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.     

Fat stores of American redstarts Setophaga ruticilla arriving at northerly breeding grounds

Many long-distance passerine migrants arrive with more fat stores than necessary to have reached their northerly breeding grounds. Researchers have argued for adaptive advantages associated with arriving with 'surplus' migratory fat, including increased breeding performance and insurance against adverse weather, and reduced food resources during the days following arrival. The objectives of this work were to document fat stores in American redstarts Setophaga ruticilla arriving to breed in northern Michigan and to test predictions associated with the hypothesis that arrival fat serves an insurance role. Results suggest that redstarts arrived in northern Michigan with fat stores sufficient to have continued migrating an additional distance greater than 1000 km. Significant yearly variation in arrival fat corresponded in part with environmental conditions measured at the breeding grounds. Birds arrived with the most fat in the year with the coolest temperatures and the lowest food abundance at the time of arrival to the site. Further, an inverse relationship between arrival fat and arrival day in males indicated that early arrivals carried more fat than later arrivals. Birds that arrived early faced cooler temperatures and lower densities of terrestrial invertebrates, and arrival fat may have provided a mechanism to overcome poor early season foraging conditions. However, our results are not entirely consistent with the hypothesis that arrival fat serves only as insurance. Arrival fat appeared important even during the most benign year of this study. Further, evidence suggests that females may have derived more benefit from arrival fat than males. These results highlight the connection between phases of a bird's annual cycle. Migrants that do well en route , arrive at breeding grounds in better condition, which may contribute to survival and reproductive success.     

Timing of arrival at breeding grounds determines spatial patterns of productivity within the population of white stork (Ciconia ciconia)

Early arrival at breeding grounds have important fitness consequences for migratory birds, both at individual and population level. The aim of this study was to investigate how the timing of arrival at the breeding territories affects the spatial patterns of reproductive success within a population of white storks (Ciconia ciconia). Data were gathered annually for ca. 200 pairs of storks breeding in central Poland between 1994 and 2011. Geostatistical analysis of data indicated that in years of delayed arrival of the population (measured by the first quartile arrival date), the reproductive output of storks was negatively autocorrelated, which indicated that there was a tendency for pairs of high breeding success to neighbour with pairs of low success. By contrast, in years when first storks returned in early dates to the breeding grounds, their reproductive success did not show any kind of spatial autocorrelation. These results suggest that delayed return of the first-arriving storks of the population may increase intensity of intra-specific competition to the level at which high-quality breeding pairs monopolize most of available resources at the expense of neighbouring low-quality pairs, which have lower reproductive success as a consequence. Such hypothesis was further supported with the analysis of nesting densities, showing that the late-arriving breeding pairs incurred greater fitness costs (or derived lower fitness benefits) while breeding in high densities comparatively to the early-arriving conspecifics.     

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.     

Environmental conditions at arrival to the wintering grounds and during spring migration affect population dynamics of barn swallows Hirundo rustica breeding in Northern Italy

Several populations of long-distance migratory birds are currently suffering steep demographic declines. The identification of the causes of such declines is difficult because population changes may be driven by events occurring in distant geographical areas during different phases of the annual life-cycle of migrants. Furthermore, wintering areas and migration routes of populations of small-sized species are still largely unknown, with few exceptions. In this paper we identified the critical phases of the annual life-cycle that most influence the population dynamics of a small passerine, the Barn Swallow Hirundo rustica. We used information on temporal dynamics of a population breeding in Northern Italy, whose wintering range and timing of migration have been recently described by miniaturised tracking dataloggers. Our results indicated that primary productivity in the wintering grounds in the month when most individuals arrive from autumn migration and primary productivity in an area that is probably a stopover site during spring migration, influenced population dynamics more than habitat conditions at the breeding grounds. By using annual variation in primary productivity at the wintering grounds and stopover sites as predictors, we replicated the observed interannual population changes with great accuracy. However, the steep decline recently suffered by the population could be replicated only by including a constant annual decline in the model, suggesting that changes in primary productivity only predicted the interannual variation around the long-term trend. Our study therefore suggests the existence of critical periods during wintering and migration that may have large impact on population fluctuations of migrant birds.     

