveys to describe the habitat. To assess how species richness and abundance of birds of different guilds were related to grazer density, season, and habitat characteristics, we used GAMM models in a spatially explicit framework. We also compared bird numbers at our main study site with a nearby non-grazed control area. Results: The number of birds in the grazed area was about twice the number in the non-grazed control area. Within the grazed area, the abundance of open-area foraging birds increased with increasing grazer density. The number of woodland-foraging birds was also positively correlated with grazer density but less so than open-area foraging birds. The number of individuals in the aerial and wetland bird guilds was not correlated with the density of grazers. Most bird species and individuals were observed on open landscapes scattered with woody patches and waterbodies, and on areas with moderate grazer density. Conclusions: Low-intensity grazing represents a potentially important management tool in creating heterogeneity in alluvial grasslands, thereby promoting suitable habitat for a diverse assemblage of bird species.

It's about Her: Male Within-Season Movements Are Related to Mate Searching in a Songbird.

AbstractIn species with resource-defense mating systems (such as most temperate-breeding songbirds), male dispersal is often considered to be limited in both frequency and spatial extent. When dispersal occurs within a breeding season, the favored explanation is ecological resource tracking. In contrast, movements of male birds associated with temporary emigration, such as polyterritoriality (i.e., defense of an additional location after attracting a female in the initial territory), are usually attributed to mate searching. We suggest that male dispersal and polyterritoriality are functionally related and that mate searching may be a unifying hypothesis for predicting the within-season movements of male songbirds. Here, we test three key predictions derived from this hypothesis in Wood Warblers (Phylloscopus sibilatrix). We collected data on the spatial behavior of 107 males between 2017 and 2019 and related male movements to a new territory (in both a dispersal and a polyterritorial context) to mating potential in the current territory. Most males dispersed from their territories within days or weeks after failing to attract a female, despite occupying territories in apparently suitable habitat. Probability of polyterritoriality by paired males increased after the peak fertile period of their mate. Males never dispersed following nest predation if the female remained to renest. Thus, our data are consistent with the hypothesis that both movement types are functionally related to mate searching.

Wood warbler population dynamics in response to mast seeding regimes in Europe.

Mast seeding is the episodic, massive production of plant seeds synchronized over large areas. The resulting superabundance of seeds represents a resource pulse that can profoundly affect animal populations across trophic levels. Following years of high seed production, the abundance of both seed consumers and their predators increase. Higher predator abundance leads to increased predation pressure across the trophic web, impacting nonseed consumers such as the wood warbler Phylloscopus sibilatrix through increased nest predation after tree mast years. Over the past 30 years, the frequency of tree seed masts has increased, while wood warbler populations have declined in several regions of Europe. We hypothesized that increasing mast frequencies may have contributed to the observed population declines by creating suboptimal breeding conditions in years after masting. We measured reproductive output in four study areas in central Europe, which was between 0.61 and 1.24 fledglings lower in the years following masting than nonmasting. For each study area, we used matrix population models to predict population trends based on the estimated reproductive output and the local mast frequencies. We then compared the predicted with the observed population trends to assess if the frequency of mast years had contributed to the population dynamics. In Wielkopolska National Park (PL) and Hessen (DE), masting occurred on average only every 4 years and populations were stable or nearly so, whereas in Jura (CH) and Bialowieza National Park (PL), masting occurred every 2 and 2.5 years, respectively, and populations were declining. The simple matrix population models predicted the relative difference among local population trends over the past 10-20 years well, suggesting that the masting frequency may partly explain regional variation in population trends. Simulations suggest that further increases in mast frequency will lead to further declines in wood warbler populations. We show that changes in a natural process, such as mast seeding, may contribute to the decline in animal populations through cascading effects.

Patterns of high-flying insect abundance are shaped by landscape type and abiotic conditions.

