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Geology topics

Mark A. Haroldson

Publications and source records attributed to Mark A. Haroldson.

At least 37 records · Page 2Linked to original sources

Agent-based models for collective animal movement: Proximity-induced state switching

Animal movement is a complex phenomenon where individual movement patterns can be influenced by a variety of factors including the animal’s current activity, available terrain and habitat, and locations of other animals. Motivated by modeling grizzly bear movement in the Greater Yellowstone Ecosystem, this article presents an agent-based model represented in a state-space framework for collective animal movement. The novel contribution of this work is a collective animal movement model that captures interactions between animals that can trigger changes in movement patterns, such as when a dominant grizzly bear may cause another subordinate bear to temporarily leave an area. The modeling framework enables learning different movement patterns through a state-space representation with particle-MCMC methods for fully Bayesian model fitting and the prediction of future animal movement behaviors.

Journal of Agricultural, Biological and Environmen

Physiological consequences of consuming low-energy foods: Herbivory coincides with a stress response in Yellowstone bears.

Meat, fruit, seeds and other high-energy bear foods are often highly localized and briefly available and understanding which factors influence bear consumption of these foods is a common focus of bear conservation and ecology. However, the most common bear foods, graminoids and forbs, are more widespread but of lower quality. We poorly understand how herbage consumption impacts bear physiology, such as endocrine system function that regulates homeostasis and stress responses. Here, we described bear diets with a novel approach, measuring the concentration of chlorophyll in bear scats (faecal chlorophyll) to index the proportion of the recent diet that was composed of leaves from graminoids and forbs. We measured faecal chlorophyll and faecal cortisol in 351 grizzly ( Ursus arctos , n = 255) and black bear ( Ursus americanus , n = 96) scats from Yellowstone National Park in 2008–2009. We compared models of faecal chlorophyll and faecal cortisol concentrations considering the effects of spatial, dietary, scat and bear-specific factors including species. Faecal chlorophyll levels were the strongest predictor of faecal cortisol in a manner that suggested an endocrine response to a low-energy diet. Both compounds were highest during the spring and early summer months, overlapping the breeding season when higher energy foods were less available. Effects of scat composition, scat weathering, bear age, bear sex, species and other factors that have previously been shown to influence faecal cortisol in bears were not important unless faecal chlorophyll was excluded from models. The top models of faecal chlorophyll suggested grazing was primarily influenced by spatial attributes, with greater grazing closer to recreational trails, implying that elevated cortisol with grazing could be a response to anthropogenic activity. Our results confirm that higher stress hormone concentrations correspond with lower quality diets in bears, particularly grazing, and that faecal chlorophyll shows promise as a metric for studying grazing behaviour and its consequences.

Wyoming

Quantifying energetic costs and defining energy landscapes experienced by grizzly bears

Animal movements are major determinants of energy expenditure and ultimately the cost–benefit of landscape use. Thus, we sought to understand those costs and how grizzly bears ( Ursus arctos ) move in mountainous landscapes. We trained captive grizzly bears to walk on a horizontal treadmill and up and down 10% and 20% slopes. The cost of moving upslope increased linearly with speed and slope angle, and this was more costly than moving horizontally. The cost of downslope travel at slower speeds was greater than the cost of traveling horizontally but appeared to decrease at higher speeds. The most efficient walking speed that minimized cost per unit distance was 1.19±0.11 m s −1 . However, grizzly bears fitted with GPS collars in the Greater Yellowstone Ecosystem moved at an average velocity of 0.61±0.28 m s −1 and preferred to travel on near-horizontal slopes at twice their occurrence. When traveling uphill or downhill, grizzly bears chose paths across all slopes that were ∼54% less steep and costly than the maximum available slope. The net costs (J kg −1 m −1 ) of moving horizontally and uphill were the same for grizzly bears, humans and digitigrade carnivores, but those costs were 46% higher than movement costs for ungulates. These movement costs and characteristics of landscape use determined using captive and wild grizzly bears were used to understand the strategies that grizzly bears use for preying on large ungulates and the similarities in travel between people and grizzly bears that might affect the risk of encountering each other on shared landscapes.

