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Mapping behavioral landscapes for animal movement: a finite mixture modeling approach

Because of its role in many ecological processes, movement of animals in response to landscape features is an important subject in ecology and conservation biology. In this paper, we develop models of animal movement in relation to objects or fields in a landscape. We take a finite mixture modeling approach in which the component densities are conceptually related to different choices for movement in response to a landscape feature, and the mixing proportions are related to the probability of selecting each response as a function of one or more covariates. We combine particle swarm optimization and an Expectation-Maximization (EM) algorithm to obtain maximum likelihood estimates of the model parameters. We use this approach to analyze data for movement of three bobcats in relation to urban areas in southern California, USA. A behavioral interpretation of the models revealed similarities and differences in bobcat movement response to urbanization. All three bobcats avoided urbanization by moving either parallel to urban boundaries or toward less urban areas as the proportion of urban land cover in the surrounding area increased. However, one bobcat, a male with a dispersal-like large-scale movement pattern, avoided urbanization at lower densities and responded strictly by moving parallel to the urban edge. The other two bobcats, which were both residents and occupied similar geographic areas, avoided urban areas using a combination of movements parallel to the urban edge and movement toward areas of less urbanization. However, the resident female appeared to exhibit greater repulsion at lower levels of urbanization than the resident male, consistent with empirical observations of bobcats in southern California. Using the parameterized finite mixture models, we mapped behavioral states to geographic space, creating a representation of a behavioral landscape. This approach can provide guidance for conservation planning based on analysis of animal movement data using statistical models, thereby linking connectivity evaluations to empirical data.

Ecological Applications↗

Movement mysteries unveiled: spatial ecology of juvenile green sea turtles

Locations of important foraging areas are not well defined for many marine species. Unraveling these mysteries is vital to develop conservation strategies for these species, many of which are threatened or endangered. Satellite-tracking is a tool that can reveal movement patterns at both broad and fine spatial scales, in all marine environments. This chapter presents records of the longest duration track of an individual juvenile green turtle (434 days) and highest number of tracking days in any juvenile green turtle study (5483 tracking days) published to date. In this chapter, we use spatial modeling techniques to describe movements and identify foraging areas for juvenile green turtles ( Chelonia mydas ) captured in a developmental habitat in south Texas, USA. Some green turtles established residency in the vicinity of their capture and release site, but most used a specific habitat feature (i.e., a jettied pass) to travel between the Gulf of Mexico and a nearby bay. Still others moved southward within the Gulf of Mexico into Mexican coastal waters, likely in response to decreasing water temperatures. These movements to waters off the coast of Mexico highlight the importance of international cooperation in restoration efforts undertaken on behalf of this imperiled species.

Texas↗

Lake sturgeon behavioral diversity in the Laurentian Great Lakes: Migratory patterns across populations and habitats

Background Characterizing the diversity of migration behaviors from the individual to the population level is essential for understanding how organisms respond to environmental variation and how these responses affect survival and habitat use. Lake sturgeon ( Acipenser fulvescens ) is a species of special concern in the Laurentian Great Lakes that are long-lived and generally classified as intermittent, adfluvial spawners. Observations of lake sturgeon movements at ecologically relevant spatiotemporal scales have shown that migration behavior often varies among individuals within the same population. However, studies on individual populations, particularly when focused only on a part of the life cycle (e.g., often spawning), provide just a partial understanding of the species’ full migratory scope and processes underlying expression of different migratory behaviors. To better understand lake sturgeon migratory diversity, we characterized and compared migratory behaviors of six lake sturgeon populations occupying environments that varied in habitat availability and connectivity in different Laurentian Great Lakes. Methods Sequence analysis combined with agglomerative hierarchical clustering and visual inspection of daily location data were used to identify distinct lake sturgeon migratory behaviors present in each population. Results Seven distinct migratory behaviors were identified based on differential patterns of lake and river use that encompass spawning and other seasonal periods. Behaviors were categorized as annual spring river, intermittent spring river, intermittent two-step, annual summer river, annual winter river, and annual interlake migrants along with river residents. The presence and frequency of migratory behaviors varied substantially among populations. Conclusions Our study demonstrated that migratory diversity is a general feature of lake sturgeon life history that may be partially shaped by habitat availability and connectivity. Given these results, we propose a conceptual model that links habitat availability and connectivity to migratory diversity and predict a positive association between them. This updated framework provides a cohesive basis for understanding lake sturgeon migratory behavior across variable ecological contexts in the Laurentian Great Lakes and will help promote future research to refute or refine the model.

