Influence of interspecific competition on mule deer birthing and rearing site selection

Ungulates often alter behavior and space use in response to interspecific competition. Despite observable changes in behavior caused by competitive interactions, research describing the effects of competition on survival or growth is lacking. We used spatial modeling to determine if habitat use by female mule deer (Odocoileus hemionus) was affected by other ungulate species prior to, during, and after parturition. We conducted our study in the Book Cliffs region of eastern Utah, USA, during 2019 and 2020. We used resource selection function (RSF) analysis to model space use of 4 ungulate species that potentially competed with mule deer: bison (Bos bison), cattle, elk (Cervus canadensis), and feral horses. We incorporated RSF models for competing species into a random forest analysis to determine if space use by mule deer was influenced by these other ungulate species. We used survival and growth data from neonate mule deer to directly assess potential negative effects of other ungulates. Habitat use by elk was an important variable in predicting use locations of mule deer during birthing and rearing. The relationship was positive, suggesting interference competition was not occurring. Survival of neonate mule deer increased as the probability of use by elk increased (hazard ratio = 0.185 ± 0.497 [SE]). Further, probability of use by elk in rearing habitat had no influence on growth of neonate mule deer from birth to 6 months of age, suggesting that exploitative competition was not occurring.     

Temporal Overlap Among Feral Horses,Cattle, and Native Ungulates at Water Sources

Feral horse (Equus ferus caballus) populations on public rangelands in the western United States threaten forage production for livestock and wildlife habitat. Interference competition between feral horses and heterospecifics at watering sources can have negative effects on livestock and wildlife. Researchers have documented altered timing and behavior of wild ungulates at water sources when horses were present. The few studies examining these interactions have infrequently occurred within areas specifically managed for feral equids and have not occurred in sites with cattle. We used motion-sensitive cameras at 8 watering sources to document watering activity patterns and construct indices of temporal overlap among feral horses, cattle, elk (Cervus canadensis), mule deer (Odocoileus hemionus), and pronghorn (Antilocapra americana) within the Adobe Town Herd Management Area in southern Wyoming, USA, between June and September 2018 and 2019. Feral horses, cattle, and pronghorn exhibited a high degree of temporal overlap (>79%) in water use, with feral horses and pronghorn exhibiting the highest estimated percent overlap (88.1%, 95% CI = 86.5–89.6%). Mule deer and elk watering activity also overlapped with horses and cattle but to a lesser degree (<55%). Feral horses spent a mean of 16.7 ± 30.5 (SD) minutes during a watering event and were present at a given water source on average 4.5 ± 6.3% and up to 34.9% of the day, which is less than reported in previous studies. Cattle spent on average 23.5 ± 44.9 minutes during a watering event, and were present on average 4.2 ± 7.7% and up to 42.4% of the day at a single water source. Results of generalized linear mixed-effects models indicated that number of conspecifics was the strongest predictor of visit duration for pronghorn and horses; hour of the day and group size of heterospecifics were informative, but less important, variables. There was no difference in peak visitation time for any species between sites of high versus low horse or cattle use. Despite temporal overlap, we did not find evidence of interference competition between feral horses, cattle, and pronghorn. We recommend future examination of interference competition and its biological consequences between introduced and native ungulates at water sources of varying size across sites, equid population levels, and livestock stocking rates. © 2020 The Wildlife Society.     

