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.

Secret of Owls

This chapter looks at the author's experience hiking in the Cumberland Forest, which stretches up from swamps and wetlands, along ridges of hemlocks and pines, into a series of canyons that contain frozen rivers and a couple of small, ice-locked lakes. For three nights, the author had heard the sounds of Vancouver Island northern pygmy-owls. It is not uncommon to hear barred owls or great horned owls in Cumberland Forest, but the unique call of a pygmy-owl is relatively rare, and they had been more active than usual. Northern pygmy-owls were once numerous on the island, but since the 1920s, their numbers have been declining fast due to logging, urban development, and other activities leading to habitat destruction and so the fragmentation of their population.

Passive acoustic monitoring effectively detects Northern Spotted Owls and Barred Owls over a range of forest conditions

Passive acoustic monitoring using autonomous recording units (ARUs) is a fast-growing area of wildlife research especially for rare, cryptic species that vocalize. Northern Spotted Owl (Strix occidentalis caurina) populations have been monitored since the mid-1980s using mark–recapture methods. To evaluate an alternative survey method, we used ARUs to detect calls of Northern Spotted Owls and Barred Owls (S. varia), a congener that has expanded its range into the Pacific Northwest and threatens Northern Spotted Owl persistence. We set ARUs at 30 500-ha hexagons (150 ARU stations) with recent Northern Spotted Owl activity and high Barred Owl density within Northern Spotted Owl demographic study areas in Oregon and Washington, and set ARUs to record continuously each night from March to July, 2017. We reviewed spectrograms (visual representations of sound) and tagged target vocalizations to extract calls from ~160,000 hr of recordings. Even in a study area with low occupancy rates on historical territories (Washington’s Olympic Peninsula), the probability of detecting a Northern Spotted Owl when it was present in a hexagon exceeded 0.95 after 3 weeks of recording. Environmental noise, mainly from rain, wind, and streams, decreased detection probabilities for both species over all study areas. Using demographic information about known Northern Spotted Owls, we found that weekly detection probabilities of Northern Spotted Owls were higher when ARUs were closer to known nests and activity centers and when owls were paired, suggesting passive acoustic data alone could help locate Northern Spotted Owl pairs on the landscape. These results demonstrate that ARUs can effectively detect Northern Spotted Owls when they are present, even in a landscape with high Barred Owl density, thereby facilitating the use of passive, occupancy-based study designs to monitor Northern Spotted Owl populations.

Early detection of rapid Barred Owl population growth within the range of the California Spotted Owl advises the Precautionary Principle

Biological invasions are most practical to manage when invasive species population densities are low. Despite a potentially narrow window of opportunity for efficient management, managers tend to delay intervention because the cost of prompt action is often high and resources are limited. The Barred Owl (Strix varia) invaded and colonized the entire range of the Northern Spotted Owl (S. occidentalis caurina), but insufficient population data contributed to delays in action until the Barred Owl posed an existential threat to the Spotted Owl. The leading edge of the Barred Owl expansion has since reached the Sierra Nevada, the core range of the California Spotted Owl (S. o. occidentalis). We conducted passive acoustic surveys within 400-ha grid cells across ~6,200 km2 in the northern Sierra Nevada and detected a 2.6-fold increase in Barred Owl site occupancy between 2017 and 2018, from 0.082 (85% confidence interval: 0.045–0.12) to 0.21 (0.14–0.28). The probability of Barred Owl site colonization increased with the amount of older forest, suggesting that Barred Owls are first occupying the preferred habitat of Spotted Owls. GPS-tagged Barred Owls (n = 10) generally displayed seasonal and interannual site fidelity over territories averaging 411 ha (range: 150–513 ha), suggesting that our occupancy estimates were not substantially upwardly biased by “double counting” individuals whose territories spanned multiple grid cells. Given the Barred Owl’s demonstrated threat to the Northern Spotted Owl, we believe our findings advise the Precautionary Principle, which posits that management actions such as invasive species removal should be taken despite uncertainties about, for example, true rates of population growth if the cost of inaction is high. In this case, initiating Barred Owl removals in the Sierra Nevada before the population grows further will likely make such action more cost-effective and more humane than if it is delayed. It could also prevent the extirpation of the California Spotted Owl from its core range.

