Using Vaginal Implant Transmitters to Aid in Capture of Mule Deer Neonates

Abstract: Estimating survival of the offspring of marked female ungulates has proven difficult in free-ranging populations yet could improve our understanding of factors that limit populations. We evaluated the feasibility and efficiency of capturing large samples (i.e., >80/yr) of neonate mule deer (Odocoileus hemionus) exclusively from free-ranging, marked adult females using vaginal implant transmitters (VITs, n = 154) and repeated locations of radiocollared females without VITs. We also evaluated the effectiveness of VITs, when used in conjunction with in utero fetal counts, for obtaining direct estimates of fetal survival. During 2003 and 2004, after we placed VIT batteries on a 12-hour duty cycle to lower electronic failure rates, the proportion that shed ≤ 3 days prepartum or during parturition was 0.623 (SE = 0.0456), and the proportion of VITs shed only during parturition was 0.447 (SE = 0.0468). Our neonate capture success rate was 0.880 (SE = 0.0359) from females with VITs shed ≤ 3 days prepartum or during parturition and 0.307 (SE = 0.0235) from radiocollared females without VITs or whose implant failed to function properly. Using a combination of techniques, we captured 275 neonates and found 21 stillborns during 2002-2004. We accounted for all fetuses at birth (i.e., live or stillborn) from 78 of the 147 females (0.531, SE = 0.0413) having winter fetal counts, and this rate was heavily dependent on VIT retention success. Deer that shed VITs prepartum were larger than deer that retained VITs to parturition, indicating a need to develop variable-sized VITs that may be fitted individually to deer in the field. We demonstrated that direct estimates of fetal and neonatal survival may be obtained from previously marked female mule deer in free-ranging populations, thus expanding opportunities for conducting field experiments. Survival estimates using VITs lacked bias that is typically associated with other neonate capture techniques. However, current vaginal implant failure rates and overall expense limit broad applicability of the technique.     

Re-Evaluating Neonatal-Age Models for Ungulates: Does Model Choice Affect Survival Estimates?

New-hoof growth is regarded as the most reliable metric for predicting age of newborn ungulates, but variation in estimated age among hoof-growth equations that have been developed may affect estimates of survival in staggered-entry models. We used known-age newborns to evaluate variation in age estimates among existing hoof-growth equations and to determine the consequences of that variation on survival estimates. During 2001–2009, we captured and radiocollared 174 newborn (≤24-hrs old) ungulates: 76 white-tailed deer (Odocoileus virginianus) in Minnesota and South Dakota, 61 mule deer (O. hemionus) in California, and 37 pronghorn (Antilocapra americana) in South Dakota. Estimated age of known-age newborns differed among hoof-growth models and varied by >15 days for white-tailed deer, >20 days for mule deer, and >10 days for pronghorn. Accuracy (i.e., the proportion of neonates assigned to the correct age) in aging newborns using published equations ranged from 0.0% to 39.4% in white-tailed deer, 0.0% to 3.3% in mule deer, and was 0.0% for pronghorns. Results of survival modeling indicated that variability in estimates of age-at-capture affected short-term estimates of survival (i.e., 30 days) for white-tailed deer and mule deer, and survival estimates over a longer time frame (i.e., 120 days) for mule deer. Conversely, survival estimates for pronghorn were not affected by estimates of age. Our analyses indicate that modeling survival in daily intervals is too fine a temporal scale when age-at-capture is unknown given the potential inaccuracies among equations used to estimate age of neonates. Instead, weekly survival intervals are more appropriate because most models accurately predicted ages within 1 week of the known age. Variation among results of neonatal-age models on short- and long-term estimates of survival for known-age young emphasizes the importance of selecting an appropriate hoof-growth equation and appropriately defining intervals (i.e., weekly versus daily) for estimating survival.     

