35446867
Quantifying full phenological event distributions reveals simultaneous advances, temporal stability and delays in spring and autumn migration timing in long‐distance migratory birds
Phenological changes in key seasonally expressed life-history traits occurring across periods of climatic and environmental change can cause temporal mismatches between interacting species, and thereby impact population and community dynamics. However, studies quantifying long-term phenological changes have commonly only measured variation occurring in spring, measured as the first or mean dates on which focal traits or events were observed. Few studies have considered seasonally paired events spanning spring and autumn or tested the key assumption that single convenient metrics accurately capture entire event distributions. We used 60 years (1955-2014) of daily bird migration census data from Fair Isle, Scotland, to comprehensively quantify the degree to which the full distributions of spring and autumn migration timing of 13 species of long-distance migratory bird changed across a period of substantial climatic and environmental change. In most species, mean spring and autumn migration dates changed little. However, the early migration phase (≤10th percentile date) commonly got earlier, while the late migration phase (≥90th percentile date) commonly got later. Consequently, species' total migration durations typically lengthened across years. Spring and autumn migration phenologies were not consistently correlated within or between years within species and hence were not tightly coupled. Furthermore, different metrics quantifying different aspects of migration phenology within seasons were not strongly cross-correlated, meaning that no single metric adequately described the full pattern of phenological change. These analyses therefore reveal complex patterns of simultaneous advancement, temporal stability and delay in spring and autumn migration phenologies, altering species' life-history structures. Additionally, they demonstrate that this complexity is only revealed if multiple metrics encompassing entire seasonal event distributions, rather than single metrics, are used to quantify phenological change. Existing evidence of long-term phenological changes detected using only one or two metrics should consequently be interpreted cautiously because divergent changes occurring simultaneously could potentially have remained undetected.
15144885
Toward a synthetic understanding of the role of phenology in ecology and evolution
Phenology affects nearly all aspects of ecology and evolution. Virtually all biological phenomena—from individual physiology to interspecific relationships to global nutrient fluxes—have annual cycles and are influenced by the timing of abiotic events. Recent years have seen a surge of interest in this topic, as an increasing number of studies document phenological responses to climate change. Much recent research has addressed the genetic controls on phenology, modelling techniques and ecosystem-level and evolutionary consequences of phenological change. To date, however, these efforts have tended to proceed independently. Here, we bring together some of these disparate lines of inquiry to clarify vocabulary, facilitate comparisons among habitat types and promote the integration of ideas and methodologies across different disciplines and scales. We discuss the relationship between phenology and life history, the distinction between organismal- and population-level perspectives on phenology and the influence of phenology on evolutionary processes, communities and ecosystems. Future work should focus on linking ecological and physiological aspects of phenology, understanding the demographic effects of phenological change and explicitly accounting for seasonality and phenology in forecasts of ecological and evolutionary responses to climate change.
448284
The effects of phenological mismatches on demography
Climate change is altering the phenology of species across the world, but what are the consequences of these phenological changes for the demography and population dynamics of species? Time-sensitive relationships, such as migration, breeding and predation, may be disrupted or altered, which may in turn alter the rates of reproduction and survival, leading some populations to decline and others to increase in abundance. However, finding evidence for disrupted relationships, or lack thereof, and their demographic effects, is difficult because the necessary detailed observational data are rare. Moreover, we do not know how sensitive species will generally be to phenological mismatches when they occur. Existing long-term studies provide preliminary data for analysing the phenology and demography of species in several locations. In many instances, though, observational protocols may need to be optimized to characterize timing-based multi-trophic interactions. As a basis for future research, we outline some of the key questions and approaches to improving our understanding of the relationships among phenology, demography and climate in a multi-trophic context. There are many challenges associated with this line of research, not the least of which is the need for detailed, long-term data on many organisms in a single system. However, we identify key questions that can be addressed with data that already exist and propose approaches that could guide future research.
46398411
How Climate Change Alters Rhythms of the Wild
The more scientists look, the more connections they see between shifts in climate and changes in animal behavior and populations. On page
 854, researchers report that while the number of dippers in a population in southern Norway fluctuated between 1978 and 1997, it followed an upward trajectory that was closely associated with an atmospheric pressure system called the North Atlantic Oscillation and the milder winter temperatures it brought. These and other findings pouring in could have huge implications for conservation.
1176350
Ecological responses to recent climate change
There is now ample evidence of the ecological impacts of recent climate change, from polar terrestrial to tropical marine environments. The responses of both flora and fauna span an array of ecosystems and organizational hierarchies, from the species to the community levels. Despite continued uncertainty as to community and ecosystem trajectories under global change, our review exposes a coherent pattern of ecological change across systems. Although we are only at an early stage in the projected trends of global warming, ecological responses to recent climate change are already clearly visible.
1190097
A globally coherent fingerprint of climate change impacts across natural systems
Causal attribution of recent biological trends to climate change is complicated because non-climatic influences dominate local, short-term biological changes. Any underlying signal from climate change is likely to be revealed by analyses that seek systematic trends across diverse species and geographic regions; however, debates within the Intergovernmental Panel on Climate Change (IPCC) reveal several definitions of a ‘systematic trend’. Here, we explore these differences, apply diverse analyses to more than 1,700 species, and show that recent biological trends match climate change predictions. Global meta-analyses documented significant range shifts averaging 6.1 km per decade towards the poles (or metres per decade upward), and significant mean advancement of spring events by 2.3 days per decade. We define a diagnostic fingerprint of temporal and spatial ‘sign-switching’ responses uniquely predicted by twentieth century climate trends. Among appropriate long-term/large-scale/multi-species data sets, this diagnostic fingerprint was found for 279 species. This suite of analyses generates ‘very high confidence’ (as laid down by the IPCC) that climate change is already affecting living systems.
4428020
Spatial patterns in species distributions reveal biodiversity change
Interpretation of global biodiversity change is hampered by a lack of information on the historical status of most species in most parts of the world. Here we show that declines and increases can be deduced from current species distributions alone, using spatial patterns of occupancy combined with distribution size. Declining species show sparse, fragmented distributions for their distribution size, reflecting the extinction process; expanding species show denser, more aggregated distributions, reflecting colonization. Past distribution size changes for British butterflies were deduced successfully from current distributions, and former distributions had some power to predict future change. What is more, the relationship between distribution pattern and change in British butterflies independently predicted distribution change for butterfly species in Flanders, Belgium, and distribution change in British rare plant species is similarly related to spatial distribution pattern. This link between current distribution patterns and processes of distribution change could be used to assess relative levels of threat facing different species, even for regions and taxa lacking detailed historical and ecological information.
