Northward range expansion in spring‐staging barnacle geese is a response to climate change and population growth,mediated by individual experience

All long‐distance migrants must cope with changing environments, but species differ greatly in how they do so. In some species, individuals might be able to adjust by learning from individual experiences and by copying others. This could greatly speed up the process of adjustment, but evidence from the wild is scarce. Here, we investigated the processes by which a rapidly growing population of barnacle geese (Branta leucopsis) responded to strong environmental changes on spring‐staging areas in Norway. One area, Helgeland, has been the traditional site. Since the mid‐1990s, an increasing number of geese stage in another area 250 km further north, Vesterålen. We collected data on goose numbers and weather conditions from 1975 to 2017 to explore the extent to which the increase in population size and a warmer climate contributed to this change in staging area use. During the study period, the estimated onset of grass growth advanced on average by 0.54 days/year in each of the two areas. The total production of digestible biomass for barnacle geese during the staging period increased in Vesterålen but remained stable in Helgeland. The goose population has doubled in size during the past 25 years, with most of the growth being accommodated in Vesterålen. The observations suggest that this dramatic increase would not have happened without higher temperatures in Vesterålen. Records of individually marked geese indicate that from the initial years of colonization onwards, especially young geese tended to switch to Vesterålen, thereby predominating in the flocks at Vesterålen. Older birds had a lower probability of switching to Vesterålen, but over the years, the probability increased for all ages. Our findings suggest that barnacle geese integrate socially learned behaviour with adjustments to individual experiences, allowing the population to respond rapidly and accurately to global change.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Central European Greylag Geese Anser anser show a shortening of migration distance and earlier spring arrival over 60 years

Global climate change can cause pronounced changes in species? migratory behaviour. Numerous recent studies have demonstrated climate‐driven changes in migration distance and spring arrival date in waterbirds, but detailed studies based on long‐term records of individual recapture or re‐sighting events are scarce. Using re‐sighting data from 430 marked individuals spanning a 60‐year period (winters 1956/1957 to 2015/2016), we assessed patterns in migration distance and spring arrival date, wintering‐site fidelity and survival in the increasing central European breeding population of Greylag Geese Anser anser. We demonstrate a long‐term decrease in migration distance, changes in the wintering range caused by winter partial short‐stopping, and the earlier arrival of geese on their breeding grounds. Greylag Geese marked on central Europe moulting grounds have not been recorded wintering in Spain since 1986 or in Tunisia and Algeria since 2004. The migration distance and spring arrival of geese indicated an effect of temperature at the breeding site and values of the NAO index. Greylag Geese migrate shorter distances and arrive earlier in milder winters. We suggest that shifts in the migratory behaviour of Central European Greylag Geese are individual temperature‐dependent decisions to take advantage of wintering grounds becoming more favourable closer to their breeding grounds, allowing birds to acquire breeding territories earlier.     

The reliance on distant resources for egg formation in high Arctic breeding barnacle geese Branta leucopsis

Breeding in the high Arctic is time constrained and animals should therefore start with their annual reproduction as early as possible. To allow for such early reproduction in migratory birds, females arrive at the breeding grounds either with body stores or they try to rapidly develop their eggs after arrival using local resources. Svalbard breeding barnacle geese Branta leucopsis have to fly non‐stop for about 1100 km from their last continental staging site to the archipelago making the transport of body stores costly. However, environmental conditions at the breeding grounds are highly unpredictable favouring residual body stores allowing for egg production after arrival on the breeding grounds. We estimated the reliance on southern continental resources, i.e. body stores for egg formation, in barnacle geese using stable isotope ratios in the geese's forage along the flyway and in their eggs. Females adopted mixed breeding strategies by using southern resources as well as local resources to varying extents for egg formation. Southern capital in lipid‐free yolk averaged 41% (range: 23–65%), early laid eggs containing more southern capital than eggs laid late in the season. Yolk lipids and albumen did not vary over time and averaged a southern capital proportion of 54% (range: 32–73%) and 47% (range: 25–88%), respectively. Our findings indicate that female geese vary the use of southern resources when synthesising their eggs and this allocation also varies among egg tissues. Their mixed and flexible use of distant and local resources potentially allows for adaptive adjustments to environmental conditions encountered at the archipelago just before breeding.     

