Mark A. Ricca

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 eagles 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.

Sexual Segregation in Roosevelt Elk: Cropping Rates and Aggression in Mixed-Sex Groups

Abstract Few studies of sexual segregation in ruminants have tested widely invoked mechanisms of segregation in mixed-sex groups. In a sexually segregated population of Roosevelt elk (Cervus elaphus roosevelti), we examined if adult males had reduced intake of forage when in mixed-sex groups and if intersexual differences in aggression caused females to avoid males. Based on a mechanistic model of forage intake, animals with lower instantaneous feed intake should have higher cropping rates. Focal animal sampling indicated that adult males and females in summer and winter had similar cropping rates in mixed-sex groups, whereas males in male-only groups had lower rates of cropping than males in mixed-sex groups. Outside the mating season, males in male groups spent proportionally less time ≤1 body length of congenders than females in female groups, and the rate of aggression ≤1 body length was higher for males. Female–female aggression was higher in mixed-sex groups that contained more males than the median proportion of males in mixed-sex groups. Female and mixed-sex groups walked away when groups of males numbering >6 were ≤50 m but did not walk away when male groups ≤50 m had ≤5 individuals. Sexual segregation was associated with behaviors of sexes in mixed-sex groups: reduced intake of forage by males and increased female–female aggression with more males.

Survival of Columbian white-tailed deer in western Oregon

Columbian white-tailed deer (Odocoileus virginianus leucurus; CWTD) are an endangered subspecies on which little demographic information exists. We determined survival rates and causes of mortality for 64 radiocollared adults from 1996 to 1998, and for 63 radiocollared neonatal fawns during the summer and fall months of 1996-2001 in Douglas County, Oregon, USA. Annual adult survival rates averaged 0.74 over 3 years, and most mortality (73%) occurred between fall and winter. Seasonal survival was lowest (0.75) for the fall-winter 1997-1998, and was ≥0.90 during all spring-summer periods. Annual and seasonal survival rates did not differ by gender. Average annual survival was 0.77 for deer in wildland areas compared with 0.66 for deer in suburban areas, but these differences were not consistent between years and seasons. Survival over the entire 3-year study was low (0.38). Eight deer died from a combination of emaciation and disease, and almost all (92%) necropsied deer were in poor body condition. Fawn survival to 7 months was low (0.14, 95% CI = 0.02-0.26) and declined most rapidly during the first 1.5 months of life. Predation (n = 21) and abandonment (n = 6) were the most frequent known causes of death for fawns. Our results suggest that CWTD may have responded to density-dependent factors during this short-term study, although the effects of other environmental or intrinsic factors cannot be ignored. Fawn survival may be insufficient to produce enough recruits for population growth and eventual range expansion.

Polycyclic aromatic hydrocarbon exposure in Steller's eiders (Polysticta stelleri) and harlequin ducks (Histronicus histronicus) in the Eastern Aleutian Islands, Alaska, USA

Seaducks may be affected by harmful levels of polycyclic aromatic hydrocarbons (PAHs) at seaports near the Arctic. As an indicator of exposure to PAHs, we measured hepatic enzyme 7-ethoxyresorufin-O-deethylase activity (EROD) to determine cytochrome P4501A induction in Stellers eiders (Polysticta stelleri) and Harlequin ducks (Histronicus histronicus) from Unalaska, Popof, and Unga Islands (AK, USA) in 2002 and 2003. We measured PAHs and organic contaminants in seaduck prey samples and polychlorinated biphenyl congeners in seaduck blood plasma to determine any relationship to EROD. Using Akaikes information criterion, species and site differences best explained EROD patterns: Activity was higher in Harlequin ducks than in Stellers eiders and higher at industrial than at nonindustrial sites. Site-specific concentrations of PAHs in blue mussels ([Mytilus trossilus] seaduck prey; PAH concentrations higher at Dutch Harbor, Unalaska, than at other sites) also was important in defining EROD patterns. Organochlorine compounds rarely were detected in prey samples. No relationship was found between polychlorinated biphenyl congeners in avian blood and EROD, which further supported inferences derived from Akaikes information criterion. Congeners were highest in seaducks from a nonindustrial or reference site, contrary to PAH patterns. To assist in interpreting the field study, 15 captive Stellers eiders were dosed with a PAH known to induce cytochrome P4501A. Dosed, captive Stellers eiders had definitive induction, but results indicated that wild Stellers eiders were exposed to PAHs or other inducing compounds at levels greater than those used in laboratory studies. Concentrations of PAHs in blue mussels at or near Dutch Harbor (approximately 1,180-5,980 ng/g) approached those found at highly contaminated sites (approximately 4,100-7,500 ng/g).

