Autumn Migration Research Articles

Stopover habitat use of coastal Pied Avocet revealed by satellite tracking and remote sensing technology

Stopover sites identification for migratory waterbirds is important in China’s coastal wetlands, which have experienced a massive loss of natural wetlands in recent decades. Taking the Pied Avocet (Recurvirostra avosetta), a common waterbird in China’s coastal zone as an example, our research focused on their stopover habitat use based on satellite tracking and remote sensing technology. Along the coast, a total of 6 critical stopover sites, duration of stay more than 14 days, were recognized during their 2022–2023 autumn and spring migrations of the 4 tagged individuals. The results showed that Pied Avocets used habitats close to the coastline and water source, at low elevation, and with sparse vegetation. They also had a certain tolerance for human interference. From the perspective of the land use and land cover type, the proportion (33.01 %) of artificial wetlands including paddy, salt pan, and mariculture has exceeded that (27.97 %) of natural wetlands such as bottomland, tidal flat, estuarine waters, and shallow water in the home ranges of the 6 stopover areas. The study emphasized that the habitat function of artificial wetland should be paid enough attention to while maintaining the natural wetland without further loss. In addition, habitat-oriented management suggestions were put forward for artificial wetlands. This study can provide data and technical support for the conservation and management of waterbirds stopovers, also having reference value for other species.

High-resolution spatiotemporal forecasting of the European crane migration

In this paper we present three different models to forecast bird migration. They are species-specific individual-based models that operate on a high spatiotemporal resolution (kilometres, 15 min-hours), as an addition to radar-based migration forecast models that currently exist. The models vary in complexity, and use GPS-tracked location, flying direction and speed, and/or wind data to forecast migration speed and direction. Our aim is to quantitatively evaluate the forecasting performance and assess which metrics improve forecasts at different ranges. We test the models through cross-validation using GPS tracks of common cranes during spring and autumn migration. Our results show that recordings of flight speed and direction improve the accuracy of forecasts on the short range (<2 h). Adding wind data at flight altitude results in consistent improvements of the forecasts across the entire range, particularly in the predicted speed. Direction forecasts are less affected by adding wind data because cranes mostly compensate for wind drift during migration. Migration in spring is more difficult to forecast than in autumn, resulting in larger errors in flight speed and direction during spring. We further find that a combination of flight behaviours – thermal soaring, gliding, and flapping – complicates the forecasts by inducing variance in flight speed and direction. Fitting those behaviours into flight optimisation models proves to be challenging, and even results in significant biases in speed forecasts in spring. We conclude that flight speed is the most difficult parameter to forecast, whereas flight direction is the most critical for practical applications of these models. Such applications could e.g., be prevention of bird strikes in aviation or with wind turbines, and public engagement with bird migration.

Differential responses to weather and land‐cover conditions explain spatial variation in winter abundance trends in a migratory bird of conservation concern

Abstract Effective conservation strategies for animal populations require knowledge of relationships between population dynamics and their environmental drivers. However, these processes often vary within animal populations, requiring site‐specific conservation planning. Given limited financial resources, identifying groups of sites with similar population dynamics can help practitioners efficiently implement conservation programs to larger areas. We evaluated spatial patterns and environmental drivers of wintering site trends in a migratory bird of conservation concern, the Greenland white‐fronted goose (Anser albifrons flavirostris). We used latent class analysis to identify trend patterns in 35 years of abundance data among 59 geographically discrete Greenland white‐fronted goose wintering sites. We developed a state‐space abundance model in a Bayesian framework to quantify the effects of weather and land‐cover conditions experienced throughout spring migration, summer breeding, autumn migration and wintering periods on variation in wintering site abundance. We identified two main patterns in Greenland white‐fronted goose abundance trends: northeastern wintering sites declined on average by 3% per year, while southwestern wintering sites declined on average by 14% per year. Differential responses to weather and habitat conditions likely explained variation among groups, as geese at southwestern wintering sites were more negatively affected by harsh weather conditions (e.g. low temperatures and high precipitation on breeding areas) and poor habitat conditions (i.e. low‐quality grasslands and croplands) on wintering areas. Future conservation efforts to improve the suitability and nutritional quality of agricultural areas, especially cereal croplands in autumn and early winter and grasslands in late winter and early spring, could potentially improve local habitat conditions, especially in the southwestern wintering sites where abundance declines were steepest. Synthesis and applications. We demonstrate the potential to delineate animal populations based on spatial patterns in population dynamics using long‐term abundance monitoring data, which are commonly the only available data for conservation practitioners. By grouping sites based on spatial patterns in local abundance trends, we can further test hypotheses about how these groups are differentially affected by changing environmental conditions. This information is important for informing efficient conservation planning over large areas when financial resources are limited.

