Spatiotemporal variation in avian migration phenology: citizen science reveals effects of climate change

A growing number of studies have documented shifts in avian migratory phenology in response to climate change, and yet there is a large amount of unexplained variation in the magnitude of those responses across species and geographic regions. We use a database of citizen science bird observations to explore spatiotemporal variation in mean arrival dates across an unprecedented geographic extent for 18 common species in North America over the past decade, relating arrival dates to mean minimum spring temperature. Across all species and geographic locations, species shifted arrival dates 0.8 days earlier for every °C of warming of spring temperature, but it was common for some species in some locations to shift as much as 3-6 days earlier per °C. Species that advanced arrival dates the earliest in response to warming were those that migrate more slowly, short distance migrants, and species with broader climatic niches. These three variables explained 63% of the interspecific variation in phenological response. We also identify a latitudinal gradient in the average strength of phenological response, with species shifting arrival earlier at southern latitudes than northern latitudes for the same degree of warming. This observation is consistent with the idea that species must be more phenologically sensitive in less seasonal environments to maintain the same degree of precision in phenological timing.     

How consistent are trends in arrival (and departure) dates of migrant birds in the UK?

We examine the first arrival and last departure dates of migrant bird species from, respectively, six and three English area bird reports. Of all 145 bird series, 50% demonstrated significantly earlier arrival in recent years, with the average advance over all species being 0.25 days/year or 12 days earlier over 50 years. Thirty percent of 67 series demonstrated significantly later departure, with the average species delay being 0.16 days/year or eight days later over 50 years. There was greater consistency between species in trends in first arrival than in last departure, with species such as sand martin Riparia riparia significantly earlier at all six sites while, for example, spotted flycatcher Muscicapa striata showed no significant change in arrival at all sites. Significant negative correlations between arrival dates and English temperatures were found for 26% of all series, but temperature effects on departures were less clear. We provide some evidence that trends in arrival dates may be masked by population declines in birds. Since migrant bird populations are in decline generally, this may suggest that the real advance in arrival dates may be greater than that reported here.     

Why is timing of bird migration advancing when individuals are not?

Recent advances in spring arrival dates have been reported in many migratory species but the mechanism driving these advances is unknown. As population declines are most widely reported in species that are not advancing migration, there is an urgent need to identify the mechanisms facilitating and constraining these advances. Individual plasticity in timing of migration in response to changing climatic conditions is commonly proposed to drive these advances but plasticity in individual migratory timings is rarely observed. For a shorebird population that has significantly advanced migration in recent decades, we show that individual arrival dates are highly consistent between years, but that the arrival dates of new recruits to the population are significantly earlier now than in previous years. Several mechanisms could drive advances in recruit arrival, none of which require individual plasticity or rapid evolution of migration timings. In particular, advances in nest-laying dates could result in advanced recruit arrival, if benefits of early hatching facilitate early subsequent spring migration. This mechanism could also explain why arrival dates of short-distance migrants, which generally return to breeding sites earlier and have greater scope for advance laying, are advancing more rapidly than long-distance migrants.     

Factors influencing plasticity in the arrival‐breeding interval in a migratory species reacting to climate change

Climate change is profoundly affecting the phenology of many species. In migratory birds, there is evidence for advances in their arrival time at the breeding ground and their timing of breeding, yet empirical studies examining the interdependence between arrival and breeding time are lacking. Hence, evidence is scarce regarding how breeding time may be adjusted via the arrival‐breeding interval to help local populations adapt to local conditions or climate change. We used long‐term data from an intensively monitored population of the northern wheatear (Oenanthe oenanthe) to examine the factors related to the length of 734 separate arrival‐to‐breeding events from 549 individual females. From 1993 to 2017, the mean arrival and egg‐laying dates advanced by approximately the same amount (~5–6 days), with considerable between‐individual variation in the arrival‐breeding interval. The arrival‐breeding interval was shorter for: (a) individuals that arrived later in the season compared to early‐arriving birds, (b) for experienced females compared to first‐year breeders, (c) as spring progressed, and (d) in later years compared to earlier ones. The influence of these factors was much larger for birds arriving earlier in the season compared to later arriving birds, with most effects on variation in the arrival‐breeding interval being absent in late‐arriving birds. Thus, in this population it appears that the timing of breeding is not constrained by arrival for early‐ to midarriving birds, but instead is dependent on local conditions after arrival. For late‐arriving birds, however, the timing of breeding appears to be influenced by arrival constraints. Hence, impacts of climate change on arrival dates and local conditions are expected to vary for different parts of the population, with potential negative impacts associated with these factors likely to differ for early‐ versus late‐arriving birds.     

