Migratory connectivity of a Neotropical migratory songbird revealed by archival light-level geolocators.

Understanding migratory connectivity is critical for interpreting population dynamics, seasonal interactions, and for the implementation of conservation strategies of migratory species. We evaluated the migratory connectivity of a Neotropical migratory songbird, the Ovenbird (Seiurus aurocapilla) using archival light-level geolocators deployed at two breeding and four nonbreeding locations while incorporating Ovenbird abundance as prior information using Bayes' Rule. We also included band recoveries submitted to the United States Geological Survey's Bird Banding Laboratory to assess connectivity of areas where geolocators were not deployed. We created a probabilistic map of origin for each capture site and mapped spring migration routes between nonbreeding and breeding locations. We found a complete separation of eastern and western populations of Ovenbirds throughout the annual cycle. Breeding Ovenbirds from western Canada spent the nonbreeding season throughout Central America and migrated through central North America during spring migration. Birds breeding in the northeastern United States were distributed throughout the central Greater Antilles in the Caribbean and migrated through eastern North America during spring migration. Fall migration routes were not included because the timing of migration coincided with fall equinox when latitudinal estimates are unreliable. However, longitudinal estimates suggest no overlap between eastern and western populations during fall migration. Ovenbirds with geolocators attached in Jamaica bred in the northeastern United States with the highest posterior probability of origin found in Massachusetts, while Ovenbirds captured in Florida and Puerto Rico bred primarily in the mid-Atlantic. Incorporating Ovenbird abundance as a prior into geolocator estimates decreased the area of origin by 90.37% ± 1.05% (mean ± SE) for the breeding season and 62.30% ± 1.69% for the nonbreeding season, compared to geolocator estimates alone. Ovenbirds exhibited strong migratory connectivity between breeding and nonbreeding season, which has important implications for various aspects of the ecology, evolution, and conservation.

Conservation through connectivity: can isotopic gradients in Africa reveal winter quarters of a migratory bird?

Conservation of migratory wildlife requires knowledge of migratory connectivity between breeding and non-breeding locations. Stable isotopes in combination with geographical isotopic patterns (isoscapes) can provide inferences about migratory connectivity. This study examines whether such an approach can be used to infer wintering areas in sub-Saharan Africa, where we lack such knowledge for many species, but where this method has not been used widely. We measured δ (2)H, δ (13)C and δ (15)N in winter-grown feathers of a breeding Swiss and Spanish population of European hoopoe Upupa epops--a typical Palaearctic-Afrotropical migrant. δ (2)H values predicted that ~70 % of the hoopoes spent the non-breeding season in the western portion of their potential winter range. This was corroborated by a shallow east-west gradient in feather-δ (2)H values of museum specimens from known African origin across the potential winter range and by the recovery of Swiss hoopoes marked with geolocators. Hoopoes categorized as from eastern versus western regions of the wintering range were further delineated spatially using feather δ (13)C and δ (15)N. δ (15)N showed no trend, whereas adults were more enriched in (13)C in the western portion of the range, with eastern adults being in addition more depleted in (13)C than eastern juveniles. This suggests that eastern juveniles may have occupied more xeric habitats than sympatric adults. We demonstrated that stable isotopes, especially δ (2)H, could only very roughly delineate the winter distribution of a trans-Saharan Palaearctic migrant restricted primarily to the Sahelian and savanna belt south of the Sahara. Further refinements of precipitation isoscapes for Africa as well the development of isoscapes for δ (13)C and δ (15)N may improve assignment of this and other migrants.

Positive relationships between association strength and phenotypic similarity characterize the assembly of mixed-species bird flocks worldwide.

