TDFCAM: A method for estimating stable isotope trophic discrimination in wild populations.

Stable isotope mixing models (SIMMs) are widely used for characterizing wild animal diets. Such models rely upon using accurate trophic discrimination factors (TDFs) to account for the digestion, incorporation, and assimilation of food. Existing methods to calculate TDFs rely on controlled feeding trials that are time-consuming, often impractical for the study taxon, and may not reflect natural variability of TDFs present in wild populations.We present TDFCAM as an alternative approach to estimating TDFs in wild populations, by using high-precision diet estimates from a secondary methodological source-in this case nest cameras-in lieu of controlled feeding trials, and provide a framework for how and when it should be applied.In this study, we evaluate the TDFCAM approach in three datasets gathered on wild raptor nestlings (gyrfalcons Falco rusticolus; peregrine falcons Falco perigrinus; common buzzards Buteo buteo) comprising contemporaneous δ13C & δ15N stable isotope data and high-quality nest camera dietary data. We formulate Bayesian SIMMs (BSIMMs) incorporating TDFs from TDFCAM and analyze their agreement with nest camera data, comparing model performance with those based on other relevant TDFs. Additionally, we perform sensitivity analyses to characterize TDFCAM variability, and identify ecological and physiological factors contributing to that variability in wild populations.Across species and tissue types, BSIMMs incorporating a TDFCAM outperformed any other TDF tested, producing reliable population-level estimates of diet composition. We demonstrate that applying this approach even with a relatively low sample size (n < 10 individuals) produced more accurate estimates of trophic discrimination than a controlled feeding study conducted on the same species. Between-individual variability in TDFCAM estimates for ∆13C & ∆15 N increased with analytical imprecision in the source dietary data (nest cameras) but was also explained by natural variables in the study population (e.g., nestling nutritional/growth status and dietary composition).TDFCAM is an effective method of estimating trophic discrimination in wild animal populations. Here, we use nest cameras as source dietary data, but this approach is applicable to any high-accuracy method of measuring diet, so long as diet can be monitored over an interval contemporaneous with a tissue's isotopic turnover rate.

Isotopic niche partitioning and individual specialization in an Arctic raptor guild.

Intra- and inter-specific resource partitioning within predator communities is a fundamental component of trophic ecology, and one proposed mechanism for how populations partition resources is through individual niche variation. The Niche Variation Hypothesis (NVH) predicts that inter-individual trait variation leads to functional trade-offs in foraging efficiency, resulting in populations composed of individual dietary specialists. The degree to which niche specialization persists within a population is plastic and responsive to fluctuating resource availability. We quantified niche overlap and tested the NVH within an Arctic raptor guild, focusing on three species that employ different foraging strategies: golden eagles (generalists); gyrfalcons (facultative specialists); and rough-legged hawks (specialists). Tundra ecosystems exhibit cyclic populations of arvicoline rodents (lemmings and voles), providing a unique system in which to examine predator diet in response to interannual fluctuations in resource availability. Using blood δ13C and δ15N values from 189 raptor nestlings on Alaska's Seward Peninsula (2014-2019), we calculated isotopic niche width and used Bayesian stable isotope mixing models (BSIMMs) to characterize individual specialization and test the NVH. Nest-level specialization estimated from stable isotopes was strongly correlated with indices of specialization based on camera trap data. We observed a high degree of isotopic niche overlap between the three species and gyrfalcons displayed a positive relationship between individual specialization and population niche width on an interannual basis consistent with the NVH. Our findings suggest plasticity in niche specialization may reduce intra- and inter-specific resource competition under dynamic ecological conditions.

Demographic partitioning of dynamic energy subsidies revealed with an Ornstein-Uhlenbeck space use model.

