Integrating data types to estimate spatial patterns of avian migration across the Western Hemisphere.

For many avian species, spatial migration patterns remain largely undescribed, especially across hemispheric extents. Recent advancements in tracking technologies and high-resolution species distribution models (i.e., eBird Status and Trends products) provide new insights into migratory bird movements and offer a promising opportunity for integrating independent data sources to describe avian migration. Here, we present a three-stage modeling framework for estimating spatial patterns of avian migration. First, we integrate tracking and band re-encounter data to quantify migratory connectivity, defined as the relative proportions of individuals migrating between breeding and nonbreeding regions. Next, we use estimated connectivity proportions along with eBird occurrence probabilities to produce probabilistic least-cost path (LCP) indices. In a final step, we use generalized additive mixed models (GAMMs) both to evaluate the ability of LCP indices to accurately predict (i.e., as a covariate) observed locations derived from tracking and band re-encounter data sets versus pseudo-absence locations during migratory periods and to create a fully integrated (i.e., eBird occurrence, LCP, and tracking/band re-encounter data) spatial prediction index for mapping species-specific seasonal migrations. To illustrate this approach, we apply this framework to describe seasonal migrations of 12 bird species across the Western Hemisphere during pre- and postbreeding migratory periods (i.e., spring and fall, respectively). We found that including LCP indices with eBird occurrence in GAMMs generally improved the ability to accurately predict observed migratory locations compared to models with eBird occurrence alone. Using three performance metrics, the eBird + LCP model demonstrated equivalent or superior fit relative to the eBird-only model for 22 of 24 species-season GAMMs. In particular, the integrated index filled in spatial gaps for species with over-water movements and those that migrated over land where there were few eBird sightings and, thus, low predictive ability of eBird occurrence probabilities (e.g., Amazonian rainforest in South America). This methodology of combining individual-based seasonal movement data with temporally dynamic species distribution models provides a comprehensive approach to integrating multiple data types to describe broad-scale spatial patterns of animal movement. Further development and customization of this approach will continue to advance knowledge about the full annual cycle and conservation of migratory birds.

Estimating updraft velocity components over large spatial scales: contrasting migration strategies of golden eagles and turkey vultures.

Soaring birds migrate in massive numbers worldwide. These migrations are complex and dynamic phenomena, strongly influenced by meteorological conditions that produce thermal and orographic uplift as the birds traverse the landscape. Herein we report on how methods were developed to estimate the strength of thermal and orographic uplift using publicly available digital weather and topography datasets at continental scale. We apply these methods to contrast flight strategies of two morphologically similar but behaviourally different species: golden eagle, Aquila chrysaetos, and turkey vulture, Cathartes aura, during autumn migration across eastern North America tracked using GPS tags. We show that turkey vultures nearly exclusively used thermal lift, whereas golden eagles primarily use orographic lift during migration. It has not been shown previously that migration tracks are affected by species-specific specialisation to a particular uplift mode. The methods introduced herein to estimate uplift components and test for differences in weather use can be applied to study movement of any soaring species.

Generation of raptor diversity in Europe: linking speciation with climate changes and the ability to migrate.

Europe holds a rich community of diurnal birds of prey, and the highest proportion of transcontinental migratory raptorial species of any landmass. This study will test the hypotheses that the high diversification of the raptor assemblage in Europe is a recent event, that closely related species sharing the same trophic niches can only coexist in sympatry during the breeding period, when food availability is higher, and finally that migration is a function of size, with the smaller species in every trophic group moving further. A consensus molecular phylogeny for the 38 regular breeding species of raptors in Europe was obtained from BirdTree (www.birdtree.org). For the same species, a trophic niche cluster dendrogram was constructed. Size and migratory strategy were introduced in the resulting phylogeny, where trophic groups were also identified. Multispecific trophic groups tended to be composed of reciprocal sister species of different sizes, while monospecific groups (n = 3) were composed of highly specialized species. Many speciation events took place recently, during the glacial cycles of the Quaternary, and size divergence among competing species may be due to character displacement. Nowadays, the smaller species in every trophic group migrate to sub-Saharan Africa. This investigation illustrates how the rich assemblage of diurnal birds of prey in Europe, more diverse and more migratory than, for instance, the North American assemblage at equivalent latitudes, has emerged recently due to the multiplication of look-alike species with similar trophic ecologies, possibly in climate refugia during cold periods.