The effect of terrain and female density on survival of neonatal white‐tailed deer and mule deer fawns

Juvenile survival is a highly variable life‐history trait that is critical to population growth. Antipredator tactics, including an animal's use of its physical and social environment, are critical to juvenile survival. Here, we tested the hypothesis that habitat and social characteristics influence coyote (Canis latrans) predation on white‐tailed deer (Odocoileus virginianus) and mule deer (O. hemionus) fawns in similar ways during the neonatal period. This would contrast to winter when the habitat and social characteristics that provide the most safety for each species differ. We monitored seven cohorts of white‐tailed deer and mule deer fawns at a grassland study site in Alberta, Canada. We used logistic regression and a model selection procedure to determine how habitat characteristics, climatic conditions, and female density influenced fawn survival during the first 8 weeks of life. Fawn survival improved after springs with productive vegetation (high integrated Normalized Difference Vegetation Index values). Fawns that used steeper terrain were more likely to survive. Fawns of both species had improved survival in years with higher densities of mule deer females, but not with higher densities of white‐tailed deer females, as predicted if they benefit from protection by mule deer. Our results suggest that topographical variation is a critical resource for neonates of many ungulate species, even species like white‐tailed deer that use more gentle terrain when older. Further, our results raise the possibility that neonatal white‐tailed fawns may benefit from associating with mule deer females, which may contribute to the expansion of white‐tailed deer into areas occupied by mule deer.     

Effectiveness of a redesigned vaginal implant transmitter in mule deer

Our understanding of factors that limit mule deer (Odocoileus hemionus) populations may be improved by evaluating neonatal survival as a function of dam characteristics under free-ranging conditions, which generally requires that both neonates and dams are radiocollared. The most viable technique facilitating capture of neonates from radiocollared adult females is use of vaginal implant transmitters (VITs). To date, VITs have allowed research opportunities that were not previously possible; however, VITs are often expelled from adult females prepartum, which limits their effectiveness. We redesigned an existing VIT manufactured by Advanced Telemetry Systems (ATS; Isanti, MN) by lengthening and widening wings used to retain the VIT in an adult female. Our objective was to increase VIT retention rates and thereby increase the likelihood of locating birth sites and newborn fawns. We placed the newly designed VITs in 59 adult female mule deer and evaluated the probability of retention to parturition and the probability of detecting newborn fawns. We also developed an equation for determining VIT sample size necessary to achieve a specified sample size of neonates. The probability of a VIT being retained until parturition was 0.766 (SE = 0.0605) and the probability of a VIT being retained to within 3 days of parturition was 0.894 (SE = 0.0441). In a similar study using the original VIT wings (Bishop et al. 2007 ), the probability of a VIT being retained until parturition was 0.447 (SE = 0.0468) and the probability of retention to within 3 days of parturition was 0.623 (SE = 0.0456). Thus, our design modification increased VIT retention to parturition by 0.319 (SE = 0.0765) and VIT retention to within 3 days of parturition by 0.271 (SE = 0.0634). Considering dams that retained VITs to within 3 days of parturition, the probability of detecting at least 1 neonate was 0.952 (SE = 0.0334) and the probability of detecting both fawns from twin litters was 0.588 (SE = 0.0827). We expended approximately 12 person-hours per detected neonate. As a guide for researchers planning future studies, we found that VIT sample size should approximately equal the targeted neonate sample size. Our study expands opportunities for conducting research that links adult female attributes to productivity and offspring survival in mule deer. © 2011 The Wildlife Society.     

Comparing Survival and Movements of Non-Urban and Urban Translocated Mule Deer

In many parts of North America, deer (Odocoileus spp.) have adapted to live in urban areas and are a source of negative human-wildlife interactions. Management strategies such as culling, immunocontraceptives, sterilization, and translocation have been implemented to manage urban deer populations. In the East Kootenay region of southern British Columbia, urban mule deer (Odocoileus hemionus) populations have been increasing, whereas non-urban mule deer populations have decreased. In 2014 a non-urban mule deer research project began in the area and in 2016 an urban deer translocation trial was approved in the same region. We fit 121 non-urban deer with global positioning system (GPS)-collars and translocated 135 urban mule deer to non-urban areas, of which 57 were fit with GPS-collars. We tested if annual survival between urban translocated (i.e., translocated) and non-urban deer differed, and if translocated deer survival increased in subsequent years after translocation. We also determined if age, body condition, release site, capture area and distance between capture and release sites affected translocated deer survival. We evaluated if translocated deer exhibited different movement behaviors than non-urban deer by comparing probability of migration, maximum net displacement, home range size, and probability of crossing a paved road. Finally, during our study we observed some translocated deer return to a municipal area after translocation and assessed if any covariates such as age, release site, or capture city could help predict this behavior. Annual survival of translocated deer was 0.48 and was significantly lower than survival of non-urban deer, which was 0.77. We observed 20 of 57 collared translocated deer return to a town after translocation. Translocated deer had larger net displacements and larger seasonal home range sizes than non-urban deer. Non-urban deer were more likely to migrate than translocated deer and crossed fewer paved roads than translocated deer. The management effectiveness of translocation to reduce urban deer densities is mixed because annual survival of translocated deer may be lower than may be acceptable to some stakeholders. Additionally, some translocated deer returned to an urban area, and the large distances traveled by deer after translocation may unintentionally spread disease. © 2020 The Wildlife Society.     

