Non-breeding conditions induce carry-over effects on survival of migratory birds.

Identifying the processes that limit populations is a foundational objective of ecology and an urgent need for conservation. For migratory animals, researchers must study individuals throughout their annual cycles to determine how environmental conditions limit demographic rates within each period of the annual cycle and also between periods through carry-over effects and seasonal interactions.1,2,3,4,5,6 Our poor understanding of the rates and causes of avian migration mortality7 hinders the identification of limiting factors and the reversal of widespread avian population declines.8,9 Here, we implement new methods to estimate apparent survival (hereafter survival) during migration directly from automated telemetry data10 in Kirtland's Warblers (Setophaga kirtlandii) and indirectly from mark-recapture data in Black-throated Blue Warblers (S. caerulescens). Previous experimental and observational studies of our focal species and other migratory songbirds have shown strong effects of Caribbean precipitation and habitat quality on food availability,11,12,13,14 body condition,12,13,14,15,16,17,18,19 migration timing,11,12,15,16,20,21,22,23 natal dispersal,24,25 range dynamics,26 reproductive success,20,22,27 and annual survival.18,19,20,23,28,29,30,31 Building on this research, we test the hypotheses that environmental conditions during the non-breeding period affect subsequent survival during spring migration and breeding. We found that reduced precipitation and environmental productivity in the non-breeding period strongly influenced survival in both species, primarily by reducing survival during spring migration. Our results indicate that climate-driven environmental conditions can carry over to affect survival in subsequent periods and thus likely play an important role in year-round population dynamics. These lethal carry-over effects may be widespread and are likely magnified by intensifying climate change.

Feather carotenoids of the American Redstart (Setophaga ruticilla) across age and sex classes and the reliability of standard color metrics to capture pigment variation.

Plumage ornaments act as important sexual signals, though the extent to which these ornaments act as honest signals-and the physiological mechanisms that maintain honesty-remain poorly understood. We studied the pigmentary basis of tail color in the American Redstart (Setophaga ruticilla), a species of songbird with sexual dichromatism and delayed plumage maturation; younger males resemble females, only replacing their yellow feathers for bright orange ones after the first breeding season. The yellow rectrices of females and young males and the orange feathers of older males largely contain the same pigments, but in vastly different proportions. Whereas the feathers of females and young males contain primarily lutein, 3'-dehydro-lutein and canary-xanthophylls, those of older males contain primarily 4-keto-carotenoids. The presence of lutein and the predominance of α-doradexanthin as 4-keto-carotenoid, a pigment with a shortened chain of conjugated double bonds compared to keto-carotenoids commonly found in red feathers, in the feathers of older males contribute to their uncommon orange hue. Since the orange coloration of the tail in the American redstart results from the combination of yellow, orange, and red pigments, this is a system where slight adjustments in the types of carotenoids deposited could significantly alter hue. Factors either work against achieving the most oxidized state in this species or the hue is maintained through stabilizing selection for a favored color. The color metrics of Carotenoid Chroma, Visible Hue, λR50 and tetrahedral θ best captured differences in pigment concentration and make-up, and are recommended in future spectrophotometric studies of carotenoid-based traits.

Museum genomics provide insight into the extinction of a specialist North American warbler species.

Museum genomics provide an opportunity to investigate population demographics of extinct species, especially valuable when research prior to extinction was minimal. The Bachman's warbler (Vermivora bachmanii) is hypothesized to have gone extinct due to loss of its specialized habitat. However, little is known about other potential contributing factors such as natural rarity or changes to connectivity following habitat fragmentation. We examined mitochondrial DNA (mtDNA) and genome-wide SNPs using specimens collected from breeding and migration sites across the range of the Bachman's warbler. We found no signals of strong population structuring across the breeding range of Bachman's warblers in both mtDNA and genome-wide SNPs. Thus, long-term population isolation did not appear to be a significant contributor to the extinction of the Bachman's warbler. Instead, our findings support the theory that Bachman's warblers underwent a rapid decline likely driven by habitat destruction, which may have been exacerbated by the natural rarity, habitat specificity and low genetic diversity of the species.

Extreme elevational migration spurred cryptic speciation in giant hummingbirds.