Differential prey use by male and female cougars in Washington

Male and female predators are often assumed to have the same effects on prey. Because of differences in body size and behavior, however, male and female predators may use different species, sexes, and ages of prey, which could have important implications for wildlife conservation and management. We tested for differential prey use by male and female cougars (Puma concolor) from 2003 to 2008 in Washington State. We predicted that male cougars would kill a greater proportion of larger and older prey (i.e., adult elk [Cervus elaphus]), whereas females would kill smaller and younger prey (i.e., elk calves, mule deer [Odocoileus hemionus]). We marked cougars with Global Positioning System (GPS) radio collars and investigated 436 predation sites. We located prey remains at 345 sites from 9 male and 9 female cougars. We detected 184 mule deer, 142 elk, and 17 remains from 4 other species. We used log-linear modeling to detect differences in species and age of prey killed among cougar reproductive classes. Solitary females and females with dependent offspring killed more mule deer than elk (143 vs. 83, P < 0.01), whereas males killed more elk than mule deer (59 vs. 41, P < 0.01). Proportionately, males killed 4 times more adult elk than did females (24% vs. 6% of kills) and females killed 2 times more adult mule deer than did males (26% vs. 15% of kills). Managers should consider the effects of sex of predator in conservation and management of ungulates, particularly when managing for sensitive species. © 2011 The Wildlife Society.     

A Simple Solar Radiation Index for Wildlife Habitat Studies

Abstract: Solar radiation is a potentially important covariate in many wildlife habitat studies, but it is typically addressed only indirectly, using problematic surrogates like aspect or hillshade. We devised a simple solar radiation index (SRI) that combines readily available information about aspect, slope, and latitude. Our SRI is proportional to the amount of extraterrestrial solar radiation theoretically striking an arbitrarily oriented surface during the hour surrounding solar noon on the equinox. Because it derives from first geometric principles and is linearly distributed, SRI offers clear advantages over aspect-based surrogates. The SRI also is superior to hillshade, which we found to be sometimes imprecise and ill-behaved. To illustrate application of our SRI, we assessed niche separation among 3 ungulate species along a single environmental axis, solar radiation, on the northern Yellowstone winter range. We detected no difference between the niches occupied by bighorn sheep (Ovis canadensis) and elk (Cervus elaphus; P = 0.104), but found that mule deer (Odocoileus hemionus) tended to use areas receiving more solar radiation than either of the other species (P < 0.001). Overall, our SRI provides a useful metric that can reduce noise, improve interpretability, and increase parsimony in wildlife habitat models containing a solar radiation component.     

Revisions of Rump Fat and Body Scoring Indices for Deer,Elk, and Moose

Abstract: Because they do not require sacrificing animals, body condition scores (BCS), thickness of rump fat (MAXFAT), and other similar predictors of body fat have advanced estimating nutritional condition of ungulates and their use has proliferated in North America in the last decade. However, initial testing of these predictors was too limited to assess their reliability among diverse habitats, ecotypes, subspecies, and populations across the continent. With data collected from mule deer (Odocoileus hemionus), elk (Cervus elaphus), and moose (Alces alces) during initial model development and data collected subsequently from free-ranging mule deer and elk herds across much of the western United States, we evaluated reliability across a broader range of conditions than were initially available. First, to more rigorously test reliability of the MAXFAT index, we evaluated its robustness across the 3 species, using an allometric scaling function to adjust for differences in animal size. We then evaluated MAXFAT, rump body condition score (rBCS), rLIVINDEX (an arithmetic combination of MAXFAT and rBCS), and our new allometrically scaled rump-fat thickness index using data from 815 free-ranging female Roosevelt and Rocky Mountain elk (C. e. roosevelti and C. e. nelsoni) from 19 populations encompassing 4 geographic regions and 250 free-ranging female mule deer from 7 populations and 2 regions. We tested for effects of subspecies, geographic region, and captive versus free-ranging existence. Rump-fat thickness, when scaled allometrically with body mass, was related to ingesta-free body fat over a 38–522-kg range of body mass (r² = 0.87; P < 0.001), indicating the technique is remarkably robust among at least the 3 cervid species of our analysis. However, we found an underscoring bias with the rBCS for elk that had >12% body fat. This bias translated into a difference between subspecies, because Rocky Mountain elk tended to be fatter than Roosevelt elk in our sample. Effects of observer error with the rBCS also existed for mule deer with moderate to high levels of body fat, and deer body size significantly affected accuracy of the MAXFAT predictor. Our analyses confirm robustness of the rump-fat index for these 3 species but highlight the potential for bias due to differences in body size and to observer error with BCS scoring. We present alternative LIVINDEX equations where potential bias from rBCS and bias due to body size are eliminated or reduced. These modifications improve the accuracy of estimating body fat for projects intended to monitor nutritional status of herds or to evaluate nutrition's influence on population demographics.     