The effects of temperature, altitude and latitude on the arrival and departure dates of the swallow Hirundo rustica in the Slovak Republic

The (barn) swallow Hirundo rustica is a traditional harbinger of spring in many countries of the Northern Hemisphere. This paper uses information on the arrival and departure dates of the swallow in the Slovak Republic for the 30 years 1961–1985 and 1996–2000. Records were taken at 19 locations throughout the Republic representing an altitude range from 105 m to 760 m. Monthly temperature data were constructed from six meteorological stations. With the use of regression techniques, trends towards later arrival, earlier departure and the effects of latitude, altitude and temperature are all apparent. Received: 25 October 2000 / Revised: 23 April 2001 / Accepted: 23 April 2001     

The effects of climate change on the phenology of winter birds in Yokohama, Japan

Observations made largely from summer breeding sites in Europe and North America have been used to document the effects of climate change on many bird species. We extend these studies by examining 23 years of observations between 1986 and 2008 of six winter bird species made by citizens at a city park in Yokohama, Japan. Bird species arrive in autumn and spend the winter in the area, before departing in the late winter or spring. On average, birds species are arriving 9 days later than in the past and are departing on average 21 days earlier, meaning that the average duration of their stay in Yokohama is about 1 month shorter now than in the past. Patterns of changes over time varied among species, but departure dates changed for more species than did arrival dates. Dates of departure and arrival were sometimes correlated with monthly average temperatures—later arrivals and earlier departures were associated with warmer temperatures. In addition, interannual variation in arrival and departure dates were strongly correlated across species, suggesting that species were responding to the same or similar environmental cues. This study provides a clear demonstration of the value of using citizens to make observations that contribute to research in climate change biology.     

Predation can select for later and more synchronous arrival times in migrating species

For migratory species, the timing of arrival at breeding grounds is an important determinant of fitness. Too early arrival at the breeding ground is associated with various costs, and we focus on one understudied cost: that migrants can experience a higher risk of predation if arriving earlier than the bulk of the breeding population. We show, using both a semi‐analytic and simulation model, that predation can select for later arrival. This is because of safety in numbers: predation risk becomes diluted if many other individuals, either con‐ or heterospecific, are already residing in the area. Predation risk dilution can also select for more synchronous arrival because deviating from the current population‐wide norm to earlier or later dates leads to higher predation risk or to failures in territory acquisition, respectively. The fact that selection for high arrival synchrony can in some cases be more important than selection for a specific date (early or late) within the season is an example of an ‘evolutionary priority effect’: whichever strategy – in this case a particular arrival time – becomes established in a population can remain stable over long periods of time; there are many possible equilibria (multiple stable states) which the population can remain at. Mixed arrival strategies are also possible under some circumstances.     