Competition theory predicts that local communities should consist of species that are more dissimilar than expected by chance. We find a strikingly different pattern in a multicontinent data set (55 presence-absence matrices from 24 locations) on the composition of mixed-species bird flocks, which are important subunits of local bird communities the world over. By using null models and randomization tests followed by meta-analysis, we find the association strengths of species in flocks to be strongly related to similarity in body size and foraging behavior and higher for congeneric compared with noncongeneric species pairs. Given the local spatial scales of our individual analyses, differences in the habitat preferences of species are unlikely to have caused these association patterns; the patterns observed are most likely the outcome of species interactions. Extending group-living and social-information-use theory to a heterospecific context, we discuss potential behavioral mechanisms that lead to positive interactions among similar species in flocks, as well as ways in which competition costs are reduced. Our findings highlight the need to consider positive interactions along with competition when seeking to explain community assembly.

Combining stable-isotope (deltaD) and band recovery data to improve probabilistic assignment of migratory birds to origin.

The recent application of stable-isotope analyses, particularly the use of stable-hydrogen-isotope (deltaD) measurements of animal tissues, has greatly improved our ability to infer geographic origins of migratory animals. However, many individual sources of error contribute to the overall error in assignment; thus likelihood-based assignments incorporating estimates of error are now favored. In addition, globally, the nature of the underlying precipitation-based deltaD isoscapes is such that longitudinal resolution is often compromised. For example, in North America, amount-weighted expected mean growing-season precipitation deltaD is similar between the boreal forest of southwestern Canada and areas of northern Quebec/Labrador and Alaska. Thus, it can often be difficult to distinguish objectively between these areas as potential origins for broadly distributed migrants using a single isotopic measurement. We developed a Bayesian framework for assigning geographic origins to migrant birds based on combined stable-isotope analysis of feathers and models of migratory directions estimated from band recovery data. We outline our method and show an example of its application for assigning origins to a population of migrant White-throated Sparrows (Zonotrichia albicollis) sampled at a Canadian Migration Monitoring Network station at Delta Marsh, Manitoba, Canada. We show that likelihood-based assignments of geographic origins can provide improved spatial resolution when models of migration direction are combined with assignments based on deltaD analysis of feathers.

A cost efficient spatially balanced hierarchical sampling design for monitoring boreal birds incorporating access costs and habitat stratification.

Predicting and mitigating impacts of climate change and development within the boreal biome requires a sound understanding of factors influencing the abundance, distribution, and population dynamics of species inhabiting this vast biome. Unfortunately, the limited accessibility of the boreal biome has resulted in sparse and spatially biased sampling, and thus our understanding of boreal bird population dynamics is limited. To implement effective conservation of boreal birds, a cost-effective approach to sampling the boreal biome will be needed. Our objective was to devise a sampling scheme for monitoring boreal birds that would improve our ability to model species-habitat relationships and monitor changes in population size and distribution. A statistically rigorous design to achieve these objectives would have to be spatially balanced and hierarchically structured with respect to ecozones, ecoregions and political jurisdictions. Therefore, we developed a multi-stage hierarchically structured sampling design known as the Boreal Optimal Sampling Strategy (BOSS) that included cost constraints, habitat stratification, and optimization to provide a cost-effective alternative to other common monitoring designs. Our design provided similar habitat and spatial representation to habitat stratification and equal-probability spatially balanced designs, respectively. Not only was our design able to achieve the desired habitat representation and spatial balance necessary to meet our objectives, it was also significantly less expensive (1.3-2.6 times less) than the alternative designs we considered. To further balance trade-offs between cost and representativeness prior to field implementation, we ran multiple iterations of the BOSS design and selected the one which minimized predicted costs while maximizing a multi-criteria evaluation of representativeness. Field implementation of the design in three vastly different regions over three field seasons showed that the approach can be implemented in a wide variety of logistical scenarios and ecological conditions. We provide worked examples and scripts to allow our approach to be implemented or adapted elsewhere. We also provide recommendations for possible future refinements to our approach, but recommend that our design now be implemented to provide unbiased information to assess the status of boreal birds and inform conservation and management actions.

Migratory connectivity in the Loggerhead Shrike (Lanius ludovicianus).