Laurentian Great Lakes↗

Ecological diffusion models are still useful in ecology

Context Connecting animal movement across scales—both among spatial scales and between individual and population level processes—is a central theme in ecology, including the sub-fields of movement and landscape ecology. However, most modeling frameworks operate at either the individual (Lagrangian) or population (Eulerian) level. Objectives We aimed to review how a partial differential equation (PDE) known as the ecological diffusion equation (EDE) offers connections among the scales of animal movement and how it can be embedded in commonly used statistical frameworks (e.g., hierarchical models) and fit to multiple types of data, such as count, presence-only, presence-absence, and telemetry data. In doing so, we also highlight emerging and promising avenues of research related to statistical EDE models. Methods We present how the EDE is derived from a simple set of first principles, can be used to represent both individual animal movement and spatiotemporal population dynamics, and can be fit to data by being applied in a hierarchical statistical framework. Finally, we review literature on recent applications and advances related to EDE models. Results In presenting the derivation of the EDE and reviewing recent advances and applications of EDE models, we established that the EDE can enable a mechanistic understanding of animal movement across scales, as well as the integration of multiple types of data. We also identified a number of topics where EDE models can be advanced to gain additional ecological insight. Conclusions Ecological diffusion is a process that emerges naturally from first principles governing animal movement and can be used to model both individual and population-level spatiotemporal dynamics. Statistical EDE models are related to other modeling frameworks, such as species distribution models, occupancy models, and step-selection analysis. The flexibility of hierarchical statistical modeling allows combining different types of data collected on movement and population dynamics, and the theoretical underpinnings of the EDE allow it to connect ecological processes occurring at fine and coarse spatial scales, as well as individual animal movement and population dynamics.

Landscape Ecology↗

Movement patterns of Brook Trout in a restored coastal stream system in southern Massachusetts

Coastal Brook Trout ( Salvelinus fontinalis ) populations are found from northern Canada to New England. The extent of anadromy generally decreases with latitude, but the ecology and movements of more southern populations are poorly understood. We conducted a 33-month acoustic telemetry study of Brook Trout in Red Brook, MA, and adjacent Buttermilk Bay (marine system) using 16 fixed acoustic receivers and surgically implanting acoustic transmitters in 84 individuals. Tagged Brook Trout used the stream, estuary (50% of individuals) and bay (10% of individuals). Movements into full sea water were brief when occurring. GAMM models revealed that transitions between habitat areas occurred most often in spring and fall. Environmental data suggest that use of the saline environment is limited by summer temperatures in the bay. Movements may also be related to moon phase. Compared to more northern coastal populations of Brook Trout, the Red Brook population appears to be less anadromous overall, yet the estuarine segment of the system may have considerable ecological importance as a food resource.

Ecology of Freshwater Fish↗

Agricultural land use shapes dispersal in white-tailed deer (Odocoileus virginianus)

