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.

Linking robust spatiotemporal datasets to assess and monitor habitat attributes of a threatened species.

Accessibility of multispectral, multitemporal imagery combined with recent advances in cloud computing and machine learning approaches have enhanced our ability to model habitat characteristics across broad spatial and temporal scales. We integrated a large dataset of known nest and roost sites of a threatened species, the Mexican spotted owl (Strix occidentalis lucida), in the southwestern USA with Landsat imagery processed using the Continuous Change Detection and Classification (CCDC) time series algorithm on Google Earth Engine. We then used maximum entropy modeling (Maxent) to classify the landscape into four 'spectral similarity' classes that reflected the degree to which 30-m pixels contained a multispectral signature similar to that found at known owl nest/roost sites and mapped spectral similarity classes from 1986-2020. For map interpretation, we used nationally consistent forest inventory data to evaluate the structural and compositional characteristics of each spectral similarity class. We found a monotonic increase of structural characteristics typically associated with owl nesting and roosting over classes of increasing similarity, with the 'very similar' class meeting or exceeding published minimum desired management conditions for owl nesting and roosting. We also found an increased rate of loss of forest vegetation typical of owl nesting and roosting since the beginning of the 21st century that can be partly attributed to increased frequency and extent of large (≥400 ha) wildfires. This loss resulted in a 38% reduction over the 35-year study period in forest vegetation most similar to that used for owl nesting and roosting. Our modelling approach using cloud computing with time series of Landsat imagery provided a cost-effective tool for landscape-scale, multidecadal monitoring of vegetative components of a threatened species' habitat. Our approach could be used to monitor trends in the vegetation favored by any other species, provided that high-quality location data such as we presented here are available.

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.

The past and future roles of competition and habitat in the range-wide occupancy dynamics of Northern Spotted Owls.

Slow ecological processes challenge conservation. Short-term variability can obscure the importance of slower processes that may ultimately determine the state of a system. Furthermore, management actions with slow responses can be hard to justify. One response to slow processes is to explicitly concentrate analysis on state dynamics. Here, we focus on identifying drivers of Northern Spotted Owl (Strix occidentalis caurina) territorial occupancy dynamics across 11 study areas spanning their geographic range and forecasting response to potential management actions. Competition with Barred Owls (Strix varia) has increased Spotted Owl territory extinction probabilities across all study areas and driven recent declines in Spotted Owl populations. Without management intervention, the Northern Spotted Owl subspecies will be extirpated from parts of its current range within decades. In the short term, Barred Owl removal can be effective. Over longer time spans, however, maintaining or improving habitat conditions can help promote the persistence of northern spotted owl populations. In most study areas, habitat effects on expected Northern Spotted Owl territorial occupancy are actually greater than the effects of competition from Barred Owls. This study suggests how intensive management actions (removal of a competitor) with rapid results can complement a slower management action (i.e., promoting forest succession).

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.

Isolation by distance versus landscape resistance: Understanding dominant patterns of genetic structure in Northern Spotted Owls (Strix occidentalis caurina).

Landscape genetics investigations examine how the availability and configuration of habitat influence genetic structure of plants and animals. We used landscape genetics to evaluate the role that forest connectivity plays in determining genetic structure of the federally-threatened Northern Spotted Owl (Strix occidentalis caurina) using genotypes of 339 Northern Spotted Owls obtained for 10 microsatellite loci. Spatial clustering analyses identified a distinct genetic cluster at the southern extent of the region examined. This cluster could not be linked to landscape connectivity patterns and suggested that post-Pleistocene processes were involved with its development rather than contemporary landscape configuration. We also compared matrices of pairwise inter-individual genetic distances with resistance distances derived from a circuit-theory based framework. Resistance distances were obtained for an idealized raster map that reflected continuous unimpeded dispersal habitat across the landscape along with five empirically-derived raster maps reflecting the 1870's, 1940's, 1986, 1994, and 2012. Resistance distances from the idealized map served as surrogates for linear geographic distances. Relative to idealized conditions, resistance distances were ~250% higher in the 1940's and ~200% higher from 1986 onward. Resistance distances from the 1870's were ~40% higher than idealized conditions. Inter-individual genetic distances were most highly correlated with resistance distances from the idealized map rather than any of the empirical maps. Two hypotheses explain our results. First, our results may reflect temporal lags between the onset of large-scale habitat alterations and their novel effects on genetic structure in long-lived species such as Northern Spotted Owls. Second, because Northern Spotted Owls disperse over long distances, our results may indicate that forest habitat has never been sufficiently fragmented to the point where connectivity was disrupted. The second hypothesis could indicate that forest management practices mandated by the Northwest Forest Plan succeeded with one of its primary goals. However, our results do not represent a complete portrayal of the status of Northern Spotted Owls given detection of significant population declines and bottlenecks in other studies. Future investigations based on computer simulations may help distinguish between hypotheses.

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.