4414217
Climate change and population declines in a long-distance migratory bird
Phenological responses to climate change differ across trophic levels, which may lead to birds failing to breed at the time of maximal food abundance. Here we investigate the population consequences of such mistiming in the migratory pied flycatcher, Ficedula hypoleuca. In a comparison of nine Dutch populations, we find that populations have declined by about 90% over the past two decades in areas where the food for provisioning nestlings peaks early in the season and the birds are currently mistimed. In areas with a late food peak, early-breeding birds still breed at the right time, and there is, at most, a weak population decline. If food phenology advances further, we also predict population declines in areas with a late food peak, as in these areas adjustment to an advanced food peak is insufficient. Mistiming as a result of climate change is probably a widespread phenomenon, and here we provide evidence that it can lead to population declines.
44236521
Ecological and Evolutionary Responses to Recent Climate Change
Ecological changes in the phenology and distribution of plants and animals are occurring in all well-studied marine, freshwater, and terrestrial groups. These observed changes are heavily biased in the directions predicted from global warming and have been linked to local or regional climate change through correlations between climate and biological variation, field and laboratory experiments, and physiological research. Range-restricted species, particularly polar and mountaintop species, show severe range contractions and have been the first groups in which entire species have gone extinct due to recent climate change. Tropical coral reefs and amphibians have been most negatively affected. Predator-prey and plant-insect interactions have been disrupted when interacting species have responded differently to warming. Evolutionary adaptations to warmer conditions have occurred in the interiors of species’ ranges, and resource use and dispersal have evolved rapidly at expanding range margins. Observed genetic shifts modulate local effects of climate change, but there is little evidence that they will mitigate negative effects at the species level.
7657509
Potential impacts of climate change on the winter distribution of Afro-Palaearctic migrant passerines
We modelled the present and future sub-Saharan winter distributions of 64 trans-Saharan migrant passerines to predict the potential impacts of climate change. These predictions used the recent ensemble modelling developments and the latest IPCC climatic simulations to account for possible methodological uncertainties. Results suggest that 37 species would face a range reduction by 2100 (16 of these by more than 50%); however, the median range size variation is −13 per cent (from −97 to +980%) under a full dispersal hypothesis. Range centroids were predicted to shift by 500±373 km. Predicted changes in range size and location were spatially structured, with species that winter in southern and eastern Africa facing larger range contractions and shifts. Predicted changes in regional species richness for these long-distance migrants are increases just south of the Sahara and on the Arabian Peninsula and major decreases in southern and eastern Africa.
206525630
Global Biodiversity: Indicators of Recent Declines
Global Biodiversity Target Missed In 2002, the Convention on Biological Diversity (CBD) committed to a significant reduction in the rate of biodiversity loss by 2010. There has been widespread conjecture that this target has not been met. Butchart et al. (p. 1164, published online 29 April) analyzed over 30 indicators developed within the CBD's framework. These indicators include the condition or state of biodiversity (e.g., species numbers, population sizes), the pressures on biodiversity (e.g., deforestation), and the responses to maintain biodiversity (e.g., protected areas) and were assessed between about 1970 and 2005. Taken together, the results confirm that we have indeed failed to meet the 2010 targets. An analysis of 30 indicators shows that the Convention on Biological Diversity’s 2010 targets have not been met. In 2002, world leaders committed, through the Convention on Biological Diversity, to achieve a significant reduction in the rate of biodiversity loss by 2010. We compiled 31 indicators to report on progress toward this target. Most indicators of the state of biodiversity (covering species’ population trends, extinction risk, habitat extent and condition, and community composition) showed declines, with no significant recent reductions in rate, whereas indicators of pressures on biodiversity (including resource consumption, invasive alien species, nitrogen pollution, overexploitation, and climate change impacts) showed increases. Despite some local successes and increasing responses (including extent and biodiversity coverage of protected areas, sustainable forest management, policy responses to invasive alien species, and biodiversity-related aid), the rate of biodiversity loss does not appear to be slowing.
4358387
UK birds are laying eggs earlier
The evidence for global climate change and for its underlying anthropogenic causes is gathering rapidly. Over the past 11 years the active growing season of plants has advanced by roughly 8 days in northern latitudes. This evidence for increased photosynthetic activity is supported by the positive trend in the amplitude of the seasonal cycle in atmospheric CO2 (ref. 2). The phenology of animal populations should also be affected by climate change, but to date there has been little evidence of this. Here we report that long-term trends in the seasonal distributions of laying dates of birds in the United Kingdom show a tendency towards earlier laying, consistent with the changes reported in growing season.
4420436
Breeding phenology and climate⃛
European amphibians and birds have been breeding consistently earlier over the past two to three decades,. These changes have been attributed to the observed trends in increasing average spring temperatures in Europe producing earlier growing seasons and increased forage availability. Here we show that variations in breeding of European amphibians and birds are influenced by variations in a natural, large-scale atmospheric phenomenon, the North Atlantic Oscillation (NAO). Our results support the proximate cause (that is, increase in spring temperatures) of the altered breeding phenology as suggested previously,, but by extending previous analyses as well as integrating data from other bird species, they also suggest that organisms with complex life histories respond to year-to-year variations in their abiotic environment.
4312140
⃛temperature and egg-laying trends
Crick et al. showed that the laying dates for many British birds have become earlier during 1971-95, which they ascribe to warmer springs resulting from global warming. We provide partial support for this view from a great tit population in Oxfordshire.
85251036
Warmer springs lead to mistimed reproduction in great tits (Parus major)
In seasonal environments, the main selection pressure on the timing of reproduction (the ultimate factor) is synchrony between offspring requirements and food availability. However, reproduction is initiated much earlier than the time of maximum food requirement of the offspring. Individuals should therefore start reproduction in response to cues (the proximate factors), available in the environment of reproductive decision making, which predict the later environment of selection. With increasing spring temperatures over the past decades, vegetation phenology has advanced, with a concomitant advancement in the reproduction of some species at higher trophic levels. However, a mismatch between food abundance and offspring needs may occur if changes in the environment of decision making do not match those in the environment of selection. Date of egg laying in a great tit (Parus major) population has not advanced over a 23–year period, but selection for early laying has intensified. We believe that this is the first documented case of an adaptive response being hampered because a changing abiotic factor affects the environment in which a reproductive decision is made differently from the environment in which selection occurs.
86659608
Phenology of British butterflies and climate change
Summary 
Data from a national butterfly monitoring scheme were analysed to test for relationships between temperature and three phenological measures, duration of flight period and timing of both first and peak appearance. First appearances of most British butterflies has advanced in the last two decades and is strongly related to earlier peak appearance and, for multibrooded species, longer flight period. Mean dates of first and peak appearance are examined in relation to Manley's central England temperatures, using regression techniques. We predict that, in the absence of confounding factors, such as interactions with other organisms and land-use change, climate warming of the order of 1 °C could advance first and peak appearance of most butterflies by 2–10 days.