Insects are of increasing conservation concern as a severe decline of both biomass and biodiversity have been reported. At the same time, data on where and when they occur in the airspace is still sparse, and we currently do not know whether their density is linked to the type of landscape above which they occur. Here, we combined data of high-flying insect abundance from six locations across Switzerland representing rural, urban and mountainous landscapes, which was recorded using vertical-looking radar devices. We analysed the abundance of high-flying insects in relation to meteorological factors, daytime, and type of landscape. Air pressure was positively related to insect abundance, wind speed showed an optimum, and temperature and wind direction did not show a clear relationship. Mountainous landscapes showed a higher insect abundance than the other two landscape types. Insect abundance increased in the morning, decreased in the afternoon, had a peak after sunset, and then declined again, though the extent of this general pattern slightly differed between landscape types. We conclude that the abundance of high-flying insects is not only related to abiotic parameters, but also to the type of landscapes and its characteristics, which, on a long-term, should be taken into account for when designing conservation measures for insects.

Estimating survival in continuous space from mark-dead-recovery data - Towards a continuous version of the multinomial dead recovery model.

Understanding spatially varying survival is crucial for understanding the ecology and evolution of migratory animals, which may ultimately help to conserve such species. We develop an approach to estimate an annual survival probability function varying continuously in geographic space, if the recovery probability is constant over space. This estimate is based on a density function over continuous geographic space and the discrete age at death obtained from dead recovery data. From the same density function, we obtain an estimate for animal distribution in space corrected for survival, i.e., migratory connectivity. This is possible, when migratory connectivity can be separated from recovery probability. In this article, we present the method how spatially and continuously varying survival and the migratory connectivity corrected for survival can be obtained, if a constant recovery probability can be assumed reasonably. The model is a stepping stone in developing a model allowing for disentangling spatially heterogeneous survival and migratory connectivity corrected for survival from a spatially heterogeneous recovery probability. We implement the method using kernel density estimates in the R-package CONSURE. Any other density estimation technique can be used as an alternative. In a simulation study, the estimators are unbiased but show edge effects in survival and migratory connectivity. Applying the method to a real-world data set of European robins Erithacus rubecula results in biologically reasonable continuous heat-maps for survival and migratory connectivity.

Different roles of concurring climate and regional land-use changes in past 40 years' insect trends.

Climate and land-use changes are main drivers of insect declines, but their combined effects have not yet been quantified over large spatiotemporal scales. We analysed changes in the distribution (mean occupancy of squares) of 390 insect species (butterflies, grasshoppers, dragonflies), using 1.45 million records from across bioclimatic gradients of Switzerland between 1980 and 2020. We found no overall decline, but strong increases and decreases in the distributions of different species. For species that showed strongest increases (25% quantile), the average proportion of occupied squares increased in 40 years by 0.128 (95% credible interval: 0.123-0.132), which equals an average increase in mean occupancy of 71.3% (95% CI: 67.4-75.1%) relative to their 40-year mean occupancy. For species that showed strongest declines (25% quantile), the average proportion decreased by 0.0660 (95% CI: 0.0613-0.0709), equalling an average decrease in mean occupancy of 58.3% (95% CI: 52.2-64.4%). Decreases were strongest for narrow-ranged, specialised, and cold-adapted species. Short-term distribution changes were associated to both climate changes and regional land-use changes. Moreover, interactive effects between climate and regional land-use changes confirm that the various drivers of global change can have even greater impacts on biodiversity in combination than alone. In contrast, 40-year distribution changes were not clearly related to regional land-use changes, potentially reflecting mixed changes in local land use after 1980. Climate warming however was strongly linked to 40-year changes, indicating its key role in driving insect trends of temperate regions in recent decades.

Estimation in the multinomial reencounter model - Where do migrating animals go and how do they survive in their destination area?

Spatial variation in survival has individual fitness consequences and influences population dynamics. Which space animals use during the annual cycle determines how they are affected by this spatial variability. Therefore, knowing spatial patterns of survival and space use is crucial to understand demography of migrating animals. Extracting information on survival and space use from observation data, in particular dead recovery data, requires explicitly identifying the observation process. We build a fully stochastic model for animals marked in populations of origin, which were found dead in spatially discrete destination areas. The model acts on the population level and includes parameters for use of space, survival and recovery probability. It is based on the division coefficient and the multinomial reencounter model. We use a likelihood-based approach, derive Restricted Maximum Likelihood-like estimates for all parameters and prove their existence and uniqueness. In a simulation study we demonstrate the performance of the model by using Bayesian estimators derived by the Markov chain Monte Carlo method. We obtain unbiased estimates for survival and recovery probability if the sample size is large enough. Moreover, we apply the model to real-world data of European robins Erithacus rubecula ringed at a stopover site. We obtain annual survival estimates for different spatially discrete non-breeding areas. Additionally, we can reproduce already known patterns of use of space for this species.

