Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

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.     

Plasma mammalian leptin analogue predicts reproductive phenology,but not reproductive output in a capital‐income breeding seaduck

To invest in energetically demanding life history stages, individuals require a substantial amount of resources. Physiological traits, particularly those related to energetics, can be useful for examining variation in life history decisions and trade‐offs because they result from individual responses to environmental variation. Leptin is a protein hormone found in mammals that is proportional to the amount of endogenous fat stores within an individual. Recently, researchers have confirmed that a mammalian leptin analogue (MLA), based on the mammalian sequence of leptin, is present with associated receptors and proteins in avian species, with an inhibitory effect on foraging and body mass gain at high circulating levels. While MLA has been both quantified and manipulated in avian species, little is currently known regarding whether plasma MLA in wild‐living species and individuals is associated with key reproductive decisions. We quantified plasma MLA in wild, Arctic‐nesting female common eiders (Somateria mollissima) at arrival on the breeding grounds and followed them to determine subsequent breeding propensity, and reproductive phenology, investment, and success. Common eiders are capital‐income breeding birds that require the accumulation of substantial fat stores to initiate laying and successfully complete incubation. We found that females with lower plasma MLA initiated breeding earlier and in a shorter period of time. However, we found no links between plasma MLA levels and breeding propensity, clutch size, or reproductive success. Although little is still known about plasma MLA, based on these results and its role in influencing foraging behaviors and condition gain, plasma MLA appears to be closely linked to reproductive timing and is therefore likely to underlie trade‐offs surrounding life history decisions.     

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.     

Pay-offs and penalties of competing migratory schedules

We relate variation in the timing of arrival by migrating birds breeding at northerly latitudes to individual differences in the prior accumulation of energy stores. Balancing starvation risks early in the season against the almost universal declining trend in reproductive prospects with advancing date is seen as an individual decision with fitness consequences. We review three studies implicating events at the staging sites or in winter in setting the individual migratory schedule. Climate change influences the timetable of a pied flycatcher (Ficedula hypoleuca) population breeding in The Netherlands and wintering in West Africa, followed since 1960. Mean air temperature in the period mid April‐mid May (arrival and laying) increased and laying date advanced by 10 days. Still, in recent years most birds did not lay early enough to maximise fitness (determined by recruitment and parental survival) whereas many parents achieved this goal in 1980–1985. As the flycatchers have not started to arrive earlier, some ecological constraint further upstream is postulated (possibly the hurdle of the crossing of Sahara and Mediterranean). The ability to follow individual migrants provides a second avenue to assess the fitness implications of migratory schedules. Thus, brightly coloured male bar‐tailed godwits (Limosa lapponica) captured in the Dutch Wadden Sea (the intermediate staging site linking a West African wintering area with breeding sites in arctic Russia) and traced with miniature radio‐transmitters did not depart early. The ‘best’ males (with bright breeding plumage) were picked up by the listening stations in Sweden 650 km further along the migratory route ten days later than the paler individuals. If early arrival confers the competitive advantage of prior occupancy but increases mortality, the ‘best’ males may be able to afford arriving later and thus avoid some of the survival costs. Return rate of the ‘bright’ males to the staging site in later seasons was indeed higher than for the ‘pale’, early males. Intensive observation of pink‐footed geese (Anser brachyrhynchus) fitted with coded neck‐collars substantiate the tight relationship between energy stores (fat) accumulated up to final departure from the final staging site (Vesterålen, N. Norway) en route to the nesting grounds (Spitsbergen) and subsequent success. The breeding outcome of individual parents (accompanied by juveniles or not) could be related to observations of body condition before departure (visual ‘abdominal profile index’). Recently, perceived conflicts with agriculture have resulted in widespread harassment by humans. The geese have: drastically shortened their stay on the Vesterålen, fail to achieve the body condition usual a decade ago and reproductive output has fallen. Although the geese are currently pioneering new staging sites, an adequate alternative has not materialised, underlining the critical role of the final take‐off site.     

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.     

