Reproductive Fate of Brucellosis-Seropositive Elk (Cervus canadensis): Implications for Disease Transmission Risk.

Brucellosis is a disease caused by the bacterium Brucella abortus that infects elk (Cervus canadensis) and cattle (Bos taurus). There is the potential for transmission from wildlife to livestock through contact with infected material shed during abortions or live births. To understand the impact of exposure on pregnancy rates we captured 30-100 elk per year from 2011 through 2020, testing their blood for serologic exposure to B. abortus. Predicted pregnancy rates for seropositive animals were 9.6% lower in prime-age (2.5-15.5 yr; 85%, 95% confidence interval [CI]: 74-91%) and 37.7% lower in old (>15.5 yr; 43%, 95% CI: 19-71%) elk as compared with seronegative animals. To understand the risk of seropositive elk shedding B. abortus bacteria and the effects of exposure on elk reproductive performance, we conducted a 5-yr longitudinal study monitoring 30 seropositive elk. We estimated the annual probability of a seropositive elk having an abortion as 0.06 (95% CI: 0.02-0.15). We detected B. abortus at three abortions and two live births, using a combination of culture and PCR testing. The predicted probability of a pregnant seropositive elk shedding B. abortus during an abortion or live birth was 0.08 (95% CI: 0.04-0.19). To understand what proportion of seropositive elk harbored live B. abortus bacteria in their tissues, we euthanized seropositive elk at the end of 5 yr of monitoring and sampled tissues for B. abortus. Assuming perfect detection, the predicted probability of a seropositive elk having B. abortus in at least one tissue was 0.18 (95% CI: 0.06-0.43). The transmission risk seropositive elk pose is mitigated by decreased pregnancy rates, low probability of abortion events, low probability of shedding at live birth events, and reasonably low probability of B. abortus in tissues.

Evaluating the summer landscapes of predation risk and forage quality for elk (Cervus canadensis).

The recovery of carnivore populations in North American has consequences for trophic interactions and population dynamics of prey. In addition to direct effects on prey populations through killing, predators can influence prey behavior by imposing the risk of predation. The mechanisms through which patterns of space use by predators are linked to behavioral response by prey and nonconsumptive effects on prey population dynamics are poorly understood. Our goal was to characterize population- and individual-level patterns of resource selection by elk (Cervus canadensis) in response to risk of wolves (Canis lupus) and mountain lions (Puma concolor) and evaluate potential nonconsumptive effects of these behavioral patterns. We tested the hypothesis that individual elk risk-avoidance behavior during summer would result in exposure to lower-quality forage and reduced body fat and pregnancy rates. First, we evaluated individuals' second-order and third-order resource selection with a used-available sampling design. At the population level, we found evidence for a positive relationship between second- and third-order selection and forage, and an interaction between forage quality and mountain lion risk such that the relative probability of use at low mountain lion risk increased with forage quality but decreased at high risk at both orders of selection. We found no evidence of a population-level trade-off between forage quality and wolf risk. However, we found substantial among-individual heterogeneity in resource selection patterns such that population-level patterns were potentially misleading. We found no evidence that the diversity of individual resource selection patterns varied predictably with available resources, or that patterns of individual risk-related resource selection translated into biologically meaningful changes in body fat or pregnancy rates. Our work highlights the importance of evaluating individual responses to predation risk and predator hunting technique when assessing responses to predators and suggests nonconsumptive effects are not operating at a population scale in this system.

Non-invasive monitoring of multiple wildlife health factors by fecal microbiome analysis.