In populations across many taxa, a large fraction of sexually mature individuals do not breed but are attempting to enter the breeding population. Such individuals, often referred to as "floaters," can play critical roles in the dynamics and stability of these populations and buffer them through periods of high adult mortality. Floaters are difficult to study, however, so we lack data needed to understand their roles in the population ecology and conservation status of many species. Here, we analyzed satellite telemetry data with a newly developed mechanistic space use model based on an Ornstein-Uhlenbeck process to help overcome the paucity of data in studying the differential habitat selection and space use of floater and territorial golden eagles Aquila chrysaetos. Our sample consisted of 49 individuals tracked over complete breeding seasons across 4 years, totaling 104 eagle breeding seasons. Modeling these data mechanistically was required to disentangle key differences in movement and particularly to separate aspects of movement driven by resource selection from those driven by use of a central place. We found that floaters generally had more expansive space use patterns and larger home ranges, as well as evidence that they partition space with territorial individuals seemingly on fine scales through differential habitat and resource selection. Floater and territorial eagle home ranges overlapped markedly, suggesting that floaters use the interstices between territories. Furthermore, floater and territorial eagles differed in how they selected for uplift variables, key components of soaring birds' energy landscape, with territorial eagles apparently better able to find and use thermal uplift. We also found relatively low individual heterogeneity in resource selection, especially among territorial individuals, suggesting a narrow realized niche for breeding individuals, which varied from the level of among-individual variation present during migration. This work furthers our understanding of floaters' potential roles in the population ecology of territorial species and suggests that conserving landscapes occupied by territorial eagles also protects floaters.

Ecological insights from three decades of animal movement tracking across a changing Arctic.

The Arctic is entering a new ecological state, with alarming consequences for humanity. Animal-borne sensors offer a window into these changes. Although substantial animal tracking data from the Arctic and subarctic exist, most are difficult to discover and access. Here, we present the new Arctic Animal Movement Archive (AAMA), a growing collection of more than 200 standardized terrestrial and marine animal tracking studies from 1991 to the present. The AAMA supports public data discovery, preserves fundamental baseline data for the future, and facilitates efficient, collaborative data analysis. With AAMA-based case studies, we document climatic influences on the migration phenology of eagles, geographic differences in the adaptive response of caribou reproductive phenology to climate change, and species-specific changes in terrestrial mammal movement rates in response to increasing temperature.

Isolation and characterization of microsatellite loci in merlins (Falco columbarius) and cross-species amplification in gyrfalcons (F. rusticolus) and peregrine falcons (F. peregrinus).

Merlins, Falco columbarius, breed throughout temperate and high latitude habitats in Asia, Europe, and North America. Like peregrine falcons, F. peregrinus, merlins underwent population declines during the mid-to-late twentieth century, due to organochlorine-based contamination, and have subsequently recovered, at least in North American populations. To better understand levels of genetic diversity and population structuring in contemporary populations and to assess the impact of the twentieth century decline, we used genomic data archived in public databases and constructed genomic libraries to isolate and characterize a suite of 17 microsatellite markers for use in merlins. We also conducted cross-amplification experiments to determine the markers' utility in peregrine falcons and gyrfalcons, F. rusticolus. These markers provide a valuable addition to marker suites that can be used to determine individual identity and conduct genetic analyses on merlins and congeners.

Bayesian stable isotope mixing models effectively characterize the diet of an Arctic raptor.

Bayesian stable isotope mixing models (BSIMMs) for δ13 C and δ15 N can be a useful tool to reconstruct diets, characterize trophic relationships, and assess spatiotemporal variation in food webs. However, use of this approach typically requires a priori knowledge on the level of enrichment occurring between the diet and tissue of the consumer being sampled (i.e. a trophic discrimination factor or TDF). Trophic discrimination factors derived from captive feeding studies are highly variable, and it is challenging to select the appropriate TDF for diet estimation in wild populations. We introduce a novel method for estimating TDFs in a wild population-a proportionally balanced equation that uses high-precision diet estimates from nest cameras installed on a subset of nests in lieu of a controlled feeding study (TDFCAM ). We tested the ability of BSIMMs to characterize diet in a free-living population of gyrfalcon Falco rusticolus nestlings by comparing model output to high-precision nest camera diet estimates. We analysed the performance of models formulated with a TDFCAM against other relevant TDFs and assessed model sensitivity to an informative prior. We applied the most parsimonious model inputs to a larger sample to analyse broad-scale temporal dietary trends. Bayesian stable isotope mixing models fitted with a TDFCAM and uninformative prior had the best agreement with nest camera data, outperforming TDFs derived from captive feeding studies. BSIMMs produced with a TDFCAM produced reliable diet estimates at the nest level and accurately identified significant temporal shifts in gyrfalcon diet within and between years. Our method of TDF estimation produced more accurate estimates of TDFs in a wild population than traditional approaches, consequently improving BSIMM diet estimates. We demonstrate how BSIMMs can complement a high-precision diet study by expanding its spatiotemporal scope of inference and recommend this integrative methodology as a powerful tool for future trophic studies.

