Kelly M. Proffitt

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

Abstract Prey behavioral responses to predation risk in wolf–ungulate–plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry-collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf- and human-predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf- and human-predation risk were similar; however, responses to human-predation risk were stronger than responses to wolf-predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public-land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.

Migration of northern Yellowstone elk: implications of spatial structuring

Abstract Migration can enhance survival and recruitment of mammals by increasing access to higher-quality forage or reducing predation risk, or both. We used telemetry locations collected from 140 adult female elk during 2000–2003 and 2007–2008 to identify factors influencing the migration of northern Yellowstone elk. Elk wintered in 2 semidistinct herd segments and migrated 10–140 km to at least 12 summer areas in Yellowstone National Park (YNP) and nearby areas of Montana. Spring migrations were delayed after winters with increased snow pack, with earlier migration in years with earlier vegetation green-up. Elk wintering at lower elevations outside YNP migrated an average of 13 days earlier than elk at higher elevations. The timing of autumn migrations varied annually, but elk left their summer ranges at about the same time regardless of elevation, wolf numbers, or distance to their wintering areas. Elk monitored for multiple years typically returned to the same summer (96% fidelity, n  =  52) and winter (61% fidelity, n  =  41) ranges. Elk that wintered at lower elevations in or near the northwestern portion of the park tended to summer in the western part of YNP (56%), and elk that wintered at higher elevations spent summer primarily in the eastern and northern parts of the park (82%). Elk did not grossly modify their migration timing, routes, or use areas after wolf restoration. Elk mortality was low during summer and migration (8 of 225 elk-summers). However, spatial segregation and differential mortality and recruitment between herd segments on the northern winter range apparently contributed to a higher proportion of the elk population wintering outside the northwestern portion of YNP and summering in the western portion of the park. This change could shift wolf spatial dynamics more outside YNP and increase the risk of transmission of brucellosis from elk to cattle north of the park.

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.

Site fidelity of Weddell seals: the effects of sex and age

Abstract Site fidelity is believed to be an important life history strategy for Weddell seals (Leptonychotes weddellii), that return to traditional breeding colonies each spring. We examined four hypotheses concerning their fidelity to these colonies: 1) fidelity is stronger to natal sites (natal fidelity) than to other sites, 2) females exhibit greater site fidelity than males, 3) site fidelity for both sexes increases with age, 4) site fidelity in adult females is related to their reproductive status and their total number of offspring. Analysis of a long-term tagging database from McMurdo Sound did not support hypotheses 1 and 2. Although animals did express fidelity to specific sites over their lifetime (χ2 tests, P < 0.05), fidelity to natal colonies was lower than to other sites (χ2 test, P < 0.05). There were no differences in site fidelity between males and females (χ2 tests, P > 0.05). Hypothesis 3 was supported. Since the probability of a returning seal occupying the same colony as the previous year increased with age among both sexes to about age 12. Finally, in support of hypothesis 4, females with a higher degree of site fidelity were more likely to both have a higher reproductive rate and return to a site where they have previously given birth.

Relative influence of human harvest, carnivores, and weather on adult female elk survival across western North America

Summary 1. Well-informed management of harvested species requires understanding how changing ecological conditions affect demography and population dynamics, information that is lacking for many species. We have limited understanding of the relative influence of carnivores, harvest, weather and forage availability on elk Cervus elaphus demography, despite the ecological and economic importance of this species. We assessed adult female survival, a key vital rate for population dynamics, from 2746 radio-collared elk in 45 populations across western North America that experience wide variation in carnivore assemblage, harvest, weather and habitat conditions. 2. Proportional hazard analysis revealed that ‘baseline’ (i.e. not related to human factors) mortality was higher with very high winter precipitation, particularly in populations sympatric with wolves Canis lupus. Mortality may increase via nutritional stress and heightened vulnerability to predation in snowy winters. Baseline mortality was unrelated to puma Puma concolor presence, forest cover or summer forage productivity. 3. Cause-specific mortality analyses showed that wolves and all carnivore species combined had additive effects on baseline elk mortality, but only reduced survival by <2%. When human factors were included, ‘total’ adult mortality was solely related to harvest; the influence of native carnivores was compensatory. Annual total mortality rates were lowest in populations sympatric with both pumas and wolves because managers reduced female harvest in areas with abundant or diverse carnivores.

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.

Changes in Elk Resource Selection and Distributions Associated With a Late-Season Elk Hunt

Abstract Changes in resource selection associated with human predation risk may alter elk distributions and availability for harvest. We used Global Positioning System data collected from telemetered female elk (Cervus elaphus) to evaluate effects of refuges (areas where hunting was prohibited), spatial variation in hunting risk, and landscape attributes on resource selection within an established Greater Yellowstone Area, USA, winter range. We also evaluated elk distributions during and outside of a late-season hunting period. Refuge areas and landscape attributes such as habitat type and snow water equivalents (SWE) affected resource selection. Elk selection for flat grasslands increased as SWE increased, likely because these areas were windswept, leaving grasses exposed for foraging. Elk distributions differed during hunting and no-hunting periods. During the hunting period, elk shifted to privately owned refuge areas and the estimated odds of elk occupying refuge areas more than doubled. Risk-driven changes in resource selection resulted in reduced availability of elk for harvest. Elk selection for areas where hunting is prohibited presents a challenge for resource managers that use hunting as a tool for managing populations and influences grazing patterns on private ranchlands.

Exploring Linkages between Abiotic Oceanographic Processes and a Top-trophic Predator in an Antarctic Ecosystem

Climatic variation affects the physical and biological components of ecosystems, and global-climate models predict enhanced sensitivity in polar regions, raising concern for Antarctic animal populations that may show direct responses to changes in sea-ice distribution and extent, or indirect responses to changes in prey distribution and abundance. Here, we show that over a 30-year period in the Ross Sea, average weaning masses of Weddell seals, Leptonychotes weddellii, varied strongly among years and were correlated to large-scale climatic and oceanographic variations. Foraging success of pregnant seals (reflected by weaning mass the following pupping season) increased during summers characterized by reduced sea-ice cover and positive phases of the southern oscillation. These results demonstrate a correlation between environmental variation and an important life history characteristic (weaning mass) of an Antarctic marine mammal. Understanding the mechanisms that link climatic variation and animal life history characteristics will contribute to understanding both population dynamics and global climatic processes. For the world’s most southerly distributed mammal species, the projected trend of increasing global climate change raises concern because increasing sea-ice trends in the Ross Sea sector of Antarctica will likely reduce populations due to reduced access to prey as expressed through declines in body condition and reproductive performance.

Integrating resource selection into spatial capture‐recapture models for large carnivores

Wildlife managers need reliable methods to estimate large carnivore densities and population trends; yet large carnivores are elusive, difficult to detect, and occur at low densities making traditional approaches intractable. Recent advances in spatial capture-recapture (SCR) models have provided new approaches for monitoring trends in wildlife abundance and these methods are particularly applicable to large carnivores. We applied SCR models in a Bayesian framework to estimate mountain lion densities in the Bitterroot Mountains of west central Montana. We incorporate an existing resource selection function (RSF) as a density covariate to account for heterogeneity in habitat use across the study area and include data collected from harvested lions. We identify individuals through DNA samples collected by (1) biopsy darting mountain lions detected in systematic surveys of the study area, (2) opportunistically collecting hair and scat samples, and (3) sampling all harvested mountain lions. We included 80 DNA...

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.