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.     

Spatial Isolation and Temporal Variation in Fitness and Condition Facilitate Divergence in a Migratory Divide

A novel migratory polymorphism evolved within the last 60 years in blackcaps (Sylvia atricapilla) breeding sympatrically in southwestern Germany. While most individuals winter in the traditional areas in the Mediterranean, a growing number of blackcaps started migrating to Britain instead. The rapid microevolution of this new strategy has been attributed to assortative mating and better physical condition of birds wintering in Britain. However, the isolating barriers as well as the physical condition of birds are not well known. In our study, we examined whether spatial isolation occurred among individuals with distinct migratory behaviour and birds with different arrival dates also differed in physical and genetic condition. We caught blackcaps in six consecutive years upon arrival on the breeding grounds and assigned them via stable isotope analysis to their wintering areas. Analysis of the vegetation structure within blackcap territories revealed different microhabitat preferences of birds migrating to distinct wintering areas. Blackcaps arriving early on the breeding grounds had higher survival rates, better body condition and higher multilocus heterozygosities than later arriving birds. We did however not find an effect of parasite infection status on arrival time. Our results suggest that early arriving birds have disproportionate effects on population dynamics. Allochrony and habitat isolation may thus act together to facilitate ongoing divergence in hybrid zones, and migratory divides in particular.     

Malaria infection and feather growth rate predict reproductive success in house martins

Carry-over effects take place when events occurring in one season influence individual performance in a subsequent season. Blood parasites (e.g. Plasmodium and Haemoproteus) have strong negative effects on the body condition of their hosts and could slow the rate of feather growth on the wintering grounds. In turn, these winter moult costs could reduce reproductive success in the following breeding season. In house martins Delichon urbica captured and studied at a breeding site in Europe, we used ptilochronology to measure growth rate of tail feathers moulted on the winter range in Africa, and assessed infection status of blood parasites transmitted on the wintering grounds. We found a negative association between haemosporidian parasite infection status and inferred growth rate of tail feathers. A low feather growth rate and blood parasite infections were related to a delay in laying date in their European breeding quarters. In addition, clutch size and the number of fledglings were negatively related to a delayed laying date and blood parasite infection. These results stress the importance of blood parasites and feather growth rate as potentially mechanisms driving carry-over effects to explain fitness differences in wild populations of migratory birds.     

Interactions between a Candidate Gene for Migration (ADCYAP1), Morphology and Sex Predict Spring Arrival in Blackcap Populations

Avian research has begun to reveal associations between candidate genes and migratory behaviors of captive birds, yet few studies utilize genotypic, morphometric, and phenological data from wild individuals. Previous studies have identified an association between ADCYAP1 polymorphism and autumn migratory behavior (restlessness, or zugunruhe), but little is known about the relationship between ADCYAP1 and spring migratory behavior. The timing of spring migration and arrival to the breeding ground are phenological traits which could be particularly favorable for establishing territories and acquiring mates, thus important to fitness and reproductive success. Here, we investigated how individual genotypic ADCYAP1 variation and phenotypic variation (wing length and shape) of blackcaps (Sylvia atricapilla) affect spring arrival date across nine natural populations in Europe. We hypothesized that longer alleles should be associated with earlier spring arrival dates and expected the effect on arrival date to be stronger for males as they arrive earlier. However, we found that longer wings were associated with earlier spring arrival to the breeding grounds for females, but not for males. Another female-specific effect indicated an interaction between ADCYAP1 allele size and wing pointedness on the response of spring arrival: greater allele size had a positive effect on spring arrival date for females with rounder wings, while a negative effect was apparent for females with more pointed wings. Also, female heterozygotes with pointed wing tips arrived significantly earlier than both homozygotes with pointed wings and heterozygotes with round wings. Stable isotope ratios (δ ² H) of a subset of blackcaps captured in Freiburg in 2011 allowed us also to assign individuals to their main overwintering areas in northwest (NW) and southwest (SW) Europe. NW males arrived significantly earlier to the Freiburg breeding site than both SW males and females in 2011. NW females had more pointed wing tips compared to SW females, but no difference in ADCYAP1 allele size was found between the different migration routes.     