The ecoevolutionary drivers of species niche expansion or contraction are critical for biodiversity but challenging to infer. Niche expansion may be promoted by local adaptation or constrained by physiological performance trade-offs. For birds, evolutionary shifts in migratory behavior permit the broadening of the climatic niche by expansion into varied, seasonal environments. Broader niches can be short-lived if diversifying selection and geography promote speciation and niche subdivision across climatic gradients. To illuminate niche breadth dynamics, we can ask how "outlier" species defy constraints. Of the 363 hummingbird species, the giant hummingbird (Patagona gigas) has the broadest climatic niche by a large margin. To test the roles of migratory behavior, performance trade-offs, and genetic structure in maintaining its exceptional niche breadth, we studied its movements, respiratory traits, and population genomics. Satellite and light-level geolocator tracks revealed an >8,300-km loop migration over the Central Andean Plateau. This migration included a 3-wk, ~4,100-m ascent punctuated by upward bursts and pauses, resembling the acclimatization routines of human mountain climbers, and accompanied by surging blood-hemoglobin concentrations. Extreme migration was accompanied by deep genomic divergence from high-elevation resident populations, with decisive postzygotic barriers to gene flow. The two forms occur side-by-side but differ almost imperceptibly in size, plumage, and respiratory traits. The high-elevation resident taxon is the world's largest hummingbird, a previously undiscovered species that we describe and name here. The giant hummingbirds demonstrate evolutionary limits on niche breadth: when the ancestral niche expanded due to evolution (or loss) of an extreme migratory behavior, speciation followed.

Springing forward: Migrating songbirds catch up with the start of spring in North America.

In temperate regions, the annual pattern of spring onset can be envisioned as a 'green wave' of emerging vegetation that moves across continents from low to high latitudes, signifying increasing food availability for consumers. Many herbivorous migrants 'surf' such resource waves, timing their movements to exploit peak vegetation resources in early spring. Although less well studied at the individual level, secondary consumers such as insectivorous songbirds can track vegetation phenology during migration as well. We hypothesized that four species of ground-foraging songbirds in eastern North America-two warblers and two thrushes-time their spring migrations to coincide with later phases of vegetation phenology, corresponding to increased arthropod prey, and predicted they would match their migration rate to the green wave but trail behind it rather than surfing its leading edge. We further hypothesized that the rate at which spring onset progresses across the continent influences bird migration rates, such that individuals adjust migration timing within North America to phenological conditions they experience en route. To test our hypotheses, we used a continent-wide automated radio telemetry network to track individual songbirds on spring migration between the U.S. Gulf Coast region and northern locations closer to their breeding grounds. We measured vegetation phenology using two metrics of spring onset, the spring index first leaf date and the normalized difference vegetation index (NDVI), then calculated the rate and timing of spring onset relative to bird detections. All individuals arrived in the southeastern United States well after local spring onset. Counter to our expectations, we found that songbirds exhibited a 'catching up' pattern: Individuals migrated faster than the green wave of spring onset, effectively closing in on the start of spring as they approached breeding areas. While surfing of resource waves is a well-documented migration strategy for herbivorous waterfowl and ungulates, individual songbirds in our study migrated faster than the green wave and increasingly caught up to its leading edge en route. Consequently, songbirds experience a range of vegetation phenophases while migrating through North America, suggesting flexibility in their capacity to exploit variable resources in spring.