Use of Bait to Increase Archery Deer Harvest in an Urban–Suburban Landscape

Abstract: Hunting is the primary tool for managing white-tailed deer (Odocoileus virginianus) populations. Effectiveness of hunting in suburban areas may be reduced due to limited hunter access to small properties, firearms-discharge laws, and public safety concerns. In Connecticut, USA, hunting over bait on private land was recently legalized to increase harvest opportunities. Our objective was to assess bow-hunter willingness to use bait and effects of bait type, hunter disturbance, time spent hunting, and property size on deer-harvest potential in a suburban landscape. We mailed a prebaiting survey in February 2002 and a postbaiting survey in February 2004 to the same group of hunters. Hunters using bait were more successful and harvested more deer than hunters using no bait. Hunters using bait on small properties observed similar numbers of deer within shooting range as hunters using bait on larger properties. Hunters using bait met their venison needs, whereas hunters using no bait did not meet their needs. Resource managers should implement strategies that increase hunter success when developing urban deer-management programs for communities.     

Space Use and Survival of White-Tailed Deer in an Exurban Landscape

Abstract: Exurban development is nonmetropolitan, residential development characterized by a human population density and average property size intermediate between suburban and rural areas. Although growth in exurban areas is outpacing that of urban, suburban, or rural landscapes, studies of deer (Odocoileus spp.) ecology in exurban areas are nonexistent. During 2003–2005, we studied space use (i.e., seasonal home-range and core-area size and habitat use relative to human dwellings) and survival of 43 female white-tailed deer (O. virginianus) in an exurban setting near Carbondale, Illinois. Deer had larger home ranges than most suburban deer populations and generally smaller home ranges than rural deer populations. When we analytically controlled for habitat use, deer exhibited a subtle avoidance of human dwellings, especially during the fawning season. The annual survival rate was among the highest reported in the literature at 0.872 (SE = 0.048). Only 5 deer (cause-specific mortality rate = 0.091) were harvested by hunters, indicating major obstacles for wildlife managers when attempting to manage deer in exurban areas using traditional hunter harvest.     

Comparison of Effects of Shed Antler Hunting and Helicopter Surveys on Ungulate Movements and Space Use

Shed antler hunting (i.e., collecting cast cervid antlers) has increased in popularity during the past decade, but little is known about how this recreational activity affects ungulate movements and space use. We placed geographic positioning system (GPS)-collars on 133 female and male bighorn sheep (Ovis canadensis), bison (Bison bison), and mule deer (Odocoileus hemionus) to quantify their movements and space use during shed antler hunts compared with those behaviors during helicopter surveys in Utah, USA, from 2012 to 2015. For each species, we calculated means and 95% confidence intervals for distance moved during 90-minute segments (16 points/day) pre-event (control, 7 consecutive days prior to event), event (1–2 days), and post-event (7 consecutive days after event) for shed antler hunts and helicopter surveys. We also compared use of space for each species during these events. Female bighorn sheep did not increase distance moved or substantially change space use during shed antler hunts and helicopter surveys. Male bighorn sheep increased distance moved 41% on average during shed antler hunts and by 2.02 times during helicopter surveys but did not change space use during those events. Female bison increased distance moved 15% on average during shed antler hunts and 30% during helicopter surveys. Mule deer increased distance moved and altered space use the most during shed antler hunts; females increased distance moved 97%, and 54% of females moved a mean distance of 742 ± 642 (SD) m (range = 9–3,778 m) outside of their home ranges during those hunts for a mean of 9.2 ± 9.4 hours (range = 1.5 to 41 hr). Male mule deer increased distance moved by 2.10 times on average during shed antler hunts, and 82% of males moved a mean distance of 1,264 ± 732 m (range = 131–3,637 m) outside of their home ranges during those hunts for a mean of 12.6 ± 7.6 hours (range = 4.5–33 hr). Our results provide timely information about how legal shed antler hunting affects movements and space use of female and male ungulates, especially mule deer, and can guide the conservation of ungulate populations and their habitat. © 2021 The Wildlife Society.     

