Weather and Sex Influence Migratory Behaviors and Habitat Selection in Mule Deer


Book Description

Seasonal migrations along elevational and latitudinal gradients allow a diversity of wildlife species to persist in the face of dramatic seasonal shifts in habitat quality. Recent work to better understand ungulate migration has produced a suite of analytical tools for identifying the mechanisms behind migration and quantifying associated behaviors. While environmental variables like weather and plant phenology are thought to be the main drivers of migration, emergent evidence suggests that anthropogenic disturbances, such as hunting, as well as intrinsic factors like age and sex can also influence migratory patterns. However, critical gaps in understanding of the influence of both the intrinsic and extrinsic factors that influence migratory behaviors remain common. We used three years (2016–2018) of GPS-collar data for a herd of migratory mule deer (Odocoileus hemionus) in south-central Wyoming to compare key migratory behaviors between sexes, identify factors that influence the timing of migration, and evaluate habitat selection in and around the autumn hunting season. Overall, migratory behaviors were sex specific. Both sexes showed high-fidelity to their migratory routes, but route fidelity was more variable in males. Males also migrated 30% farther on average, spent 14% more time on stopover sites, and took twice as long in spring and 44% longer in autumn to complete migration. Additionally, we found that weather, mainly precipitation events (i.e., snowfall), not hunting disturbance, was the main driver of autumn migration, increasing the likelihood of migration in both males and females by 14%. Our findings suggest that response to disturbance from hunting season is sex-specific. The hunted sex (males) more intensively sought refugia, whereas the less-hunted sex continued to use areas with good forage. Overall, our findings contribute to a broader understanding of migration ecology and the factors that influence migration and habitat selection of wild ungulates in western landscapes.







Advantages of Habitat Selection and Sexual Segregation in Mule and White-tailed Deer


Book Description

I studied sexual segregation in mule deer (Odocoileus hemionus) and white-tailed deer (Q. virginianus) in different environments and at different population densities to test the hypothesis that sexual segregation occurs in ungulates as the result of different reproductive strategies; females select habitat and behave in manners primarily designed to promote offspring survival, while adult males act primarily to maximize energy stores prior to the rut. Data collected for both species were consistent with this hypothesis. Mule deer segregated both socially and spatially. Compared to males, female mule deer used areas that were closer to water, supported superior browse species, and provided greater security for offspring from coyotes. Locally available resources enabled females to minimize movements, the apparent consequence of which was the depletion of forb biomass, the major diet class for both sexes. Males ranged further than females, a strategy that provided a diet high in forbs and high in quality. The occurrence of mixed-sex groups in highly productive areas suggested that males responded to forage availability and not females perse, and that the effects of localized grazing pressure by females may competitively exclude males from areas. Sexual segregation was not maintained by intersexual aggression in either mule or white-tailed deer. White tailed deer segregated socially, but demonstrated broad spatial overlap. During the peak of segregation (June-October), male behaviors were consistent with a pre-rut energy saving and forage seeking strategy; males used larger areas and made longer short term movements than females, but were less active overall and were relatively more active at night. Females were less mobile and foraged in smaller areas than males, particularly during fawn rearing periods (June-October). I also tested the hypothesis that male white-tailed deer segregate and range widely to scout potential mates and rivals. Based upon comparisons of areas used by radio-collared males and females during the peak periods of segregation (June-September) with areas used during the rut, this hypothesis was rejected.




