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.     

The influence of climate on the timing and rate of spring bird migration

Ecological processes are changing in response to climatic warming. Birds, in particular, have been documented to arrive and breed earlier in spring and this has been attributed to elevated spring temperatures. It is not clear, however, how long-distance migratory birds that overwinter thousands of kilometers to the south in the tropics cue into changes in temperature or plant phenology on northern breeding areas. We explored the relationships between the timing and rate of spring migration of long-distance migratory birds, and variables such as temperature, the North Atlantic Oscillation (NAO) and plant phenology, using mist net capture data from three ringing stations in North America over a 40-year period. Mean April/May temperatures in eastern North America varied over a 5°C range, but with no significant trend during this period. Similarly, we found few significant trends toward earlier median capture dates of birds. Median capture dates were not related to the NAO, but were inversely correlated to spring temperatures for almost all species. For every 1°C increase in spring temperature, median capture dates of migratory birds averaged, across species, one day earlier. Lilac (Syringa vulgaris) budburst, however, averaged 3 days earlier for every 1°C increase in spring temperature, suggesting that the impact of temperature on plant phenology is three times greater than on bird phenology. To address whether migratory birds adjust their rate of northward migration to changes in temperature, we compared median capture dates for 15 species between a ringing station on the Gulf Coast of Louisiana in the southern USA with two stations approximately 2,500 km to the north. The interval between median capture dates in Louisiana and at the other two ringing stations was inversely correlated with temperature, with an average interval of 22 days, that decreased by 0.8 days per 1°C increase in temperature. Our results suggest that, although the onset of migration may be determined endogenously, the timing of migration is flexible and can be adjusted in response to variation in weather and/or phenology along migration routes.     

Extreme spring conditions in the Arctic delay spring phenology of long-distance migratory songbirds

Arctic regions are warming rapidly, with extreme weather events increasing in frequency, duration, and intensity just as in other regions. Many studies have focused on how shifting seasonality in environmental conditions affects vegetation phenology, while far fewer have examined how the breeding phenology of arctic fauna responds. We studied two species of long-distance migratory songbirds, Lapland longspurs, Calcarius lapponicus, and white-crowned sparrows, Zonotrichia leucophrys gambelii, across five consecutive breeding seasons in northern Alaskan tundra. We aimed to understand how spring environmental conditions affected breeding cycle phenology, including the timing of arrival on breeding grounds, territory establishment, and clutch initiation. Spring temperatures, precipitation, and snow-free dates differed significantly among years, with 2013 characterized by unusually late snow cover. In response, we found a significant delay in breeding-cycle phenology for both study species in 2013 relative to other study years: the first bird observed was delayed by 6–10 days, with mean arrival by 3–6 days, territory establishment by 6–13 days, and clutch initiation by 4–10 days. Further, snow cover, temperature, and precipitation during the territory establishment period were important predictors of clutch initiation dates for both species. These findings suggest that Arctic-breeding passerine communities may have the flexibility required to adjust breeding phenology in response to the increasingly extreme and unpredictable environmental conditions—although future generations may encounter conditions that exceed their current range of phenological flexibility.     

What prevents phenological adjustment to climate change in migrant bird species? Evidence against the “arrival constraint” hypothesis

Phenological studies have demonstrated changes in the timing of seasonal events across multiple taxonomic groups as the climate warms. Some northern European migrant bird populations, however, show little or no significant change in breeding phenology, resulting in synchrony with key food sources becoming mismatched. This phenological inertia has often been ascribed to migration constraints (i.e. arrival date at breeding grounds preventing earlier laying). This has been based primarily on research in The Netherlands and Germany where time between arrival and breeding is short (often as few as 9 days). Here, we test the arrival constraint hypothesis over a 15-year period for a U.K. pied flycatcher (Ficedula hypoleuca) population where laying date is not constrained by arrival as the period between arrival and breeding is substantial and consistent (average 27 ± 4.57 days SD). Despite increasing spring temperatures and quantifiably stronger selection for early laying on the basis of number of offspring to fledge, we found no significant change in breeding phenology, in contrast with co-occurring resident blue tits (Cyanistes caeruleus). We discuss possible non-migratory constraints on phenological adjustment, including limitations on plasticity, genetic constraints and competition, as well as the possibility of counter-selection pressures relating to adult survival, longevity or future reproductive success. We propose that such factors need to be considered in conjunction with the arrival constraint hypothesis.     