4365256
Adjustment to climate change is constrained by arrival date in a long-distance migrant bird
Spring temperatures in temperate regions have increased over the past 20 years, and many organisms have responded to this increase by advancing the date of their growth and reproduction. Here we show that adaptation to climate change in a long-distance migrant is constrained by the timing of its migratory journey. For long-distance migrants climate change may advance the phenology of their breeding areas, but the timing of some species’ spring migration relies on endogenous rhythms that are not affected by climate change. Thus, the spring migration of these species will not advance even though they need to arrive earlier on their breeding grounds to breed at the appropriate time. We show that the migratory pied flycatcher Ficedula hypoleuca has advanced its laying date over the past 20 years. This temporal shift has been insufficient, however, as indicated by increased selection for earlier breeding over the same period. The shift is hampered by its spring arrival date, which has not advanced. Some of the numerous long-distance migrants will suffer from climate change, because either their migration strategy is unaffected by climate change, or the climate in breeding and wintering areas are changing at different speeds, preventing adequate adaptation.
129692224
Observed changes in seasons: An overview
Within the last decade the study of phenology has taken on a new legitimacy in the area of climate change research. A growing literature reveals that a change in the timing of natural events is occurring in a wide range of locations and affecting a wide range of species. Changes in spring have been those most commonly reported, with the emphasis on an advance in spring linked to an increase in temperature. Detection of change in autumn is hampered by a smaller pool of available data, events that are harder to define (such as leaf coloration), and various influencing environmental factors triggering autumnal phases. Despite this, the general pattern may be towards a delay in autumn. Plant, animal and abiotic responses, especially in spring, are quite similar. Thus, it would appear that winter is being squeezed at both ends, and this effect, of increasing the growing season, should become more pronounced in the face of predicted global warming. Copyright © 2002 Royal Meteorological Society.
23594616
Shifts in phenology due to global climate change: the need for a yardstick
Climate change has led to shifts in phenology in many species distributed widely across taxonomic groups. It is, however, unclear how we should interpret these shifts without some sort of a yardstick: a measure that will reflect how much a species should be shifting to match the change in its environment caused by climate change. Here, we assume that the shift in the phenology of a species' food abundance is, by a first approximation, an appropriate yardstick. We review the few examples that are available, ranging from birds to marine plankton. In almost all of these examples, the phenology of the focal species shifts either too little (five out of 11) or too much (three out of 11) compared to the yardstick. Thus, many species are becoming mistimed due to climate change. We urge researchers with long-term datasets on phenology to link their data with those that may serve as a yardstick, because documentation of the incidence of climate change-induced mistiming is crucial in assessing the impact of global climate change on the natural world.
34408962
Shifting plant phenology in response to global change.
Plants are finely tuned to the seasonality of their environment, and shifts in the timing of plant activity (i.e. phenology) provide some of the most compelling evidence that species and ecosystems are being influenced by global environmental change. Researchers across disciplines have observed shifting phenology at multiple scales, including earlier spring flowering in individual plants and an earlier spring green-up' of the land surface revealed in satellite images. Experimental and modeling approaches have sought to identify the mechanisms causing these shifts, as well as to make predictions regarding the consequences. Here, we discuss recent advances in several fields that have enabled scaling between species responses to recent climatic changes and shifts in ecosystem productivity, with implications for global carbon cycling.
85900389
Trophic level asynchrony in rates of phenological change for marine, freshwater and terrestrial environments
Recent changes in the seasonal timing (phenology) of familiar biological events have been one of the most conspicuous signs of climate change. However, the lack of a standardized approach to analysing change has hampered assessment of consistency in such changes among different taxa and trophic levels and across freshwater, terrestrial and marine environments. We present a standardized assessment of 25 532 rates of phenological change for 726 UK terrestrial, freshwater and marine taxa. The majority of spring and summer events have advanced, and more rapidly than previously documented. Such consistency is indicative of shared large scale drivers. Furthermore, average rates of change have accelerated in a way that is consistent with observed warming trends. Less coherent patterns in some groups of organisms point to the agency of more local scale processes and multiple drivers. For the first time we show a broad scale signal of differential phenological change among trophic levels; across environments advances in timing were slowest for secondary consumers, thus heightening the potential risk of temporal mismatch in key trophic interactions. If current patterns and rates of phenological change are indicative of future trends, future climate warming may exacerbate trophic mismatching, further disrupting the functioning, persistence and resilience of many ecosystems and having a major impact on ecosystem services.
41962528
Autumn, the neglected season in climate change research.
Autumn remains a relatively neglected season in climate change research in temperate and arctic ecosystems. This neglect occurs despite the importance of autumn events, including leaf senescence, fruit ripening, bird and insect migration, and induction of hibernation and diapause. Changes in autumn phenology alter the reproductive capacity of individuals, exacerbate invasions, allow pathogen amplification and higher disease-transmission rates, reshuffle natural enemy-prey dynamics, shift the ecological dynamics among interacting species, and affect the net productivity of ecosystems. We synthesize some of our existing understanding of autumn phenology and identify five areas ripe for future climate change research. We provide recommendations to address common pitfalls in autumnal research as well as to support the conservation and management of vulnerable ecosystems and taxa.
38378636
Timing of autumn bird migration under climate change: advances in long–distance migrants, delays in short–distance migrants
As a response to increasing spring temperature in temperate regions in recent years, populations of many plant and animal species, including migratory birds, have advanced the seasonal start of their reproduction or growth. However, the effects of climate changes on subsequent events of the annual cycle remain poorly understood. We investigated long–term changes in the timing of autumn migration in birds, a key event in the annual cycle limiting the reproductive period. Using data spanning a 42–year period, we analysed long–term changes in the passage of 65 species of migratory birds through Western Europe. The autumn passage of migrants wintering south of the Sahara has advanced in recent years, presumably as a result of selection pressure to cross the Sahel before its seasonal dry period. In contrast, migrants wintering north of the Sahara have delayed autumn passage. In addition, species with a variable rather than a fixed number of broods per year have delayed passage, possibly because they are free to attempt more broods. Recent climate changes seem to have a simple unidirectional effect on the seasonal onset of reproduction, but complex and opposing effects on the timing of subsequent events in the annual cycle, depending on the ecology and life history of a species. This complicates predictions of overall effects of global warming on avian communities.
83657002
Can we detect change in the phenology of winter migrant birds in the UK?: Phenology of winter migrants
Although there is very strong evidence for an advance in the spring arrival dates of migrating birds, there are few data with which to examine the changes, if any, in the dates of winter migration. In this paper we examine data on the first arrivals and last departures of winter visitors from two UK county bird reports. A small number of changes in phenology and some relationships with climatic variables were present. However, results were often contradictory or inconclusive. As data such as these are highly variable we recommend a more thorough examination for climatic impact where records exist.
131183878
Climate Change Science: Adapt, Mitigate, or Ignore?
Global warming due to increased greenhouse gas emissions poses the most severe problem for governments today. A risk analysis forecasts increased flooding and coastline vulnerability in the United Kingdom and the world. The seriousness of economic damage depends on future global emission scenarios. The U.K. government is planning to reduce carbon dioxide emissions in the United Kingdom by 60% by 2050. Global agreement and sustained action to limit carbon dioxide emissions is needed.