Wyoming

Thermal constraints on energy balance, behaviour and spatial distribution of grizzly bears

1. Heat dissipation limit theory posits that energy available for growth and reproduction in endotherms is limited by their ability to dissipate heat. In mammals, endogenous heat production increases markedly during gestation and lactation, and thus female mammals may be subject to greater thermal constraints on energy expenditure than males. Such constraints likely have important implications for behaviour and population performance in a warming climate. 2. We used a mechanistic simulation model based on the first principles of heat and mass transfer to study thermal constraints on activity (both timing and intensity) of captive female grizzly bears Ursus arctos in current and future climate scenarios. We then quantified the relative importance of regulatory behaviours for maintaining heat balance using GPS telemetry locations of lactating versus non-lactating female bears from Yellowstone National Park, and assessed the degree to which costs of thermoregulation constrained the distribution of sampled bears in space and time. 3. Lactating female bears benefitted considerably more from behavioural cooling mechanisms (e.g. partial submersion in cool water or bedding on cool substrate) than non-lactating females in our simulations; the availability of water for thermoregulation increased the number of hours during which lactating females could be active by up to 60% under current climatic conditions and by up to 43% in the future climate scenario. Moreover, even in the future climate scenario, lactating bears were able to achieve heat balance 24 hr/day by thermoregulating behaviourally when water was available to facilitate cooling. 4. The most important predictor of female grizzly bear distribution in Yellowstone, regardless of reproductive status, was elevation. However, variables associated with the thermal environment were relatively more important for predicting the distribution of lactating than non-lactating female bears. 5. Our results suggest that the costs of heat dissipation, which are modulated by climate, may impose constraints on the behaviour and energetics of large endotherms like grizzly bears, and that access to water for cooling will likely be an increasingly important driver of grizzly bear distribution in Yellowstone as the climate continues to warm.

Wyoming

A reassessment of Chao2 estimates for population monitoring of grizzly bears in the Greater Yellowstone Ecosystem

