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.

Unraveling Adaptive Evolutionary Divergence at Microgeographic Scales.

AbstractStriking examples of local adaptation at fine geographic scales are increasingly being documented in natural populations. However, the relative contributions made by natural selection, phenotype-dependent dispersal (when individuals disperse with respect to a habitat preference), and mate preference in generating and maintaining microgeographic adaptation and divergence are not well studied. Here, we develop quantitative genetics models and individual-based simulations (IBSs) to uncover the evolutionary forces that possibly drive microgeographic divergence. We also perform Bayesian estimation of the parameters in our IBS using empirical data on habitat-specific variation in bill morphology in the island scrub-jay (Aphelocoma insularis) to apply our models to a natural system. We find that natural selection and phenotype-dependent dispersal can generate the patterns of divergence we observe in the island scrub-jay. However, mate preference for a mate with similar bill morphology, even though observed in the species, does not play a significant role in driving divergence. Our modeling approach provides insights into phenotypic evolution occurring over small spatial scales relative to dispersal ranges, suggesting that adaptive divergence at microgeographic scales may be common across a wider range of taxa than previously thought. Our quantitative genetic models help to inform future theoretical and empirical work to determine how selection, habitat preference, and mate preference contribute to local adaptation and microgeographic divergence.

Climate-mediated population dynamics of a migratory songbird differ between the trailing edge and range core.

Understanding the demographic drivers of range contractions is important for predicting species' responses to climate change; however, few studies have examined the effects of climate change on survival and recruitment across species' ranges. We show that climate change can drive trailing edge range contractions through the effects on apparent survival, and potentially recruitment, in a migratory songbird. We assessed the demographic drivers of trailing edge range contractions using a long-term demography dataset for the black-throated blue warbler (Setophaga caerulescens) collected across elevational climate gradients at the trailing edge and core of the breeding range. We used a Bayesian hierarchical model to estimate the effect of climate change on apparent survival and recruitment and to forecast population viability at study plots through 2040. The trailing edge population at the low-elevation plot became locally extinct by 2017. The local population at the mid-elevation plot at the trailing edge gradually declined and is predicted to become extirpated by 2040. Population declines were associated with warming temperatures at the mid-elevation plot, although results were more equivocal at the low-elevation plot where we had fewer years of data. Population density was stable or increasing at the range core, although warming temperatures are predicted to cause population declines by 2040 at the low-elevation plot. This result suggests that even populations within the geographic core of the range are vulnerable to climate change. The demographic drivers of local population declines varied between study plots, but warming temperatures were frequently associated with declining rates of population growth and apparent survival. Declining apparent survival in our study system is likely to be associated with increased adult emigration away from poor-quality habitats. Our results suggest that demographic responses to warming temperatures are complex and dependent on local conditions and geographic range position, but spatial variation in population declines is consistent with the climate-mediated range shift hypothesis. Local populations of black-throated blue warblers near the warm-edge range boundary at low latitudes and low elevations are likely to be the most vulnerable to climate change, potentially leading to local extirpation and range contractions.

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.

Differential survival throughout the full annual cycle of a migratory bird presents a life-history trade-off.