An ecological vulnerability index to assess impacts of offshore wind facilities on migratory songbirds

Abstract As offshore wind (OSW) energy expands globally, migratory songbirds are at risk of mortality from collisions with turbine blades, though the magnitude of this threat and which species are most vulnerable, remains poorly understood. Ecological vulnerability indices are commonly used to assess species' susceptibility to harmful factors, with results used to direct scarce research and monitoring resources to species showing relatively high vulnerability. These indices are based on the traits that elevate a species risk to adverse impacts (sensitivity), the overlap in occurrence between a species and the potentially harmful agent (exposure) and the influence of this exposure on the species' local or global persistence (resilience). We modified ecological vulnerability indices for seabirds to assess vulnerability of migratory songbirds to OSW related mortality. As a pertinent case study, we considered songbirds that fly across the Northwest Atlantic during their autumn migration. We utilized readily available information on each species' migratory behaviour, life history, and conservation status to calculate an index score that could range from 1 (lowest vulnerability) to 125 (highest vulnerability). We found scores of 3 to 55.2 for the 101 songbird species evaluated, with New World warblers (Parulidae) over‐represented among the highest scoring species. We found the scores to be sensitive to uncertainty in index components, highlighting the importance of considering scoring uncertainty when evaluating ecological vulnerability indices. Finally, we found that for seven of the top 10 highest scoring species, modest improvements in population trends had the potential to lower the scores substantially. Synthesis and applications. Our methodology is readily applicable to other regions where offshore wind (OSW) development is planned and songbird migration is common, allowing research and monitoring activities to be targeted to species most likely to be negatively affected by OSW facility encounters.

Brain gene expression reveals pathways underlying nocturnal migratory restlessness

Migration is one of the most extreme and energy demanding life history strategies to have evolved in the animal kingdom. In birds, champions of long-distance migrations, the seasonal emergence of the migratory phenotype is characterised by rapid physiological and metabolic remodelling, including substantial accumulation of fat stores and increases in nocturnality. The molecular underpinnings and brain adaptations to seasonal migrations remain poorly understood. Here, we exposed Common quails (Coturnix coturnix) to controlled changes in day length to simulate southward autumn migration, and then blocked the photoperiod until birds entered the non-migratory wintering phase. We first performed de novo RNA-Sequencing from selected brain samples (hypothalamus) collected from birds at a standardised time at night, either in a migratory state (when restlessness was highest and at their body mass peak), or in a non-migratory state and conducted differential gene expression and functional pathways analyses. We found that the migratory state was associated with up-regulation of a few, yet functionally well defined, gene expression networks implicated in fat trafficking, protein and carbohydrate metabolism. Further analyses that focused on candidate genes (apolipoprotein H or APOH, lysosomal associated membrane protein-2 or LAMP2) from samples collected during the day or night across the entire study population suggested differences in the expression of these genes depending on the time of the day with the largest expression levels being found in the migratory birds sampled at night. We also found that expression of APOH was positively associated with levels of nocturnal activity in the migratory birds; such an association was absent within the non-migratory birds. Our results provide novel experimental evidence revealing that hypothalamic changes in expression of apolipoprotein pathways, which regulate the circulatory transport of lipids, are likely key regulatory activators of nocturnal migratory movements. Our study paves the way for performing deeper functional investigations on seasonal molecular remodelling underlying the development of the migratory phenotype.

