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Mark A. Haroldson

Publications and source records attributed to Mark A. Haroldson.

At least 19 recordsLinked to original sources
USGS2026

An energetic tradeoff best explains parturition timing in grizzly bears

Timing of grizzly bear ( Ursus arctos ) parturition during hibernation has been explained by ancestral traits (delayed implantation, altricial young, obligate maternal denning), but the ultimate driver underlying precise timing has not been fully explored. Capitalizing on an observed latitudinal increase in denning duration among four populations in interior North America, we tested two alternative hypotheses. First, that birth timing results from a physiological cue that synchronizes implantation with the onset of hibernation, allowing females to forgo reproduction should they lack adequate fat stores. Alternatively, that parturition is optimally timed relative to den exit to balance an energetic tradeoff between minimizing lactation time to protect the mother and maximizing developmental time to increase cub survival. Using parturition dates previously predicted from accelerometer data (27 Dec–28 Feb), we classified 115 females according to apparent litter survival when first visually observed after den exit: 57% successful (with cubs), 22% unsuccessful (alone), and 21% unknown (not observed). The number of days between birth and den exit showed no association with latitude ( p = 0.29). It averaged 103 days among successful females but only 77 days among unsuccessful females ( p < 0.001) owing to later births and earlier exit. With each increasing degree of latitude, birth date increased by 1.0 and number of days between den entry and birth increased by 2.5 ( p < 0.001). Implantation dates were not centered on den entry dates ( p < 0.001). These results supported the energetic tradeoff hypothesis and suggested natural selection has favored a consistent number of days between parturition and den exit under average body conditions and shifts toward later or earlier births for females with lower or higher levels of bodily stored energy, respectively. This flexible tradeoff may support resilience to climate change and present a possible mechanism explaining reduced natality and cub survival in high-density populations.

Alaska, British Columbia, Idaho, Montana, Wyoming↗
USGS2026

Detecting occurrence and timing of grizzly bear parturition based on anomalies in accelerometer data

Documenting natality of brown (grizzly) bears ( Ursus arctos ) is an important component of many population monitoring and research programs. Brown bears give birth during hibernation, so observation of litters is generally not feasible until females exit dens, and detection of litters can be compromised by poor observability. Using triaxial accelerometer activity data stored on radiocollars, we developed a technique that predicts births by the presence of brief upsurges in activity likely associated with postnatal maternal behaviors. We developed presence criteria with a training sample of known parturient females ( n = 22) to detect anomalies within time-series of daily motion-counts (i.e., activity readings >0 measured at 10-min intervals) during 25 December–7 March. To test performance, we applied criteria to a blinded sample of activity data obtained from female grizzly bears (captured and collared during 2012–2022) in 4 populations in interior North America ( n = 295). We assigned predicted status and compared assignments to reproductive status at first visual observation. The true positive rate was 91% ( n = 47 females observed with cubs of the year) and the false-positive rate was 17% ( n = 65 females observed with older offspring). Births were predicted for 50% of females with unknown reproductive status ( n = 48), 21% of females observed without offspring ( n = 114), and 10% of subadult females considered too young to reproduce ( n = 21). Dates of predicted births varied from 27 December to 28 February. Our anomaly detection technique was successful at estimating parturition events, and despite some error, indicated that a number of litters were born but not observed, presumably due to mortality in or shortly after den emergence. This technique provides an additional tool for supplementing visual observations for natality estimation and population modeling, provided that potential biases stemming from increased detection rates are considered.

Alaska, British Columbia, Idaho, Montana↗
USGS2025

Bioclimatic, demographic, and anthropogenic correlates of grizzly bear activity patterns in the Greater Yellowstone Ecosystem