Migration path annotation: cross-continental study of migration-flight response to environmental conditions.

Understanding the movements of animals is pivotal for understanding their ecology and predicting their survival in the face of rapid global changes to climate, land use, and habitats, thus facilitating more effective habitat management. Migration by flying animals is an extreme form of movement that may be especially influenced by weather. With satellite telemetry studies, and the growing availability of information about the Earth's weather and land surface conditions, many data are collected that can advance our understanding about the mechanisms that shape migrations. We present the track annotation approach for movement data analysis using information about weather from the North American Reanalysis data set, a publicly available, regional, high-resolution model-observation hybrid product, and about topography, from a publicly available high-resolution digital elevation model (DEM). As a case study, we present the analysis of the response to environmental conditions in three contrasting populations of Turkey Vultures (Cathartes aura) across North America, tracked with a three-dimensional GPS-based sensor. Two populations in the east and west coasts of the United States responded similarly to weather, indicating use of both slope and thermal soaring. Continental-interior, "Plains populations," exhibited a different migratory pattern primarily indicative of thermal soaring. These differences help us understand the constraints and behaviors of soaring migrants. The track annotation approach allowed large-scale comparative study of movement in an important migratory species, and will enable similar studies at local to global scales.

Inclement weather forces stopovers and prevents migratory progress for obligate soaring migrants.

BACKGROUND: Migrating birds experience weather conditions that change with time, which affect their decision to stop or resume migration. Soaring migrants are especially sensitive to changing weather conditions because they rely on the availability of environmental updrafts to subsidize flight. The timescale that local weather conditions change over is on the order of hours, while stopovers are studied at the daily scale, creating a temporal mismatch. METHODS: We used GPS satellite tracking data from four migratory Turkey Vulture (Cathartes aura) populations, paired with local weather data, to determine if the decision to stopover by migrating Turkey Vultures was in response to changing local weather conditions. We analyzed 174 migrations of 34 individuals from 2006 to 2019 and identified 589 stopovers based on variance of first passage times. We also investigated if the extent of movement activity correlated with average weather conditions experienced during a stopover, and report general patterns of stopover use by Turkey Vultures between seasons and across populations. RESULTS: Stopover duration ranged from 2 h to more than 11 days, with 51 % of stopovers lasting < 24 h. Turkey Vultures began stopovers immediately in response to changes in weather variables that did not favor thermal soaring (e.g., increasing precipitation fraction and decreasing thermal updraft velocity) and their departure from stopovers was associated with improvements in weather that favored thermal development. During stopovers, proportion of activity was negatively associated with precipitation but was positively associated with temperature and thermal updraft velocity. CONCLUSIONS: The rapid response of migrating Turkey Vultures to changing weather conditions indicates weather-avoidance is one of the major functions of their stopover use. During stopovers, however, the positive relationship between proportion of movement activity and conditions that promote thermal development suggests not all stopovers are used for weather-avoidance. Our results show that birds are capable of responding rapidly to their environment; therefore, for studies interested in external drivers of weather-related stopovers, it is essential that stopovers be identified at fine temporal scales.

Conserving migratory land birds in the new world: do we know enough?