The Diet of Eleonora’s Falcons (Falco eleonorae) during the Autumn Migration of Passerine Birds across the Aegean Sea

Every year, several hundred million birds cross the Mediterranean on their migration from Eurasia to their wintering quarters in Africa. As many migrants travel at night or at high altitudes, direct observations of bird migration are difficult and thus our information about migrating species, numbers and timing is incomplete. An indirect way to assess autumn migration is the analysis of prey remains of Eleonora’s Falcons (Falco eleonorae). These falcons breed in large colonies on islands in the Mediterranean and on the Canary Islands. Many migrants have to pass these islands on their flight to their African wintering quarters. Eleonora’s Falcons appear to be adapted to the autumn bird migration and raise their young between August and October, when migrating birds are abundant. When nestlings have to be fed, falcons exclusively hunt small birds of 10 to 150 g body mass, whereas they prey mostly on aerial invertebrates (Coleoptera, Hymenoptera, Diptera, Orthoptera, Hemiptera, Odonata, Lepidoptera) from November to July. We studied Eleonora’s Falcons from 1965 to 2001 on a rocky islet, north of Crete, which harboured a colony of about 200 breeding pairs. In 1969, 1971, 1977, and 1988 we systematically monitored and collected the pluckings and cached food items in 22 to 36 nest sites each year. Pluckings were systematically analysed later in Germany using a reference collection of bird feathers for identification. In total, we determined more than 111 prey species (mostly Passerines) comprising more than 13,450 individuals. The top 12 prey species were: Willow Warbler (27.8% of all prey items), Red-backed Shrike (10.7%), Spotted Flycatcher (9.9%), Whinchat (8.8%), Common Whitethroat (5.1%), Wood Warbler (3.8), Tree Pipit (2.9%), Icterine Warbler (2.5%), Greater Short-toed Lark (2.5%), Northern Wheatear (1.8%), Common Nightingale (1.6%), and European Pied Flycatcher (1.5%). Eleonora’s Falcons are selective hunters to some degree; thus, the phenology and abundance data derived from the plucking analyses are biased towards slow-flying species or smaller birds (only up to a body mass of 150 g). When the young falcons develop and grow, food demand increases concomitantly. Comparing the total weight of prey over time indicates a correlation with food demand and in consequence with the number of prey items brought to the nest sites by the falcons.

Shoreline Translocation during Road Expansion Was Successful for Most Waterbirds but Not for Waders

Coastal wetlands are one of the most threatened ecosystems due to, firstly, their relative rarity and, secondly, the strong human interest in these coastal sites for infrastructure development, and recreation. These coastal wetlands also serve as important migration stopover sites for a range of waterbirds. There is great international interest in mitigating the negative effects of human land development and in restoring degraded habitats. I evaluated the response of one waterbird community to the mitigation action implemented during road expansion in an important waterbird coastal habitat in central Norway. Using a novel mitigation method, the shoreline was moved seawards to allow space for a continued shoreline habitat and the extended road. By weekly monitoring the waterbird community during spring and autumn migration periods (n = 7 of each), I found similar biodiversity, evenness, and abundance of geese, dabbling ducks, diving ducks, gulls, and waders when data for the whole wetland were used. However, after construction waders were partially displaced from the zone closest to the road to a zone further away. For some groups of birds, shoreline translocation can be a fruitful mitigation action aiming to reduce the negative effects of infrastructure expansion in coastal areas.

The first satellite tracking data on the migration of the common cuckoo Cuculus canorus (Cuculiformes, Cuculidae) from Southern Siberia (Khakassia, Russia)

In early June and July 2018 and 2019, during the reproduction period, we captured and equipped with satellite transmitters (5.3 g PinPoint GPS ARGOS 120) four males of the common cuckoo Cuculus canorus canorus (Linnaeus, 1758) in the vicinity of the city of Abakan in Khakassia, Southern Siberia. In one male, it was possible to track the route of autumn migration up to the border of the Democratic Republic of the Congo with Zambia (10 819 km), in another male to Western Iran. The routes of the other two birds were interrupted for unknown reasons in the Pavlodar region of Kazakhstan and in the Novosibirsk region. Satellite data confirmed our assumption that the cuckoos breeding in the territory of Southern Siberia migrate in autumn to the southwest via Kazakhstan, Central Asian countries (Uzbekistan, Turkmenistan), Iran, and Saudi Arabia to the southern part of Africa.

Contrasting seasonal responses to wind in migrating songbirds on a globally important flyway.