Effect of global warming on the timing of migration and breeding of passerine birds in the 20th century

Long-term monitoring of the dates of arrival, breeding, and autumn migration in 25 passerine bird species on the Kurshskaya (Courland) Spit, the Baltic Sea, has shown that spring migration and nesting in most species wintering in Europe or Africa have shifted to earlier dates in the past two decades, whereas the dates of autumn migration in most species studied have not changed significantly. In 16 bird species, a significant negative correlation of the timing of arrival and breeding with the average spring air temperature and the North Atlantic Oscillation index (NAO) in February and March was revealed. In years with early and warm springs, birds arrived at the spit and nested considerably earlier than in years with cold springs. The dates of autumn migration in most species studied largely depended on the timing of nesting but not on weather conditions in autumn. The data obtained indicate that the main factor responsible for long-term changes in the timing of arrival, nesting, and autumn migrations of passerine birds in the Baltic Region is climate fluctuations that led to considerable changes in thermal conditions in the Northern Hemisphere in the 20th century. The hypothesis is proposed that recent climate warming has caused changes in the timing of not only the arrival of birds in Europe but also of their spring migrations from Africa. Further changes in the dates of passerine bird arrival and breeding in the Palearctic in subsequent years will largely depend on the dynamics of winter and spring air temperatures in the Northern Hemisphere, whereas the timing of autumn migrations will be determined mainly by the dates of their arrival and nesting.     

Spring migration phenology of birds in the Northern Prairie region is correlated with local climate change

ABSTRACT In apparent response to recent periods of global warming, some migratory birds now arrive earlier at stopover sites and breeding grounds. However, the effects of this warming on arrival times vary among locations and species. Migration timing is generally correlated with temperature, with earlier arrival during warm years than during cold years, so local variation in climate change might produce different effects on migration phenology in different geographic regions. We examined trends in first spring arrival dates (FADs) for 44 species of common migrant birds in South Dakota (1971–2006) and Minnesota (1964–2005) using observations compiled by South Dakota and Minnesota Ornithologists’ Unions. We found significant trends in FAD over time for 20 species (18 arriving earlier and two later) in South Dakota and 16 species (all earlier) in Minnesota. Of these species, 10 showed similar significant trends for both states. All 10 of these species exhibited significantly earlier arrival, and all were early spring migrants, with median FADs before 10 April in both states. Eighteen of the 44 species showed significant negative correlations of FADs with either winter (December–February) or spring (arrival month plus previous month) temperatures in one or both states. Interestingly, spring temperatures in both South Dakota and Minnesota did not warm significantly from 1971–2006, but winter temperatures in both states warmed significantly over the same time period. This suggests that the warmer winters disproportionately affected early spring migrants, especially those associated with aquatic habitats (seven of the 10 species showing significantly earlier spring arrival in both states). The stronger response to climate change by early spring migrants in our study is consistent with the results of several other studies, and suggests that migrants, especially early migrants, are capable of responding to local temperature conditions experienced on wintering grounds or along the migration route.     

Climate change and bird phenology: a long-term study in the Iberian Peninsula

Many studies in recent years have demonstrated long‐term temporal trends in biological parameters that can only be explained by climate change. Bird phenology has received great attention, as it studies one of the most conspicuous, popular, and easily observable phenomena in nature. There are many studies of long‐term changes in spring arrival dates, most of which concur with earlier records from the last few decades. However, few data are available for autumn departures or length of stays. Furthermore, existing data offer an equivocal picture. In this study, we analysed a huge database of about 44 000 records for five trans‐Saharan bird species (Ciconia ciconia, Cuculus canorus, Apus apus, Hirundo rustica and Luscinia megarhynchos). Data were collected from over 1300 sites around Spain during the period 1944–2004. Common spring arrival patterns were found in all species. Spring arrival dates have tended to advance since the mid‐1970s. Current dates are similar to those from the 1940s (except for C. ciconia). Thus, the advance of spring migration over the last three decades could be seen as a return to the initial timing of arrival dates, after abnormally delayed arrivals during the 1970s. A strong negative relationship with temperature in Spain at arrival time was observed in all species. A negative relationship with the Sahel Index (a measurement of precipitation in the African Sahel area during the rainy season) for the previous year was also found in C. canorus, A. apus and H. rustica. Regarding autumn departures, all species showed common interdecadal fluctuations, but only H. rustica is leaving earlier Spain at present. All species departed earlier in years that had higher temperatures during their reproductive period. However, only for H. rustica the relation between Spanish temperatures at departure time and the last sightings of individuals was significant. A heterogeneous temporal response for the length of stay was also found: C. ciconia increased, A. apus did not change and H. rustica decreased its stay. This is the first study, based on an extensive bird phenology observational network covering a large region, that shows the most complete and thorough analysis available for the Mediterranean region.     