32845937
Is the detection of the first arrival date of migrating birds influenced by population size? A case study of the red-backed shrike Lanius collurio
Abstract Many analyses do not consider the problems associated with the effects of population size on encounter recording. Population size could impact on the detection of bird arrival time as there is a higher probability of observing earlier arrival when the population size is greater and the song activity of birds is increased, as occurs with a larger population. As a case study, we have analysed data on the red-backed shrike Lanius collurio collected in Western Poland during 1983–2000. In this period the red-backed shrike’s return to its breeding sites became significantly earlier whilst the contemporary population size increased significantly. To eliminate linear trends through time we have worked on the standardised residuals from regression of both arrival time and population size on year. The correlation between arrival time and population size residuals was significantly negative, further supporting the link between detection and population size. This finding suggests that, in studies of avian migration and its changes over time, the relationship between arrival date and population size needs to be considered.
83056604
Arrival and Departure Dates
Publisher Summary This chapter reviews the evidence indicating that the arrival and departure dates of migratory birds have changed, and the relationship of these changes to climate variability. The longest continuous time series available for analysis exceed 250 years in length. All these data are observations of first arrivals of bird species that were well known to people of rural communities. In time series approaching or exceeding 100 years, it becomes readily evident that the variation of arrival times around the overall average or trend often shows periods of constant direction of change and then a shift in the direction. Correlating or regressing the arrival dates with a suitable weather (often temperature) variable often reveals a strong relationship between arrival and temperature, but a substantial part of the variation remains unexplained. A proportion of unexplained variation arises from the fact that the local temperature that is used in correlation and regression analyses is, at best, only a proxy of the weather along the migration route prior to arrival at the observation locality. Responses determining departure date probably depend on the individual species. For some species, advancement in spring may result directly in earlier departure in autumn while in others longer summer and later autumn may lead to later departure. Apparently, the direction of response depends on the importance of summer and winter seasons in the species' complete life history strategy.
55931289
Characterizing bird migration phenology using data from standardized monitoring at bird observatories
Long-term data from standardized monitoring programmes at bird observatories are becoming increasingly available. These data are frequently used for detecting changes in the timing of bird migration that may relate to recent climate change. We present an overview of problematic issues in the analysis of these data, and review approaches to and methods for characterizing bird migration phenology and its change over time. Methods are illustrated and briefly compared using autumn data on garden warbler Sylvia borin from a standardized mist-netting programme at Lista bird observatory, southern Norway. Bird migration phenology is usually characterized rather coarsely using a small number of sample statistics such as mean, median and selected quantiles. We present 2 alternative approaches. Smoothing methods describe the within-season pattern in the data at an arbitrary level of detail, while fitting a parametric seasonal distribution curve offers a coarse description of migration phenology relatively robust to sampling effects. Various methods for analyzing linear trends in the timing of bird migration are reviewed and discussed. Exploratory studies using long-term data gathered at bird observatories can yield more detailed insight into the phenomenon of bird migration and how phenologies relate to climate. Methodological advances are needed, particularly in order to better characterize the shape of phenological distributions and separate between sampling effects and 'true' phenology.
25311569
North Atlantic Oscillation and timing of spring migration in birds
Migrant birds have been trapped on the island of Helgoland (southeastern North Sea) since 1909, with methods and sampling effort remaining unchanged throughout the last four decades. In 12 short/medium–distance migrants and 12 long–distance migrants (23 passerines plus the European woodcock) sample sizes were sufficient to calculate mean spring passage (msp) times and to relate these to climate change. All but one species, passing Helgoland en route to their breeding areas (mainly in Scandinavia), show a trend towards earlier msp–time, which is significant in 7 short/medium–distance migrants and 10 long–distance migrants. The msp–times advanced by 0.05–0.28 days per year, short/medium–distance migrants not differing from long–distance migrants. In 23 out of the 24 species, earlier msp–times coincide with local warmer msp–temperatures (significantly in 11 and 7 species of the two groups, respectively). Even more striking is the relation to a large–scale phenomenon, the North Atlantic Oscillation (NAO), during the last four decades. Again, in 23 out of the 24 species, an earlier msp-time coincides with higher NAO indices (significantly in 9 and 12 species, respectively). The NAO index can also explain differences and similarities in spring migration strategies, as well as migration routes within Europe.
86094907
Examining the total arrival distribution of migratory birds
This paper reports on the total distribution of spring migration timing of willow warbler, chiffchaff and pied flycatcher at locations in the UK, Germany, Russia and Finland. This is the first time that high-quality data based on known-effort monitoring has been examined on a continental scale. First arrival dates, commonly reported in the literature, were positively correlated with mean arrival dates although they would not make good predictors of the latter. At all locations, at least one aspect of the arrival distribution of each species had got significantly earlier in recent years. The trend towards earliness was associated with warmer local temperatures and more positive winter North Atlantic Oscillation index. In years that were early, the arrival distribution became more elongated and skewed. Researchers should now investigate the consequences of earlier arrival on current and future bird populations.
43570362
Climate change and unequal phenological changes across four trophic levels: constraints or adaptations?
1. Climate change has been shown to affect the phenology of many organisms, but interestingly these shifts are often unequal across trophic levels, causing a mismatch between the phenology of organisms and their food. 2. We consider two alternative hypotheses: consumers are constrained to adjust sufficiently to the lower trophic level, or prey species react more strongly than their predators to reduce predation. We discuss both hypotheses with our analyses of changes in phenology across four trophic levels: tree budburst, peak biomass of herbivorous caterpillars, breeding phenology of four insectivorous bird species and an avian predator. 3. In our long-term study, we show that between 1988 and 2005, budburst advanced (not significantly) with 0.17 d yr(-1), while between 1985 and 2005 both caterpillars (0.75 d year(-1)) and the hatching date of the passerine species (range for four species: 0.36-0.50 d year(-1)) have advanced, whereas raptor hatching dates showed no trend. 4. The caterpillar peak date was closely correlated with budburst date, as were the passerine hatching dates with the peak caterpillar biomass date. In all these cases, however, the slopes were significantly less than unity, showing that the response of the consumers is weaker than that of their food. This was also true for the avian predator, for which hatching dates were not correlated with the peak availability of fledgling passerines. As a result, the match between food demand and availability deteriorated over time for both the passerines and the avian predators. 5. These results could equally well be explained by consumers' insufficient responses as a consequence of constraints in adapting to climate change, or by them trying to escape predation from a higher trophic level, or both. Selection on phenology could thus be both from matches of phenology with higher and lower levels, and quantifying these can shed new light on why some organisms do adjust their phenology to climate change, while others do not.
