An introduction to decision science for conservation.

Biodiversity conservation decisions are difficult, especially when they involve differing values, complex multidimensional objectives, scarce resources, urgency, and considerable uncertainty. Decision science embodies a theory about how to make difficult decisions and an extensive array of frameworks and tools that make that theory practical. We sought to improve conceptual clarity and practical application of decision science to help decision makers apply decision science to conservation problems. We addressed barriers to the uptake of decision science, including a lack of training and awareness of decision science; confusion over common terminology and which tools and frameworks to apply; and the mistaken impression that applying decision science must be time consuming, expensive, and complex. To aid in navigating the extensive and disparate decision science literature, we clarify meaning of common terms: decision science, decision theory, decision analysis, structured decision-making, and decision-support tools. Applying decision science does not have to be complex or time consuming; rather, it begins with knowing how to think through the components of a decision utilizing decision analysis (i.e., define the problem, elicit objectives, develop alternatives, estimate consequences, and perform trade-offs). This is best achieved by applying a rapid-prototyping approach. At each step, decision-support tools can provide additional insight and clarity, whereas decision-support frameworks (e.g., priority threat management and systematic conservation planning) can aid navigation of multiple steps of a decision analysis for particular contexts. We summarize key decision-support frameworks and tools and describe to which step of a decision analysis, and to which contexts, each is most useful to apply. Our introduction to decision science will aid in contextualizing current approaches and new developments, and help decision makers begin to apply decision science to conservation problems.

Las decisiones sobre la conservación de la biodiversidad son difíciles de tomar, especialmente cuando involucran diferentes valores, objetivos multidimensionales complejos, recursos limitados, urgencia y una incertidumbre considerable. Las ciencias de la decisión incorporan una teoría sobre cómo tomar decisiones difíciles y una variedad extensa de marcos de trabajo y herramientas que transforman esa teoría en práctica. Buscamos mejorar la claridad conceptual y la aplicación práctica de las ciencias de la decisión para ayudar al órgano decisorio a aplicar estas ciencias a los problemas de conservación. Nos enfocamos en las barreras para la aceptación de las ciencias de la decisión, incluyendo la falta de capacitación y de conciencia por estas ciencias; la confusión por la terminología común y cuáles herramientas y marcos de trabajo aplicar; y la impresión errónea de que la aplicación de estas ciencias consume tiempo y debe ser costosa y compleja. Para asistir en la navegación de la literatura extensa y dispar de las ciencias de la decisión, aclaramos el significado de varios términos comunes: ciencias de la decisión, teoría de la decisión, análisis de decisiones, toma estructurada de decisiones y herramientas de apoyo para las decisiones. La aplicación de las ciencias de la decisión no tiene que ser compleja ni debe llevar mucho tiempo; de hecho, todo comienza con saber cómo pensar detenidamente en los componentes de una decisión mediante el análisis de decisiones (es decir, definir el problema, producir objetivos, desarrollar alternativas, estimar consecuencias y realizar compensaciones). Lo anterior se logra de mejor manera mediante la aplicación de una estrategia prototipos rápidos. En cada paso, las herramientas de apoyo para las decisiones pueden proporcionar visión y claridad adicionales, mientras que los marcos de apoyo para las decisiones (p.ej.: gestión de amenazas prioritarias y planeación sistemática de la conservación) pueden asistir en la navegación de los diferentes pasos de un análisis de decisiones para contextos particulares. Resumimos los marcos de trabajo y las herramientas más importantes de apoyo para las decisiones y describimos el paso, y el contexto, del análisis de decisiones para el que es más útil aplicarlos. Nuestra introducción a las ciencias de la decisión apoyará en la contextualización de las estrategias actuales y los nuevos desarrollos, y ayudarán al órgano decisorio a comenzar a aplicar estas ciencias en los problemas de conservación.

Demographic risk assessment for a harvested species threatened by climate change: polar bears in the Chukchi Sea.