During spring migration, nocturnal migrants attempt to minimize their travel time to reach their breeding grounds early. However, how they behave and respond to unfavourable conditions during their springtime travels is much less understood. In this study, we reveal the effects of atmospheric factors on nocturnal bird migration under adverse conditions during spring and autumn, based on one of the most detailed bird migration studies globally, using radar data from 13 deployments over a period of seven years (2014-2020) in the Levant region. Using ERA5 reanalysis data, we found that migratory birds maintain similar ground speeds in both autumn and spring migrations, but during spring, when encountering unfavourable winds, they put more effort into maintaining their travel speed by increasing self-powered airspeed by 18%. Moreover, we report for the first time that spring migrants showed less selectivity to wind conditions and migrated even under unfavourable headwind and crosswind conditions. Interestingly, we discovered that temperature was the most important weather parameter, such that warm weather substantially increased migration intensities in both seasons. Our results enhance our understanding of bird migration over the Levant region, one of the world's largest and most important migration flyways, and the factors controlling it. This information is essential for predicting bird migration, which-especially under the ongoing anthropogenic changes-is of high importance.

Age differences in primary moult of the Black-headed Gull Chroicocephalus ridibundus

Flight feather moult is an energetically expensive stage of the annual cycle of birds. Its timing is adjusted to other important time- and energy-demanding activities, including migration. In the vast majority of migratory birds, primary moult occurs before or after migration or moulting is suspended during migration. The Black-headed Gull is an exception: it starts moulting flight feathers at the onset of the autumn migration. The course of primary moult in this species is still poorly understood due to the difficulty in obtaining sufficient data from the entire period of flight feather replacement. In this paper, we employed digital photos taken in the field to estimate the start, variation in the start date and duration of primary moult in adult and immature Black-headed Gulls, by using the Underhill–Zucchini likelihood moult model. On average, immatures started their moult on 7 June, 25 days earlier than adults, but their moult lasted 5 days longer. Consequently, there was a 20-day difference in the end of moult – that is, on average it occurred on 16 September and 6 October in immature and adult gulls, respectively. Adults showed greater variability in the mean primary moult start date than immatures, as the beginning of flight feather replacement may depend on breeding success (earlier after breeding failure) and on breeding phenology. The overlap of primary moult and migration produces a trade-off in resource allocation between those two processes, which leads to a low number of simultaneously growing primaries and a decrease in the growth rate of the two external, heaviest flight feathers. Existing reports of a later completion of primary moult in the first half of November in this species are based on the faulty assumption that the outermost primary is always the longest, which is not the case in 34% of individuals.

[On the origin of the name Girifalco, an Italian town in a hotspot of the autumn migration of soaring raptors]

[On the origin of the name Girifalco, an Italian town in a hotspot of the autumn migration of soaring raptors]

Quantifying nocturnal thrush migration using sensor data fusion between acoustics and vertical‐looking radar

AbstractStudying nocturnal bird migration is challenging because direct visual observations are difficult during darkness. Radar has been the means of choice to study nocturnal bird migration for several decades, but provides limited taxonomic information. Here, to ascertain the feasibility of enhancing the taxonomic resolution of radar data, we combined acoustic data with vertical‐looking radar measurements to quantify thrush (Family: Turdidae) migration. Acoustic recordings, collected in Helsinki between August and October of 2021–2022, were used to identify likely nights of high and low thrush migration. Then, we built a random forest classifier that used recorded radar signals from those nights to separate all migrating passerines across the autumn migration season into thrushes and non‐thrushes. The classifier had a high overall accuracy (≈0.82), with wingbeat frequency and bird size being key for separation. The overall estimated thrush autumn migration phenology was in line with known migratory patterns and strongly correlated (Pearson correlation coefficient ≈0.65) with the phenology of the acoustic data. These results confirm how the joint application of acoustic and vertical‐looking radar data can, under certain migratory conditions and locations, be used to quantify ‘family‐level’ bird migration.

Constitutive immune function is not associated with fuel stores in spring migrating passerine birds.