128511871
The Migration Ecology of Birds
This book presents an up-to-date, detailed and thorough review of the most fascinating ecological findings of bird migration. It deals with all aspects of this absorbing subject, including the problems of navigation and vagrancy, the timing and physiological control of migration, the factors that limit their populations, and more. Author, Ian Newton, reveals the extraordinary adaptability of birds to the variable and changing conditions across the globe, including current climate change. This adventurous book places emphasis on ecological aspects, which have received only scant attention in previous publications. Overall, the book provides the most thorough and in-depth appraisal of current information available, with abundant tables, maps and diagrams, and many new insights. Written in a clear and readable style, this book appeals not only to migration researchers in the field and Ornithologists, but to anyone with an interest in this fascinating subject.Hot ecological aspects include: various types of bird movements, including dispersal and nomadism, and how they relate to food supplies and other external conditions. This book contains numerous tables, maps and diagrams, a glossary, and a bibliography of more than 2,700 references. It is written by an active researcher with a distinguished career in avian ecology, including migration research.
15077499
Challenging claims in the study of migratory birds and climate change
Recent shifts in phenology in response to climate change are well established but often poorly understood. Many animals integrate climate change across a spatially and temporally dispersed annual life cycle, and effects are modulated by ecological interactions, evolutionary change and endogenous control mechanisms. Here we assess and discuss key statements emerging from the rapidly developing study of changing spring phenology in migratory birds. These well‐studied organisms have been instrumental for understanding climate‐change effects, but research is developing rapidly and there is a need to attack the big issues rather than risking affirmative science. Although we agree poorly on the support for most claims, agreement regarding the knowledge basis enables consensus regarding broad patterns and likely causes. Empirical data needed for disentangling mechanisms are still scarce, and consequences at a population level and on community composition remain unclear. With increasing knowledge, the overall support (‘consensus view’) for a claim increased and between‐researcher variability in support (‘expert opinions') decreased, indicating the importance of assessing and communicating the knowledge basis. A proper integration across biological disciplines seems essential for the field's transition from affirming patterns to understanding mechanisms and making robust predictions regarding future consequences of shifting phenologies.
24691780
Detours in bird migration.
Bird migration routes often follow detours where passages across ecological barriers are reduced in extent. This occurs in spite of the fact that long barrier crossings are within the birds' potential flight range capacity. Long-distance flights are associated with extra energy costs for transport of the heavy fuel loads required. This paper explores how important the fuel transport costs, estimated on the basis of flight mechanics, may be to explain detours for birds migrating by flapping flight. Maximum detours in relation to expanse of the barrier are predicted for cases where birds travel along the detour by numerous short flights and small fuel reserves, divide the detour into a limited number of flight steps, and where a reduced barrier passage is included in the detour. The principles for determining the optimum route, often involving a shortcut across part of the barrier, are derived. Furthermore, the effects of differences in fuel deposition rates and in transport costs for the profitability of detours are briefly considered. An evaluation of a number of observed and potential detours in relation to the general predictions of maximum detours, indicates that reduction of fuel transport costs may well be a factor of widespread importance for the evolution of detours in bird migration at wide ecological barriers.
128348981
Animal Migration: A Synthesis
THEME 1: THE EVOLUTION OF MIGRATION THEME 2: HOW TO MIGRATE THEME 3: MIGRATION IN TIME AND SPACE THEME 4: BROADER CONTEXTS
59053937
Long-distance migration: evolution and determinants
Long distance migration has evolved in many organisms moving through different media and using various modes of locomotion and transport. Migration continues to evolve or become suppressed as shown by ongoing dynamic and rapid changes of migration patterns. This great evolutionary flexibility may seem surprising for such a complex attribute as migration. Even if migration in most cases has evolved basically as a strategy to maximise fitness in a seasonal environment, its occurrence and extent depend on a multitude of factors. We give a brief overview of different factors (e.g. physical, geographical, historical, ecological) likely to facilitate and/or constrain the evolution of long distance migration and discuss how they are likely to affect migration. The basic driving forces for migration are ecological and biogeographic factors like seasonality, spatiotemporal distributions of resources, habitats, predation and competition. The benefit of increased resource availability will be balanced by costs associated with the migratory process in terms of time (incl. losses of prior occupancy advantages), energy and mortality (incl. increased exposure to parasites). Furthermore, migration requires genetic instructions (allowing substantial room for learning in some of the traits) about timing, duration and distance of migration as well as about behavioural and physiological adaptations (fuelling, organ flexibility, locomotion, use of environmental transport etc) and control of orientation and navigation. To what degree these costs and requirements put constraints on migration often depends on body size according to different scaling relationships. From this expos it is clear that research on migration warrants a multitude of techniques and approaches for a complete as possible understanding of a very complex evolutionary syndrome. In addition, we also present examples of migratory distances in a variety of taxons. In recent years new techniques, especially satellite radio telemetry, provide new information of unprecedented accuracy about journeys of individual animals, allowing re-evaluation of migration, locomotion and navigation theories. (Less)
56051257
Travelling through a warming world: climate change and migratory species.
Long-distance migrations are among the wonders of the natural world, but this multi- taxon review shows that the characteristics of species that undertake such movements appear to make them particularly vulnerable to detrimental impacts of climate change. Migrants are key com- ponents of biological systems in high latitude regions, where the speed and magnitude of climate change impacts are greatest. They also rely on highly productive seasonal habitats, including wet- lands and ocean upwellings that, with climate change, may become less food-rich and predictable in space and time. While migrants are adapted to adjust their behaviour with annual changes in the weather, the decoupling of climatic variables between geographically separate breeding and non- breeding grounds is beginning to result in mistimed migration. Furthermore, human land-use and activity patterns will constrain the ability of many species to modify their migratory routes and may increase the stress induced by climate change. Adapting conservation strategies for migrants in the light of climate change will require substantial shifts in site designation policies, flexibility of management strategies and the integration of forward planning for both people and wildlife. While adaptation to changes may be feasible for some terrestrial systems, wildlife in the marine ecosystem may be more dependent on the degree of climate change mitigation that is achievable.