The Yellowstone Ecosystem Subcommittee (YES) asked the Interagency Grizzly Bear Study Team (IGBST) to re-assess a technique used in annual population estimation and trend monitoring of grizzly bears in the Greater Yellowstone Ecosystem (GYE). This technique is referred to as the Chao2 approach and estimates the number of females with cubs-of-the-year (hereafter, females with cubs) and, in association with other demographic data, is used by the IGBST to produce annual population estimates. Females with cubs are an easily recognizable population segment, and trends for this reproductive segment of the population are assumed to be representative of trend for the entire population. The overarching objective of the analyses presented in this report was to provide a more accurate representation of the GYE grizzly bear population using the current methodologies in place. Specifically, we addressed two limitations of the current Chao2 approach: 1) underestimation bias associated with a distance criterion used to differentiate annual sightings of females with cubs into unique individuals and 2) limitations of the model-averaging approach to effectively distinguish among potential future population trajectories (decline, stability, and growth). The first issue addressed in this report is the underestimation bias associated with the rule set that Knight et al. (1995) developed to differentiate sightings of females with cubs into unique individuals (i.e., unique family groups). The rule set was originally designed to be conservative by reducing the risk of identifying more females with cubs than actually existed, primarily through use of a distance criterion of 30 km to separate sightings of unique females. This approach resulted in an underestimation bias, and previous research demonstrated that this bias increases with increasing number of females with cubs. Using location data from radio-marked females with cubs, we evaluated alternative distance criteria by simulating scenarios with varying numbers of true females with cubs and sightings. Findings from these analyses demonstrate that bias in estimates of females with cubs can be substantially reduced by changing the 30-km distance criterion in the rule set to 16 km, which produced relatively unbiased estimates. Findings also indicate, however, the importance of adaptability with regard to the distance criteria because of the complex relationships and biases among the various parameters involved in estimation of unique females with cubs. The total number of annual sightings and the true number of females with cubs play particularly important roles. Whereas these analyses remind us that there is no perfect approach to estimating the number of females with cubs from sightings under various scenarios, they provide us with new tools to determine when and how to adapt the monitoring program. The second issue we were tasked to investigate was the potential for improvement of the technique referred to as model-averaging, which serves to smooth relatively high variation in annual estimates. This technique was chosen by YES as the basis for monitoring the Yellowstone grizzly bear population, as described in the 2016 Conservation Strategy. This choice was made in part because the technique has been well documented and population estimates derived from counts of females with cubs are conservative. Using simulations of population trends, we demonstrate why the model-averaging technique currently used cannot distinguish between plausible future trend scenarios. As a suitable alternative to model averaging, we propose the use of generalized additive models (GAMs). Using a suite of simulated trend dynamics relevant to management, we demonstrate GAM performance for tracking trends in females with cubs within the context of the annual monitoring program. We demonstrate the ability to not only document directional changes in population trend but also patterns of stabilization or resiliency after such changes. Furthermore, the proposed monitoring framework provides objective measures useful for early detection of directional changes in trend. The new framework is flexible, allowing retrospective analysis of Chao2-based estimates and future applications to time series of other population metrics, such as vital rates. The aforementioned updates provide us with new tools to determine when and how to adapt the monitoring program. Within the context of current monitoring protocols and effort, and considering the full suite of simulations presented in this report and previous studies, the IGBST plans to incorporate the following changes to the population monitoring protocol: 1) modify the distance criterion, starting with 16 km under current sampling conditions and 2) revise the population monitoring framework using GAMs as the basis for smoothing of annual estimates and detecting trends and changes in trend. Implementation of the 16-km distance criterion combined with use of GAM techniques would affect some of the population metrics (e.g., annual population size and uncertainty, population trend, mortality rates) used to inform management responses. A primary consideration is that the 16-km distance criterion results in total population estimates derived from the Chao2 estimates that are greater than those we have reported in the past. This increase is due to a change in the implementation of the technique and more accurately represents the number of females with cubs in the GYE grizzly bear population. Additionally, interpretation of retrospective trend patterns may change due to the combination of a different distance criterion and enhanced trend monitoring based on the GAM approach we present here. Implementation will require relatively minor changes in the monitoring protocols described in Appendices B and C of the 2016 Conservation Strategy. Finally, we note that the IGBST has ongoing investigations into the merits of an Integrated Population Model (IPM), for which annual Chao2-based estimates are important input data. The IGBST plans to continue those investigations using the 16-km distance criterion to derive Chao2 estimates.

Idaho, Montana, Wyoming

Brown Bear (Ursus arctos; North America)

This chapter comprises the following sections: names, taxonomy, subspecies and distribution, descriptive notes, habitat, movements and home range, activity patterns, feeding ecology, reproduction and growth, behavior, parasites and diseases, status in the wild, and status in captivity.

Book chapter

Potential for recreational restrictions to reduce grizzly bear–caused human injuries

