Migratory birds, some small and light weight as matchboxes, engage in seasonal inter-continental journeys in order to take advantage of the long summer days and abundance of food at northern latitudes to breed and raise their young, and then escape the harsh winters by migrating to lower latitudes. This thesis deals with two important aspects of migration, the routes taken during migration and the birds’ behaviour at stopovers. The migratory routes are for many species unknown, whole or in part, and this is especially true for species that migrate nocturnally. At stopovers birds replenish fuel reserves that powers migratory flight, and studying how birds utilise stopovers is important in order to understand how migration is organised. In this thesis I have used modern tracking technology to study both continental wide movements of thrush nightingales (Luscinia luscinia) and common rosefinches (Carpodacus erythrinus) using small light-level geolocators, and smaller scale movements at a single stopover site of garden warblers (Sylvia borin) using miniature radio-transmitters. I have also studied the fuelling behaviour of garden warblers during autumn migration in the field and in the lab, and great reed warblers (Acrocephalus arundinaceus) at a stopover site on Crete during spring migration after the Sahara crossing. The thesis discusses the significance of several aspects of migration shown by the birds that would have been very difficult to detect without the aid of modern tracking technology, such as loop migration, prolonged stops during migration, multiple wintering sites, and nocturnal relocations at stopover sites. Studies carried out at stopover sites also show that garden warblers and great reed warblers can attain large fuel loads even at sites where they have no barrier to cross and this might be a result of good foraging conditions. The thesis also highlights the importance of combining different techniques when studying stopover behaviour to get reliable estimates on stopover durations and fuel deposition rates as well as the importance of choosing sites preferred by birds when planning stopover studies. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 4: Manuscript. Paper 5: Manuscript.</p>

Master of Science / Department of Biology / Brett K. Sandercock / The migratory period is a critical time for birds. Population declines in long-distance migratory birds have been associated with trophic mismatches among climate change, timing of food availability, and timing of migratory movements. Studies on migratory songbirds have been limited to eastern North America and Europe, and migration strategies of birds may differ along the Pacific flyway. We evaluated the stopover ecology of five species of migratory songbirds at a coastal site in northern California. We found variation in changes in timing of spring migration, but consistent changes in timing of autumn migration over a 22-year period. Timing of spring migration advanced for Swainson’s Thrushes (Catharus ustulatus) and Yellow Warblers (Setophaga petechia), was compressed for Pacific-slope Flycatchers (Empidonax difficilis) and Wilson’s Warblers (Cardellina pusilla), but was protracted for Orange-crowned Warblers (Oreothlypis celata). In contrast, timing of autumn migration was delayed for Pacific-slope Flycatchers, Orange-crowned Warblers, and Yellow Warblers, but was protracted for Wilson’s Warblers. Warm, wet conditions were associated with advanced spring migration, and warm, dry conditions were associated with delayed autumn migration. Changes in timing of migration related to climate conditions were strongest for Pacific-slope Flycatchers and Orange-crowned Warblers. Stopover duration of our five study species was longer than songbirds in eastern flyways, and on average, was longer in spring than autumn. Pacific-slope Flycatchers and Yellow Warblers had longer stopovers in spring, whereas Swainson’s Thrushes and Wilson’s Warblers stopped had longer stopovers in autumn. Birds captured at low body mass had longer stopovers in spring and autumn. Migratory birds in western North America may have different migration strategies because of differences in climate and geography, but more study is needed to discover migratory routes and compare our results to other stopover sites along the Pacific flyway. Understanding differences in migration strategies of different populations of the same species are important in directing conservation efforts, especially in light of ongoing climate change.