Fecal microbial biomarkers represent a less invasive alternative for acquiring information on wildlife populations than many traditional sampling methodologies. Our goal was to evaluate linkages between fecal microbiome communities in Rocky Mountain elk (Cervus canadensis) and four host factors including sex, age, population, and physical condition (body-fat). We paired a feature-selection algorithm with an LDA-classifier trained on elk differential bacterial abundance (16S-rRNA amplicon survey) to predict host health factors from 104 elk microbiomes across four elk populations. We validated the accuracy of the various classifier predictions with leave-one-out cross-validation using known measurements. We demonstrate that the elk fecal microbiome can predict the four host factors tested. Our results show that elk microbiomes respond to both the strong extrinsic factor of biogeography and simultaneously occurring, but more subtle, intrinsic forces of individual body-fat, sex, and age-class. Thus, we have developed and described herein a generalizable approach to disentangle microbiome responses attributed to multiple host factors of varying strength from the same bacterial sequence data set. Wildlife conservation and management presents many challenges, but we demonstrate that non-invasive microbiome surveys from scat samples can provide alternative options for wildlife population monitoring. We believe that, with further validation, this method could be broadly applicable in other species and potentially predict other measurements. Our study can help guide the future development of microbiome-based monitoring of wildlife populations and supports hypothetical expectations found in host-microbiome theory.

Sex-Specific Elk Resource Selection during the Anthrax Risk Period.

Anthrax, caused by the spore-forming bacterium Bacillus anthracis, is a zoonosis affecting animals and humans globally. In the United States, anthrax outbreaks occur in wildlife and livestock, with frequent outbreaks in native and exotic wildlife species in Texas, livestock outbreaks in the Dakotas, and sporadic mixed outbreaks in Montana. Understanding where pathogen and host habitat selection overlap is essential for anthrax management. Resource selection and habitat use of ungulates may be sex-specific and lead to differential anthrax exposure risks across the landscape for males and females. We evaluated female elk (Cervus canadensis) resource selection in the same study areas as male elk in a previous anthrax risk study to identify risk of anthrax transmission to females and compare transmission risk between females and males. We developed a generalized linear mixed-effect model to estimate resource selection for female elk in southwest Montana during the June to August anthrax transmission risk period. We then predicted habitat selection of female and male elk across the study area and compared selection with the distribution of anthrax risk to identify spatial distributions of potential anthrax exposure for the male and female elk. Female and male elk selected different resources during the anthrax risk period, which resulted in different anthrax exposure areas for females and males. The sex-specific resource selection and habitat use could infer different areas of risk for anthrax transmission, which can improve anthrax and wildlife management and have important public health and economic implications.

Elk migration influences the risk of disease spillover in the Greater Yellowstone Ecosystem.

Wildlife migrations provide important ecosystem services, but they are declining. Within the Greater Yellowstone Ecosystem (GYE), some elk Cervus canadensis herds are losing migratory tendencies, which may increase spatiotemporal overlap between elk and livestock (domestic bison Bison bison and cattle Bos taurus), potentially exacerbating pathogen transmission risk. We combined disease, movement, demographic and environmental data from eight elk herds in the GYE to examine the differential risk of brucellosis transmission (through aborted foetuses) from migrant and resident elk to livestock. For both migrants and residents, we found that transmission risk from elk to livestock occurred almost exclusively on private ranchlands as opposed to state or federal grazing allotments. Weather variability affected the estimated distribution of spillover risk from migrant elk to livestock, with a 7%-12% increase in migrant abortions on private ranchlands during years with heavier snowfall. In contrast, weather variability did not affect spillover risk from resident elk. Migrant elk were responsible for the majority (68%) of disease spillover risk to livestock because they occurred in greater numbers than resident elk. On a per-capita basis, however, our analyses suggested that resident elk disproportionately contributed to spillover risk. In five of seven herds, we estimated that the per-capita spillover risk was greater from residents than from migrants. Averaged across herds, an individual resident elk was 23% more likely than an individual migrant elk to abort on private ranchlands. Our results demonstrate links between migration behaviour, spillover risk and environmental variability, and highlight the utility of integrating models of pathogen transmission and host movement to generate new insights about the role of migration in disease spillover risk. Furthermore, they add to the accumulating body of evidence across taxa that suggests that migrants and residents should be considered separately during investigations of wildlife disease ecology. Finally, our findings have applied implications for elk and brucellosis in the GYE. They suggest that managers should prioritize actions that maintain spatial separation of elk and livestock on private ranchlands during years when snowpack persists into the risk period.

Wave-like Patterns of Plant Phenology Determine Ungulate Movement Tactics.