Colony attendance and at‐sea distribution of thin‐billed prions during the early breeding season

Procellariiform seabirds have extreme life histories; they are very long‐lived, first breed when relatively old, lay single egg clutches, both incubation and chick‐rearing are prolonged and chicks exhibit slow growth. The early part of the breeding season is crucial, when pair bonds are re‐established and partners coordinate their breeding duties, but is a difficult period to study in burrow‐nesting species. Miniature geolocators (Global Location Sensors or GLS loggers) now offer a way to collect data on burrow attendance, as well as determine at‐sea movements. We studied the early breeding season in thin‐billed prions Pachyptila belcheri breeding at New Island, Falkland Islands. Males and females arrived back at the colony at similar times, with peak arrival in the last days of September. However, males spent more time on land during the pre‐laying period, presumably defending and maintaining the burrow and maximising mating opportunities. Males departed later than females, and carried out a significantly shorter pre‐laying exodus. Males took on the first long incubation shift, whereas females returned to sea soon after egg laying. During the pre‐laying exodus and incubation, males and females travelled at similar speeds (> 250 km d^?1) and were widely distributed over large areas of the Patagonian Shelf. Inter‐annual differences in oceanographic conditions were stronger during the incubation than during the pre‐laying exodus and were matched by stronger differences in distribution. The study thus suggests that extended trips and flexible distribution enable thin‐billed prions to meet the high energy demands of egg production and incubation despite low productivity in waters around the colony during the early summer.     

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.     

Delayed timing of breeding as a cost of reproduction

Timing of breeding is a trait with considerable individual variation, often closely linked to fitness because of seasonal declines in reproduction. The drivers of this variation have received much attention, but how reproductive costs may influence the timing of subsequent breeding has been largely unexplored. We examined a population of northern wheatears Oenanthe oenanthe to compare three groups of individuals that differed in their timing of breeding termination and reproductive effort to investigate how these factors may carry over to influence reproductive timing and reproductive output in the following season. Compared to females that bred successfully, females that put in less effort and terminated breeding early due to nest failure tended to arrive and breed earlier in year 2 (mean advancement = 2.2 and 3.3 d respectively). Females that spent potentially more effort and terminated breeding later due to production of a replacement clutch after nest failure, arrived later than other females in year 2. Reproductive output (number of fledglings) in year 2 differed between the three groups as a result of group‐level differences in the timing of breeding in combination with the general seasonal decline in reproductive output. Our study shows that the main cost of reproduction was apparent in the timing of arrival and breeding in this migratory species. Hence, reproductive costs can arise through altered timing of breeding since future reproductive success (including adult survival) is often dependent on the timing of breeding in seasonal systems.     

Age‐specific variation in relationship between moult and pre‐migratory fuelling in Wood Sandpipers Tringa glareola in southern Africa

Trans‐equatorial avian migrants tend to breed, moult and migrate – the main energy‐requiring events in their lifecycle – at different times. Little is known about the relationship between wing moult and pre‐migratory fuelling in waders on their non‐breeding grounds, where time is less constrained than during their brief high‐latitude breeding season. We determined age‐related strategies of Wood Sandpipers Tringa glareola to balance the energetic demands of primary moult against pre‐migratory fuelling in southern Africa by analysing body mass and primary moult at first capture of 1721 birds mist‐netted in 1972–96 at waterbodies in Zimbabwe. Adults moulted all their primaries in August–December, but immatures underwent a supplemental moult of varying numbers of outer primaries in December–April, close to departure. We used locally weighted linear regression to estimate trends in Wood Sandpiper body mass from 1 July to 1 May. They maintained low mass from arrival in July–September to February–early March. Adults fuelled from 10 February to 1 May at a mean rate of 0.25 g/day (sd = 0.16). Most adults (98%) began fuelling 10–75 days after completing primary moult. Immatures fuelled from 4 March to 13 April at 0.24 g/day (sd = 0.14). They used varying strategies depending on their condition: a brief gap between moult and fuelling; an overlap of these processes near departure, leading to slower fuelling; or skipping fuelling altogether and staying in southern Africa for a ‘gap year’. Immatures moulting three or five outer primaries fuelled more slowly than post‐moult birds. Immatures moulting four outer primaries started fuelling 3 weeks later but at a higher rate than did post‐moult birds of this group. In post‐moult immatures, the later they ended moult, the later and faster they fuelled. The heaviest adults and immatures using all moult patterns accumulated fuel loads of c. 50% of lean body mass, and could potentially cross 2397–4490 km to reach the Great Rift Valley in one non‐stop flight. Immatures were more flexible in the timing and extent of moult and in the timing and rate of fuelling than adults. This flexibility enables inexperienced Wood Sandpipers to cope with inter‐annual differences in feeding conditions at Africa's ephemeral inland waterbodies.     

