Geomagnetic information modulates nocturnal migratory restlessness but not fueling in a long distance migratory songbird

Geomagnetic cues have been shown to influence migratory orientation and migratory fuelling in night‐migratory songbird species. Here, we used captive‐bred northern wheatears Oenanthe oenanthe from the southern Norwegian population to show that other aspects of the birds’ migratory program can be influenced by magnetic cues as well. We observed that the amount of migratory restlessness increased strongly with progression of the migratory season when the birds were kept constantly in the magnetic field of northern Germany, but the amount of migratory restlessness decreased when the magnetic field changed along the birds’ natural flyway are simulated. Thus, the Earth's magnetic field can also act as a ‘signpost’ cue for fine‐tuning the spatio‐temporal course of migration.     

Wind conditions experienced during the day predict nocturnal restlessness in a migratory songbird

A variety of methods have been used to study the relationship between wind conditions and departure decisions of migrant birds at stopover sites. These methods are either costly or suffer from inaccuracy in determining whether or not an individual has resumed migration. Here we present a novel and simple approach to studying the relationship between wind conditions and departure likelihood. Northern Wheatears Oenanthe oenanthe caught during stopover were temporarily caged to measure their nocturnal migratory restlessness, which is an accurate proxy for their individual departure likelihood. We then related the degree of nocturnal restlessness to wind conditions prevailing at the time of capture. Confirming the general pattern from previous studies of departure, the intensity of nocturnal migratory restlessness, and hence departure likelihood, increased with increasing wind support towards the migratory goal. This suggests that approximating the propensity to depart by measuring nocturnal migratory restlessness is a reliable way to study the effect that wind conditions experienced during stopover has on the departure decision of migrants. Our study also shows that nocturnal migrants possess the ability to use information gathered during the day for their departure decisions at night. Because measuring migratory restlessness is straightforward and inexpensive, our approach is ideally suited to test hypotheses regarding spatio‐temporal variation in wind selectivity in migrating birds.     

Potential age differences in the migratory behaviour of a nocturnal songbird migrant during autumn and spring

In migratory songbirds, older individuals are thought to be more efficient migrants than younger individuals. Age‐specific differences in migratory efficiency have been reported mainly in respect of arrival timing, energy stores, rate of energy accumulation, departure behaviour, and departure direction. Yet, these traits were rarely assessed simultaneously in a single species. We further lack information whether age‐specific differences in behavioural traits present in autumn still manifest to the same degree in spring. Here we used the northern wheatear Oenanthe oenanthe, a long‐distance nocturnal songbird migrant, and investigated age‐specific differences in energy stores at capture (autumn: 1059 birds/spring: 803 birds), rate of energy accumulation (168/147), nocturnal departure timing (126/105), and departure direction (94/77) for both seasons. We found that in autumn, young northern wheatears departed significantly later in the night than older birds. This difference was not observed in spring. The resulting advance in nocturnal departure timing by young birds from autumn to spring may be due to learning based on experience gained during autumn, and/or to selective disappearance of those young individuals showing late departure times during autumn. We found no age‐specific difference in any of the other migratory traits investigated. To get a better understanding of age effects in songbird migrants, we need to study the potential adjustments of migratory behaviour within the individual over its life time. By individually tracking songbirds throughout their lifetime, we could start estimating whether a certain migratory decision (fuelling, departure, orientation) translates into higher (or lower) fitness, and whether individuals adjust their migratory behaviour based on learning from ‘wrong decisions’.     

A separate circadian oscillator controls nocturnal migratory restlessness in the songbird Sylvia borin

When confined to a cage, migratory songbirds exhibit nocturnal migratory restlessness (also called Zugunruhe) during the spring and autumn migratory periods, even though these birds are exclusively diurnal during the remainder of the year. Zugunruhe, which has been demonstrated to be under the direct control of a circannual timer, is characterized by a stereotypic "wing-whirring" behavior while the bird is perched. To elucidate the role played by the circadian system in the regulation of Zugunruhe, the authors studied the activity of garden warblers (Sylvia borin), long-distance nocturnal migrants, under skeleton photoperiods of different lengths and under constant dim light. In 11.5D:1L:10.5D:1L skeleton photoperiods, the authors found that Zugunruhe free-ran in a substantial proportion of birds, while their normal daily activities (e.g., feeding and preening) remained synchronized to 24 h. Some birds expressing Zugunruhe under constant dim light continued to show 2 distinct bouts of activity: one corresponding to daily activities, the other to wing-whirring. In some cases, these 2 bouts crossed while free-running with different periods. Birds expressing Zugunruhe also had significantly longer free-running periods than birds that did not. The study data suggest that the seasonal appearance of Zugunruhe is the result of the interactions of at least 2 circadian oscillators and that it is the phase relationship of these 2 oscillators that determines when nocturnal migratory restlessness is expressed. Furthermore, these data are consistent with the previously proposed internal coincidence hypothesis as a model for the ontogeny of circannual rhythms.     