84017846
Population limitation in migrants
Unlike resident bird species, the population sizes of migratory species can be influenced by conditions in more than one part of the world. Changes in the numbers of migrant birds, either long-term or year-to-year, may be caused by changes in conditions in the breeding or wintering areas or both. The strongest driver of numerical change is provided in whichever area the per capita effects of adverse factors on survival or fecundity are greatest. Examples are given of some species whose numbers have changed in association with conditions in breeding areas, and of others whose numbers have changed in association with conditions in wintering areas. In a few such species, the effects of potential limiting factors have been confirmed locally by experiment. In theory, population sizes might also be limited by severe competition at restricted stopover sites, where bird densities are often high and food supplies heavily depleted, but (with one striking exception) the evidence is as yet no more than suggestive. In some species, habitats occupied in wintering and migration areas, and their associated food supplies, can influence the body condition, migration dates and subsequent breeding success of migrants. Body reserves accumulated in spring by large waterfowl serve for migration and for subsequent breeding, and females with the largest reserves are most likely to produce young. Hence, the conditions experienced by individuals in winter in one region can affect their subsequent breeding success in another region. Such effects are apparent at the level of the individual and at the level of the population. Similarly, the numbers of young produced in one region could, through density-dependent processes, affect subsequent overall mortality in another region. Events in breeding, migration and wintering areas are thus interlinked in their effects on bird numbers. Although in the last 30–40 years the numbers of some tropical wintering birds have declined in western Europe and others in eastern North America, the causes seem to differ. In Europe, declines have mainly involved species that winter in the arid savannas of tropical Africa, which have suffered from the effects of drought and increasing desertification. In several species, annual fluctuations in numbers and adult survival rates were correlated with annual fluctuations in rainfall, and by implication in winter food supplies. In North America, by contrast, numerical declines have affected many species that breed and winter in forest, especially those eastern species favouring the forest interior. Declines have been attributed ultimately to human-induced changes in the breeding range, particularly forest fragmentation, which have led to increases in the densities of nest predators and parasitic cowbirds. These in turn are thought to have caused declines in the breeding success of some neotropical migrants, which is now too low to offset the usual adult mortality, but as yet convincing evidence is available for only a minority of species. The breeding rates and population changes of some migratory species have been influenced by natural changes in the availability of defoliating caterpillars. In other species, tropical deforestation is likely to have played the major role in population decline, and if recent rates of tropical deforestation continue, it is likely to affect an increasing range of migratory species in the future. Not all such species are likely to be affected adversely by deforestation, however, and some may benefit from the resulting habitat changes.
24131751
Animal migration: is there a common migratory syndrome?
Ornithologists, and especially northern hemisphere ornithologists, have traditionally thought of migration as an annual return movement of populations between regular breeding and non-breeding grounds. Problems arise because selection does not ordinarily act on populations and because organisms of many taxa (including birds) are clearly migrants, but fail to undertake movements of the kind described. There are also extensive return movements that are not migratory. I propose that it is more useful to think of migration as a syndrome of behavioral and other traits that function together within individuals, and that such a syndrome provides a common ground across taxa from aphids to albatrosses. Large-scale return movements of populations are one outcome of the syndrome. Similar behavioral and physiological traits serve both to define migration and to provide a test for it. I use two insect (Hemipteran) examples to illustrate migratory syndromes and to demonstrate that, in many migrants, behavior and physiology correlate with life history and morphological traits to form syndromes at two levels. I then compare the two Hemipterans with migration in birds, butterflies, and fish to assess the question of whether there are migratory syndromes in common between these diverse migrants. Syndromes are more similar at the level of behavior than when morphology and life history traits are included. Recognizing syndromes leads to important evolutionary questions concerning migration strategies, trade-offs, the maintenance of genetic variance and the responses of migratory syndromes to both similar and different selective regimes.
8261593
Avian migration phenology and global climate change
There is mounting evidence that global climate change has extended growing seasons, changed distribution patterns, and altered the phenology of flowering, breeding, and migration. For migratory birds, the timing of arrival on breeding territories and over-wintering grounds is a key determinant of reproductive success, survivorship, and fitness. But we know little of the factors controlling earlier passage in long-distance migrants. Over the past 30 years in Oxfordshire, U.K., the average arrival and departure dates of 20 migrant bird species have both advanced by 8 days; consequently, the overall residence time in Oxfordshire has remained unchanged. The timing of arrival has advanced in relation to increasing winter temperatures in sub-Saharan Africa, whereas the timing of departure has advanced after elevated summer temperatures in Oxfordshire. This finding demonstrates that migratory phenology is quite likely to be affected by global climate change and links events in tropical winter quarters with those in temperate breeding areas.
4359363
Land–atmosphere coupling and climate change in Europe
Increasing greenhouse gas concentrations are expected to enhance the interannual variability of summer climate in Europe and other mid-latitude regions, potentially causing more frequent heatwaves. Climate models consistently predict an increase in the variability of summer temperatures in these areas, but the underlying mechanisms responsible for this increase remain uncertain. Here we explore these mechanisms using regional simulations of recent and future climatic conditions with and without land–atmosphere interactions. Our results indicate that the increase in summer temperature variability predicted in central and eastern Europe is mainly due to feedbacks between the land surface and the atmosphere. Furthermore, they suggest that land–atmosphere interactions increase climate variability in this region because climatic regimes in Europe shift northwards in response to increasing greenhouse gas concentrations, creating a new transitional climate zone with strong land–atmosphere coupling in central and eastern Europe. These findings emphasize the importance of soil-moisture–temperature feedbacks (in addition to soil-moisture–precipitation feedbacks) in influencing summer climate variability and the potential migration of climate zones with strong land–atmosphere coupling as a consequence of global warming. This highlights the crucial role of land–atmosphere interactions in future climate change.
15586973
Climate change threatens European conservation areas
Europe has the world's most extensive network of conservation areas. Conservation areas are selected without taking into account the effects of climate change. How effectively would such areas conserve biodiversity under climate change? We assess the effectiveness of protected areas and the Natura 2000 network in conserving a large proportion of European plant and terrestrial vertebrate species under climate change. We found that by 2080, 58 ± 2.6% of the species would lose suitable climate in protected areas, whereas losses affected 63 ± 2.1% of the species of European concern occurring in Natura 2000 areas. Protected areas are expected to retain climatic suitability for species better than unprotected areas (P<0.001), but Natura 2000 areas retain climate suitability for species no better and sometimes less effectively than unprotected areas. The risk is high that ongoing efforts to conserve Europe's biodiversity are jeopardized by climate change. New policies are required to avert this risk.
130802377
Rapid Advance of Spring Arrival Dates in Long-Distance Migratory Birds
Several bird species have advanced the timing of their spring migration in response to recent climate change. European short-distance migrants, wintering in temperate areas, have been assumed to be more affected by change in the European climate than long-distance migrants wintering in the tropics. However, we show that long-distance migrants have advanced their spring arrival in Scandinavia more than short-distance migrants. By analyzing a long-term data set from southern Italy, we show that long-distance migrants also pass through the Mediterranean region earlier. We argue that this may reflect a climate-driven evolutionary change in the timing of spring migration.
23695341
Spring arrival response to climate change in birds: a case study from eastern Europe
This paper analyses the dependence of the first spring arrival dates of short/medium- and long-distance migrant bird species on climate warming in eastern Europe. The timing of arrival of the selected species at the observation site correlates with the North Atlantic Oscillation (NAO) index, air temperature, atmospheric pressure, precipitation and wind characteristics. A positive correlation of fluctuations in winter and spring air temperatures with variations in the NAO index has been established in eastern Europe. Positive winter NAO index values are related to earlier spring arrival of birds in the eastern Baltic region and vice versa—arrival is late when the NAO index is negative. The impact of climate warming on the bird’s life cycle depends on local or regional climate characteristics. We tested the hypothesis that differences in climate indices between North Africa and Europe can influence the timing of spring arrival. Our results support the hypothesis that differences in first spring arrival dates between European populations occur after individuals cross the Sahara. We assume that the endogenous programme of migration control in short/medium-distance migrants synchronises with the changing environment on their wintering grounds and along their migration routes, whereas in long-distance migrants it is rather with environmental changes in the second part of their migratory route in Europe. Our results strongly indicate that the mechanism of dynamic balance in the interaction between the endogenous regulatory programme and environmental factors determines the pattern of spring arrival, as well as migration timing.