Animals exhibit a diversity of movement tactics [1]. Tracking resources that change across space and time is predicted to be a fundamental driver of animal movement [2]. For example, some migratory ungulates (i.e., hooved mammals) closely track the progression of highly nutritious plant green-up, a phenomenon called "green-wave surfing" [3-5]. Yet general principles describing how the dynamic nature of resources determine movement tactics are lacking [6]. We tested an emerging theory that predicts surfing and the existence of migratory behavior will be favored in environments where green-up is fleeting and moves sequentially across large landscapes (i.e., wave-like green-up) [7]. Landscapes exhibiting wave-like patterns of green-up facilitated surfing and explained the existence of migratory behavior across 61 populations of four ungulate species on two continents (n = 1,696 individuals). At the species level, foraging benefits were equivalent between tactics, suggesting that each movement tactic is fine-tuned to local patterns of plant phenology. For decades, ecologists have sought to understand how animals move to select habitat, commonly defining habitat as a set of static patches [8, 9]. Our findings indicate that animal movement tactics emerge as a function of the flux of resources across space and time, underscoring the need to redefine habitat to include its dynamic attributes. As global habitats continue to be modified by anthropogenic disturbance and climate change [10], our synthesis provides a generalizable framework to understand how animal movement will be influenced by altered patterns of resource phenology.

Evaluating wildlife translocations using genomics: A bighorn sheep case study.

Wildlife restoration often involves translocation efforts to reintroduce species and supplement small, fragmented populations. We examined the genomic consequences of bighorn sheep (Ovis canadensis) translocations and population isolation to enhance understanding of evolutionary processes that affect population genetics and inform future restoration strategies. We conducted a population genomic analysis of 511 bighorn sheep from 17 areas, including native and reintroduced populations that received 0-10 translocations. Using the Illumina High Density Ovine array, we generated datasets of 6,155 to 33,289 single nucleotide polymorphisms and completed clustering, population tree, and kinship analyses. Our analyses determined that natural gene flow did not occur between most populations, including two pairs of native herds that had past connectivity. We synthesized genomic evidence across analyses to evaluate 24 different translocation events and detected eight successful reintroductions (i.e., lack of signal for recolonization from nearby populations) and five successful augmentations (i.e., reproductive success of translocated individuals) based on genetic similarity with the source populations. A single native population founded six of the reintroduced herds, suggesting that environmental conditions did not need to match for populations to persist following reintroduction. Augmentations consisting of 18-57 animals including males and females succeeded, whereas augmentations of two males did not result in a detectable genetic signature. Our results provide insight on genomic distinctiveness of native and reintroduced herds, information on the relative success of reintroduction and augmentation efforts and their associated attributes, and guidance to enhance genetic contribution of augmentations and reintroductions to aid in bighorn sheep restoration.

Characterizing population and individual migration patterns among native and restored bighorn sheep (Ovis canadensis).

Migration evolved as a behavior to enhance fitness through exploiting spatially and temporally variable resources and avoiding predation or other threats. Globally, landscape alterations have resulted in declines to migratory populations across taxa. Given the long time periods over which migrations evolved in native systems, it is unlikely that restored populations embody the same migratory complexity that existed before population reductions or regional extirpation.We used GPS location data collected from 209 female bighorn sheep (Ovis canadensis) to characterize population and individual migration patterns along elevation and geographic continuums for 18 populations of bighorn sheep with different management histories (i.e., restored, augmented, and native) across the western United States.Individuals with resident behaviors were present in all management histories. Elevational migrations were the most common population-level migratory behavior. There were notable differences in the degree of individual variation within a population across the three management histories. Relative to native populations, restored and augmented populations had less variation among individuals with respect to elevation and geographic migration distances. Differences in migratory behavior were most pronounced for geographic distances, where the majority of native populations had a range of variation that was 2-4 times greater than restored or augmented populations. Synthesis and applications. Migrations within native populations include a variety of patterns that translocation efforts have not been able to fully recreate within restored and augmented populations. Theoretical and empirical research has highlighted the benefits of migratory diversity in promoting resilience and population stability. Limited migratory diversity may serve as an additional factor limiting demographic performance and range expansion. We suggest preserving native systems with intact migratory portfolios and a more nuanced approach to restoration and augmentation in which source populations are identified based on a suite of criteria that includes matching migratory patterns of source populations with local landscape attributes.