Roosevelt Elk Density and Social Segregation: Foraging Behavior and Females Avoiding Larger Groups of Males

Abstract Intersexual social segregation at small spatial scales is prevalent in ruminants that are sexually dimorphic in body size. Explaining social segregation, however, from hypotheses of how intersexual size differences affects the foraging process of males and females has had mixed results. We studied whether body size influences on forage behavior, intersexual social incompatibility or both might influence social segregation in a population of Roosevelt elk (Cervus elaphus roosevelti) that declined 40% over 5 y. Most males and females in the population occurred in the same forage patches, meadows, but occupied different parts of meadows and most groups were overwhelming comprised of one sex. The extent of segregation varied slightly with changing elk density. Cropping rate, our surrogate of forage ingestion, of males in mixed-sex groups differed from males in male-only groups at high, but not low, elk density. In a prior study of intersexual social interactions it was shown that females avoided groups containing ≥6 males. Therefore, we predicted that females should avoid parts of meadows where groups of males ≥6 were prevalent. Across the 5 y of study this prediction held because ≤5% of all females were found in parts of meadows where median aggregation sizes of males were ≥6. Social segregation was coupled to body size influences on forage ingestion at high density and social incompatibility was coupled to social segregation regardless of elk density.

Wildfire, climate, and invasive grass interactions negatively impact an indicator species by reshaping sagebrush ecosystems

Significance The Great Basin of western North America is larger than 75% of countries worldwide and is comprised mostly of a “sagebrush sea” threatened by a novel disturbance cycle of wildfire and annual grass invasion. The greater sage-grouse is a sagebrush-obligate species whose populations generally track declines in sagebrush, and is highly influential in shaping state and national land-use policy. Using three decades of sage-grouse population count, wildfire, and climate data within a modeling framework that allowed for variable postfire recovery of sagebrush, we provide quantitative evidence that links long-term declines of sage-grouse to chronic effects of wildfire. Projected declines may be slowed or halted by targeting fire suppression in remaining areas of intact sagebrush with high densities of breeding sage-grouse. Iconic sagebrush ecosystems of the American West are threatened by larger and more frequent wildfires that can kill sagebrush and facilitate invasion by annual grasses, creating a cycle that alters sagebrush ecosystem recovery post disturbance. Thwarting this accelerated grass–fire cycle is at the forefront of current national conservation efforts, yet its impacts on wildlife populations inhabiting these ecosystems have not been quantified rigorously. Within a Bayesian framework, we modeled 30 y of wildfire and climatic effects on population rates of change of a sagebrush-obligate species, the greater sage-grouse, across the Great Basin of western North America. Importantly, our modeling also accounted for variation in sagebrush recovery time post fire as determined by underlying soil properties that influence ecosystem resilience to disturbance and resistance to invasion. Our results demonstrate that the cumulative loss of sagebrush to direct and indirect effects of wildfire has contributed strongly to declining sage-grouse populations over the past 30 y at large spatial scales. Moreover, long-lasting effects from wildfire nullified pulses of sage-grouse population growth that typically follow years of higher precipitation. If wildfire trends continue unabated, model projections indicate sage-grouse populations will be reduced to 43% of their current numbers over the next three decades. Our results provide a timely example of how altered fire regimes are disrupting recovery of sagebrush ecosystems and leading to substantial declines of a widespread indicator species. Accordingly, we present scenario-based stochastic projections to inform conservation actions that may help offset the adverse effects of wildfire on sage-grouse and other wildlife populations.

Body weight and rumen-reticulum capacity in tule elk and mule deer

Abstract The relationship between body size and rumen–reticulum capacity among conspecific individuals is predicted to be isometric. We examined whether the relationship between body weight and rumen–reticulum capacity was isometric in adult male and female tule elk (Cervus elaphus nannodes) and in adult female mule deer (Odocoileus hemionus). We detected no effect of sex on this relationship in elk, and the slope of the regression was 1.0 for one measure of rumen–reticulum capacity and <1.0 for another. Among deer, the slope of the relationship was <1.0 in one measure of rumen–reticulum capacity, and we detected no relationship with the other.