AIM: We combine genetic and stable isotope data to quantify migration patterns in Loggerhead Shrike (Lanius ludovicianus), a species of conservation concern in North America, to assess how connectivity differs and impacts population evolution, ecology, and conservation. LOCATION: We sampled shrikes across the majority of their nonbreeding range, from the Atlantic Coast to the western United States east of the Rocky Mountains and throughout Mexico. METHODS: Our study used a Bayesian framework using δ2Hf from a breeding season origin feather and nuclear genetic microsatellite markers to distinguish between co-occurring migratory and nonmigratory individuals on the wintering grounds and, for migrants, to assign individuals to a breeding ground origin and genetic group. RESULTS: Migratory shrikes were present throughout the nonbreeding range but the proportion differed among sample areas. Four main wintering areas were identified. Connectivity ranged from weakly negative in birds wintering on the Atlantic Coast to strongly positive between wintering grounds in the southwestern United States and Mexico and northwestern breeding populations. Connectivity was weakest in L. l. migrans, and strongest in L. l. mexicanus and L. l. excubitorides. Although believed to be nonmigratory, long-distance movements of individuals were observed in L. ludovicianus and L. l. mexicanus. Our data support a pattern of chain migration, again most notable in the western half of the species nonbreeding range, and differential migration based on age. MAIN CONCLUSIONS: Our study provides of one such of the first quantitative measures of migratory connectivity and is among the first studies of a short-distance migratory passerine in North America. The higher migratory connectivity among western, versus eastern populations, and less severe population declines attributable to habitat loss or reproductive success, may result in more localized and/or less severe limiting factors for western populations and more severe on the Atlantic coast and Mississippi Alluvial Valley wintering grounds.

Multi-Isotopic (δ2H, δ13C, δ15N) Tracing of Molt Origin for Red-Winged Blackbirds Associated with Agro-Ecosystems.

We analyzed stable-hydrogen (δ2H), carbon (δ13C) and nitrogen (δ 15N) isotope ratios in feathers to better understand the molt origin and food habits of Red-winged Blackbirds (Agelaius phoeniceus) near sunflower production in the Upper Midwest and rice production in the Mid-South of the United States. Outer primary feathers were used from 661 after-second-year (ASY) male blackbirds collected in Minnesota, Montana, North Dakota and South Dakota (spring collection), and Arkansas, Louisiana, Mississippi, Missouri and Texas (winter collection). The best-fit model indicated that the combination of feather δ2H, δ13C and δ15N best predicted the state of sample collections and thus supported the use of this approach for tracing molt origins in Red-winged Blackbirds. When considering only birds collected in spring, 56% of birds were classified to their collection state on the basis of δ2H and δ13C alone. We then developed feather isoscapes for δ13C based upon these data and for δ2H based upon continental patterns of δ2H in precipitation. We used 81 birds collected at the ten independent sites for model validation. The spatially-explicit assignment of these 81 birds to the δ2H isoscape resulted in relatively high rates (~77%) of accurate assignment to collection states. We also modeled the spatial extent of C3 (e.g. rice, sunflower) and C4 (corn, millet, sorghum) agricultural crops grown throughout the Upper Midwest and Mid-South United States to predict the relative use of C3- versus C4-based foodwebs among sampled blackbirds. Estimates of C3 inputs to diet ranged from 50% in Arkansas to 27% in Minnesota. As a novel contribution to blackbird conservation and management, we demonstrate how such feather isoscapes can be used to predict the molt origin and interstate movements of migratory blackbirds for subsequent investigations of breeding biology (e.g. sex-specific philopatry), agricultural depredation, feeding ecology, physiology of migration and sensitivity to environmental change.

A Continent-Wide Migratory Divide in North American Breeding Barn Swallows (Hirundo rustica).