Native forage mediates influence of irrigated agriculture on migratory behaviour of elk.

Ungulates migrate to maximize nutritional intake when forage varies seasonally. Populations of ungulates often include both migratory and non-migratory individuals, but the mechanisms driving individual differences in migratory behaviour are not well-understood. We quantified associations between hypothesized drivers of partial migration and the likelihood of migration for individual ungulates that experienced a range of environmental conditions and anthropogenic influences. We evaluated the effects of forage variation, conspecific density, and human land uses on migratory behaviour of 308 adult female elk in 16 herds across western Montana. We found irrigated agriculture on an individual's winter range reduced migratory behaviour, but individuals were more likely to migrate away from irrigated agricultural areas if better forage was available elsewhere or if they experienced high conspecific density on their winter range. When the forage available during the summer growing season varied predictably between years, elk were more likely to migrate regardless of whether they had access to irrigated agriculture. Our study shows that predictable availability of beneficial native forage can encourage migration even for ungulates with irrigated agriculture on their winter range. Perturbations that can affect the forage available to ungulates include wildfires, timber harvest, livestock grazing and changing weather patterns. If these or other disturbances negatively affect forage on summer ranges of migrants, or if they cause forage to vary unpredictably across space and time, our results suggest migratory behaviour may decline as a result.

EFFECTS OF BRUCELLOSIS SEROLOGIC STATUS ON PHYSIOLOGY AND BEHAVIOR OF ROCKY MOUNTAIN ELK ( CERVUS CANADENSIS NELSONI) IN SOUTHWESTERN MONTANA, USA.

Brucellosis, caused by bacteria in the genus Brucella, is an infectious zoonosis affecting animals and humans worldwide. Free-ranging Rocky Mountain elk ( Cervus canadensis nelsoni) and bison ( Bison bison) in the Greater Yellowstone Ecosystem (areas of southwestern Montana, eastern Idaho, and northwestern Wyoming, US) are the self-sustaining reservoirs of bovine brucellosis ( Brucella abortus) and elk are considered the primary source of livestock infections. It has been hypothesized that Brucella-exposed elk might have different physiologic status (pregnancy rates and body condition) and migration behaviors than would healthy elk. Here we tested the effects of brucellosis serologic status on pregnancy rates and winter ingesta free body fat of 100 female elk in southwestern Montana. We also evaluated the effects of serologic status on two characteristics of spring migration behavior, migration types (migrant, mixed migrant, resident, disperser, nomad, and undetermined type) and timing (start and end dates and duration). The migration behaviors were quantified using a model-driven approach based on the relative net squared displacement. We detected a significant difference (P=0.003) in pregnancy rates between seropositive and seronegative elk, with about a 30% drop in seropositive individuals. However, we did not detect differences in body fat between seropositive and seronegative elk or differences in either migration type or timing of spring migration. These results confirmed that the major effect of brucellosis in free-ranging elk is associated with reproduction.

Respiratory pathogens and their association with population performance in Montana and Wyoming bighorn sheep populations.