82311959
The impact of climate change on birds: Impact of climate change on birds
Weather is of major importance for the population dynamics of birds, but the implications of climate change have only recently begun to be addressed. There is already compelling evidence that birds have been affected by recent climate changes. This review suggests that although there is a substantial body of evidence for changes in the phenology of birds, particularly of the timing of migration and of nesting, the consequences of these responses for a species’ population dynamics is still an area requiring in-depth research. The potential for phenological miscuing (responding inappropriately to climate change, including a lack of response) and for phenological disjunction (in which a bird species becomes out of synchrony with its environment) are beginning to be demonstrated, and are also important areas for further research. The study of climatically induced distributional change is currently at a predictive modelling stage, and will need to develop methods for testing these predictions. Overall, there is a range of intrinsic and extrinsic factors that could potentially inhibit adaptation to climate change and these are a high priority for research.
84831198
Quantifying the robustness of first arrival dates as a measure of avian migratory phenology
As the climate changes, many long-term studies have shown that the timing of bird migration is shifting, increasing the need for reliable measures of migratory phenology. Ideally, daily counts of birds at a site are used to calculate the mean arrival date (MAD) but, as this approach is not always possible and is very labour-intensive, simpler metrics such as first arrival date (FAD) have commonly been used. Here, we examine the relationship between FAD and MAD in 28 summer migrant bird species over a 42-year period (1970–2011) at Portland Bird Observatory, UK. Although significant correlations between FAD and MAD were detected, relationships were weak, particularly in long-distance migrants. We suggest that FAD, although a simple and straightforward measure, is not particularly robust as a proxy for overall migratory phenology at a population level.
84920348
Variable shifts in spring and autumn migration phenology in North American songbirds associated with climate change.
Monitoring studies find that the timing of spring bird migration has advanced in recent decades, especially in Europe. Results for autumn migration have been mixed. Using data from Powdermill Nature Reserve, a banding station in western Pennsylvania, USA, we report an analysis of migratory timing in 78 songbird species from 1961 to 2006. Spring migration became significantly earlier over the 46-year period, and autumn migration showed no overall change. There was much variation among species in phenological change, especially in autumn. Change in timing was unrelated to summer range (local vs. northern breeders) or the number of broods per year, but autumn migration became earlier in neotropical migrants and later in short-distance migrants. The migratory period for many species lengthened because late phases of migration remained unchanged or grew later as early phases became earlier. There was a negative correlation between spring and autumn in long-term change, and this caused dramatic adjustments in the amount of time between migrations: the intermigratory periods of 10 species increased or decreased by > 15 days. Year-to-year changes in timing were correlated with local temperature (detrended) and, in autumn, with a regional climate index (detrended North Atlantic Oscillation). These results illustrate a complex and dynamic annual cycle in songbirds, with responses to climate change differing among species and migration seasons.
84018737
Do changes in climate patterns in wintering areas affect the timing of the spring arrival of trans‐Saharan migrant birds?
The life cycles of plants and animals are changing around the world in line with the predictions originated from hypotheses concerning the impact of global warming and climate change on biological systems. Commonly, the search for ecological mechanisms behind the observed changes in bird phenology has focused on the analysis of climatic patterns from the species breeding grounds. However, the ecology of bird migration suggests that the spring arrival of long-distance migrants (such as trans-Saharan birds) is more likely to be influenced by climate conditions in wintering areas given their direct impact on the onset of migration and its progression. We tested this hypothesis by analysing the first arrival dates (FADs) of six trans-Saharan migrants (cuckoo Cuculus canorus ,s wiftApus apus, hoopoe Upupa epops ,s wallowHirundo rustica, house martin Delichon urbica and nightingale Luscinia megarhynchos), in a western Mediterranean area since from 1952 to 2003. By means of multiple regression analyses, FADs were analysed in relation to the monthly temperature and precipitation patterns of five African climatic regions south of the Sahara where species are thought to overwinter and from the European site from where FADs were collected. We obtained significant models for five species explaining 9‐41% of the variation in FADs. The interpretation of the models suggests that: (1) The climate in wintering quarters, especially the precipitation, has a stronger influence on FADs than that in the species’ potential European breeding grounds. (2) The accumulative effects of climate patterns prior to migration onset may be of considerable importance since those climate variables that served to summarize climate patterns 12 months prior to the onset of migration were selected by final models. (3) Temperature and precipitation in African regions are likely to affect departure decision in the species studied through their indirect effects on food availability and the build-up of reserves for migration. Our results concerning the factors that affect the arrival times of trans-Saharan migrants indicate that the effects of climate change are more complex than previously suggested, and that these effects might have an interacting impact on species ecology, for example by reversing ecological pressures during species’ life cycles.
83690370
Climatic connectivity between Africa and Europe may serve as a basis for phenotypic adjustment of migration schedules of trans-Saharan migratory birds
Most European migratory birds wintering in sub-Saharan Africa have anticipated arrival to the breeding areas over the past decades. This phenological change may be ultimately caused by warming of the Northern Hemisphere via evolutionary changes or phenotypic plasticity in migration behavior. First arrival dates are negatively predicted by temperatures upon arrival to the breeding grounds. This seems puzzling, because migrants should be unable to predict weather conditions at long range. Migrants can enjoy diverse fitness benefits from early arriving. However, if weather conditions at destination cannot be predicted, early arrival can also entail severe costs. If meteorological conditions in Europe during breeding covary with those in sub-Saharan Africa during late winter, long-distance migrants may have a clue to predict meteorological conditions in their breeding areas while they are still in Africa and adjust their migration schedule consequently, an idea that has never been tested. We analyzed the correlation between March–April temperature anomalies (Tan) in Europe and February Tan in the Sahel and sub-Sahel, where long-distance migrants winter or stop-over. Tan in Africa negatively predicted Tan in Europe, the association being particularly strong (unsigned effect size, zr>0.35) for eastern Sahel and northern and eastern Europe, where the risks of early arrival may be larger. However, the strength of the correlations between Tan in the two continents has declined during the last 25 years; thus, possibly, partly compromising adaptive mechanisms of adjustment of migration. The existence of such climatic connectivity leads to several predictions, including that positive Tan in Africa should delay arrival. Consistent with this prediction, we found that first arrival dates of seven long-distance migratory species positively covaried with February Tan in Africa. Thus, while wintering, migrants might be able to predict meteorological conditions at the beginning of the breeding season, and phenotypically adjust migration schedules to optimally tune arrival date.