Plasticity of diel activity rhythms may be a key element for adaptations of wildlife populations to changing environmental conditions. In the last decades, grizzly bears Ursus arctos in the Greater Yellowstone Ecosystem (GYE) have experienced notable environmental fluctuations, including changes in availability of food sources and severe droughts. Although substantial research has been conducted on grizzly bear diets, space use, and demographic parameters, studies on factors that may influence their diel activity patterns are lacking. We investigated diel activity of grizzly bears in the GYE as a function of anthropogenic landscape modification, maximum daily ambient temperature, drought severity, and bear density. Specifically, we used accelerometry readings of 169 bears (39 females, 130 males) from 2009 to 2022 to compute three complementary activity measures, hourly intensity of activity, daily active minutes, and active bout length, each used as a response variable within a Bayesian modeling framework. Grizzly bears generally exhibited bimodal diel activity, with crepuscular peaks and slight variations across seasons. Females with young (i.e. cubs or yearlings) were an exception, with more pronounced diurnal activity patterns, possibly as a strategy to avoid infanticide by dominant males. Landscape modification and maximum ambient temperature were the factors most strongly associated with activity patterns of grizzly bears, with greater nocturnality observed in lone females and males as these factors increased. Females with young were comparatively less affected. The GYE is changing because of increasing land development, human recreation pressures, and effects of climate change. Given their greater diurnal activity compared with other cohorts, female grizzly bears with dependent offspring may be more constrained in their ability to modify activity patterns. Our findings add to a growing body of research emphasizing the importance of the temporal dimension of wildlife behavior as a critical factor in assessing species adaptability and vulnerability in a changing world.

Idaho, Montana, Wyoming↗
USGS2025

Exploring the exploitation of migratory moths by radio-marked grizzly bears in Wyoming

Grizzly bear ( Ursus arctos ) feeding on migratory army cutworm moths ( Euxoa auxiliaris ) was first documented by the Interagency Grizzly Bear Study Team (IGBST) during the early 1980s in the southeastern portion of the Greater Yellowstone Ecosystem (GYE). Since those initial observations, use of this seasonally available food resource by grizzly bears has grown substantially. As of 2023, we have documented 4,754 observations of grizzly bears feeding or digging at high-elevation talus slopes. We used those records to identify 36 unique moth sites in the GYE, assessed geographic characteristics of these sites, and documented chronology and frequency of use by grizzly bears. We used occurrences of radio-collared grizzly bears to identify a sample of bears that foraged at moth sites and investigated characteristics of fidelity, duration, diel activity, and movement patterns. Grizzly bears exhibited high fidelity to specific moth sites within and across years. Bears showed approximately 50% reductions in movement metrics while using sites, albeit with increased activity patterns. Estimates of feeding metrics by females exceeded those of males in intensity and duration of both use and daily activity. Given increasing human impacts and interest in observing this feeding interaction, quantifying metrics that describe the spatial and temporal patterns of moth site use by grizzly bears could be beneficial for future management. Further analyses are needed to fully examine the relationship between caloric influences of moth use on grizzly bear demographic rates.

Idaho, Montana, Wyoming↗
USGS2025

The influence of human presence and footprint on animal space use in US national parks

Given the importance of protected areas for biodiversity, the growth of visitation to many areas has raised concerns about the effects of humans on wildlife. In 2020, the COVID-19 pandemic led to temporary closure of national parks in the United States, offering a pseudonatural experiment to tease apart the effects of permanent infrastructure and transient human presence on animals. We compiled GPS tracking data from 229 individuals of 10 mammal species in 14 parks and used third-order hierarchical resource selection functions to evaluate the influence of the human footprint on animal space use in 2019 and 2020. Averaged across all parks and species, animals avoided the human footprint, whether the park was open or closed. However, although animals in remote areas showed consistent avoidance, on average those in more developed areas switched from avoidance to selection when protected areas were closed. Findings varied across species: some responded consistently negatively to the footprint (wolves, mountain goats), some positively (mule deer, red fox) and others had a strong exposure-mediated response (elk, mountain lion). Furthermore, some species responded more strongly to the park closure (black bear, moose). This study advances our understanding of complex interactions between recreation and wildlife in protected areas.

western United States↗
USGS2025

Female and male grizzly bears differ in their responses to low-intensity recreation in a protected area