Migratory birds may either upregulate their immune system during migration as they might encounter novel pathogens or downregulate their immune system as a consequence of trade-offs with the resource costs of migration. Support for the latter comes not least from a study that reports a positive correlation in autumn migrating birds between fuel stores and parameters of innate and acquired immune function, that is, energy-exhausted migrants appear to have lowered immune function. However, to our knowledge, no study has tested whether this pattern exists in spring migrating birds, which may face other trade-offs than autumn migrants. Here, we investigate if in spring there is a relationship between fuel stores and microbial-killing ability, a measure of innate immune function, and total immunoglobulin (IgY), a measure of acquired immune function, in four migrating bird species: chaffinches (Fringilla coelebs), dunnocks (Prunella modularis), song thrushes (Turdus philomelos) and northern wheatears (Oenanthe oenanthe). Our findings indicate no significant correlation between fuel stores and either microbial killing ability or IgY levels when considering all species collectively. When analysing species separately, we found a significant negative correlation between fuel stores and microbial-killing ability in chaffinches and a positive correlation between fuel stores and IgY levels in wheatears. In song thrushes, but not in any of the other species, there was a significant negative correlation between relative arrival date and microbial-killing ability and between arrival date and IgY levels. Sex did not affect immune function in any of the species. Our study suggests that the relationship between immune function and fuel stores may be different during spring migration compared to autumn migration. Differences in the speed of migration or pathogen pressure may result in different outcomes of the resource trade-off between investment in immune function and migration among the seasons.

Seasonal patterns and processes of migration in a long-distance migratory bird: energy or time minimization?

Optimal migration theory prescribes adaptive strategies of energy, time or mortality minimization. To test alternative hypotheses of energy- and time-minimization migration we used multisensory data loggers that record time-resolved flight activity and light for positioning by geolocation in a long-distance migratory shorebird, the little ringed plover, Charadrius dubius. We could reject the hypothesis of energy minimization based on a relationship between stopover duration and subsequent flight time as predicted for a time minimizer. We found seasonally diverging slopes between stopover and flight durations in relation to the progress (time) of migration, which follows a time-minimizing policy if resource gradients along the migration route increase in autumn and decrease in spring. Total flight duration did not differ significantly between autumn and spring migration, although spring migration was 6% shorter. Overall duration of autumn migration was longer than that in spring, mainly owing to a mid-migration stop in most birds, when they likely initiated moult. Overall migration speed was significantly different between autumn and spring. Migratory flights often occurred as runs of two to seven nocturnal flights on adjacent days, which may be countering a time-minimization strategy. Other factors may influence a preference for nocturnal migration, such as avoiding flight in turbulent conditions, heat stress and diurnal predators.

Population size and nesting peculiarities of the black-headed gull Chroicocephalus ridibundus (Linnaeus, 1766) on the territory of water treatment facilities

Background. Today, the black-headed gull inhabits man-made areas of wastewater treatment facilities (WTF) to comensate for the the reduction of natural aquatic and wetland habitats. Over the last decade, a nearly tenfold increase in its population has been recorded, despite a low reproduction rate. This fact indicates the lack of stability in the bird population, necessitating thorough research. Materials and Methods. The analysis of the population size and biological charac­teristics of black-headed gulls involved censuses and observations at the WTF of the city of Kharkiv using conventional methods during the spring-summer periods of 2020–2021 and 2023. Results. The population of the black-headed gull reached its peak in the third decade of May 2020 (2637 individuals) and 2023 (2124 individuals), as well as in the second decade of May 2021 (3949 individuals). The maximum nesting density was observed on sludge sites (SS) of wastewater treatment facilities that are most similar to natural habitats, where dried mud alternates with water patches and vegetation at the bottom and around the perimeter (Type V): 236.7±26.7 pairs/ha in 2020 and 242.9±28.5 pairs/ha in 2021. The majority of nests were found in the first decade of May 2021 and the third decade of May 2020. The black-headed gull forms mixed-species, occasionally monospecific subcolonies. Nesting in association with it were: Sterna hirundo, Anas platyrhynchos, Spatula clypeata, Vanellus vanellus, Charadrius dubius, Himantopus himantopus, Fulica atra, Gallinula chloropus, and Aythya ferina. The size of the complete black-headed gull clutch was 3.1±0.4 eggs (n = 190). The egg dimensions were 50.9±2.1 ´ 36.2±1.1, with a mass of 34.6±2.8 g. Regarding shell coloration, five types of background colors were identified. Mass egg laying occurred from the third decade of April to the first decade of May, constituting 40.3 % (n = 993) in 2020 and 62.3 % (n = 1757) in 2021. The egg-laying period extended from April to July, with the latest non-incubated clutches recorded in the first decade of July (3.07.2020). The first chicks were found in nests in the first decade of May, while mass hatching occurred in the second half of May. The latest registration dates of the birds on nesting territories were in the second decade of July for 2020 and 2023, and the first decade of August for 2022. The nesting season duration varied from 125 to 140 days in different years. Autumn migration commenced with summer relocations and concluded from late October to early November. The reproductive success – the percentage of nestlings that fledged and successfully achieved flight – constituted 29.2 % (n = 2404 of laid eggs) in 2020, and 15.5 % (n = 6138) in 2021. The majority of offspring perished due to changes in water levels (prolonged rainfall or industrial wastewater discharge), predation, disturbance factors, etc. Conclusion. The colonial nesting of the black-headed gull creates favorable conditions for the habitation and reproduction of various bird species, including rare ones, which is essential for their conservation.