En las regiones templadas, el patrón anual de inicio de la primavera puede concebirse como una "ola verde" de vegetación emergente que se desplaza por los continentes desde las latitudes bajas a las altas, lo que significa una mayor disponibilidad de alimento para los consumidores. Muchos herbívoros migratorios "surfean" estas olas de recursos, programando sus movimientos para aprovechar los picos de vegetación a principios de primavera. Aunque menos estudiados a nivel de individuo, los consumidores secundarios, como las aves terrestres insectívoras, también pueden seguir la fenología de la vegetación durante la migración. Hipotetizamos es que cuatro especies de aves terrestres que se alimentan en el suelo en el este de Norteamérica - dos reinitas y dos zorzales - programan sus migraciones primaverales para que coincidan con las fases más tardías de la fenología de la vegetación, que se corresponden con un aumento de artrópodos, y predijimos que sincronizarian su ritmo de migración con la ola verde, pero que irían detrás de ella en lugar de surfear su borde delantero. También hipotetizamos que el ritmo al que avanza la primavera en el continente influye en las tasas de migración de las aves, de modo que los individuos ajustan la fecha de migración dentro de Norteamérica a las condiciones fenológicas que experimentan en ruta. Para comprobar nuestras hipótesis, utilizamos una red automatizada de radiotelemetría a escala continental para seguir individuos en su migración primaveral entre la región de la costa del Golfo de EEUU y las localidades septentrionales más cercanas a sus zonas de cría. Medimos la fenología de la vegetación utilizando dos métricas del inicio de la primavera, el índice de la fecha de la primera hoja primaveral y el índice de vegetación de diferencia normalizada (NDVI), luego calculamos la tasa y el tiempo de la aparaciòn de la primavera relativo a las detecciones de aves. Todos los individuos llegaron al sureste de EEUU bastante después del inicio de la primavera local. Contrario a nuestras expectativas, descubrimos que las aves terrestres mostraron un patrón de Carrera para "ponerse al día": los individuos migraron frente a la ola verde del inicio de la primavera, acercándose efectivamente al inicio de la primavera a medida que llegaban a las zonas de cría. Mientras que el surfing de las olas de recursos es una estrategia migratoria bien documentada para las aves acuáticas herbívoras y los ungulados, los individuos de aves terrestres de nuestro estudio migraron más rápido que la ola verde y alcanzaron cada vez más el borde delantero en ruta. En consecuencia, las aves terrestres experimentan una serie de fases fenológicas de la vegetación mientras migran a través de Norteamérica, lo que sugiere flexibilidad en su capacidad para explotar recursos variables en primavera.

The role of tropical rainfall in driving range dynamics for a long-distance migratory bird.

Predicting how the range dynamics of migratory species will respond to climate change requires a mechanistic understanding of the factors that operate across the annual cycle to control the distribution and abundance of a species. Here, we use multiple lines of evidence to reveal that environmental conditions during the nonbreeding season influence range dynamics across the life cycle of a migratory songbird, the American redstart (Setophaga ruticilla). Using long-term data from the nonbreeding grounds and breeding origins estimated from stable hydrogen isotopes in tail feathers, we found that the relationship between annual survival and migration distance is mediated by precipitation, but only during dry years. A long-term drying trend throughout the Caribbean is associated with higher mortality for individuals from the northern portion of the species' breeding range, resulting in an approximate 500 km southward shift in breeding origins of this Jamaican population over the past 30 y. This shift in connectivity is mirrored by changes in the redstart's breeding distribution and abundance. These results demonstrate that the climatic effects on demographic processes originating during the tropical nonbreeding season are actively shaping range dynamics in a migratory bird.

Low-coverage whole genome sequencing for highly accurate population assignment: Mapping migratory connectivity in the American Redstart (Setophaga ruticilla).

Understanding the geographic linkages among populations across the annual cycle is an essential component for understanding the ecology and evolution of migratory species and for facilitating their effective conservation. While genetic markers have been widely applied to describe migratory connections, the rapid development of new sequencing methods, such as low-coverage whole genome sequencing (lcWGS), provides new opportunities for improved estimates of migratory connectivity. Here, we use lcWGS to identify fine-scale population structure in a widespread songbird, the American Redstart (Setophaga ruticilla), and accurately assign individuals to genetically distinct breeding populations. Assignment of individuals from the nonbreeding range reveals population-specific patterns of varying migratory connectivity. By combining migratory connectivity results with demographic analysis of population abundance and trends, we consider full annual cycle conservation strategies for preserving numbers of individuals and genetic diversity. Notably, we highlight the importance of the Northern Temperate-Greater Antilles migratory population as containing the largest proportion of individuals in the species. Finally, we highlight valuable considerations for other population assignment studies aimed at using lcWGS. Our results have broad implications for improving our understanding of the ecology and evolution of migratory species through conservation genomics approaches.