Long-distance migrations are among the most physically demanding feats animals perform. Understanding the potential costs and benefits of such behaviour is a fundamental question in ecology and evolution. A hypothetical cost of migration should be outweighed by higher productivity and/or higher annual survival, but few studies on migratory species have been able to directly quantify patterns of survival throughout the full annual cycle and across the majority of a species' range. Here, we use telemetry data from 220 migratory Egyptian vultures Neophron percnopterus, tracked for 3,186 bird months and across approximately 70% of the species' global distribution, to test for differences in survival throughout the annual cycle. We estimated monthly survival probability relative to migration and latitude using a multi-event capture-recapture model in a Bayesian framework that accounted for age, origin, subpopulation and the uncertainty of classifying fates from tracking data. We found lower survival during migration compared to stationary periods (ß = -0.816; 95% credible interval: -1.290 to -0.318) and higher survival on non-breeding grounds at southern latitudes (<25°N; ß = 0.664; 0.076-1.319) compared to on breeding grounds. Survival was also higher for individuals originating from Western Europe (ß = 0.664; 0.110-1.330) as compared to further east in Europe and Asia, and improved with age (ß = 0.030; 0.020-0.042). Anthropogenic mortalities accounted for half of the mortalities with a known cause and occurred mainly in northern latitudes. Many juveniles drowned in the Mediterranean Sea on their first autumn migration while there were few confirmed mortalities in the Sahara Desert, indicating that migration barriers are likely species-specific. Our study advances the understanding of important fitness trade-offs associated with long-distance migration. We conclude that there is lower survival associated with migration, but that this may be offset by higher non-breeding survival at lower latitudes. We found more human-caused mortality farther north, and suggest that increasing anthropogenic mortality could disrupt the delicate migration trade-off balance. Research to investigate further potential benefits of migration (e.g. differential productivity across latitudes) could clarify how migration evolved and how migrants may persist in a rapidly changing world.

Ecological and Social Factors Constrain Spatial and Temporal Opportunities for Mating in a Migratory Songbird.

Many studies of sexual selection assume that individuals have equal mating opportunities and that differences in mating success result from variation in sexual traits. However, the inability of sexual traits to explain variation in male mating success suggests that other factors moderate the strength of sexual selection. Extrapair paternity is common in vertebrates and can contribute to variation in mating success and thus serves as a model for understanding the operation of sexual selection. We developed a spatially explicit, multifactor model of all possible female-male pairings to test the hypothesis that ecological (food availability) and social (breeding density, breeding distance, and the social mate's nest stage) factors influence an individual's opportunity for extrapair paternity in a socially monogamous bird, the black-throated blue warbler, Setophaga caerulescens. A male's probability of siring extrapair young decreased with increasing distance to females, breeding density, and food availability. Males on food-poor territories were more likely to sire extrapair young, and these offspring were produced farther from the male's territory relative to males on food-abundant territories. Moreover, males sired extrapair young mostly during their social mates' incubation stage, especially males on food-abundant territories. This study demonstrates how ecological and social conditions constrain the spatial and temporal opportunities for extrapair paternity that affect variation in mating success and the strength of sexual selection in socially monogamous species.

Spatial and temporal drivers of avian population dynamics across the annual cycle.

Untangling the spatial and temporal processes that influence population dynamics of migratory species is challenging, because changes in abundance are shaped by variation in vital rates across heterogeneous habitats and throughout the annual cycle. We developed a full-annual-cycle, integrated, population model and used demographic data collected between 2011 and 2014 in southern Indiana and Belize to estimate stage-specific vital rates of a declining migratory songbird, the Wood Thrush (Hylocichla mustelina). Our primary objective was to understand how spatial and temporal variation in demography contributes to local and regional population growth. Our full-annual-cycle model allowed us to estimate (1) age-specific, seasonal survival probabilities, including latent survival during both spring and autumn migration, and (2) how the relative contribution of vital rates to population growth differed among habitats. Wood Thrushes in our study populations experienced the lowest apparent survival rates during migration and apparent survival was lower during spring migration than during fall migration. Both mortality and high dispersal likely contributed to low apparent survival during spring migration. Population growth in high-quality habitat was most sensitive to variation in fecundity and apparent survival of juveniles during spring migration, whereas population growth in low-quality sites was most sensitive to adult apparent breeding-season survival. These results elucidate how full-annual-cycle vital rates, particularly apparent survival during migration, interact with spatial variation in habitat quality to influence population dynamics in migratory species.

Seasonal survival estimation for a long-distance migratory bird and the influence of winter precipitation.