Timing of breeding carries over to influence migratory departure in a songbird: an automated radiotracking study

1. Determining how events interact across stages of the annual cycle is critical for understanding the factors that affect individual fitness. However, there is currently little information detailing how breeding events influence migratory behaviour. 2. Using an automated digital telemetry array and an isolated island-breeding population of Savannah sparrows Passerculus sandwichensis, we provide the first direct evidence that the timing of breeding events carries over to influence the timing of migration in a songbird and assess for the first time how weather conditions on the breeding grounds also affect departure dates. 3. Date of migratory departure between September and October was strongly influenced by date of breeding completion in adults and fledging date in juveniles from June to July. 4. With respect to weather, adults departed during the first half of high-pressure systems, while juveniles departed throughout the entirety of high-pressure systems (including rainy evenings on the western edge of systems). 5. By combining both ecological and weather data, we could explain almost all variation in departure date for adults (95%), but weather conditions were not a good predictor of departure date for juveniles. 6. Our results provide strong evidence that the timing of breeding events is an important driver of migration timing and that exact departure dates are fine-tuned according to local weather conditions in adults, but not in juveniles.     

Rainfall and habitat interact to affect the condition of a wintering migratory songbird in The Bahamas

On the subtropical and tropical wintering grounds of migratory birds, variation in moisture levels and habitat can influence the availability of food resources and subsequently impact overwintering birds. Using stable carbon isotopes in blood samples as a measure of moisture, we assessed the interactive effects of rainfall, vegetation, and moisture on the demographics and condition of Prairie Warblers (Setophaga discolor) wintering in The Bahamas. Carbon isotopes in Prairie Warbler blood were more depleted in taller, wetter habitats; we additionally detected novel temporal effects of rainfall on isotope values. During a winter with more rainfall, most birds maintained mass and pectoral muscle regardless of the habitat type occupied. In a winter with less rainfall, birds lost mass and pectoral muscle, and this effect was more pronounced in birds with enriched isotope values and birds that occupied drier, shorter habitat. Prairie Warblers exhibited strong patterns of sexual habitat segregation with males disproportionately observed in areas with taller vegetation and females in shorter vegetation. During the drier winter, older males had better maintenance of pectoral muscle compared to females and younger individuals. Also in the drier winter, daily rainfall patterns explained more of the variation in body condition compared to the date of capture; pectoral muscle was best explained by recent precipitation (during the previous 30 days), while size‐corrected mass was more a function of longer‐term (90‐day) rainfall and habitat moisture. Our findings along with other studies suggest that Prairie Warblers and other migratory birds are sensitive to interactions between annual variation in winter rainfall, within‐season daily rainfall patterns, and habitat quality. Increasing drought and habitat loss in the Caribbean may be having a negative impact on wintering bird populations. To best conserve Nearctic–Neotropical migratory passerines in the region, we recommend prioritizing the protection of the least drought‐prone wintering areas.     

Winter body condition in relation to age,sex and plumage ornamentation in a migratory songbird

Winter body condition may play important roles in the life history of migratory birds, but it is difficult to estimate. We used the growth rate of winter‐grown tail feathers of Collared Flycatchers Ficedula albicollis as an indicator of winter body condition, comparing this trait between age classes and sexes and relating it to plumage ornamentation (forehead and wing patch sizes). Adults and males were in better nutritional condition during winter, as indicated by their faster tail feather growth rates, than were yearlings and females, respectively, which could indicate differences in individual quality and foraging ability with age, or age‐ and sex‐related winter habitat segregation. However, feather growth rate was related neither to the size of the winter‐grown forehead patch nor to the size of the summer‐grown wing patch, suggesting weak condition‐dependence for the winter‐grown ornament and complex life‐history consequences for the summer‐grown ornament.     

Continent-wide variation in feather colour of a migratory songbird in relation to body condition and moulting locality

Understanding the causes of variation in feather colour in free-living migratory birds has been challenging owing to our inability to track individuals during the moulting period when colours are acquired. Using stable-hydrogen isotopes to estimate moulting locality, we show that the carotenoid-based yellow-orange colour of American redstart (Setophaga ruticilla) tail feathers sampled on the wintering grounds in Central America and the Caribbean is related to the location where feathers were grown the previous season across North America. Males that moulted at southerly latitudes were more likely to grow yellowish feathers compared with males that moulted more orange-red feathers further north. Independent samples obtained on both the breeding and the wintering grounds showed that red chroma-an index of carotenoid content-was not related to the mean daily feather growth rate, suggesting that condition during moult did not influence feather colour. Thus, our results support the hypothesis that feather colour is influenced by ecological conditions at the locations where the birds moulted. We suggest that these colour signals may be influenced by geographical variation in diet related to the availability of carotenoids.     