Songbirds initiate migratory flights synchronously relative to civil dusk.

BACKGROUND: Each spring and fall billions of songbirds depart on nocturnal migrations across the globe. Theory suggests that songbirds should depart on migration shortly after sunset to maximize their potential for nightly flight duration or to time departure with the emergence of celestial cues needed for orientation and navigation. Although captive studies have found that songbirds depart during a narrow window of time after sunset, observational studies have found that wild birds depart later and more asynchronously relative to sunset than predicted. METHODS: We used coded radio tags and automated radio-telemetry to estimate the time that nearly 400 individuals from nine songbird species departed their breeding or wintering grounds across North America. We also assessed whether each species was most likely beginning long-distance migratory flights at departure or instead first making non-migratory regional flights. We then explored variation in nocturnal departure time by post-departure movement type, species, age, sex, and season. RESULTS: We found that 90% of individuals from species that were likely initiating long-distance migratory flights departed within 69 min of civil dusk, regardless of species, season, age, or sex. By contrast, species that likely first made non-migratory regional movements away from the migratory destination departed later and more asynchronously throughout the night. Regardless of post-departure movement type, 98% of individuals departed after civil dusk but otherwise showed no preference in relation to twilight phase. CONCLUSIONS: Although the presence of celestial orientation cues at civil dusk may set a starting point for departure each night, the fact that species likely beginning long-distance migration departed earlier and more synchronously relative to civil dusk than those first making non-migratory regional movements is consistent with the hypothesis that departing promptly after civil dusk functions to maximize the potential for nightly flight duration and distance. By studying the onset of migration, our study provides baseline information about departure decisions that may enhance our understanding of departure timing throughout migration.

Atmospheric pressure predicts probability of departure for migratory songbirds.

BACKGROUND: Weather can have both delayed and immediate impacts on animal populations, and species have evolved behavioral adaptions to respond to weather conditions. Weather has long been hypothesized to affect the timing and intensity of avian migration, and radar studies have demonstrated strong correlations between weather and broad-scale migration patterns. How weather affects individual decisions about the initiation of migratory flights, particularly at the beginning of migration, remains uncertain. METHODS: Here, we combine automated radio telemetry data from four species of songbirds collected at five breeding and wintering sites in North America with hourly weather data from a global weather model. We use these data to determine how wind profit, atmospheric pressure, precipitation, and cloud cover affect probability of departure from breeding and wintering sites. RESULTS: We found that the probability of departure was related to changes in atmospheric pressure, almost completely regardless of species, season, or location. Individuals were more likely to depart on nights when atmospheric pressure had been rising over the past 24 h, which is predictive of fair weather over the next several days. By contrast, wind profit, precipitation, and cloud cover were each only informative predictors of departure probability in a single species. CONCLUSIONS: Our results suggest that individual birds actively use weather information to inform decision-making regarding the initiation of departure from the breeding and wintering grounds. We propose that birds likely choose which date to depart on migration in a hierarchical fashion with weather not influencing decision-making until after the departure window has already been narrowed down by other ultimate and proximate factors.

Migratory birds with delayed spring departure migrate faster but pay the costs.

Migratory birds that experience poor overwintering conditions are often late to arrive at the breeding grounds, which is known to depress individual fitness. Despite the importance of this carryover effect, few studies have investigated how individuals can modify migratory behaviors en route to reduce delays on arrival and whether accelerating migration incurs survival costs. To examine this, we used Motus Wildlife Tracking System to track individual American redstarts (Setophaga ruticilla) as they migrated from wintering grounds in Southwest Jamaica through Florida en route to their breeding areas. We leveraged long-term data on spring departure timing and breeding latitude to quantify the relative departure dates (early vs. delayed) of tagged individuals, which we then related to individual migration rates and apparent annual survival. Compared to those initiating migration earlier, individuals that departed relatively late (10-day delay) migrated at a 43% faster rate, which decreased their annual survival by 6.3%. Our results are consistent with the hypothesis that spring migrants use speed to compensate for departure delays despite incurring survival costs. This compensatory behavior may potentially underly differential survival during spring migration and may be particularly widespread across short-lived migratory birds generally considered time-constrained.

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.