Climate change threatens global biodiversity. Many species vulnerable to climate change are important to humans for nutritional, cultural, and economic reasons. Polar bears Ursus maritimus are threatened by sea-ice loss and represent a subsistence resource for Indigenous people. We applied a novel population modeling-management framework that is based on species life history and accounts for habitat loss to evaluate subsistence harvest for the Chukchi Sea (CS) polar bear subpopulation. Harvest strategies followed a state-dependent approach under which new data were used to update the harvest on a predetermined management interval. We found that a harvest strategy with a starting total harvest rate of 2.7% (˜85 bears/yr at current abundance), a 2:1 male-to-female ratio, and a 10-yr management interval would likely maintain subpopulation abundance above maximum net productivity level for the next 35 yr (approximately three polar bear generations), our primary criterion for sustainability. Plausible bounds on starting total harvest rate were 1.7-3.9%, where the range reflects uncertainty due to sampling variation, environmental variation, model selection, and differing levels of risk tolerance. The risk of undesired demographic outcomes (e.g., overharvest) was positively related to harvest rate, management interval, and projected declines in environmental carrying capacity; and negatively related to precision in population data. Results reflect several lines of evidence that the CS subpopulation has been productive in recent years, although it is uncertain how long this will last as sea-ice loss continues. Our methods provide a template for balancing trade-offs among protection, use, research investment, and other factors. Demographic risk assessment and state-dependent management will become increasingly important for harvested species, like polar bears, that exhibit spatiotemporal variation in their response to climate change.

The dilemma of pest suppression in the conservation of endangered species.

In the conservation of endangered species, suppression of a population of one native species to benefit another poses challenges. Examples include predator control and nest parasite reduction. Less obvious is the control of blood-feeding arthropods. We conducted a case study of the effect of native black flies (Simulium spp.) on reintroduced Whooping Cranes (Grus americana). Our intent was to provide a science-driven approach for determining the effects of blood-feeding arthropods on endangered vertebrates and identifying optimal management actions for managers faced with competing objectives. A multiyear experiment demonstrated that black flies reduce nest success in cranes by driving incubating birds off their nests. We used a decision-analytic approach to develop creative management alternatives and evaluate trade-offs among competing objectives. We identified 4 management objectives: establish a self-sustaining crane population, improve crane well-being, maintain native black flies as functional components of the ecosystem, and minimize costs. We next identified potential management alternatives: do nothing, suppress black flies, force crane renesting to occur after the activity period of black flies, relocate releases of cranes, suppress black flies and relocate releases, or force crane renesting and relocate releases. We then developed predictions on constructed scales of 0 (worst-performing alternative) to 1 (best-performing alternative) to indicate how alternative actions performed in terms of management objectives. The optimal action depended on the relative importance of each objective to a decision maker. Only relocating releases was a dominated alternative, indicating that it was not optimal regardless of the relative importance of objectives. A rational decision maker could choose any other management alternative we considered. Recognizing that decisions involve trade-offs that must be weighed by decision makers is crucial to identifying alternatives that best balance multiple management objectives. Given uncertainty about the population dynamics of blood-feeding arthropods, an adaptive management approach could offer substantial benefits.

El Dilema de la Eliminación de Plagas en la Conservación de Especies en Peligro Resumen En la conservación de especies en peligro, la eliminación de la población de una especie nativa para beneficiar a otra representa un reto. Los ejemplos incluyen el control de depredadores y la reducción de parásitos de nido. El control de artrópodos hematófagos es menos obvio. Realizamos un estudio de caso sobre el efecto que las moscas negras nativas (Simulium spp.) tienen sobre las grullas trompeteras (Grus americana) reintroducidas. Nuestra intención era proporcionar una estrategia científica para determinar los efectos que tienen los artrópodos hematófagos sobre los vertebrados en peligro de extinción e identificar las acciones de manejo óptimo para los administradores que enfrentan objetivos en competencia. Un experimento multianual demostró que las moscas negras reducen el éxito de anidación de las grullas al ahuyentar a las aves incubadoras de sus nidos. Usamos una estrategia analítica de decisión para desarrollar alternativas creativas de manejo y para evaluar las compensaciones entre los objetivos en competencia. Identificamos cuatro objetivos de manejo: establecer una población autosustentable de grullas, mejorar el bienestar de las grullas, mantener a las moscas negras nativas como componentes funcionales del ecosistema, y minimizar los costos. Después, identificamos las alternativas potenciales de manejo: no realizar acciones, eliminar a las moscas, obligar a que la re-anidación de las grullas ocurra después del periodo de actividad de las moscas, reubicar las liberaciones de las grullas, eliminar a las moscas y reubicar las liberaciones, o forzar la re-anidación de las grullas y reubicar las liberaciones. Finalmente, desarrollamos predicciones sobre escalas construidas de 0 (la alternativa con el peor desempeño) a 1 (la alternativa con el mejor desempeño) para indicar el éxito de las acciones alternativas con respecto a los objetivos de manejo. La acción óptima dependió de la importancia relativa que cada objetivo tenía para un responsable de la conservación. Sólo la reubicación de las liberaciones fue una alternativa dominada, lo que indica que no era óptima a pesar de la importancia relativa de sus objetivos. Un encargado razonable podría entonces elegir de entre cualquier otra de las alternativas de manejo que consideramos. El reconocimiento de las decisiones que involucran compensaciones que deben ser sopesadas por los encargados de la conservación es importante para la identificación de alternativas que balancean de mejor manera los objetivos múltiples del manejo. Dada la incertidumbre sobre las dinámicas poblacionales de los artrópodos hematófagos, una estrategia de manejo adaptativo podría ofrecer beneficios sustanciales.