Experimental dissociation of individual quality, food and timing of breeding effects on double-brooding in a migratory songbird

Annual reproductive success in many species is influenced by the number of breeding attempts within a season. Although previous studies have shown isolated effects of female quality, food, and timing of breeding on the probability of female birds producing second broods, to our knowledge, none have tested the relative importance of multiple factors and their interactions using simultaneous manipulations within populations of free-living birds. In this study, we show that individual quality and timing of breeding interact to affect the probability of double-brooding in female mountain bluebirds (Sialia currucoides). High-quality females (those that naturally initiated clutches early in the season) were more likely to double-brood, regardless of whether their hatching date was advanced or delayed, whereas later breeding, lower quality females were much less likely to double-brood when their first attempt was delayed. This indicates that annual fecundity of poorer quality (or younger) female bluebirds may be more sensitive to seasonal variation in environmental conditions. In addition, birds that were provided with supplemental food throughout first breeding attempts were more likely to double-brood in one of the study years, suggesting that female bluebirds may be energetically limited in their capacity to initiate a second brood. Females that had their first brood delayed also had a shorter inter-brood interval and were moulting fewer feathers during second broods compared to controls, while females in better condition showed more advanced moult in second breeding attempts. Taken together, our results demonstrate the combined effects of age- or individual quality-mediated energetic trade-offs between current and future reproduction, and between investments in offspring and self-maintenance, on annual fecundity of female birds.     

Associations between malaria and MHC genes in a migratory songbird

Malaria parasites are a widespread and species-rich group infecting many wild populations of mammals, birds and reptiles. Studies on humans have demonstrated that genetic factors play a key role in the susceptibility and outcome of malaria infections. Until the present study, it has not been examined whether genetic variation in hosts is important for the outcome of malaria infections in natural avian populations. We investigated associations between major histocompatibility complex (MHC) genes and prevalence of three different avian malaria parasites (Haemoproteus payevskyi (GRW1), Plasmodium sp. (GRW2) and Plasmodium sp. (GRW4)) in a long-term study of great reed warblers Acrocephalus arundinaceus. We hypothesized that the MHC genes could either give full protection against a malaria infection, or confer protection against lethal malaria and direct the infection towards being milder. We found a positive association between numbers of MHC class I alleles (a measure of level of heterozygosity) and prevalence of the GRW2 parasite, suggesting the latter scenario. There was also a positive association between a specific MHC allele (B4b), previously shown to be under frequency-dependent selection in the study population, and prevalence of GRW2. These associations suggest that individuals carrying either a large number of MHC alleles or a specific MHC allele are protected against lethal malaria infections.     

Effects of post-breeding moult and energetic condition on timing of songbird migration into the tropics

Each autumn billions of songbirds migrate between the temperate zone and tropics, but little is known about how events on the breeding grounds affect migration to the tropics. Here, we use light level geolocators to track the autumn migration of wood thrushes Hylocichla mustelina and test for the first time if late moult and poor physiological condition prior to migration delays arrival on the winter territory. Late nesting thrushes postponed feather moult, and birds with less advanced moult in August were significantly farther north on 10 October while en route to the tropics. Individuals in relatively poor energetic condition in August (high β-Hydroxybutyrate, low triglyceride, narrow feather growth bars) passed into the tropics significantly later in October. However, late moult and poor pre-migratory condition did not result in late arrival on the winter territory because stopover duration was highly variable late in migration. Although carry-over effects from the winter territory to spring migration may be strong in migratory songbirds, our study suggests that high reproductive effort late in the season does not impose time constraints that delay winter territory acquisition.     

Spring nocturnal migration of Reed Warblers Acrocephalus scirpaceus: departure, landing and body condition

Nocturnal migration of Reed Warblers Acrocephalus scirpaceus was studied by trapping with 'high nets' on the Courish Spit (Eastern Baltic) during spring 1998–2000. In spring, Reed Warblers left the stopover site between 45 and 240 min after sunset (median 84 min), although 85% of birds took off between 45 and 120 min after sunset. Birds did not arrive until the fifth hour after sunset; 67% of birds ended their nocturnal flights in the penultimate hour before sunrise, i.e. at dawn. At the moment of migratory departure, the average Reed Warbler body mass was 12.79 ± 0.66 g ( n  = 60). Average body mass of birds ending migratory flight was 11.69 ± 0.67 g ( n  = 18). The difference was highly significant. However, more than half of the birds completed migratory flights with a considerable fuel load, and some even had energy stores sufficient for a migratory flight on the next night. The spring migratory strategy of Reed Warblers over Central and Northern Europe probably includes a succession of short migratory flights (4–6 h) during several subsequent nights with 1-day stopovers.     

