Invader removal triggers competitive release in a threatened avian predator.

Changes in the distribution and abundance of invasive species can have far-reaching ecological consequences. Programs to control invaders are common but gauging the effectiveness of such programs using carefully controlled, large-scale field experiments is rare, especially at higher trophic levels. Experimental manipulations coupled with long-term demographic monitoring can reveal the mechanistic underpinnings of interspecific competition among apex predators and suggest mitigation options for invasive species. We used a large-scale before-after control-impact removal experiment to investigate the effects of an invasive competitor, the barred owl (Strix varia), on the population dynamics of an iconic old-forest native species, the northern spotted owl (Strix occidentalis caurina). Removal of barred owls had a strong, positive effect on survival of sympatric spotted owls and a weaker but positive effect on spotted owl dispersal and recruitment. After removals, the estimated mean annual rate of population change for spotted owls stabilized in areas with removals (0.2% decline per year), but continued to decline sharply in areas without removals (12.1% decline per year). The results demonstrated that the most substantial changes in population dynamics of northern spotted owls over the past two decades were associated with the invasion, population expansion, and subsequent removal of barred owls. Our study provides experimental evidence of the demographic consequences of competitive release, where a threatened avian predator was freed from restrictions imposed on its population dynamics with the removal of a competitively dominant invasive species.

Range-wide sources of variation in reproductive rates of northern spotted owls.

We conducted a range-wide investigation of the dynamics of site-level reproductive rate of northern spotted owls using survey data from 11 study areas across the subspecies geographic range collected during 1993-2018. Our analytical approach accounted for imperfect detection of owl pairs and misclassification of successful reproduction (i.e., at least one young fledged) and contributed further insights into northern spotted owl population ecology and dynamics. Both nondetection and state misclassification were important, especially because factors affecting these sources of error also affected focal ecological parameters. Annual probabilities of site occupancy were greatest at sites with successful reproduction in the previous year and lowest for sites not occupied by a pair in the previous year. Site-specific occupancy transition probabilities declined over time and were negatively affected by barred owl presence. Overall, the site-specific probability of successful reproduction showed substantial year-to-year fluctuations and was similar for occupied sites that did or did not experience successful reproduction the previous year. Site-specific probabilities for successful reproduction were very small for sites that were unoccupied the previous year. Barred owl presence negatively affected the probability of successful reproduction by northern spotted owls in Washington and California, as predicted, but the effect in Oregon was mixed. The proportions of sites occupied by northern spotted owl pairs showed steep, near-monotonic declines over the study period, with all study areas showing the lowest observed levels of occupancy to date. If trends continue it is likely that northern spotted owls will become extirpated throughout large portions of their range in the coming decades.

Conspecific and congeneric interactions shape increasing rates of breeding dispersal of northern spotted owls.

Breeding dispersal, the movement from one breeding territory to another, is rare for philopatric species that evolved within relatively stable environments, such as the old-growth coniferous forests of the Pacific Northwest. Although dispersal is not inherently maladaptive, the consequences of increased dispersal on population dynamics in populations whose historical dispersal rates are low could be significant, particularly for a declining species. We examined rates and possible causes of breeding dispersal based on a sample of 4,118 northern spotted owls (Strix occidentalis caurina) monitored in seven study areas over 28 yr, 1990-2017, in Oregon and Washington, USA. Using a multistate mark-resight analysis, we investigated the potential impacts of an emergent congeneric competitor (barred owl Strix varia) and forest alteration (extrinsic factors), and social and individual conditions (intrinsic factors) on 408 successive and 1,372 nonsuccessive dispersal events between years. The annual probability of breeding dispersal increased for individual owls that had also dispersed in the previous year and decreased for owls on territories with historically high levels of reproduction. Intrinsic factors including pair status, prior reproductive success, and experience at a site, were also associated with breeding dispersal movements. The percent of monitored owls dispersing each year increased from ˜7% early in the study to ˜25% at the end of the study, which coincided with a rapid increase in numbers of invasive and competitively dominant barred owls. We suggest that the results presented here can inform spotted owl conservation efforts as we identify factors contributing to changing rates of demographic parameters including site fidelity and breeding dispersal. Our study further shows that increasing rates of breeding dispersal associated with population declines contribute to population instability and vulnerability of northern spotted owls to extinction, and the prognosis is unlikely to change unless active management interventions are undertaken.

Integrating count and detection-nondetection data to model population dynamics.

There is increasing need for methods that integrate multiple data types into a single analytical framework as the spatial and temporal scale of ecological research expands. Current work on this topic primarily focuses on combining capture-recapture data from marked individuals with other data types into integrated population models. Yet, studies of species distributions and trends often rely on data from unmarked individuals across broad scales where local abundance and environmental variables may vary. We present a modeling framework for integrating detection-nondetection and count data into a single analysis to estimate population dynamics, abundance, and individual detection probabilities during sampling. Our dynamic population model assumes that site-specific abundance can change over time according to survival of individuals and gains through reproduction and immigration. The observation process for each data type is modeled by assuming that every individual present at a site has an equal probability of being detected during sampling processes. We examine our modeling approach through a series of simulations illustrating the relative value of count vs. detection-nondetection data under a variety of parameter values and survey configurations. We also provide an empirical example of the model by combining long-term detection-nondetection data (1995-2014) with newly collected count data (2015-2016) from a growing population of Barred Owl (Strix varia) in the Pacific Northwest to examine the factors influencing population abundance over time. Our model provides a foundation for incorporating unmarked data within a single framework, even in cases where sampling processes yield different detection probabilities. This approach will be useful for survey design and to researchers interested in incorporating historical or citizen science data into analyses focused on understanding how demographic rates drive population abundance.