Spatiotemporal distribution of Eurasian Spoonbills (Platalea leucorodia) outside the Carpathian Basin – the results of the Hungarian colour-ringing project based on twenty years (2003–2023)

Abstract The Eurasian Spoonbill (Platalea leucorodia) is a migratory waterbird. We used the confirmed observations of colour-ringed individuals (2,735 specimens) of the Hungarian population collected outside the Carpathian Basin between May 2003 and February 2023. 546 Spoonbills occurred in 28 countries. They used mainly the Central Mediterranean Flyway, however, some individuals were seen along the East Atlantic Flyway or the East Mediterranean Flyway. Three individuals were observed north of the Alps, too. A small proportion of Spoonbills crossed the Sahara and they occurred in the Sahel zone, between Sudan in the east and Senegal and Mauritania in the west. A few individuals were observed on Saharan wetlands in Algeria during winter. The most important destination for the Hungarian (Pannonian) population during the migration is Tunisia, where many of them spend the winter, primarily in the tidal area of the Gulf of Gabes. The first adults reached North Africa (Tunisia) on 28 July during their southward migration, while the last ones stayed there until 23 April. The first observation of juveniles in North Africa (Tunisia) happened on 31 August. Immature, 2- and 3-cyold birds spend the summer in unknown numbers in Tunisia. The majority of Pannonian Spoonbills migrate through the Balkans and Italy and some stay there to winter, mainly in Italy. Some of the immatures spent the summer in Italy. In southern Europe, the peak of the spring migration was in March, and the peak of the autumn migration was in September. There was also a difference in the migration of different age groups: in Southern Europe, the migration peak of adult birds falls between March and April, while most of the immatures were observed in May and June during northward migration. The peak of the southward migration in all age groups was observed in September. In the southern part of Europe, there may still be adults migrating north on 13 May, while other adults may already migrate southward on 15 May. The earliest juvenile migrating south was documented on 30 June in Italy.

Year-round carryover effects are driven by migration phenology for Hirundo rustica (Barn Swallow) wintering in West Africa

Abstract Recently, population declines have been reported for many migratory birds. Because of complex life cycles, determining the causes for such declines is often difficult. Thus, migratory birds are of special conservation interest. We studied the migratory behavior of Hirundo rustica (Barn Swallow) tagged with solar geolocators and determined carryover effects during the entire annual cycle from one breeding season to the next. We used a partial least squares path model (PLS-PM) to disentangle migratory and breeding events that occur in chronological order. In addition, we controlled for broad environmental conditions in the wintering grounds (NDVI and latitude) and the specific molting habitat (δ13C). We did not find a carryover effect from reproduction investment in the attachment year to breeding success in the subsequent year. Individuals that invested more in reproduction departed earlier from the breeding colonies, but this in turn did not affect the onset of autumn migration. Thus, the premigratory period should be acting as a buffer stage counteracting any previous carryover effects from reproduction investment. On the other hand, we found a long-lasting domino effect from the onset of autumn migration to subsequent breeding success, consistent with the notion of a migratory race. Specifically, individuals which started earlier in the autumn migration arrived earlier to the wintering grounds, started earlier in the spring migration arrived earlier to the breeding colonies, and had a higher breeding success. We highlight that the premigratory period (i.e., the time elapsed between departure from breeding areas and the onset of autumn migration) should be important for the life cycle of migratory species, but it has been frequently overlooked.