Separating Components of Variation in Survival of Mule Deer in Colorado

ABSTRACT Survival is an important parameter for understanding population dynamics of mule deer (Odocoileus hemionus) and other large herbivores. To understand long-term dynamics it is important to separate sampling and biological process variation in survival. Moreover, knowledge of correlations in survival across space and between young and adults can provide more informed predictions of survival in unsampled areas. We estimated survival of fawn, yearling, and adult mule deer from 4 spatially separated regions of Colorado, USA, from 1997 to 2008. We also estimated process variance in survival across time for each age and site using Markov chain Monte Carlo (MCMC) methods. Finally, we estimated correlations in survival among sites and ages with MCMC methods. Average winter fawn survival was 0.721 (SD = 0.024) for the 4 regions. Average winter adult female survival was 0.935 (SD = 0.007). Annual adult female survival ranged from 0.803 (SD = 0.017) to 0.900 (SD = 0.028) for the 4 regions, excluding hunting mortality. The correlation between fawn and adult female survival was high, 0.563 (SD = 0.253). Correlations in winter fawn survival were higher between populations at the same latitude than they were for populations to the north and south. We used survival estimates from our analysis to inform prior distributions for a Bayesian population dynamics model from one population in Colorado and compared that model to one with noninformative prior distributions. Population models including informative prior distributions based on our results performed better than those noninformative prior distributions on survival, providing more biologically defensible results when data were sparse. Knowledge of process distributions of survival can help wildlife managers better predict future population status and understand the likely range of survival rates.     

Effect of Net-Gun Capture on Survival of Mule Deer

Capture techniques to deploy radio-collars often risk mortality and injury to the animal. Capture-induced mortality can affect population sizes but also introduces bias in survival estimates based on data from captured animals. In recent years, a large-scale research and monitoring project in Utah, USA, has involved capturing and radio-collaring hundreds of mule deer (Odocoileus hemionus), a species of great interest in large parts of North America. Our objective was to investigate how the survival rates of these mule deer were affected by capture and handling. During winters of 2014–2018, an experienced capture crew net-gunned and fitted 1,805 animals with global positioning system (GPS)-collars. We estimated survival rates during the first 6 weeks after capture using Cox proportional hazard regression, and compared the survival rates of animals that were captured in a particular year to those of animals that were not captured but fitted with a GPS-collar in a previous year. We used a model selection framework to evaluate how long survival rates of captured animals were different from those of animals that were not captured. Our results indicated that weekly survival rates of captured animals were 0.985 ± 0.003 (SE), 0.988 ± 0.002, and 0.990 ± 0.001 in weeks 1, 2 and 3, respectively. Weekly survival rates of captured deer during weeks 4–6 were 0.993 ± 0.001, the same as those of deer that were not captured at the same time. Furthermore, post-capture survival rates were positively influenced by body size and negatively influenced by age. We conclude that the mortality resulting from helicopter capture was low but recommend comparing newly captured and previously captured individuals to examine what proportion of observed mortality is likely capture-related. © 2020 The Wildlife Society.     

Does small-perimeter fencing inhibit mule deer or pronghorn use of water developments?