Implicit decision framing as an unrecognized source of confusion in endangered species classification.

Legal classification of species requires scientific and values-based components, and how those components interact depends on how people frame the decision. Is classification a negotiation of trade-offs, a decision on how to allocate conservation efforts, or simply a comparison of the biological status of a species to a legal standard? The answers to problem-framing questions such as these influence decision making in species classifications. In our experience, however, decision makers, staff biologists, and stakeholders often have differing perspectives of the decision problem and assume different framings. In addition to differences between individuals, in some cases it appears individuals themselves are unclear about the decision process, which contributes to regulatory paralysis, litigation, and a loss of trust by agency staff and the public. We present 5 framings: putting species in the right bin, doing right by the species over time, saving the most species on a limited budget, weighing extinction risk against other objectives, and strategic classification to advance conservation. These framings are inspired by elements observed in current classification practices. Putting species in the right bin entails comparing a scientific status assessment with policy thresholds and accounting for potential misclassification costs. Doing right by the species adds a time dimension to the classification decision, and saving the most species on a limited budget classifies a suite of species simultaneously. Weighing extinction risk against other objectives would weigh ecological or socioeconomic concerns in classification decisions, and strategic classification to advance conservation would make negotiation a component of classification. We view these framings as a means to generate thought, discussion, and movement toward selection and application of explicit classification framings. Being explicit about the decision framing could lead decision makers toward more efficient and defensible decisions, reduce internal confusion and external conflict, and support better collaboration between scientists and policy makers.

Population size and stopover duration estimation using mark-resight data and Bayesian analysis of a superpopulation model.

We present a novel formulation of a mark-recapture-resight model that allows estimation of population size, stopover duration, and arrival and departure schedules at migration areas. Estimation is based on encounter histories of uniquely marked individuals and relative counts of marked and unmarked animals. We use a Bayesian analysis of a state-space formulation of the Jolly-Seber mark-recapture model, integrated with a binomial model for counts of unmarked animals, to derive estimates of population size and arrival and departure probabilities. We also provide a novel estimator for stopover duration that is derived from the latent state variable representing the interim between arrival and departure in the state-space model. We conduct a simulation study of field sampling protocols to understand the impact of superpopulation size, proportion marked, and number of animals sampled on bias and precision of estimates. Simulation results indicate that relative bias of estimates of the proportion of the population with marks was low for all sampling scenarios and never exceeded 2%. Our approach does not require enumeration of all unmarked animals detected or direct knowledge of the number of marked animals in the population at the time of the study. This provides flexibility and potential application in a variety of sampling situations (e.g., migratory birds, breeding seabirds, sea turtles, fish, pinnipeds, etc.). Application of the methods is demonstrated with data from a study of migratory sandpipers.

Demography of an apex predator at the edge of its range: impacts of changing sea ice on polar bears in Hudson Bay.