Background Dispersal is a fundamental process to animal population dynamics and gene flow. In white-tailed deer (WTD; Odocoileus virginianus ), dispersal also presents an increasingly relevant risk for the spread of infectious diseases. Across their wide range, WTD dispersal is believed to be driven by a suite of landscape and host behavioral factors, but these can vary by region, season, and sex. Our objectives were to (1) identify dispersal events in Wisconsin WTD and determine drivers of dispersal rates and distances, and (2) determine how landscape features (e.g., rivers, roads) structure deer dispersal paths. Methods We developed an algorithmic approach to detect dispersal events from GPS collar data for 590 juvenile, yearling, and adult WTD. We used statistical models to identify host and landscape drivers of dispersal rates and distances, including the role of agricultural land use, the traversability of the landscape, and potential interactions between deer. We then performed a step selection analysis to determine how landscape features such as agricultural land use, elevation, rivers, and roads affected deer dispersal paths. Results Dispersal predominantly occurred in juvenile males, of which 64.2% dispersed, with dispersal events uncommon in other sex and age classes. Juvenile male dispersal probability was positively associated with the proportion of the natal range that was classified as agricultural land use, but only during the spring. Dispersal distances were typically short (median 5.77 km, range: 1.3–68.3 km), especially in the fall. Further, dispersal distances were positively associated with agricultural land use in potential dispersal paths but negatively associated with the number of proximate deer in the natal range. Lastly, we found that, during dispersal, juvenile males typically avoided agricultural land use but selected for areas near rivers and streams. Conclusion Land use—particularly agricultural—was a key driver of dispersal rates, distances, and paths in Wisconsin WTD. In addition, our results support the importance of deer social environments in shaping dispersal behavior. Our findings reinforce knowledge of dispersal ecology in WTD and how landscape factors—including major rivers, roads, and land-use patterns—structure host gene flow and potential pathogen transmission.

Wisconsin↗

Flexible characterization of animal movement pattern using net squared displacement and a latent state model

Background Characterizing the movement patterns of animals is an important step in understanding their ecology. Various methods have been developed for classifying animal movement at both coarse (e.g., migratory vs. sedentary behavior) and fine (e.g., resting vs. foraging) scales. A popular approach for classifying movements at coarse resolutions involves fitting time series of net-squared displacement (NSD) to models representing different conceptualizations of coarse movement strategies (i.e., migration, nomadism, sedentarism, etc.). However, the performance of this method in classifying actual (as opposed to simulated) animal movements has been mixed. Here, we develop a more flexible method that uses the same NSD input, but relies on an underlying discrete latent state model. Using simulated data, we first assess how well patterns in the number of transitions between modes of movement and the duration of time spent in a mode classify movement strategies. We then apply our approach to elucidate variability in the movement strategies of eight giant tortoises ( Chelonoidis sp.) using a multi-year (2009–2014) GPS dataset from three different Galapagos Islands. Results With respect to patterns of time spent and the number of transitions between modes, our approach out-performed previous efforts to distinguish among migration, dispersal, and sedentary behavior. We documented marked inter-individual variation in giant tortoise movement strategies, with behaviors indicating migration, dispersal, nomadism and sedentarism, as well as hybrid behaviors such as “exploratory residence”. Conclusions Distilling complex animal movement into discrete modes remains a fundamental challenge in movement ecology, a problem made more complex by the ever-longer duration, ever-finer resolution, and gap-ridden trajectories recorded by GPS devices. By clustering into modes, we derived information on the time spent within one mode and the number of transitions between modes which enabled finer differentiation of movement strategies over previous methods. Ultimately, the techniques developed here address limitations of previous approaches and provide greater insights with respect to characterization of movement strategies across scales by more fully utilizing long-term GPS telemetry datasets.

Movement Ecology↗

Diffusion modeling reveals effects of multiple release sites and human activity on a recolonizing apex predator

Background Reintroducing predators is a promising conservation tool to help remedy human-caused ecosystem changes. However, the growth and spread of a reintroduced population is a spatiotemporal process that is driven by a suite of factors, such as habitat change, human activity, and prey availability. Sea otters ( Enhydra lutris ) are apex predators of nearshore marine ecosystems that had declined nearly to extinction across much of their range by the early 20th century. In Southeast Alaska, which is comprised of a diverse matrix of nearshore habitat and managed areas, reintroduction of 413 individuals in the late 1960s initiated the growth and spread of a population that now exceeds 25,000. Methods Periodic aerial surveys in the region provide a time series of spatially-explicit data to investigate factors influencing this successful and ongoing recovery. We integrated an ecological diffusion model that accounted for spatially-variable motility and density-dependent population growth, as well as multiple population epicenters, into a Bayesian hierarchical framework to help understand the factors influencing the success of this recovery. Results Our results indicated that sea otters exhibited higher residence time as well as greater equilibrium abundance in Glacier Bay, a protected area, and in areas where there is limited or no commercial fishing. Asymptotic spread rates suggested sea otters colonized Southeast Alaska at rates of 1–8 km/yr with lower rates occurring in areas correlated with higher residence time, which primarily included areas near shore and closed to commercial fishing. Further, we found that the intrinsic growth rate of sea otters may be higher than previous estimates suggested. Conclusions This study shows how predator recolonization can occur from multiple population epicenters. Additionally, our results suggest spatial heterogeneity in the physical environment as well as human activity and management can influence recolonization processes, both in terms of movement (or motility) and density dependence.