Strategies animals use to navigate human-dominated landscapes frequently mimic anti-predator responses employed by prey species. Understanding how large carnivores respond to outdoor recreation is important for conservation, particularly in protected areas with preservation mandates. Visitation to Yellowstone National Park doubled from 1980 to 2015, increasing the need to examine potential changes in behavior of grizzly bears ( Ursus arctos ) in relation to human recreation sites (trails, backcountry campsites). We developed integrated step-selection functions to explore how recreation sites influenced the movement rate and selection by male and female grizzly bears. Further, we tested whether time of day (diurnal, crepuscular, nocturnal) and restrictions to human access (i.e., restricted, unrestricted) modified bear responses and then compared behaviors based on proximity to recreation sites. Male grizzly bears used trails to travel during crepuscular and nocturnal hours and exhibited more pronounced behavior in restricted areas compared with unrestricted areas, suggesting recreation in unrestricted areas influenced the behavior of male bears. In contrast, female bears varied their movement rate and selection of trails in restricted areas much more than in unrestricted areas, suggesting females may make security tradeoffs between male bears and people. Both sexes used trails, likely as energetically efficient travel corridors; however, our analyses did not indicate that bears spent time near backcountry campsites. The sex-based differences in selection and movement patterns associated with trails and campsites suggest a single management approach for recreation may not equally benefit all bears. Recreation impacts on wildlife are complex to characterize and predict, but simultaneously modeling movement and selection provides a more comprehensive assessment of strategies animals use to navigate perceived risk.

Montana, Wyoming↗
USGS2025

The diel niche of brown bears: Constraints on adaptive capacity in human-modified landscapes

D Diel activity rhythms, representing the behavioral pattern of the sleep–wake cycle, may be adjusted by wildlife in response to changes in environmental conditions. An increase in nocturnality is typically recognized as an adaptive strategy to segregate from humans and mitigate heat stress. Numerous studies have investigated spatial patterns and habitat use of large carnivores in human-modified landscapes, but little research has examined their activity rhythms. We compiled Global Positioning System data (2004–2022) for 139 brown bears Ursus arctos from six populations across Europe, representing a human-modified landscape, and the Greater Yellowstone Ecosystem, U.S.A., representing a landscape with limited human impact, which we used to calculate hourly movement rates as an activity proxy. Using a Bayesian approach to model the temporal autocorrelation of activity data, we tested if the extent of nocturnality in brown bears is modulated by intensity of human encroachment, accounting for primary productivity and maximum ambient temperature. All bear populations exhibited a predominantly bimodal, crepuscular pattern of activity, although Yellowstone bears were proportionally more crepuscular and diurnal. Whereas the effect of primary productivity was variable, all European populations became more nocturnal in response to higher human encroachment and reduced diurnal and crepuscular activity at higher summer temperatures, decreasing overall diel activity levels. Yellowstone bears displayed the greatest shift towards nocturnality among all populations in response to increasing human encroachment, and increased nocturnal activity to compensate for lower diurnal and crepuscular activity at higher summer temperatures. Our research indicates that European bears in human-modified landscapes may be reaching a limit in the behavioral plasticity they can manifest in their activity patterns, being already constrained into increased nocturnality. Our findings enhance the understanding of brown bear adaptive capacity to accommodate future changes, such as urbanization and increasing temperatures, to the ecosystems they inhabit.

Ecography↗
USGS2025

Identifying presence or absence of grizzly and polar bear cubs from the movements of adult females with machine learning

Background Information on reproductive success is crucial to understanding population dynamics but can be difficult to obtain, particularly for species that birth while denning. For grizzly ( Ursus arctos ) and polar bears ( U. maritimus ), den visits are impractical because of safety and logistical considerations. Reproduction is typically documented through direct observation, which can be difficult, costly, and often occurs long after den departure. Reproduction could be documented remotely, however, from post-denning movement data if discernable differences exist between females with and without cubs. Methods We trained support vector machines (SVMs) with eight variables derived from telemetry data of female grizzly (2000–2022) and polar bears (1985–2016) with or without cubs during seven periods with lengths ranging from 5 to 60 days starting at den departure. We assessed SVM classification accuracy by withholding two samples (one cub-present, one cub-absent), training SVMs with the remaining data, predicting classification of the withheld samples, and repeating this process for each sample combination. Additionally, we evaluated how classification accuracy for grizzly bears was influenced by sample size, length of the post-departure period, and frequency of standardized location estimates. Results Accuracy of predicting cub presence or absence was 87% for grizzly bears with only 5 days of post-departure data and increased to a maximum of 92% with 20 days of data. For polar bears, accuracy was 86% at 5 days post-departure and increased to a maximum of 93% at 50 days. Classification accuracy for grizzly bears increased from 76 to 90% when sample size increased from 10 to 30 bears while holding period length constant (30 days) but did not increase at larger sample sizes. When sample size was held constant, increasing the length of the post-departure period did not affect classification accuracy markedly. Conclusion Presence or absence of grizzly and polar bear cubs can be identified with high accuracy even when SVM models are trained with limited data. Detecting cub presence or absence remotely could improve estimates of reproductive success and litter survival, enhancing our understanding of factors affecting cub recruitment.

