Hall Sawyer

Winter Habitat Selection of Mule Deer Before and During Development of a Natural Gas Field

Abstract Increased levels of natural gas exploration, development, and production across the Intermountain West have created a variety of concerns for mule deer (Odocoileus hemionus) populations, including direct habitat loss to road and well-pad construction and indirect habitat losses that may occur if deer use declines near roads or well pads. We examined winter habitat selection patterns of adult female mule deer before and during the first 3 years of development in a natural gas field in western Wyoming. We used global positioning system (GPS) locations collected from a sample of adult female mule deer to model relative frequency or probability of use as a function of habitat variables. Model coefficients and predictive maps suggested mule deer were less likely to occupy areas in close proximity to well pads than those farther away. Changes in habitat selection appeared to be immediate (i.e., year 1 of development), and no evidence of well-pad acclimation occurred through the course of the study; rather, mule deer selected areas farther from well pads as development progressed. Lower predicted probabilities of use within 2.7 to 3.7 km of well pads suggested indirect habitat losses may be substantially larger than direct habitat losses. Additionally, some areas classified as high probability of use by mule deer before gas field development changed to areas of low use following development, and others originally classified as low probability of use were used more frequently as the field developed. If areas with high probability of use before development were those preferred by the deer, observed shifts in their distribution as development progressed were toward less-preferred and presumably less-suitable habitats.

Identifying and prioritizing ungulate migration routes for landscape‐level conservation

As habitat loss and fragmentation increase across ungulate ranges, identifying and prioritizing migration routes for conservation has taken on new urgency. Here we present a general framework using the Brownian bridge movement model (BBMM) that: (1) provides a probabilistic estimate of the migration routes of a sampled population, (2) distinguishes between route segments that function as stopover sites vs. those used primarily as movement corridors, and (3) prioritizes routes for conservation based upon the proportion of the sampled population that uses them. We applied this approach to a migratory mule deer (Odocoileus hemionus) population in a pristine area of southwest Wyoming, USA, where 2000 gas wells and 1609 km of pipelines and roads have been proposed for development. Our analysis clearly delineated where migration routes occurred relative to proposed development and provided guidance for on-the-ground conservation efforts. Mule deer migration routes were characterized by a series of stopover sites where deer spent most of their time, connected by movement corridors through which deer moved quickly. Our findings suggest management strategies that differentiate between stopover sites and movement corridors may be warranted. Because some migration routes were used by more mule deer than others, proportional level of use may provide a reasonable metric by which routes can be prioritized for conservation. The methods we outline should be applicable to a wide range of species that inhabit regions where migration routes are threatened or poorly understood.

Stopover ecology of a migratory ungulate

1. Birds that migrate long distances use stopover sites to optimize fuel loads and complete migration as quickly as possible. Stopover use has been predicted to facilitate a time-minimization strategy in land migrants as well, but empirical tests have been lacking, and alternative migration strategies have not been considered. 2. We used fine-scale movement data to evaluate the ecological role of stopovers in migratory mule deer Odocoileus hemionus- a land migrant whose fitness is strongly influenced by energy intake rather than migration speed. 3. Although deer could easily complete migrations (range 18-144 km) in several days, they took an average of 3 weeks and spent 95% of that time in a series of stopover sites that had higher forage quality than movement corridors. Forage quality of stopovers increased with elevation and distance from winter range. Mule deer use of stopovers corresponded with a narrow phenological range, such that deer occupied stopovers 44 days prior to peak green-up, when forage quality was presumed to be highest. Mule deer used one stopover for every 5·3 and 6·7 km travelled during spring and autumn migrations, respectively, and used the same stopovers in consecutive years. 4. Study findings indicate that stopovers play a key role in the migration strategy of mule deer by allowing individuals to migrate in concert with plant phenology and maximize energy intake rather than speed. Our results suggest that stopover use may be more common among non-avian taxa than previously thought and, although the underlying migration strategies of temperate ungulates and birds are quite different, stopover use is important to both. 5. Exploring the role of stopovers in land migrants broadens the scope of stopover ecology and recognizes that the applied and theoretical benefits of stopover ecology need not be limited to avian taxa.

