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.     

How many juvenile Teal Anas crecca reach the wintering grounds? Flyway-scale survival rate inferred from wing age-ratios

Autumn postnuptial migration is critical in the dabbling duck annual cycle, when first-year birds in particular suffer high losses to natural and hunting mortality. Mortality rates in this age-class are generally unknown in Europe where winter ringing predominates. We used data from large-scale wing collections from hunters in Finland, Denmark and France to test the prediction that juvenile proportions among killed Teal (Anas crecca) would decline with distance along the flyway. As expected, this proportion decreased from 89% in Northern Finland to 58% in Western France. Potential biases linked with age determination from the wings, differential migration of age-classes, relative susceptibility to different forms hunting and gradual improvement of juvenile survival as they learn to avoid hunters could not explain the observed decline of juveniles in the shot population. This pattern was therefore considered to be genuine, the result of the cumulative depletion of first-years along the flyway, likely through hunting. On this assumption, combined with known adult monthly survival rates during August–November (94.2%), monthly juvenile survival rate was estimated at 52.8%, i.e. 14.7% (range 13.9–15.4% based on extreme values of adult survival) amongst Scandinavian juveniles reaching wintering quarters in Western France. Despite lack of precision in such estimates based on relative proportions, there is little doubt about the magnitude of autumn juvenile mortality and its consequences for the population dynamics of Teal. Lack of correlations between annual proportions of juveniles in the hunting bag and an index of Teal breeding success in Finland may result from such high and variable inter-annual mortality.     

Rapid 'evolution' of migratory behaviour in the introduced house finch of eastern North America

The house finch (Carpodacus mexicanus) of eastern North America was introduced onto Long Island, New York, around 1940. The source is presumed to be southern California, where ca. 80% of individuals are completely sedentary. The eastern population has become migratory: by the early 1960s, 36% of eastern house finches were performing migratory movements (more than 80 km from their banding site) and that proportion has fluctuated between 28% and 54% in succeeding years. The movements of birds banded during the breeding season and recovered in winter were strongly orientated toward the south-west, and the same pattern was evident in the earliest recoveries (1958 to 1966); recoveries of birds banded during winter and recovered in the breeding season were orientated toward the north-east. The average distance of migration has continued to increase logarithmically. Areas colonized later, as the range expanded, were characterized by initial long migration distances and high proportions of migrants, suggesting that these traits have evolved in the eastern population. Eastern house finches are partial migrants: not all individuals migrate, and birds that migrate some winters remain in breeding areas in others. Younger birds exhibit a stronger tendency to migrate. A very few western (including southern California) house finches moved long distances, but they did so in directions consistent with seasonal migration, indicating that the machinery subserving migratory behaviour pre-existed in the parent population.     

Seasonal climatic niche and migration movements of Double‐crested Cormorants

Avian migrants are challenged by seasonal adverse climatic conditions and energetic costs of long‐distance flying. Migratory birds may track or switch seasonal climatic niche between the breeding and non‐breeding grounds. Satellite tracking enables avian ecologists to investigate seasonal climatic niche and circannual movement patterns of migratory birds. The Double‐crested Cormorant (Nannopterum auritum, hereafter cormorant) wintering in the Gulf of Mexico (GOM) migrates to the Northern Great Plains and Great Lakes and is of economic importance because of its impacts on aquaculture. We tested the climatic niche switching hypothesis that cormorants would switch climatic niche between summer and winter because of substantial differences in climate between the non‐breeding grounds in the subtropical region and breeding grounds in the northern temperate region. The ordination analysis of climatic niche overlap indicated that cormorants had separate seasonal climatic niche consisting of seasonal mean monthly minimum and maximum temperature, seasonal mean monthly precipitation, and seasonal mean wind speed. Despite non‐overlapping summer and winter climatic niches, cormorants appeared to be subjected to similar wind speed between winter and summer habitats and were consistent with similar hourly flying speed between winter and summer. Therefore, substantial differences in temperature and precipitation may lead to the climatic niche switching of fish‐eating cormorants, a dietary specialist, between the breeding and non‐breeding grounds.     