Migratory bird needs must be met during four phases of the year: breeding season, fall migration, wintering, and spring migration; thus, management may be needed during all four phases. The bulk of research and management has focused on the breeding season, although several issues remain unsettled, including the spatial extent of habitat influences on fitness and the importance of habitat on the breeding grounds used after breeding. Although detailed investigations have shed light on the ecology and population dynamics of a few avian species, knowledge is sketchy for most species. Replication of comprehensive studies is needed for multiple species across a range of areas, Information deficiencies are even greater during the wintering season, when birds require sites that provide security and food resources needed for survival and developing nutrient reserves for spring migration and, possibly, reproduction. Research is needed on many species simply to identify geographic distributions, wintering sites, habitat use, and basic ecology. Studies are complicated, however, by the mobility of birds and by sexual segregation during winter. Stable-isotope methodology has offered an opportunity to identify linkages between breeding and wintering sites, which facilitates understanding the complete annual cycle of birds. The twice-annual migrations are the poorest-understood events in a bird's life. Migration has always been a risky undertaking, with such anthropogenic features as tall buildings, towers, and wind generators adding to the risk. Species such as woodland specialists migrating through eastern North America have numerous options for pausing during migration to replenish nutrients, but some species depend on limited stopover locations. Research needs for migration include identifying pathways and timetables of migration, quality and distribution of habitats, threats posed by towers and other tall structures, and any bottlenecks for migration. Issues such as human population growth, acid deposition, climate change, and exotic diseases are global concerns with uncertain consequences to migratory birds and even less-certain remedies. Despite enormous gaps in our understanding of these birds, research, much of it occurring in the past 30 years, has provided sufficient information to make intelligent conservation efforts but needs to expand to handle future challenges.

Migratory patterns and settlement areas revealed by remote sensing in an endangered intra-African migrant, the Black Harrier (Circus maurus).

Annual movements have been widely described for birds migrating across the Americas and between Eurasia and Africa, yet relatively little information exists for intra-African migrants. Identifying the areas used throughout a species annual cycle by understanding migratory patterns and settlement areas during breeding and non-breeding seasons is essential for conservation initiatives. Here, we describe for the first time, the migratory patterns and settlement areas of an endangered raptor endemic to Southern Africa, the Black Harrier (Circus maurus). From 2008 to 2015, thirteen breeding adult Black Harriers were trapped in south-western South Africa and fitted either with a GPS-GSM or with a PTT tracker device. Adults were monitored for 365 ± 198 days (range: 56-819 days) revealing great individual variability in annual movements. Most Black Harriers performed an unusual West-East migration from their breeding areas, but routes of all migrating individuals covered the entire southern land area of South Africa and Lesotho. The distance travelled averaged 814 ± 324 km, but unlike many other species, migrants travelled faster during post-breeding (i.e. austral summer) (207.8 ± 113.2 km.day-1) than during pre-breeding (i.e. austral winter/spring) migrations (143.8 ± 32.2 km.day-1). Although most marked individuals displayed movements similar to those that bred following pre-breeding migrations, only two of thirteen were confirmed as breeders the year after being tagged. This suggests that individuals may sometimes take a sabbatical year in reproduction, although this requires confirmation. Most tagged birds died on migration or during the non-breeding season. Adults frequently returned to the same non-breeding settlement areas, and often used up to 3 different locations an average of about 200 km apart. On the other hand, there was wide variation in distance between subsequent reproductive events. We discuss the implications of our study for the conservation of Black Harriers and more broadly for intra-African bird migrants.

Seasonal micro-migration in a farm-island population of striated caracaras (Phalcoboenus australis) in the Falkland Islands.