Influence of well pad activity on winter habitat selection patterns of mule deer.

Abstract Conversion of native winter range into producing gas fields can affect the habitat selection and distribution patterns of mule deer (Odocoileus hemionus). Understanding how levels of human activity influence mule deer is necessary to evaluate mitigation measures and reduce indirect habitat loss to mule deer on winter ranges with natural gas development. We examined how 3 types of well pads with varying levels of vehicle traffic influenced mule deer habitat selection in western Wyoming during the winters of 2005–2006 and 2006–2007. Well pad types included producing wells without a liquids gathering system (LGS), producing wells with a LGS, and well pads with active directional drilling. We used 36,699 Global Positioning System locations collected from a sample (n  =  31) of adult (>1.5-yr-old) female mule deer to model probability of use as a function of traffic level and other habitat covariates. We treated each deer as the experimental unit and developed a population-level resource selection function for each winter by averaging coefficients among models for individual deer. Model coefficients and predictive maps for both winters suggested that mule deer avoided all types of well pads and selected areas further from well pads with high levels of traffic. Accordingly, impacts to mule deer could probably be reduced through technology and planning that minimizes the number of well pads and amount of human activity associated with them. Our results suggested that indirect habitat loss may be reduced by approximately 38–63% when condensate and produced water are collected in LGS pipelines rather than stored at well pads and removed via tanker trucks. The LGS seemed to reduce long-term (i.e., production phase) indirect habitat loss to wintering mule deer, whereas drilling in crucial winter range created a short-term (i.e., drilling phase) increase in deer disturbance and indirect habitat loss. Recognizing how mule deer respond to different types of well pads and traffic regimes may improve the ability of agencies and industry to estimate cumulative effects and quantify indirect habitat losses associated with different development scenarios.

Linking anti-predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide-ranging, active predators should induce prey behavioural responses but not major, population- or community-level consequences. We evaluated the non-consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high-risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20-fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.

Habitat Selection of Rocky Mountain Elk in a Nonforested Environment

Abstract Recent expansions by Rocky Mountain elk (Cervus elaphus) into nonforested habitats across the Intermountain West have required managers to reconsider the traditional paradigms of forage and cover as they relate to managing elk and their habitats. We examined seasonal habitat selection patterns of a hunted elk population in a nonforested high-desert region of southwestern Wyoming, USA. We used 35,246 global positioning system locations collected from 33 adult female elk to model probability of use as a function of 6 habitat variables: slope, aspect, elevation, habitat diversity, distance to shrub cover, and distance to road. We developed resource selection probability functions for individual elk, and then we averaged the coefficients to estimate population-level models for summer and winter periods. We used the population-level models to generate predictive maps by assigning pixels across the study area to 1 of 4 use categories (i.e., high, medium-high, medium-low, or low), based on quartiles of the predictions. Model coefficients and predictive maps indicated that elk selected for summer habitats characterized by higher elevations in areas of high vegetative diversity, close to shrub cover, northerly aspects, moderate slopes, and away from roads. Winter habitat selection patterns were similar, except elk shifted to areas with lower elevations and southerly aspects. We validated predictive maps by using 528 locations collected from an independent sample of radiomarked elk (n = 55) and calculating the proportion of locations that occurred in each of the 4 use categories. Together, the high- and medium-high use categories of the summer and winter predictive maps contained 92% and 74% of summer and winter elk locations, respectively. Our population-level models and associated predictive maps were successful in predicting winter and summer habitat use by elk in a nonforested environment. In the absence of forest cover, elk seemed to rely on a combination of shrubs, topography, and low human disturbance to meet their thermal and hiding cover requirements.