Winter rainfall predicts phenology in widely separated populations of a migrant songbird

Climate change is affecting behaviour and phenology in many animals. In migratory birds, weather patterns both at breeding and at non-breeding sites can influence the timing of spring migration and breeding. However, variation in responses to weather across a species range has rarely been studied, particularly among populations that may winter in different locations. We used prior knowledge of migratory connectivity to test the influence of weather from predicted non-breeding sites on bird phenology in two breeding populations of a long-distance migratory bird species separated by 3,000 km. We found that winter rainfall showed similar associations with arrival and egg-laying dates in separate breeding populations on an east–west axis: greater rainfall in Jamaica and eastern Mexico was generally associated with advanced American redstart (Setophaga ruticilla) phenology in Ontario and Alberta, respectively. In Ontario, these patterns of response could largely be explained by changes in the behaviour of individual birds, i.e., phenotypic plasticity. By explicitly incorporating migratory connectivity into responses to climate, our data suggest that widely separated breeding populations can show independent and geographically specific associations with changing weather conditions. The tendency of individuals to delay migration and breeding following dry winters could result in population declines due to predicted drying trends in tropical areas and the tight linkage between early arrival/breeding and reproductive success in long-distance migrants.     

Migratory patterns and connectivity of two North American grassland bird species

Effective management and conservation of migratory bird populations require knowledge and incorporation of their movement patterns and space use throughout the annual cycle. To investigate the little‐known migratory patterns of two grassland bird species, we deployed 180 light‐level geolocators on Grasshopper Sparrows (Ammodramus savannarum) and 29 Argos‐GPS tags on Eastern Meadowlarks (Sturnella magna) at Konza Prairie, Kansas, USA, and six US Department of Defense (DoD) installations distributed across the species' breeding ranges. We analyzed location data from 34 light‐level geolocators and five Argos‐GPS tags attached for 1 year to Grasshopper Sparrows and Eastern Meadowlarks, respectively. Grasshopper Sparrows were present on the breeding grounds from mid‐April through early October, substantially longer than previously estimated, and migrated on average ~2,500 km over ~30 days. Grasshopper Sparrows exhibited strong migratory connectivity only at a continental scale. The North American Great Lakes region likely serves as a migratory divide for Midwest and East Coast Grasshopper Sparrows; Midwest populations (Kansas, Wisconsin, and North Dakota; n = 13) largely wintered in Texas or Mexico, whereas East Coast populations (Maryland and Massachusetts, n = 20) wintered in the northern Caribbean or Florida. Our data from Eastern Meadowlarks provided evidence for a diversity of stationary and short‐ and long‐distance migration strategies. By providing the most extensive examination of the nonbreeding movement ecology for these two North American grassland bird species to date, we refine information gaps and provide key insight for their management and conservation.     

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.     

Challenging a 15‐year‐old claim: The North Atlantic Oscillation index as a predictor of spring migration phenology of birds