Alaska↗

An integrated path for spatial capture–recapture and animal movement modeling

Ecologists and conservation biologists increasingly rely on spatial capture–recapture (SCR) and movement modeling to study animal populations. Historically, SCR has focused on population-level processes (e.g., vital rates, abundance, density, and distribution), whereas animal movement modeling has focused on the behavior of individuals (e.g., activity budgets, resource selection, migration). Even though animal movement is clearly a driver of population-level patterns and dynamics, technical and conceptual developments to date have not forged a firm link between the two fields. Instead, movement modeling has typically focused on the individual level without providing a coherent scaling from individual- to population-level processes, whereas SCR has typically focused on the population level while greatly simplifying the movement processes that give rise to the observations underlying these models. In our view, the integration of SCR and animal movement modeling has tremendous potential for allowing ecologists to scale up from individuals to populations and advancing the types of inferences that can be made at the intersection of population, movement, and landscape ecology. Properly accounting for complex animal movement processes can also potentially reduce bias in estimators of population-level parameters, thereby improving inferences that are critical for species conservation and management. This introductory article to the Special Feature reviews recent advances in SCR and animal movement modeling, establishes a common notation, highlights potential advantages of linking individual-level (Lagrangian) movements to population-level (Eulerian) processes, and outlines a general conceptual framework for the integration of movement and SCR models. We then identify important avenues for future research, including key challenges and potential pitfalls in the developments and applications that lie ahead.

Ecology↗

Application of isoscapes to determine geographic origin of terrestrial wildlife for conservation and management

Accounting for migration and connectivity of mobile species across the annual cycle can present challenges for conservation and management efforts. The use of stable isotope approaches to examine the movements and ecology of wildlife has been widespread over the past two decades. Hydrogen stable isotope ( δ 2 H) composition, in particular, has been frequently used to provide insight into the origin of migratory species, although isotopes of other elements are sometimes used. These intrinsic markers can yield valuable information about distributions of wildlife on a broad scale, with reduced labor and expense compared to tracking and telemetry. Many of the applications of isotopes to migratory species to date have addressed connectivity and origin, and studies in support of conservation biology are less common. In addition, there are few guides for how to best employ these methods for management. Therefore, we provide an overview for the wildlife conservation and management community on how stable isotope methods may be applied to conservation problems and a primer on the process for assigning geographic origins to terrestrial wildlife. We also discuss best practices for employing environmental isoscapes (isotopic distributions across landscapes), rescaling functions, and the assumptions required for assignment to origin while highlighting emerging issues in the modeling process. Finally, we provide example applications to illustrate these principles, and we explore strengths and limitations of this approach in a conservation context.

Biological Conservation↗

Social influences complement environmental cues to stimulate migrating juvenile salmon

Background The large-scale seasonal migrations undertaken by many species require complex navigational and timing decisions. Animals migrating in groups might benefit from collective decision making, especially if the environment has large local variation rather than smooth gradients in, for example, salinity or temperature, or is unpredictable, or if the migrants cannot rely on individually acquired information. We focus on juvenile salmon whose downstream migration is timed to match suitable conditions for growth and survival at sea. While the environmental and physiological factors that influence the timing of migration have been well studied, the influence of social interactions on migration timing is poorly understood. Method We compiled data on two species of juvenile salmon, collected at traps over 19 years, during their downstream seaward migration in three rivers in Washington state along with relevant environmental data. We developed state space statistical models to estimate the influence of hatchery-produced salmon in stimulating the downstream migration of wild salmon, while also incorporating potential environmental stimuli. Results Our results are consistent with the “pied-piper” hypothesis that large numbers of migrating hatchery-origin salmon provide a social cue stimulating migration of co-occurring wild salmon. The increase in the number of hatchery salmon counted at the trap was a strong predictor of the increase in wild sub-yearling Chinook salmon in the Dungeness, Puyallup, and Skagit rivers and yearling coho salmon in the Puyallup and Skagit rivers. Migration timing was also associated with abiotic factors related to temperature, river flow, and time of year. Conclusions Our findings highlight the potential for social cues to affect migration timing of downstream migrating salmon, in concert with environmental factors. Incorporating social information into timing decisions may allow animals to benefit from collective decision-making strategies and better time their migrations. Moreover, understanding the effects of large-scale hatchery releases on wild salmon migration may provide valuable insights for planning the timing and duration of hatchery releases.