Changes in the abundance and distribution of wildlife populations are common consequences of historic and contemporary climate change. Some Arctic marine mammals, such as the polar bear (Ursus maritimus), may be particularly vulnerable to such changes due to the loss of Arctic sea ice. We evaluated the impacts of environmental variation on demographic rates for the Western Hudson Bay (WH), polar bear subpopulation from 1984 to 2011 using live-recapture and dead-recovery data in a Bayesian implementation of multistate capture-recapture models. We found that survival of female polar bears was related to the annual timing of sea ice break-up and formation. Using estimated vital rates (e.g., survival and reproduction) in matrix projection models, we calculated the growth rate of the WH subpopulation and projected population responses under different environmental scenarios while accounting for parametric uncertainty, temporal variation, and demographic stochasticity. Our analysis suggested a long-term decline in the number of bears from 1185 (95% Bayesian credible interval [BCI] = 993-1411) in 1987 to 806 (95% BCI = 653-984) in 2011. In the last 10 yr of the study, the number of bears appeared stable due to temporary stability in sea ice conditions (mean population growth rate for the period 2001-2010 = 1.02, 95% BCI = 0.98-1.06). Looking forward, we estimated long-term growth rates for the WH subpopulation of ~1.02 (95% BCI = 1.00-1.05) and 0.97 (95% BCI = 0.92-1.01) under hypothetical high and low sea ice conditions, respectively. Our findings support previous evidence for a demographic linkage between sea ice conditions and polar bear population dynamics. Furthermore, we present a robust framework for sensitivity analysis with respect to continued climate change (e.g., to inform scenario planning) and for evaluating the combined effects of climate change and management actions on the status of wildlife populations.

Planning for ex situ conservation in the face of uncertainty.

Ex situ conservation strategies for threatened species often require long-term commitment and financial investment to achieve management objectives. We present a framework that considers the decision to adopt ex situ management for a target species as the end point of several linked decisions. We used a decision tree to intuitively represent the logical sequence of decision making. The first decision is to identify the specific management actions most likely to achieve the fundamental objectives of the recovery plan, with or without the use of ex-situ populations. Once this decision has been made, one decides whether to establish an ex situ population, accounting for the probability of success in the initial phase of the recovery plan, for example, the probability of successful breeding in captivity. Approaching these decisions in the reverse order (attempting to establish an ex situ population before its purpose is clearly defined) can lead to a poor allocation of resources, because it may restrict the range of available decisions in the second stage. We applied our decision framework to the recovery program for the threatened spotted tree frog (Litoria spenceri) of southeastern Australia. Across a range of possible management actions, only those including ex situ management were expected to provide >50% probability of the species' persistence, but these actions cost more than use of in situ alternatives only. The expected benefits of ex situ actions were predicted to be offset by additional uncertainty and stochasticity associated with establishing and maintaining ex situ populations. Naïvely implementing ex situ conservation strategies can lead to inefficient management. Our framework may help managers explicitly evaluate objectives, management options, and the probability of success prior to establishing a captive colony of any given species.

Female gonadal hormones and reproductive behaviors as key determinants of successful reproductive output of breeding whooping cranes (Grus americana).

Reproductive success of endangered whooping cranes (Grus americana) maintained ex situ is poor. As part of an effort to identify potential causes of poor reproductive success in a captive colony, we used non-invasive endocrine monitoring to assess gonadal and adrenal steroids of bird pairs with various reproductive outcomes and evaluated the relationships of hormones and behaviors to reproductive performance. Overall, reproductively successful (i.e., egg laying) females had significantly higher mean estrogen levels but lower mean progestogen concentrations than did unsuccessful females. Other hormones, including glucocorticoids and androgens, were not significantly different between successful and unsuccessful individuals. Observations of specific behaviors such as unison calling, marching, and the number of copulation attempts, along with overall time spent performing reproductive behaviors, were significantly higher in successful pairs. Our findings indicate that overall reproductive performance of whooping crane pairs is linked to female gonadal hormone excretion and reproductive behaviors, but not to altered adrenal hormone production.

Demography of a reintroduced population: moving toward management models for an endangered species, the Whooping Crane.