Turkey lies on one of major migratory routes between Palearctic and Afrotropical regions.Despite its importance for many species, few studies exist on bird migration over Turkey. In this study, autumn migration dynamics and stopover behaviour of Willow Warbler (Phylloscopus trochilus), a small insectivorous passerine, was documented and analyzed at Manyas KuScenneti National Park (northwestern Turkey). Birds were mistnetted, ringed, measured, weighed and fat-scored from mid August in 2002 and end of August in 2003 to end of October in both years. Totally 543 and 929 Willow Warblers were ringed in 2002 and 2003 respectively. For 2002 and 2003 respectively, fat score values (mean&plusmn / SE) are 4.63&plusmn / 0.06 and 3.84&plusmn / 0.05 and body mass values are 11.38&plusmn / 0.07 and 10.37&plusmn / 0.05 grams for birds captured the first time.Fat scores in 2003 show a bimodal distribution with peaks of T2 and T5, indicating populations or age classes with different migratory strategies.The number of retraps constituted 9.2-12.1 % of the total in 2002 and 2003. In both years,minimum stopover length ranged from 1 to 15 or 16 days with a median of 5 days. The majority of the retraps put on significant fat in both years. Retraps continued to put on weight for up to two weeks after they arrived. In this first ever study documenting passerine migration at Manyas KuScenneti National Park, it was revealed that such wetlands provide crucial stopover habitat for many migrant passerines, enables them to gain necessary fat loads before crossing two ecological barriers,the Mediterranean Sea and the Sahara.

The mallard (Anas platyrhynchos) is the most numerous and widespread duck in the northern hemisphere and a model species in ecology and harvest management. Migration is a crucial life stage for many birds and understanding the drivers of migration has important implications for conservation biology and assessment of animal population responses to global changes. Furthermore, mallard migration is a fundamental determinant of the epidemiology of many diseases of major relevance for both animal and human health. For example, it is the reservoir host for influenza A viruses (IAV), a widespread zoonosis causing mortality and economic damage. Improved knowledge of mallard behaviour during migration and the impacts of infection in mallards is needed to determine the role of wild birds in global IAV dynamics. This thesis focuses on mallard stopover ecology, an explicitly important part of the annual life cycle that is not well understood. The study area was southern Öland, SE Sweden, where mallard stopover behaviour was scrutinized by a combination of telemetry and ringing data analyses. Specifically, habitat preferences, movements, and emigration decisions were studied in-depth. Potential effects of low pathogenic avian influenza (LPAIV) infection on movement parameters were also investigated. Radio-tracking revealed that stopover mallards adhered to a strict diel pattern, in which they spent the days resting along the coast, visited crop fields at dawn and dusk, and foraged on inland water bodies during the darkest night hours. Notably, the importance of residual maize, as well as small ephemeral wetlands on the unique alvar steppe habitat that predominates on Öland, was previously unknown. LPAIV infection status did not affect movement behaviour, highlighting the possible risk of spread of IAV from wild mallards to poultry along the migratory flyway. Through capture-mark-recapture modelling, it was confirmed that weather, particularly wind direction, was the most important determinant of departure from the stopover site. In contrast, the body condition of departing mallards was less crucial. Taken together, the research presented in this thesis contributes to improved knowledge about mallard stopover ecology and its role in LPAIV disease dynamics.

<div> <div> <div> <p>Billions of birds migrate annually between breeding and wintering habitats, following transient resources. Though a majority of time is spent in breeding and wintering habitats, habitats used during the migratory periods are especially important for migrating birds. Migration and stopovers, where birds rest and refuel before continuing a migratory journey, are critical points in a bird’s annual lifecycle, and are important ecologically, socially, and economically. Populations of migratory birds are declining on a global scale, however, and proper management is vital to their persistence in an urbanizing environment. Indiana in the Midwestern United States is an important area in which to study stopover ecology of migratory birds, as it is a fragmented forest- urban-agricultural matrix almost entirely managed through private ownership. In this dissertation, I studied three questions of stopover ecology within the landscape context of the Midwestern United States, primarily using weather surveillance radar and eBird citizen science data. </p> <p><br></p><p>First, I studied spatiotemporal changes over an 11-year period (autumn 2005-2016 and spring 2006-2017) in densities of nocturnally migrating birds at two radar stations in Indiana. I found that mean density of migratory birds stopping over in Indiana declined by approximately 6.8% annually, but variability in stopover site use increased over the same period. This is consistent with other work completed on continental scales, and highlights the need for further conservation of migratory birds. Second, I studied patterns of stopover site use in Indiana during spring 2016- 2017 and autumn 2015-2016, identifying landscape and local factors associated with those patterns. I used both traditional land cover characteristics and a novel approach using human socioeconomic measures to describe these patterns, and found that socioeconomics, particularly the size of a housing unit, were among the most important predictors of migratory bird density in Indiana. The results from this study suggest that migratory birds are utilizing urban habitats, which are known to contain several novel hazards for birds, but that migratory birds will benefit greatly from interdisciplinary work focusing on urban habitats. Third, I explored a novel method of using weather surveillance radar and eBird citizen science data in combination with each other, to see if both measures provided similar estimates of bird abundances during stopover. Though I found no correlation between the two, I argue that eBird and radar still provide important and complementary insights for the field of migration ecology. Finally, I provide guidelines for private landowners in Indiana on management for declining populations of migratory birds. </p> </div> </div> </div>