Niche dynamics suggest ecological factors influencing migration in an insectivorous owl.

Seasonal migration is a widespread phenomenon undertaken by myriad organisms, including birds. Competing hypotheses about ultimate drivers of seasonal migration in birds contrast relative resource abundances at high latitudes ("southern home hypothesis") against avoidance of winter resource scarcity ("dispersal-migration hypothesis"). However, direct tests of these competing hypotheses have been rare and to date limited to historical biogeographic reconstructions. Here we derive novel predictions about the dynamics of individual niches from each hypothesis and provide a framework for evaluating support for these competing hypotheses using contemporary environmental and behavioral data. Using flammulated owls (Psiloscops flammeolus) as a model, we characterized year-round occupied niche dynamics using high-resolution global positioning system tracking and remote-sensed environmental data. We also compared occupied niche dynamics to counterfactual niches using simulated alternative nonmigratory strategies. Owl occupied mean niche was conserved among seasons, whereas niche variance was generally higher during migratory periods. Simulated year-round residents in Mexico would have experienced putatively more productive niches than migrants. These findings provide ecological support for the "dispersal-migration" hypothesis in which winter resource scarcity is the primary driver of migration rather than summer resource abundances.

Impact of cocoa agricultural intensification on bird diversity and community composition.

To meet the growing demand for chocolate, cocoa (Theobroma cacao) agriculture is expanding and intensifying. Although this threatens tropical forests, cocoa sustainability initiatives largely overlook biodiversity conservation. To inform these initiatives, we analyzed how cocoa agriculture affects bird diversity at farm and landscape scales with a meta-analysis of 23 studies. We extracted 214 Hedges' g* comparisons of bird diversity and 14 comparisons of community similarity between a forest baseline and 4 farming systems that cover an intensification gradient in landscapes with high and low forest cover, and we summarized 119 correlations between cocoa farm features and bird diversity. Bird diversity declined sharply in low shade cocoa. Cocoa with >30% canopy cover from diverse trees retained bird diversity similar to nearby primary or mature secondary forest but held a different community of birds. Diversity of endemic species, frugivores, and insectivores (agriculture avoiders) declined, whereas diversity of habitat generalists, migrants, nectarivores, and granivores (agriculture associates) increased. As forest decreased on the landscape, the difference in bird community composition between forest and cocoa also decreased, indicating agriculture associates replaced agriculture avoiders in forest patches. Our results emphasize the need to conserve forested landscapes (land sparing) and invest in mixed-shade agroforestry (land sharing) because each strategy benefits a diverse and distinct biological community.

Impacto de la Intensificación Agrícola del Cacao sobre la Diversidad y Composición de la Comunidad de Aves Resumen Para responder a la demanda creciente de chocolate, el cultivo de cacao (Theobroma cacao) se ha expandido e intensificado. Aunque esto es una amenaza para los bosques tropicales, las iniciativas de cacao sustentable en gran medida pasan por alto la conservación de la biodiversidad. Para proporcionar información a estas iniciativas, analizamos como la agricultura del cacao afecta a la diversidad de aves a escala de rancho y de paisaje mediante un metaanálisis de 23 estudios. Extrajimos 214 comparaciones de Hedges g* de la diversidad de aves y 14 comparaciones de la similitud de comunidades entre una línea de base de bosque y 4 sistemas de cultivo que cubren un gradiente de intensificación en paisajes con cobertura de bosque alta a baja, y sintetizamos 119 correlaciones entre características de cultivos de cacao y la diversidad de aves. La diversidad de aves declinó claramente en cultivos con poca sombra. Cultivos con >30% de cobertura de diversos árboles retuvieron una diversidad de aves similar a la de bosques primarios o maduros cercanos, pero presentaron una comunidad diferente. La diversidad de especies endémicas, frugívoras e insectívoras (evasoras de agricultura) declinó, mientras que la diversidad de generalistas de hábitat, migrantes, nectarívoras y granívoras (asociadas a agricultura) incrementó. A medida que decreció el bosque en el paisaje, la diferencia en la composición de la comunidad de aves entre bosque y cacao también decreció, lo que indica que las especies asociadas a la agricultura reemplazaron a las evasoras de la agricultura en los fragmentos de bosque. Nuestros resultados enfatizan la necesidad de conservar paisajes boscosos (conservación de tierras) e invertir en agroforestería de sombra mixta (compartición de tierras) porque cada estrategia beneficia a una comunidad biológica diversa y distinta.