Wheat Vs. Wild


Book Description

Converting wildlife habitat to agricultural monoculture is one of the greatest drivers of habitat loss and can be a severe threat to many wildlife species. However, because mule deer (Odocoileus hemionus) typically live in rugged, less arable landscapes, little is known about how they use habitat fragmented by agriculture. Using Global Positioning System (GPS) data from a partially migratory herd of 62 adult female mule deer from 2018-2022, we examined migration strategies and modeled habitat selection at three spatial scales in southeastern Washington, which is a mosaic of mostly privately owned cereal grain agriculture, shrubland, grassland, and restored cropland in the Conservation Reserve Program (CRP). A minority of the mule deer population was migratory (30%), with relatively short and fast migrations. During migration, they primarily selected for shrubland and grassland in both fall and spring. Mule deer used various components of the landscape throughout the year but selected grassland as their highest-ranked habitat year-round when selecting for home ranges within the study area (second order) and in winter when selecting habitat within their home ranges (third order). During summer, mule deer focused on shrubland and forested habitats within their home range, likely because it provided increased security and thermal cover for fawn-rearing. In the winter, they selected agricultural habitats to a greater degree, which might reflect the growing winter wheat (Triticum aestivum) providing more nutrition during that time. Survival from birth to recruitment into the adult population can greatly influence population dynamics of wild ungulates like mule deer and is affected by both nutrition and cover found in high-quality habitats. We used the GPS data from our population to create resource selection function models during parturition and early fawn-rearing and for fawn bed site characteristics (fourth order). We searched for fawns and fawn bed sites in 2021 and 2022 and used a parturition model to predict birthing events of collared females for which we were unable to find fawns in 2018-2022. For parturition, fawn-rearing habitat, and fawn bed sites, vegetation types that provided more vertical structure, including shrubs and trees, were consistently selected and agriculture avoided. Both intact (i.e., never plowed) shrubland and CRP shrubland were equally selected. These vegetation types provided vertical and horizontal concealment cover for fawns that were also important characteristics of fawn bed site selection. Our research suggests that to promote sustainable populations of mule deer in a landscape dominated by agriculture, managers and landowners could increase the prevalence of quality shrubs and trees where possible to improve habitat and continue support for the CRP program that provides economic incentive to restore croplands for the benefit of wildlife like mule deer.




Population Ecology and Summer Habitat Selection of Mule Deer in the White Mountains


Book Description

Changes in landscape composition have the potential to negatively influence animal populations through shifts in dominant plant communities, loss of important forage items, or changes in structural components of habitat. In the western United States, expansion of woodland vegetation into shrub dominated communities is of concern, particularly with regard to animal populations reliant on robust sagebrush and shrub vegetation. Once established, trees can out-compete shrubs and herbaceous plants resulting in declines in abundance and diversity of shrub-forb vegetation, which female mule deer are reliant on during summer months to meet nutritional demands and to provide hiding cover for young. As a result, shifts in the distribution of pinyon-juniper woodland and increases in tree densities could negatively affect mule deer population. The study had two primary objectives, (1) to determine summer habitat composition of female mule deer in the White Mountains of California and eastern Nevada, and assess implications of pinyon-juniper expansion on habitat availability, and (2) evaluate the status of the population relative to nutritional carrying capacity and determined the influence of habitat and precipitation on demographic rates. I used mixed-effects logistic regression to model summer resource selection and demographic rates of female mule deer from 2005 to 2008. Summer resource selection was modeled at two spatial scales and among three behavioral periods, related to foraging, resting, and parturition. Summer habitat consisted of sites with high productivity, greater shrub abundance, and greater proximity to riparian areas. Deer avoided high levels of tree cover at all spatial and temporal scales, but they selected areas with low to intermediate tree cover during resting periods and during parturition. Moreover, mule deer avoided areas of productive shrub-forb vegetation (riparian and shrub NDVI), when surrounded by stands of high level pinyon-juniper cover, otherwise those vegetation types were strongly selected. During parturition female mule deer selected habitat that maximized hiding cover for newborns (greater shrub densities and structural cover), while still providing foraging opportunities (greater NDVI and shrub cover). Females underutilized certain areas that contained optimal forage such as riparian corridors, high AET sites, higher elevation shrub communities, and selected areas with low to moderate tree cover, suggesting some trade-off between minimizing predation risk for offspring, and maximizing foraging opportunities. Demographic rates (body condition, survival, fetal rates, and index of recruitment) of female mule deer were sensitive to changes in resource availability resulting from variation in precipitation or habitat composition and suggestive of a population regulated to a greater degree by bottom-up processes, and likely nearing nutritional carrying capacity. Moreover, I identified a strong negative effect of pinyon-juniper cover on annual survival, only during periods of drought, otherwise individuals were able to maintain relatively high survival regardless of habitat composition. These results suggest that in productive years mule deer are able to inhabit areas of varying levels of pinyon-juniper cover with little effect on survival, and only during the drought years are negative effects evident. Results from this study emphasize the importance of productive shrub and forb vegetation to mule deer inhabiting semi-arid regions. Maintaining areas with low-to-intermediate tree cover, where there is still abundant shrub understory and sufficient concealment cover, may be beneficial to mule deer populations. Nevertheless, the strong influence of resource availability on the population suggests that conversion of sagebrush-steppe communities into large stands of PJ dominated woodlands would likely reduce the quality and abundance of available habitat for mule deer in the Great Basin.