Sex‐specific patterns of fuelling and pre‐breeding body moult of Little Stints Calidris minuta in South Africa

The timing and duration of each stage of the life of a long‐distance migrant bird are constrained by time and resources. If the parental roles of males and females differ, the timing of other life stages, such as moult or pre‐migratory fuelling, may also differ between the sexes. Little is known about sexual differences for species with weak sexual dimorphism, but DNA‐sexing enables fresh insights. The Little Stint Calidris minuta is a monomorphic long‐distance migrant wader breeding in the Arctic tundra. Males compete for territories and perform elaborate aerial displays. Females produce two clutches a season. Each sex may be a bigamist and incubate one nest a season, each with a different partner. We expect that these differences in breeding behaviour entail different preparations for breeding by males and females, so we aimed to determine whether Little Stints showed any sex differences in their strategies for pre‐breeding moult and pre‐migratory fuelling at their non‐breeding grounds in South Africa. We used body moult records, wing length and body mass of 241 DNA‐sexed Little Stints that we caught and ringed between 27 January and 29 April in 2008–2018 at two neighbouring wetlands in North West Province, South Africa. For each individual we assessed the percentage of breeding plumage on its upperparts and took blood samples for DNA‐sexing. We calculated an adjusted Body Moult Index and an adjusted Wing Coverts Moult Index, then used the Underhill–Zucchini moult model to estimate the start dates and the rate of body moult in males and females. We estimated the changes in the sex ratio of the local population during their stay in South Africa, and also estimated the timing and rate of pre‐migratory fuelling and the potential flight ranges for males and females. The males started body moult on average on 7 February and the females on 12 February, but the sexes did not differ in their timing of wing covert moult, which started on average on 10 February. In January to mid‐February, males constituted c. 57% of the population, but their proportion declined afterwards, indicating an earlier departure than females. We estimated that both sexes began pre‐migratory fuelling on average on 15 March. The sexes did not differ in fuelling rate, but most females stayed at the non‐breeding site longer than the males, and thus accumulated more fuel and had longer potential flight ranges. These patterns of moult and fuelling suggest sex differences in preparations for breeding. We suggest that the males depart from South Africa earlier but with smaller fuel loads than the females to establish breeding territories before the females arrive. We conclude that for each sex the observed trade‐offs between fuelling and moult at the non‐breeding grounds are precursors to different migration strategies, which in turn are adaptations for their different roles in reproductive behaviour.     

A pre‐breeding immune challenge delays reproduction in the female dark‐eyed junco Junco hyemalis

Precise timing of life‐history transitions in predictably changing environments is hypothesized to aid in individual survival and reproductive success, by appropriately matching an animal's physiology and behavior with prevailing environmental conditions. Therefore, it is imperative for individuals to time energetically costly life‐history stages (i.e. reproduction) so they overlap with seasonal peaks in food abundance and quality. Female lifetime reproductive fitness is affected by several factors that influence energy balance, including arrival date, timing of egg production, and energetic condition. Therefore, any extra energetic costs during reproduction may negatively affect timing of egg production, and ultimately a female's fitness. For example, mounting an immunological response elicits a high energetic cost, and this transfer of resources towards cell and immune system maintenance could have direct negative effects on reproductive timing. In order to determine whether an immune challenge delays onset of breeding (i.e. egg production), we administered either a humoral immune challenge (keyhole limpet hemocyanin (KLH)) (treatment) or physiological saline (control) to free‐living female dark‐eyed juncos Junco hyemalis in the period immediately prior to egg‐laying (～4 weeks). We found that KLH‐injected females artificially delayed clutch initiation when compared to control females. These data help to refine our understanding of how free‐living birds allocate resources between reproduction and self‐maintenance processes during the critical pre‐laying period of the annual cycle.     