In 2011, 2 hikers were killed by grizzly bears (Ursus arctos) in separate incidents on backcountry trails in Hayden Valley, Yellowstone National Park, USA (YNP). Hayden Valley provides prime habitat for grizzly bears and is known to have high densities of bears. During 1970–2017, 23% (10 of 44) of all backcountry grizzly bear–inflicted human injuries and fatalities in YNP occurred in the valley even though it comprises only 1% of the park. In addition, 3 of the last 5 fatal bear attacks in the park occurred in the valley. We evaluated retrospectively whether restrictions and closures on visitor recreational activity would have prevented many of these injuries. We considered prohibitions on recreational activity during seasons when bears forage for specific high-quality foods; potential closures that coincided with the times of day and year bears were most active in the valley; and visitor use restrictions that would have prevented the most common human behaviors associated with grizzly bear–caused human injuries. The food-based closure that may have prevented the most human injuries occurred during middle to late summer when bears scavenge bison (Bison bison) carcasses that result from annual rutting behavior of bison in the valley. However, safety precautions such as hiking in groups of >=3, remaining on maintained trails, and carrying bear spray would likely reduce the frequency of bear-inflicted human injuries more than most food-based seasonal closures. Our analyses provide broadly applicable findings regarding use of visitor behavior restrictions and seasonal closures to reduce the risk of bear-inflicted human injuries.

Wyoming

Primarily resident grizzly bears respond to late-season elk harvest

Autumn ungulate hunting in the Greater Yellowstone Ecosystem carries the risk of hunter–grizzly bear ( Ursus arctos ) conflict and creates a substantial challenge for managers. For Grand Teton National Park, Wyoming, USA, a key information need is whether increased availability of elk ( Cervus canadensis ) carcasses during a late autumn (Nov–Dec) harvest within the national park attracts grizzly bears and increases the potential for conflict with hunters. Using a robust design analysis with 6 primary sampling periods during 2014–2015, we tested the hypothesis that the elk harvest resulted in temporary movements of grizzly bears into the hunt areas, thus increasing bear numbers. We detected 31 unique individuals (6 F, 25 M) through genetic sampling and retained 26 encounter histories for analysis. Markovian movement models had more support than a null model of no temporary movement. Contrary to our research hypothesis, temporary movements into the study area occurred between the July–August (no hunt; N̄ 2014–2015 = 5) and September–October (no hunt; N̄ 2014–2015 = 24) primary periods each year, rather than during the transition from September–October (no hunt) to November–December (hunt; N̄ 2014–2015 = 15). A post hoc analysis indicated that September–October population estimates were biased high by detections of transient bears. Grizzly bear presence during the elk hunt was limited to approximately 15 resident bears that specialized in accessing elk carcasses. The late timing of the elk hunt likely moderated the effect of carcasses as a food attractant because it coincides with the onset of hibernation. From a population response perspective, the current timing of the elk harvest likely represents a scenario of low relative risk of hunter–bear conflicts. The risk of hunter–grizzly bear encounters remains, but may be more a function of factors that operate at the level of individual bears and hunters, such as hunter movements and bear responses to olfactory cues.

Wyoming

Estimating distemper virus dynamics among wolves and grizzly bears using serology and Bayesian state‐space models

Many parasites infect multiple hosts, but estimating the transmission across host species remains a key challenge in disease ecology. We investigated the within and across host species dynamics of canine distemper virus (CDV) in grizzly bears ( Ursus arctos ) and wolves ( Canis lupus ) of the Greater Yellowstone Ecosystem (GYE). We hypothesized that grizzly bears may be more likely to be exposed to CDV during outbreaks in the wolf population because grizzly bears often displace wolves while scavenging carcasses. We used serological data collected from 1984 to 2014 in conjunction with Bayesian state‐space models to infer the temporal dynamics of CDV. These models accounted for the unknown timing of pathogen exposure, and we assessed how different testing thresholds and the potential for testing errors affected our conclusions. We identified three main CDV outbreaks (1999, 2005, and 2008) in wolves, which were more obvious when we used higher diagnostic thresholds to qualify as seropositive. There was some evidence for increased exposure rates in grizzly bears in 2005, but the magnitude of the wolf effect on bear exposures was poorly estimated and depended upon our prior distributions. Grizzly bears were exposed to CDV prior to wolf reintroduction and during time periods outside of known wolf outbreaks, thus wolves are only one of several potential routes for grizzly bear exposures. Our modeling approach accounts for several of the shortcomings of serological data and is applicable to many wildlife disease systems, but is most informative when testing intervals are short. CDV circulates in a wide range of carnivore species, but it remains unclear whether the disease persists locally within the GYE carnivore community or is periodically reintroduced from distant regions with larger host populations.

