Age and sex effects on spring migration timing in great reed warblers: early start allows older males to arrive first

Migratory birds travel vast distances and the timing of migratory flights can affect survival and the ability to reproduce. For Neotropical migrant songbirds, early spring departure from wintering sites, early arrival to the breeding grounds and higher reproductive success have been related to the use of suitable habitats and environmental conditions during the non-breeding season. However, how migratory strategies are shaped by winter habitat choice is largely unknown due to the general inability to track birds from specific wintering habitats to stopovers or breeding destinations. We assessed how winter habitat (native forest vs. shade-grown coffee plantations) relates to spring departure date and migration pace in Swainson's Thrush Catharus ustulatus. We also determined the effect of departure date and total migration duration on the arrival date of birds detected near or within their breeding range. We used a novel application of Motus radiotelemetry arrays to track individuals from their wintering grounds in the Andes of South America along their migratory journey to North America. We found variation in migratory strategies between habitats, with birds wintering in native forest departing later than birds in coffee. We present isotopic evidence for native forest being of higher quality than shade-coffee for Swainson's Thrush and hypothesize that moister conditions in forest, as shown by stable isotope (δ13 C) analysis of thrush whole blood, provides favourable pre-migratory conditions allowing birds to delay departure from wintering grounds. Habitat, between-site and -year variation in departure date, suggests that birds made facultative adjustments to winter habitat quality and environmental conditions. Independent of habitat, birds that departed later migrated faster and this pattern was maintained along the migration route (n=44). Migrating earlier and slower or later and faster was unlikely to result in significant differences in arrival time to breeding destinations. Our findings reveal underappreciated complexity in migratory decisions by long-distance migrants that contrast with the current paradigm of earlier departures and arrival from optimal habitats. The next step is to understand the relative fitness benefits of early versus late schedules or whether each strategy is an equally good response to experienced conditions.

Numerous factors influence the timing of spring migration in birds, yet the relative importance of intrinsic and extrinsic variables on migration initiation remains unclear. To test for interactions among weather, migration distance, parasitism, and physiology in determining spring departure date, we used the Dark-eyed Junco (Junco hyemalis) as a model migratory species known to harbor diverse and common haemosporidian parasites. Prior to spring migration departure from their wintering grounds in Indiana, USA, we quantified the intrinsic variables of fat, body condition (i.e., mass ~ tarsus residuals), physiological stress (i.e., ratio of heterophils to lymphocytes), cellular immunity (i.e., leukocyte composition and total count), migration distance (i.e., distance to the breeding grounds) using stable isotopes of hydrogen from feathers, and haemosporidian parasite intensity. We then attached nanotags to determine the timing of spring migration departure date using the Motus Wildlife Tracking System. We used additive Cox proportional hazard mixed models to test how risk of spring migratory departure was predicted by the combined intrinsic measures, along with meteorological predictors on the evening of departure (i.e., average wind speed and direction, relative humidity, and temperature). Model comparisons found that the best predictor of spring departure date was average nightly wind direction and a principal component combining relative humidity and temperature. Juncos were more likely to depart for spring migration on nights with largely southwestern winds and on warmer and drier evenings (relative to cooler and more humid evenings). Our results indicate that weather conditions at take-off are more critical to departure decisions than the measured physiological and parasitism variables.

The long-term advance in the timing of bird spring migration in the Northern Hemisphere is associated with global climate change. The extent to which changes in bird phenology reflect responses to weather conditions in the wintering or breeding areas, or during migration, however, remains to be elucidated. We analyse the relationships between the timing of spring migration of 9 species of trans-Saharan migratory birds across the, Mediterranean, and thermal and precipitation anomalies in the main wintering areas south of the Sahara Desert and in North African stopover areas. Median migration dates were collected on the island of Capri (southern Italy) by standardized mist-netting during 1981 to 2004. High temperatures in sub-Saharan Africa (Sahel and Gulf of Guinea) prior to northward migration (February and March) were associated with advanced migration. Moreover, birds migrated earlier when winter rainfall in North Africa was more abundant. The relationships between relevant meteorological variables and timing of migration were remarkably consistent among species, suggesting a coherent response to the same extrinsic stimuli. All these results were obtained while statistically controlling for the long-term trend towards the earlier timing of spring migration across the Mediterranean that has been documented in previous analyses of the same dataset, a trend that was confirmed by the present analyses. In conclusion, our results suggest that thermal conditions in the wintering quarters, as well as rainfall in North African stopover areas, can influence interannual variation in migration phenology of trans-Saharan migratory birds, although the ecological mechanisms that causally link meteorological conditions to the timing of migration remain a matter of speculation.