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

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

Insect-related differences in growth of birch and pine at northern treeline in Swedish Lapland

The tree-ring patterns of birch and pine in the Torneträsk area of Swedish Lapland were compared. In order to dendrochronologically identify years of major insect attacks on birch, pine being unaffected by these outbreaks, served as control. The sharp growth increase in birch following the historically confirmed, heavy outbreak in 1954–55 is discussed and evidence pointing to possible earlier outbreaks is presented.     

Conservation of migrants on their breeding grounds

This paper addresses the question of how to diagnose the causes of a population decline of a migratory bird so that conservation action can be taken. Diagnosis may be through study of habitats or population factors. Different approaches are illustrated with examples from the more threatened migratory species which breed in Europe. Gaps in knowledge of how to make conservation diagnoses and of the circumstances of some of Europe's few migratory and globally endangered birds are shown.     

Polygyny in the Willow Warbler Phylloscopus trochilus in Swedish Lapland

The breeding sex ratio of Willow Warblers Phylloscopus trochilus in a subalpine birch forest was strongly female biased. Very few territorial males remained unmated and most became polygynous. Bigyny was the rule, but some males probably had three females. Male assistance at secondary nests varied and was probably dependent on temporal distance between the females' nests. There was no significant reduction in reproductive output for these females. Variation in the sex ratio of breeding birds can be explained, at least partly, by variation in the timing of male settlement.     

Natural selection for earlier male arrival to breeding grounds through direct and indirect effects in a migratory songbird

For migratory birds, the earlier arrival of males to breeding grounds is often expected to have fitness benefits. However, the selection differential on male arrival time has rarely been decomposed into the direct effect of male arrival and potential indirect effects through female traits. We measured the directional selection differential on male arrival time in the pied flycatcher (Ficedula hypoleuca) using data from 6 years and annual number of fledglings as the fitness proxy. Using structural equation modeling, we were able to take into account the temporal structure of the breeding cycle and the hierarchy between the examined traits. We found directional selection differentials for earlier male arrival date and earlier female laying date, as well as strong selection differential for larger clutch size. These selection differentials were due to direct selection only as indirect selection for these traits was nonsignificant. When decomposing the direct selection for earlier male arrival into direct and indirect effects, we discovered that it was almost exclusively due to the direct effect of male arrival date on fitness and not due to its indirect effects via female traits. In other words, we showed for the first time that there is a direct effect of male arrival date on fitness while accounting for those effects that are mediated by effects of the social partner. Our study thus indicates that natural selection directly favored earlier male arrival in this flycatcher population.     

Prevalence and intensity of excretion of coccidial oocysts in migrating passerines on Helgoland

Prevalence and intensity of excretion of coccidial oocysts in actively migrating passerines (Turdus philomelos, Fringilla coelebs, Sylvia borin, Phoenicurus phoenicurus) were assessed in spring and autumn 2001 during stopovers on Helgoland. All examined species showed low prevalences of oocysts in spring (28–33%) and high prevalences in autumn migration (66–92%). In autumn, there are significant species-specific differences in prevalence and, partially, in intensity of excretion. Ground-feeding Chaffinches most commonly and most intensively shed oocysts (maximal 2,7×10⁶ oocysts per g fresh faeces) indicating that feeding style may influence the parasite load. No significant correlation between parasite load, age, and body condition (fat depots, breast muscle shape, body mass) was found. The connection between feeding style, age, migration, immunocompetence, and survival of passerine hosts on one hand and the particularities of coccidial pathogenicity on the other are discussed.     

Diurnal variability in the detection of waders on their breeding grounds

When censusing breeding waders in large upland areas, the time of day has a major influence on the numbers of birds detected.     

Lipid and water depletion in migrating passerines following passage over the Gulf of Mexico

Lipid depletion is currently believed to be the primary factor limiting flight duration of migrating birds in North America, while the influence of water loss is thought to be small. Three migrating species of passerines, wood thrush (Hylocichla mustelina), Swainson's thrush (Catharus ustulatus), and summer tanager (Piranga rubra) were captured during the 1993 spring migration just after crossing the Gulf of Mexico and examined for lipid and water depletion. The redwinged blackbird (Agelaius phoeniceus), a winter resident, was examined for comparison. Although some migrants had low levels of fat, most were not seriously fat-depleted and had much higher lipid levels than red-winged blackbirds. Migrants appeared dehydrated, usually having less than 60% body water and significantly less water than the blackbirds. Recaptured wood thrushes had significantly higher mass than when initially captured. Many of these birds were replenishing significant amounts of water, while the acquisition of lipid mass was dependent on the amount of time the birds spent on the study site. Migrants were significantly fatter but had significantly less water mass when captured during unfavorable weather than under favorable conditions. It appears that both dehydration and lipid depletion are major physiological constraints on migrating passerines.     