The reintroduction of threatened and endangered species is now a common method for reestablishing populations. Typically, a fundamental objective of reintroduction is to establish a self-sustaining population. Estimation of demographic parameters in reintroduced populations is critical, as these estimates serve multiple purposes. First, they support evaluation of progress toward the fundamental objective via construction of population viability analyses (PVAs) to predict metrics such as probability of persistence. Second, PVAs can be expanded to support evaluation of management actions, via management modeling. Third, the estimates themselves can support evaluation of the demographic performance of the reintroduced population (e.g., via comparison with wild populations). For each of these purposes, thorough treatment of uncertainties in the estimates is critical. Recently developed statistical methods (namely, hierarchical Bayesian implementations of state-space models) allow for effective integration of different types of uncertainty in estimation. We undertook a demographic estimation effort for a reintroduced population of endangered Whooping Cranes with the purpose of ultimately developing a Bayesian PVA for determining progress toward establishing a self-sustaining population, and for evaluating potential management actions via a Bayesian PVA-based management model. We evaluated individual and temporal variation in demographic parameters based upon a multi-state, mark-recapture model. We found that survival was relatively high across time and varied little by sex. There was some indication that survival varied by release method. Survival was similar to that observed in the wild population. Although overall reproduction in this reintroduced population is poor, birds formed social pairs when relatively young, and once a bird was in a social pair, it had a nearly 50% chance of nesting the following breeding season. Also, once a bird had nested, it had a high probability of nesting again. These results are encouraging, considering that survival and reproduction have been major challenges in past reintroductions of this species. The demographic estimates developed will support construction of a management model designed to facilitate exploration of management actions of interest, and will provide critical guidance in future planning for this reintroduction. An approach similar to what we describe could be usefully applied to many reintroduced populations.

Effects of population density and environmental conditions on life-history prevalence in a migratory fish.

Individual variation in life-history traits can have important implications for the ability of populations to respond to environmental variability and change. In migratory animals, flexibility in the timing of life-history events, such as juvenile emigration from natal areas, can influence the effects of population density and environmental conditions on habitat use and population dynamics. We evaluated the functional relationships between population density and environmental covariates and the abundance of juveniles expressing different life-history pathways in a migratory fish, Chinook salmon (Oncorhynchus tshawytscha), in the Wenatchee River basin in Washington State, USA. We found that the abundance of younger emigrants from natal streams was best described by an accelerating or near-linear function of spawners, whereas the abundance of older emigrants was best described by a decelerating function of spawners. This supports the hypothesis that emigration timing varies in response to density in natal areas, with younger-emigrating life-history pathways comprising a larger proportion of emigrants when densities of conspecifics are high. We also observed positive relationships between winter stream discharge and abundance of younger emigrants, supporting the hypothesis that habitat conditions can also influence the prevalence of different life-history pathways. Our results suggest that early emigration, and a resultant increase in the use of downstream rearing habitats, may increase at higher population densities and with greater winter precipitation. Winter precipitation is projected to increase in this system due to climate warming. Characterizing relationships between life-history prevalence and environmental conditions may improve our understanding of species habitat requirements and is a first step in understanding the dynamics of species with diverse life-history strategies. As environmental conditions change-due to climate change, management, or other factors-resultant life-history changes are likely to have important demographic implications that will be challenging to predict when life-history diversity is not accounted for in population models.

Do predators control prey species abundance? An experimental test with brown treesnakes on Guam.

The effect of predators on the abundance of prey species is a topic of ongoing debate in ecology; the effect of snake predators on their prey has been less debated, as there exists a general consensus that snakes do not negatively influence the abundance of their prey. However, this viewpoint has not been adequately tested. We quantified the effect of brown treesnake (Boiga irregularis) predation on the abundance and size of lizards on Guam by contrasting lizards in two 1-ha treatment plots of secondary forest from which snakes had been removed and excluded vs. two 1-ha control plots in which snakes were monitored but not removed or excluded. We removed resident snakes from the treatment plots with snake traps and hand capture, and snake immigration into these plots was precluded by electrified snake barriers. Lizards were sampled in all plots quarterly for a year following snake elimination in the treatment plots. Following the completion of this experiment, we used total removal sampling to census lizards on a 100-m2 subsample of each plot. Results of systematic lizard population monitoring before and after snake removal suggest that the abundance of the skink, Carlia ailanpalai, increased substantially and the abundance of two species of gekkonids, Lepidodactylus lugubris and Hemidactylus frenatus, also increased on snake-free plots. No treatment effect was observed for the skink Emoia caeruleocauda. Mean snout-vent length of all lizard species only increased following snake removal in the treatment plots. The general increase in prey density and mean size was unexpected in light of the literature consensus that snakes do not control the abundance of their prey species. Our findings show that, at least where alternate predators are lacking, snakes may indeed affect prey populations.