Hatching phenology is lagging behind an advancing snowmelt pattern in a high-alpine bird.

To track peaks in resource abundance, temperate-zone animals use predictive environmental cues to rear their offspring when conditions are most favourable. However, climate change threatens the reliability of such cues when an animal and its resource respond differently to a changing environment. This is especially problematic in alpine environments, where climate warming exceeds the Holarctic trend and may thus lead to rapid asynchrony between peaks in resource abundance and periods of increased resource requirements such as reproductive period of high-alpine specialists. We therefore investigated interannual variation and long-term trends in the breeding phenology of a high-alpine specialist, the white-winged snowfinch, Montifringilla nivalis, using a 20-year dataset from Switzerland. We found that two thirds of broods hatched during snowmelt. Hatching dates positively correlated with April and May precipitation, but changes in mean hatching dates did not coincide with earlier snowmelt in recent years. Our results offer a potential explanation for recently observed population declines already recognisable at lower elevations. We discuss non-adaptive phenotypic plasticity as a potential cause for the asynchrony between changes in snowmelt and hatching dates of snowfinches, but the underlying causes are subject to further research.

High turn-over rates at the upper range limit and elevational source-sink dynamics in a widespread songbird.

The formation of an upper distributional range limit for species breeding along mountain slopes is often based on environmental gradients resulting in changing demographic rates towards high elevations. However, we still lack an empirical understanding of how the interplay of demographic parameters forms the upper range limit in highly mobile species. Here, we study apparent survival and within-study area dispersal over a 700 m elevational gradient in barn swallows (Hirundo rustica) by using 15 years of capture-mark-recapture data. Annual apparent survival of adult breeding birds decreased while breeding dispersal probability of adult females, but not males increased towards the upper range limit. Individuals at high elevations dispersed to farms situated at elevations lower than would be expected by random dispersal. These results suggest higher turn-over rates of breeding individuals at high elevations, an elevational increase in immigration and thus, within-population source-sink dynamics between low and high elevations. The formation of the upper range limit therefore is based on preference for low-elevation breeding sites and immigration to high elevations. Thus, shifts of the upper range limit are not only affected by changes in the quality of high-elevation habitats but also by factors affecting the number of immigrants produced at low elevations.

Spatio-temporal variation in the wintering associations of an alpine bird.

Many animals make behavioural changes to cope with winter conditions, being gregariousness a common strategy. Several factors have been invoked to explain why gregariousness may evolve during winter, with individuals coming together and separating as they trade off the different costs and benefits of living in groups. These trade-offs may, however, change over space and time as a response to varying environmental conditions. Despite its importance, little is known about the factors triggering gregarious behaviour during winter and its change in response to variation in weather conditions is poorly documented. Here, we aimed at quantifying large-scale patterns in wintering associations over 23 years of the white-winged snowfinch Montifringilla nivalis nivalis. We found that individuals gather in larger groups at sites with harsh wintering conditions. Individuals at colder sites reunite later and separate earlier in the season than at warmer sites. However, the magnitude and phenology of wintering associations are ruled by changes in weather conditions. When the temperature increased or the levels of precipitation decreased, group size substantially decreased, and individuals stayed united in groups for a shorter time. These results shed light on factors driving gregariousness and points to shifting winter climate as an important factor influencing this behaviour.

Grazer density and songbird counts in a restored conservation area.

Grazing by large herbivores is increasingly used as a management tool in European nature reserves. The aim is usually to support an open but heterogeneous habitat and its corresponding plant and animal communities. Previous studies showed that birds may profit from grazing but that the effect varies among bird species. Such studies often compared bird counts among grazed areas with different stocking rates of herbivores. Here, we investigated how space use of Konik horses and Highland cattle is related to bird counts in a recently restored conservation area with a year-round natural grazing management. We equipped five horses and five cattle with GPS collars and correlated the density of their GPS positions on the grazed area with the density of bird observations from winter through the breeding season. We found that in the songbirds of our study site, both the overall density of bird individuals and the number of species increased with increasing density of GPS positions of grazers. Correlations of bird density with horse density were similar to correlations with cattle density. Of the eight most common songbird species observed in our study area, the Eurasian Skylark and the Common Starling had the clearest positive correlations with grazer density, while the Blackbird showed a negative correlation. Skylarks and Starlings in our study area thus seem to profit from year-round natural grazing by a mixed group of horses and cattle.

