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.

Barred owls and landscape attributes influence territory occupancy of northern spotted owls.

We used multi-season occupancy analyses to model 2 fates of northern spotted owl territories in relation to habitat amount, habitat fragmentation, and the presence of barred owls in Washington State, USA, 1989-2005. Local colonization is the probability a territory unoccupied by a spotted owl in year i would be occupied in year i + 1, and local extinction is the probability a territory that was occupied by a spotted owl in year i would be unoccupied in year i + 1. We found a negative relationship between local extinction probability and amount of late-seral forest edge. We found a negative relationship between colonization probability and the number of late-seral forest patches (higher fragmentation), and a negative relationship between colonization probability and the amount of non-habitat within 600 m of a spotted owl territory center (Akaike weight = 0.59). The presence of barred owls was positively related to extinction probability and negatively related to detection probability of spotted owls. The negative relationship between presence of barred owls and detectability of spotted owls indicated that spotted owls could be modifying their calling behavior in the presence of barred owls. The positive relationship between barred owl detections and local extinction probability suggests that because of competition with barred owls, spotted owls are being displaced. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

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

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.

Organochlorine contaminants in fishes from coastal waters west of Amukta Pass, Aleutian Islands, Alaska, USA.

Organochlorines were examined in liver and stable isotopes in muscle of fishes from the western Aleutian Islands, Alaska, in relation to islands or locations affected by military occupation. Pacific cod (Gadus macrocephalus), Pacific halibut (Hippoglossus stenolepis), and rock greenling (Hexagrammos lagocephalus) were collected from nearshore waters at contemporary (decommissioned) and historical (World War II) military locations, as well as at reference locations. Total (Sigma) polychlorinated biphenyls (PCBs) dominated the suite of organochlorine groups (SigmaDDTs, Sigmachlordane cyclodienes, Sigmaother cyclodienes, and Sigmachlorinated benzenes and cyclohexanes) detected in fishes at all locations, followed by SigmaDDTs and Sigmachlordanes; dichlorodiphenyldichloroethylene (p,p'DDE) composed 52 to 66% of SigmaDDTs by species. Organochlorine concentrations were higher or similar in cod compared to halibut and lowest in greenling; they were among the highest for fishes in Arctic or near Arctic waters. Organochlorine group concentrations varied among species and locations, but SigmaPCB concentrations in all species were consistently higher at military locations than at reference locations. Moreover, all organochlorine group concentrations were higher in halibut from military locations than those from reference locations. A wide range of molecular weight organochlorines was detected at all locations, which implied regional or long-range transport and deposition, as well as local point-source contamination. Furthermore, a preponderance of higher-chlorinated PCB congeners in fishes from contemporary military islands implied recent exposure. Concentrations in all organochlorine groups increased with delta15N enrichment in fishes, and analyses of residual variation provided further evidence of different sources of SigmaPCBs and p,p'DDE among species and locations.

Bald eagles and sea otters in the Aleutian Archipelago: indirect effects of trophic cascades.

Because sea otters (Enhydra lutris) exert a wide array of direct and indirect effects on coastal marine ecosystems throughout their geographic range, we investigated the potential influence of sea otters on the ecology of Bald Eagles (Haliaeetus leucocephalus) in the Aleutian Islands, Alaska, USA. We studied the diets, productivity, and density of breeding Bald Eagles on four islands during 1993-1994 and 2000-2002, when sea otters were abundant and scarce, respectively. Bald Eagles depend on nearshore marine communities for most of their prey in this ecosystem, so we predicted that the recent decline in otter populations would have an indirect negative effect on diets and demography of Bald Eagles. Contrary to our predictions, we found no effects on density of breeding pairs on four islands from 1993-1994 to 2000-2002. In contrast, diets and diet diversity of Bald Eagles changed considerably between the two time periods, likely reflecting a change in prey availability resulting from the increase and subsequent decline in sea otter populations. The frequency of sea otter pups, rock greenling (Hexagammus lagocephalus), and smooth lumpsuckers (Aptocyclus ventricosus) in the eagle's diet declined with corresponding increases in Rock Ptarmigan (Lagopus mutus), Glaucous-winged Gulls (Larus glaucescens), Atka mackerel (Pleurogrammus monopterygius), and various species of seabirds during the period of the recent otter population decline. Breeding success and productivity of Bald Eagles also increased during this time period, which may be due to the higher nutritional quality of avian prey consumed in later years. Our results provide further evidence of the wide-ranging indirect effects of sea otter predation on nearshore marine communities and another apex predator, the Bald Eagle. Although the indirect effects of sea otters are widely known, this example is unique because the food-web pathway transcended five species and several trophic levels in linking one apex predator to another.

Sources of organochlorine contaminants and mercury in seabirds from the Aleutian archipelago of Alaska: inferences from spatial and trophic variation.