Integrated animal movement and spatial capture-recapture models: Simulation, implementation, and inference.

Over the last decade, spatial capture-recapture (SCR) models have become widespread for estimating demographic parameters in ecological studies. However, the underlying assumptions about animal movement and space use are often not realistic. This is a missed opportunity because interesting ecological questions related to animal space use, habitat selection, and behavior cannot be addressed with most SCR models, despite the fact that the data collected in SCR studies - individual animals observed at specific locations and times - can provide a rich source of information about these processes and how they relate to demographic rates. We developed SCR models that integrated more complex movement processes that are typically inferred from telemetry data, including a simple random walk, correlated random walk (i.e., short-term directional persistence), and habitat-driven Langevin diffusion. We demonstrated how to formulate, simulate from, and fit these models with standard SCR data using data-augmented Bayesian analysis methods. We evaluated their performance through a simulation study, in which we varied the detection, movement, and resource selection parameters. We also examined different numbers of sampling occasions and assessed performance gains when including auxiliary location data collected from telemetered individuals. Across all scenarios, the integrated SCR movement models performed well in terms of abundance, detection, and movement parameter estimation. We found little difference in bias for the simple random walk model when reducing the number of sampling occasions from T = 25 to T = 15. We found some bias in movement parameter estimates under several of the correlated random walk scenarios, but incorporating auxiliary location data improved parameter estimates and significantly improved mixing during model fitting. The Langevin movement model was able to recover resource selection parameters from standard SCR data, which is particularly appealing because it explicitly links the individual-level movement process with habitat selection and population density. We focused on closed population models, but the movement models developed here can be extended to open SCR models. The movement process models could also be easily extended to accommodate additional "building blocks" of random walks, such as central tendency (e.g., territoriality) or multiple movement behavior states, thereby providing a flexible and coherent framework for linking animal movement behavior to population dynamics, density, and distribution.

Modeling spatiotemporal abundance and movement dynamics using an integrated spatial capture-recapture movement model.

Animal movement is a fundamental ecological process affecting the survival and reproduction of individuals, the structure of populations, and the dynamics of communities. Methods to quantify animal movement and spatiotemporal abundances, however, are generally separate and therefore omit linkages between individual-level and population-level processes. We describe an integrated spatial capture-recapture (SCR) movement model to jointly estimate (1) the number and distribution of individuals in a defined spatial region and (2) movement of those individuals through time. We applied our model to a study of polar bears (Ursus maritimus) in a 28,125 km2 survey area of the eastern Chukchi Sea, USA in 2015 that incorporated capture-recapture and telemetry data. In simulation studies, the model provided unbiased estimates of movement, abundance, and detection parameters using a bivariate normal random walk and correlated random walk movement process. Our case study provided detailed evidence of directional movement persistence for both male and female bears, where individuals regularly traversed areas larger than the survey area during the 36-day study period. Scaling from individual- to population-level inferences, we found that densities varied from <0.75 bears/625 km2 grid cell/day in nearshore cells to 1.6-2.5 bears/grid cell/day for cells surrounded by sea ice. Daily abundance estimates ranged from 53 to 69 bears, with no trend across days. The cumulative number of unique bears that used the survey area increased through time due to movements into and out of the area, resulting in an estimated 171 individuals using the survey area during the study (95% credible interval 124-250). Abundance estimates were similar to a previous multiyear integrated population model using capture-recapture and telemetry data (2008-2016; Regehr et al., Scientific Reports 8:16780, 2018). Overall, the SCR-movement model successfully quantified both individual- and population-level space use, including the effects of landscape characteristics on movement, abundance, and detection, while linking the movement and abundance processes to directly estimate density within a prescribed spatial region and temporal period. Integrated SCR-movement models provide a generalizable approach to incorporate greater movement realism into population dynamics and link movement to emergent properties including spatiotemporal densities and abundances.