Relationship between arrival timing and breeding success of intra‐tropical migratory Fork‐tailed Flycatchers (Tyrannus savana)

For migratory birds, early arrival at breeding areas has many benefits, such as acquisition of better territories and mates. This strategy has been found in numerous species breeding at north‐temperate latitudes, but has not been yet reported for intra‐tropical migratory species. We evaluated the relationship between arrival date, initiation of breeding, and breeding success of Fork‐tailed Flycatchers (Tyrannus savana) breeding in southeastern Brazil and overwintering in northern South America. We color‐banded adult flycatchers during three breeding seasons and searched for them during the following breeding seasons. We also monitored nests from construction until either failure or fledging of young. We found that: (1) male Fork‐tailed Flycatchers arrived at the breeding site earlier than females, (2) males that arrived earlier had greater breeding success, and (3) nests where eggs were laid earlier in the breeding season were more likely to be successful than those where eggs were laid later. Male Fork‐tailed Flycatchers appeared to benefit from early arrival at a tropical breeding site, potentially mediated by their ability to acquire a high‐quality territory and mate as early as possible, and by the ability of their mate to begin breeding as early as possible. Breeding success for female Fork‐tailed Flycatchers may be determined primarily by a combination of the arrival date of their mate and how quickly they can begin breeding. Our results suggest that protandry occurs in an intra‐tropical migratory bird and that early arrival of males and early initiation of reproduction by females results in greater reproductive success. A better understanding of the underlying mechanisms that control the timing of migration and reproduction of this and other intra‐tropical migratory species is important for evaluating the challenges they face in light of current and future rapid environmental changes.     

Individually tracked geese follow peaks of temperature acceleration during spring migration

Many migratory herbivores seem to follow the flush of plant growth during migration in order to acquire the most nutrient‐rich plants. This has also been hypothesized for arctic‐breeding geese, but so far no test of this so‐called green wave hypothesis has been performed at the individual level. During four years, a total of 30 greater white‐fronted geese Anser albifrons albifrons was tracked using GPS transmitters, of which 13 yielded complete spring migration tracks. From those birds we defined stopover sites and related the date of arrival at each of these stopovers to temperature sum (growing degree days, GDD), snow cover, accumulated photoperiod and latitude. We found that geese arrived at spring stopovers close to the peak in GDD jerk; the ‘jerk’ is the third derivative, or the rate of change in acceleration, and GDD jerk maxima therefore represent the highest acceleration of daily temperature per site. Day of snow melt also correlated well with the observed arrival of the geese. Factors not closely related to onset of spring, i.e. accumulated photoperiod and latitude, yielded poorer fits. A comparison with published data revealed that the GDD jerk occurs 1–2 weeks earlier than the onset of spring derived from NDVI, and probably represents the very start of spring growth. Our data therefore suggest that white‐fronted geese track the front of the green wave in spring.     

Early arrival is not associated with more extra‐pair fertilizations in a long‐distance migratory bird

When assessing the benefits of early arrival date of migratory birds, a hidden and often ignored component of males’ fitness is the higher chance of early‐arriving birds to obtain extra‐pair fertilizations. Here we investigated how extra‐pair paternity might affect the relationship between male arrival date and number of fertilizations in a model study system, the European pied flycatcher Ficedula hypoleuca. For this purpose, we sampled and genotyped breeding pairs, unpaired males and offspring (including embryos from unhatched eggs when possible) of a Dutch pied flycatcher population. Detailed information on arrival date of males, egg laying date of their social mates and nest success was also recorded. Early‐arriving males had early‐laying females and males with early‐laying females had a higher probability of siring extra‐pair eggs and obtain more fertilizations. However, male arrival date alone did not correlate with the probability to gain extra‐pair paternity and neither to the amount of fertilized eggs. Both early‐ and late‐arriving males had a higher probability of losing paternity in their own nest compared to birds with an intermediate arrival date. Finally, late‐arriving males were more likely to remain unpaired but, interestingly, a few of these birds obtained paternity via extra‐pair copulations. Because earlier arrival date did not lead to more extra‐pair fertilizations and because such relationship seems to be driven mainly by the female's laying date, we conclude that the contribution of extra‐pair paternity to the overall fitness benefits of early male arrival date is relatively small.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

Potential for an Arctic‐breeding migratory bird to adjust spring migration phenology to Arctic amplification

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long‐distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.     

Eggs brought in from afar: Svalbard‐breeding pink‐footed geese can fly their eggs across the Barents Sea

Many Arctic‐breeding waterbirds are thought to bring nutrients for egg production from southern latitudes to allow early breeding. It has proved problematic to quantify the extent of such capital breeding and identify whether nutrients for egg production are brought in from nearby or from afar. Before reaching their breeding grounds on Svalbard, pink‐footed geese Anser brachyrhynchus fly ～ 1100 km across the Barents Sea from Norway. Using abdominal profile indexing (API) we scored body stores in individually marked geese just prior to migration from the northernmost staging area in Norway to Svalbard, followed by their breeding success on their non‐breeding grounds in autumn. In productive breeding years leading to a high (> 13.8%) proportion of juveniles in the autumn population, there was a positive relationship between female API and number of young produced, suggesting that the geese are at least partial capital breeders. Moreover, focusing on the geographic origin of proteins used in egg synthesis and measuring nitrogen stable isotope ratios in pink‐footed geese's eggs and food sources in Norway and Svalbard, we identified that capital breeding in this species is ～ 50% on average but may potentially amount to as much as 100%, notably in females laying early. About 60% of this protein capital is carried in well‐developed follicles across the Barents Sea, the remainder likely being stored in muscle tissues. Conditions on the wintering grounds and migratory stopover sites can have profound effects on an individual's fitness but the here presented link between the use of migratory stopover sites and breeding performance is particularly noteworthy. Apparently, some individuals accept the putative costs of carrying body stores over large distances to the breeding grounds. The data also highlights considerable variation in the reliance on capital for breeding, suggesting substantial individual scope to adjust breeding strategy to changing environmental conditions.     