Fence-Line Contact Between Wild and Farmed Cervids in Colorado: Potential for Disease Transmission

Abstract: Direct and indirect contact between wild and farmed cervids along perimeter fences may play a role in transmission of diseases like chronic wasting disease (CWD), but no studies have quantified such interactions. At 9 high-fenced commercial elk (Cervus elaphus) farms in Colorado, USA, during October 2003 to January 2005, we used animal-activated video to estimate rates of fence-line use by wild cervids, rates of direct contact between farmed and wild cervids, and probability of direct contact when wild cervids were present. We recorded 8-fold-more wild elk per unit time than mule deer (Odocoileus hemionus) at fence lines. Depending on site, we recorded 0.66 to 46.90 wild elk per 1,000 hours of camera monitoring. We documented 77 interactions between wild and farmed elk involving naso-oral contact and no contact between wild mule deer and farmed elk. Rate of direct contact ranged from 0.00 to 1.92 direct contacts per 1,000 hours of camera monitoring among sites. Given recorded presence of wild elk, estimated probability of observing direct contact during a 2-minute video recording ranged from 0.00 to 0.11 among sites. Risk of direct contact was about 3.5 times greater for single woven-wire fence compared with offset electric fence attached to a single woven-wire fence. We observed no direct contact through double woven-wire fences. Because little is currently known about infection rates associated with infection mechanisms, we cannot infer a level of CWD infection risk from our results, but some form of double fencing should reduce potential for direct and indirect transmission of disease into or out of elk farms.     

Effects of Season and Scale on Response of Elk and Mule Deer to Habitat Manipulation

Abstract: Manipulation of forest habitat via mechanical thinning or prescribed fire has become increasingly common across western North America. Nevertheless, empirical research on effects of those activities on wildlife is limited, although prescribed fire in particular often is assumed to benefit large herbivores. We evaluated effects of season and spatial scale on response of Rocky Mountain elk (Cervus elaphus) and mule deer (Odocoileus hemionus) to experimental habitat manipulation at the Starkey Experimental Forest and Range in northeastern Oregon, USA. From 2001 to 2003, 26 densely stocked stands of true fir (Abies spp.) and Douglas-fir (Pseudotsuga menziesii) were thinned and burned whereas 27 similar stands were left untreated to serve as experimental controls. We used location data for elk and mule deer collected during spring (1 Apr-14 Jun) and summer (15 Jun-31 Aug) of 1999–2006 to compare use of treated and untreated stands and to model effects of environmental covariates on use of treated stands. In spring, elk selected burned stands and avoided control stands within the study area (second-order selection; large scale). Within home ranges (third-order selection; small scale), however, elk did not exhibit selection. In addition, selection of treatment stands by elk in spring was not strongly related to environmental covariates. Conversely, in summer elk selected control stands and either avoided or used burned stands proportional to their availability at the large scale; patterns of space use within home ranges were similar to those observed in spring. Use of treatment stands by elk in summer was related to topography, proximity to roads, stand size and shape, and presence of cattle, and a model of stand use explained 50% of variation in selection ratios. Patterns of stand use by mule deer did not change following habitat manipulation, and mule deer avoided or used all stand types proportional to their availability across seasons and scales. In systems similar to Starkey, manipulating forest habitat with prescribed fire might be of greater benefit to elk than mule deer where these species are sympatric, and thus maintaining a mixture of burned and unburned (late successional) habitat might provide better long-term foraging opportunities for both species than would burning a large proportion of a landscape.     