Wildlife water development can be an important habitat management strategy in western North America for many species, including both pronghorn (Antilocapra americana) and mule deer (Odocoileus hemionus). In many areas, water developments are fenced (often with small-perimeter fencing) to exclude domestic livestock and feral horses. Small-perimeter exclosures could limit wild ungulate use of fenced water sources, as exclosures present a barrier pronghorn and mule deer must negotiate to gain access to fenced drinking water. To evaluate the hypothesis that exclosures limit wild ungulate access to water sources, we compared use (photo counts) of fenced versus unfenced water sources for both pronghorn and mule deer between June and October 2002–2008 in western Utah. We used model selection to identify an adequate distribution and best approximating model. We selected a zero-inflated negative binomial distribution for both pronghorn and mule deer photo counts. Both pronghorn and mule deer photo counts were positively associated with sampling time and average daily maximum temperature in top models. A fence effect was present in top models for both pronghorn and mule deer, but mule deer response to small-perimeter fencing was much more pronounced than pronghorn response. For mule deer, we estimated that presence of a fence around water developments reduced photo counts by a factor of 0.25. We suggest eliminating fencing of water developments whenever possible or fencing a big enough area around water sources to avoid inhibiting mule deer. More generally, our results provide additional evidence that water development design and placement influence wildlife use. Failure to account for species-specific preferences will limit effectiveness of management actions and could compromise research results. © 2011 The Wildlife Society.     

Survival of White-Tailed Deer Fawns in Southern Illinois

Abstract: Survival of white-tailed deer (Odocoileus virginianus) fawns has been quantified throughout much of North America. However, few studies have assessed the influence of intrinsic factors (e.g., fawn age and birth mass) and habitat on fawn survival. During 2002-2004, we captured and radiocollared 166 fawns in southern Illinois, USA, to estimate survival rates, determine causes of mortality, and identify factors influencing fawn survival. We used a known fates model in program MARK to estimate survival rates and compare explanatory models based on Akaike's Information Criterion corrected for small sample sizes (AIC_c). We developed 2 candidate sets of a priori models to quantify factors influencing fawn survival: model set 1 included intrinsic factors and model set 2 focused on habitat variables. We recorded 64 mortalities and the overall survival rate was 0.59 (95% CI = 0.51-0.68). Predation was the leading source of mortality (64%) and coyotes (Canis latrans) were the most prominent predator. For model set 1, model {S_{age X year}} had the lowest AIC_c value suggesting that the age at mortality varied among capture years. For model set 2, model {S_{landscape+forest}} had the lowest AIC_c value and indicated that areas inhabited by surviving fawns were characterized by a few large (i.e., > 5 ha) irregular forest patches adjacent to several small nonforest patches, and survival areas also contained more edge habitat than mortality areas. Due to the magnitude of coyote predation, survival areas could have represented landscapes where coyotes were less effective at locating and capturing fawns when compared to mortality areas. This study was the first account of macrohabitat characteristics directly influencing fawn survival. Wildlife managers can use this information to determine how habitat management activities may affect deer populations.     

Effectiveness of Partial Sedation to Reduce Stress in Captured Mule Deer

Information garnered from the capture and handling of free-ranging animals helps advance understanding of wildlife ecology and can aid in decisions on wildlife management. Unfortunately, animals may experience increased levels of stress, injuries, and death resulting from captures (e.g., exertional myopathy, trauma). Partial sedation is a technique proposed to alleviate stress in animals during capture, yet efficacy of partial sedation for reducing stress and promoting survival post-capture remains unclear. We evaluated the effects of partial sedation on physiological, biochemical, and behavioral indicators of acute stress and probability of survival post-capture for mule deer (Odocoileus hemionus) that were captured via helicopter net-gunning in the eastern Greater Yellowstone Ecosystem, Wyoming, USA. We administered 10–30 mg of midazolam and 15 mg of azaperone intramuscularly (IM) to 32 mule deer in 2016 and 53 mule deer in 2017, and maintained a control group (captured but not sedated) of 38 mule deer in 2016 and 54 mule deer in 2017. To evaluate indicators of acute stress, we measured heart rate, blood-oxygen saturation, body temperature, respiration rate, and levels of serum cortisol. We recorded number of kicks and vocalizations of deer during handling and evaluated behavior during release. We also measured levels of fecal glucocorticoids as an indicator of baseline stress. Midazolam and azaperone did not reduce physiological, biochemical, or behavioral indicators of acute stress or influence probability of survival post-capture. Mule deer that were administered midazolam and azaperone, however, were more likely to hesitate, stumble or fall, and walk during release compared with individuals in the control group, which were more likely to trot, stot, or run without stumbling or falling. Our findings suggest that midazolam (10–30 mg IM) and azaperone (15 mg IM) may not yield physiological or demographic benefits for captured mule deer as previously assumed and may pose adverse effects that can complicate safety for captured animals, including drug-induced lethargy. Although we failed to find efficacy of midazolam and azaperone as a method for reducing stress in captured mule deer, the efficacy of midazolam and azaperone or other combinations of partial sedatives in reducing stress may depend on the dose of tranquilizer, study animal, capture setting, and how stress is defined. © 2020 The Wildlife Society.     