Sexual Segregation in Vertebrates


Book Description

Males and females of many species can, and do, live separately for long periods of time. This sexual segregation is widespread and can be on social, spatial or habitat scales. An understanding of sexual segregation is important in the explanation of life history and social preference, population dynamics and the conservation of rare species. Sexual Segregation in Vertebrates explores the reasons why this behaviour has evolved and what factors contribute to it.




Mule and Black-tailed Deer of North America


Book Description

Developed in co-operation with U.S. Department of Agriculture, Forest Service.




Habitat Use and Migration Ecology of Mule Deer in Developing Gas Fields of Western Wyoming


Book Description

Increased levels of energy development across the intermountain West have created a variety of wildlife and habitat management concerns. Because many of the energy resources in the region occur in shrub-dominated basins (e.g., Powder River, Piceance, Great Divide, and Green River basins), management concerns have focused on native shrub communities and associated species, including mule deer (Odocoileus hemionus). Two of the more pressing concerns are how mule deer respond when critical habitats (e.g., winter range) are impacted by development and how their migration routes can be identified and prioritized for conservation. To address the first, I examined how three types of natural gas well pads with varying levels of vehicle traffic influenced the winter habitat selection patterns of mule deer in western Wyoming. My results showed that mule deer avoided all types of well pads and selected areas further from well pads that received high levels of traffic. Accordingly, impacts to mule deer could likely be reduced through technology and planning that minimizes the number of well pads and amount of human activity associated with them. To address the migration concerns, I developed a quantitative framework that uses global positioning system (GPS) data and the Brownian bridge movement model (BBMM) to: (1) provide a probabilistic estimate of the migration routes of a sampled population, (2) distinguish between route segments that function as stopover sites versus those used primarily as movement corridors, and (3) prioritize routes for conservation based upon the proportion of the sampled population that uses them. 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. These 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. Although stopovers appeared to be a prominent feature of mule deer migration routes, the explicit study of stopovers (i.e., stopover ecology) has been limited to avian species. To assess whether stopover ecology was relevant to mule deer, I again used fine-scale GPS data and BBMMs to quantify a suite of stopover characteristics and examine the ecological role of stopovers in the seasonal migrations of mule deer. Mule deer utilized a series of stopover sites in both spring and fall migrations, across a range of migration distances (18-144 km). Overall, mule deer used 1.9 and 1.5 stopovers for every 10 km increase in migration distance during spring and fall migrations, respectively. Stopovers had higher quality forage compared to movement corridors, and forage quality increased with elevation, presumably because of delayed phenology along the altitudinal migration route. Stopovers likely play a key role in the migration strategy of mule deer by allowing them to migrate in concert with vegetative phenology and optimize their foraging during migration. My results suggest stopovers were a critical component in the altitudinal migrations of mule deer and that conservation of stopover sites may improve efforts aimed at sustaining migratory mule deer populations.