Effect of archer density on elk pregnancy rates and conception dates

Archery hunting in Oregon has increased dramatically over the past 2 decades. At the same time, spring juvenile to adult female ratios of Rocky Mountain elk (Cervus elaphus) have been declining. This has raised concern that archery seasons may be disrupting elk breeding and contributing to the decline in recruitment. Two mechanisms could contribute to reduced juvenile:female ratios: 1) reduced pregnancy rates, and 2) delayed conception dates because of human disturbance during the rut. We varied the number of archery hunters at the Starkey Experimental Forest and Range over 13 years to evaluate effects of archer density on reproduction of elk. Archer densities were maintained at high densities during 4 years , low densities during 3 years , and no archers during 6 years. We determined pregnancy status, age, kidney fat index (KFI), lactation status, and fetus conception dates for 622 female elk harvested in December. We found pregnancy rate differences of 0.105, 0.080, and 0.021 between high and no archer density years (P = 0.004), high and low archer density years (P = 0.054), and low and no archer density years (P = 0.616), respectively. Conception dates were 4 days later for high archer density compared to low archer density (P = 0.006), but did not differ between high and no archer years (2 days; P = 0.136) or between low and no archer years (2 days; P = 0.108). We compared generalized linear model estimates of pregnancy rates and determined pregnancy rates for 28% of the lactating female elk to be affected by high archer density, whereas archer densities had no significant affect on pregnancy rate estimates for non-lactating females. We found no difference in conception dates among archer densities when comparing model estimates. Our results suggest that archer density and its interaction with nutritional condition of elk influence pregnancy rates of lactating females with low KFI levels; however, the effect of archer density alone does not explain the magnitude of decline in juvenile to female ratios observed in Oregon. © 2012 The Wildlife Society.     

Influence of Summer and Autumn Nutrition on Body Condition and Reproduction in Lactating Mule Deer

ABSTRACT Recent work suggests that availability and quality of forage in late summer and early autumn, a time when female ungulates face multiple energetic demands, is critical to reproduction in wild ungulates. Therefore, we examined direct links between nutritional quality of diets, body condition, and reproduction of lactating mule deer. Using captive mule deer, we tested the hypothesis that females consuming diets with lower digestible energy (DE; kJ/g) would have lower DE intake rates (DEI; MJ/day), have less body fat and muscle, have later estrus cycles, and have lower pregnancy and twinning rates. Deer fed lower DE diets had lower DEI during summer and autumn. In turn, deer with lower DEI, regardless of diet DE, had lower body mass, body fat, and muscle thickness. When nutritional quality of diets began to decline earlier in the summer, relationships between food quality, DEI, and body condition were stronger. Although DEI did not influence estrus date for deer that became pregnant before 21 December, deer with lower DEI had a lower probability of becoming pregnant and had a lower probability of producing twins. Measures of body condition in October (i.e., body mass, body fat, and muscle depth) predicted pregnancy and twinning rates in mule deer. Serum concentration of hormones leptin and Insulin Growth Factor 1 were not good predictors of body condition or reproduction. These findings suggest that managers concerned with productivity of mule deer populations should consider focusing on assessing and improving quality of forage available in summer and autumn.     

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.     