36722043
The North Atlantic Oscillation: Past, present, and future
The climate of the Atlantic sector exhibits considerable variability on a wide range of time scales. A substantial portion is associated with the North Atlantic Oscillation (NAO), a hemispheric meridional oscillation in atmospheric mass with centers of action near Iceland and over the subtropical Atlantic. NAO-related impacts on winter climate extend from Florida to Greenland and from northwestern Africa over Europe far into northern Asia. Over the last 3 decades, the phase of the NAO has been shifting from mostly negative to mostly positive index values. Much remains to be learned about the mechanisms that produce such low frequency changes in the North Atlantic climate, but it seems increasingly likely that human activities are playing a significant role.
6415789
Assortative Mating as a Mechanism for Rapid Evolution of a Migratory Divide
There have been numerous recent observations of changes in the behavior and dynamics of migratory bird populations, but the plasticity of the migratory trait and our inability to track small animals over large distances have hindered investigation of the mechanisms behind migratory change. We used habitat-specific stable isotope signatures to show that recently evolved allopatric wintering populations of European blackcaps Sylvia atricapilla pair assortatively on their sympatric breeding grounds. Birds wintering further north also produce larger clutches and fledge more young. These findings describe an important process in the evolution of migratory divides, new migration routes, and wintering quarters. Temporal segregation of breeding is a way in which subpopulations of vertebrates may become isolated in sympatry.
83755661
THE TIMING OF BIRDS‘ BREEDING SEASONS
Summary 
Examination of survival rdtes of nestlings and fledglings of some species show that there is a strong tendency for those young which are hatched earliest in the season to have the greatest chance of surviving to breed. Since natural selection so strongly favours parents who leave many surviving young, the question arises as to why other birds breed later than the date at which they could most successfully raise their young. 
 
It is suggested that the food supply for the breeding females immediately prior to the breeding season may limit their ability to form eggs and the females may thus not be able to lay at the time which would result in young being in the nest at the best time for raising them, but as soon after this time as the female is able to produce her eggs. Not all species are likely to be prevented, by food shortage, from breeding at the best time for raising young and the groups of birds most likely to be affected are discussed.
1837288
Heritability of arrival date in a migratory bird.
The behaviour of long-distance migratory birds is assumed to partly be under the influence of genes, as demonstrated by selection experiments. Furthermore, competition for early arrival among males may lead to condition-dependent migration associated with fitness benefits of early arrival achieved by individuals in prime condition. Here I present field data on the repeatability and the heritability of arrival date in a trans-equatorial migratory bird, the barn swallow Hirundo rustica, and I test for a genetic correlation between arrival date and the expression of a condition-dependent secondary sexual character. The repeatability was statistically significant and the heritability of arrival date was estimated to be 0.54 (s.e. = 0.15). There was no significant evidence of this estimate being inflated by environmental or maternal condition during rearing. Arrival date and migration are condition dependent in the barn swallow, with males with the most exaggerated secondary sexual characters also arriving the earliest. There was a significant genetic correlation between arrival date and tail length in male barn swallows, providing indirect evidence for a genetic basis of this condition dependence. Given the high level of heritability, arrival date could readily respond to selection caused by environmental change.
46383630
Patterns of spring arrival dates differ in two hirundines
We examined the first arrival dates in the UK over a 56 yr period of 2 hirundines, sand martin Riparia riparia and barn swallow Hirundo rustica, in relation to temperatures along migration routes and at destination. Changes in arrival dates have been much greater for sand martin than for barn swallow, and the arrival order of the 2 species now appears to have reversed. There appear to be 2 reasons for this: a greater response to temperatures in SW Europe and a different response to temperature in recent years, leading to earlier arrival at the same temperature than formerly. We argue that this may involve evolutionary change, but conclude that it is difficult to be definite about this without information on individual animals.
84676984
Timing of Autumn Migration and Its Relation to Winter Distribution in Dark‐Eyed Juncos
Migratory Dark-eyed Juncos in eastern North America migrate southward in autumn and tend to segregate in the winter range according to sex and age. North to south, the most abundant classes are young males, adult males, young females, and adult females. Because adults tend to dominate young at winter feeding sites, this distribution appears at first to conflict with the view that dominance interactions are responsible for differential avian migrations and that dominant individuals remain nearest the breeding range. However, if young juncos establish winter residency at earlier dates than adults, a prior residence effect might make them dominant. This hypothesis requires that young arrive at wintering sites earlier than adults and that residents of all classes arrive earlier than their transient counterparts. We tested these predictions by comparing first-capture dates ofjuncos at Bloomington, Indiana, which is north of the latitudinal midline of the junco's winter range. During 13 autumns, individuals were classed as local residents or as transients and were sexed and aged. Contrary to predictions, residents were not caught earlier than transients. Further, adults were caught earlier, not later, than young among transients and probably also among residents. Thus, settlement of the winter range does not proceed from north to south, and dominance established through prior residence cannot account for the concentration of 1st-yr males in the northern part of the winter range. The fact that some sex-age classes tend to winter south of others predicts that at a northern capture site the classes that migrate farthest should be commoner among transients than among residents. This expectation was fulfilled. In addition, the median capture dates of the sex-age classes were arranged approximately according to the north-to-south order of their distribution, indicating that classes with the farthest to travel passed through (transients) or settled (residents) earliest in autumn. We conclude that comparison of autumn migration schedules of transient and resident passerine birds at a single location can yield considerable information about the dynamics of settlement of the entire winter range, including possible information about differences in destination of subsets of tran- sients.
26431428
Climate and the complexity of migratory phenology: sexes, migratory distance, and arrival distributions
The intra- and inter-season complexity of bird migration has received limited attention in climatic change research. Our phenological analysis of 22 species collected in Chicago, USA, (1979–2002) evaluates the relationship between multi-scalar climate variables and differences (1) in arrival timing between sexes, (2) in arrival distributions among species, and (3) between spring and fall migration. The early migratory period for earliest arriving species (i.e., short-distance migrants) and earliest arriving individuals of a species (i.e., males) most frequently correlate with climate variables. Compared to long-distance migrant species, four times as many short-distance migrants correlate with spring temperature, while 8 of 11 (73%) of long-distance migrant species’ arrival is correlated with the North Atlantic Oscillation (NAO). While migratory phenology has been correlated with NAO in Europe, we believe that this is the first documentation of a significant association in North America. Geographically proximate conditions apparently influence migratory timing for short-distance migrants while continental-scale climate (e.g., NAO) seemingly influences the phenology of Neotropical migrants. The preponderance of climate correlations is with the early migratory period, not the median of arrival, suggesting that early spring conditions constrain the onset or rate of migration for some species. The seasonal arrival distribution provides considerable information about migratory passage beyond what is apparent from statistical analyses of phenology. A relationship between climate and fall phenology is not detected at this location. Analysis of the within-season complexity of migration, including multiple metrics of arrival, is essential to detect species’ responses to changing climate as well as evaluate the underlying biological mechanisms.