Arrival fat and reproductive performance in a long-distance passerine migrant

Long-distance passerine migrants deposit substantial fat stores to fuel their migratory journey. Many of those migratory birds arrive at their northerly breeding grounds with larger fat stores than were necessary to reach their breeding area. Both male and female American Redstarts ( Setophaga ruticilla) arrived to breed in Michigan's Upper Peninsula with fat, and females arrived with more fat than males in 2 out of 3 years. We test the hypothesis that migrants arriving at the breeding grounds with more body fat have higher reproductive success than birds arriving with little or no fat. Females, and to a lesser extent males, that arrive with fat experience gains in reproductive performance as evidenced by increased clutch size, egg volume, and nestling mass. The results have implications for understanding how events occurring during one phase of the annual cycle influence survival and/or reproductive performance in subsequent phases.     

Temporal patterns of arrival from migration as a response to competition for breeding space

Birds arriving from spring migration have higher chances of securing a territory if they reach the breeding area before their companions. However, an early arrival may compromise their chances of surviving until reproduction begins. Here, we study how evolutionarily stable responses to these opposite constraints may shape patterns of arrival in a population of migratory birds, with the help of a simple evolutionary algorithm. We show that, initially, an increasing level of competition for breeding territories generates a massive peak of early arrivals. However, when the level of competition becomes very high, the temporal distribution of arrivals may take different shapes, depending on the modalities of competition. If individuals trying to usurp territories have relatively high chances of success, a second peak of arrival emerges at the very end of the arrival period. Birds arrive then in two successive waves. By contrast, if these chances are low, a single early peak of arrival is still present. Many individuals do however continuously arrive afterwards, at a regular and increasing tempo, until reproduction begins. The prior residence effect, that rules competition between resident birds and their challengers, is thus expected to influence the whole distribution of arrival dates on breeding grounds. This distribution will, in turn, affect the ability of the population to respond to changing environmental conditions.     

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.     

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.     

Travel schedules to the high arctic: barnacle geese trade-off the timing of migration with accumulation of fat deposits

On their way from the wintering area to the breeding grounds in Spitsbergen, barnacle geese Branta leucopsis stage on islands off the coast of Norway. The aim of this study was to describe when the geese migrate in relation to the body stores deposited and explore questions related to the concept of optimal migration schedules and on the possible mechanisms involved. We estimated fat stores by repeated assessments of the abdominal profile index of individually marked females throughout staging. Reproductive success was derived from observations of the same individuals later in the annual cycle. Females arriving late, or with low fat stores at arrival, achieved higher fat deposition rates, probably by spending more time foraging. But they were unable to match final fat scores of birds that arrived earlier or with larger fat stores. Reproductive success was correlated with the timing of migration and individuals departing at intermediate dates achieved highest success. The exact date of peak reproductive success depended on the size of fat stores accumulated, such that low-quality birds (depositing less fat) benefited most from an early departure to the breeding grounds. Observations in the breeding colonies showed that these birds did not initiate a nest earlier but they spent a longer time in Spitsbergen before settling. The length of stay in Norway was close to the prediction derived from an optimisation model relating spring events to eventual breeding success. Poorest performing birds stayed longer than expected, perhaps depositing more fat to avoid the risk of starvation. Two possible mechanisms of the timing of migration were contrasted and it seemed that the geese departed for migration as soon as they were unable to accumulate any more fat stores.     

Social cues facilitate habitat selection: American redstarts establish breeding territories in response to song

For migratory songbirds nesting in northern temperate forests, a short breeding season demands that males rapidly establish territories. Because critical insect and vegetation resources are unavailable during spring arrival, we suggest that conspecifics serve as settlement cues for males new to a local population. To test conspecific attraction, we conducted playback experiments with American redstarts Setophaga ruticilla. Experimental results demonstrate that song playbacks strongly attract conspecifics, recruiting an average of 4.2 additional males per plot; adult males new to our sites increased, while yearling males failed to respond. Yearlings arrived 6 to 10 days later than adults, raising the possibility that yearlings responded to songs of early arriving adults rather than to playbacks. Our work indicates that conspecific attraction is an important mechanism for breeding habitat selection in an established population of a migratory forest songbird, but the effect is moderated by age, reproductive experience and arrival timing.     