La creciente necesidad de que, tanto los científicos como los técnicos medioambientales, conozcan los diversos aspectos de la ecología de los animales para proponer acciones de conservación adecuadas, ha promovido el desarrollo de técnicas y herramientas que permitan desentrañar algunas de las incógnitas que envuelven a un nuevo paradigma emergente; la "ecología del movimiento", enfocada a estudiar todo tipo de movimientos realizados por todo tipo de organismos. Poco a poco, y sobre todo en las últimas décadas, las nuevas tecnologías de seguimiento remoto aplicadas al estudio del movimiento animal han permitido llevar a cabo nuevas y modernas investigaciones con una amplia gama de enfoques científicos especializados para estudiar los diferentes movimientos llevados a cabo por grupos específicos de organismos. En las aves, la aplicación de estas técnicas ha aportado multitud de nueva información sobre el uso del hábitat y del espacio y, sobre todo, ha permitido el seguimiento de las especies a lo largo de sus movimientos migratorios y en las áreas de invernada, aportando gran cantidad de información en estas etapas de su ciclo biológico muy desconocidas hasta la aplicación de estas técnicas. El Cernícalo primilla (Falco naumanni) es una pequeña rapaz migratoria que estuvo muy amenazada debido a un fuerte declive de sus poblaciones ocurrido a finales del siglo pasado, llegando a extinguirse en muchas localidades a lo largo de su rango de distribución; la situación llevo a que la especie estuviera catalogada como en “peligro” y fue objeto de numerosas investigaciones. Sin embargo, varios aspectos sobre su biología básica seguían siendo desconocidos. Por ejemplo, aunque el conocimiento sobre la migración y la invernada había mejorado en los últimos años con la utilización de geolocalizadores, esta técnica puede generar grandes errores (hasta unos pocos cientos de kilómetros) en la estimación de la ubicación de un ave y no es posible mapear las rutas e inquirir su fenología migratoria con precisión. Por otro lado, la especie ha sufrido recientemente de nuevo una fuerte regresión, con registros de reducciones en poblaciones reproductoras de más del 50% de los efectivos poblacionales en solo una generación en algunas áreas de reproducción de la Península Ibérica (especialmente durante las temporadas reproductivas de 2017 y 2018). Los investigadores señalan que la principal causa de las repentinas disminuciones que se han producido en los últimos años puede deberse a los eventos que ocurren en las zonas de invernada que ocupa la especie y de las cuales no existe información detallada; sin embargo existen otros aspectos de su declive, actual y pasado, que siguen siendo controvertidos, p.ej., las posibles afecciones que causa la pérdida de hábitat asociado a la construcción de infraestructuras en hábitat prioritarios para la especie, que hemos investigado recientemente. En este trabajo de investigación, además, aportamos nuevos datos sobre la duración correcta de las migraciones del Cernícalo primilla, de las rutas migratorias exactas, de los posibles efectos de las condiciones climáticas (p.ej., el viento) durante su migración y hemos estudiado el uso del hábitat y las zonas de invernada de la especie en el Sahel. Por último, con nuestras investigaciones con un enfoque multiespecífico (trabajando con el Águila calzada Hieraaetus pennatus y el Aguilucho cenizo Circus pygargus), hemos podido describir mejor el hábitat de las rapaces en sus zonas de invernada y definir los puntos clave hacia los que focalizar los esfuerzos para conservar, no solo al Cernícalo primilla, sino a muchas rapaces migratorias transaharianas.

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