BACKGROUND: The extent to which seasonal changes in food availability affect small-scale movements in free-ranging populations of birds of prey is relatively little studied. Here we describe a seasonal "micro-migration" of a farm-island population of striated caracaras (Phalcoboenus australis) in the Falkland Islands in response to seasonal changes in the availability of seabird carcasses. We banded more than 450 individuals on Saunders Island, deployed archival and satellite GPS data loggers on 17 individuals, and monitored movements within and between two feeding areas on Saunders Island, a "marine-subsidized" site near seabird colonies and an anthropogenic "human-subsidized" farm site 16 km to the southeast. RESULTS: During 67 observation days between 2010 and 2015, resightings of 312 banded caracaras were greater at the marine-subsidized site during austral summer than winter, and the total daily resightings varied significantly between spring versus summer, summer versus winter, autumn versus spring, and autumn versus winter. Resightings were higher at the human-subsidized site in austral winter than summer and the total daily resightings varied significantly across all bi-seasonal comparisons. Resightings indicated that at least 12 of 197 birds (6.1%) moved between the human- and marine-subsidized sites at least once during the same winter, 15 of 335 birds (4.5%) did so in spring, none of 164 birds did so in summer, and 16 of 297 birds (5.4%) did so in autumn. Individuals fitted with archival GPS data loggers at the marine-subsidized site in summer maintained highly localized 95% kernel core areas (0.55 ± 0.12 km2 [mean ± SD]), whereas those at the human-subsidized site in winter maintained larger 95% kernel core areas (3.8 ± 4.6 km2). Two of 6 satellite-tagged individuals that summered at known caracara breeding colonies 80 km WNW of Saunders Island were subsequently resighted in winter at the human-subsidized site. CONCLUSION: Our results suggest that seasonal shifts in food resource availability drive seasonal micro-migrations in a farm-island population of striated caracaras, and that farm sites can be critical in providing nutritional resources for caracaras when naturally occurring marine-subsidized resources become less available. Our results have important implications for striated caracara spatial ecology and conservation, as increased winter survival could improve the status of this globally Near-Threatened population.

Prevalence of west Nile virus antibodies in a breeding population of American Kestrels (Falco sparverius) in Pennsylvania.

West Nile virus (WNV) has been identified in nearly 300 species of wild birds, including raptors, in North America since its introduction in New York City in 1999. American Kestrels (Falco sparverius) are susceptible to WNV infection, and the numbers of these birds have declined along the Atlantic coast in recent years. We examined the population biology and WNV exposure of kestrels breeding in the area surrounding Hawk Mountain Sanctuary in Kempton, Pennsylvania, USA. The reproductive biology of kestrels in this area was studied from 1992 until 2004. The number of kestrels breeding in nestboxes in 2004 was only 44% of the 6-yr mean observed prior to 1999. During the 2004 nesting season (study period: 8 June through 22 July 2004), adult kestrels were trapped near the site of their nestboxes. Blood samples were obtained, and serum antibodies specific to WNV were quantified using a plaque reduction neutralization test. Of 22 birds tested, 21 exhibited serum antibodies to WNV, suggesting that most (95%) of the adult kestrels in the population had been exposed to WNV.

Is pre-breeding prospecting behaviour affected by snow cover in the irruptive snowy owl? A test using state-space modelling and environmental data annotated via Movebank.

BACKGROUND: Tracking individual animals using satellite telemetry has improved our understanding of animal movements considerably. Nonetheless, thorough statistical treatment of Argos datasets is often jeopardized by their coarse temporal resolution. State-space modelling can circumvent some of the inherent limitations of Argos datasets, such as the limited temporal resolution of locations and the lack of information pertaining to the behavioural state of the tracked individuals at each location. We coupled state-space modelling with environmental characterisation of modelled locations on a 3-year Argos dataset of 9 breeding snowy owls to assess whether searching behaviour for breeding sites was affected by snow cover and depth in an arctic predator that shows a lack of breeding site fidelity. RESULTS: The state-space modelling approach allowed the discrimination of two behavioural states (searching and moving) during pre-breeding movements. Tracked snowy owls constantly switched from moving to searching behaviour during pre-breeding movements from mid-March to early June. Searching events were more likely where snow cover and depth was low. This suggests that snowy owls adapt their searching effort to environmental conditions encountered along their path. CONCLUSIONS: This modelling technique increases our understanding of movement ecology and behavioural decisions of individual animals both locally and globally according to environmental variables.