British Columbia, Washington↗

The environmental-data automated track annotation (Env-DATA) system: linking animal tracks with environmental data

The movement of animals is strongly influenced by external factors in their surrounding environment such as weather, habitat types, and human land use. With advances in positioning and sensor technologies, it is now possible to capture animal locations at high spatial and temporal granularities. Likewise, scientists have an increasing access to large volumes of environmental data. Environmental data are heterogeneous in source and format, and are usually obtained at different spatiotemporal scales than movement data. Indeed, there remain scientific and technical challenges in developing linkages between the growing collections of animal movement data and the large repositories of heterogeneous remote sensing observations, as well as in the developments of new statistical and computational methods for the analysis of movement in its environmental context. These challenges include retrieval, indexing, efficient storage, data integration, and analytical techniques.

Movement Ecology↗

Flying with the wind: Scale dependency of speed and direction measurements in modelling wind support in avian flight

Background : Understanding how environmental conditions, especially wind, influence birds' flight speeds is a prerequisite for understanding many important aspects of bird flight, including optimal migration strategies, navigation, and compensation for wind drift. Recent developments in tracking technology and the increased availability of data on large-scale weather patterns have made it possible to use path annotation to link the location of animals to environmental conditions such as wind speed and direction. However, there are various measures available for describing not only wind conditions but also the bird's flight direction and ground speed, and it is unclear which is best for determining the amount of wind support (the length of the wind vector in a bird’s flight direction) and the influence of cross-winds (the length of the wind vector perpendicular to a bird’s direction) throughout a bird's journey. Results : We compared relationships between cross-wind, wind support and bird movements, using path annotation derived from two different global weather reanalysis datasets and three different measures of direction and speed calculation for 288 individuals of nine bird species. Wind was a strong predictor of bird ground speed, explaining 10-66% of the variance, depending on species. Models using data from different weather sources gave qualitatively similar results; however, determining flight direction and speed from successive locations, even at short (15 min intervals), was inferior to using instantaneous GPS-based measures of speed and direction. Use of successive location data significantly underestimated the birds' ground and airspeed, and also resulted in mistaken associations between cross-winds, wind support, and their interactive effects, in relation to the birds' onward flight. Conclusions : Wind has strong effects on bird flight, and combining GPS technology with path annotation of weather variables allows us to quantify these effects for understanding flight behaviour. The potentially strong influence of scaling effects must be considered and implemented in developing sampling regimes and data analysis.

Movement Ecology↗

When to be discrete: The importance of time formulation in understanding animal movement

Animal movement is essential to our understanding of population dynamics, animal behavior, and the impacts of global change. Coupled with high-resolution biotelemetry data, exciting new inferences about animal movement have been facilitated by various specifications of contemporary models. These approaches differ, but most share common themes. One key distinction is whether the underlying movement process is conceptualized in discrete or continuous time. This is perhaps the greatest source of confusion among practitioners, both in terms of implementation and biological interpretation. In general, animal movement occurs in continuous time but we observe it at fixed discrete-time intervals. Thus, continuous time is conceptually and theoretically appealing, but in practice it is perhaps more intuitive to interpret movement in discrete intervals. With an emphasis on state-space models, we explore the differences and similarities between continuous and discrete versions of mechanistic movement models, establish some common terminology, and indicate under which circumstances one form might be preferred over another. Counter to the overly simplistic view that discrete- and continuous-time conceptualizations are merely different means to the same end, we present novel mathematical results revealing hitherto unappreciated consequences of model formulation on inferences about animal movement. Notably, the speed and direction of movement are intrinsically linked in current continuous-time random walk formulations, and this can have important implications when interpreting animal behavior. We illustrate these concepts in the context of state-space models with multiple movement behavior states using northern fur seal ( Callorhinus ursinus ) biotelemetry data.