Age- and sex-dependent spring arrival dates of Eastern Kingbirds

ABSTRACT.   Among migratory passerines, the first birds to arrive on the breeding grounds are usually older males. Early arrival by older birds may be driven by experience, age-dependent changes in body condition, age-dependent access to resources during the nonbreeding period, or latitudinal segregation by age. Males may arrive earlier than females (protandry) because males maintain better condition due to greater access to resources during the winter, or because selection favors early arriving males that acquire the best territories or experience enhanced mating opportunities. During a 4-yr study (2004–2007) in Oregon, we found that older Eastern Kingbirds ( Tyrannus tyrannus ), regardless of sex, arrived nearly a week before younger birds and that males arrived about 5 d before females. Age- and sex-dependent arrival dates do not appear to be related to differences in body condition, social dominance in winter, or latitudinal segregation, and protandry is unrelated to the ability of early-arriving males to acquire high-quality territories. Instead, we propose that young birds have less to gain from early arrival because of their probable inability to displace experienced birds from prime territories and that protandry evolved due to enhanced mating opportunities for early arriving males that arise from the high rates of extra-pair matings in our population of Eastern Kingbirds.     

Advancement of spring arrival in a long-term study of a passerine bird: sex,age and environmental effects

In migratory birds, mistimed arrival might have negative consequences for individual fitness, causing population declines. This may happen if arrival time is not synchronized with breeding time, especially when earlier springs favour earlier reproduction. We studied spring arrival time to the breeding areas in a pied flycatcher Ficedula hypoleuca population in southern Norway during a 30-year period (1985–2014). We investigated trends in arrival both for the entire population and for different population fractions (e.g. early vs. late arrivals). We also studied sex and age class differences, along with repeatability of arrival. Finally, we explored how arrival is influenced by environmental conditions at the areas birds use throughout the year, using mixed-effects models and quantile regressions with individual-based data. Spring arrival advanced over five days, at a similar rate through the entire population. Males and adult birds arrived earlier than females and yearlings. Arrival was significantly repeatable for males and females. Birds arrived earlier in years with high temperature and rainfall at the breeding grounds, and low NDVI both on the Iberian Peninsula and in central Europe. Later fractions of the population showed a steeper response to these environmental variables. This intra-population heterogeneity in the responses to the environment probably stems from a combination between the different selection pressures individuals are subject to and their age-related experience. Our results highlight the importance of studying how migration phenology is affected by the environment not only on the breeding grounds but also on the other areas birds use throughout the year.     

Arrival timing and hematological parameters in Gray Catbirds (Dumetella carolinensis)

Early arrival at the breeding grounds for migratory birds is associated with greater reproductive success. According to the condition-dependent arrival hypothesis, only those individuals in superior physiological condition are able to bear the costs (e.g., poor environmental conditions, limited food availability) of early arrival. Condition has usually been measured in terms of energy reserves or mass but other physiological measures of condition such as hematocrit and immune function have been gaining attention. We examined several measures of condition and their association with date of first capture in Gray Catbirds (Dumetella carolinensis) arriving at breeding grounds in northeastern Pennsylvania. Earlier arrivals had higher hematocrit and H/L ratios and lower lymphocyte counts. Arrival date was also negatively associated with fat score. Fat score was positively related to hematocrit, total number of leukocytes, and number of lymphocytes, but the other hematological parameters were not associated with traditional measures of condition (keel score, fat score, or a body condition index). Our results provide some support for the condition-dependent arrival hypothesis and suggest that there may be immunological differences between early- and late-arriving birds.     

Early arriving males wait longer for a mate than later arrivals: the case of a migratory monogamous passerine bird species

Individuals that arrive earlier on the breeding grounds may obtain many advantages but they also have to spend time waiting for a mate. I studied the waiting times of male red-breasted flycatchers Ficedula parva, a small, migratory, sexually dichromatic passerine bird species under natural conditions (Bia?owie?a National Park, Poland) in relation to year, arrival time, age of male and morphological parameters. The length of waiting time was dependent on males’ arrival time. The males which arrived later waited a shorter time for females than earlier arrivals. In some years older males spent more time waiting for mates than younger males, but in other years they waited for shorter times. A significant interaction between age of male and year was also observed. Despite the significantly earlier arrival of older males, the waiting time was not related to male age. The waiting time was also not related to body biometric parameters of the male. Despite waiting longer, early male red-breasted flycatchers have an advantage over later arrivals given this greater chance of mating.     