Ecology and Evolution

Potential paths for male-mediated gene flow to and from an isolated grizzly bear population

For several decades, grizzly bear populations in the Greater Yellowstone Ecosystem (GYE) and the Northern Continental Divide Ecosystem (NCDE) have increased in numbers and range extent. The GYE population remains isolated and although effective population size has increased since the early 1980s, genetic connectivity between these populations remains a long-term management goal. With only ~110 km distance separating current estimates of occupied range for these populations, the potential for gene flow is likely greater now than it has been for many decades. We sought to delineate potential paths that would provide the opportunity for male-mediated gene flow between the two populations. We first developed step-selection functions to generate conductance layers using ecological, physical, and anthropogenic landscape features associated with non-stationary GPS locations of 124 male grizzly bears (199 bear-years). We then used a randomized shortest path (RSP) algorithm to estimate the average number of net passages for all grid cells in the study region, when moving from an origin to a destination node. Given habitat characteristics that were the basis for the conductance layer, movements follow certain grid cell sequences more than others and the resulting RSP values thus provide a measure of movement potential. Repeating this process for 100 pairs of random origin and destination nodes, we identified paths for three levels of random deviation (θ) from the least-cost path. We observed broad-scale concordance between model predictions for paths originating in the NCDE and those originating in the GYE for all three levels of movement exploration. Model predictions indicated that male grizzly bear movement between the ecosystems could involve a variety of routes, and verified observations of grizzly bears outside occupied range supported this finding. Where landscape features concentrated paths into corridors (e.g., because of anthropogenic influence), they typically followed neighboring mountain ranges, of which several could serve as pivotal stepping stones. The RSP layers provide detailed, spatially explicit information for land managers and organizations working with land owners to identify and prioritize conservation measures that maintain or enhance the integrity of potential areas conducive to male grizzly bear dispersal.

Montana

Diet and macronutrient optimization in wild ursids: A comparison of grizzly bears with sympatric and allopatric black bears

When fed ad libitum, ursids can maximize mass gain by selecting mixed diets wherein protein provides 17 ± 4% of digestible energy, relative to carbohydrates or lipids. In the wild, this ability is likely constrained by seasonal food availability, limits of intake rate as body size increases, and competition. By visiting locations of 37 individuals during 274 bear-days, we documented foods consumed by grizzly ( Ursus arctos ) and black bears ( Ursus americanus ) in Grand Teton National Park during 2004–2006. Based on published nutritional data, we estimated foods and macronutrients as percentages of daily energy intake. Using principal components and cluster analyses, we identified 14 daily diet types. Only 4 diets, accounting for 21% of days, provided protein levels within the optimal range. Nine diets (75% of days) led to over-consumption of protein, and 1 diet (3% of days) led to under-consumption. Highest protein levels were associated with animal matter (i.e., insects, vertebrates), which accounted for 46–47% of daily energy for both species. As predicted: 1) daily diets dominated by high-energy vertebrates were positively associated with grizzly bears and mean percent protein intake was positively associated with body mass; 2) diets dominated by low-protein fruits were positively associated with smaller-bodied black bears; and 3) mean protein was highest during spring, when high-energy plant foods were scarce, however it was also higher than optimal during summer and fall. Contrary to our prediction: 4) allopatric black bears did not exhibit food selection for high-energy foods similar to grizzly bears. Although optimal gain of body mass was typically constrained, bears usually opted for the energetically superior trade-off of consuming high-energy, high-protein foods. Given protein digestion efficiency similar to obligate carnivores, this choice likely supported mass gain, consistent with studies showing monthly increases in percent body fat among bears in this region.