Movement Ecology↗

Mapping migratory flyways in Asia using dynamic Brownian bridge movement models

Background Identifying movement routes and stopover sites is necessary for developing effective management and conservation strategies for migratory animals. In the case of migratory birds, a collection of migration routes, known as a flyway, is often hundreds to thousands of kilometers long and can extend across political boundaries. Flyways encompass the entire geographic range between the breeding and non-breeding areas of a population, species, or a group of species, and they provide spatial frameworks for management and conservation across international borders. Existing flyway maps are largely qualitative accounts based on band returns and survey data rather than observed movement routes. In this study, we use satellite and GPS telemetry data and dynamic Brownian bridge movement models to build upon existing maps and describe waterfowl space use probabilistically in the Central Asian and East Asian-Australasian Flyways. Results Our approach provided new information on migratory routes that was not easily attainable with existing methods to describe flyways. Utilization distributions from dynamic Brownian bridge movement models identified key staging and stopover sites, migration corridors and general flyway outlines in the Central Asian and East Asian-Australasian Flyways. A map of space use from ruddy shelducks depicted two separate movement corridors within the Central Asian Flyway, likely representing two distinct populations that show relatively strong connectivity between breeding and wintering areas. Bar-headed geese marked at seven locations in the Central Asian Flyway showed heaviest use at several stopover sites in the same general region of high-elevation lakes along the eastern Qinghai-Tibetan Plateau. Our analysis of data from multiple Anatidae species marked at sites throughout Asia highlighted major movement corridors across species and confirmed that the Central Asian and East Asian-Australasian Flyways were spatially distinct. Conclusions The dynamic Brownian bridge movement model improves our understanding of flyways by estimating relative use of regions in the flyway while providing detailed, quantitative information on migration timing and population connectivity including uncertainty between locations. This model effectively quantifies the relative importance of different migration corridors and stopover sites and may help prioritize specific areas in flyways for conservation of waterbird populations.

Movement Ecology↗

Is there a single best estimator? selection of home range estimators using area- under- the-curve

Background Global positioning system (GPS) technology for monitoring home range and movements of wildlife has resulted in prohibitively large sample sizes of locations for traditional estimators of home range. We used area-under-the-curve to explore the fit of 8 estimators of home range to data collected with both GPS and concurrent very high frequency (VHF) technology on a terrestrial mammal, the Florida panther Puma concolor coryi , to evaluate recently developed and traditional estimators. Results Area-under-the-curve was the highest for Florida panthers equipped with Global Positioning System (GPS) technology compared to VHF technology. For our study animal, estimators of home range that incorporated a temporal component to estimation performed better than traditional first- and second-generation estimators. Conclusions Comparisons of fit of home range contours with locations collected would suggest that use of VHF technology is not as accurate as GPS technology to estimate size of home range for large mammals. Estimators of home range collected with GPS technology performed better than those estimated with VHF technology regardless of estimator used. Furthermore, estimators that incorporate a temporal component (third-generation estimators) appeared to be the most reliable regardless of whether kernel-based or Brownian bridge-based algorithms were used and in comparison to first- and second-generation estimators. We defined third-generation estimators of home range as any estimator that incorporates time, space, animal-specific parameters, and habitat. Such estimators would include movement-based kernel density, Brownian bridge movement models, and dynamic Brownian bridge movement models among others that have yet to be evaluated.