Persistent organochlorine compounds and mercury (Hg) have been detected in numerous coastal organisms of the Aleutian archipelago of Alaska, yet sources of these contaminants are unclear. We collected glaucous-winged gulls, northern fulmars, and tufted puffins along a natural longitudinal gradient across the western and central Aleutian Islands (Buldir, Kiska, Amchitka, Adak), and an additional 8 seabird species representing different foraging and migratory guilds from Buldir Island to evaluate: 1) point source input from former military installations, 2) westward increases in contaminant concentrations suggestive of distant source input, and 3) effects of trophic status (delta15N) and carbon source (delta13C) on contaminant accumulation. Concentrations of Sigma polychlorinated biphenyls (PCBs) and most chlorinated pesticides in glaucous-winged gulls consistently exhibited a 'U'-shaped pattern of high levels at Buldir and the east side of Adak and low levels at Kiska and Amchitka. In contrast, concentrations of Sigma PCBs and chlorinated pesticides in northern fulmars and tufted puffins did not differ among islands. Hg concentrations increased westward in glaucous-winged gulls and were highest in northern fulmars from Buldir. Among species collected only at Buldir, Hg was notably elevated in pelagic cormorants, and relatively high Sigma PCBs were detected in black-legged kittiwakes. Concentrations of Sigma PCBs, dichlorodiphenyldichloroethylene (p,p' DDE), and Hg were positively correlated with delta15N across all seabird species, indicating biomagnification across trophic levels. The east side of Adak Island (a former military installation) was a likely point source of Sigma PCBs and p,p' DDE, particularly in glaucous-winged gulls. In contrast, elevated levels of these contaminants and Hg, along with PCB congener and chlorinated pesticide compositional patterns detected at Buldir Island indicated exposure from distant sources influenced by a combination of atmospheric-oceanic processes and the migratory movements of seabirds.

Environmental contaminants in bald eagle eggs from the Aleutian archipelago.

We collected 136 fresh and unhatched eggs from bald eagle (Haliaeetus leucocephalus) nests and assessed productivity on eight islands in the Aleutian archipelago, 2000 to 2002. Egg contents were analyzed for a broad spectrum of organochlorine (OC) contaminants, mercury (Hg), and stable isotopes of carbon (delta13C) and nitrogen (delta15N). Concentrations of polychlorinated biphenyls (SigmaPCBs), p,p'-dichlorodiphenyldichloroethylene (DDE), and Hg in bald eagle eggs were elevated throughout the archipelago, but the patterns of distribution differed among the various contaminants. Total PCBs were highest in areas of past military activities on Adak and Amchitka Islands, indicating local point sources of these compounds. Concentrations of DDE and Hg were higher on Amchitka Island, which was subjected to much military activity during World War II and the middle of the 20th century. Concentrations of SigmaPCBs also were elevated on islands with little history of military activity (e.g., Amlia, Tanaga, Buldir), suggesting non-point sources of PCBs in addition to point sources. Concentrations of DDE and Hg were highest in eagle eggs from the most western Aleutian Islands (e.g., Buldir, Kiska) and decreased eastward along the Aleutian chain. This east-to-west increase suggested a Eurasian source of contamination, possibly through global transport and atmospheric distillation and/or from migratory seabirds. Eggshell thickness and productivity of bald eagles were normal and indicative of healthy populations because concentrations of most contaminants were below threshold levels for effects on reproduction. Contrary to our predictions, contaminant concentrations were not correlated with stable isotopes of carbon (delta13C) or nitrogen (delta15N) in eggs. These latter findings indicate that contaminant concentrations were influenced more by point sources and geographic location than trophic status of eagles among the different islands.

Interactive effects of prey and p,p'-DDE on burrowing owl population dynamics.

We used population models to explore the effects of the organochlorine contaminant p,p'-DDE and fluctuations in vole availability on the population dynamics of Burrowing Owls (Athene cunicularia). Previous work indicated an interaction between low biomass of voles in the diet and moderate levels of p,p'-DDE in Burrowing Owl eggs that led to reproductive impairment. We constructed periodic and stochastic matrix models that incorporated three vole population states observed in the field: average, peak, and crash years. We modeled varying frequencies of vole crash years and a range of impairment of owl demographic rates in vole crash years. Vole availability had a greater impact on owl population growth rate than did reproductive impairment if vole populations peaked and crashed frequently. However, this difference disappeared as the frequency of vole crash years declined to once per decade. Fecundity, the demographic rate most affected by p,p'-DDE, had less impact on population growth rate than adult or juvenile survival. A life table response experiment of time-invariant matrices for average, peak, and crash vole conditions showed that low population growth under vole crash conditions was due to low adult and juvenile survival rates, whereas the extremely high population growth under vole peak conditions was due to increased fecundity. Our results suggest that even simple models can provide useful insights into complex ecological interactions. This is particularly valuable when temporal or spatial scales preclude manipulative experimental work in the field or laboratory.

Changes in productivity and contaminants in bald eagles nesting along the lower Columbia River, USA.

Previous studies documented poor productivity of bald eagles (Haliaeetus leucocephalus) in the lower Columbia River (LCR), USA, and elevated p,p'-dichlorodiphenyldichloroethylene (DDE), polychlorinated biphenyls (PCBs), dioxins, and furans in eagle eggs. From 1994 to 1995, we collected partially incubated eggs at 19 of 43 occupied territories along the LCR and compared productivity and egg contaminants to values obtained in the mid-1980s. We found higher productivity at new nesting sites along the river, yet productivity at 23 older breeding territories remained low and was not different (p = 0.713) between studies. Eggshell thickness at older territories had not improved (p = 0.404), and eggshells averaged 11% thinner than shells measured before dichlorodiphenyltrichloroethane use. Decreases in DDE (p = 0.022) and total PCBs (p = 0.0004) in eggs from older breeding areas occurred between study periods. Productivity was not correlated to contaminants, but DDE, PCBs, and dioxin-like chemicals exceeded estimated no-effect values. Some dioxin-like contaminants in eggs were correlated to nest location, with highest concentrations occurring toward the river's mouth where productivity was lowest. Although total productivity increased due to the success of new nesting pairs in the region, egg contaminants remain high enough to impair reproduction at older territories and, over time, may alter productivity of new pairs nesting near the river's mouth.