Importance of artificial stopover sites through avian migration flyways: a landfill‐based assessment with the White Stork Ciconia ciconia

Open landfills seem to be playing an increasing role as target feeding areas for several species, not only in their breeding areas or during the winter, but also during the migration period. Evaluating the extent to which landfill sites are used by migrants is crucial to understanding their role in driving stopover decisions during migration, and in the potential health risks linked to feeding on refuse. The aim of this study was to evaluate the role of two open landfills located just before (France) and after (Spain) the East‐Atlantic flyway enters Iberia through the western Pyrenees as potentially important stopover sites for the White Stork populations moving along this route. Overall, we detected that these sites were used by storks that had been ringed from many western European breeding populations, mainly during the migration period, but also in winter. The mean distance between the stork breeding/ringing origin and the landfill sites increased from summer to winter, suggesting that storks breeding further away pass through Iberia later in the season, reflecting population‐specific timing of migration. During the autumn migration period (August–September), the first encountered landfill in France was estimated to be used by c. 1200 storks, and the other in Spain by 4000 storks. Our study hence contributes to a better understanding of the current and potentially hazardous role played by landfill sites in White Stork ecology, which is essential in order to provide management recommendations, and to evaluate the consequences of proposed open landfill closures in Europe.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

Diffuse, short and slow migration among Blue Tits

The knowledge of migration systems in long-distance regular migrants is in many cases extensive. Our understanding of the migratory characteristics of partial migrants, on the other hand, is far more rudimentary. We investigated migratory characteristics of partially migratory Blue Tits Cyanistes caeruleus using ringing recoveries of Swedish birds, to answer questions about geographic migration patterns, age-specific migrations, migration speeds and synchrony of movements. Median migration distance of Swedish Blue Tits was 82 km, with a main autumn direction in the sector between S and W (large directional scatter). Northerly and southerly populations did not differ in migration directions or distances, suggesting chain migration to be the general pattern. A larger proportion of adult Blue Tits remained near the breeding grounds during winter than was the case for juveniles. Some of the migrating birds (17%) seemed not to return in spring but stayed to breed closer to the winter area. Swedish Blue Tits show an exceptionally slow migration speed (median 13 km/day), among the slowest speeds recorded for any migrant bird. The Blue Tit represents an extreme case of diffuse, short and slow bird migration.     

Demographic and spatial responses of resident bird populations to the arrival of migratory birds within an urban environment

Long-distance migration allows many bird species to overcome the severe climatic changes that occur in seasonal environments. Migration is highly demanding, and given its cyclical nature, we currently know that it has substantial effects on the population parameters of migratory birds during both breeding and wintering seasons. However, the potential effects of the presence of migratory birds in their wintering grounds on populations of resident birds have remain largely unexplored. Here, we propose the hypothesis that migratory birds negatively affect the habitat occupancy and population abundance of resident birds because of the arrival of numerous individuals during the most limiting months of the year. Here, we studied different species of migratory and resident birds that coexist during winter in an urban ecological reserve located within Mexico City. We used single-species multiseason occupancy models, two-species occupancy models, and distance sampling techniques to evaluate changes in occupancy and population density of resident bird species during three consecutive winters. We found an aggregation pattern between two resident species (Psaltriparus minimus and Thryomanes bewickii) with three migratory warblers (Cardellina pusilla, Setophaga coronata and Setophaga townsendi). Thus, our results provide evidence of the formation of mixed-species flocks in our study area. We also conclude that resident birds experience different demographic and behavioral processes during winter that not necessarily result from interspecific interactions with migratory birds.     

Geolocators map the wintering grounds of threatened Lesser Kestrels in Africa

Aim  To identify the wintering grounds of the threatened western European Lesser Kestrels to focus conservation efforts in those areas.
Location  Huelva Province, southern Spain, as breeding range, and western Africa (Senegal and Mauritania), as wintering range.
Methods  We used archival light level geolocators (1.5 g) to map the wintering areas and determine some characteristics of the migratory journeys of 20 adult Lesser Kestrels from the Iberian Peninsula tagged in 2007.
Results  Thirteen geolocators were recovered the following breeding season (2008) after attachment in 2007. Four recovered geolocators provided useful data. According to kernel density analyses, kestrels wintered near the Senegal River (border between Mauritania and Senegal). Pre-nuptial migration took longer than the post-nuptial migration, which may be the consequence of a loop migration.
Main conclusions  Geolocators have solved a crucial conservation question (i.e. the winter destination of western European Lesser kestrels), and these devices have thus proved useful to determine the location of the winter quarters of small sized migratory species. Our data indicate that European Lesser Kestrels winter in West Africa, in accordance with previous suggestions based on scattered observations during the winter months. This valuable information should serve to focus conservation efforts both in northern Senegal and southern Mauritania. Large roosts gathering thousands of lesser kestrels had been recorded in these areas over the years, but there was no previous confirmation of individuals staying all winter long. Specific and sustained protection of the roost sites, where the birds may be most vulnerable, should be sought in conjunction with local authorities.     

