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For various temperate ungulate species, recent research has highlighted the potential for spring vegetation phenology ("green‐up") to influence individual condition, with purported benefits to population productivity. However, few studies have been able to measure the benefit on vital rates directly, and fewer still have investigated the comparative influence of other phenological periods on ungulate vital rates. In this study, we tracked phenological changes throughout the duration of the growing season and examined how their timing affected the probability of pregnancy in an ungulate population. We did this for elk (Cervus canadensis) across the Greater Yellowstone Ecosystem (GYE) by sampling 1106 adult females in winter at 25 sites over a 13‐year period and assessing sources of variation in pregnancy using a Bayesian hierarchical model. Pregnancy rates were generally high across the GYE (82.4%), and the primary influences on probability of pregnancy were the timing of vegetation senescence ("brown‐down") in autumn and exposure to the reproductive disease brucellosis. Earlier forage brown‐down in fall negatively influenced the probability of pregnancy of elk aged 6–9 years by an estimated 17.2% within the range (ca. 32 days) of observed brown‐down end dates. While summer habitat quality has been inferred to influence elk pregnancy previously, our findings specify the key influence of foraging conditions later in the seasonal cycle, immediately before the breeding season. The reproductive disease brucellosis was also an important factor, reducing the probability of pregnancy by 12.4% in elk in the 6‐ to 9‐year age class. Because pregnancy was tested before most disease‐induced abortions occur, the apparent mechanism for this effect is a prolonged reduction in fertility beyond the period of initial exposure in which fetal mortality is typically expected. Our results prompt greater scrutiny of the combined effects of late‐season phenology and disease on reproductive rates and population productivity in temperate ungulates. [ABSTRACT FROM AUTHOR]

Human activity and associated landscape modifications alter the movements of animals with consequences for populations and ecosystems worldwide. Species performing long‐distance movements are thought to be particularly sensitive to human impact. Despite the increasing anthropogenic pressure, it remains challenging to understand and predict animals' responses to human activity. Here we address this knowledge gap using 1206 Global Positioning System movement trajectories of 815 individuals from 14 red deer (Cervus elaphus) and 14 elk (Cervus canadensis) populations spanning wide environmental gradients, namely the latitudinal range from the Alps to Scandinavia in Europe, and the Greater Yellowstone Ecosystem in North America. We measured individual‐level movements relative to the environmental context, or movement expression, using the standardized metric Intensity of Use, reflecting both the directionality and extent of movements. We expected movement expression to be affected by resource (Normalized Difference Vegetation Index, NDVI) predictability and topography, but those factors to be superseded by human impact. Red deer and elk movement expression varied along a continuum, from highly segmented trajectories over relatively small areas (high intensity of use), to directed transitions through restricted corridors (low intensity of use). Human activity (Human Footprint Index, HFI) was the strongest driver of movement expression, with a steep increase in Intensity of Use as HFI increased, but only until a threshold was reached. After exceeding this level of impact, the Intensity of Use remained unchanged. These results indicate the overall sensitivity of Cervus movement expression to human activity and suggest a limitation of plastic responses under high human pressure, despite the species also occurring in human‐dominated landscapes. Our work represents the first comparison of metric‐based movement expression across widely distributed populations of a deer genus, contributing to the understanding and prediction of animals' responses to human activity. [ABSTRACT FROM AUTHOR]

Conversion of land for settlements and agriculture is increasing globally and can influence wildlife space use. However, there is limited research to identify the thresholds of land‐use change that incur wildlife avoidance and how these thresholds might vary across levels of selection.We evaluated multi‐level avoidance thresholds of elk Cervus canadensis impacted by residential development and irrigated agriculture across the Greater Yellowstone Ecosystem in Idaho, Montana and Wyoming. Using GPS data from 765 elk in 21 herds, we estimated habitat selection in relation to development and agriculture at three levels (home range selection, within home range selection and movement path selection). Next, using individual selection covariates and associated measures of land‐use availability, we used functional‐response models to evaluate how selection varied based on availability, and in turn, to estimate avoidance thresholds.We found individual and level‐specific variation in elk responses to environmental factors. Elk exhibited stronger responses (either selection or avoidance) when selecting home range locations (i.e. second‐order selection) than when selecting areas within home ranges (i.e. third‐order selection) or selecting movement paths (i.e. fourth‐order selection). Importantly, elk avoidance of development and agriculture changed as the amount of land in these categories changed. Across all levels of selection elk exhibited neutral selection for human development at low levels of availability (<1.1%–2.2% developed) but avoided areas that were >1.1%–2.2% developed. Conversely, elk selected positively for irrigated agriculture at low to moderate levels of availability (<52.0%–66.2% agriculture) but exhibited neutral selection in areas that were >52.0%–66.2% agriculture.Synthesis and applications. Elk avoidance of low levels of human development suggests conservation efforts such as restrictions on future development or conservation easements could focus on areas that are still below 2% developed. Additionally, because elk selection was strongest at the landscape scale, conservation actions that are based on information about the overall landscape structure may be most impactful. Our results highlight the importance of understanding variability in wildlife habitat selection at multiple levels, particularly in relation to land‐use change, and highlight how functional response modelling can help inform landscape conservation. [ABSTRACT FROM AUTHOR]