Conservation of migratory animals requires information about seasonal survival rates. Identifying factors that limit populations, and the portions of the annual cycle in which they occur, are critical for recognizing and reducing potential threats. However, such data are lacking for virtually all migratory taxa. We investigated patterns and environmental correlates of annual, oversummer, overwinter, and migratory survival for adult male Kirtland's warblers (Setophaga kirtlandii), an endangered, long-distance migratory songbird. We used Cormack-Jolly-Seber models to analyze two mark-recapture datasets: 2006-2011 on Michigan breeding grounds, and 2003-2010 on Bahamian wintering grounds. The mean annual survival probability was 0.58 ± 0.12 SE. Monthly survival probabilities during the summer and winter stationary periods were relatively high (0.963 ± 0.005 SE and 0.977 ± 0.002 SE, respectively). Monthly survival probability during migratory periods was substantially lower (0.879 ± 0.05 SE), accounting for ~44% of all annual mortality. March rainfall in the Bahamas was the best-supported predictor of annual survival probability and was positively correlated with apparent annual survival in the subsequent year, suggesting that the effects of winter precipitation carried over to influence survival probability of individuals in later seasons. Projection modeling revealed that a decrease in Bahamas March rainfall >12.4% from its current mean could result in negative population growth in this species. Collectively, our results suggest that increased drought during the non-breeding season, which is predicted to occur under multiple climate change scenarios, could have important consequences on the annual survival and population growth rate of Kirtland's warbler and other Neotropical-Nearctic migratory bird species.

Climate, demography and lek stability in an Amazonian bird.

Lekking is a rare, but iconic mating system where polygynous males aggregate and perform group displays to attract females. Existing theory postulates that demographic and environmental stability are required for lekking to be an evolutionarily viable reproductive strategy. However, we lack empirical tests for the hypotheses that lek stability is facilitated by age-specific variation in demographic rates, and by predictable, abundant resources. To address this knowledge gap, we use multistate models to examine how two demographic elements of lek stability-male survival and recruitment-vary with age, social status and phase of the El Niño Southern Oscillation (ENSO) in a Neotropical frugivorous bird, the wire-tailed manakin (Pipra filicauda). Our results show that demographic and environmental conditions were related to lek stability in the Ecuadorean Amazon. Apparent annual survival probability of territorial males was higher than that of non-territorial floaters, and recruitment probability increased as males progressed in an age-graded queue. Moreover, annual survival of territorial males and body condition of both floaters and territory holders were higher following years with El Niño conditions, associated with reduced rainfall and probably higher fruit production in the northern Neotropics, and lower after years with wet, La Niña conditions that predominated our study. Recruitment probabilities varied annually, independent of ENSO phase, and increased over our study period, but the annual mean number of territorial males per lek declined. Our results provide empirical support for hypothesized demographic and environmental drivers of lek dynamics. This study also suggests that climate-mediated changes in resource availability can affect demography and subsequent lek stability in a relatively buffered, lowland rainforest.

Adaptive divergence despite strong genetic drift: genomic analysis of the evolutionary mechanisms causing genetic differentiation in the island fox (Urocyon littoralis).

The evolutionary mechanisms generating the tremendous biodiversity of islands have long fascinated evolutionary biologists. Genetic drift and divergent selection are predicted to be strong on islands and both could drive population divergence and speciation. Alternatively, strong genetic drift may preclude adaptation. We conducted a genomic analysis to test the roles of genetic drift and divergent selection in causing genetic differentiation among populations of the island fox (Urocyon littoralis). This species consists of six subspecies, each of which occupies a different California Channel Island. Analysis of 5293 SNP loci generated using Restriction-site Associated DNA (RAD) sequencing found support for genetic drift as the dominant evolutionary mechanism driving population divergence among island fox populations. In particular, populations had exceptionally low genetic variation, small Ne (range = 2.1-89.7; median = 19.4), and significant genetic signatures of bottlenecks. Moreover, islands with the lowest genetic variation (and, by inference, the strongest historical genetic drift) were most genetically differentiated from mainland grey foxes, and vice versa, indicating genetic drift drives genome-wide divergence. Nonetheless, outlier tests identified 3.6-6.6% of loci as high FST outliers, suggesting that despite strong genetic drift, divergent selection contributes to population divergence. Patterns of similarity among populations based on high FST outliers mirrored patterns based on morphology, providing additional evidence that outliers reflect adaptive divergence. Extremely low genetic variation and small Ne in some island fox populations, particularly on San Nicolas Island, suggest that they may be vulnerable to fixation of deleterious alleles, decreased fitness and reduced adaptive potential.