Mechanical and structural adaptations to migration in the flight feathers of a Palaearctic passerine

Current avian migration patterns in temperate regions have been developed during the glacial retreat and subsequent colonization of the ice‐free areas during the Holocene. This process resulted in a geographic gradient of greater seasonality as latitude increased that favoured migration‐related morphological and physiological (co)adaptations. Most evidence of avian morphological adaptations to migration comes from the analysis of variation in the length and shape of the wings, but the existence of intra‐feather structural adjustments has been greatly overlooked despite their potential to be under natural selection. To shed some light on this question, we used data from European robins Erithacus rubecula overwintering in Campo de Gibraltar (Southern Iberia), where sedentary robins coexist during winter with conspecifics showing a broad range of breeding origins and, hence, migration distances. We explicitly explored how wing length and shape, as well as several functional (bending stiffness), developmental (feather growth rate) and structural (size and complexity of feather components) characteristics of flight feathers, varied in relation to migration distance, which was estimated from the hydrogen stable isotope ratios of the summer‐produced tail feathers. Our results revealed that migration distance not only favoured longer and more concave wings, but also promoted primaries with a thicker dorsoventral rachis and shorter barb lengths, which, in turn, conferred more bending stiffness to these feathers. We suggest that these intra‐feather structural adjustments could be an additional, largely unnoticed, adaptation within the avian migratory syndrome that might have the potential to evolve relatively quickly to facilitate the occupation of seasonal environments.     

African Odyssey project – satellite tracking of black storks Ciconia nigra breeding at a migratory divide

The African Odyssey project focuses on studying the migration of the black stork Ciconia nigra breeding at a migratory divide. In 1995–2001, a total of 18 black storks breeding in the Czech Republic were equipped with satellite (PTT) and VHF transmitters. Of them, 11 birds were tracked during at least one migration season and three birds were tracked repeatedly. The birds migrated either across western or eastern Europe to spend the winter in tropical west or east Africa, respectively. One of the juveniles made an intermediate route through Italy where it was shot during the first autumn migration. The mean distance of autumn migration was 6,227 km. The eastern route was significantly longer than the western one (7,000 km and 5,667 km respectively). Important stopover sites were discovered in Africa and Israel. Wintering areas were found from Mauritania and Sierra Leone in the west to Ethiopia and Central African Republic in the east and south. One of the storks migrating by the eastern migration route surprisingly reached western Africa. Birds that arrived early in the wintering areas stayed longer than those arriving later. On the average, birds migrating via the western route spent 37 d on migration compared to 80 d for birds migrating via the eastern route. The mean migration speed in the autumn was 126 km/d and the fastest stork flew 488 km/d when crossing the Sahara. The repeatedly tracked storks showed high winter site fidelity.     

Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration

Migration during spring is usually faster than during autumn because of competition for breeding territories. In some cases, however, the costs and benefits associated with the environment can lead to slower spring migration, but examples are quite rare. We compared seasonal migration strategies of the endangered Baltic population of the dunlin Calidris alpina schinzii using light‐level geolocator data from 26 individuals breeding in Finland. Autumn migration was faster, with individuals showing a ‘jump’ and ‘skipping’ migration strategy characterised by fewer stationary periods, shorter total stopping time and faster flight. Spring migration was slower, with individuals using a ‘skipping’ strategy. The duration of migration was longer for early departing birds during spring but not during autumn suggesting that early spring migrants are prevented from arriving to the breeding areas or that fueling conditions are worse on the stopover sites for early arriving individuals. Dunlins showed high migratory connectivity. All individuals had one long staging at the Wadden Sea in the autumn after which half of the individuals flew 4500 km non‐stop to Banc d’Arguin, Mauritania. The other half stopped briefly on the Atlantic coast on their way to Mauritania. One bird wintered on the coast of Portugal. Nine individuals that carried geolocators for two years were site faithful to their final non‐breeding sites. Based on the strategies during the non‐breeding period we identified, Baltic dunlin may be especially vulnerable to rapid environmental changes at the staging and non‐breeding areas. Consequently, the preservation of the identified non‐breeding areas is important for their conservation.     