A framework for understanding semi-permeable barrier effects on migratory ungulates

Summary 1. Impermeable barriers to migration can greatly constrain the set of possible routes and ranges used by migrating animals. For ungulates, however, many forms of development are semi-permeable, and making informed management decisions about their potential impacts to the persistence of migration routes is difficult because our knowledge of how semi-permeable barriers affect migratory behaviour and function is limited. 2. Here, we propose a general framework to advance the understanding of barrier effects on ungulate migration by emphasizing the need to (i) quantify potential barriers in terms that allow behavioural thresholds to be considered, (ii) identify and measure behavioural responses to semi-permeable barriers and (iii) consider the functional attributes of the migratory landscape (e.g. stopovers) and how the benefits of migration might be reduced by behavioural changes. 3. We used global position system (GPS) data collected from two subpopulations of mule deer Odocoileus hemionus to evaluate how different levels of gas development influenced migratory behaviour, including movement rates and stopover use at the individual level, and intensity of use and width of migration route at the population level. We then characterized the functional landscape of migration routes as either stopover habitat or movement corridors and examined how the observed behavioural changes affected the functionality of the migration route in terms of stopover use. 4. We found migratory behaviour to vary with development intensity. Our results suggest that mule deer can migrate through moderate levels of development without any noticeable effects on migratory behaviour. However, in areas with more intensive development, animals often detoured from established routes, increased their rate of movement and reduced stopover use, while the overall use and width of migration routes decreased. 5. Synthesis and applications. In contrast to impermeable barriers that impede animal movement, semi-permeable barriers allow animals to maintain connectivity between their seasonal ranges. Our results identify the mechanisms (e.g. detouring, increased movement rates, reduced stopover use) by which semi-permeable barriers affect the functionality of ungulate migration routes and emphasize that the management of semi-permeable barriers may play a key role in the conservation of migratory ungulate populations.

Estimating habitat selection when GPS fix success is less than 100

Inferences about habitat selection by animals derived from sequences of relocations obtained with global positioning system (GPS) collars can be influenced by GPS fix success. Environmental factors such as dense canopy cover or rugged terrain can reduce GPS fix success, making subsequent modeling problematic if fix success depends on the selected habitat. Ignoring failed fix attempts may affect estimates of model coefficients and lead to incorrect conclusions about habitat selection. Here, we present a habitat selection model that accounts for missing locations due to habitat-induced data losses, called a resource selection function (RSF) for GPS fix success. The models formulation is similar to adjusting estimates of probability of occupancy when detection is less than 100% in patch occupancy sampling. We demonstrate use of the model with GPS data collected from an adult female mule deer (Odocoileus hemionus) and discuss how to analyze data from multiple animals. In the simulations presented, our habitat selection model was generally unbiased for GPS data sets missing up to 50% of the locations.

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.

Large herbivores surf waves of green-up during spring

The green wave hypothesis (GWH) states that migrating animals should track or ‘surf’ high-quality forage at the leading edge of spring green-up. To index such high-quality forage, recent work proposed the instantaneous rate of green-up (IRG), i.e. rate of change in the normalized difference vegetation index over time. Despite this important advancement, no study has tested the assumption that herbivores select habitat patches at peak IRG. We evaluated this assumption using step selection functions parametrized with movement data during the green-up period from two populations each of bighorn sheep, mule deer, elk, moose and bison, totalling 463 individuals monitored 1–3 years from 2004 to 2014. Accounting for variables that typically influence habitat selection for each species, we found seven of 10 populations selected patches exhibiting high IRG—supporting the GWH. Nonetheless, large herbivores selected for the leading edge, trailing edge and crest of the IRG wave, indicating that other mechanisms (e.g. ruminant physiology) or measurement error inherent with satellite data affect selection for IRG. Our evaluation indicates that IRG is a useful tool for linking herbivore movement with plant phenology, paving the way for significant advancements in understanding how animals track resource quality that varies both spatially and temporally.