The interacting effects of food, spring temperature, and global climate cycles on population dynamics of a migratory songbird.

Although long-distance migratory songbirds are widely believed to be at risk from warming temperature trends, species capable of attempting more than one brood in a breeding season could benefit from extended breeding seasons in warmer springs. To evaluate local and global factors affecting population dynamics of the black-throated blue warbler (Setophaga caerulescens), a double-brooded long-distance migrant, we used Pradel models to analyze 25 years of mark-recapture data collected in New Hampshire, USA. We assessed the effects of spring temperature (local weather) and the El Niño Southern Oscillation index (a global climate cycle), as well as predator abundance, insect biomass, and local conspecific density on population growth in the subsequent year. Local and global climatic conditions affected warbler populations in different ways. We found that warbler population growth was lower following El Niño years (which have been linked to poor survival in the wintering grounds and low fledging weights in the breeding grounds) than La Niña years. At a local scale, populations increased following years with warm springs and abundant late-season food, but were unaffected by spring temperature following years when food was scarce. These results indicate that the warming temperature trends might have a positive effect on recruitment and population growth of black-throated blue warblers if food abundance is sustained in breeding areas. In contrast, potential intensification of future El Niño events could negatively impact vital rates and populations of this species.

Context-dependent seed dispersal by a scatter-hoarding corvid.

Corvids (crows, jays, magpies and nutcrackers) are important dispersers of large-seeded plants. Studies on captive or supplemented birds suggest that they flexibly adjust their scatter-hoarding behaviour to the context of social dynamics and relative seed availability. Because many corvid-dispersed trees show high annual variation in seed production, context-dependent foraging can have strong effects on natural corvid scatter-hoarding behaviour. We investigated how seed availability and social dynamics affected scatter-hoarding in the island scrub jays (Aphelocoma insularis). We quantified rates of scatter-hoarding behaviour and territorial defence of 26 colour-marked birds over a three-year period with variable acorn crops. We tested whether caching parameters were correlated with variation in annual seed production of oaks as predicted by the predator dispersal hypothesis, which states that caching rates and distances should vary with seed abundance in ways that benefit tree fitness. We also tested whether antagonistic interactions with conspecifics would affect scatter-hoarding adversely, as found in experimental studies. Caching behaviour varied with acorn availability. Caching distances correlated positively with annual acorn crop size, increasing by as much as 40% between years. Caching rates declined over time in years with small acorn crops, but increased when crops were large. Acorn foraging and caching rates were also negatively correlated with rates of territorial aggression. Overall foraging rates, however, were not associated with aggression, suggesting that reduced dispersal rates were not simply due to time constraints. Our field results support laboratory findings that caching rates and distances by scatter-hoarding corvids are context-dependent. Furthermore, our results are consistent with predictions of the predator dispersal hypothesis and suggest that large seed crops and social interactions among scatter-hoarders affect dispersal benefits for oaks and other masting tree species.

Experimental food supplementation reveals habitat-dependent male reproductive investment in a migratory bird.