Novel step selection analyses on energy landscapes reveal how linear features alter migrations of soaring birds.

Human modification of landscapes includes extensive addition of linear features, such as roads and transmission lines. These can alter animal movement and space use and affect the intensity of interactions among species, including predation and competition. Effects of linear features on animal movement have seen relatively little research in avian systems, despite ample evidence of their effects in mammalian systems and that some types of linear features, including both roads and transmission lines, are substantial sources of mortality. Here, we used satellite telemetry combined with step selection functions designed to explicitly incorporate the energy landscape (el-SSFs) to investigate the effects of linear features and habitat on movements and space use of a large soaring bird, the golden eagle Aquila chrysaetos, during migration. Our sample consisted of 32 adult eagles tracked for 45 spring and 39 fall migrations from 2014 to 2017. Fitted el-SSFs indicated eagles had a strong general preference for south-facing slopes, where thermal uplift develops predictably, and that these areas are likely important aspects of migratory pathways. el-SSFs also provided evidence that roads and railroads affected movement during both spring and fall migrations, but eagles selected areas near roads to a greater degree in spring compared to fall and at higher latitudes compared to lower latitudes. During spring, time spent near linear features often occurred during slower-paced or stopover movements, perhaps in part to access carrion produced by vehicle collisions. Regardless of the behavioural mechanism of selection, use of these features could expose eagles and other soaring species to elevated risk via collision with vehicles and/or transmission lines. Linear features have previously been documented to affect the ecology of terrestrial species (e.g. large mammals) by modifying individuals' movement patterns; our work shows that these effects on movement extend to avian taxa.

Local meteorological conditions reroute a migration.

For migrating animals, realized migration routes and timing emerge from hundreds or thousands of movement decisions made along migration routes. Local weather conditions along migration routes continually influence these decisions, and even relatively small changes in en route weather may cumulatively result in major shifts in migration patterns. Here, we analysed satellite tracking data to score a discrete navigation decision by a large migratory bird as it navigated a high-latitude, 5000 m elevation mountain range to understand how those navigational decisions changed under different weather conditions. We showed that wind conditions in particular areas along the migration pathway drove a navigational decision to reroute a migration; conditions encountered predictably resulted in migrants routing either north or south of the mountain range. With abiotic conditions continuing to change globally, simple decisions, such as the one described here, might additively emerge into new, very different migration routes.

Migration corridors of adult Golden Eagles originating in northwestern North America.

There has been increasing concern for Golden Eagle (Aquila chrysaetos) populations in North America due to current and future projections of mortality risk and habitat loss from anthropogenic sources. Identification of high-use movement corridors and bottlenecks for the migratory portion of the eagle population in western North America is an important first step to help habitat conservation and management efforts to reduce the risk of eagle mortality. We used dynamic Brownian Bridge movement models to estimate utilization distributions of adult eagles migrating across the western North America and identified high-use areas by calculating the overlap of individuals on population and regional levels. On a population level, the Rocky Mountain Front from east-central British Columbia to central Montana and southwestern Yukon encompassed the most used migration corridors with our study extent for both spring and fall. Regional analysis on a 100 x 200 km scale revealed additional moderate and high-level use corridors in the central British Columbia plateaus. Eagles were more dispersed in the spring until their routes converged in southern Alberta. High-use fall corridors extended farther south into central Wyoming. Knowledge of these high-use areas can aid in conservation and site planning to help maintain and enhance migratory Golden Eagle populations in western North America.