Many migrant bird species that breed in the Northern Hemisphere show advancement in spring arrival dates. The North Atlantic Oscillation (NAO) index is one of the climatic variables that have been most often investigated and shown to be correlated with these changes in spring arrival. Although the NAO is often claimed to be a good predictor or even to have a marked effect on interannual changes in spring migration phenology of Northern Hemisphere breeding birds, the results on relations between spring migration phenology and NAO show a large variety, ranging from no, over weak, to a strong association. Several factors, such as geographic location, migration phase, and the NAO index time window, have been suggested to partly explain these observed differences in association. A combination of a literature meta‐analysis, and a meta‐analysis and sliding time window analysis of a dataset of 23 short‐ and long‐distance migrants from the constant‐effort trapping garden at Helgoland, Germany, however, paints a completely different picture. We found a statistically significant overall effect size of the NAO on spring migration phenology (coefficient = ?0.14, SE = 0.054), but this on average only explains 0%–6% of the variance in spring migration phenology across all species. As such, the value and biological meaning of the NAO as a general predictor or explanatory variable for climate change effects on migration phenology of birds, seems highly questionable. We found little to no definite support for previously suggested factors, such as geographic location, migration phenology phase, or the NAO time window, to explain the heterogeneity in correlation differences. We, however, did find compelling evidence that the lack of accounting for trends in both time series has led to strongly inflated (spurious) correlations in many studies (coefficient = ?0.13, SE = 0.019).     

Using bioacoustics to examine shifts in songbird phenology

Monitoring patterns in biodiversity and phenology have become increasingly important given accelerating levels of anthropogenic change. Long‐term monitoring programs have reported earlier occurrence of spring activity, reflecting species response to climate change. Although tracking shifts in spring migration represents a valuable approach to monitoring community‐level consequences of climate change, robust long‐term observations are challenging and costly. Audio recordings and metrics of bioacoustic activity could provide an effective method for monitoring changes in songbird activity and broader biotic interactions. We used 3 years of spring and fall recordings at six sites in Glacier Bay National Park, Alaska, an area experiencing rapid warming and glacial retreat, to examine the utility of bioacoustics to detect changes in songbird phenology. We calculated the Acoustic Complexity Index (ACI), an algorithm representing an index of bird community complexity. Abrupt changes in ACI values from winter to spring corresponded to spring transition, suggesting that ACI may be an effective, albeit coarse metric to detect the arrival of migrating songbirds. The first peak in ACI shifted from April 16 to April 11 from 2012 to 2014. Changes in ACI were less abrupt in the fall due to weather events, suggesting spring recordings are better suited to indicate phenology. To ensure changes in ACI values were detecting real changes in songbird activity, we explored the relationship between ACI and song of three species: varied thrush   (Ixoreus naevius), Pacific wren (Troglodytes pacificus), and ruby‐crowned kinglet (Regulus calendula). ACI was positively related to counts of all species, but most markedly with song of the varied thrush, the most common species in our recordings and a known indicator of forest ecosystem health. We conclude that acoustic recordings paired with bioacoustic indices may be a useful method of monitoring shifts in songbird communities due to climate change and other sources of anthropogenic disturbance.     

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.     

Changes in spring arrival of Nearctic-Neotropical migrants attributed to multiscalar climate

Climate-related changes associated with the California marine ecosystem have been documented; however, there are no studies assessing changes in terrestrial vertebrate phenology on the Pacific coast of western North America. We analyze the spring phenology of 21 Nearctic-Neotropical migratory songbird species in central and northern CA. Using observational and banding data at multiple sites, we evaluate evidence for a change in arrival timing being linked to either nonclimatic or multiscalar climatic explanations. Using correlation analysis, of the 13 species with a significant ( P <0.10) change in arrival, the arrival timing of 10 species (77%) is associated with both temperature and a large-scale climate oscillation index (El Niño Southern Oscillation, ENSO; North Atlantic Oscillation, NAO; and/or Pacific Decadal Oscillation, PDO) at least at one location. Eight of the 13 species (62%) are advancing their migratory timing. All species for which spring arrival is associated with climate at multiple locations are exhibiting changes ( n =5) and all species lacking evidence for association between migration phenology and climate ( n =3) exhibit no change. Migrants tend to arrive earlier in association with warmer temperatures, positive NAO indices, and stronger ENSO indices. Twelve species negatively correlate ( P ≤0.05) with local or regional temperature at least at one location; five species negatively correlate with ENSO. Eleven species' arrival is correlated ( P ≤0.05) with NAO; 10 are negatively associated. After an exhaustive literature search, this is apparently the first documentation of an association between NAO and migratory phenology in western North America.     