Wyoming

Detecting grizzly bear use of ungulate carcasses using global positioning system telemetry and activity data

Global positioning system (GPS) wildlife collars have revolutionized wildlife research. Studies of predation by free-ranging carnivores have particularly benefited from the application of location clustering algorithms to determine when and where predation events occur. These studies have changed our understanding of large carnivore behavior, but the gains have concentrated on obligate carnivores. Facultative carnivores, such as grizzly/brown bears ( Ursus arctos ), exhibit a variety of behaviors that can lead to the formation of GPS clusters. We combined clustering techniques with field site investigations of grizzly bear GPS locations ( n = 732 site investigations; 2004–2011) to produce 174 GPS clusters where documented behavior was partitioned into five classes (large-biomass carcass, small-biomass carcass, old carcass, non-carcass activity, and resting). We used multinomial logistic regression to predict the probability of clusters belonging to each class. Two cross-validation methods—leaving out individual clusters, or leaving out individual bears—showed that correct prediction of bear visitation to large-biomass carcasses was 78–88%, whereas the false-positive rate was 18–24%. As a case study, we applied our predictive model to a GPS data set of 266 bear-years in the Greater Yellowstone Ecosystem (2002–2011) and examined trends in carcass visitation during fall hyperphagia (September–October). We identified 1997 spatial GPS clusters, of which 347 were predicted to be large-biomass carcasses. We used the clustered data to develop a carcass visitation index, which varied annually, but more than doubled during the study period. Our study demonstrates the effectiveness and utility of identifying GPS clusters associated with carcass visitation by a facultative carnivore.

Greater Yellowstone Ecosystem

Multiple estimates of effective population size for monitoring a long-lived vertebrate: An application to Yellowstone grizzly bears

Effective population size ( N e ) is a key parameter for monitoring the genetic health of threatened populations because it reflects a population's evolutionary potential and risk of extinction due to genetic stochasticity. However, its application to wildlife monitoring has been limited because it is difficult to measure in natural populations. The isolated and well-studied population of grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem provides a rare opportunity to examine the usefulness of different N e estimators for monitoring. We genotyped 729 Yellowstone grizzly bears using 20 microsatellites and applied three single-sample estimators to examine contemporary trends in generation interval (GI), effective number of breeders ( N b ) and N e during 1982–2007. We also used multisample methods to estimate variance ( N eV ) and inbreeding N e ( N eI ). Single-sample estimates revealed positive trajectories, with over a fourfold increase in N e (≈100 to 450) and near doubling of the GI (≈8 to 14) from the 1980s to 2000s. N eV (240–319) and N eI (256) were comparable with the harmonic mean single-sample N e (213) over the time period. Reanalysing historical data, we found N eV increased from ≈80 in the 1910s–1960s to ≈280 in the contemporary population. The estimated ratio of effective to total census size ( N e /N c ) was stable and high (0.42–0.66) compared to previous brown bear studies. These results support independent demographic evidence for Yellowstone grizzly bear population growth since the 1980s. They further demonstrate how genetic monitoring of N e can complement demographic-based monitoring of N c and vital rates, providing a valuable tool for wildlife managers.