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 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.     

MULE DEER SURVIVAL AMONG ADJACENT POPULATIONS IN SOUTHWEST IDAHO

Abstract: We investigated survival and cause-specific mortality of mule deer (Odocoileus hemionus) on 3 distinct winter ranges in southwest Idaho from 1992 to 1997 to identify demographic variation and potential limiting factors based on a sample of 447 radiocollared deer. During winters 1995–1996 and 1996–1997, we modeled overwinter fawn mortality based on early winter mass, sex, activity, and habitat use variables. Annual survival rates of adult mule deer varied among the 3 adjacent study areas (χ₂² = 10.93, P = 0.004). Overwinter deer survival also varied among study areas (χ₂² = 8.00, P = 0.018), and the study area X year, study area X sex, and study area X age interactions were all significant (P ≤ 0.018). Overwinter survival differences among the study areas were not consistent over time or among sexes and ages of deer. Winter malnutrition was the main cause of mortality for both adults and fawns during the severe winter of 1992–1993, when overall survival was low. Excluding harvest, predation was the major proximate cause of deer mortality during 1993–97 when overall survival was higher. The probability of winter fawn mortality increased with lower mass (χ₂¹ = 7.38, P = 0.007), being male (χ₂¹ = 5.61, P = 0.018), smaller group sizes (χ₂¹ = 3.62, P = 0.057), and using steeper slopes (χ₂¹ = 3.05, P = 0.081). Smaller group sizes and use of steep slopes corresponded to conditions where predators were more successful. Our findings suggest that coyote (Canis latrans) predation was largely compensatory whereas mountain lion (Puma concolor) predation was apparently independent of animal condition and dependent more on deer habitat use. Early winter fawn mass was a better predictor of overwinter fawn survival than a suite of winter resource use variables, lending further support for use of fawn mass to predict winters where fawn mortality may be high. No single population in this study could be used to make reliable inferences regarding deer survival in the other populations. Survival rate measurements should be used cautiously to make inferences in populations where survival has not been directly measured.     

Semi‐parametric analysis of overdispersed count and metric data with varying follow‐up times: Asymptotic theory and small sample approximations

Count data are common endpoints in clinical trials, for example magnetic resonance imaging lesion counts in multiple sclerosis. They often exhibit high levels of overdispersion, that is variances are larger than the means. Inference is regularly based on negative binomial regression along with maximum‐likelihood estimators. Although this approach can account for heterogeneity it postulates a common overdispersion parameter across groups. Such parametric assumptions are usually difficult to verify, especially in small trials. Therefore, novel procedures that are based on asymptotic results for newly developed rate and variance estimators are proposed in a general framework. Moreover, in case of small samples the procedures are carried out using permutation techniques. Here, the usual assumption of exchangeability under the null hypothesis is not met due to varying follow‐up times and unequal overdispersion parameters. This problem is solved by the use of studentized permutations leading to valid inference methods for situations with (i) varying follow‐up times, (ii) different overdispersion parameters, and (iii) small sample sizes.     