Range-wide migration corridors and non-breeding areas of a northward expanding Afro-Palaearctic migrant,the European Bee-eater Merops apiaster

Across their ranges, different populations of migratory species often use separate routes to migrate between breeding and non-breeding grounds. Recent changes in climate and land-use have led to breeding range expansions in many species but it is unclear whether these populations also establish new migratory routes, non-breeding sites and migration phenology. Thus, we compared the migration patterns of European Bee-eaters Merops apiaster from two established western (n = 5) and eastern (n = 6) breeding populations in Europe, with those from a newly founded northern population (n = 19). We aimed to relate the breeding populations to the two known non-breeding clusters in Africa, and to test for similarities of migration routes and timing between the old and new populations. Western Bee-eaters used the western flyway to destinations in West Africa; the eastern birds uniformly headed south to southern African non-breeding sites, confirming a complete separation in time and space between these long-established populations. The recently founded northern population, however, also used a western corridor, but crossed the Mediterranean further east than the western population and overwintered mainly in a new non-breeding area in southern Congo/northern Angola. The migration routes and the new non-breeding range overlapped only slightly with the western, but not with the eastern, population. In contrast, migration phenology appeared to differ between the western and both the northern and the eastern populations, with tracked birds from the western population migrating 2–4 weeks earlier. The northern population thus shares some spatial traits with western Bee-eaters, but similar phenology only with eastern population. This divergence highlights the adjustments in the timing of migration to local environmental conditions in newly founded populations, and a parallel establishment of new breeding and non-breeding sites.     

Seasonal abundance and habitat use of bird species in and around Wondo Genet Forest,south‐central Ethiopia

The habitat use and seasonal migratory pattern of birds in Ethiopia is less explored as compared to diversity studies. To this end, this study aimed at investigating the patterns of distribution related to seasonality and the effect of habitat characteristics (elevation, slope, and average vegetation height) on habitat use of birds of Wondo Genet Forest Patch. A stratified random sampling design was used to assess the avian fauna across the four dominant habitat types found in the study area: natural forest, wooded grassland, grassland, and agroforestry land. A point transect count was employed to investigate avian species richness and abundance per habitat type per season. Ancillary data, such as elevation above sea level, latitude and longitude, average vegetation height, and percent slope inclination, were recorded with a GPS and clinometers per plot. A total of 33 migratory bird species were recorded from the area, of which 20 species were northern (Palearctic) migrants while 13 were inter‐African migrants. There was a significant difference in the mean abundance of migratory bird species between dry and wet seasons (t = 2.13, p = .038, df = 44). The variation in mean abundance per plot between the dry and wet seasons in the grassland habitat was significant (t = 2.35, p = .051, df = 7). In most habitat types during both dry and wet seasons, omnivore birds were the most abundant. While slope was a good predictor for bird species abundance in the dry season, altitude and average vegetation height accounted more in the wet season. The patch of forest and its surrounding is an important bird area for migratory, endemic, and global threatened species. Hence, it is conservation priority area, and the study suggests that conservation coupled with ecotourism development is needed for its sustainability.     

Climate warming,ecological mismatch at arrival and population decline in migratory birds

