Clark S. Rushing

Quantifying drivers of population dynamics for a migratory bird throughout the annual cycle.

Worldwide, migratory species are undergoing rapid declines but understanding the factors driving these declines is hindered by missing information about migratory connectivity and the lack of data to quantify environmental processes across the annual cycle. Here, we combined range-wide information about migratory connectivity with global remote-sensing data to quantify the relative importance of breeding and non-breeding environmental processes to persistent long-term population declines of a migratory songbird, the wood thrush (Hylocichla mustelina). Consistent with theoretical predictions about population limitation of migratory birds, our results suggest that habitat loss and climate have contributed to the observed declines in wood thrush breeding abundance, yet the relative importance of breeding versus non-breeding factors is population-specific. For example, high-abundance core breeding populations appear to be more limited by habitat loss, whereas low-abundance, peripheral populations appear to be limited by climate-driven seasonal interactions. Further, our analysis indicates that the relative impact of breeding habitat loss is at least three to six times greater than the impact of equivalent non-breeding habitat loss and therefore the steepest regional declines have likely been driven by the loss of breeding habitat. These results underscore the need for population-specific conservation strategies implemented throughout the annual cycle to reverse long-term declines.

Assessing migratory connectivity for a long‐distance migratory bird using multiple intrinsic markers

Patterns of migratory connectivity are a vital yet poorly understood component of the ecology and evolution of migratory birds. Our ability to accurately characterize patterns of migratory connectivity is often limited by the spatial resolution of the data, but recent advances in probabilistic assignment approaches have begun pairing stable isotopes with other sources of data (e.g., genetic and mark-recapture) to improve the accuracy and precision of inferences based on a single marker. Here, we combine stable isotopes and geographic variation in morphology (wing length) to probabilistically assign Wood Thrush (Hylocichla mustilena) captured on the wintering grounds to breeding locations. In addition, we use known-origin samples to validate our model and assess potentially important impacts of isotopic and morphological covariates (age, sex, and breeding location). Our results show that despite relatively high levels of mixing across their breeding and nonbreeding ranges, moderate levels of migratory connectivity exist along an east-west gradient. In addition, combining stable isotopes with geographic variation in wing length improved the precision of breeding assignments by 10% and 37% compared to assignments based on isotopes alone or wing length alone, respectively. These results demonstrate that geographical variation in morphological traits can greatly improve estimates of migratory connectivity when combined with other intrinsic markers (e.g., stable isotopes or genetic data). The wealth of morphological data available from museum specimens across the world represents a tremendously valuable, but largely untapped, resource that is widely applicable for quantifying patterns of migratory connectivity.

Integrating microorganism and macroorganism dispersal: modes, techniques and challenges with particular focus on co-dispersal

ABSTRACT Whatever their size and the ecosystem they live in, all organisms may disperse at some stage of their life cycle. Dispersal dynamics are to a varying extent dependent on organismal size, life history, ecological niche, survival capacities and phylogeny. Moves towards a synthesis in dispersal ecology have focused primarily on vertebrates and higher plants, yet recent studies suggest that the dispersal of microorganisms and macroorganisms has much more in common than previously assumed. The dispersal of one organism enables co-dispersal for many others, smaller in size. There is an increasing need for a more integrated approach to study dispersal within the context of organismal interactions and their environments. Such an approach is facilitated by recent developments of powerful indirect techniques that enable tracking of microorganisms and macroorganisms over multiple spatial and temporal scales. Likewise, dispersal modelling and theoretical models of the consequences of dispersal can inspire empirical studies across the entire size spectrum. Simultaneously studying the relationships between dispersal of microorganisms and macroorganisms, and accounting for dispersal through time and space, will allow us to better understand the functioning and dynamics of communities and ecosystems, and to make better predictions of future dispersal patterns, changes in biodiversity and connectivity.

Winter habitat quality but not long‐distance dispersal influences apparent reproductive success in a migratory bird

Long-distance breeding and natal dispersal play central roles in many ecological and evolutionary processes, including gene flow, population dynamics, range expansion, and individual responses to fluctuating biotic and abiotic conditions. However, the relative contribution of long-distance dispersal to these processes depends on the ability of dispersing individuals to successfully reproduce in their new environment. Unfortunately, due to the difficulties associated with tracking dispersal in the field, relatively little is known about its reproductive consequences. Furthermore, because reproductive success is influenced by a variety of processes, disentangling the influence of each of these processes is critical to understanding the direct consequences of dispersal. In this study, we used stable hydrogen and carbon isotopes to estimate long-distance dispersal and winter territory quality in a migratory bird, the American Redstart (Setophaga ruticilla). We then applied Aster life-history models to quantify the strength of influence of these factors on apparent reproductive success. We found no evidence that male or female reproductive success was lower for long-distance dispersers relative to non-dispersing individuals. In contrast, carry-over effects from the winter season did influence male, but not female, reproductive success. Use of Aster models further revealed that for adult males, winter territory quality influenced the number of offspring produced whereas for yearling males, high-quality winter territories were associated with higher mating and nesting success. These results suggest that although long-distance natal and breeding dispersal carry no immediate reproductive cost for American Redstarts, reproductive success in this species may ultimately be limited by the quality of winter habitat.

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.