A growing body of evidence shows that some ungulates alternate between migratory and nonmigratory behaviors over time. Yet it remains unclear whether such short‐term behavioral changes can help explain reported declines in ungulate migration worldwide, as opposed to long‐term demographic changes. Furthermore, advances in tracking technology reveal that a simple distinction between migration and nonmigration may not sufficiently describe all individual behaviors. To better understand the dynamics and drivers of ungulate switching behavior, we investigated 14 years of movement data from 361 elk in 20 herds across the Greater Yellowstone Ecosystem (GYE). First, we categorized yearly individual behaviors using a clustering algorithm that identified similar migratory tactics across a continuum of behaviors. Then, we tested seven hypotheses to explain why some ungulates switch behaviors, and we evaluated how behavioral changes affected the proportions of different behaviors across the system. We identified four distinct behavioral tactics: residents (4.8% of elk‐years), short‐distance migrants (53.7%), elevational migrants (21.9%) and long‐distance migrants (19.6%). Of the 20 herds, 18 were partially migratory, and 5 had all four movement tactics present. We observed switches between migratory tactics in all sets of consecutive years during our study period, with an average of 22.5% of individual elk changing movement tactics from one year to the next. Elk in herds with higher movement tactic diversity were significantly more likely to switch tactics and often responded more effectively to adverse environmental conditions, compared to those in herds with low movement tactic diversity. During our study period, switching increased the prevalence of both short‐ and long‐distance migrants, decreased the prevalence of elevational migrants, and had no effect on the prevalence of residents. Our findings suggest that rather than contributing to the declining migratory behavior found in the GYE, switching behavior may enable greater resiliency to continuously changing environmental and anthropogenic conditions. [ABSTRACT FROM AUTHOR]

Novel approaches to quantifying density and distributions could help biologists adaptively manage wildlife populations, particularly if methods are accurate, consistent, cost‐effective, rapid, and sensitive to change. Such approaches may also improve research on interactions between density and processes of interest, such as disease transmission across multiple populations. We assess how satellite imagery, unmanned aerial system (UAS) imagery, and Global Positioning System (GPS) collar data vary in characterizing elk density, distribution, and count patterns across times with and without supplemental feeding at the National Elk Refuge (NER) in the US state of Wyoming. We also present the first comparison of satellite imagery data with traditional counts for ungulates in a temperate system. We further evaluate seven different aggregation metrics to identify the most consistent and sensitive metrics for comparing density and distribution across time and populations. All three data sources detected higher densities and aggregation locations of elk during supplemental feeding than non‐feeding at the NER. Kernel density estimates (KDEs), KDE polygon areas, and the first quantile of interelk distances detected differences with the highest sensitivity and were most highly correlated across data sources. Both UAS and satellite imagery provide snapshots of density and distribution patterns of most animals in the area at lower cost than GPS collars. While satellite‐based counts were lower than traditional counts, aggregation metrics matched those from UAS and GPS data sources when animals appeared in high contrast to the landscape, including brown elk against new snow in open areas. UAS counts of elk were similar to traditional ground‐based counts on feed grounds and are the best data source for assessing changes in small spatial extents. Satellite, UAS, or GPS data can provide appropriate data for assessing density and changes in density from adaptive management actions. For the NER, where high elk densities are beneath controlled airspace, GPS collar data will be most useful for evaluating how management actions, including changes in the dates of supplemental feeding, influence elk density and aggregation across large spatial extents. Using consistent and sensitive measures of density may improve research on the drivers and effects of density within and across a wide range of species. [ABSTRACT FROM AUTHOR]