Populations of most North American aerial insectivores have undergone steep population declines over the past 40 years but the relative importance of factors operating on breeding, wintering, or stopover sites remains unknown. We used archival light-level geolocators to track the phenology, movements and winter locations of barn swallows (Hirdundo rustica; n = 27) from populations across North America to determine their migratory connectivity. We identified an east-west continental migratory divide for barn swallows with birds from western regions (Washington State, USA (n = 8) and Saskatchewan, Canada (n = 5)) traveling shorter distances to wintering areas ranging from Oregon to northern Colombia than eastern populations (Ontario (n = 3) and New Brunswick (n = 10), Canada) which wintered in South America south of the Amazon basin. A single swallow from a stable population in Alabama shared a similar migration route to eastern barn swallows but wintered farther north in northeast Brazil indicating a potential leap frog pattern migratory among eastern birds. Six of 9 (67%) birds from the two eastern populations and Alabama underwent a loop migration west of fall migration routes including around the Gulf of Mexico travelling a mean of 2,224 km and 722 km longer on spring migration, respectively. Longer migration distances, including the requirement to cross the Caribbean Sea and Gulf of Mexico and subsequent shorter sedentary wintering periods, may exacerbate declines for populations breeding in northeastern North America.

Tracking natal dispersal in a coastal population of a migratory songbird using feather stable isotope (δ2H, δ34S) tracers.

Adult birds tend to show high fidelity to their breeding territory or disperse over relatively short distances. Gene flow among avian populations is thus expected to occur primarily through natal dispersal. Although natal dispersal is a critical demographic process reflecting the area over which population dynamics take place, low recapture rates of birds breeding for the first time have limited our ability to reliably estimate dispersal rates and distances. Stable isotope approaches can elucidate origins of unmarked birds and so we generated year- and age-specific δ2H and δ34S feather isoscapes (ca. 180 000 km2) of coastal-breeding Ovenbirds (Seiurus aurocapilla) and used bivariate probability density functions to assign the likely natal areas of 35 males recruited as first-year breeders into a population located in northwestern New Brunswick, Canada. Most individuals (80-94% depending on the magnitude of an age correction factor used; i.e. 28-33 out of 35) were classified as residents (i.e. fledged within our study area) and estimated minimum dispersal distances of immigrants were between 40 and 240 km. Even when considering maximum dispersal distances, the likely origin of most first-year breeders was<200 km from our study area. Our method identified recruitment into our population from large geographic areas with relatively few samples whereas previous mark-recapture based methods have required orders of magnitude more individuals to describe dispersal at such geographic scales. Natal dispersal movements revealed here suggest the spatial scale over which many population processes are taking place and we suggest that conservation plans aiming to maintain populations of Ovenbirds and ecologically-similar species should consider management units within 100 or at most 200 km of target breeding populations.

Identifying the African wintering grounds of hybrid flycatchers using a multi-isotope (δ2H, δ13C, δ15N) assignment approach.

Migratory routes and wintering grounds can have important fitness consequences, which can lead to divergent selection on populations or taxa differing in their migratory itinerary. Collared (Ficedula albicollis) and pied (F. hypoleuca) flycatchers breeding in Europe and wintering in different sub-Saharan regions have distinct migratory routes on the eastern and western sides of the Sahara desert, respectively. In an earlier paper, we showed that hybrids of the two species did not incur reduced winter survival, which would be expected if their migration strategy had been a mix of the parent species' strategies potentially resulting in an intermediate route crossing the Sahara desert to different wintering grounds. Previously, we compared isotope ratios and found no significant difference in stable-nitrogen isotope ratios (δ15N) in winter-grown feathers between the parental species and hybrids, but stable-carbon isotope ratios (δ13C) in hybrids significantly clustered only with those of pied flycatchers. We followed up on these findings and additionally analyzed the same feathers for stable-hydrogen isotope ratios (δ2H) and conducted spatially explicit multi-isotope assignment analyses. The assignment results overlapped with presumed wintering ranges of the two species, highlighting the efficacy of the method. In contrast to earlier findings, hybrids clustered with both parental species, though most strongly with pied flycatcher.