Climate is changing at a fast pace, causing widespread, profound consequences for living organisms. Failure to adjust the timing of life-cycle events to climate may jeopardize populations by causing ecological mismatches to the life cycle of other species and abiotic factors. Population declines of some migratory birds breeding in Europe have been suggested to depend on their inability to adjust migration phenology so as to keep track of advancement of spring events at their breeding grounds. In fact, several migrants have advanced their spring arrival date, but whether such advancement has been sufficient to compensate for temporal shift in spring phenophases or, conversely, birds have become ecologically mismatched, is still an unanswered question, with very few exceptions. We used a novel approach based on accumulated winter and spring temperatures (degree-days) as a proxy for timing of spring biological events to test if the progress of spring at arrival to the breeding areas by 117 European migratory bird species has changed over the past five decades. Migrants, and particularly those wintering in sub-Saharan Africa, now arrive at higher degree-days and may have therefore accumulated a ‘thermal delay’, thus possibly becoming increasingly mismatched to spring phenology. Species with greater ‘thermal delay’ have shown larger population decline, and this evidence was not confounded by concomitant ecological factors or by phylogenetic effects. These findings provide general support to the largely untested hypotheses that migratory birds are becoming ecologically mismatched and that failure to respond to climate change can have severe negative impacts on their populations. The novel approach we adopted can be extended to the analysis of ecological consequences of phenological response to climate change by other taxa.     

Overtaking on migration: does longer distance migration always incur a penalty?

For many migratory bird species, the latitudinal range of the winter distribution spans thousands of kilometres, thus encompassing considerable variation in individual migration distances. Pressure to winter near breeding areas is thought to be a strong driver of the evolution of migration patterns, as individuals undertaking a shorter migration are generally considered to benefit from earlier arrival on the breeding grounds. However, the influence of migration distance on timing of arrival is difficult to quantify because of the large scales over which individuals must be tracked. Using a unique dataset of individually‐marked Icelandic black‐tailed godwits Limosa limosa islandica tracked throughout the migratory range by a network of hundreds of volunteer observers, we quantify the consequences of migrating different distances for the use of stop‐over sites and timing of arrival in Iceland. Modelling of potential flight distances and tracking of individuals from across the winter range shows that individuals wintering further from the breeding grounds must undertake a stop‐over during spring migration. However, despite travelling twice the distance and undertaking a stop‐over, individuals wintering furthest from the breeding grounds are able to overtake their conspecifics on spring migration and arrive earlier in Iceland. Wintering further from the breeding grounds can therefore be advantageous in migratory species, even when this requires the use of stop‐over sites which lengthen the migratory journey. As early arrival on breeding sites confers advantages for breeding success, the capacity of longer distance migrants to overtake conspecifics is likely to influence the fitness consequences of individual migration strategies. Variation in the quality of wintering and stopover sites throughout the range can therefore outweigh the benefits of wintering close to the breeding grounds, and may be a primary driver of the evolution of specific migration routes and patterns.     

Length polymorphisms at two candidate genes explain variation of migratory behaviors in blackpoll warblers (Setophaga striata)

Migratory behaviors such as the timing and duration of migration are genetically inherited and can be under strong natural selection, yet we still know very little about the specific genes or molecular pathways that control these behaviors. Studies in candidate genes Clock and Adcyap1 have revealed that both of these loci can be significantly correlated with migratory behaviors in birds, though observed relationships appear to vary across species. We investigated geographic genetic structure of Clock and Adcyap1 in four populations of blackpoll warblers (Setophaga striata), a Neotropical–Nearctic migrant that exhibits geographic variation in migratory timing and duration across its boreal breeding distribution. Further, we used data on migratory timing and duration, obtained from light‐level geolocator trackers to investigate candidate genotype–phenotype relationships at the individual level. While we found no geographic structure in either candidate gene, we did find evidence that candidate gene lengths are correlated with five of the six migratory traits. Maximum Clock allele length was significantly and negatively associated with spring arrival date. Minimum Adcyap1 allele length was significantly and negatively associated with spring departure date and positively associated with fall arrival date at the wintering grounds. Additionally, we found a significant interaction between Clock and Adcyap1 allele lengths on both spring and fall migratory duration. Adcyap1 heterozygotes also had significantly shorter migration duration in both spring and fall compared to homozygotes. Our results support the growing body of evidence that Clock and Adcyap1 allele lengths are correlated with migratory behaviors in birds.     