Evidence for breeding of Megadyptes penguins in the North Island at the time of human arrival

The arrival of humans in New Zealand around 750 years ago resulted in widespread faunal extinctions including the endemic Waitaha penguin (Megadyptes waitaha). Previously thought to have only bred on coastal South Island and Stewart Island, recent genetic reanalysis of prehistoric large penguin bones from the lower North Island indicates that the Waitaha penguin may have been a common resident. Here we synthesise previous studies and present new palaeontological and archaeological evidence to suggest that the Waitaha penguin was probably breeding in the lower North Island at the time of human arrival, and did not represent vagrant individuals from more southerly breeding colonies. The elimination of breeding Megadyptes from the North Island would add to the already significant avifaunal losses from New Zealand, of which the North Island suffered the greatest biodiversity loss after the arrival of humans.     

Individual migration patterns of Eurasian golden plovers Pluvialis apricaria breeding in Swedish Lapland; examples of cold spell‐induced winter movements

Tracking studies normally focus on long‐distance migrants, meaning that our understanding about short‐distance migration remains limited. In this study, we present the first individual tracks of the Eurasian golden plover Pluvialis apricaria, a short‐distance migrant, which were tracked from a Scandinavian breeding population using geolocators. In addition, golden plovers are known for their cold spell‐induced winter movements, and this study provides some first individual tracking data on this type of movements. In three cases the plovers spent the winter in NW Europe and in four cases they departed during winter from NW Europe to spend the rest of the winter in Iberia or Morocco (one bird that was tracked during two subsequent migration cycles moved to Iberia in the first winter but remained in NW Europe during the second winter). The four winter departures were associated with a cold spell in NW Europe during which maximum temperatures dropped to freezing. Cold spell‐induced winter movements were notably long and fast. The birds that remained at their NW European wintering site did not experience such cold spell. However, the plovers did not always move in response to freezing temperatures, as demonstrated by the individual that was tracked for a second season, when it experienced four cold spells at its wintering site in NW France without leaving. Little information was obtained about spring migration, but one bird had a prominent counter‐clockwise loop migration pattern through E Europe. Due to their cold spell winter movements, golden plovers exhibit great flexibility in migration patterns, resulting in a notably large spread in final wintering areas.     

Fuel reserves affect migratory orientation of thrushes and sparrows both before and after crossing an ecological barrier near their breeding grounds

Fat reserves influence the orientation of migrating songbirds at ecological barriers, such as expansive water crossings. Upon encountering a body of water, fat migrants usually cross the barrier exhibiting 'forward' migration in a seasonally appropriate direction. In contrast, lean birds often exhibit temporary 'reverse' orientation away from the water, possibly to lead them to suitable habitats for refueling. Most examples of reverse orientation are restricted to autumn migration and, in North America, are largely limited to transcontinental migrants prior to crossing the Gulf of Mexico. Little is known about the orientation of lean birds after crossing an ecological barrier or on the way to their breeding grounds. We examined the effect of fat stores on migratory orientation of both long- and short-distance migrants before and after a water crossing near their breeding grounds; Catharus thrushes (Swainson's and gray-cheeked thrushes, C. ustulatus and C. minimus ) and white-throated sparrows Zonotrichia albicollis were tested for orientation at the south shore of Lake Ontario during spring and autumn. During both spring and autumn, fat birds oriented in a seasonally appropriate, forward direction. Lean thrushes showed a tendency for reverse orientation upon encountering water in the spring and axial, shoreline orientation after crossing water in the autumn. Lean sparrows were not consistently oriented in any direction during either season. The responses of lean birds may be attributable to their stopover ecology and seasonally-dependent habitat quality.