Pinyon and Juniper Encroachment into Sagebrush Ecosystems Impacts Distribution and Survival of Greater Sage-Grouse☆,☆☆

ABSTRACT In sagebrush (Artemisia spp.) ecosystems, encroachment of pinyon (Pinus spp.) and juniper (Juniperus spp.; hereafter, “pinyon-junipeR&Rdquo;) trees has increased dramatically since European settlement. Understanding the impacts of this encroachment on behavioral decisions, distributions, and population dynamics of greater sage-grouse (Centrocercus urophasianus) and other sagebrush obligate species could help benefit sagebrush ecosystem management actions. We employed a novel two-stage Bayesian model that linked avoidance across different levels of pinyon-juniper cover to sage-grouse survival. Our analysis relied on extensive telemetry data collected across 6 yr and seven subpopulations within the Bi-State Distinct Population Segment (DPS), on the border of Nevada and California. The first model stage indicated avoidance behavior for all canopy cover classes on average, but individual grouse exhibited a high degree of heterogeneity in avoidance behavior of the lowest cover class (e.g., scattered isolated trees). The second stage modeled survival as a function of estimated avoidance parameters and indicated increased survival rates for individuals that exhibited avoidance of the lowest cover class. A post hoc frailty analysis revealed the greatest increase in hazard (i.e., mortality risk) occurred in areas with scattered isolated trees consisting of relatively high primary plant productivity. Collectively, these results provide clear evidence that local sage-grouse distributions and demographic rates are influenced by pinyon-juniper, especially in habitats with higher primary productivity but relatively low and seemingly benign tree cover. Such areas may function as ecological traps that convey attractive resources but adversely affect population vital rates. To increase sage-grouse survival, our model predictions support reducing actual pinyon-juniper cover as low as 1.5%, which is lower than the published target of 4.0%. These results may represent effects of pinyon-juniper cover in areas with similar ecological conditions to those of the Bi-State DPS, where populations occur at relatively high elevations and pinyon-juniper is abundant and widespread.

Integrating spatially explicit indices of abundance and habitat quality: an applied example for greater sage-grouse management.

Summary Predictive species distributional models are a cornerstone of wildlife conservation planning. Constructing such models requires robust underpinning science that integrates formerly disparate data types to achieve effective species management. Greater sage‐grouse Centrocercus urophasianus, hereafter ‘sage‐grouse’ populations are declining throughout sagebrush‐steppe ecosystems in North America, particularly within the Great Basin, which heightens the need for novel management tools that maximize the use of available information. Herein, we improve upon existing species distribution models by combining information about sage‐grouse habitat quality, distribution and abundance from multiple data sources. To measure habitat, we created spatially explicit maps depicting habitat selection indices (HSI) informed by >35 500 independent telemetry locations from >1600 sage‐grouse collected over 15 years across much of the Great Basin. These indices were derived from models that accounted for selection at different spatial scales and seasons. A region‐wide HSI was calculated using the HSI surfaces modelled for 12 independent subregions and then demarcated into distinct habitat quality classes. We also employed a novel index to describe landscape patterns of sage‐grouse abundance and space use (AUI). The AUI is a probabilistic composite of the following: (i) breeding density patterns based on the spatial configuration of breeding leks and associated trends in male attendance; and (ii) year‐round patterns of space use indexed by the decreasing probability of use with increasing distance to leks. The continuous AUI surface was then reclassified into two classes representing high and low/no use and abundance. Synthesis and applications. Using the example of sage‐grouse, we demonstrate how the joint application of indices of habitat selection, abundance and space use derived from multiple data sources yields a composite map that can guide effective allocation of management intensity across multiple spatial scales. As applied to sage‐grouse, the composite map identifies spatially explicit management categories within sagebrush steppe that are most critical to sustaining sage‐grouse populations as well as those areas where changes in land use would likely have minimal impact. Importantly, collaborative efforts among stakeholders guide which intersections of habitat selection indices and abundance and space use classes are used to define management categories. Because sage‐grouse are an umbrella species, our joint‐index modelling approach can help target effective conservation for other sagebrush obligate species and can be readily applied to species in other ecosystems with similar life histories, such as central‐placed breeding.

PCB exposure in sea otters and harlequin ducks in relation to history of contamination by the Exxon Valdez oil spill.

Exposure to contaminants other than petroleum hydrocarbons could confound interpretation of Exxon Valdez oil spill effects on biota at Prince William Sound, Alaska. Hence, we investigated polychlorinated biphenyls (PCBs) in blood of sea otters and harlequin ducks sampled during 1998. PCB concentrations characterized by lower chlorinated congeners were highest in sea otters from the unoiled area, whereas concentrations were similar among harlequin ducks from the oiled and unoiled area. Blood enzymes often elevated by xenobiotics were not related to PCB concentrations in sea otters. Only sea otters from the unoiled area had estimated risk from PCBs, and PCB composition or concentrations did not correspond to reported lower measures of population performance in sea otters or harlequin ducks from the oiled area. PCBs probably did not influence limited sea otter or harlequin duck recovery in the oiled area a decade after the spill.