Novel methods reveal shifts in migration phenology of barn swallows in South Africa

Many migratory bird species, including the barn swallow (Hirundo rustica), have advanced their arrival date at Northern Hemisphere breeding grounds, showing a clear biotic response to recent climate change. Earlier arrival helps maintain their synchrony with earlier springs, but little is known about the associated changes in phenology at their non-breeding grounds. Here, we examine the phenology of barn swallows in South Africa, where a large proportion of the northern European breeding population spends its non-breeding season. Using novel analytical methods based on bird atlas data, we show that swallows first arrive in the northern parts of the country and gradually appear further south. On their north-bound journey, they leave South Africa rapidly, resulting in mean stopover durations of 140 days in the south and 180 days in the north. We found that swallows are now leaving northern parts of South Africa 8 days earlier than they did 20 years ago, and so shortened their stay in areas where they previously stayed the longest. By contrast, they did not shorten their stopover in other parts of South Africa, leading to a more synchronized departure across the country. Departure was related to environmental variability, measured through the Southern Oscillation Index. Our results suggest that these birds gain their extended breeding season in Europe partly by leaving South Africa earlier, and thus add to scarce evidence for phenology shifts in the Southern Hemisphere.     

Changes in migration phenology and biometrical traits of Reed,Marsh and Sedge Warblers

Global environmental processes like climate change could severely affect population level migratory behaviour of long range migrant birds. We analyzed changes in migration phenology and biometrics of three closely-related long-distance migrant Acrocephalus species. We used the records of 12 063 Sedge, 12 913 Reed, and 5 409 Marsh Warblers caught and ringed between 1989–2009, at a Hungarian stopover site. Quantile regressions were used to analyse the changes in spring and autumn migration phenology. Median spring arrival date of Sedge and Reed Warblers shifted 6.5 and 7.5 days earlier, respectively. Autumn arrival of all species shifted one (Reed and Marsh Warblers) or two (Sedge Warbler) weeks later. Mean body mass of adult Reed and Marsh Warblers decreased in spring (by 0.3 and 0.2 grams, respectively) and in autumn (by 0.8 and 0.2 grams, respectively) while body mass of adult Sedge Warblers decreased only in autumn (by 0.4 grams). Mean wing length of all species increased significantly (range of change: 0.6–1 mm). Despite the fact that the studied species are closely related, all three have remarkably different migration strategies. However, similar patterns can be observed in the studied parameters, indicating that global processes may have general effects on these species, albeit through markedly different mechanisms.     

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.     

Drought reshuffles plant phenology and reduces the foraging benefit of green‐wave surfing for a migratory ungulate

To increase resource gain, many herbivores pace their migration with the flush of nutritious plant green‐up that progresses across the landscape (termed “green‐wave surfing”). Despite concerns about the effects of climate change on migratory species and the critical role of plant phenology in mediating the ability of ungulates to surf, little is known about how drought shapes the green wave and influences the foraging benefits of migration. With a 19 year dataset on drought and plant phenology across 99 unique migratory routes of mule deer (Odocoileus hemionus) in western Wyoming, United States, we show that drought shortened the duration of spring green‐up by approximately twofold (2.5 weeks) and resulted in less sequential green‐up along migratory routes. We investigated the possibility that some routes were buffered from the effects of drought (i.e., routes that maintained long green‐up duration irrespective of drought intensity). We found no evidence of drought‐buffered routes. Instead, routes with the longest green‐up in non‐drought years also were the most affected by drought. Despite phenological changes along the migratory route, mule deer closely followed drought‐altered green waves during migration. Migrating deer did not experience a trophic mismatch with the green wave during drought. Instead, the shorter window of green‐up caused by drought reduced the opportunity to accumulate forage resources during rapid spring migrations. Our work highlights the synchronization of phenological events as an important mechanism by which climate change can negatively affect migratory species by reducing the temporal availability of key food resources. For migratory herbivores, climate change poses a new and growing threat by altering resource phenology and diminishing the foraging benefit of migration.