Repeated sampling of individuals reveals impact of tropical and temperate habitats on microbiota of a migratory bird.

Migratory animals experiencing substantial change in diet and habitat across the annual cycle may have corresponding shifts in host-associated microbial diversity. Using automated telemetry and radio tags to recapture birds, we examined gut microbiota structure in the same population and often same individual of Kirtland's Warblers (Setophaga kirtlandii) initially sampled on their wintering grounds in The Bahamas and subsequently resampled within their breeding territories in Michigan, USA. Initial sampling occurred in March and April and resampling occurred in May, June and early July. The composition of the most abundant phyla and classes of the warblers' microbiota is similar to that of other migratory birds. However, we detected notable variation in abundance and diversity of numerous bacterial taxa, including a decrease in microbial richness and significant differences in microbial communities when comparing the microbiota of birds first captured in The Bahamas to that of birds recaptured in Michigan. This is observed at the individual and population level. Furthermore, we found that 22 bacterial genera exhibit heightened abundance within specific sampling periods and are probably associated with diet and environmental change. Finally, we described a small, species-specific shared microbial profile that spans multiple time periods and environments within the migratory cycle. Our research highlights that the avian gut microbiota is dynamic over time, most significantly impacted by changing environments associated with migration. These results support the need for full annual cycle monitoring of migratory bird microbiota to improve understanding of seasonal host movement ecologies and response to recurrent physiological stressors.

Hatching date influences winter habitat occupancy: Examining seasonal interactions across the full annual cycle in a migratory songbird.

Birds experience a sequence of critical events during their life cycle, and past events can subsequently determine future performance via carry-over effects. Events during the non-breeding season may influence breeding season phenology or productivity. Less is understood about how events during the breeding season affect individuals subsequently in their life cycle. Using stable carbon isotopes, we examined carry-over effects throughout the annual cycle of prairie warblers (Setophaga discolor), a declining Nearctic-Neotropical migratory passerine bird. In drier winters, juvenile males that hatched earlier at our study site in Massachusetts, USA, occupied wetter, better-quality winter habitat in the Caribbean, as indicated by depleted carbon isotope signatures. For juveniles that were sampled again as adults, repeatability in isotope signatures indicated similar winter habitat occupancy across years. Thus, hatching date of juvenile males appears to influence lifetime winter habitat occupancy. For adult males, reproductive success did not carry over to influence winter habitat occupancy. We did not find temporally consecutive "domino" effects across the annual cycle (breeding to wintering to breeding) or interseasonal, intergenerational effects. Our finding that a male's hatching date can have a lasting effect on winter habitat occupancy represents an important contribution to our understanding of seasonal interactions in migratory birds.

Variable tropical moisture and food availability underlie mixed winter space-use strategies in a migratory songbird.

Identifying environmental correlates driving space-use strategies can be critical for predicting population dynamics; however, such information can be difficult to attain for small mobile species such as migratory songbirds. We combined radio-telemetry and high-resolution GPS tracking to examine space-use strategies under different moisture gradients for wood thrush (Hylocichla mustelina). We explored the role moisture plays in driving food abundance and, in turn, space-use strategies at a wintering site in Belize across 3 years. Individuals occupying drier habitats experienced lower food abundance and poorer body condition. Using data from our radio-tracked study population and GPS tracking from across five breeding populations, we detected low rates of overwinter site persistence across the wood thrush wintering range. Contrary to expectations, individuals in wetter habitats were more likely to engage in permanent mid-winter relocations, up to 148 km. We suggest facultative movements are instead a condition-dependent strategy that enables wintering wood thrush to locate alternative habitat as food availability declines throughout the dry season. Increased aridity is predicted across the wintering range of wood thrush, and future research should delve deeper into understanding how moisture impacts within and between season space-use dynamics and its ultimate impact on the population dynamics of this declining species.