Macro-scale relationship between body mass and timing of bird migration

Clarifying migration timing and its link with underlying drivers is fundamental to understanding the evolution of bird migration. However, previous studies have focused mainly on environmental drivers such as the latitudes of seasonal distributions and migration distance, while the effect of intrinsic biological traits remains unclear. Here, we compile a global dataset on the annual cycle of migratory birds obtained by tracking 1531 individuals and 177 populations from 186 species, and investigate how body mass, a key intrinsic biological trait, influenced timings of the annual cycle using Bayesian structural equation models. We find that body mass has a strong direct effect on departure date from non-breeding and breeding sites, and indirect effects on arrival date at breeding and non-breeding sites, mainly through its effects on migration distance and a carry-over effect. Our results suggest that environmental factors strongly affect the timing of spring migration, while body mass affects the timing of both spring and autumn migration. Our study provides a new foundation for future research on the causes of species distribution and movement.

Fluctuations in population size and migration of two species of crossbills in relation to seed crop size of spruce and pine: asymmetric importance of alternative food resources

Bird species depending on seeds from masting trees face large temporal and spatial variation in food availability and perform nomadic or irruptive movements to locate food. In northern Europe, the Common Crossbill (Loxia curvirostra) is specialized on seeds from Norway Spruce (Picea abies) which usually has peak years followed by crop failure. Common Crossbills breed in mast years and emigrate when spruce seeds are no longer available. On the other hand, the Parrot Crossbill (L. pytyopsittacus) is specialized on seeds from Scots Pine (Pinus sylvestris) in which seed production is more stable from year to year. The Parrot Crossbill is therefore thought to be more sedentary. However, both species may switch food if there is low abundance of their 'own' seed type, but little is known about the relative importance of crop size of the two tree species on breeding population size and migration of the two crossbill species. Here, we analyse time series (up to 36 years) of crossbills and crop size of spruce and pine in Norway and Sweden to investigate this. We found that breeding population sizes of both species were positively related to spruce crop size. Surprisingly, pine crop size had no positive influence, not even on the Parrot Crossbill. Emigration of both crossbill species (based on autumn migration counts in southern Sweden) increased when there were small seed crops of spruce, but pine crop size had no influence, not even on the Parrot Crossbill. There was no influence of population size on emigration. These results suggest that alternative food resources had an asymmetric effect, only influencing the Parrot Crossbill. This asymmetry explains why the two crossbill species often have synchronous eruptions despite being specialized on different tree species with different seed crop size dynamics.

Migration increases mitochondrial oxidative capacity without increasing reactive oxygen species emission in a songbird.

Birds remodel their flight muscle metabolism prior to migration to meet the physiological demands of migratory flight, including increases in both oxidative capacity and defence against reactive oxygen species. The degree of plasticity mediated by changes in these mitochondrial properties is poorly understood but may be explained by two non-mutually exclusive hypotheses: variation in mitochondrial quantity or in individual mitochondrial function. We tested these hypotheses using yellow-rumped warblers (Setophaga coronata), a Nearctic songbird which biannually migrates 2000-5000 km. We predicted higher flight muscle mitochondrial abundance and substrate oxidative capacity, and decreased reactive oxygen species emission in migratory warblers captured during autumn migration compared with a short-day photoperiod-induced non-migratory phenotype. We assessed mitochondrial abundance via citrate synthase activity and assessed isolated mitochondrial function using high-resolution fluororespirometry. We found 60% higher tissue citrate synthase activity in the migratory phenotype, indicating higher mitochondrial abundance. We also found 70% higher State 3 respiration (expressed per unit citrate synthase) in mitochondria from migratory warblers when oxidizing palmitoylcarnitine, but similar H2O2 emission rates between phenotypes. By contrast, non-phosphorylating respiration was higher and H2O2 emission rates were lower in the migratory phenotype. However, flux through electron transport system complexes I-IV, II-IV and IV was similar between phenotypes. In support of our hypotheses, these data suggest that flight muscle mitochondrial abundance and function are seasonally remodelled in migratory songbirds to increase tissue oxidative capacity without increasing reactive oxygen species formation.