Glucocorticoid levels are linked to lifetime reproductive success and survival of adult barn owls.

Glucocorticoid hormones, such as corticosterone, are crucial in regulating daily life metabolism and energy expenditure, as well as promoting short-term physiological and behavioural responses to unpredictable environmental challenges. Therefore, glucocorticoids are considered to mediate trade-offs between survival and reproduction. Relatively little is known about how selection has shaped glucocorticoid levels. We used 15 years of capture-recapture and dead recovery data combined with 13 years of corticosterone and breeding success data taken on breeding barn owls (Tyto alba) to investigate such trade-offs. We found that survival was positively correlated with stress-induced corticosterone levels in both sexes, whereas annual and lifetime reproductive success (i.e. the sum of young successfully fledged during the entire reproductive career) was positively correlated with both baseline and stress-induced corticosterone levels in females only. Our results suggest that, in the barn owl, the stress-induced corticosterone response is a good proxy for adult survival and lifetime reproductive success. However, selection pressure appears to act differently on corticosterone levels of males and females.

Potential sex-dependent effects of weather on apparent survival of a high-elevation specialist.

Mountain ecosystems are inhabited by highly specialised and endemic species which are particularly susceptible to climatic changes. However, the mechanisms by which climate change affects species population dynamics are still largely unknown, particularly for mountain birds. We investigated how weather variables correlate with survival or movement of the white-winged snowfinch Montifringilla nivalis, a specialist of high-elevation habitat. We analysed a 15-year (2003-2017) mark-recapture data set of 671 individuals from the Apennines (Italy), using mark-recapture models. Mark-recapture data allow estimating, forgiven time intervals, the probability that individuals stay in the study area and survive, the so called apparent survival. We estimated annual apparent survival to be around 0.44-0.54 for males and around 0.51-0.64 for females. Variance among years was high (range: 0.2-0.8), particularly for females. Apparent survival was lower in winter compared to summer. Female annual apparent survival was negatively correlated with warm and dry summers, whereas in males these weather variables only weakly correlated with apparent survival. Remarkably, the average apparent survival measured in this study was lower than expected. We suggest that the low apparent survival may be due to recent changes in the environment caused by global warming. Possible, non-exclusive mechanisms that potentially also could explain sexual differential apparent survival act via differential breeding dispersal, hyperthermia, weather-dependent food availability, and weather-dependent trade-off between reproduction and self-maintenance. These results improve our current understanding of the mechanisms driving population dynamics in high-elevation specialist birds, which are particularly at risk due to climate change.

Integrating stable isotopes, parasite, and ring-reencounter data to quantify migratory connectivity-A case study with Barn Swallows breeding in Switzerland, Germany, Sweden, and Finland.

Ecosystems around the world are connected by seasonal migration. The migrant animals themselves are influenced by migratory connectivity through effects on the individual and the population level. Measuring migratory connectivity is notoriously difficult due to the simple requirement of data conveying information about the nonbreeding distribution of many individuals from several breeding populations. Explicit integration of data derived from different methods increases the precision and the reliability of parameter estimates. We combine ring-reencounter, stable isotope, and blood parasite data of Barn Swallows Hirundo rustica in a single integrated model to estimate migratory connectivity for three large scale breeding populations across a latitudinal gradient from Central Europe to Scandinavia. To this end, we integrated a non-Markovian multistate mark-recovery model for the ring-reencounter data with normal and binomial mixture models for the stable isotope and parasite data. The integration of different data sources within a mark-recapture modeling framework enables the most precise quantification of migratory connectivity on the given broad spatial scale. The results show that northern-breeding populations and Southern Africa as well as southern-breeding populations and Western-Central Africa are more strongly connected through Barn Swallow migration than central European breeding populations with any of the African wintering areas. The nonbreeding distribution of Barn Swallows from central European breeding populations seems to be a mixture of those populations breeding further north and south, indicating a migratory divide.