Drivers of Departure and Prospecting in Dispersing Juvenile Red Kites (Milvus milvus)

In April 2011, the British Ornithologist’s Union’s annual conference took place at the University of Leicester. The theme of this year’s conference was the ecology and conservation of migratory birds, and talks ranged from those focussed on either breeding, migration or wintering ecology, to those linking two or more of these and trying to build a more complete picture of the complex problems faced by long distance migrants. Talks also covered recent technological advances, which are increasingly allowing us intriguing new insights, particularly into the seasonal movements and wintering areas of some of the most declining species. Finally, talks examined the policy challenges associated with protecting migratory species and how best to tackle flyway-scale conservation of migratory birds.

Global climate change is having a major impact on the viability of species. Whether a particular population will survive or go extinct will depend on its potential to adapt to new environmental conditions. In migratory birds, the timing of spring migration is one of the major determinants of the timing of reproduction and as spring temperatures increase, we expect birds to return earlier to their breeding grounds. In accord with this expectation, the timing of spring arrival has advanced in many bird populations within the last 30 years. Yet, despite the presence of genetic variation in the timing of migration and fitness advantages in birds arriving early we still lack conclusive evidence for evolutionary change. One reason for this may be that weather conditions en route, particular the direction and speed of winds, have an important influence on individual migration speed. Moreover, there is strong evidence that physical condition may modify the timing of departure from the wintering grounds, migration speed and ultimately the timing of arrival and breeding. We also expect nestling condition and maternal effects to influence spring migration timing. Long-term population studies monitoring individual arrival in combination with experimental approaches will be needed to tease apart genetic and different environmental effects on the traits determining migration timing. This knowledge will be crucial for predicting adaptive responses of the timing of spring arrival in the future.

BackgroundThe arrival of many species of migrant passerine in the European spring has shifted earlier over recent decades, attributed to climate change and rising temperatures in Europe and west Africa. Few studies have shown the effects of climate change in both hemispheres though many long-distance migrants use wintering grounds which span Africa. The migrants’ arrival in Europe thus potentially reflects a combination of the conditions they experience across Africa. We examine if the timing of spring migration of a long-distance migrant, the Willow Warbler, is related to large-scale climate indices across Africa and Europe.MethodsUsing data from daily mistnetting from 1 April to 15 May in 1982–2017 at Bukowo (Poland, Baltic Sea coast), we developed an Annual Anomaly metric (AA, in days) to estimate how early or late Willow Warblers arrive each spring in relation to their multi-year average pattern. The Willow Warblers’ spring passage advanced by 5.4 days over the 36 years. We modelled AA using 14 potential explanatory variables in multiple regression models. The variables were the calendar year and 13 large-scale indices of climate in Africa and Europe averaged over biologically meaningful periods of two to four months during the year before spring migration.ResultsThe best model explained 59% of the variation in AA with seven variables: Northern Atlantic Oscillation (two periods), Indian Ocean Dipole, Southern Oscillation Index, Sahel Precipitation Anomaly, Scandinavian Index and local mean temperatures. The study also confirmed that a long-term trend for Willow Warblers to arrive earlier in spring continued up to 2017.DiscussionOur results suggest that the timing of Willow Warbler spring migration at the Baltic Sea coast is related to a summation of the ecological conditions they had encountered over the previous year during breeding, migration south, wintering in Africa and migration north. We suggest these large-scale climate indices reflect ecological drivers for phenological changes in species with complex migration patterns and discuss the ways in which each of the seven climate indices could be related to spring migration at the Baltic Sea coast.

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.

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.

Arrival timing in spring may be mediated by conditions experienced during migratory stopovers or staging areas, but our knowledge about their impact on migration timing and reproduction is limited. We explored the role of stopover duration on spring migration timing and successful incubation in cackling geese (Branta hutchinsii), which breed at high arctic latitudes where climate change effects are more pronounced. To track migration phenology and incubation duration, 236 light-level geolocators were deployed on cackling geese during the breeding period at Baffin Island, Nunavut, Canada between 2016 and 2018. Using data available for spring migration in the year following tag deployment (25 tags retrieved), we found that most geese had long, coastal stopovers (8–39 days) before crossing Hudson Bay on the last leg of their spring migration to their Baffin Island breeding area. We show that longer stopover durations at these Hudson Bay Lowland sites were associated with successful completion of incubation (a proxy for breeding success). Although spending more time at the stopover led to a later arrival date at the breeding ground, longer stopovers may increase the energy stores necessary for reproduction in these capital breeders. Stopover duration did not influence the incubation interval (number of days between arrival date at the breeding ground and start of incubation). Lastly, we found that the temperature at stopovers influenced migration timing, with higher temperatures resulting in earlier arrival at the breeding ground. Overall, our results demonstrate that conditions and behaviour at distant stopovers (1700–3000 km away) have important influence on timing and breeding success once birds arrive at their Arctic breeding sites. Therefore, our understanding of climate change impacts on these Arctic-breeding geese must also include the influence of en route conditions