Embracing Complexity and Context to Improve Science Communication

Members of the public play a primary role in successful implementation of wildlife management plans, making communication between scientists and the public a vital component of wildlife management. Although there is substantial public interest in the health of ungulate populations, stakeholder perspectives can vary widely, rendering a single approach to communication ineffective. To improve science communication, we characterized perspectives regarding issues negatively affecting mule deer (Odocoileus hemionus) in Wyoming, USA. We used Q methodology, a mixed quantitative-qualitative approach where participants ranked a series of statements followed by semi-structured interviews, to identify shared perspectives. We interviewed individuals (n = 37) representing prominent stakeholder groups (e.g., ranchers, hunters, conservation non-profits) in Wyoming. We identified 3 perspectives (52% of variance explained) that captured shared views regarding what factors are negatively affecting mule deer: bottom-up (n = 17 participants; 26% variance), human contributions (n = 9; 14% variance), and top-down (n = 8; 12% variance) perspectives. Most participants shared the idea that mule deer are being negatively affected, but participants diverged in views as to the primary issues. Perspectives ranged from being focused on bottom-up factors (e.g., habitat fragmentation, condition of winter ranges) to top-down factors (e.g., predation, disease) to factors focused on human contributions (e.g., human activity, public and political interests). Based on how participants diverged in perspectives and their interest in mule deer management, we discuss opportunities for scientists to improve communication by incorporating ecological complexity and nuance, moving towards a 2-way dialogue of communication, and sharing their own first-hand experiences in future communications with stakeholders. © 2021 The Authors. The Journal of Wildlife Management published by Wiley Periodicals LLC on behalf of The Wildlife Society.     

Competition for safe real estate,not food,drives density‐dependent juvenile survival in a large herbivore

Density‐dependent competition for food reduces vital rates, with juvenile survival often the first to decline. A clear prediction of food‐based, density‐dependent competition for large herbivores is decreasing juvenile survival with increasing density. However, competition for enemy‐free space could also be a significant mechanism for density dependence in territorial species. How juvenile survival is predicted to change across density depends critically on the nature of predator–prey dynamics and spatial overlap among predator and prey, especially in multiple‐predator systems. Here, we used a management experiment that reduced densities of a generalist predator, coyotes, and specialist predator, mountain lions, over a 5‐year period to test for spatial density dependence mediated by predation on juvenile mule deer in Idaho, USA. We tested the spatial density‐dependence hypothesis by tracking the fate of 251 juvenile mule deer, estimating cause‐specific mortality, and testing responses to changes in deer density and predator abundance. Overall juvenile mortality did not increase with deer density, but generalist coyote‐caused mortality did, but not when coyote density was reduced experimentally. Mountain lion‐caused mortality did not change with deer density in the reference area in contradiction of the food‐based competition hypothesis, but declined in the treatment area, opposite to the pattern of coyotes. These observations clearly reject the food‐based density‐dependence hypothesis for juvenile mule deer. Instead, our results provide support for the spatial density‐dependence hypothesis that competition for enemy‐free space increases predation by generalist predators on juvenile large herbivores.     

Landscape-scale habitat characteristics and neonatal white-tailed deer survival

Landscape-level habitat characteristics affect neonatal white-tailed deer (Odocoileus virginianus) survival. Little is known, however, about how changes in maternal habitat use after parturition affect neonate survival. We quantified survival rates and determined if neonate survival to 8 weeks was affected by weekly maternal habitat use in the agricultural Glaciated Plains (GP) and forest-grassland Ozark (OZ) eco-regions of Missouri, USA. We captured 127 pregnant female deer during 2015–2017, and fitted each with a global positioning system (GPS) radio-collar and vaginal implant transmitter (VIT). We captured 226 neonatal deer during 2015–2017, fitted each with an expandable radio-collar, and monitored survival status daily. We estimated weekly maternal home ranges and calculated habitat metrics within these home ranges. We used the Kaplan-Meier estimator to calculate 8-week survival estimates and Cox proportional hazards models to investigate the influence of habitat metrics on neonate survival. The 8-week survival estimates were 0.43 (95% CI = 0.35–0.54) and 0.47 (95% CI = 0.38–0.57) in the GP and OZ, respectively. Both of these survival estimates were lower than expected but particularly so in the GP because it is dominated by agricultural fields, a land cover type typically associated with high survival. Neonate survival in the GP was negatively correlated with the amount of edge and forest patch size within maternal home ranges. In the OZ, female neonate survival was positively correlated with birth mass, male neonate survival was not affected by birth mass, and survival of both sexes was negatively correlated with grassland patch density. We suspect these habitat metrics were related to predator searching efficiency and abundance. In the highly fragmented GP, predators might be able to easily search the largest cover habitat patches, whereas in the more contiguous OZ landscape, where cover habitat patch sizes were > 10 times the size of patches in the GP, large patches might be difficult for predators to search efficiently. Therefore, we recommend managers consider the larger landscape context when making habitat management decisions to increase white-tailed deer population productivity. © 2019 The Wildlife Society.     