Relationship between arrival date, hatching date and breeding success of the white stork (Ciconia ciconia) in Slovakia

Changes in the spring arrival dates of migrant birds have been reported from a range of locations and many authors have focused on long-term trends and their relationship to temperature and other climatic events. Perhaps more importantly, changed arrival dates may have consequences for the breeding dates of birds which strongly influence breeding success. In this paper we take the opportunity provided by a monitoring scheme of the white stork (Ciconia ciconia) to examine several features of the timing of arrival and breeding in relation to chick production in Slovakia during the period 1978–2002. First arrival dates ranged from 5^th March to 30^th April, and hatching dates varied between 26^th April and 8^th July. Generally, early arriving pairs started breeding earlier and a shorter interval between the arrival of the first partner and breeding, expressed here as hatching date, resulted in higher breeding success.     

Temperature‐related increases in grass growth and greater competition for food drive earlier migrational departure of wintering Whooper Swans

Understanding the departure decisions of migratory birds is critical for determining how changing climatic conditions will influence subsequent arrival times on the breeding grounds. A long‐term dataset (1972–2008) of Whooper Swan Cygnus cygnus departure dates from a wintering site in Ireland was used to assess the factors determining the timing of migration. Early and late migrating swans showed different departure patterns. Earlier wintering ground departure was more pronounced for the first 50% of the population than the last 10% of departing individuals. Earlier departure was associated with an increase in February temperatures at the wintering site for all departure phases except the date when the last individual departed. The date by which the first 50% of Swans had departed was earlier with increasing numbers of wintering Swans, suggesting that competition on the wintering grounds may further influence the timing of departure. The results also suggested that departure is mediated by the influence of spring temperature on food resources, with increased February grass growth in warmer years enabling earlier departure of migrating Swans. To determine why arrival dates in the breeding ground have altered, environmental conditions in the wintering grounds must be taken into account.     

Long-term trends in the arrival dates of spring migrants

Data on the spring arrival dates of 23 species of migrants in Leicestershire over a 50-year period are presented. Chiffchaff, Sand Martin, Blackcap and Sedge Warbler showed a significant trend towards earlier arrival over the period, while Tree Pipit, Cuckoo, Whinchat, Whitethroat and Garden Warbler showed a significant trend towards later arrival. Fifteen species arrived noticeably earlier in the 1940s, a period of warm springs, while several species showed earlier arrivals in the 1980s. A number of species showed later arrival dates in the 1960s and 1970s, when April temperatures were colder than average. Several species showed significant correlations between arrival date and temperature. Arrival dates of the earliest species were much more variable than those arriving later, while species arriving in the second half of April showed a generally synchronous arrival. The results are discussed in the context of global warming.     

Flexibility in phenology and habitat use act as buffers to long‐term population declines in UK passerines