Effects of large-scale removal of coyotes on pronghorn and mule deer productivity and abundance

We tested the hypothesis that predation by coyotes (Canis latrans) impacts pronghorn (Antilocapra americana) and mule deer (Odocoileus hemionus) populations. We did so by examining the effects of coyote removal on pronghorn and mule deer populations within 12 large areas (>10,500 km²) located in Wyoming and Utah during 2007 and 2008. Pronghorn productivity (fawn to adult female ratio) and abundance were positively correlated with the number of coyotes removed and removal effort (hours spent hunting coyotes from aircraft) although the correlation between pronghorn productivity and removal effort was not statistically significant (P = 0.08). Mule deer productivity and abundance were not correlated with either the number of coyotes removed or removal effort. Coyote removal conducted during the winter and spring provided greater benefit than removals conducted during the prior fall or summer. Our results suggest that coyote removal conducted over large areas increases fawn survival and abundance of pronghorn but not mule deer. © 2011 The Wildlife Society.     

Influence of Landscape Composition and Structure on Antler Size of White-Tailed Deer

Abstract: Cause for spatial variation in phenotypic quality of white-tailed deer (Odocoileus virginianus) populations is of great interest to wildlife managers. Relating phenotypic variation of populations to large-scale land-use patterns may provide insight into why populations exhibit spatial variation and elucidate how management can influence population phenotype. We used an information-theoretic approach to relate average antler size of 203 deer populations to composition and structure of the habitat occupied by those populations. We used interspersion, edge, and diversity indices to represent habitat structure and percentage of area in vegetation types to represent habitat composition. Landscape composition was a better predictor of deer population antler size than was landscape structure. Percentages of the management unit in agriculture, pasture, and pine forest were variables commonly found in the region-specific set of best models. Model-averaged estimates of agriculture and pasture parameters were always positive and estimates of pine forest parameters were always negative, which suggests that land-use types that promote growth of early successional herbaceous plants will positively influence antler size and, most likely, body growth and reproduction of white-tailed deer populations. Conversely, our findings suggest landscapes dominated by pine forests did not provide optimal amounts of quality forages for white-tailed deer. Pine forest effects should be mitigated using a combination of increased harvest to lower deer density and silvicultural practices like thinning, prescribed burning, and selective herbicide applications that stimulate growth of high-quality forages beneath the forest canopy to improve deer phenotypic quality.     

Effects of Joint Space Use and Group Membership on Contact Rates Among White-Tailed Deer

Abstract: Establishment and spread of infectious diseases are controlled by the frequency of contacts among hosts. Although managers can estimate transmission coefficients from the relationship between disease prevalence and age or time, they may wish to quantify or compare contact rates before a disease is established or while it is at very low prevalence. Our objectives were to quantify direct and indirect contacts rates among white-tailed deer (Odocoileus virginianus) and to compare these measures of contact rate with simpler measures of joint space use. We deployed Global Positioning System (GPS) collars on 23 deer near Carbondale, Illinois, USA, from 2002 to 2005. We used location data from the GPS collars to measure pairwise rates of direct and indirect contact, based on a range of proximity criteria and time lags, as well as volume of intersection (VI) of kernel utilization distributions. We analyzed contact rates at a given distance criterion and time lag using mixed-model logistic regression. Direct contact rates increased with increasing VI and were higher in autumn—spring than in summer. After accounting for VI, the estimated odds of direct contact during autumn—spring periods were 5.0–22.1-fold greater (depending on the proximity criterion) for pairs of deer in the same social group than for between-group pairs, but for direct contacts during summer the within:between-group odds ratio did not differ significantly from 1. Indirect contact rates also increased with VI, but the effects of both season and pair-type were much smaller than for direct contacts and differed little as the time lag increased from 1–30 days. These results indicate that simple measures of joint space use are insufficient indices of direct contact because group membership can substantially increase contacts at a given level of joint space use. With indirect transmission, however, group membership had a much smaller influence after accounting for VI. Relationships between contact rates and season, VI, and pair-type were generally robust to changes in the proximity criterion defining a contact, and patterns of indirect contacts were affected little by the choice of time lag from 1–30 days. The use of GPS collars provides a framework for testing hypotheses about the form of contact networks among large mammals and comparing potential direct and indirect contact rates across gradients of ecological factors, such as population density or landscape configuration.