Alaska, Idaho, Montana, Wyoming↗
USGS2025

A summary of grizzly bear distribution in the lower-48 US states in 2024

Understanding the distribution of grizzly bear populations in the lower-48 states, is important for their conservation and management, and for public safety. Previously, our research teams working in grizzly bear ecosystems in the lower-48 states used varying methods to estimate distribution of grizzly bear populations. In the Greater Yellowstone Ecosystem (GYE) and Northern Continental Divide Ecosystem (NCDE), zonal analysis and ordinary kriging were applied to an array of grid cells with or without verified presence of grizzly bears, however the parameters of the methods varied between the two ecosystems. In the Cabinet-Yaak Ecosystem (CYE) and the Selkirk Ecosystem (SE), population distribution was mapped as the Recovery Zone plus “bears outside of Recovery Zone” areas (Allen 2011). Additionally, the U.S. Fish and Wildlife Service developed a method for estimating areas where grizzly bears “may be present” to help agencies or prospective applicants evaluate whether or not proposed actions may affect grizzly bears (U.S. Fish and Wildlife Service 2020). Since the mid 2010s, cooperating agencies have collaborated in documenting and maintaining a database of verified outlier observations that occur between or well outside of grizzly bear Recovery Zones and these data inform the “may be present” mapping.

lower 48 states↗
USGS2024

A unified approach to long-term population monitoring of grizzly bears in the Greater Yellowstone Ecosystem

Long-term wildlife research and monitoring programs strive to maintain consistent data collections and analytical methods. Incorporating new techniques is important but can render data sets incongruent and limit their potential to discern trends in demographic parameters. Integrated population models (IPMs) can address these limitations by combining data sources that may span different periods into a unified statistical framework while providing a holistic view of population dynamics. We developed an IPM in a Bayesian framework for grizzly bears ( Ursus arctos ) in the Greater Yellowstone Ecosystem. We coupled demographic data with multiple, independent population count data to link annual changes in abundance with vital rates over 4 decades (1983–2023). Abundance increased threefold from an estimated 270 individuals in 1984 to 1030 individuals in 2023. Parameter estimates indicated survival of bears ≥2 years of age was high, contributing to robust population growth during the 1980s (λ = 1.023 [50 % interquartile range = 0.993–1.082]) and 1990s (λ = 1.064 [1.023–1.103]). A slowing of population growth started around 2000 (2000s: λ = 1.030 [0.989–1.068]) and continued into the 2010s (λ = 1.021 [0.985–1.057]), due primarily to reductions in survival of bears <2 years of age. These findings corroborate previous research that identified density-dependent effects as a likely cause. The IPM framework provided greater certainty and understanding regarding the dynamic demographic characteristics of the population and serves as a powerful monitoring tool for this long-lived species. Implementation of the IPM allows timely dissemination of demographic data to help inform adaptive management strategies and policy decisions necessary for the continued management and conservation of this population. This robust and flexible monitoring system allows scientists to investigate the effects of a changing ecosystem on population dynamics, incorporate new data sources and statistical models, and respond to changes in monitoring needs for the population. We highlight the efficacy of the IPM in estimating and tracking demographic parameters for a long-lived species, while accommodating shifts in monitoring techniques and data collections typical of long-term wildlife conservation programs worldwide.

Idaho, Montana, Wyoming↗
USGS2024

Integrating ecological value and charismatic species habitats to prioritize habitats for conservation: A case study from Greater Yellowstone

Expanding human pressure has reduced natural habitats globally and motivated strategies to conserve remaining natural habitats. Decisions about conservation on private lands, however, are typically made by local stakeholders who are motivated by the elements of nature they most highly value. Thus, national prioritization for conservation should be complemented by local analysis of species or habitats that most influence local landowner decisions. We demonstrate within the Greater Yellowstone Ecosystem how quantitative mapping of wildlife species that are highly valued by local residents can be integrated with indices of ecosystem integrity to prioritize private lands for conservation. We found that natural vegetation cover (NVC) comprised 81% of the private lands. Some watersheds have lost 6% of NVC since 2001 and developed lands now cover >40% of their areas. Locations high in ecological value, elk habitat, and grizzly habitat occurred in different biophysical settings. Consequently, only 2% of the NVC supports high levels of all three biodiversity measures and 26% of this area was within conservation easements. The remaining areas of high biodiversity value that are unprotected are priorities for conservation. We suggest that national-scale conservation planning will be most effective on private lands if additional within-ecoregion analyses are done on the elements of biodiversity that are most valued by local people.