Ecological responses to environmental change are wide‐ranging, from alterations in the timing of life‐history events to range and population changes. Explaining the variation across species in these responses is essential for identifying vulnerable species and for developing adequate conservation or mitigation strategies. Using population trend data from the UK Breeding Bird Survey, this study examined the association between long‐term population trends (1994–2007) and phenological, life‐history and resource‐use traits of UK passerine species. Phenology, as well as productivity and resource use were significantly associated with long‐term population trends. Average laying date and first clutch laying period were key predictors, with higher population growth rates associated with earlier laying dates and longer laying periods. This suggests that flexibility in the duration of reproductive periods buffers species against environmental changes. Average laying period was particularly important for migrant species. Flexibility in laying dates for these species is constrained by their arrival dates; mean change in arrival date over a twenty‐five year period strongly predicted population trends amongst migrant species. Besides the key role phenological flexibility plays in buffering population declines, we also showed that more productive, generalist species were less likely to have declining populations than species with specialized habitat requirements, particularly those associated with farmland and urban areas and those reliant on highly seasonal food items (i.e. invertebrate eaters). These results underscore the need for a multi‐faceted approach to understanding the mechanisms governing population trends. Additionally, species’ sensitivity to environmental change is likely to depend on interactions between species‐specific phenology, habitat and resource‐use traits.     

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long‐term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg‐laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year‐to‐year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival‐breeding interval (earlier breeding). Finally, mark–recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.     

Relaxed selection when you least expect it: why declining bird populations might fail to respond to phenological mismatches

Ongoing climate change threatens to cause mismatches between the phenology of many organisms and their resources. Populations of migratory birds may need to undergo ‘evolutionary rescue’ if resource availability moves to earlier dates in the year, as shifted arrival dates at the breeding grounds may be required for persistence under new environmental schedules. Here we show a counterintuitive process that can reduce the strength of selection for early arrival when the resource peaks earlier. This happens when two processes combine to determine selection for early arrival: breeding success is higher if a bird does not miss the resource peak, but this occurs together with a ‘zero‐sum game’ where birds acquire good territories ahead of their competitors if they arrive early. The latter process can relax if the population has experienced a recent decline. Therefore, climate change can have two opposing effects: its direct effect on breeding success strengthens selection for early arrival, but this combines with an indirect effect of relaxed selection due to population declines, if territoriality is a significant determinant of population dynamics and fitness. We show that the latter process can predominate, and this can cause a failure for a population to adapt to a new schedule under changing environmental conditions.     

Birth dates vary with fixed and dynamic maternal features,offspring sex,and extreme climatic events in a high‐latitude marine mammal

Reproductive synchrony tends to be widespread in diverse species of plants and animals, especially at higher latitudes. However, for long‐lived mammals, birth dates for different individuals can vary by weeks within a population. A mother's birth timing can reveal useful information about her reproductive abilities and have important implications for the characteristics and survival of her offspring. Despite this, our current knowledge of factors associated with variation in birth dates is modest. We used long‐term data for known‐age Weddell seals in Antarctica and a Bayesian hierarchical modeling approach to study how birth dates varied with fixed and temporally varying features of mothers, whether sex allocation varied with birth timing, and annual variation in birth dates. Based on birth dates for 4465 pups born to 1117 mothers aged 4–31, we found that diverse features of mothers were associated with variation in birth dates. Maternal identity was the most important among these. Unlike most studies, which have reported that birth dates occur earlier as mothers age, we found that birth dates progressively occurred earlier in the year in the early part of a mother's reproductive life, reached a minimum at age 16, and then occurred later at later ages. Birth dates were positively related to a mother's age at primiparity and recent reproductive effort. The earliest birth dates were for pups born to prime‐age mothers who did not reproduce in the previous year but began reproduction early in life, suggesting that females in the best condition gave birth earlier than others. If so, our finding that male pups tended to be born earlier than females provides support for the Trivers–Willard sex‐allocation model. Average birth dates were quite consistent across years, except for 2 years that had notable delays and occurred during the period when massive icebergs were present and disrupted the ecosystem.     

