Estimates of species' vital rates and an understanding of the factors affecting those parameters over time and space can provide crucial information for management and conservation. We used mark–recapture, reproductive output, and territory occupancy data collected during 1985–2013 to evaluate population processes of Northern Spotted Owls (Strix occidentalis caurina) in 11 study areas in Washington, Oregon, and northern California, USA. We estimated apparent survival, fecundity, recruitment, rate of population change, and local extinction and colonization rates, and investigated relationships between these parameters and the amount of suitable habitat, local and regional variation in meteorological conditions, and competition with Barred Owls (Strix varia). Data were analyzed for each area separately and in a meta-analysis of all areas combined, following a strict protocol for data collection, preparation, and analysis. We used mixed effects linear models for analyses of fecundity, Cormack-Jolly-Seber open population models for analyses of apparent annual survival (ϕ), and a reparameterization of the Jolly-Seber capture–recapture model (i.e. reverse Jolly-Seber; RJS) to estimate annual rates of population change (λRJS) and recruitment. We also modeled territory occupancy dynamics of Northern Spotted Owls and Barred Owls in each study area using 2-species occupancy models. Estimated mean annual rates of population change (λ) suggested that Spotted Owl populations declined from 1.2% to 8.4% per year depending on the study area. The weighted mean estimate of λ for all study areas was 0.962 (± 0.019 SE; 95% CI: 0.925–0.999), indicating an estimated range-wide decline of 3.8% per year from 1985 to 2013. Variation in recruitment rates across the range of the Spotted Owl was best explained by an interaction between total winter precipitation and mean minimum winter temperature. Thus, recruitment rates were highest when both total precipitation (29 cm) and minimum winter temperature (−9.5°C) were lowest. Barred Owl presence was associated with increased local extinction rates of Spotted Owl pairs for all 11 study areas. Habitat covariates were related to extinction rates for Spotted Owl pairs in 8 of 11 study areas, and a greater amount of suitable owl habitat was generally associated with decreased extinction rates. We observed negative effects of Barred Owl presence on colonization rates of Spotted Owl pairs in 5 of 11 study areas. The total amount of suitable Spotted Owl habitat was positively associated with colonization rates in 5 areas, and more habitat disturbance was associated with lower colonization rates in 2 areas. We observed strong declines in derived estimates of occupancy in all study areas. Mean fecundity of females was highest for adults (0.309 ± 0.027 SE), intermediate for 2-yr-olds (0.179 ± 0.040 SE), and lowest for 1-yr-olds (0.065 ± 0.022 SE). The presence of Barred Owls and habitat covariates explained little of the temporal variation in fecundity in most study areas. Climate covariates occurred in competitive fecundity models in 8 of 11 study areas, but support for these relationships was generally weak. The fecundity meta-analysis resulted in 6 competitive models, all of which included the additive effects of geographic region and annual time variation. The 2 top-ranked models also weakly supported the additive negative effects of the amount of suitable core area habitat, Barred Owl presence, and the amount of edge habitat on fecundity. We found strong support for a negative effect of Barred Owl presence on apparent survival of Spotted Owls in 10 of 11 study areas, but found few strong effects of habitat on survival at the study area scale. Climate covariates occurred in top or competitive survival models for 10 of 11 study areas, and in most cases the relationships were as predicted; however, there was little consistency among areas regarding the relative importance of specific climate covariates. In contrast, meta-analysis results suggested that Spotted Owl survival was higher across all study areas when the Pacific Decadal Oscillation (PDO) was in a warming phase and the Southern Oscillation Index (SOI) was negative, with a strongly negative SOI indicative of El Niño events. The best model that included the Barred Owl covariate (BO) was ranked 4th and also included the PDO covariate, but the BO effect was strongly negative. Our results indicated that Northern Spotted Owl populations were declining throughout the range of the subspecies and that annual rates of decline were accelerating in many areas. We observed strong evidence that Barred Owls negatively affected Spotted Owl populations, primarily by decreasing apparent survival and increasing local territory extinction rates. However, the amount of suitable owl habitat, local weather, and regional climatic patterns also were related to survival, occupancy (via colonization rate), recruitment, and, to a lesser extent, fecundity, although there was inconsistency in regard to which covariates were important for particular demographic parameters or across study areas. In the study areas where habitat was an important source of variation for Spotted Owl demographics, vital rates were generally positively associated with a greater amount of suitable owl habitat. However, Barred Owl densities may now be high enough across the range of the Northern Spotted Owl that, despite the continued management and conservation of suitable owl habitat on federal lands, the long-term prognosis for the persistence of Northern Spotted Owls may be in question without additional management intervention. Based on our study, the removal of Barred Owls from the Green Diamond Resources (GDR) study area had rapid, positive effects on Northern Spotted Owl survival and the rate of population change, supporting the hypothesis that, along with habitat conservation...