Climate and density influence annual survival and movement in a migratory songbird

Assessing the drivers of survival across the annual cycle is important for understanding when and how population limitation occurs in migratory animals. Density‐dependent population regulation can occur during breeding and nonbreeding periods, and large‐scale climate cycles can also affect survival throughout the annual cycle via their effects on local weather and vegetation productivity. Most studies of survival use mark–recapture techniques to estimate apparent survival, but true survival rates remain obscured due to unknown rates of permanent emigration. This is especially problematic when assessing annual survival of migratory birds, whose movement between breeding attempts, or breeding dispersal, can be substantial. We used a multistate approach to examine drivers of annual survival and one component of breeding dispersal (habitat‐specific movements) in a population of American redstarts (Setophaga ruticilla) over 11 years in two adjacent habitat types. Annual survival displayed a curvilinear relation to the Southern Oscillation Index, with lower survival during La Niña and El Niño conditions. Although redstart density had no impact on survival, habitat‐specific density influenced local movements between habitat types, with redstarts being less likely to disperse from their previous year's breeding habitat as density within that habitat increased. This finding was strongest in males and may be explained by conspecific attraction influencing settlement decisions. Survival was lowest in young males, but movement was highest in this group, indicating that apparent survival rates were likely biased low due to permanent emigration. Our findings demonstrate the utility of examining breeding dispersal in mark–recapture studies and complement recent work using spatially explicit models of dispersal probability to obtain greater accuracy in survival estimates.     

A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

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.     

Individual condition,but not fledging phenology,carries over to affect post-fledging survival in a Neotropical migratory songbird

Migratory animals face severe time and energy constraints during their annual cycle. These constraints may be exacerbated in young animals by conditions experienced during development that can affect both phenotype and phenology. For young migratory songbirds, the period between fledging and autumn migration, the post-fledging period, is believed to represent a time of intense selective pressure. However, there has yet to be a study that has assessed post-fledging survival for the entirety of the post-fledging period, probably due to the challenge of following juveniles as they move broadly across the landscape (tens to hundreds of kilometres). To overcome this challenge, we used an automated radiotelemetry array spanning 60 000 km² in southern Ontario, Canada, and miniature digital radiotelemetry tags to track 216 juvenile Barn Swallows Hirundo rustica continuously from fledging to migration. We hypothesized that young that fledged in better condition and earlier in the breeding season would have higher survival relative to birds fledging in poorer condition, because they have more energy to deal with resource constraints, and that early-fledging birds would depart on migration earlier than late-fledging birds because there is probably a fixed period of time required post-fledging to prepare for migration. We found that average cumulative apparent survival was 42% and that condition in the nest was a strong positive predictor of post-fledging apparent survival. We also found that birds that fledged earlier in the season departed on migration earlier in the autumn relative to late-fledging birds. Contrary to our prediction, average apparent survival was equal for early- and late-fledging birds. Our results suggest that factors during development that promote better nestling condition are critical for predicting future apparent survival prior to migration. Differences in annual apparent survival between early- and late-fledging songbirds, as commonly observed, may be driven by events occurring at later stages of the annual cycle.     

Quantifying non‐breeding season occupancy patterns and the timing and drivers of autumn migration for a migratory songbird using Doppler radar

For seasonal migrants, non‐breeding regions can play different roles in the ecology of their annual cycles: as stopover habitat, overwintering habitat, or as a combination in which some individuals stop‐over and others over‐winter. Such functional variations can lead to variation in occupancy dynamics and migration phenology to these different regions. In this study, we used data from archived Doppler weather surveillance radar to compare site‐occupancy and movement dynamics of a migratory songbird (tree swallow Tachycineta bicolor) between two non‐breeding areas: southeastern Louisiana and central peninsular Florida, USA. Specifically, in each area, we 1) quantified long‐term (1996–2012) non‐breeding season occupancy dynamics, 2) quantified variation in timing of autumn migration, and 3) tested which climate variables along their respective flyways were best correlated with variation in dates of arrival. Additionally, we cross‐validated the dynamics from archived radar with data from eBird, a large‐scale citizen science database that provides an independent measure of avian occupancy. We found strong and significant correlations between radar‐estimated and eBird‐estimated occupancy dynamics in both Louisiana and Florida. Long‐term Louisiana occupancy dynamics conformed to our hypothesis that this region acts as a combined stopover and overwintering region whereas Florida occupancy dynamics were akin to a traditional winter region. Arrival to Louisiana during the study period was much earlier and took place over a much shorter arrival window than did arrival to Florida, which showed much more gradual arrival over the course of several months. At both sites, annual variation in mean arrival date was best explained by the amount of precipitation along the lower portions of their respective migration flyways.