Predicting the spatial distribution of wintering golden eagles to inform full annual cycle conservation in western North America.

Wildlife conservation strategies focused on one season or population segment may fail to adequately protect populations, especially when a species' habitat preferences vary among seasons, age-classes, geographic regions, or other factors. Conservation of golden eagles (Aquila chrysaetos) is an example of such a complex scenario, in which the distribution, habitat use, and migratory strategies of this species of conservation concern vary by age-class, reproductive status, region, and season. Nonetheless, research aimed at mapping priority use areas to inform management of golden eagles in western North America has typically focused on territory-holding adults during the breeding period, largely to the exclusion of other seasons and life-history groups. To support population-wide conservation planning across the full annual cycle for golden eagles, we developed a distribution model for individuals in a season not typically evaluated-winter-and in an area of the interior western U.S. that is a high priority for conservation of the species. We used a large GPS-telemetry dataset and library of environmental variables to develop a machine-learning model to predict spatial variation in the relative intensity of use by golden eagles during winter in Wyoming, USA, and surrounding ecoregions. Based on a rigorous series of evaluations including cross-validation, withheld and independent data, our winter-season model accurately predicted spatial variation in intensity of use by multiple age- and life-history groups of eagles not associated with nesting territories (i.e., all age classes of long-distance migrants, and resident non-adults and adult "floaters", and movements of adult territory holders and their offspring outside their breeding territories). Important predictors in the model were wind and uplift (40.2% contribution), vegetation and landcover (27.9%), topography (14%), climate and weather (9.4%), and ecoregion (8.7%). Predicted areas of high-use winter habitat had relatively low spatial overlap with nesting habitat, suggesting a conservation strategy targeting high-use areas for one season would capture as much as half and as little as one quarter of high-use areas for the other season. The majority of predicted high-use habitat (top 10% quantile) occurred on private lands (55%); lands managed by states and the Bureau of Land Management (BLM) had a lower amount (33%), but higher concentration of high-use habitat than expected for their area (1.5-1.6x). These results will enable those involved in conservation and management of golden eagles in our study region to incorporate spatial prioritization of wintering habitat into their existing regulatory processes, land-use planning tasks, and conservation actions.

Demographic implications of lead poisoning for eagles across North America.

Lead poisoning occurs worldwide in populations of predatory birds, but exposure rates and population impacts are known only from regional studies. We evaluated the lead exposure of 1210 bald and golden eagles from 38 US states across North America, including 620 live eagles. We detected unexpectedly high frequencies of lead poisoning of eagles, both chronic (46 to 47% of bald and golden eagles, as measured in bone) and acute (27 to 33% of bald eagles and 7 to 35% of golden eagles, as measured in liver, blood, and feathers). Frequency of lead poisoning was influenced by age and, for bald eagles, by region and season. Continent-wide demographic modeling suggests that poisoning at this level suppresses population growth rates for bald eagles by 3.8% (95% confidence interval: 2.5%, 5.4%) and for golden eagles by 0.8% (0.7%, 0.9%). Lead poisoning is an underappreciated but important constraint on continent-wide populations of these iconic protected species.

Linking behavioral states to landscape features for improved conservation management.

A central theme for conservation is understanding how animals differentially use, and are affected by change in, the landscapes they inhabit. However, it has been challenging to develop conservation schemes for habitat-specific behaviors.Here we use behavioral change point analysis to identify behavioral states of golden eagles (Aquila chrysaetos) in the Sonoran and Mojave Deserts of the southwestern United States, and we identify, for each behavioral state, conservation-relevant habitat associations.We modeled behavior using 186,859 GPS points from 48 eagles and identified 2,851 distinct segments comprising four behavioral states. Altitude above ground level (AGL) best differentiated behavioral states, with two clusters of short-distance movement behaviors characterized by low AGL (state 1 AGL = 14 m (median); state 2 AGL = 11 m) and two associated with longer-distance movement behaviors and characterized by higher AGL (state 3 AGL = 108 m; state 4 AGL = 450 m).Behaviors such as perching and low-altitude hunting were associated with short-distance movements in updraft-poor environments, at higher elevations, and over steeper and more north-facing terrain. In contrast, medium-distance movements such as hunting and transiting were over gentle and south-facing slopes. Long-distance transiting occurred over the desert habitats that generate the best updraft.This information can guide management of this species, and our approach provides a template for behavior-specific habitat associations for other species of management concern.

Modeling spatial variation in density of golden eagle nest sites in the western United States.