Solving a migration riddle using isoscapes: house martins from a Dutch village winter over West Africa.

BACKGROUND: The ability to connect breeding, stopover and wintering locations of populations of migratory birds greatly enhances our understanding of the phenomenon of migration and improves our chances of effectively conserving these species. Among Palearctic-Afrotropical migratory species, aerial insectivores like the house martin (Delichon urbicum) are sensitive to factors influencing the availability of flying insects, and have declined in recent decades. The strict aerial behaviour of martins severely limits ring recoveries on wintering grounds and so there is a dearth of information on where European breeding populations over-winter in Africa, and the relative effects of population regulation on breeding vs. wintering grounds. We used a newly developed multi-isotope (δ(2)H, δ(13)C, δ(15)N) feather isoscape for Africa together with inferences from summarized ring return data based on longitude, to assign winter origins to birds captured at a breeding colony in The Netherlands. PRINCIPAL FINDINGS: Based on isotopic analyses of winter-grown martin feathers, we used a likelihood-based assignment approach to describe potential wintering locations where molt occurred of individual house martins from a Dutch colony by assigning them to four potential isotopically distinct clusters in Africa. We found the overwhelming majority of Dutch martins were assigned to a geographical cluster associated with West Africa. CONCLUSIONS/SIGNIFICANCE: The existence of strong isotopic gradients and patterns in African foodwebs that support migratory wildlife allows for the spatial assignment of tissues grown there. The assignment of Dutch house martins to wintering grounds primarily in West Africa was in strong agreement with independent and indirect methods used to infer winter origins of this species based on the association between the Normalized Difference Vegetation Index (NDVI) in Africa and population patterns in Italy and the United Kingdom. These confirmatory data-sets underscore the importance of suitable habitats in West Africa to the conservation of migratory aerial insectivores and other species.

Linking hydrogen (δ2H) isotopes in feathers and precipitation: sources of variance and consequences for assignment to isoscapes.

BACKGROUND: Tracking small migrant organisms worldwide has been hampered by technological and recovery limitations and sampling bias inherent in exogenous markers. Naturally occurring stable isotopes of H (δ(2)H) in feathers provide an alternative intrinsic marker of animal origin due to the predictable spatial linkage to underlying hydrologically driven flow of H isotopes into foodwebs. This approach can assess the likelihood that a migrant animal originated from a given location(s) within a continent but requires a robust algorithm linking H isotopes in tissues of interest to an appropriate hydrological isotopic spatio-temporal pattern, such as weighted-annual rainfall. However, a number of factors contribute to or alter expected isotopic patterns in animals. We present results of an extensive investigation into taxonomic and environmental factors influencing feather δ(2)H patterns across North America. PRINCIPAL FINDINGS: Stable isotope data were measured from 544 feathers from 40 species and 140 known locations. For δ(2)H, the most parsimonious model explaining 83% of the isotopic variance was found with amount-weighted growing-season precipitation δ(2)H, foraging substrate and migratory strategy. CONCLUSIONS/SIGNIFICANCE: This extensive H isotopic analysis of known-origin feathers of songbirds in North America and elsewhere reconfirmed the strong coupling between tissue δ(2)H and global hydrologic δ(2)H patterns, and accounting for variance associated with foraging substrate and migratory strategy, can be used in conservation and research for the purpose of assigning birds and other species to their approximate origin.

Advances in linking wintering migrant birds to their breeding-ground origins using combined analyses of genetic and stable isotope markers.