Respiratory disease caused by Mycoplasma ovipneumoniae and Pasteurellaceae poses a formidable challenge for bighorn sheep (Ovis canadensis) conservation. All-age epizootics can cause 10-90% mortality and are typically followed by multiple years of enzootic disease in lambs that hinders post-epizootic recovery of populations. The relative frequencies at which these epizootics are caused by the introduction of novel pathogens or expression of historic pathogens that have become resident in the populations is unknown. Our primary objectives were to determine how commonly the pathogens associated with respiratory disease are hosted by bighorn sheep populations and assess demographic characteristics of populations with respect to the presence of different pathogens. We sampled 22 bighorn sheep populations across Montana and Wyoming, USA for Mycoplasma ovipneumoniae and Pasteurellaceae and used data from management agencies to characterize the disease history and demographics of these populations. We tested for associations between lamb:ewe ratios and the presence of different respiratory pathogen species. All study populations hosted Pasteurellaceae and 17 (77%) hosted Mycoplasma ovipneumoniae. Average lamb:ewe ratios for individual populations where both Mycoplasma ovipneumoniae and Pasteurellaceae were detected ranged from 0.14 to 0.40. However, average lamb:ewe ratios were higher in populations where Mycoplasma ovipneumoniae was not detected (0.37, 95% CI: 0.27-0.51) than in populations where it was detected (0.25, 95% CI: 0.21-0.30). These findings suggest that respiratory pathogens are commonly hosted by bighorn sheep populations and often reduce recruitment rates; however ecological factors may interact with the pathogens to determine population-level effects. Elucidation of such factors could provide insights for management approaches that alleviate the effects of respiratory pathogens in bighorn sheep. Nevertheless, minimizing the introduction of novel pathogens from domestic sheep and goats remains imperative to bighorn sheep conservation.

Assessing respiratory pathogen communities in bighorn sheep populations: Sampling realities, challenges, and improvements.

Respiratory disease has been a persistent problem for the recovery of bighorn sheep (Ovis canadensis), but has uncertain etiology. The disease has been attributed to several bacterial pathogens including Mycoplasma ovipneumoniae and Pasteurellaceae pathogens belonging to the Mannheimia, Bibersteinia, and Pasteurella genera. We estimated detection probability for these pathogens using protocols with diagnostic tests offered by a fee-for-service laboratory and not offered by a fee-for-service laboratory. We conducted 2861 diagnostic tests on swab samples collected from 476 bighorn sheep captured across Montana and Wyoming to gain inferences regarding detection probability, pathogen prevalence, and the power of different sampling methodologies to detect pathogens in bighorn sheep populations. Estimated detection probability using fee-for-service protocols was less than 0.50 for all Pasteurellaceae and 0.73 for Mycoplasma ovipneumoniae. Non-fee-for-service Pasteurellaceae protocols had higher detection probabilities, but no single protocol increased detection probability of all Pasteurellaceae pathogens to greater than 0.50. At least one protocol resulted in an estimated detection probability of 0.80 for each pathogen except Mannheimia haemolytica, for which the highest detection probability was 0.45. In general, the power to detect Pasteurellaceae pathogens at low prevalence in populations was low unless many animals were sampled or replicate samples were collected per animal. Imperfect detection also resulted in low precision when estimating prevalence for any pathogen. Low and variable detection probabilities for respiratory pathogens using live-sampling protocols may lead to inaccurate conclusions regarding pathogen community dynamics and causes of bighorn sheep respiratory disease epizootics. We recommend that agencies collect multiples samples per animal for Pasteurellaceae detection, and one sample for Mycoplasma ovipneumoniae detection from at least 30 individuals to reliably detect both Pasteurellaceae and Mycoplasma ovipneumoniae at the population-level. Availability of PCR diagnostic tests to wildlife management agencies would improve the ability to reliably detect Pasteurellaceae in bighorn sheep populations.

Assessing the importance of demographic parameters for population dynamics using Bayesian integrated population modeling.

To successfully respond to changing habitat, climate or harvest, managers need to identify the most effective strategies to reverse population trends of declining species and/or manage harvest of game species. A classic approach in conservation biology for the last two decades has been the use of matrix population models to determine the most important vital rates affecting population growth rate (λ), that is, sensitivity. Ecologists quickly realized the critical role of environmental variability in vital rates affecting λ by developing approaches such as life-stage simulation analysis (LSA) that account for both sensitivity and variability of a vital rate. These LSA methods used matrix-population modeling and Monte Carlo simulation methods, but faced challenges in integrating data from different sources, disentangling process and sampling variation, and in their flexibility. Here, we developed a Bayesian integrated population model (IPM) for two populations of a large herbivore, elk (Cervus canadensis) in Montana, USA. We then extended the IPM to evaluate sensitivity in a Bayesian framework. We integrated known-fate survival data from radio-marked adults and juveniles, fecundity data, and population counts in a hierarchical population model that explicitly accounted for process and sampling variance. Next, we tested the prevailing paradigm in large herbivore population ecology that juvenile survival of neonates <90 d old drives λ using our Bayesian LSA approach. In contrast to the prevailing paradigm in large herbivore ecology, we found that adult female survival explained more of the variation in λ than elk calf survival, and that summer and winter elk calf survival periods were nearly equivalent in importance for λ. Our Bayesian IPM improved precision of our vital rate estimates and highlighted discrepancies between count and vital rate data that could refine population monitoring, demonstrating that combining sensitivity analysis with population modeling in a Bayesian framework can provide multiple advantages. Our Bayesian LSA framework will provide a useful approach to addressing conservation challenges across a variety of species and data types.