Environmental factors influencing the timing of bird migration frequently affect age and sex groups differently. We applied discriminant functions to determine the sex of adult Song Thrushes ringed during spring migration in 1968–2019 at Hel on the Polish Baltic coast. We aimed to determine any differences between the sexes in the timing of spring migration and changes in protandry, and to identify any relationships between migration timing and temperatures at the wintering grounds. We analysed relationships between the date of migration and the year and monthly mean minimum temperatures in December–February by sex, using multiple quantile regression for q10, q50 and q90 of spring passage. On average males migrated six days before females. Spring protandry increased over 1968–2019 and with warm winters. The sexes responded differently to winter temperatures, likely because of the species’ mating strategy. For males the warmer the winter, especially in February, the earlier spring migration began and the later it ended, prolonging its duration. The influence of February’s temperature on the beginning of males’ migration is likely explained by selective pressure for their early arrival to establish territories. The later end of their arrival with a warm winter might reflect more low-quality males surviving, and migrating later than high-quality males, than with cold winters. For females December and January temperatures had no effect on the beginning and the end of spring migration; but the warmer the February, the earlier migration started and the later it ended. Females probably benefitted from improved conditions in a warm February in two ways: early-arriving females maximized their breeding success with a chance to raise a second brood, and late-arriving females maximized their chances of surviving migration by prolonging fuelling and stopovers, which extended females’ arrivals.

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

The question “Which factors govern the timing of migration in birds?” has fascinated researchers for a long time. It was initially assumed that avian migration is triggered by environmental factors, such as ambient temperature and food availability. Later laboratory experiments in various avian species convincingly showed that timing of spring migration is mainly governed by daylength (photoperiod) and is controlled by circannual rhythms. As a result, the concept that environmental factors (air temperature, precipitation, food availability) have no significant impact on timing of spring migration generally took hold. However, in recent decades more and more data has become available showing that the timing of spring migration in many bird species has significantly changed. These data allow the formulation of a novel concept of regulation mechanisms of timing of spring migration which accounts not only for photoperiodic and endogenous control, but also for the already mentioned extrinsic factors. Studies of endocrine control of spring migratory disposition showed that features of endocrine mechanisms governing the onset of spring migration depend on speciesspecific migratory strategies and the stability of environmental conditions in winter quarters and on migratory routes. It is becoming clear precisely which endocrine mechanisms are involved in adjusting migratory behaviour to variation of the local environment. In recent years, progress has also been made in finding genetic mechanisms controlling the timing of spring migration.

Climate change has advanced the breeding dates of many bird species, but for few species we know whether this advancement is sufficient to track the advancement of the underlying levels of the food chain. For the long‐distance migratory pied flycatcher Ficedula hypoleuca the advancement in breeding time has been insufficient to maintain the synchrony with their main food sources. The timing of arrival in the breeding areas from their African wintering grounds is likely to constrain the advancement of breeding date. We hypothesise that this is because in Africa they cannot predict the advancement of spring in their breeding habitat. However, long‐distance migrants may advance their arrival time by migrating faster when circumstances en route are favourable. In this study we show that both arrival and breeding date depend on temperatures at their main North African staging grounds, as well as on temperature at the breeding grounds. Male arrival and average laying date were not correlated, but the positive effect of temperature in North Africa on breeding dates suggests that breeding date is indeed constrained by arrival of females. Long‐distance migrants thus are able to adjust arrival and hence breeding by faster spring migration, but the degree of adjustment is probably limited as timing schedules in spring are tight. Furthermore, as climate change is affecting temperatures differently along the migratory flyway and the breeding areas, it is unlikely that arrival dates are advanced at the same rate as the timing of breeding should advance, given the advancement of the underlying levels of the food chain.

Speciation Associated with Shifts in Migratory Behavior in an Avian Radiation.