Near and Deer


Book Description

Outdoor recreation extends human influence on landscapes beyond built environments but is often thought to be compatible with wildlife conservation. Human capability as a highly efficient predator creates a strong selective force on wildlife, analogous to natural predation risk, regardless of trophic level. Behavioral responses have been observed across taxa but linking these changes in behavior to changes at the population-level represents an important step in understanding and mitigating the impact of our everyday presence on ecosystems. Here we present results of the habitat selection exhibited by ten mule deer (Odocoileus hemionus) in response to habitat and human factors and the possible influences on density. Mule deer abundance is generally declining throughout their range and disturbance on winter range presents a possible limiting factor. Winter range is geographically constrained, has increased movement costs associated with snow, and reduced forage quality. Pinyon-juniper forest on winter range has been managed to provide better habitat for ungulates. Disturbance associated with human activity could further limit deer winter habitat. Mule deer on our study area show differential habitat use and movement characteristics with time of day, and intensity of human disturbance. Habitat selection for mule deer was modelled using integrated step selection analysis (iSSA). Mule deer increasingly selected for forest land cover as human presence increased. Deer showed different movement responses with increasing human presence depending on the time of day. Changes to habitat selection, and movement patterns have important implications for habitat management of winter range and potential costs to deer populations. Pairing our movement data with camera data, we found evidence for a low density of mule deer on our winter range compared to winter ranges across Colorado possibly caused by human recreation. Low densities of mule deer has implications for the role density-dependent effects play in modifying the impact of recreation. Our study provides critical information on the role human recreation plays in modifying the effectiveness of habitat management on deer winter ranges and possible scenarios this may scale up to impact mule deer populations.




Movements, Seasonal Home Ranges, and Resource Selection of Mule Deer on Navajo Nation Lands


Book Description

Mule deer (Odocoileus hemionus) populations have declined throughout their range in the western United States since the 1980s. Habitat loss, overgrazing, disease, and predation contribute to the decline of mule deer populations. Navajo Nation, the largest federally recognized Indian tribe in the United States, encompassing 71,000 km2 in New Mexico, Arizona, and Utah, has experienced a 49% decline in mule deer over the past decade. Given knowledge of space use is an important component to recovery plans, my objectives were to (1) classify each deer as a migrant, resident, disperser, or nomad; (2) determine dates and durations of deer classified as migrants including means and ranges of spring and fall migration; (3) quantify distances traveled during spring and fall migration; (4) estimate sizes of seasonal home ranges and core ranges for migratory mule deer; and (5) develop resource selection functions. GPS collars were placed on 99 mule deer (79 F, 20 M) during 2018-2020. Movements were analyzed using net-squared displacement for individuals with >6 mo of data. Movement trajectories (n=108) from 67 unique mule deer were analyzed to determine whether they were migrants or non-migrants. An ANOVA was performed to determine whether sex, season (i.e., spring or fall), strategy (i.e., short-distance or long-distance migrant), or an interaction between migration duration and migration distance. Seasonal home ranges were defined using 95% kernel density estimates (KDE), and core ranges with 50% KDE. An ANOVA was performed to determine whether sex, migration strategy (i.e., short-distance or long-distance), or season (i.e., winter or summer) affected seasonal home range and core range size. I modeled third-order resource selection functions (RSF) following a use-availability design. Seventy-four percent (n=50) of mule deer were long-distance migrants, 18% (n=12) were short-distance migrants, and 6% (n=5) were non-migrants. Minimum, maximum, and mean distances traveled during migration were 2.6 km, 68.3 km, and 17.7 ± 1.1 km, respectively. Seasonal home ranges, core range, and resource selection were quantified for 84 (15 M, 69 F) trajectories from 63 mule deer. Mean female and male summer home ranges were 9.1 ± 11.1 km2 and 7.4 ± 6.6 km2, respectively, and mean female and male winter home ranges were 15.6 ± 21.5 km2 and 16.1 ± 7.5 km2, respectively. Season had a significant effect on home range size (F1,158=28.02, P