Breeding phenology and winter activity predict subsequent breeding success in a trans-global migratory seabird

Inter-seasonal events are believed to connect and affect reproductive performance (RP) in animals. However, much remains unknown about such carry-over effects (COEs), in particular how behaviour patterns during highly mobile life-history stages, such as migration, affect RP. To address this question, we measured at-sea behaviour in a long-lived migratory seabird, the Manx shearwater (Puffinus puffinus) and obtained data for individual migration cycles over 5 years, by tracking with geolocator/immersion loggers, along with 6 years of RP data. We found that individual breeding and non-breeding phenology correlated with subsequent RP, with birds hyperactive during winter more likely to fail to reproduce. Furthermore, parental investment during one year influenced breeding success during the next, a COE reflecting the trade-off between current and future RP. Our results suggest that different life-history stages interact to influence RP in the next breeding season, so that behaviour patterns during winter may be important determinants of variation in subsequent fitness among individuals.     

Seasonal migration of male red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis>) in southern Sweden and consequences for management

Red deer Cervus elaphus is a highly appreciated and intensively managed game species throughout Europe. A common management objective is a sustainable harvest of large trophies. In southern Sweden, management has mainly aimed at preserving the nominate subspecies C. elaphus elaphus. Seasonal migration of red deer males may, however, complicate both harvest management as well as conservation efforts. I used individually identified male red deer in southern Sweden to observe distance travelled from rutting areas to areas used by males in summer and winter. Adult males were identified by antler shape and photo-documented during rut. Photos from the rut were compared to trophies of deer harvested or found dead, to found cast antlers and to stags photographed during summer. From 1969 to 2007, a distance between rutting ground and summer/winter quarters was established for 96 identified stags. An average distance of 14 km and a maximum distance of 47 km were recorded between rut and summer/winter observations. The seasonal migration of males increases the risk of overexploitation of males with harvest in both rutting areas and wintering areas. Harvest management and conservation efforts may fail if males seasonally migrate outside the management unit. The results suggest that seasonal migration must be considered in harvest management and conservation and that there is a need for a regulation of male harvest. Furthermore, the study stresses that the success in deer management of single hunting units, may be largely dependent on the harvest policies in the near surroundings as well as in areas tenths of kilometres away, suggesting that a successful management must rely on co-operation and co-ordination on a landscape scale.     

Seasonal migration strategies of Common Ringed Plovers Charadrius hiaticula

How individual birds schedule their movements and use different sites during the non‐breeding season are fundamental issues in avian migration ecology, and studies have often revealed strong seasonal variation in such strategies. Using geolocators we tracked Common Ringed Plovers Charadrius hiaticula from northern Norway to West Africa and back to assess whether there were differences in migratory speed, duration and stopover use between autumn and spring migration and whether birds used multiple sites during the non‐breeding season. Although the pace of migration was similar between autumn and spring, the length of flight bouts and duration of the preceding stopovers were positively correlated only in autumn. Four of five birds showed a marked southward movement in mid‐winter.     

Seasonal differences in energy expenditure,flight characteristics and spatial utilization of Dalmatian Pelicans Pelecanus crispus in Greece

Animals typically adjust their behaviour to their changing environment throughout the annual cycle, modulating key processes such as the timing of breeding and the onset of migration. Such behavioural changes are commonly manifested in the movements and the energetic balance of individuals in relation to their species‐specific physiological characteristics, habitat attributes and the environmental properties of their distribution ranges. We used GPS and acceleration data collected using transmitters on free‐ranging birds to quantify annual movement patterns and estimate energy expenditure of the Dalmatian Pelican Pelecanus crispus, a large, soaring avian species which performs short‐distance migration and spends its entire annual cycle in mid‐latitudes. To assess the representativeness of our results, the transmitter effect was also tested. We found that daily trends in the overall dynamic body acceleration (ODBA; a proxy for energy expenditure) differed among seasons, with the highest values occurring during spring and the lowest during winter. Long inter‐lake flights were very rare in winter, and the number of flights and ODBA during spring was higher than during summer, suggesting greater motivation to move in spring. Although transmitters may have affected the birds, as none of the tagged birds bred, we found seasonal differences in behaviour and activity level. The observed patterns in differences in activity levels, long‐distance flights and flight characteristics between seasons suggest an annual rhythm of energy expenditure. These findings allow a better understanding of bird phenology, specifically regarding adaptations to wintering in a cold climate by reducing movement‐driven energy expenditure. Finally, the identification of periods with high and low energy expenditure may guide future conservation efforts by adjusting conservation plans in accordance with changing needs during the annual cycle.     