Linking landscape-scale differences in forage to ungulate nutritional ecology.

Understanding how habitat and nutritional condition affect ungulate populations is necessary for informing management, particularly in areas experiencing carnivore recovery and declining ungulate population trends. Variations in forage species availability, plant phenological stage, and the abundance of forage make it challenging to understand landscape-level effects of nutrition on ungulates. We developed an integrated spatial modeling approach to estimate landscape-level elk (Cervus elaphus) nutritional resources in two adjacent study areas that differed in coarse measures of habitat quality and related the consequences of differences in nutritional resources to elk body condition and pregnancy rates. We found no support for differences in dry matter digestibility between plant samples or in phenological stage based on ground sampling plots in the two study areas. Our index of nutritional resources, measured as digestible forage biomass, varied among land cover types and between study areas. We found that altered plant composition following fires was the biggest driver of differences in nutritional resources, suggesting that maintaining a mosaic of fire history and distribution will likely benefit ungulate populations. Study area, lactation status, and year affected fall body fat of adult female elk. Elk in the study area exposed to lower summer range nutritional resources had lower nutritional condition entering winter. These differences in nutritional condition resulted in differences in pregnancy rate, with average pregnancy rates of 89% for elk exposed to higher nutritional resources and 72% for elk exposed to lower nutritional resources. Summer range nutritional resources have the potential to limit elk pregnancy rate and calf production, and these nutritional limitations may predispose elk to be more sensitive to the effects of harvest or predation. Wildlife managers should identify ungulate populations that are nutritionally limited and recognize that these populations may be more impacted by recovering carnivores or harvest than populations inhabiting more productive summer habitats.

Elk Resource Selection and Implications for Anthrax Management in Montana.

Anthrax, caused by the spore-forming bacterium Bacillus anthracis, is a zoonotic disease that affects humans and animals throughout the world. In North America, anthrax outbreaks occur in livestock and wildlife species. Vaccine administration in wildlife is untenable; the most effective form of management is surveillance and decontamination of carcasses. Successful management is critical because untreated carcasses can create infectious zones increasing risk for other susceptible hosts. We studied the bacterium in a re-emerging anthrax zone in southwest Montana. In 2008, a large anthraxepizootic primarily affected a domestic bison (Bison bison) herd and the male segment of a free-ranging elk (Cervus elaphus) herd in southwestern Montana. Following the outbreak, we initiated a telemetry study on elk to evaluate resource selection during the anthrax season to assist with anthrax management. We used a mixed effects generalized linear model (GLM) to estimate resource selection by male elk, and we mapped habitat preferences across the landscape. We overlaid preferred habitats on ecological niche model-based estimates of B. anthracis presence. We observed significant overlap between areas with a high predicted probability of male elk selection and B. anthracis potential. These potentially risky areas of elk and B. anthracis overlap were broadly spread over public and private lands. Future outbreaks in the region are probable, and this analysis identified the spatial extent of the risk area in the region, which can be used to prioritize anthrax surveillance.

Mapping Resource Selection Functions in Wildlife Studies: Concerns and Recommendations.