Manipulating the Alpha Level Cannot Cure Significance Testing.

We argue that making accept/reject decisions on scientific hypotheses, including a recent call for changing the canonical alpha level from p = 0.05 to p = 0.005, is deleterious for the finding of new discoveries and the progress of science. Given that blanket and variable alpha levels both are problematic, it is sensible to dispense with significance testing altogether. There are alternatives that address study design and sample size much more directly than significance testing does; but none of the statistical tools should be taken as the new magic method giving clear-cut mechanical answers. Inference should not be based on single studies at all, but on cumulative evidence from multiple independent studies. When evaluating the strength of the evidence, we should consider, for example, auxiliary assumptions, the strength of the experimental design, and implications for applications. To boil all this down to a binary decision based on a p-value threshold of 0.05, 0.01, 0.005, or anything else, is not acceptable.

Early returning long-distance migrant males do pay a survival cost.

Timing of return to the breeding area presumably optimizes breeding output in migrants. How timing affects the other components of fitness - survival, has been comparatively little studied. Returning too early in spring is expected to be associated with high mortality in insectivorous migrants when weather conditions are still unsuitable. Yet, males in particular arrive early to get access to the best territories which have been suggested to cause arrival before it is optimal for their survival. For the outward migration in autumn, timing is presumably less directly associated with reproduction and fitness and how it might affect survival is not well understood. We use data of eight songbird species ringed across Denmark to investigate how timing of return migration in spring and departure migration in autumn close to the breeding areas affects survival for short- and long-distance migrants. Further, we compare survival optimum to the timing of males and females at a stopover site in Denmark in three sexually dimorphic, protandric species. We find a clear relationship between return migration and survival which differs between short- and long-distance migrants: Survival decreases with date for short-distance migrants and a bell-shaped relationship, with low survival for earliest and latest individuals, for long-distance migrants. In protandric species, the majority of males return before survival is optimal, whereas females on average return close to the survival optimum. The pattern of survival in relation to autumn timing is less clear, although a similar bell-shaped relationship is suggested for long-distance migrants. Our findings support the predicted mortality consequences of too early return to the breeding grounds and also that selection for early return in males leads to suboptimal migration timing regarding survival.

The earth is flat (p > 0.05): significance thresholds and the crisis of unreplicable research.

The widespread use of 'statistical significance' as a license for making a claim of a scientific finding leads to considerable distortion of the scientific process (according to the American Statistical Association). We review why degrading p-values into 'significant' and 'nonsignificant' contributes to making studies irreproducible, or to making them seem irreproducible. A major problem is that we tend to take small p-values at face value, but mistrust results with larger p-values. In either case, p-values tell little about reliability of research, because they are hardly replicable even if an alternative hypothesis is true. Also significance (p ≤ 0.05) is hardly replicable: at a good statistical power of 80%, two studies will be 'conflicting', meaning that one is significant and the other is not, in one third of the cases if there is a true effect. A replication can therefore not be interpreted as having failed only because it is nonsignificant. Many apparent replication failures may thus reflect faulty judgment based on significance thresholds rather than a crisis of unreplicable research. Reliable conclusions on replicability and practical importance of a finding can only be drawn using cumulative evidence from multiple independent studies. However, applying significance thresholds makes cumulative knowledge unreliable. One reason is that with anything but ideal statistical power, significant effect sizes will be biased upwards. Interpreting inflated significant results while ignoring nonsignificant results will thus lead to wrong conclusions. But current incentives to hunt for significance lead to selective reporting and to publication bias against nonsignificant findings. Data dredging, p-hacking, and publication bias should be addressed by removing fixed significance thresholds. Consistent with the recommendations of the late Ronald Fisher, p-values should be interpreted as graded measures of the strength of evidence against the null hypothesis. Also larger p-values offer some evidence against the null hypothesis, and they cannot be interpreted as supporting the null hypothesis, falsely concluding that 'there is no effect'. Information on possible true effect sizes that are compatible with the data must be obtained from the point estimate, e.g., from a sample average, and from the interval estimate, such as a confidence interval. We review how confusion about interpretation of larger p-values can be traced back to historical disputes among the founders of modern statistics. We further discuss potential arguments against removing significance thresholds, for example that decision rules should rather be more stringent, that sample sizes could decrease, or that p-values should better be completely abandoned. We conclude that whatever method of statistical inference we use, dichotomous threshold thinking must give way to non-automated informed judgment.