An enduring problem in avian ecology and conservation is linking breeding and wintering grounds of migratory species. As migratory species and populations vary in the degree to which individuals from distinct breeding locales mix on stop-over sites and wintering grounds, establishing migratory connectivity informs our understanding of population demography and species management. We present a new Bayesian approach for inferring breeding grounds of wintering birds of unknown origins in North America. We incorporate prior information from analysis of genetic markers into geographic origin assignment based upon stable-hydrogen isotope analysis of feathers (δ(2)H(f)), using the Loggerhead Shrike (Lanius ludovicianus). Likely geographic origins derived from analyses of DNA microsatellites were used as priors for Bayesian analyses in which birds were assigned to a breeding-ground origin using their δ(2)H(f) values. As with most applications of Bayesian methods, our approach greatly improved the results (i.e. decreased the size of the potential area of origin). Area of origin decreased by 3 to 5-fold on average, but ranged up to a 10-fold improvement. We recommend this approach in future studies of migratory connectivity and suggest that our methodology could be applied more broadly to the study of dispersal, sources of productivity of migratory populations, and a range of evolutionary phenomena.

Disease dynamics and bird migration--linking mallards Anas platyrhynchos and subtype diversity of the influenza A virus in time and space.

The mallard Anas platyrhynchos is a reservoir species for influenza A virus in the northern hemisphere, with particularly high prevalence rates prior to as well as during its prolonged autumn migration. It has been proposed that the virus is brought from the breeding grounds and transmitted to conspecifics during subsequent staging during migration, and so a better understanding of the natal origin of staging ducks is vital to deciphering the dynamics of viral movement pathways. Ottenby is an important stopover site in southeast Sweden almost halfway downstream in the major Northwest European flyway, and is used by millions of waterfowl each year. Here, mallards were captured and sampled for influenza A virus infection, and positive samples were subtyped in order to study possible links to the natal area, which were determined by a novel approach combining banding recovery data and isotopic measurements (δ(2)H) of feathers grown on breeding grounds. Geographic assignments showed that the core natal areas of studied mallards were in Estonia, southern and central Finland, and northwestern Russia. This study demonstrates a clear temporal succession of latitudes of natal origin during the course of autumn migration. We also demonstrate a corresponding and concomitant shift in virus subtypes. Acknowledging that these two different patterns were based in part upon different data, a likely interpretation worth further testing is that the early arriving birds with more proximate origins have different influenza A subtypes than the more distantly originating late autumn birds. If true, this knowledge would allow novel insight into the origins and transmission of the influenza A virus among migratory hosts previously unavailable through conventional approaches.

A method for investigating population declines of migratory birds using stable isotopes: origins of harvested lesser scaup in North America.

BACKGROUND: Elucidating geographic locations from where migratory birds are recruited into adult breeding populations is a fundamental but largely elusive goal in conservation biology. This is especially true for species that breed in remote northern areas where field-based demographic assessments are logistically challenging. METHODOLOGY/FINDINGS: Here we used hydrogen isotopes (deltaD) to determine natal origins of migrating hatch-year lesser scaup (Aythya affinis) harvested by hunters in the United States from all North American flyways during the hunting seasons of 1999-2000 (n = 412) and 2000-2001 (n = 455). We combined geospatial, observational, and analytical data sources, including known scaup breeding range, deltaD values of feathers from juveniles at natal sites, models of deltaD for growing-season precipitation, and scaup band-recovery data to generate probabilistic natal origin landscapes for individual scaup. We then used Monte Carlo integration to model assignment uncertainty from among individual deltaD variance estimates from birds of known molt origin and also from band-return data summarized at the flyway level. We compared the distribution of scaup natal origin with the distribution of breeding population counts obtained from systematic long-term surveys. CONCLUSIONS/SIGNIFICANCE: Our analysis revealed that the proportion of young scaup produced in the northern (above 60 degrees N) versus the southern boreal and Prairie-Parkland region was inversely related to the proportions of breeding adults using these regions, suggesting that despite having a higher relative abundance of breeding adults, the northern boreal region was less productive for scaup recruitment into the harvest than more southern biomes. Our approach for evaluating population declines of migratory birds (particularly game birds) synthesizes all available distributional data and exploits the advantages of intrinsic isotopic markers that link individuals to geography.