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.

Ontogenetic shifts from social to experiential learning drive avian migration timing.

Migrating animals may benefit from social or experiential learning, yet whether and how these learning processes interact or change over time to produce observed migration patterns remains unexplored. Using 16 years of satellite-tracking data from 105 reintroduced whooping cranes, we reveal an interplay between social and experiential learning in migration timing. Both processes dramatically improved individuals' abilities to dynamically adjust their timing to track environmental conditions along the migration path. However, results revealed an ontogenetic shift in the dominant learning process, whereby subadult birds relied on social information, while mature birds primarily relied on experiential information. These results indicate that the adjustment of migration phenology in response to the environment is a learned skill that depends on both social context and individual age. Assessing how animals successfully learn to time migrations as environmental conditions change is critical for understanding intraspecific differences in migration patterns and for anticipating responses to global change.

Age-structured Jolly-Seber model expands inference and improves parameter estimation from capture-recapture data.

Understanding the influence of individual attributes on demographic processes is a key objective of wildlife population studies. Capture-recapture and age data are commonly collected to investigate hypotheses about survival, reproduction, and viability. We present a novel age-structured Jolly-Seber model that incorporates age and capture-recapture data to provide comprehensive information on population dynamics, including abundance, age-dependent survival, recruitment, age structure, and population growth rates. We applied our model to a multi-year capture-recapture study of polar bears (Ursus maritimus) in western Hudson Bay, Canada (2012-2018), where management and conservation require a detailed understanding of how polar bears respond to climate change and other factors. In simulation studies, the age-structured Jolly-Seber model improved precision of survival, recruitment, and annual abundance estimates relative to standard Jolly-Seber models that omit age information. Furthermore, incorporating age information improved precision of population growth rates, increased power to detect trends in abundance, and allowed direct estimation of age-dependent survival and changes in annual age structure. Our case study provided detailed evidence for senescence in polar bear survival. Median survival estimates were lower (<0.95) for individuals aged <5 years, remained high (>0.95) for individuals aged 7-22 years, and subsequently declined to near zero for individuals >30 years. We also detected cascading effects of large recruitment classes on population age structure, which created major shifts in age structure when these classes entered the population and then again when they reached prime breeding ages (10-15 years old). Overall, age-structured Jolly-Seber models provide a flexible means to investigate ecological and evolutionary processes that shape populations (e.g., via senescence, life expectancy, and lifetime reproductive success) while improving our ability to investigate population dynamics and forecast population changes from capture-recapture data.

Bias, precision, and parameter redundancy in complex multistate models with unobservable states.

Multistate mark-recapture models with unobservable states can yield unbiased estimators of survival probabilities in the presence of temporary emigration (i.e., in cases where some individuals are temporarily unavailable for capture). In addition, these models permit the estimation of transition probabilities between states, which may themselves be of interest; for example, when only breeding animals are available for capture. However, parameter redundancy is frequently a problem in these models, yielding biased parameter estimates and influencing model selection. Using numerical methods, we examine complex multistate mark-recapture models involving two observable and two unobservable states. This model structure was motivated by two different biological systems: one involving island-nesting albatross, and another involving pond-breeding amphibians. We found that, while many models are theoretically identifiable given appropriate constraints, obtaining accurate and precise parameter estimates in practice can be difficult. Practitioners should consider ways to increase detection probabilities or adopt robust design sampling in order to improve the properties of estimates obtained from these models. We suggest that investigators interested in using these models explore both theoretical identifiability and possible near-singularity for likely parameter values using a combination of available methods.

Sampling design considerations for demographic studies: a case of colonial seabirds.