Environmental factors can shape reproductive investment strategies and influence the variance in male mating success. Environmental effects on extrapair paternity have traditionally been ascribed to aspects of the social environment, such as breeding density and synchrony. However, social factors are often confounded with habitat quality and are challenging to disentangle. We used both natural variation in habitat quality and a food supplementation experiment to separate the effects of food availability-one key aspect of habitat quality-on extrapair paternity (EPP) and reproductive success in the black-throated blue warbler, Setophaga caerulescens. High natural food availability was associated with higher within-pair paternity (WPP) and fledging two broods late in the breeding season, but lower EPP. Food-supplemented males had higher WPP leading to higher reproductive success relative to controls, and when in low-quality habitat, food-supplemented males were more likely to fledge two broods but less likely to gain EPP. Our results demonstrate that food availability affects trade-offs in reproductive activities. When food constraints are reduced, males invest in WPP at the expense of EPP. These findings imply that environmental change could alter how individuals allocate their resources and affect the selective environment that drives variation in male mating success.

Non-breeding season habitat quality mediates the strength of density-dependence for a migratory bird.

Our understanding of when natural populations are regulated during their annual cycle is limited, particularly for migratory species. This information is needed for parametrizing models that can inform management and conservation. Here, we use 14 years of data on colour-marked birds to investigate how conspecific density and habitat quality during the tropical non-breeding period interact to affect body condition and apparent annual survival of a long-distance migratory songbird, the American redstart (Setophagaruticilla). Body condition and survival of birds in high-quality mangrove habitat declined as density increased. By contrast, body condition improved and survival did not vary as density increased in adjacent, lower quality scrub habitat, although mean condition and survival were almost always lower than in mangrove. High rainfall enhanced body condition in scrub but not in mangrove, suggesting factors such as food availability outweighed consequences of crowding in lower quality habitat. Thus, survival of overwintering redstarts in mangrove habitat, disproportionately males,appears to be regulated by a crowding mechanism based on density-dependent resource competition. Survival of individuals in scrub, mostly females, appears to be limited by density-independent environmental factors but not regulated by crowding. The contrasting effects of density and food limitation on individuals overwintering in adjacent habitats illustrate the complexity of processes operating during the non-breeding period for migratory animals, and emphasize the need for long-term studies of animals in multiple habitats and throughout their annual cycles.

An open-population hierarchical distance sampling model.

Modeling population dynamics while accounting for imperfect detection is essential to monitoring programs. Distance sampling allows estimating population size while accounting for imperfect detection, but existing methods do not allow for estimation of demographic parameters. We develop a model that uses temporal correlation in abundance arising from underlying population dynamics to estimate demographic parameters from repeated distance sampling surveys. Using a simulation study motivated by designing a monitoring program for Island Scrub-Jays (Aphelocoma insularis), we investigated the power of this model to detect population trends. We generated temporally autocorrelated abundance and distance sampling data over six surveys, using population rates of change of 0.95 and 0.90. We fit the data generating Markovian model and a mis-specified model with a log-linear time effect on abundance, and derived post hoc trend estimates from a model estimating abundance for each survey separately. We performed these analyses for varying numbers of survey points. Power to detect population changes was consistently greater under the Markov model than under the alternatives, particularly for reduced numbers of survey points. The model can readily be extended to more complex demographic processes than considered in our simulations. This novel framework can be widely adopted for wildlife population monitoring.

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.

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.

Black-throated blue warblers (Setophaga caerulescens) exhibit diet flexibility and track seasonal changes in insect availability.