Florida↗

Metapopulation viability of an endangered shorebird depends on dispersal and human-created habitats: Piping plovers ( Charadrius melodus ) and prairie rivers

Background Many species are distributed as metapopulations in dynamic landscapes, where habitats change through space and time. Individuals locate habitat through dispersal, and the relationship between a species and landscape characteristics can have profound effects on population persistence. Despite the importance of connectivity in dynamic environments, few empirical studies have examined temporal variability in dispersal or its effect on metapopulation dynamics. In response to this knowledge gap, we studied the dispersal, demography, and viability of a metapopulation of an endangered, disturbance-dependent shorebird. We examined three subpopulations of piping plovers ( Charadrius melodus ) on the lower Platte and Missouri rivers from 2008–2013. High flow events from an upstream dam on the Missouri River in 2010 and 2011 allowed us to assess the effect of total habitat loss and the subsequent creation of new habitat associated with a large disturbance at one ‘natural’ study location. The other two sites within the metapopulation, which were maintained by anthropogenic activities (e.g., mining, development, habitat restoration), were largely unaffected by this disturbance, resulting in a controlled natural experiment. Results High flow events were associated with increased emigration, decreased immigration, and decreased survival in the subpopulation that experienced high flows. Following the high flow event, immigration into that subpopulation increased. Dispersal rates among subpopulations were negatively correlated with distance. The metapopulation had a low probability of extinction over 100 years (0%) under the current disturbance interval and associated dispersal and survival rates. However, persistence depended on relatively stable, human-created habitats, not the dynamic, natural habitat (47.7 % extinction probability for this subpopulation). Conclusions We found that functional connectivity, as measured by the rate of dispersal among subpopulations, increased as a result of the high flow event in our study metapopulation. Plovers also increased reproductive output following this event. Although the study metapopulation had a low overall probability of extinction, metapopulation persistence depended on anthropogenically created habitats that provided a small but stable source of nesting habitat and dispersers through time. However, all subpopulations remained small, even if persistent, making them individually vulnerable to extinction through stochastic events. Given the highly dynamic nature of habitat availability in this system, maintaining several subpopulations within the metapopulation and stable sources of habitat will be critical, and this species will likely remain conservation-reliant.

Nebraska↗

Demographic outcomes of diverse migration strategies assessed in a metapopulation of tundra swans

Background Migration is a prominent aspect of the life history of many avian species, but the demographic consequences of variable migration strategies have only infrequently been investigated, and rarely when using modern technological and analytical methods for assessing survival, movement patterns, and long-term productivity in the context of life history theory. We monitored the fates of 50 satellite-implanted tundra swans (Cygnus columbianus) over 4 years from five disparate breeding areas in Alaska, and used known-fate analyses to estimate monthly survival probability relative to migration distance, breeding area, migratory flyway, breeding status, and age. We specifically tested whether migratory birds face a trade-off, whereby long-distance migrants realize higher survival rates at the cost of lower productivity because of reduced time on breeding areas relative to birds that migrate shorter distances and spend more time on breeding areas. Results Annual migration distances varied significantly among breeding areas (1020 to 12720 km), and were strongly negatively correlated with time spent on breeding areas (r = −0.986). Estimates of annual survival probability varied by wintering area (Pacific coast, Alaska Peninsula, and Eastern seaboard) and ranged from 0.79 (95%CI: 0.70–0.88) to 1.0, depending on criteria used to discern mortalities from radio failures. We did not find evidence for a linear relationship between migration distance and survival as swans from the breeding areas with the shortest and longest migration distances had the highest survival probabilities. Survival was lower in the first year post-marking than in subsequent years, but there was not support for seasonal differences in survival. Productivity varied among breeding populations and was generally inversely correlated to survival, but not migration distance or time spent on breeding areas. Conclusions Tundra swans conformed to a major tenet of life history theory, as populations with the highest survival generally had the lowest productivity. The lack of a uniform relationship between time spent on breeding areas and productivity, or time spent on wintering areas and survival, indicates that factors other than temporal investment dictate demographic outcomes in this species. The tremendous diversity of migration strategies we identify in Alaskan tundra swans, without clear impacts on survival, underscores the ability of this species to adapt to different environments and climatic regimes. Keywords: Cygnus columbianus, Known fate, Life history, Metapopulation, Migration distance, Productivity, Satellite telemetry, Survival, Transmitter effects, Tundra swan

Movement Ecology↗