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.     

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.     

Spring arrival along a migratory divide of sympatric blackcaps (Sylvia atricapilla)

The recent formation of a migratory divide in the blackcap (Sylvia atricapilla) involves sympatrically breeding birds migrating to different overwintering quarters. Within the last 50 years, a novel migratory strategy has evolved resulting in an increasing proportion of birds now wintering in Britain instead of migrating to the traditional sites in the Mediterranean area. This rapid microevolution has been attributed to allochronic spring arrival of migrants from the respective wintering quarters leading to assortative mating. Moreover, blackcaps wintering in Britain may experience fitness advantages owing to improved local wintering conditions. We used stable hydrogen isotope signatures (δD) to scrutinize the degree of temporal segregation of blackcaps upon spring arrival and to test for carry-over effects in body condition associated with the disparate wintering environments. Although we found that migrants from Britain arrive significantly earlier on German breeding grounds than migrants from the Mediterranean region, we also found a considerable overlap in arrival times. In a resampling model, the mean probability of assortative mating of birds wintering in Britain is ≤28% in both years. These results suggest that allochrony alone is not a strong isolating barrier between the two subpopulations. Migrants from both wintering locations did not differ in terms of body mass, mass-tarsus residuals or mass-tarsus ratio and arrived in a similar reproductive disposition. Thus, blackcaps wintering in Britain do not gain an apparent fitness advantage on spring migration due to carry-over effects in body condition. Future studies should explore additional factors such as differences in song quality and habitat that might contribute to the rapid microevolution of the blackcap.     

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

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

Ecophysiological aspects of rapid population growth in a novel migratory blackcap (Sylvia atricapilla) population: an experimental approach

Summary Blackcaps (Sylvia atricapilla) that breed in central Europe have usually migrated to Mediterranean or African wintering grounds. In the past several decades, a portion of this breeding population has started migrating to the British Isles to overwinter and this population has increased dramatically. Several factors, including higher annual survivorship (due to supplemental feeding and reduced migratiry distance), assortative mating, and enhanced reproductive success may be involved in this rapid population growth. As part of an intensive, long-term study of this population, we tested the hypothesis that the differences in photoperiod experienced by British-wintering versus Mediterranean-wintering blackcaps might lead to relatively early vernal (i.e., migratory and/or reproductive) physiological condition in members of the former group. We found that birds exposed to photoperiodic conditions that simulated migration to Britain to overwinter generally initiated vernal migratory activity earlier than birds held under conditions simulating migration to traditional wintering areas in central Spain. This difference, coupled with the shorter migratory distance to the British Isles, leads to significantly earlier estimated arrival dates for blackcaps that winter in Britain compared to central Spain. Bimodality in arrival times suggests that assortative mating on central European breeding grounds might occur between members of the different wintering populations. Males exposed to British-winter photoperiods showed significantly earlier testicular development than males kept under Spanish-winter photoperiods. Early arrival on the breeding grounds, coupled with accelerated reproductive condition, should lead to a relatively early reproductive effort, perhaps increasing average reproductive success. In general, these results support the hypothesis that differences in photoperiod on the wintering grounds may play an important role in the dynamic state of this population.     

Effects of territory competition and climate change on timing of arrival to breeding grounds: a game-theory approach

Phenology is an important part of life history that is gaining increased attention because of recent climate change. We use game theory to model phenological adaptation in migratory birds that compete for territories at their breeding grounds. We investigate how the evolutionarily stable strategy (ESS) for the timing of arrival is affected by changes in the onset of spring, the timing of the resource peak, and the season length. We compare the ESS mean arrival date with the environmental optimum, that is, the mean arrival date that maximizes fitness in the absence of competition. When competition is strong, the ESS mean arrival date responds less than the environmental optimum to shifts in the resource peak but more to changes in the onset of spring. Increased season length may not necessarily affect the environmental optimum but can still advance the ESS mean arrival date. Conversely, shifting a narrow resource distribution may change the environmental optimum without affecting the ESS mean arrival date. The ESS mean arrival date and the environmental optimum may even shift in different directions. Hence, treating phenology as an evolutionary game rather than an optimization problem fundamentally changes what we predict to be an adaptive response to environmental changes.     