A general theory of avian migratory connectivity.

Birds exhibit a remarkable array of seasonal migrations. Despite much research describing migratory behaviour, the underlying forces driving how a species' breeding and wintering populations redistribute each year, that is, migratory connectivity, remain largely unknown. Here, we test the hypothesis that birds migrate in a way that minimises energy expenditure while considering intraspecific competition for energy acquisition, by developing a modelling framework that simulates an optimal redistribution of individuals between breeding and wintering areas. Using 25 species across the Americas, we find that the model accurately predicts empirical migration patterns, and thus offers a general explanation for migratory connectivity based on first ecological and energetic principles. Our model provides a strong basis for exploring additional processes underlying the ecology and evolution of migration, but also a framework for predicting how migration impacts local adaptation across seasons and how environmental change may affect population dynamics in migratory species.

Characterization of the Plasmodium and Haemoproteus parasite community in temperate-tropical birds during spring migration.

Animal movements, especially avian migration, can be a mechanism for the large-scale dispersal and geographic range expansion of parasites. The host-parasite relationships among birds during migration have yet to be fully explored. We characterized the haemosporidian parasite lineages in passerines during spring migration on the Texas coast of the Gulf of Mexico, and identified associations among wintering origin (US, Central America, South America) and foraging height (canopy, understory, ground) and infection status. We examined 743 samples representing 52 species of 10 families over six years, 2014-2019. We used PCR and DNA sequencing of the haemosporidian cytB gene from avian blood samples to determine infection status with the genera Plasmodium and Haemoproteus and characterize the lineages of blood parasites. We found an overall haemosporidian infection prevalence of 48.4% among neotropical migrant and Texas wintering birds. Among families, Icterids had the highest prevalence (75%, 24 individuals, 4 species sampled) whereas Parulids had the lowest prevalence (38.4%, 177 individuals, 18 species sampled). Among infected birds, Plasmodium spp. infections were more common than Haemoproteus spp. infections in species that winter in Central America compared to those that winter in the US or South America. Similarly, among infected birds, Plasmodium spp. infections were more common than Haemoproteus spp. infections in species that forage on the ground or in the understory compared to those that forage in the canopy. Infected birds harbored 65 different haemosporidian lineages (71% Plasmodium; 29% Haemoproteus) of which 17 lineages have never previously been reported and six lineages were documented for the first time in North America, having been previously detected only in Central or South America. These data are consistent with the premise that intercontinental parasite dispersal may be facilitated by passerine birds. Future studies focused on surveillance, the probability of establishment of parasite lineages, and the use of individual bird tracking methods to understand infection dispersion over time will allow a more comprehensive understanding of changing avian host-haemosporidian relationships.

Habitat loss on the breeding grounds is a major contributor to population declines in a long-distance migratory songbird.

Many migratory species are declining and for most, the proximate causes of their declines remain unknown. For many long-distance Neotropical migratory songbirds, it is assumed that habitat loss on breeding or non-breeding grounds is a primary driver of population declines. We integrated data collected from tracking technology, community science and remote sensing data to quantify migratory connectivity (MC), population trends and habitat loss. We quantified the correlation between forest change throughout the annual cycle and population declines of a long-distance migratory songbird, the Connecticut warbler (Oporornis agilis, observed decline: -8.99% yr-1). MC, the geographic link between populations during two or more phases of the annual cycle, was stronger between breeding and autumn migration routes (MC = 0.24 ± 0.23) than between breeding and non-breeding locations (MC = -0.2 ± 0.14). Different Connecticut warbler populations tended to have population-specific fall migration routes but overlapped almost completely within the northern Gran Chaco ecoregion in South America. Cumulative forest loss within 50 km of breeding locations and the resulting decline in the largest forested patch index was correlated more strongly with population declines than forest loss on migratory stopover regions or on wintering locations in South America, suggesting that habitat loss during the breeding season is a driver of observed population declines for the Connecticut warbler. Land-use practices that retain large, forested patches within landscapes will likely benefit breeding populations of this declining songbird, but further research is needed to help inform land-use practices across the full annual cycle to minimize the impacts to migratory songbirds and abate ongoing population declines.