Social status, forest disturbance, and Barred Owls shape long-term trends in breeding dispersal distance of Northern Spotted Owls

Dispersal among breeding sites in territorial animals (i.e. breeding dispersal) is driven by numerous selection pressures, including competition and spatiotemporal variation in habitat quality. The scale and trend of dispersal movements over time may signal changing conditions within the population or on the landscape. We examined 2,158 breeding dispersal events from 694 male and 608 female individually marked Northern Spotted Owls (Strix occidentalis caurina) monitored over 28 yr on 7 study areas to assess the relative importance of individual (sex, experience), reproductive (annual productivity, mate availability), and environmental (forest alteration, presence of competitor) sources of variation in breeding dispersal distance. Median breeding dispersal distance was 3.17 km, with 99% of all breeding dispersal events <37 km. Mean annual dispersal distances increased by 2.43 km in Oregon and 9.40 km in Washington between 1990 and 2017, which coincided with increases in annual detections of nonnative Barred Owl (S. varia). Frequency of breeding dispersal events, both among and within individuals, also increased over time. Female owls moved farther than males (median of 3.26 and 3.10 km, respectively), and birds with less experience (territory tenure) moved farther than those with more experience. Owls that were single in the year prior to dispersal moved 13–31% farther than those paired prior to dispersal. The greatest environmental change occurring over the course of our study was the expansion of Barred Owl populations. Breeding dispersal distance was positively related to Barred Owls in the study area and disturbance within the originating territory. While it appears that social factors continue to be important drivers of breeding dispersal distance in Spotted Owls, increased competition from Barred Owls and habitat alteration have a contributing effect. Increased breeding dispersal distances should be of concern for conservation efforts and considered in population monitoring because changing dispersal behavior may lead to higher rates of mortality and/or emigration from historical study areas.

Barred Owls reduce occupancy and breeding propensity of Northern Spotted Owl in a Washington old-growth forest

Protected lands like national parks are important refuges for threatened and endangered species as environmental pressures on wildlife and their habitats increase. The Northern Spotted Owl (Strix occidentalis caurina), a species designated as threatened under the Endangered Species Act, occurs on public lands throughout the western United States including Mount Rainier National Park (MRNP), Washington. With virtually no history of timber harvest or large forest disturbance within MRNP boundaries since the park’s creation in 1899, MRNP provides an ideal place to evaluate potential impacts of climate change and invasive Barred Owls (Strix varia) on the Northern Spotted Owl. We used a multi-state, multi-season occupancy model to investigate how Northern Spotted Owl occupancy dynamics and breeding propensity are related to the presence of Barred Owls, local and regional weather, and habitat characteristics at MRNP from 1997 to 2016. Historical occupancy of Northern Spotted Owl breeding territories in MRNP has declined by 50% in the last 20 yr, and territory occupancy by breeding Northern Spotted Owls also decreased, reaching a low of 25% in 2016. Occupancy rates were higher on territories with steeper terrain and breeding rates were lower when Barred Owls were detected within historical territories. Our results also indicated that breeding propensity was higher when early nesting season temperatures during March and April were higher. In addition, the ability to detect breeding Northern Spotted Owls decreased when Barred Owls were present in the territory. Habitat variables from LiDAR were not correlated with Northern Spotted Owl occupancy dynamics, likely reflecting the dominance of old-growth forest in this protected park. This study illustrates the strong relationship between Barred Owls and Northern Spotted Owl demographics and breeding site selection in a landscape where habitat loss by timber harvest and fire has not occurred.