A rare study from the wintering grounds provides insight into the costs of malaria infection for migratory birds

Malaria parasites can have strong effects on the population dynamics and evolution of migratory bird species. In many species, parasite transmission occurs on the wintering grounds, but studies to determine the consequences of infection have taken place during the breeding season, when malaria parasites circulate at chronic levels. We examined the predictors of malarial infections for great reed warblers during the northern winter in Africa, where active parasite transmission is thought to occur and naïve individuals experience acute infections. Counter to expectations, we found that winter infection intensities were lower than those encountered on the breeding grounds. One potential explanation is that reduced immune function during breeding allows parasites to persist at higher chronic intensities. We found no relationships between the incidence or intensity of infection on condition (as measured by scaled mass index, plasma metabolites, and feather corticosterone), spring migration departure dates, or home range sizes. We also tested a prediction of the Hamilton–Zuk hypothesis and found that male ornament (song) quality was unrelated to parasitic infection status. Overall, our results provide the first evidence that long‐distance migrants captured on their wintering grounds are in the chronic stage of infection, and suggest that winter studies may fare no better than breeding studies at determining the costs of acute malarial infection for great reed warblers.     

Circannual variation in blood parasitism in a sub‐Saharan migrant passerine bird,the garden warbler

Knowing the natural dynamics of pathogens in migratory birds is important, for example, to understand the factors that influence the transport of pathogens to and their transmission in new geographical areas, whereas the transmission of other pathogens might be restricted to a specific area. We studied haemosporidian blood parasites of the genera Plasmodium, Haemoproteus and Leucocytozoon in a migratory bird, the garden warbler Sylvia borin. Birds were sampled in spring, summer and early autumn at breeding grounds in Sweden, on migration at Capri, Italy and on arrival and departure from wintering staging areas in West Africa: mapping recoveries of garden warblers ringed in Fennoscandia and Capri showed that these sites are most probably on the migratory flyway of garden warblers breeding at Kvismaren. Overall, haemosporidian prevalence was 39%, involving 24 different parasite lineages. Prevalence varied significantly over the migratory cycle, with relatively high prevalence of blood parasites in the population on breeding grounds and at the onset of autumn migration, followed by marked declines in prevalence during migration both on spring and autumn passage. Importantly, we found that when examining circannual variation in the different lineages, significantly different prevalence profiles emerged both between and within genera. Our results suggest that differences in prevalence profiles are the result of either different parasite transmission strategies or coevolution between the host and the various parasite lineages. When separating parasites into common vs. rare lineages, we found that two peaks in the prevalence of rare parasites occur; on arrival at Swedish breeding grounds, and after the wintering period in Africa. Our results stress the importance of appropriate taxonomic resolution when examining host‐parasite interactions, as variation in prevalence both between and within parasite genera can show markedly different patterns.     

The migration of the great snipe Gallinago media: intriguing variations on a grand theme

The migration of the great snipe Gallinago media was previously poorly known. Three tracks in 2010 suggested a remarkable migratory behaviour including long and fast overland non‐stop flights. Here we present the migration pattern of Swedish male great snipes, based on 19 individuals tracked by light‐level geolocators in four different years. About half of the birds made stopover(s) in northern Europe in early autumn. They left the breeding area 15 d earlier than those which flew directly to sub‐Sahara, suggesting two distinct autumn migration strategies. The autumn trans‐Sahara flights were on average 5500 km long, lasted 64 h, and were flown at ground speeds of 25 m s^?1 (90 km h^?1). The arrival in the Sahel zone of west Africa coincided with the wet season there, and the birds stayed for on average three weeks. The birds arrived at their wintering grounds around the lower stretches of the Congo River in late September and stayed for seven months. In spring the great snipes made trans‐Sahara flights of similar length and speed as in autumn, but the remaining migration through eastern Europe was notably slow. All birds returned to the breeding grounds within one week around mid‐May. The annual cycle was characterized by relaxed temporal synchronization between individuals during the autumn–winter period, with maximum variation at the arrival in the wintering area. Synchronization increased in spring, with minimum time variation at arrival in the breeding area. This suggests that arrival date in the breeding area is under strong stabilizing selection, while there is room for more flexibility in autumn and arrival to the wintering area. The details of the fast non‐stop flights remain to be elucidated, but the identification of the main stopover and wintering areas is important for future conservation work on this red‐listed bird species.     

