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-...

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The effects of habitat, climate, and Barred Owls on long-term demography of Northern Spotted Owls

The federally threatened northern spotted owl (Strix occidentalis caurina) is the focus of intensive conservation efforts that have led to much forested land being reserved as habitat for the owl and associated wildlife species throughout the Pacific Northwest of the United States. Recently, however, a relatively new threat to spotted owls has emerged in the form of an invasive competitor: the congeneric barred owl (S. varia). As barred owls have rapidly expanded their populations into the entire range of the northern spotted owl, mounting evidence indicates that they are displacing, hybridizing with, and even killing spotted owls. The range expansion by barred owls into western North America has made an already complex conservation issue even more contentious, and a lack of information on the ecological relationships between the 2 species has hampered recovery efforts for northern spotted owls. We investigated spatial relationships, habitat use, diets, survival, and reproduction of sympatric spotted owls and barred owls in western Oregon, USA, during 2007–2009. Our overall objective was to determine the potential for and possible consequences of competition for space, habitat, and food between these previously allopatric owl species. Our study included 29 spotted owls and 28 barred owls that were radio-marked in 36 neighboring territories and monitored over a 24-month period. Based on repeated surveys of both species, the number of territories occupied by pairs of barred owls in the 745-km2 study area (82) greatly outnumbered those occupied by pairs of spotted owls (15). Estimates of mean size of home ranges and core-use areas of spotted owls (1,843 ha and 305 ha, respectively) were 2–4 times larger than those of barred owls (581 ha and 188 ha, respectively). Individual spotted and barred owls in adjacent territories often had overlapping home ranges, but interspecific space sharing was largely restricted to broader foraging areas in the home range with minimal spatial overlap among core-use areas. We used an information-theoretic approach to rank discrete-choice models representing alternative hypotheses about the influence of forest conditions, topography, and interspecific interactions on species-specific patterns of nighttime resource selection. Spotted owls spent a disproportionate amount of time foraging on steep slopes in ravines dominated by old (>120 yr) conifer trees. Barred owls used available forest types more evenly than spotted owls, and were most strongly associated with patches of large hardwood and conifer trees that occupied relatively flat areas along streams. Spotted and barred owls differed in the relative use of old conifer forest (greater for spotted owls) and slope conditions (steeper slopes for spotted owls), but we found no evidence that the 2 species differed in their use of young, mature, and riparian-hardwood forest types. Mean overlap in proportional use of different forest types between individual spotted owls and barred owls in adjacent territories was 81% (range = 30–99%). The best model of habitat use for spotted owls indicated that the relative probability of a location being used was substantially reduced if the location was within or in close proximity to a core-use area of a barred owl. We used pellet analysis and measures of food-niche overlap to determine the potential for dietary competition between spatially associated pairs of spotted owls and barred owls. We identified 1,223 prey items from 15 territories occupied by spotted owls and 4,299 prey items from 24 territories occupied by barred owls. Diets of both species were dominated by nocturnal mammals, but diets of barred owls included many terrestrial, aquatic, and diurnal prey species that were rare or absent in diets of spotted owls. Northern flying squirrels (Glaucomys sabrinus), woodrats (Neotoma fuscipes, N. cinerea), and lagomorphs (Lepus americanus, Sylvilagus bachmani) were primary prey for both owl species, accounting for 81% and 49% of total dietary biomass for spotted owls and barred owls, respectively. Mean dietary overlap between pairs of spotted and barred owls in adjacent territories was moderate (42%; range = 28–70%). Barred owls displayed demographic superiority over spotted owls; annual survival probability of spotted owls from known-fate analyses (0.81, SE = 0.05) was lower than that of barred owls (0.92, SE = 0.04), and pairs of barred owls produced an average of 4.4 times more young than pairs of spotted owls over a 3-year period. We found a strong, positive relationship between seasonal (6-month) survival probabilities of both species and the proportion of old (>120 yr) conifer forest within individual home ranges, which suggested that availability of old forest was a potential limiting factor in the competitive relationship between these 2 species. The annual number of young produced by spotted owls increased linearly with increasing distance from a territory center of a pair of barred owls, and all spotted owls that attempted to nest within 1.5 km of a nest used by barred owls failed to successfully produce young. We identified strong associations between the presence of barred owls and the behavior and fitness potential of spotted owls, as shown by changes in movements, habitat use, and reproductive output of spotted owls exposed to different levels of spatial overlap with territorial barred owls. When viewed collectively, our results support the hypothesis that interference competition with barred owls for territorial space can constrain the availability of critical resources required for successful recruitment and reproduction of spotted owls. Availability of old forests and associated prey species appeared to be the most strongly limiting factors in the competitive relationship between these species, indicating that further loss of these conditions can lead to increases in competitive pressure. Our findings have broad implications for the conservation of spotted owls, as they suggest that spatial heterogeneity in vital rates may not arise solely because of differences among territories in the quality or abundance of forest habitat, but also because of the spatial distribution of a newly established competitor. Experimental removal of barred owls could be used to test this hypothesis and determine whether localized control of barred owl numbers is an ecologically practical and socio-politically acceptable management tool to consider in conservation strategies for spotted owls. © 2013 The Wildlife Society

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Competitive interactions and resource partitioning between northern spotted owls and barred owls in western Oregon

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.

https://www.fws.gov/oregonfwo/Species/Data/NorthernSpottedOwl/BarredOwl/Documents/DuggerEtAl2011.pdf

Transient dynamics of invasive competition: barred owls, spotted owls, habitat, and the demons of competition present.