Adaptive management has emerged as the prevailing approach for combining environmental research and management to advance science and policy. Adaptive management, as originally formulated by Carl Walters in 1986, depends on the use of Bayesian models to provide a framework to accumulate knowledge. The emergence of ecological forecasting using the Bayesian framework has provided robust tools and supports a new approach to informing adaptive management, which can be particularly useful in developing policy for managing infectious disease in wildlife. We used the potential infection of elk populations with chronic wasting disease in the Jackson Valley of Wyoming and the National Elk Refuge as a model system to show how Bayesian forecasting can support adaptive management in anticipation of management challenges. The core of our approach resembles the sex‐ and age‐structured, discrete time models used to support management decisions on elk harvest throughout western North America. Our model differs by including stages for CWD‐infected and unaffected animals. We used data on population counts, sex and age classification, and CWD testing, as well as results from prior research, in a Bayesian statistical framework to predict model parameters and the number of animals in each age, sex, and disease stage over time. Initial forecasts suggested CWD may reach a mean prevalence in the population of 12%, but uncertainty in this forecast is large and we cannot rule out a mean forecasted prevalence as high as 20%. Using recruitment rates observed during the last two decades, the model predicted that a CWD prevalence of 7% in females would cause the population growth rate (λ) to drop below 1, resulting in population declines even when female harvest was zero. The primary value of this ecological forecasting approach is to provide a framework to assimilate data with understanding of disease processes to enable continuous improvement in understanding the ecology of CWD and its management. [ABSTRACT FROM AUTHOR]

Supplemental feeding of wildlife is a common practice often undertaken for recreational or management purposes, but it may have unintended consequences for animal health. Understanding cryptic effects of diet supplementation on the gut microbiomes of wild mammals is important to inform conservation and management strategies. Multiple laboratory studies have demonstrated the importance of the gut microbiome for extracting and synthesizing nutrients, modulating host immunity, and many other vital host functions, but these relationships can be disrupted by dietary perturbation. The well-described interplay between diet, the microbiome, and host health in laboratory and human systems highlights the need to understand the consequences of supplemental feeding on the microbiomes of free-ranging animal populations. This study describes changes to the gut microbiomes of wild elk under different supplemental feeding regimes. We demonstrated significant cross-sectional variation between elk at different feeding locations and identified several relatively low-abundance bacterial genera that differed between fed versus unfed groups. In addition, we followed four of these populations through mid-season changes in supplemental feeding regimes and demonstrated a significant shift in microbiome composition in a single population that changed from natural forage to supplementation with alfalfa pellets. Some of the taxonomic shifts in this population mirrored changes associated with ruminal acidosis in domestic livestock. We discerned no significant changes in the population that shifted from natural forage to hay supplementation, or in the populations that changed from one type of hay to another. Our results suggest that supplementation with alfalfa pellets alters the native gut microbiome of elk, with potential implications for population health. [ABSTRACT FROM AUTHOR]

Wildlife aggregation patterns can influence disease transmission. However, limited research evaluates the influence of anthropogenic and natural factors on aggregation. Many managers would like to reduce wildlife contact rates, driven by aggregation, to limit disease transmission. We develop a novel analytical framework to quantify how management activities such as supplemental feeding and hunting versus weather drive contact rates while accounting for correlated contacts. We apply the framework to the National Elk Refuge (NER), Wyoming, USA, where the probable arrival of chronic wasting disease (CWD) has magnified concerns.We used a daily proximity index to measure contact rates among 68 global positioning system collared elk from 2016 to 2019. We modelled contact rates as a function of abiotic weather‐related effects, anthropogenic effects and aggregation from the prior day. The winter of 2017–2018 had greater natural forage availability and little snow, which led to a rare non‐feeding year on the NER and provided a unique opportunity to evaluate the effect of feeding on contact rates relative to other conditions.Supplemental feeding was the strongest predictor of aggregation, and contact rates were 2.6 times larger while feeding occurred compared to the baseline rate (0.34 and 0.13, respectively). Snow‐covered area was the second strongest predictor of contact rates highlighting the importance of abiotic factors to elk aggregation, but this effect had half the strength of feeding. These results are the first to show, even in animals that congregate naturally, how greatly supplemental feeding amplifies aggregation. Contact rates were also 23% lower during times when elk hunting was active (0.10) compared to the baseline.Synthesis and applications. Supplemental feeding increased contacts between elk well above the natural effects of weather, even after accounting for correlated movement expected in wintering ungulates. Similarly, differences in hunting season timing with adjacent areas led to an increase in contacts, suggesting an additional management option for reducing aggregation. The analytical framework presented supports the evaluation of temporally varying management actions that influence aggregation broadly and can be easily implemented whether the interest in changing aggregation is related to reduction of disease transmission, human–wildlife conflict or inter‐species competition. [ABSTRACT FROM AUTHOR]