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.     

Cougar Prey Selection in a White-Tailed Deer and Mule Deer Community

Abstract Widespread mule deer (Odocoilus hemionous) declines coupled with white-tailed deer (O. virginianus) increases prompted us to investigate the role of cougar (Puma concolor) predation in a white-tailed deer, mule deer, and cougar community in northeast Washington, USA. We hypothesized that cougars select for and disproportionately prey on mule deer in such multiple-prey communities. We estimated relative annual and seasonal prey abundance (prey availability) and documented 60 cougar kills (prey usage) from 2002 to 2004. White-tailed deer and mule deer comprised 72% and 28% of the total large prey population and 60% and 40% of the total large prey killed, respectively. Cougars selected for mule deer on an annual basis (α_md = 0.63 vs. α_wt = 0.37; P = 0.066). We also detected strong seasonal selection for mule deer with cougars killing more mule deer in summer (α_md = 0.64) but not in winter (α_md = 0.53). Cougars showed no seasonal selection for white-tailed deer despite their higher relative abundance. The mean annual kill interval of 6.68 days between kills varied little by season (winter = 7.0 days/kill, summer = 6.6 days/kill; P = 0.78) or prey species (white-tailed deer = 7.0 days/kill, mule deer = 6.1 days/kill; P = 0.58). Kill locations for both prey species occurred at higher elevations during summer months (summer = 1,090 m, winter = 908 m; P = 0.066). We suspect that cougars are primarily subsisting on abundant white-tailed deer during winter but following these deer to higher elevations as they migrate to their summer ranges, resulting in a greater spatial overlap between cougars and mule deer and disproportionate predation on mule deer.     

Numerical and Demographic Responses of Pumas to Changes in Prey Abundance: Testing Current Predictions

ABSTRACT Information on factors affecting population size of pumas (Puma concolor) can be important because their principal prey over most of the western United States are valued big game species (e.g., mule deer [Odocoileus hemionus], elk [Cervus elaphus], and bighorn sheep [Ovis canadensis]). Based on the hypothesis that puma numbers are limited by their food supply, puma populations should track changes in prey abundance by growing exponentially with increases in prey and by declining with a lag response when prey decreases. Additional predictions proposed by researchers are that body mass of pumas, female productivity, kitten survival, and adult survival should decrease after a prey decline. We used a 15-year database from a hunted population of pumas in southern Idaho and northwestern Utah to test these predictions. During the 15-year time span of the database, a major decline in mule deer abundance occurred. Estimates of puma numbers and demographic characteristics came from intensive capture and radiocollaring efforts. We calculated kitten and adult survival with MICROMORT software. We found that adult puma numbers increased exponentially at r = 0.07 during a period of increasing mule deer numbers. Four years after the mule deer abundance declined, puma numbers decreased at a rate of r = −0.06. Body mass of female pumas was lower after the decline in puma numbers (42.6 ± SE = 1.2 kg, n = 40 vs. 40.1 ± 0.64 kg, n = 34, t = 5.06, P = 0.045). Kitten survival was less after the decline in deer abundance (0.573 ± 0.016, n = 30 vs. 0.856 ± 0.015, n = 25, Z = 2.40, P < 0.01). Survival of resident females was significantly less after the decline in puma numbers (0.783 ± 0.03 vs. 0.929 ± 0.019, U = 55.0, P = 0.009). Female productivity did not differ before or after the decline in deer abundance. Our results supported the majority of the predictions concerning the impact of changing deer abundance, which supported the hypothesis that the abundance of mule deer limited our population of pumas.