African vultures don't follow migratory herds: scavenger habitat use is not mediated by prey abundance.

The ongoing global decline in vulture populations raises major conservation concerns, but little is known about the factors that mediate scavenger habitat use, in particular the importance of abundance of live prey versus prey mortality. We test this using data from the Serengeti-Mara ecosystem in East Africa. The two hypotheses that prey abundance or prey mortality are the main drivers of vulture habitat use provide alternative predictions. If vultures select areas based only on prey abundance, we expect tracked vultures to remain close to herds of migratory wildebeest regardless of season. However, if vultures select areas where mortality rates are greatest then we expect vultures to select the driest regions, where animals are more likely to die of starvation, and to be attracted to migratory wildebeest only during the dry season when wildebeest mortality is greatest. We used data from GSM-GPS transmitters to assess the relationship between three vulture species and migratory wildebeest in the Mara-Serengeti ecosystem. Results indicate that vultures preferentially cluster around migratory herds only during the dry season, when herds experience their highest mortality. Additionally during the wet season, Ruppell's and Lappet-faced vultures select relatively dry areas, based on Normalized Difference Vegetation Index, whereas White-backed vultures preferred wetter areas during the wet season. Differences in habitat use among species may mediate coexistence in this scavenger guild. In general, our results suggest that prey abundance is not the primary driver of avian scavenger habitat use. The apparent reliance of vultures on non-migratory ungulates during the wet season has important conservation implications for vultures in light of on-going declines in non-migratory ungulate species and use of poisons in unprotected areas.

Environmental drivers of variability in the movement ecology of turkey vultures (Cathartes aura) in North and South America.

Variation is key to the adaptability of species and their ability to survive changes to the Earth's climate and habitats. Plasticity in movement strategies allows a species to better track spatial dynamics of habitat quality. We describe the mechanisms that shape the movement of a long-distance migrant bird (turkey vulture, Cathartes aura) across two continents using satellite tracking coupled with remote-sensing science. Using nearly 10 years of data from 24 satellite-tracked vultures in four distinct populations, we describe an enormous amount of variation in their movement patterns. We related vulture movement to environmental conditions and found important correlations explaining how far they need to move to find food (indexed by the Normalized Difference Vegetation Index) and how fast they can move based on the prevalence of thermals and temperature. We conclude that the extensive variability in the movement ecology of turkey vultures, facilitated by their energetically efficient thermal soaring, suggests that this species is likely to do well across periods of modest climate change. The large scale and sample sizes needed for such analysis in a widespread migrant emphasizes the need for integrated and collaborative efforts to obtain tracking data and for policies, tools and open datasets to encourage such collaborations and data sharing.

Why birds with deferred sexual maturity are sedentary on islands: a systematic review.

BACKGROUND: Island faunas have played central roles in the development of evolutionary biology and ecology. Birds are among the most studied organisms on islands, in part because of their dispersal powers linked to migration. Even so, we lack of information about differences in the movement ecology of island versus mainland populations of birds. METHODOLOGY/PRINCIPAL FINDINGS: Here we present a new general pattern indicating that large birds with deferred sexual maturity are sedentary on islands, and that they become so even when they are migratory on the mainland. Density-dependent variation in the age at first breeding affects the survivorship of insular populations and this, in turn, affects the movement ecology of large birds. Because density-dependent variation in the age of first breeding is critical to the long-term survival of small isolated populations of long-lived species, migratory forms can successfully colonize islands only if they become sedentary once there. Analyses of the movement ecology of continental and insular populations of 314 species of raptors, 113 species of Ciconiiformes and 136 species of passerines, along with individual-based population simulations confirm this prediction. CONCLUSIONS: This finding has several consequences for speciation, colonization and survival of small isolated population of species with deferred sexual maturity.