Body condition of shorebirds upon arrival at their Siberian breeding grounds

Arctic breeding shorebirds carry substantial body stores on their long-distance migrations from their non-breeding grounds. Upon arrival at the breeding area the remains of these stores can be used for egg formation, insurance against poor feeding conditions or rebuilding organs. We quantified body condition (body mass, total body water, lean body mass and fat mass estimated using the deuterium dilution method) in seven shorebird species caught upon arrival in the Siberian Arctic. Arrival condition was compared with incubation condition in a subset of species. After correction for structural size, body mass was significantly lower at arrival than during incubation in most of the species (but 3–18% above lean mass). Fat index (fat mass/lean mass) varied between 5.1 and 13.2%. Fat stores were estimated to enable survival for 0.6 days for the smallest and 2.5 days for the largest species. We discuss possible functions of arrival stores: insurance, egg-formation or rebuilding organs. G. H. Visser: Deceased.     

Photoperiodic response may facilitate adaptation to climatic change in long-distance migratory birds

Recent climatic change is causing spring events in northern temperate regions to occur earlier in the year. As a result, migratory birds returning from tropical wintering sites may arrive too late to take full advantage of the food resources on their breeding grounds. Under these conditions, selection will favour earlier spring arrival that could be achieved by overwintering closer to the breeding grounds. However, it is unknown how daylength conditions at higher latitudes will affect the timing of life cycle stages. Here, we show in three species of Palaearctic-African migratory songbirds that a shortening of migration distance induces an advancement of springtime activities. Birds exposed to daylengths simulating migration to and wintering in southern Europe considerably advanced their spring migratory activity and testicular development. This response to the novel photoperiodic environment will enable birds wintering further north to advance spring arrival and to start breeding earlier. Thus, phenotypic flexibility in response to the photoperiod may reinforce selection for shorter migration distance if spring temperatures continue to rise.     

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.     

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.     

Prevalence of hematozoa in overwintering American redstarts (Setophaga ruticilla): no evidence for local transmission

We examined American redstarts (Setophaga ruticilla) for protozoan blood parasites on their wintering grounds to determine whether transmission of these parasites occurs prior to spring migration. A total of 73 blood smears from 37 birds were examined for presence and intensity of infection. Thirty-six birds were sampled in the fall, soon after arriving from northern breeding grounds, and the spring prior to departure. Two (5%) of the samples collected in the fall were positive for Haemoproteus fringillae and one (3%) had detectable infections of Trypanosoma avium. Individuals infected with H. fringillae were hatching year redstarts sampled in September and October. Intensity of infection was 78 and < 1 infected erythrocytes per 10,000 erythrocytes, respectively. None of the birds had detectable infections when resampled prior to spring migration the following March.     

Divorce and asynchronous arrival in common terns, Sterna hirundo

We investigated which of three hypotheses (better option, incompatibility or asynchronous arrival) best explains divorce in the common tern. One partner did not return the next year in 18.5% of 150 pairs. Among the 106 pairs in which both mates returned, the divorce rate was 18.9%. We found no significant differences in: breeding performance or condition in relation to the probability of divorce; quality of previous mates and new mates, mean age in relation to pair bond status; breeding success before and after divorce nor did this differ from breeding success of reunited pairs. Hence the better option and incompatibility hypotheses were not supported. However, divorce was more likely in pairs in which mates arrived asynchronously on the breeding grounds, supporting the asynchronous arrival hypothesis. Median arrival asynchrony for divorced pairs was 7.5 days and for reunited pairs 2 days; mates arriving more than 16 days apart always split up. About 20% of divorced birds lost breeding status in the year of divorce, probably as a consequence of their late arrival. Our results suggest that terns search for a new mate as soon as they arrive on the breeding grounds and that mates remain faithful to each other to avoid the costs of searching for a new partner. Thus, synchrony in arrival facilitates pair bond maintenance rather than asynchrony promoting divorce, since divorce appears to be a side-effect of asynchrony and not an active decision. Copyright 1999 The Association for the Study of Animal Behaviour.     