For the purposes of making many informed conservation decisions, the main goal for data collection is to assess population status and allow prediction of the consequences of candidate management actions. Reducing the bias and variance of estimates of population parameters reduces uncertainty in population status and projections, thereby reducing the overall uncertainty under which a population manager must make a decision. In capture-recapture studies, imperfect detection of individuals, unobservable life-history states, local movement outside study areas, and tag loss can cause bias or precision problems with estimates of population parameters. Furthermore, excessive disturbance to individuals during capture-recapture sampling may be of concern because disturbance may have demographic consequences. We address these problems using as an example a monitoring program for Black-footed Albatross (Phoebastria nigripes) and Laysan Albatross (Phoebastria immutabilis) nesting populations in the northwestern Hawaiian Islands. To mitigate these estimation problems, we describe a synergistic combination of sampling design and modeling approaches. Solutions include multiple capture periods per season and multistate, robust design statistical models, dead recoveries and incidental observations, telemetry and data loggers, buffer areas around study plots to neutralize the effect of local movements outside study plots, and double banding and statistical models that account for band loss. We also present a variation on the robust capture-recapture design and a corresponding statistical model that minimizes disturbance to individuals. For the albatross case study, this less invasive robust design was more time efficient and, when used in combination with a traditional robust design, reduced the standard error of detection probability by 14% with only two hours of additional effort in the field. These field techniques and associated modeling approaches are applicable to studies of most taxa being marked and in some cases have individually been applied to studies of birds, fish, herpetofauna, and mammals.

Linking demographic and food-web models to understand management trade-offs.

Alternatives in ecosystem-based management often differ with respect to trade-offs between ecosystem values. Ecosystem or food-web models and demographic models are typically employed to evaluate alternatives, but the approaches are rarely integrated to uncover conflicts between values. We applied multistate models to a capture-recapture dataset on common guillemots Uria aalge breeding in the Baltic Sea to identify factors influencing survival. The estimated relationships were employed together with Ecopath-with-Ecosim food-web model simulations to project guillemot survival under six future scenarios incorporating climate change. The scenarios were based on management alternatives for eutrophication and cod fisheries, issues considered top priority for regional management, but without known direct effects on the guillemot population. Our demographic models identified prey quantity (abundance and biomass of sprat Sprattus sprattus) as the main factor influencing guillemot survival. Most scenarios resulted in projections of increased survival, in the near (2016-2040) and distant (2060-2085) future. However, in the scenario of reduced nutrient input and precautionary cod fishing, guillemot survival was projected to be lower in both future periods due to lower sprat stocks. Matrix population models suggested a substantial decline of the guillemot population in the near future, 24% per 10 years, and a smaller reduction, 1.1% per 10 years, in the distant future. To date, many stakeholders and Baltic Sea governments have supported reduced nutrient input and precautionary cod fishing and implementation is underway. Negative effects on nonfocal species have previously not been uncovered, but our results show that the scenario is likely to negatively impact the guillemot population. Linking model results allowed identifying trade-offs associated with management alternatives. This information is critical to thorough evaluation by decision-makers, but not easily obtained by food-web models or demographic models in isolation. Appropriate datasets are often available, making it feasible to apply a linked approach for better-informed decisions in ecosystem-based management.

Integrated Population Modeling Provides the First Empirical Estimates of Vital Rates and Abundance for Polar Bears in the Chukchi Sea.

Large carnivores are imperiled globally, and characteristics making them vulnerable to extinction (e.g., low densities and expansive ranges) also make it difficult to estimate demographic parameters needed for management. Here we develop an integrated population model to analyze capture-recapture, radiotelemetry, and count data for the Chukchi Sea subpopulation of polar bears (Ursus maritimus), 2008-2016. Our model addressed several challenges in capture-recapture studies for polar bears by including a multievent structure reflecting location and life history states, while accommodating state uncertainty. Female breeding probability was 0.83 (95% credible interval [CRI] = 0.71-0.90), with litter sizes of 2.18 (95% CRI = 1.71-2.82) for age-zero and 1.61 (95% CRI = 1.46-1.80) for age-one cubs. Total adult survival was 0.90 (95% CRI = 0.86-0.92) for females and 0.89 (95% CRI = 0.83-0.93) for males. Spring on-ice densities west of Alaska were 0.0030 bears/km2 (95% CRI = 0.0016-0.0060), similar to 1980s-era density estimates although methodological differences complicate comparison. Abundance of the Chukchi Sea subpopulation, derived by extrapolating density from the study area using a spatially-explicit habitat metric, was 2,937 bears (95% CRI = 1,552-5,944). Our findings are consistent with other lines of evidence suggesting the Chukchi Sea subpopulation has been productive in recent years, although it is uncertain how long this will continue given sea-ice loss due to climate change.