Long‐term phenological shifts and intra‐specific differences in migratory change in the willow warbler Phylloscopus trochilus

Climate change can influence many aspects of avian phenology and especially migratory shifts and changes in breeding onset receive much research interest in this context. However, changes in these different life‐cycle events in birds are often investigated separately and by means of ringing records of mixed populations. In this long‐term study on the willow warbler Phylloscopus trochilus, we investigated timing of spring and autumn migration in conjunction with timing of breeding. We made distinction among individuals with regard to age, sex, juvenile origin and migratory phase. The data set comprised 22‐yr of ringing records and two temporally separated data sets of egg‐laying dates and arrival of the breeding population close to the ringing site. The results reveal an overall advancement consistent in most, but not all, phenological events. During spring migration, early and median passage of males and females became earlier by between 4.4 to 6.3 d and median egg‐laying dates became earlier by 5 d. Male arrival advanced more, which may lead to an increase in the degree of protandry in the future. Among breeding individuals, only female arrival advanced in timing. In autumn, adults and locally hatched juvenile females did not advanced median passage, but locally hatched juvenile males appeared 4.2 d earlier. Migrating juvenile males and females advanced passage both in early and median migratory phase by between 8.4 to 10.1 d. The dissimilarities in the response between birds of different age, sex and migratory phase emphasize that environmental change may elicit intra‐specific selection pressures. The overall consistency of the phenological change in spring, autumn and egg‐laying, coupled with the unchanged number of days between median spring and autumn migration in adults, indicate that the breeding area residence has advanced seasonally but remained temporally constant.     

Late Holocene extinction of Puerto Rican native land mammals

West Indian land mammals have suffered the most severe extinctions of any Holocene mammal faunas. However, 'last-occurrence' dates based on radiometric or robust stratigraphic data remain unavailable for most West Indian species, making it impossible to identify factors responsible for these extinctions. Here, we present new radiometric dates from archaeological and palaeontological sites on Puerto Rico, the only Greater Antillean island to have lost all native land mammals. Although it has been suggested that these species died out earlier than other West Indian mammals, we demonstrate that Puerto Rican mammal last-occurrence dates are in close agreement with those from other Antillean islands, as several species in fact persisted for millennia following Amerindian arrival. Echimyid rodents and nesophontid 'island-shrews' were still present on Puerto Rico approximately 1000 years BP, and probably became extinct following European arrival. The large (13kg) heptaxodontid rodent Elasmodontomys obliquus also appears to have survived for over 2000 years after Amerindian colonization, suggesting that at least some large West Indian mammals became extinct in protracted pre-European 'sitzkrieg'-style events rather than 'blitzkrieg'-style overkill.     

Long-term trends in first arrival and first egg laying dates of some migrant and resident bird species in northern Italy

Climate change is affecting the phenology of seasonal events in Europe and the Northern Hemisphere, as shown by several studies of birds’ timing of migration and reproduction. Here, we analyse the long-term (1982–2006) trends of first arrival dates of four long-distance migratory birds [swift (Apus apus), nightingale (Luscinia megarhynchos), barn swallow (Hirundo rustica), and house martin (Delichon urbicum)] and first egg laying dates of two migrant (swift, barn swallow) and two resident species [starling (Sturnus vulgaris), Italian sparrow (Passer italiae)] at a study site in northern Italy. We also addressed the effects of local weather (temperature and precipitation) and a climate index (the North Atlantic Oscillation, NAO) on the interannual variability of phenological events. We found that the swift and the barn swallow significantly advanced both arrival and laying dates, whereas all other species did not show any significant temporal trend in either arrival or laying date. The earlier arrival of swifts was explained by increasing local temperatures in April, whereas this was not the case for arrival dates of swallows and first egg laying dates of both species. In addition, arrival dates of house martins were earlier following high NAO winters, while nightingale arrival was earlier when local spring rainfall was greater. Finally, Italian sparrow onset of reproduction was anticipated by greater spring rainfall, but delayed by high spring NAO anomalies, and swift’s onset of reproduction was anticipated by abundant rainfall prior to reproduction. There were no significant temporal trends in the interval between onset of laying and arrival in either the swift or the barn swallow. Our findings therefore indicate that birds may show idiosyncratic responses to climate variability at different spatial scales, though some species may be adjusting their calendar to rapidly changing climatic conditions.