Using demographic attributes from long-term monitoring data to delineate natural population structure

Summary The concept of ‘natural’ populations is a foundation of modern ecology and conservation, with a large body of theoretical literature using these discrete demographic units to understand population dynamics and prioritize conservation strategies. To date, there are currently no objective methods for empirically delineating large-scale population boundaries using demographic data. We present a novel approach for using large-scale, citizen-science monitoring data to quantify geographic structure in trend and abundance and identify distinct natural populations. We demonstrate this approach by delineating populations of eight passerine species using data collected as part of the North American Breeding Bird Survey. Our approach was able to identify geographic structure in both trend and abundance and to delineate distinct populations for all eight species. An independent validation of three species indicated this demographic variation was reflected in underlying vital rates. Synthesis and applications. Natural populations are biologically based alternatives to the traditional geographically defined units that can improve the ability of researchers and managers to quantify spatial variation in population dynamics. Our analysis of natural population structure in breeding songbirds demonstrates that species can show substantial geographic variation in population attributes and underlying demography. We recommend managers define spatial units using natural populations when setting regional population objectives for both single and multispecies conservation plans.

Feather corticosterone levels are related to age and future body condition, but not to subsequent fitness, in a declining migratory songbird

In migratory species, breeding and non-breeding locations are geographically separate, yet the effects of conditions from one stage may carry over to affect a subsequent stage. Ideally, to understand the mechanisms and implications of ‘carry-over effects’, one would need to follow individuals throughout the year, quantify potential environmental causal factors and physiological mediators during multiple life-history stages, and measure downstream fitness. Owing to current limitations of tracking technology, this is impossible for small, long-distance migrants, so indirect methods to characterize carry-over effects are required. Corticosterone (CORT) is a suspected physiological mediator of carry-over effects, but when collected from blood it provides only a physiological snapshot at that point in time. When extracted from feathers, however, feather corticosterone (CORTf) provides a measure of responses to stressors from previous, and longer, time periods. We collected feathers grown during two life-history stages (post-breeding and subsequent wintering) from individuals of two age classes of a rapidly declining migratory songbird, the cerulean warbler (Setophaga cerulea), on their breeding grounds and quantified CORTf concentrations. We then monitored reproduction and survival of individuals and analysed relationships among CORTf and age, body condition and future fitness. Compared with older males, second-year males had higher CORTf concentrations during both stages. When controlling for age and year, body condition at capture was positively related to CORTf concentrations from winter (especially for older birds). However, we found no relationships between CORTf and fitness (as defined by reproduction and survival). Thus, elevated CORT may represent a beneficial physiological response (e.g. hyperphagia prior to migration), particularly for certain life-history stages, and may mediate the condition in which individuals transition between stages. But for those birds that survive migration, subsequent fitness is likely determined by more recent events and local conditions (i.e. on breeding grounds), which have the potential to counteract conditions from the winter.

Habitat features and long‐distance dispersal modify the use of social information by a long‐distance migratory bird

The processes by which individuals select breeding sites have important consequences for individual tness as well as population- and community-dynamics. Although there is increasing evidence that many animal species use information acquired from conspecics to assess the suitability of potential breeding sites, little is known about how the use of this social information is modified by biotic and abiotic conditions. We used an automated playback experiment to simulate two types of social information, post-breeding public information and pre-breeding location cues, to determine the relative importance of these cues for breeding site selection by a migratory songbird, the American redstart (Setophaga ruticilla). In addition, we used stable hydrogen isotopes to determine the dispersal status of individuals that responded to our experimental treatments and quantify whether long-distance dispersers use different social cues to select breeding sites compared to philopatric individuals. We found that points that received pre-breeding location cue treatments were signi cantly more likely to be settled by redstarts than control points that received no playback. However, we found no evidence the redstarts used post-breeding public information gathered during one season to select breeding sites the following year. Breeding site habitat structure was also a strong predictor of settlement probability, indicating that redstarts modi ed the use of social information based on habitat cues. Furthermore, stable hydrogen isotope signatures from individuals that responded to location cue treatments suggest that long-distance dispersers may rely more heavily on these cues than local recruits. Collectively, these results indicate that redstarts use multiple sources of information to select breeding sites, which could buffer individuals from selecting suboptimal sites when they breed in unfamiliar locations or when habitat quality becomes decoupled from social cues.

Incorporating breeding abundance into spatial assignments on continuous surfaces

Abstract Determining the geographic connections between breeding and nonbreeding populations, termed migratory connectivity, is critical to advancing our understanding of the ecology and conservation of migratory species. Assignment models based on stable isotopes historically have been an important tool for studying migratory connectivity of small‐bodied species, but the low resolution of these assignments has generated interest into combining isotopes with other sources in information. Abundance is one of the most appealing data sources to include in isotope‐based assignments, but there are currently no statistical methods or guidelines for optimizing the contribution of stable isotopes and abundance for inferring migratory connectivity. Using known‐origin stable‐hydrogen isotope samples of six Neotropical migratory bird species, we rigorously assessed the performance of assignment models that differentially weight the contribution of the isotope and abundance data. For two species with adequate sample sizes, we used Pareto optimality to determine the set of models that simultaneously minimized both assignment error rate and assignment area. We then assessed the ability of the top models from these two species to improve assignments of the remaining four species compared to assignments based on isotopes alone. We show that the increased precision of models that include abundance is often offset by a large increase in assignment error. However, models that optimally weigh the abundance data relative to the isotope data can result in higher precision and, in some cases, lower error than models based on isotopes alone. The top models, however, depended on the distribution of relative breeding abundance, with patchier distributions requiring stronger downweighting of abundance, and we present general guidelines for future studies. These results confirm that breeding abundance can be an important source of information for studies investigating broad‐scale movements of migratory birds and potentially other taxa.