Increase in protandry over time in a long‐distance migratory bird

Protandry is a widespread life‐history phenomenon describing how males precede females at the site or state of reproduction. In migratory birds, protandry has an important influence on individual fitness, the migratory syndrome, and phenological response to climate change. Despite its significance, accurate analyses on the dynamics of protandry using data sets collected at the breeding site, are lacking. Basing our study on records collected during two time periods, 1979 to 1988 and 2006 to 2016, we aim to investigate protandry dynamics over 38 years in a breeding population of willow warblers (Phylloscopus trochilus). Change in the timing of arrival was analyzed in males and females, and protandry (number of days between male and female arrival) was investigated both at population level and within breeding pairs. Our results show advancement in the arrival time at the breeding site in both sexes, but male arrival has advanced to a greater extent, leading to an increase in protandry both at the population level and within breeding pairs. We did not observe any change in sex ratio that could explain the protandry increase, but pronounced temperature change has occurred and been reported in the breeding area and along the migratory route. Typically, natural selection opposes too early arrival in males, but given warmer springs, this counteracting force may be relaxing, enabling an increase in protandry. We discuss whether our results suggest that climate change has induced sex‐specific effects, if these could be evolutionary and whether the timing of important life‐history stages such as arrival at the breeding site may change at different rates in males and females following environmental shifts.     

Carry‐over effects and compensation: late arrival on non‐breeding grounds affects wing moult but not plumage or schedules of departing bar‐tailed godwits Limosa lapponica baueri

In the annual cycle of migratory birds, temporal and energetic constraints can lead to carry‐over effects, in which performance in one life history stage affects later stages. Bar‐tailed godwits Limosa lapponica baueri, which achieve remarkably high pre‐migratory fuel loads, undertake the longest non‐stop migratory flights yet recorded, and breed during brief high‐latitude summers, may be particularly vulnerable to persistent effects of disruptions to their rigidly‐timed annual routines. Using three years of non‐breeding data in New Zealand, we asked how arrival timing after a non‐stop flight from Alaska (>11 000 km) affected an individual godwit's performance in subsequent flight feather moult, contour feather moults, and migratory departure. Late arrival led to later wing moult, but godwits partially compensated for delayed moult initiation by increasing moult rate and decreasing the total duration of moult. Delays in arrival and wing moult up to 34–37 d had no apparent effect on an individual's migratory departure or extent of breeding plumage at departure, both of which were extraordinarily consistent between years. Thus, ‘errors’ in timing early in the non‐breeding season were essentially corrected in New Zealand prior to spring migration. Variation in migration timing also had no apparent effect on an individual's likelihood of returning the following season. The bar‐tailed godwits’ rigid maintenance of plumage and spring migration schedules, coupled with high annual survival, imply a surprising degree of flexibility to address unforeseen circumstances in the annual cycle.     

Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration

Migration during spring is usually faster than during autumn because of competition for breeding territories. In some cases, however, the costs and benefits associated with the environment can lead to slower spring migration, but examples are quite rare. We compared seasonal migration strategies of the endangered Baltic population of the dunlin Calidris alpina schinzii using light‐level geolocator data from 26 individuals breeding in Finland. Autumn migration was faster, with individuals showing a ‘jump’ and ‘skipping’ migration strategy characterised by fewer stationary periods, shorter total stopping time and faster flight. Spring migration was slower, with individuals using a ‘skipping’ strategy. The duration of migration was longer for early departing birds during spring but not during autumn suggesting that early spring migrants are prevented from arriving to the breeding areas or that fueling conditions are worse on the stopover sites for early arriving individuals. Dunlins showed high migratory connectivity. All individuals had one long staging at the Wadden Sea in the autumn after which half of the individuals flew 4500 km non‐stop to Banc d’Arguin, Mauritania. The other half stopped briefly on the Atlantic coast on their way to Mauritania. One bird wintered on the coast of Portugal. Nine individuals that carried geolocators for two years were site faithful to their final non‐breeding sites. Based on the strategies during the non‐breeding period we identified, Baltic dunlin may be especially vulnerable to rapid environmental changes at the staging and non‐breeding areas. Consequently, the preservation of the identified non‐breeding areas is important for their conservation.     