Biological Conservation↗
USGS2023

Grizzly bear responses to restrictions of recreation in Yellowstone National Park

Avoiding humans will be more difficult and energetically costly for animals as outdoor recreation increases and people venture farther into wildland areas that provide high-quality habitat for wildlife. Restricting human access can be an attractive management tool to mitigate effects of human recreation activities on wildlife; however, the efficacy of such measures is rarely assessed. In 1982, Yellowstone National Park identified areas important to grizzly bears ( Ursus arctos ) to help protect critical grizzly bear habitat and reduce the likelihood of human injuries by bears. Referred to as bear management areas (BMAs), human access is restricted in these areas for 2–8 months each year, with timing and type of restrictions varying by area. We examined 2 datasets to evaluate grizzly bear selection of BMAs and differences of bear density in BMAs and non-BMAs. First, we used 17 years of recent global positioning system telemetry data for grizzly bears to assess their selection of BMAs during periods when human access was allowed, and when access was restricted. We used step-selection functions to test the hypothesis that bears spend time in places that allow them to avoid people and select quality food sources. There was support that grizzly bears differentially select for BMAs regardless of whether human access was restricted at the time, compared with areas outside BMAs, and that selection changed with sex and season. Only males during the summer and hyperphagic seasons changed their selection of BMAs based on whether access restrictions were in place, and overall, male bears preferred unrestricted BMAs (BMAs without restrictions in place). Females preferentially selected BMAs regardless of whether the area had access restrictions in place only during the mating season. Individuals varied widely in their preference for BMAs and access restrictions. Bears likely choose to spend time in BMAs based on available food resources rather than restrictions to human access. Supporting this interpretation, our analyses indicated that a greater proportion of BMA in an area was associated with higher densities of grizzly bear. Thus, restrictions to human access likely help reduce the potential for human–bear interactions, accomplishing one of the original objectives for establishing the BMAs.

Idaho, Montana, Wyoming↗
USGS2023

Grizzly bear movement models predict habitat use for nearby populations

Conservation planning and decision-making can be enhanced by ecological models that reliably transfer to times and places beyond those where models were developed. Transferrable models can be especially helpful for species of conservation concern, such as grizzly bears ( Ursus arctos ). Currently, only four grizzly bear populations remain in the contiguous United States. We evaluated transferability of previously derived individual-based, integrated step selection functions (iSSFs) developed from GPS-collared grizzly bears in the Northern Continental Divide Ecosystem by applying them within the nearby Selkirk (SE), Cabinet-Yaak (CYE), and Greater Yellowstone Ecosystems (GYE). We simulated 100 replicates of 5000 steps for each iSSF in each ecosystem, summarized relative use into 10 equal-area classes for each sex, and overlaid GPS locations from bears in the SE, CYE, and GYE on resulting maps. Spearman rank correlations between numbers of locations and class rank were ≥ 0.96 within each study area, indicating models were highly predictive of grizzly bear space use in these nearby populations. Assessment of models using smaller subsets of data in space and time demonstrated generally high predictive accuracy for females. Although generally high across space and time, predictive accuracy for males was low within some watersheds and in summer within the SE and CYE, potentially due to seasonal effects, vegetation, and food assemblage differences. Altogether, these results demonstrated high transferability of our models to landscapes in the Northern Rocky Mountains, suggesting they may be used to evaluate habitat suitability and connectivity throughout the region to benefit conservation planning.