Predicting the spatial distribution of animals is an important and widely used tool with applications in wildlife management, conservation, and population health. Wildlife telemetry technology coupled with the availability of spatial data and GIS software have facilitated advancements in species distribution modeling. There are also challenges related to these advancements including the accurate and appropriate implementation of species distribution modeling methodology. Resource Selection Function (RSF) modeling is a commonly used approach for understanding species distributions and habitat usage, and mapping the RSF results can enhance study findings and make them more accessible to researchers and wildlife managers. Currently, there is no consensus in the literature on the most appropriate method for mapping RSF results, methods are frequently not described, and mapping approaches are not always related to accuracy metrics. We conducted a systematic review of the RSF literature to summarize the methods used to map RSF outputs, discuss the relationship between mapping approaches and accuracy metrics, performed a case study on the implications of employing different mapping methods, and provide recommendations as to appropriate mapping techniques for RSF studies. We found extensive variability in methodology for mapping RSF results. Our case study revealed that the most commonly used approaches for mapping RSF results led to notable differences in the visual interpretation of RSF results, and there is a concerning disconnect between accuracy metrics and mapping methods. We make 5 recommendations for researchers mapping the results of RSF studies, which are focused on carefully selecting and describing the method used to map RSF studies, and relating mapping approaches to accuracy metrics.

Grizzly bear predation links the loss of native trout to the demography of migratory elk in Yellowstone.

The loss of aquatic subsidies such as spawning salmonids is known to threaten a number of terrestrial predators, but the effects on alternative prey species are poorly understood. At the heart of the Greater Yellowstone ecosystem, an invasion of lake trout has driven a dramatic decline of native cutthroat trout that migrate up the shallow tributaries of Yellowstone Lake to spawn each spring. We explore whether this decline has amplified the effect of a generalist consumer, the grizzly bear, on populations of migratory elk that summer inside Yellowstone National Park (YNP). Recent studies of bear diets and elk populations indicate that the decline in cutthroat trout has contributed to increased predation by grizzly bears on the calves of migratory elk. Additionally, a demographic model that incorporates the increase in predation suggests that the magnitude of this diet shift has been sufficient to reduce elk calf recruitment (4-16%) and population growth (2-11%). The disruption of this aquatic-terrestrial linkage could permanently alter native species interactions in YNP. Although many recent ecological changes in YNP have been attributed to the recovery of large carnivores--particularly wolves--our work highlights a growing role of human impacts on the foraging behaviour of grizzly bears.

An evaluation of density-dependent and density-independent influences on population growth rates in Weddell seals.

Much of the existing literature that evaluates the roles of density-dependent and density-independent factors on population dynamics has been called into question in recent years because measurement errors were not properly dealt with in analyses. Using state-space models to account for measurement errors, we evaluated a set of competing models for a 22-year time series of mark-resight estimates of abundance for a breeding population of female Weddell seals (Leptonychotes weddellii) studied in Erebus Bay, Antarctica. We tested for evidence of direct density dependence in growth rates and evaluated whether equilibrium population size was related to seasonal sea-ice extent and the Southern Oscillation Index (SOI). We found strong evidence of negative density dependence in annual growth rates for a population whose estimated size ranged from 438 to 623 females during the study. Based on Bayes factors, a density-dependence-only model was favored over models that also included environmental covariates. According to the favored model, the population had a stationary distribution with a mean of 497 females (SD = 60.5), an expected growth rate of 1.10 (95% credible interval = 1.08-1.15) when population size was 441 females, and a rate of 0.90 (95% credible interval = 0.87-.93) for a population of 553 females. A model including effects of SOI did receive some support and indicated a positive relationship between SOI and population size. However, effects of SOI were not large, and including the effect did not greatly reduce our estimate of process variation. We speculate that direct density dependence occurred because rates of adult survival, breeding, and temporary emigration were affected by limitations on per capita food resources and space for parturition and pup-rearing. To improve understanding of the relative roles of various demographic components and their associated vital rates to population growth rate, mark-recapture methods can be applied that incorporate both environmental covariates and the seal abundance estimates that were developed here. An improved understanding of why vital rates change with changing population abundance will only come as we develop a better understanding of the processes affecting marine food resources in the Southern Ocean.