In order to contribute to conservation planning efforts for golden eagles (Aquila chrysaetos) in the western U.S., we developed nest site models using >6,500 nest site locations throughout a >3,483,000 km2 area of the western U.S. We developed models for twelve discrete modeling regions, and estimated relative density of nest sites for each region. Cross-validation showed that, in general, models accurately estimated relative nest site densities within regions and sub-regions. Areas estimated to have the highest densities of breeding golden eagles had from 132-2,660 times greater densities compared to the lowest density areas. Observed nest site densities were very similar to those reported from published studies. Large extents of each modeling region consisted of low predicted nest site density, while a small percentage of each modeling region contained disproportionately high nest site density. For example, we estimated that areas with relative nest density values <0.3 represented from 62.8-97.8% ([Formula: see text] = 82.5%) of each modeling area, and those areas contained from 14.7-30.0% ([Formula: see text] = 22.1%) of the nest sites. In contrast, areas with relative nest density values >0.5 represented from 1.0-12.8% ([Formula: see text] = 6.3%) of modeling areas, and those areas contained from 47.7-66.9% ([Formula: see text] = 57.3%) of the nest sites. Our findings have direct application to: 1) large-scale conservation planning efforts, 2) risk analyses for land-use proposals such as recreational trails or wind power development, and 3) identifying mitigation areas to offset the impacts of human disturbance.

Early Life Conditions and Immune Defense in Nestling Swainson's Hawks.

The quality of perinatal conditions directly influences the physical and immunological development of nestlings, yet it is inherently variable across space and time. Long-term breeding data for a population of Swainson's hawks (Buteo swainsoni) in northern California show a continuum of territory occupancy and productivity values of individual territories and nests. Here we explore effects of variation among territories on immune system development. We hypothesize that nestlings benefitting from favorable conditions will invest in stronger immune systems, a trait with long-term benefits. We used two immunological assays, a bactericidal assay and a hemolytic-complement activity assay, with leukocyte differentials (heterophil∶lymphocyte ratio) to evaluate the constitutive innate immune system. We examined whether early brood-rearing conditions (i.e., number of siblings, hatch date, endoparasite prevalence) were associated with immunological development. Linear mixed-effects models indicated a positive relationship between extended territory occupancy history-an index of habitat quality-and nestling immune function during years with poorer reproduction. There was no association during an exceptionally good reproductive year. Hence, at least under some circumstances, nestling environments or territory characteristics may affect immune function of nestlings. Our study contributes to the growing body of evidence highlighting the importance of facultative allocation to immune traits using long-term demographic data of a top avian predator.

Conservation planning for species recovery under the Endangered Species Act: A case study with the Northern Spotted Owl.

The northern spotted owl (Strix occidentalis caurina) was listed as threatened under the U.S. Endangered Species Act (ESA) in 1990. We applied modern spatial conservation theory and models to evaluate several candidate critical habitat networks, and sought an efficient conservation solution that encompassed the highest value lands for spotted owl recovery rather than maximizing the total area of potential critical habitat. We created a map of relative habitat suitability, which served as input to the spatial conservation prioritization program Zonation. We used the spatially-explicit individual-based population model HexSim to estimate and compare simulated spotted owl population outcomes among a suite of candidate critical habitat networks that varied in size and spatial arrangement under alternative scenarios of future habitat suitability and barred owl (S. varia) effects. We evaluated simulated spotted owl population outcomes, including total population size, and extinction and quasi-extinction likelihoods for 108 combinations of candidate critical habitat networks by habitat change by barred owl scenarios, both range-wide and within 11 distinct portions of the owl's range. Barred owl encounter rates and the amount and suitability of habitat had substantial effects on simulated spotted owl populations. When barred owl encounter rates were high, changes in the amount and suitability of habitat had minimal impacts on population performance. Under lowered barred owl encounter rates, candidate critical habitat networks that included most existing high suitability habitat supported a high likelihood of long-term population persistence. Barred owls are currently the primary driving force behind poor population performance of NSOs; however, our models demonstrated that a sufficient area of high suitability habitat remains essential for recovery when effects of barred owls can be reduced. The modeling approach we employed is sufficiently flexible to incorporate new information about spotted owls as it becomes available and could likely be applied to conservation planning for other species.

MIGRATION PATTERNS, USE OF STOPOVER AREAS, AND AUSTRAL SUMMER MOVEMENTS OF SWAINSON'S HAWKS.

From 1995-1998, we tracked movements of adult Swainson's Hawks (Buteo swainsoni) using satellite telemetry to characterize migration, important stopover areas, and austral summer movements. We tagged 46 hawks from July - September on their nesting grounds in seven U.S. states and two Canadian provinces. Swainson's Hawks basically followed three routes south on a broad front, converged along the east coast of central Mexico, and followed a concentrated corridor to a communal austral summer area in central Argentina. North of 20° N, southward and northward tracks differed little for individuals from east of the Continental Divide but differed greatly (up to 1700 km) for individuals from west of the Continental Divide. Hawks left the breeding grounds mid-August to mid-October; departure dates did not differ by location, year, or sex. South migration lasted 42 to 98 days, and north migration took 51 to 82 days. On south migration, 36% of the Swainson's Hawks departed the nesting grounds nearly 3 weeks earlier than the other radio marked hawks and made stopovers 9.0 - 26.0 days long in seven separate areas, mainly in the southern Great Plains, southern Arizona and New Mexico, and north-central Mexico. The austral period lasted 76 to 128 days. All Swainson's Hawks used a core area in central Argentina within 23% of the 738800 km2 austral summer range where they frequently moved long distances (up to 1600 km). Conservation of Swainson's Hawks must be an international effort that considers habitats used during nesting and non-nesting seasons including migration stopovers.