Idaho, Montana, Washington, Wyoming↗
USGS2023

Genetic architecture and evolution of color variation in American black bears

Color variation is a frequent evolutionary substrate for camouflage in small mammals, but the underlying genetics and evolutionary forces that drive color variation in natural populations of large mammals are mostly unexplained. The American black bear, Ursus americanus ( U. americanus ), exhibits a range of colors including the cinnamon morph, which has a similar color to the brown bear, U. arctos , and is found at high frequency in the American southwest. Reflectance and chemical melanin measurements showed little distinction between U. arctos and cinnamon U. americanus individuals. We used a genome-wide association for hair color as a quantitative trait in 151 U. americanus individuals and identified a single major locus (p < 10 −13 ). Additional genomic and functional studies identified a missense alteration (R153C) in Tyrosinase-related protein 1 ( TYRP1 ) that likely affects binding of the zinc cofactor, impairs protein localization, and results in decreased pigment production. Population genetic analyses and demographic modeling indicated that the R153C variant arose 9.36 kya in a southwestern population where it likely provided a selective advantage, spreading both northwards and eastwards by gene flow. A different TYRP1 allele, R114C, contributes to the characteristic brown color of U. arctos but is not fixed across the range.

Current Biology↗
USGS2023

Evidence for density-dependent effects on body composition of a large omnivore in a changing Greater Yellowstone Ecosystem

Understanding the density-dependent processes that drive population demography in a changing world is critical in ecology, yet measuring performance–density relationships in long-lived mammalian species demands long-term data, limiting scientists' ability to observe such mechanisms. We tested performance–density relationships for an opportunistic omnivore, grizzly bears ( Ursus arctos , Linnaeus, 1758) in the Greater Yellowstone Ecosystem, with estimates of body composition (lean body mass and percent body fat) serving as indicators of individual performance over two decades (2000–2020) during which time pronounced environmental changes have occurred. Several high-calorie foods for grizzly bears have mostly declined in recent decades (e.g., whitebark pine [ Pinus albicaulis , Engelm, 1863]), while increasing human impacts from recreation, development, and long-term shifts in temperatures and precipitation are altering the ecosystem. We hypothesized that individual lean body mass declines as population density increases (H1), and that this effect would be more pronounced among growing individuals (H2). We also hypothesized that omnivory helps grizzly bears buffer energy intake from changing foods, with body fat levels being independent from population density and environmental changes (H3). Our analyses showed that individual lean body mass was negatively related to population density, particularly among growing-age females, supporting H1 and partially H2. In contrast, population density or sex had little effect on body fat levels and rate of accumulation, indicating that sufficient food resources were available on the landscape to accommodate successful use of shifting food sources, supporting H3. Our results offer important insights into ecological feedback mechanisms driving individual performances within a population undergoing demographic and ecosystem-level changes. However, synergistic effects of continued climate change and increased human impacts could lead to more extreme changes in food availability and affect observed population resilience mechanisms. Our findings underscore the importance of long-term studies in protected areas when investigating complex ecological relationships in an increasingly anthropogenic world.

Idaho, Montana, Wyoming↗
USGS2023

Enhancements to population monitoring of Yellowstone grizzly bears

In the Greater Yellowstone Ecosystem, counts of female grizzly bears ( Ursus arctos ) with cubs-of-the-year (females with cubs) from systematic aerial surveys and opportunistic ground sightings are combined with demographic data to derive annual population estimates. We addressed 2 limitations to the monitoring approach. As part of a rule set, a conservative distance of > 30 km currently is used as a threshold to assign sightings to unique females with cubs, resulting in underestimation bias. Using telemetry locations of females with cubs collected during 1997–2019, we created 1,000 data sets for each of 5 levels of simulated number of females with cubs, simulated sightings by selecting among these locations, and evaluated the classification performance of alternative distance criteria (12–30 km). Under all scenarios, 12–16-km criteria maximized classification performance and minimized estimation bias; the 16-km criterion was optimal for current conditions and sampling efforts. Our second objective was to test generalized additive models (GAMs) as a flexible trend analysis technique. We simulated 1,000 time series for each of 10 scenarios (10, 15, and 20% decline over periods of 5, 10, and 15 yrs, plus stability), applied GAMs, and assessed metrics associated with the posterior distribution of the instantaneous rate of change. We detected declines among > 99.6% of replicates under the 15 and 20% decline scenarios and in 84.7–94.7% of replicates under the 10% decline scenario. From decline onset to first detection, periods ranged from 3.7 (20% decline over 5 yrs) to 11.1 (10% decline over 15 yrs), with 3.9–8.8 years mean duration of detection events. The GAM approach allows detection of directional changes in population trend, including early warning metrics, and stabilization after such changes. Retrospective application of the 16-km distance criterion and GAMs resulted in higher population estimates and growth rates than are reported using current methods.

Wyoming↗