Longer wings for faster springs - wing length relates to spring phenology in a long-distance migrant across its range.

In migratory birds, morphological adaptations for efficient migratory flight often oppose morphological adaptations for efficient behavior during resident periods. This includes adaptations in wing shape for either flying long distances or foraging in the vegetation and in climate-driven variation of body size. In addition, the timing of migratory flights and particularly the timely arrival at local breeding sites is crucial because fitness prospects depend on site-specific phenology. Thus, adaptations for efficient long-distance flights might be also related to conditions at destination areas. For an obligatory long-distance migrant, the common nightingale, we verified that wing length as the aerodynamically important trait, but not structural body size increased from the western to the eastern parts of the species range. In contrast with expectation from aerodynamic theory, however, wing length did not increase with increasing migration distances. Instead, wing length was associated with the phenology at breeding destinations, namely the speed of local spring green-up. We argue that longer wings are beneficial for adjusting migration speed to local conditions for birds breeding in habitats with fast spring green-up and thus short optimal arrival periods. We suggest that the speed of spring green-up at breeding sites is a fundamental variable determining the timing of migration that fine tune phenotypes in migrants across their range.

A bird distribution model for ring recovery data: where do the European robins go?

For the study of migratory connectivity, birds have been individually marked by metal rings for more than 100 years. The resulting ring recovery data have been compiled in numerous bird migration atlases. However, estimation of what proportion of a particular population is migrating to which region is confounded by spatial heterogeneity in ring recovery probability. We present a product multinomial model that enables quantifying the continent-wide distribution of different bird populations during different seasons based on ring recovery data while accounting for spatial heterogeneity of ring recovery probability. We applied the model to an example data set of the European robin Erithacus rubecula. We assumed that ring recovery probability was equal between different groups of birds and that survival probability was constant. Simulated data indicate that violation of the assumption of constant survival did not affect our estimated bird distribution parameters but biased the estimates for recovery probability. Posterior predictive model checking indicated a good general model fit but also revealed lack of fit for a few groups of birds. This lack of fit may be due to between-group differences in the spatial distribution on smaller scales within regions. We found that 48% of the Scandinavian robins, but only 31% of the central European robins, wintered in northern Africa. The remaining parts of both populations wintered in southern and central Europe. Therefore, a substantial part of the Scandinavian population appears to leap over individuals from the central European population during migration. The model is applied to summary tables of numbers of ringed and recovered birds. This allows us to handle very large data sets as, for example, those presented in bird migration atlases.

Equal nonbreeding period survival in adults and juveniles of a long-distant migrant bird.

In migrant birds, survival estimates for the different life-history stages between fledging and first breeding are scarce. First-year survival is shown to be strongly reduced compared with annual survival of adult birds. However, it remains unclear whether the main bottleneck in juvenile long-distant migrants occurs in the postfledging period within the breeding ranges or en route. Quantifying survival rates during different life-history stages and during different periods of the migration cycle is crucial to understand forces driving the evolution of optimal life histories in migrant birds. Here, we estimate survival rates of adult and juvenile barn swallows (Hirundo rusticaL.) in the breeding and nonbreeding areas using a population model integrating survival estimates in the breeding ranges based on a large radio-telemetry data set and published estimates of demographic parameters from large-scale population-monitoring projects across Switzerland. Input parameters included the country-wide population trend, annual productivity estimates of the double-brooded species, and year-to-year survival corrected for breeding dispersal. Juvenile survival in the 3-week postfledging period was low (S = 0.32; SE = 0.05), whereas in the rest of the annual cycle survival estimates of adults and juveniles were similarly high (S > 0.957). Thus, the postfledging period was the main survival bottleneck, revealing the striking result that nonbreeding period mortality (including migration) is not higher for juveniles than for adult birds. Therefore, focusing future research on sources of variation in postfledging mortality can provide new insights into determinants of population dynamics and life-history evolution of migrant birds.