Forage plants of an Arctic‐nesting herbivore show larger warming response in breeding than wintering grounds,potentially disrupting migration phenology

During spring migration, herbivorous waterfowl breeding in the Arctic depend on peaks in the supply of nitrogen‐rich forage plants, following a “green wave” of grass growth along their flyway to fuel migration and reproduction. The effects of climate warming on forage plant growth are expected to be larger at the Arctic breeding grounds than in temperate wintering grounds, potentially disrupting this green wave and causing waterfowl to mistime their arrival on the breeding grounds. We studied the potential effect of climate warming on timing of food peaks along the migratory flyway of the Russian population of barnacle geese using a warming experiment with open‐top chambers. We measured the effect of 1.0–1.7°C experimental warming on forage plant biomass and nitrogen concentration at three sites along the migratory flyway (temperate wintering site, temperate spring stopover site, and Arctic breeding site) during 2 months for two consecutive years. We found that experimental warming increased biomass accumulation and sped up the decline in nitrogen concentration of forage plants at the Arctic breeding site but not at temperate wintering and stop‐over sites. Increasing spring temperatures in the Arctic will thus shorten the food peak of nitrogen‐rich forage at the breeding grounds. Our results further suggest an advance of the local food peak in the Arctic under 1–2°C climate warming, which will likely cause migrating geese to mistime their arrival at the breeding grounds, particularly considering the Arctic warms faster than the temperate regions. The combination of a shorter food peak and mistimed arrival is likely to decrease goose reproductive success under climate warming by reducing growth and survival of goslings after hatching.     

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.     

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.     

The onset of spring and timing of migration in two arctic nesting goose populations: the pink‐footed goose Anser bachyrhynchus and the barnacle goose Branta leucopsis

An earlier onset of spring has been recorded for many parts of Eurasia in recent decades. This has consequences for migratory species, both in changing the conditions encountered by individuals on reaching migratory sites and in affecting cues regulating the timing of migration where decisions to migrate are influenced by local environmental variables. Here we examine the timing of spring migration for two arctic goose populations, the pink‐footed goose Anser brachyrhynchus (during 1990–2003) and barnacle goose Branta leucopsis (during 1982–2003), which both breed on Svalbard. The satellite‐derived Normalised Difference Vegetation Index (NDVI) was used to express the onset of spring at their wintering and spring staging sites. Pink‐footed geese use several sites during spring migration, ranging from the southernmost wintering areas in Belgium to two spring staging areas in Norway, and distances between sites used along the flyway are relatively short. There was a positive correlation in the onset of spring between neighbouring sites, and the geese migrated earlier in early springs. Barnacle geese, on the other hand, have a long overseas crossing from their wintering grounds in Britain to spring staging areas in Norway. Although spring advanced in both regions, there was no corresponding correlation in the timing of onset of spring between their wintering and spring staging sites, and little evidence for barnacle geese migrating earlier over the whole study period. Hence, where geese can use spring conditions at one site as an indicator of the conditions they might encounter at the next, they have responded quickly to the advancement of spring, whereas in a situation where they cannot predict, they have not yet responded, despite the advancement of spring in the spring staging area.     

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.     

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.     

Starting over: experimental effects of breeding delay on reproductive success in early‐arriving male American redstarts

Birds that arrive and breed early often have higher reproductive success than late individuals, either as a consequence of timing‐specific advantages (the timing hypothesis) or because these individuals and/or their resources are of higher quality (the quality hypothesis). In this study, we examined the potential influence of several factors affecting reproductive success by experimentally delaying breeding of early‐arriving male American redstarts Setophaga ruticilla, a species for which early male arrival is strongly related to increased reproductive success. Our manipulation involved the capture, holding, and release of males following pairing and territory establishment, resulting in the majority of subjects (67%) losing their initial mate (47%) or mate and territory (20%) and forcing them to start over approximately 12 d after their initial arrival. Males forced to start over (i.e. those losing their first territory and/or mate) did not experience any decrease in body condition, nor did their reproductive behaviour differ from that of early‐arriving control males. We found that naturally early‐arriving but experimentally manipulated males suffered reduced fledging success in comparison to early‐arriving males that bred early or late, but equivalent success in comparison to males that arrived and bred late. Based on our results, we propose that the relationship between early arrival and higher reproductive success in this species is mediated not simply by individual male quality or absolute arrival timing alone, but rather some other aspect of resource quality is likely important. We discuss and present evidence for two alternative explanations under the quality hypothesis: female quality and territory quality. To our knowledge, our study is the first to investigate the effects of experimentally delaying male breeding time, strengthening previous correlational evidence for resource quality as a potentially important selective agent driving early arrival in migratory birds.