How hazardous is the Sahara Desert crossing for migratory birds? Indications from satellite tracking of raptors

We investigated the risk associated with crossing the Sahara Desert for migrating birds by evaluating more than 90 journeys across this desert by four species of raptors (osprey Pandion haliaetus, honey buzzard Pernis apivorus, marsh harrier Circus aeruginosus and Eurasian hobby Falco subbuteo) recorded by satellite telemetry. Forty per cent of the crossings included events of aberrant behaviours, such as abrupt course changes, slow travel speeds, interruptions, aborted crossings followed by retreats from the desert and failed crossings due to death, indicating difficulties for the migrants. The mortality during the Sahara crossing was 31 per cent per crossing attempt for juveniles (first autumn migration), compared with only 2 per cent for adults (autumn and spring combined). Mortality associated with the Sahara passage made up a substantial fraction (up to about half for juveniles) of the total annual mortality, demonstrating that this passage has a profound influence on survival and fitness of migrants. Aberrant behaviours resulted in late arrival at the breeding grounds and an increased probability of breeding failure (carry-over effects). This study also demonstrates that satellite tracking can be a powerful method to reveal when and where birds are exposed to enhanced risk and mortality during their annual cycles.     

The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration

The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d(-1)) than during spring (233 km d(-1)). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.     

Chukar Seasonal Survival and Probable Causes of Mortality

ABSTRACT Chukars (Alectoris chukar) have been introduced throughout the world. Despite this widespread distribution, limited information regarding seasonal survival, probable causes of mortality, and other basic life history characteristics is available to manage this harvested species. We estimated the probable cause of mortality for chukars with radiotransmitters by examining evidence at kill sites. We used model selection to evaluate influences of seasonal effects (fall raptor migration, peak migration, and reproductive period), demographic effects (age and sex), radio weight, and year on survival of chukars in western Utah, USA, by using a known-fate model in Program MARK and 2 years of telemetry data. We captured and randomly fitted 125 individual chukars with 2 different-sized radiotransmitters (97 F, 20 M, 8 sex undetermined). Model selection results showed our top 3 models accounted for 99% of Akaike's Information Criterion weight, and each one had seasonal and year effects. Two-week survival estimates were lower during peak raptor migration in both years and significantly (P < 0.05) so in year 2 (2-week S = 0.87, 95% CI = 0.77–0.94) compared with other year 2 intervals (2-week S > 0.91). Annual survival was lower in 2005 (S = 0.03, 95% CI = 0.01–0.09) compared with 2006 (S = 0.19, 95% CI = 0.12–0.31). We documented 95 deaths and classified 56% unknown, 33% avian predation, 8% hunter harvest, and 3% mammalian predation. Our research suggests that predation on chukars is substantial during the peak fall raptor migratory period and that the hunting take under current regulations is relatively small and likely compensatory.     

The scheduling of molt in migratory birds

Summary We model the yearly cycle of small migratory birds to explain the variation in scheduling of complete molt, in particular why some birds molt immediately after breeding on the breeding grounds (summer molt) whereas others migrate to their wintering grounds before molt is initiated (winter molt). We employ the method of dynamic programming, because of its suitability for modelling life history traits. Feather quality and latitude entered the model as state variables and were assumed to affect survival rate and reproductive success. Migration and molt were assumed to be associated with increased mortality risks. By changing the parameters in the model we were able to generate most existing molt patterns, including summer and winter molt, biannual (summer and winter) molt, and molt migration. Our model suggests that the scheduling of molt is basically a result of a trade-off between having a high feather quality during breeding versus during the non-breeding period. A high impact of feather quality on survival rate in combination with low costs of molt resulted in biannual molt. Winter molt became more likely as the survival rateper se increased. A low seasonal amplitude in survival rate is a prerequisite for the occurrence of molt migration. Molt duration, migration costs and reproductive successper se were found to have no impact on the timing of molt. We also investigated the effect of benefits from prior occupancy at breeding and winter grounds.