The recent range expansion of Barred Owls (Strix varia) into the Pacific Northwest, where the species now co‐occurs with the endemic Northern Spotted Owl (Strix occidentalis caurina), resulted in a unique opportunity to investigate potential competition between two congeneric, previously allopatric species. The primary criticism of early competition research was the use of current species' distribution patterns to infer past processes; however, the recent expansion of the Barred Owl and the ability to model the processes that result in site occupancy (i.e., colonization and extinction) allowed us to address the competitive process directly rather than inferring past processes through current patterns. The purpose of our study was to determine whether Barred Owls had any negative effects on occupancy dynamics of nesting territories by Northern Spotted Owls and how these effects were influenced by habitat characteristics of Spotted Owl territories. We used single‐species, multi‐season occupancy models and covariates quantifying Barred Owl detections and habitat characteristics to model extinction and colonization rates of Spotted Owl pairs in southern Oregon, USA. We observed a strong, negative association between Barred Owl detections and colonization rates and a strong positive effect of Barred Owl detections on extinction rates of Spotted Owls. We observed increased extinction rates in response to decreased amounts of old forest at the territory core and higher colonization rates when old‐forest habitat was less fragmented. Annual site occupancy for pairs reflected the strong effects of Barred Owls on occupancy dynamics with much lower occupancy rates predicted for territories where Barred Owls were detected. The strong Barred Owl and habitat effects on occupancy dynamics of Spotted Owls provided evidence of interference competition between the species. These effects increase the importance of conserving large amounts of contiguous, old‐forest habitat to maintain Northern Spotted Owls in the landscape.

High survival and breeding philopatry was previously confirmed for the Adélie penguin (Pygoscelis adeliae) during a period of stable environmental conditions. However, movements of breeding adults as a result of an unplanned natural experiment within a four-colony meta-population provided interesting insights into this species’ population dynamics. We used multistate mark-recapture models to investigate apparent survival and dispersal of breeding birds in the southwestern Ross Sea during 12 breeding seasons (1996–2007). The natural experiment was facilitated by the temporary grounding of two immense icebergs that (i) erected a veritable fence separating colonies and altering migration routes and (ii) added additional stress by trapping extensive sea ice in the region during 5 of 12 y. Colony size varied by orders of magnitude, allowing investigation of apparent survival and dispersal rates in relation to both environmental conditions and colony size within this meta-population. Apparent survival was lowest for the smallest colony (4,000 pairs) and similar for the medium (45,000 pairs) and large colonies (155,000 pairs), despite increased foraging effort expended by breeders at the largest colony. Dispersal of breeding birds was low (<1%), except during years of difficult environmental conditions when movements increased, especially away from the smallest colony (3.5%). Decreased apparent survival at the smallest colony could reflect differences in migration chronology and winter habitat use compared with the other colonies, or it may reflect increased permanent emigration to colonies outside this meta-population. Contrary to current thought, breeding penguins are not always philopatric. Rather, stressful conditions can significantly increase dispersal rates.

We investigated life history responses to extreme variation in physical environmental conditions during a long-term demographic study of Adélie penguins at 3 colonies representing 9% of the world population and the full range of breeding colony sizes. Five years into the 14-year study (1997-2010) two very large icebergs (spanning 1.5 latitude degrees in length) grounded in waters adjacent to breeding colonies, dramatically altering environmental conditions during 2001 - 2005. This natural experiment allowed us to evaluate the relative impacts of expected long-term, but also extreme, short-term climate perturbations on important natural history parameters that can regulate populations. The icebergs presented physical barriers, not just to the penguins but to polynya formation, which profoundly increased foraging effort and movement rates, while reducing breeding propensity and productivity, especially at the smallest colony. We evaluated the effect of a variety of environmental parameters during breeding, molt, migration and wintering periods during years with and without icebergs on penguin breeding productivity, chick mass, and nesting chronology. The icebergs had far more influence on the natural history parameters of penguins than any of the other environmental variables measured, resulting in population level changes to metrics of reproductive performance, including delays in nesting chronology, depressed breeding productivity, and lower chick mass. These effects were strongest at the smallest, southern-most colony, which was most affected by alteration of the Ross Sea Polynya during years the iceberg was present. Additionally, chick mass was negatively correlated with colony size, supporting previous findings indicating density-dependent energetic constraints at the largest colony. Understanding the negative effects of the icebergs on the short-term natural history of Adélie penguins, as well as their response to long-term environmental variation, are important

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. Significance Invasive species can cause extinctions of native species and widespread biodiversity loss. Invader removal is a common management response, but the use of long-term field experiments to characterize effectiveness of removals in benefitting impacted native species is rare. We used a large-scale removal experiment to investigate the demographic response of a threatened native species, the northern spotted owl, to removal of an invasive competitor species, the barred owl. Removal of barred owls had a strong, positive effect on survival of spotted owls, which arrested long-term population declines of spotted owls. The results demonstrate that the long-term persistence of spotted owls will depend heavily on reducing the negative impacts of barred owls while simultaneously addressing other threats, such as habitat loss.