SPATIAL DEMOGRAPHIC MODELS TO INFORM CONSERVATION PLANNING OF GOLDEN EAGLES IN RENEWABLE ENERGY LANDSCAPES.

Spatial demographic models can help guide monitoring and management activities targeting at-risk species, even in cases where baseline data are lacking. Here, we provide an example of how site-specific changes in land-use and other anthropogenic stressors can be incorporated into a spatial demographic model to investigate effects on population dynamics of Golden Eagles (Aquila chrysaetos). Our study focused on a population of Golden Eagles exposed to risks associated with rapid increases in renewable energy development in southern California, USA. We developed a spatially-explicit, individual-based simulation model that integrated empirical data on demography of Golden Eagles with spatial data on the arrangement of nesting habitats, prey resources, and planned renewable energy development sites. Our model permitted simulated eagles of different stage-classes to disperse, establish home ranges, acquire resources, prospect for breeding sites, and reproduce. The distribution of nesting habitats, prey resources, and threats within each individual's home range influenced movement, reproduction, and survival. We used our model to explore potential effects of alternative disturbance scenarios, and proposed conservation strategies, on the future distribution and abundance of Golden Eagles in the study region. Results from our simulations suggest that probable increases in mortality associated with renewable energy infrastructure (e.g., collisions with wind-turbines and vehicles, electrocution on power poles) could have negative consequences for population trajectories, but that site-specific conservation actions could reduce the magnitude of negative impacts. Our study demonstrates the use of a flexible and expandable modeling framework to incorporate spatially dependent processes when determining relative risks of proposed management options to Golden Eagles and their habitats.

Estimating fat content in barred owls (Strix varia) with predictive models developed from direct measures of proximate body composition.

Body condition indices and related metrics can help assess habitat quality and other ecological processes, and ideally, these metrics are based on measures of lipids directly extracted from the species of interest. In recent decades, barred owls (Strix varia) have become a species of conservation concern as they invaded older forests of the US Pacific Northwest, and caused population declines of the closely related and federally threatened northern spotted owl (Strix occidentalis caurina). A simple and effective measure of barred owl body condition could help to understand how habitat quality varies within their new range, which in turn can inform their management and other aspects of their ecology. Using 77 barred owl carcasses collected during experimental removals in Washington and Oregon, USA, we measured the amount of lipid in each specimen with proximate body composition analysis. We then fit and compared (with adjusted R2 values) alternative linear regression models to estimate the percent lipids in dry mass of the owls based on morphometric body condition indices, a qualitative fat score of subcutaneous breast fat, sex and the time of year females were collected (relative to egg production). Adjusted R2 values for all models ranged from 0.49 to 0.87, with the best model including mass divided by foot-pad length, fat score, sex and the time of year a female was collected. Most models generated comparable estimates of percent lipids at a population level and we provided correction factors to apply these models when used with live barred owls, allowing for site-specific comparisons of body condition among individuals inhabiting a diversity of environmental conditions.

Three-dimensional partitioning of resources by congeneric forest predators with recent sympatry.

Coexistence of ecologically similar species can be maintained by partitioning along one or more niche axes. Three-dimensional structural complexity is central to facilitating resource partitioning between many forest species, but is underrepresented in field-based studies. We examined resource selection by sympatric northern spotted owls (Strix occidentalis caurina), a threatened species under the US Endangered Species Act, and nonnative barred owls (S. varia) in western Oregon, USA to explore the relative importance of canopy heterogeneity, vertical complexity of forest, and abiotic features to resource selection and identify potential differences that may facilitate long-term coexistence. We predicted that within home range selection of understory densities, measured with airborne lidar, would differ between species based on proportional differences in arboreal and terrestrial prey taken by each owl species. We used discrete choice models and telemetry data from 41 spotted owls and 38 barred owls monitored during 2007-2009 and 2012-2015. Our results suggested that while both species used tall canopy areas more often than low canopy areas, spotted owls were more commonly found in areas with lower tree cover, more developed understory, and steeper slopes. This is the first evidence of fine-scale partitioning based on structural forest properties by northern spotted owls and barred owls.

Post-fledging survival of northern goshawks: the importance of prey abundance, weather, and dispersal.

Effective wildlife conservation strategies require an understanding of how fluctuating environmental conditions affect sensitive life stages. As part of a long-term study, we examined post-fledging and post-independence survival of 89 radio-marked juvenile Northern Goshawks (Accipiter gentilis) produced from 48 nests in northern Arizona, USA, during 1998-2001. Information-theoretic methods were used to examine within- and among-year variation in survival relative to environmental (prey abundance, weather), territory (hatching date, brood size), and individual (gender, body mass) sources of variation. The results support age- and cohort-specific differences in survival that were best explained by behaviors occurring at distinct stages of juvenile development, annual changes in the density of primary bird and mammal prey species, and gender-related differences in body mass. Survival between fledging and independence increased linearly with age and varied among annual cohorts of radio-marked juveniles from 0.81 (95% CI = 0.60-0.93) to 1.00 (95% CI = 0.95-1.00) in association with annual differences in prey density; the slope coefficient for the additive effect of prey density on survival was 1.12 (95% CI = 0.06-2.19). Survival declined to 0.71 (95% CI = 0.60-0.93) shortly after juveniles initiated dispersal (weeks 8-12 post-fledging) and moved to more open habitats at lower elevations. Survival was not closely associated with weather or territory-level parameters. A comparison of the predictions of environmental-, territory-, and individual-based models of survival demonstrated that food availability was the primary factor limiting juvenile survival. This finding indicates that forest management prescriptions designed to support abundant prey populations while providing forest structural conditions that allow goshawks to access their prey within breeding areas should benefit juvenile survival.