Migration is an effective behavioral strategy for prolonging access to seasonal resources and may be a resilient strategy for ungulates experiencing changing climatic conditions. In the Greater Yellowstone Ecosystem (GYE), elk are the primary ungulate, with approximately 20,000 individuals migrating to exploit seasonal gradients in forage while also avoiding energetically costly snow conditions. How climate‐induced changes in plant phenology and snow accumulation are influencing elk migration timing is unknown. We present the most complete record of elk migration across the GYE, spanning 9 herds and 414 individuals from 2001 to 2017, to evaluate the drivers of migration timing and test for temporal shifts. The timing of elk departure from winter range involved a trade‐off between current and anticipated forage conditions, while snow melt governed summer range arrival date. Timing of elk departure from summer range and arrival on winter range were both influenced by snow accumulation and exposure to hunting. At the GYE scale, spring and fall migration timing changed through time, most notably with winter range arrival dates becoming almost 50 days later since 2001. Predicted herd‐level changes in migration timing largely agreed with observed GYE‐wide changes—except for predicted winter range arrival dates which did not reflect the magnitude of change detected in the elk telemetry data. Snow melt, snow accumulation, and spring green‐up dates all changed through time, with different herds experiencing different rates and directions of change. We conclude that elk migration is plastic, is a direct response to environmental cues, and that these environmental cues are not changing in a consistent manner across the GYE. The impacts of changing elk migration timing on predator–prey dynamics, carnivore–livestock conflict, disease ecology, and harvest management across the GYE are likely to be significant and complex. [ABSTRACT FROM AUTHOR]

Formally protected areas are an important component of wildlife conservation, but face limitations in their effectiveness for migratory species. Improved stewardship of working lands around protected areas is one solution for conservation planning, but private working lands are vulnerable to development. In the Greater Yellowstone Ecosystem (GYE), ungulates such as elk (Cervus canadensis) use both protected areas and private lands throughout their annual migrations. We studied patterns of landownership, protection, and conservation challenges within the ranges of migratory elk in the GYE. We used GPS data from 1088 elk in 26 herds to define herd-level seasonal ranges, and extracted covariates related to landownership and protection, land use, and human infrastructure and development. All elk herds used land encompassing >1 ownership type. Most elk herds (92.3 % of herds, n = 24) used the highest proportion of private land in the winter (mean = 36.2 % private land). Most elk herds' winter ranges contained the highest building densities (mean = 1.24 buildings/km2), fence densities (mean = 1.02 km fence/km2), and cattle grazing (mean = 1.9 cows/km2), compared to migratory and summer ranges. Out of all ranges, 36.5 % of ranges did not have any zoning regulations, indicating the potential for future development. Our results show that elk in the GYE rely on habitat outside of protected areas, and face landscape-scale conservation challenges such as habitat fragmentation from human development, particularly in winter ranges. Future conservation strategies for wildlife in this system need to encompass coordination across both public and private land to ensure migratory connectivity. [ABSTRACT FROM AUTHOR]

COVID-19 lockdowns in early 2020 reduced human mobility, providing an opportunity to disentangle its effects on animals from those of landscape modifications. Using GPS data, we compared movements and road avoidance of 2300 terrestrial mammals (43 species) during the lockdowns to the same period in 2019. Individual responses were variable with no change in average movements or road avoidance behavior, likely due to variable lockdown conditions. However, under strict lockdowns 10-day 95th percentile displacements increased by 73%, suggesting increased landscape permeability. Animals' 1-hour 95th percentile displacements declined by 12% and animals were 36% closer to roads in areas of high human footprint, indicating reduced avoidance during lockdowns. Overall, lockdowns rapidly altered some spatial behaviors, highlighting variable but substantial impacts of human mobility on wildlife worldwide.

Providing food to wildlife during periods when natural food is limited results in aggregations that may facilitate disease transmission. This is exemplified in western Wyoming where institutional feeding over the past century has aimed to mitigate wildlife-livestock conflict and minimize winter mortality of elk ( Cervus canadensis ). Here we review research across 23 winter feedgrounds where the most studied disease is brucellosis, caused by the bacterium Brucella abortus Traditional veterinary practices (vaccination, test-and-slaughter) have thus far been unable to control this disease in elk, which can spill over to cattle. Current disease-reduction efforts are being guided by ecological research on elk movement and density, reproduction, stress, co-infections and scavengers. Given the right tools, feedgrounds could provide opportunities for adaptive management of brucellosis through regular animal testing and population-level manipulations. Our analyses of several such manipulations highlight the value of a research-management partnership guided by hypothesis testing, despite the constraints of the sociopolitical environment. However, brucellosis is now spreading in unfed elk herds, while other diseases (e.g. chronic wasting disease) are of increasing concern at feedgrounds. Therefore experimental closures of feedgrounds, reduced feeding and lower elk populations merit consideration.This article is part of the theme issue 'Anthropogenic resource subsidies and host-parasite dynamics in wildlife'., (© 2018 The Authors.)