Blood parasites in migrating wood‐warblers (Parulidae): effects on refueling,energetic condition,and migration timing

Blood parasites can negatively affect energetic condition, arrival date, and reproductive performance of breeding birds, yet their impact on migrating birds is poorly understood. We quantified haematozoan infection and examined its relationship to migration timing, energetic condition, and refueling rate (plasma triglyceride concentration) for three species of wood‐warblers (Parulidae) migrating through northwestern Ohio, USA from 11 April–3 June of 2004 and 2005. Prevalence of infection for magnolia Dendroica magnolia (36.3%; n=383), yellow‐rumped D. coronata (58.3%; n=518), and yellow warblers D. petechia (12.7%; n=205) was approximately two times higher than reported in other migration studies and similar to infection rates found in breeding areas. Probability and intensity of infection were higher for later migrating yellow‐rumped warblers, especially in younger (second‐year) birds. Furthermore, we found a negative trend between intensity of infection and energetic condition for second‐year yellow‐rumped warblers. Haematozoan infection was not associated with refueling rate for any species. Our results indicate that for some migrants, especially younger birds, probability and intensity of haematozoan infection are related to later migration timing and reduced energetic condition during spring migration, and thus may be important determinants of subsequent reproductive performance in breeding areas.     

Connecting the dots: Stopover strategies of an intercontinental migratory songbird in the context of the annual cycle

The phases of the annual cycle for migratory species are inextricably linked. Yet, less than five percent of ecological studies examine seasonal interactions. In this study, we utilized stable hydrogen isotopes to geographically link individual black‐and‐white warblers (Mniotilta varia) captured during spring migration with breeding destinations to understand a migrant's stopover strategy in the context of other phases of the annual cycle. We found that stopover strategy is not only a function of a bird's current energetic state, but also the distance remaining to breeding destination and a bird's time‐schedule, which has previously been linked to habitat conditions experienced in the preceding phase of the annual cycle. Birds in close proximity to their breeding destination accumulate additional energy reserves prior to arrival on the breeding grounds, as reflected by higher migratory condition upon arrival, higher refueling rates measured via blood plasma metabolites, and longer stopover durations compared to birds migrating to breeding destinations farther from the stopover site. However, late birds near their breeding destination were more likely to depart on the day of arrival (i.e., transients), and among birds that stopped over at the site, the average duration of stopover was almost half the time of early conspecifics, suggesting late birds are trying to catch‐up with the overall time‐schedule of migration for optimal arrival time on the breeding grounds. In contrast, birds with long distances remaining to breeding destinations were more likely to depart on the day of arrival and primarily used stopover to rest before quickly resuming migration, adopting similar strategies regardless of a bird's time‐schedule. Our study demonstrates that migrants adjust their en route strategies in relation to their time‐schedule and distance remaining to their breeding destination, highlighting that strategies of migration should be examined in the context of other phases of the annual cycle.     

Patterns of Haemoproteus beckeri parasitism in the gray catbird (Dumatella carolinensis) during the breeding season

We determined the prevalence and intensity of blood parasites in breeding gray catbirds (Dumatella carolinensis) at Killbuck Wildlife Area in Wayne and Holmes Counties, Ohio (USA) from June through August 2000. Of 98 catbirds sampled, 40 (40.8%) had detectable infections of Haemoproteus beckeri. Overall prevalence of H. beckeri in this population is high relative to that reported in earlier blood parasite surveys of both breeding and migrant catbirds. Mean intensity of H. beckeri infection did not vary significantly between young and old birds or among sampling periods. We found no effect of age on prevalence or intensity of H. beckeri infection. Older birds were not more likely to be infected than younger birds, despite longer exposure to arthropod vectors. Prevalence varied significantly with season and was highest in June and lowest in August. This pattern also was observed in older birds sampled repeatedly. This seasonal variation may reflect both newly acquired infections and chronic infections relapsing in response to hormonal changes associated with breeding. Evidence of transmission was observed in the single hatching year bird that lacked detectable infection in early summer, but demonstrated a very high intensity infection in late summer. These observations provide supportive evidence that hematozoa infections are acquired on the breeding grounds during the first year of life and relapse during the breeding season in subsequent years.     