Changes in leaf phenology from warming spring and autumn temperatures have lengthened the temperate zone growing "green" season and breeding window for migratory birds in North America. However, the fitness benefits of an extended breeding season will depend, in part, on whether species have sufficient dietary flexibility to accommodate seasonal changes in prey availability. We used fecal DNA metabarcoding to test the hypothesis that seasonal changes in the diets of the insectivorous, migratory black-throated blue warbler (Setophaga caerulescens) track changes in the availability of arthropod prey at the Hubbard Brook Experimental Forest, New Hampshire, USA. We examined changes across the breeding season and along an elevation gradient encompassing a 2-week difference in green season length. From 98 fecal samples, we identified 395 taxa from 17 arthropod orders; 242 were identified to species, with Cecrita guttivitta (saddled prominent moth), Theridion frondeum (eastern long-legged cobweaver), and Philodromus rufus (white-striped running crab spider) occurring at the highest frequency. We found significant differences in diet composition between survey periods and weak differences among elevation zones. Variance in diet composition was highest late in the season, and diet richness and diversity were highest early in the season. Diet composition was associated with changes in prey availability surveyed over the green season. However, several taxa occurred in diets more or less than expected relative to their frequency of occurrence from survey data, suggesting that prey selection or avoidance sometimes accompanies opportunistic foraging. This study demonstrates that black-throated blue warblers exhibit diet flexibility and track seasonal changes in prey availability, which has implications for migratory bird responses to climate-induced changes in insect communities with longer green seasons.

Hierarchical distance-sampling models to estimate population size and habitat-specific abundance of an island endemic.

Population size and habitat-specific abundance estimates are essential for conservation management. A major impediment to obtaining such estimates is that few statistical models are able to simultaneously account for both spatial variation in abundance and heterogeneity in detection probability, and still be amenable to large-scale applications. The hierarchical distance-sampling model of J. A. Royle, D. K. Dawson, and S. Bates provides a practical solution. Here, we extend this model to estimate habitat-specific abundance and rangewide population size of a bird species of management concern, the Island Scrub-Jay (Aphelocoma insularis), which occurs solely on Santa Cruz Island, California, USA. We surveyed 307 randomly selected, 300 m diameter, point locations throughout the 250-km2 island during October 2008 and April 2009. Population size was estimated to be 2267 (95% CI 1613-3007) and 1705 (1212-2369) during the fall and spring respectively, considerably lower than a previously published but statistically problematic estimate of 12 500. This large discrepancy emphasizes the importance of proper survey design and analysis for obtaining reliable information for management decisions. Jays were most abundant in low-elevation chaparral habitat; the detection function depended primarily on the percent cover of chaparral and forest within count circles. Vegetation change on the island has been dramatic in recent decades, due to release from herbivory following the eradication of feral sheep (Ovis aries) from the majority of the island in the mid-1980s. We applied best-fit fall and spring models of habitat-specific jay abundance to a vegetation map from 1985, and estimated the population size of A. insularis was 1400-1500 at that time. The 20-30% increase in the jay population suggests that the species has benefited from the recovery of native vegetation since sheep removal. Nevertheless, this jay's tiny range and small population size make it vulnerable to natural disasters and to habitat alteration related to climate change. Our results demonstrate that hierarchical distance-sampling models hold promise for estimating population size and spatial density variation at large scales. Our statistical methods have been incorporated into the R package unmarked to facilitate their use by animal ecologists, and we provide annotated code in the Supplement.

Variance in within-pair reproductive success influences the opportunity for selection annually and over the lifetimes of males in a multibrooded songbird.

In socially monogamous species, male reproductive success consists of "within-pair" offspring produced with their socially paired mate(s), and "extra-pair" offspring produced with additional females throughout the population. Both reproductive pathways offer distinct opportunities for selection in wild populations, as each is composed of separate components of mate attraction, female fecundity, and paternity allocation. Identifying key sources of variance and covariance among these components is a crucial step toward understanding the reproductive strategies that males use to maximize fitness both annually and over their lifetimes. We use 16 years of complete reproductive data from a population of black-throated blue warblers (Setophaga caerulescens) to partition variance in male annual and lifetime reproductive success, and thereby identify if the opportunity for selection varies over the lifetimes of individual males and what reproductive strategies likely favor maximum lifetime fitness. The majority of variance in male reproduction was attributable to within-pair success, but the specific effects of individual components of variance differed between total annual and total lifetime reproductive success. Positive overall lifetime covariance between within-pair and extra-pair components indicates that males able to maximize within-pair success, particularly with double-brooding females, likely achieve higher overall lifetime fitness via both within-pair and extra-pair reproductive pathways.