In Scotland, UK, beavers became extinct about 400 years ago. Currently, two wild populations are present in Scotland on a trial basis, and the case for their full reintroduction is currently being considered by Scottish ministers. Beavers are widely considered ‘ecosystem engineers’. Indeed, beavers have large impacts on the environment, fundamentally change ecosystems, and create unusual habitats, often considered unique. In this review, we investigate the mechanisms by which beavers act as ecosystem engineers, and then discuss the possible impacts of beavers on the biodiversity of Scotland.
A meta-analysis of published studies on beavers’ interactions with biodiversity was conducted, and the balance of positive and negative interactions with plants, invertebrates, amphibians, reptiles, birds, and mammals recorded.
The meta-analysis showed that, overall, beavers have an overwhelmingly positive influence on biodiversity. Beavers’ ability to modify the environment means that they fundamentally increase habitat heterogeneity. As beavers are central-place foragers that feed only in close proximity to watercourses, their herbivory is unevenly spread in the landscape. In addition, beaver ponds and their associated unique successional stages increase habitat heterogeneity both spatially and temporally. Beavers also influence the ecosystems through the creation of a variety of features such as dams and lodges, important habitat features such as standing dead wood (after inundation), an increase in woody debris, and a graded edge between terrestrial and aquatic habitats that is rich in structural complexity.
In Scotland, a widespread positive influence on biodiversity is expected, if beavers are widely reintroduced. For instance, beaver activity should provide important habitat for the otter Lutra lutra, great crested newt Triturus cristatus and water vole Arvicola amphibious, all species of conservation importance.
Beavers are most likely to have detrimental impacts on certain woodland habitats and species of conservation importance, such as the Atlantic hazelwood climax community and aspen Populus tremula woodland. A lack of woodland regeneration caused by high deer abundance could lead to habitat degradation or loss. These are also of particular importance due to the variety of associated dependent species of conservation interest, such as lichen communities in Atlantic hazelwoods.

The impacts of beavers Castor spp. on biodiversity and the ecological basis for their reintroduction to Scotland, UK

In this paper, we modify dynamic occupancy models developed for detection-nondetection data to allow for the dependence of local vital rates on neighborhood occupancy, where neighborhood is defined very flexibly. Such dependence of occupancy dynamics on the status of a relevant neighborhood is pervasive, yet frequently ignored. Our framework permits joint inference about the importance of neighborhood effects and habitat covariates in determining colonization and extinction rates. Our specific motivation is the recent expansion of the Barred Owl (Strix varia) in western Oregon, USA, over the period 1990–2010. Because the focal period was one of dramatic range expansion and local population increase, the use of models that incorporate regional occupancy (sources of colonists) as determinants of dynamic rate parameters is especially appropriate. We began our analysis of 21 years of Barred Owl presence/nondetection data in the Tyee Density Study Area (TDSA) by testing a suite of six models that varied only i...

https://www.fs.fed.us/pnw/pubs/journals/pnw_2012_yackulic.pdf

Neighborhood and habitat effects on vital rates: expansion of the Barred Owl in the Oregon Coast Ranges

During the past century, Barred Owls (Strix varia) expanded their range from forests east of the Great Plains to forests throughout most of central and western North America. Here in Part I, I map more than 12,500 records of Barred Owls in their expanded range from the earliest records to the present, draw the species' distribution in its expanded range and infer the general timing and flow of the range expansion. Evidently, Barred Owls originally traveled across the northern Great Plains via the forested riparian corridors of the Missouri, Yellowstone and Musselshell rivers into east-central Montana by 1873. From there, they accessed western forests in southwestern Montana (1909), moved to northwestern Montana (1922) and then expanded their range in two general directions. They moved north and east to northern Alberta (1934) and Saskatchewan (1948) where they apparently encountered other Barred Owls coming westward from Manitoba. They also moved north to northern British Columbia (1943), southeastern Alaska (1967) and Northwest Territories (1977), and west and south to Washington (1965), Idaho (1968), Oregon (1972) and California (1976). In Part II (Livezey, in press), I explore what prevented Barred Owls from expanding their range westward during recent millennia and what allowed them to do so during the past century.

https://bioone.org/journals/The-American-Midland-Naturalist/volume-161/issue-1/0003-0031-161.1.49/Range-Expansion-of-Barred-Owls-Part-I--Chronology-and/10.1674/0003-0031-161.1.49.pdf

Range Expansion of Barred Owls, Part I: Chronology and Distribution

In Part I (Livezey, 2009), I presented the chronology and distribution of the range expansion of Barred Owls (Strix varia) from the late 1800s to the present. Here I explore what had prevented Barred Owls from expanding their range westward during recent millennia and what allowed them to do so during the past century. Using strength-of-evidence analysis, I evaluate the plausibility of the five ecological or behavioral changes proposed in the literature to have facilitated the range expansion. From this evaluation, three of these changes appear to be implausible, one appears to be plausible after modifying its location, and one appears to be very plausible. For the very plausible one, I score seven ecological changes that may have affected it using five strength-of-evidence criteria. Overall, it appears the historical lack of trees in the Great Plains acted as a barrier to the range expansion and recent increases in forests broke down this barrier. Increases in forest distribution along the Misso...

Kent B. Livezey

Papers

Range Expansion of Barred Owls, Part I: Chronology and Distribution

Range Expansion of Barred Owls, Part II: Facilitating Ecological Changes