Individual repeatability in laying behaviour does not support the migratory carry‐over effect hypothesis of egg‐size dimorphism in Eudyptes penguins

Penguins of the genus Eudyptes are unique among birds in that their first‐laid A‐egg is 54–85% the mass of their second‐laid B‐egg. Although the degree of intra‐clutch egg‐size dimorphism varies greatly among the seven species of the genus, obligate brood reduction is typical of each, with most fledged chicks resulting from the larger B‐egg. Many authors have speculated upon why Eudyptes penguins have evolved and maintained a highly dimorphic 2‐egg clutch, and why it is the first‐laid egg that is so much smaller than the second, but only recently has a testable, proximate mechanism been proposed. In most species of Eudyptes penguins females appear to initiate egg‐formation at sea during return migration to breeding colonies. In macaroni penguins E. chrysolophus, females with a shorter pre‐laying interval ashore (and thus presumably greater overlap between migration and egg‐formation) lay more dimorphic eggs, suggesting a physiological conflict may constrain growth of the earlier‐initiated A‐egg. This migratory carry‐over effect hypothesis (MCEH) was tested in eastern rockhopper penguins E. chrysocome filholi on Campbell Island, New Zealand, by recording the arrival and lay dates, body sizes, and egg masses of transponder‐tagged females over two years. Females with longer pre‐laying intervals laid less dimorphic clutches, as predicted by the MCEH. However, repeated measures of individual females revealed that within‐individual variation in egg‐size dimorphism between years was unrelated to within‐individual variation in pre‐laying interval. Egg masses, and to a lesser extent egg‐size dimorphism, were highly repeatable traits related to body size and body mass. These results and a detailed consideration of the MCEH suggest that egg‐size dimorphism in Eudyptes penguins is unlikely to be caused by a migratory carry‐over effect.     

Does winter region affect spring arrival time and body mass of king eiders in northern Alaska?

Events during the non-breeding season may affect the body condition of migratory birds and influence performance during the following breeding season. Migratory birds nesting in the Arctic often rely on endogenous nutrients for reproductive efforts, and are thus potentially subject to such carry-over effects. We tested whether king eider (Somateria spectabilis) arrival time and body mass upon arrival at breeding grounds in northern Alaska were affected by their choice of a winter region in the Bering Sea. We captured birds shortly after arrival on breeding grounds in early June 2002–2006 at two sites in northern Alaska and determined the region in which individuals wintered using satellite telemetry or stable isotope ratios of head feathers. We used generalized linear models to assess whether winter region explained variation in arrival body mass among individuals by accounting for sex, site, annual variation, and the date a bird was captured. We found no support for our hypothesis that either arrival time or arrival body mass of king eiders differed among winter regions. We conclude that wintering in different regions in the Bering Sea is unlikely to have reproductive consequences for king eiders in our study areas.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Temperature‐correlated shifts in the timing of egg‐laying in House Finches Haemorhous mexicanus

Although temperature‐correlated shifts in the timing of egg‐laying have been documented in numerous bird species, the vast majority of species examined to date have been those that breed in Europe and have an animal‐based diet during breeding. However, given that the timing of breeding can be driven, either in the proximate or in the ultimate sense, by seasonal fluctuations in food availability, the relationship between temperature and laying may differ with diet. Here, we report on patterns of reproductive timing in House Finches Haemorhous mexicanus, a North American species that breeds on a primarily seed‐based diet. Analysing nest records from House Finches in California spanning more than a century, we found that egg‐laying occurred significantly earlier in warmer springs. We also found that although the timing of egg‐laying does not show long‐term changes in most of California, in the hottest region of the state (the southeast desert basin) it has advanced significantly.     

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.     

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.     

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.     

Protandrous migration and variation in morphological characters in Emberiza buntings at an East Asian stopover site

The effect of the timing of spring migration on reproductive success differs between the sexes. As a consequence, various sex‐specific tactics relating to the timing of migration have evolved in migratory avian groups. Various hypotheses have been proposed to explain differential migration to breeding or wintering grounds, and inter‐ and intrasexual size differences are often considered one of the proximate mechanisms. We investigated arrival patterns in the spring by individuals of each sex, sexual size dimorphism and related morphological variables, and the relationship between size variation and arrival date in five bunting species that passed through an East Asian migratory flyway stopover site in 2006–08. Males of all the study species arrived before females, and significant sexual dimorphism was observed. Several morphological characters, including total length, wing‐length and tail‐length, contributed to the size variation. Although larger males arrived earlier, there was no relationship between arrival date and size in females. Our study confirmed that East Asian buntings display a discriminated protandrous migration pattern at the stopover site as well as at the breeding grounds. This is consistent with the view that larger body size in males is favoured due to its association with early arrival to help ensure access to the best resources and hence enhanced mating success.