Presence of Plasmodium and Haemoproteus in breeding prothonotary warblers (Protonotaria citrea: Parulidae): temporal and spatial trends in infection prevalence

Prothonotary warbler (Protonotaria citrea) has shown a long-term decline in abundance in the United States. As a long-range migrant, these warblers are exposed to parasites in both tropical and temperate regions. The focus of this study was to use molecular techniques to examine the temporal prevalence patterns of heamosopridian parasites Plasmodium and Haemoproteus in breeding prothonotary warblers. The prevalence (presence or absence) of Plasmodium and Haemoproteus species was assayed using primer sets for the cytochrome b gene of the mitochondrial DNA. Blood samples were obtained from 187 adult prothonotary warblers collected at 3 central Virginia, U.S.A., breeding sites. The relationship between haemosporidian parasite infections and reproductive success also was examined. We found that 71% of captured prothonotary warblers were infected with haemosporidian parasites, specifically, with 36% prevalence for Haemoproteus spp. and 44% prevalence for Plasmodium spp., during the 2008 breeding season; for both parasites, prevalence increased throughout the season. We found significant variation in haemosporidian parasite prevalence across the breeding season that was strongly site specific. Conversely, we found no significant effects of haemosporidian parasite infections on the reproductive success of prothonotary warblers. This is in sharp contrast to recent reports suggesting considerable effects of these parasites on the reproductive success of wild birds.     

Body composition of north and southbound migratory blackcaps is influenced by the lay‐of‐the‐land ahead

The annual migration of small birds depends on the optimal management of time and energy. Since refueling at stopovers between flights consumes most of the birds’ time and energy, selection of food‐rich sites, and timely departure therefrom are likely crucial to success. We examined this concept quantifying body composition of 200 migrating blackcaps, Sylvia atricapilla, in Eilat, Israel, using dual‐energy x‐ray absorptiometry and generated a model to predict body composition as it changes with body mass (m_b). We then back‐calculated body composition of > 20 000 blackcaps ringed between 1984 and 2005, and tested the hypothesis that the amount of fuel that a bird stores determines the length of its stopover. We predicted that 1) if time‐constrained in spring, birds at the stopover site carry less than a maximum fuel load, but 2) if not time‐constrained, as in autumn, their fuel load is much higher than in spring. We found the change in body composition of blackcaps to be biphasic and correlated with increasing m_b. At m_b < ? 17.8 g, increasing m_b is due to increasing lean mass (m_l), while at m_b > ? 17.8 g increasing m_b results from increasing fat mass (m_f), which is accompanied by decreasing m_l. Body composition of blackcaps at a spring stopover site indicates that blackcaps leave stopovers as soon as they regain functionality of their digestive systems, but before laying down much m_f. In autumn blackcaps arrive with fuel stores much larger than in spring. For these birds, the Eilat stopover apparently serves to complete fat accumulation before crossing the deserts ahead. We conclude that in spring, the decision to depart is not determined by the bird's fuel stores, especially when early arrival at the breeding site, and therefore time, is of the essence. In autumn, accumulating enough fuel to ensure successful crossing of the deserts ahead probably dictates stopover time.     

Survival of the fattest: body stores on migration and survival in red knots Calidris canutus islandica

Severe summer weather in Greenland and Arctic Canada in 1972 and 1974 caused very poor breeding success and elevated adult mortality in red knots Calidris canutus islandica. We show that those individual knots that are known to have survived these summers were in better than average nutritional condition shortly before departure from their late spring staging area in west Iceland. The condition index of previously banded or subsequently recovered birds captured in Iceland was positively related to the number of summers they were known to have survived. Body stores carried from the last spring staging area to the breeding grounds appear to offer Arctic-breeding shorebirds significant selective advantages: they are used for physical transformation from migration to breeding condition, and in years when weather is difficult may enable survival after arrival on the breeding grounds.