Three-Phase Fuel Deposition in a Long-Distance Migrant,the Red Knot (Calidris canutus piersmai), before the Flight to High Arctic Breeding Grounds

Refuelling by migratory birds before take-off on long flights is generally considered a two-phase process, with protein accumulation preceding rapid fat deposition. The first phase expresses the demands for a large digestive system for nutrient storage after shrinkage during previous flights, the second phase the demands for fat stores to fuel the subsequent flight. At the last staging site in northward migration, this process may include expression of selection pressures both en route to and after arrival at the breeding grounds, which remains unascertained. Here we investigated changes in body composition during refuelling of High Arctic breeding red knots (Calidris canutus piersmai) in the northern Yellow Sea, before their flight to the tundra. These red knots followed a three-phase fuel deposition pattern, with protein being stored in the first and last phases, and fat being deposited mainly in the second phase. Thus, they did not shrink nutritional organs before take-off, and even showed hypertrophy of the nutritional organs. These suggest the build up of strategic protein stores before departure to cope with a protein shortage upon arrival on the breeding grounds. Further comparative studies are warranted to examine the degree to which the deposition of stores by migrant birds generally reflects a balance between concurrent and upcoming environmental selection pressures.     

Early arrival, initiation of nesting, and social status: an experimental study of breeding female red-winged blackbirds

Red-winged blackbirds (Agelaius phoeniceus) arrive at breedingsites several months before nests are constructed. Males inthis highly polygynous species presumably return early to defenddesirable territories. Females, however, also begin to arrivealmost two months before nesting is attempted. Early returnto breeding sites could enhance reproductive success by increasinga female's social status and thereby allowing earlier nesting.I measured the effect of experimentally delayed arrival on thetiming of nesting, reproductive success, and social status offemales. Birds were captured as they arrived in early springand detained in an aviary. These experimental females were laterreleased at their capture sites before control females beganconstruction of nests; controls had arrived during the sameperiod as experimentals but were not detained. Experimentalfemales nested, on average, more than a week later than controls,although I could detect no effect of timing on reproductivesuccess during this study. By manipulating arrival date, butnot covariates of arrival time such as age or experience, thisresult indicates that timing of arrival directly influencedlaying date. Delayed females were subordinate to control femaleswith the same mate. This decline in social status may have beenresponsible for the delay in nesting and could have fitnessconsequences for females in some years, because the offspringof earlier nesting, dominant birds are more likely to receivemale parental care in this population.     

Fat reserves of migrating passerines at arrival on the breeding grounds in Swedish Lapland

Mist-net capture data taken during 6 years (1988–1990 and 1992–1994) of field work were used to describe the arrival sequences and fat loads of nine species of migratory passerines which breed in a near-Arctic environment in Swedish Lapland. Long-distance migrants arrived with significantly larger mean fat reserves than did medium- and short-distance migrants. Long-distance migrants carried fat loads at arrival which corresponded to potential remaining flight distances between 242 and 500 km. When females and males arrived on the breeding grounds simultaneously, females carried significantly larger energy reserves than did males in seven out of nine species. In contrast, when the sexes showed a significant difference in median arrival date (two out of nine species), there was no difference in mean fat load carried into the breeding area. A relationship was found between the migratory habits and foraging ecology of each species and the amount of fat reserves at arrival, suggesting that species-specific migratory distances and feeding habits determine the amount of fat that is needed during the transition period between migration and onset of breeding. The short growing season in the study area restricts the time available for breeding and moult, and large energy reserves at arrival may speed up the breeding schedule to counteract possible time constraints. Overloading at the last stopover site during spring migration may be an adaptation allowing birds to cope with a restricted time frame for breeding and moult at high latitudes.     

Parasite assemblages distinguish populations of a migratory passerine on its breeding grounds

We attempted to establish migratory connectivity patterns for American redstarts Setophaga ruticilla between their breeding and wintering grounds by characterizing the composition of their haematozoan parasite assemblages in different parts of their range. We detected significant but limited geographic structuring of haematozoan parasite lineages across the breeding range of the redstart. We found that redstarts from the south-eastern (SE) region of the breeding range had a significantly different haematozoan parasite assemblage compared with populations sampled throughout the rest of the breeding range. Evidence using stable isotopes from feathers previously demonstrated that redstarts from the SE of the breeding range also have a unique and separate wintering range. Thus, although two methods of estimating migratory connectivity have now both shown the SE US breeding sub-population of redstarts to be distinct from other populations on both the breeding and wintering grounds, conclusive migratory connectivity for this species as a whole could not be established.