Idaho, Montana, Wyoming

Density dependence, whitebark pine, and vital rates of grizzly bears

Understanding factors influencing changes in population trajectory is important for effective wildlife management, particularly for populations of conservation concern. Annual population growth of the grizzly bear ( Ursus arctos ) population in the Greater Yellowstone Ecosystem, USA has slowed from 4.2–7.6% during 1983–2001 to 0.3–2.2% during 2002–2011. Substantial changes in availability of a key food source and bear population density have occurred. Whitebark pine ( Pinus albicaulis ), the seeds of which are a valuable but variable fall food for grizzly bears, has experienced substantial mortality primarily due to a mountain pine beetle ( Dendroctonus ponderosae ) outbreak that started in the early 2000s. Positive growth rates of grizzly bears have resulted in populations reaching high densities in some areas and have contributed to continued range expansion. We tested research hypotheses to examine if changes in vital rates detected during the past decade were more associated with whitebark pine decline or, alternatively, increasing grizzly bear density. We focused our assessment on known-fate data to estimate survival of cubs-of-the-year (cubs), yearlings, and independent bears (≥2 yrs), and reproductive transition of females from having no offspring to having cubs. We used spatially and temporally explicit indices for grizzly bear density and whitebark pine mortality as individual covariates. Models indicated moderate support for an increase in survival of independent male bears over 1983–2012, whereas independent female survival did not change. Cub survival, yearling survival, and reproductive transition from no offspring to cubs all changed during the 30-year study period, with lower rates evident during the last 10–15 years. Cub survival and reproductive transition were negatively associated with an index of grizzly bear density, indicating greater declines where bear densities were higher. Our analyses did not support a similar relationship for the index of whitebark pine mortality. The results of our study support the interpretation that slowing of population growth during the last decade was associated more with increasing grizzly bear density than the decline in whitebark pine. Grizzly bear density and its potential effect on vital rates and population trajectory warrant consideration for management of the grizzly bear population in the Greater Yellowstone Ecosystem.

Idaho, Montana, Wyoming

Influence of whitebark pine decline on fall habitat use and movements of grizzly bears in the Greater Yellowstone Ecosystem

When abundant, seeds of the high-elevation whitebark pine (WBP; Pinus albicaulis ) are an important fall food for grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem. Rates of bear mortality and bear/human conflicts have been inversely associated with WBP productivity. Recently, mountain pine beetles ( Dendroctonus ponderosae ) have killed many cone-producing WBP trees. We used fall (15 August–30 September) Global Positioning System locations from 89 bear years to investigate temporal changes in habitat use and movements during 2000–2011. We calculated Manly–Chesson (MC) indices for selectivity of WBP habitat and secure habitat (≥500 m from roads and human developments), determined dates of WBP use, and documented net daily movement distances and activity radii. To evaluate temporal trends, we used regression, model selection, and candidate model sets consisting of annual WBP production, sex, and year. One-third of sampled grizzly bears had fall ranges with little or no mapped WBP habitat. Most other bears (72%) had a MC index above 0.5, indicating selection for WBP habitats. From 2000 to 2011, mean MC index decreased and median date of WBP use shifted about 1 week later. We detected no trends in movement indices over time. Outside of national parks, there was no correlation between the MC indices for WBP habitat and secure habitat, and most bears (78%) selected for secure habitat. Nonetheless, mean MC index for secure habitat decreased over the study period during years of good WBP productivity. The wide diet breadth and foraging plasticity of grizzly bears likely allowed them to adjust to declining WBP. Bears reduced use of WBP stands without increasing movement rates, suggesting they obtained alternative fall foods within their local surroundings. However, the reduction in mortality risk historically associated with use of secure, high-elevation WBP habitat may be diminishing for bears residing in multiple-use areas.

Idaho, Montana, Wyoming

Re-evaluation of Yellowstone grizzly bear population dynamics not supported by empirical data: response to Doak & Cutler

Doak and Cutler critiqued methods used by the Interagency Grizzly Bear Study Team (IGBST) to estimate grizzly bear population size and trend in the Greater Yellowstone Ecosystem. Here, we focus on the premise, implementation, and interpretation of simulations they used to support their arguments. They argued that population increases documented by IGBST based on females with cubs-of-the-year were an artifact of increased search effort. However, we demonstrate their simulations were neither reflective of the true observation process nor did their results provide statistical support for their conclusion. They further argued that survival and reproductive senescence should be incorporated into population projections, but we demonstrate their choice of extreme mortality risk beyond age 20 and incompatible baseline fecundity led to erroneous conclusions. The conclusions of Doak and Cutler are unsubstantiated when placed within the context of a thorough understanding of the data, study system, and previous research findings and publications.

Conservation Letters