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.     

Evaluating risks associated with capture and handling of mule deer for individual-based,long-term research

Capture and handling techniques for individual-based, long-term research that tracks the life history of animals by recapturing the same individuals for several years has vastly improved study inferences and our understanding of animal ecology. Yet there are corresponding risks to study animals associated with physical trauma or capture myopathy that can occur during or following capture events. Rarely has empirical evidence existed to guide decisions associated with understanding the magnitude of capture-related risks, how to reduce these risks when possible, and implications for mortality censoring and survival estimates. We used data collected from 2,399 capture events of mule deer (Odocoileus hemionus) via helicopter net-gunning to compare daily survival probabilities within a 10-week period centered on a capture event and evaluated how animal age, nutritional condition (body fat), and various handling methods influenced survival before, during, and following a capture event. Direct mortality resulting from capture efforts was 1.59%. Mean daily survival was 0.9993 ± 0.0001 (SE) during the 5-week pre-capture window, was depressed the day of capture at 0.9841 ± 0.0004, and rebounded to 0.9990 ± 0.0008 during the 5-week post-capture window. Neither capture nor handling had a detectable effect on post-capture survival, including handling time ($\overline{x}$ = 13.30 ± 1.87 min), capture time of year (i.e., Dec or Mar), tooth extraction, and the number of times an animal had been recaptured (2–17 times). Although mortality rate was slightly elevated during capture (resulting from physical trauma associated with capture), age and nutritional condition did not influence the probability of mortality during a capture event. Following a capture event, nutritional condition influenced survival; however, that relationship was consistent with expected effects of nutritional condition on winter survival and independent of capture and handling. Overall survival rates 5 weeks before capture and 5 weeks after capture were not different. A specified window of time with depressed survival following capture and handling was not evident, which contradicts the implementation of a predetermined window often used by researchers and managers for censoring mortalities that occur after capture. Previous notions that censorship of all mortality data in the 2 weeks following capture is unwarranted and risks removal of meaningful data. With previous evidence guiding our protocols for capture (e.g., reduced chase time) and handling (e.g., temperature mitigation), low direct mortality and almost undetectable indirect mortality post capture reinforces the efficacy of helicopter net-gunning for capture and recapture of mule deer in long-term, individual-based studies.     

Assessing the Helicopter and Net Gun as a Capture Technique for White-Tailed Deer

Abstract The helicopter and net gun is a technique used to capture white-tailed deer (Odocoileus virginianus) and is useful in a variety of habitat types and at various population densities with the ability to be highly selective. During capture, deer may sustain injuries or even die as a result of capture and handling, and may also be prone to capture myopathy. Therefore, our objectives were to determine 1) type and frequency of injuries sustained during the helicopter and net-gun capture, and 2) the effects of capture on survival of radiocollared deer. We captured 3,350 white-tailed deer from 1998 to 2005 using a net gun fired from a helicopter on 5 southern Texas, USA, ranches. Additionally, we captured 51 yearling males and 49 mature (≥4 yr of age) males and fitted them with radiocollars to monitor their survival. We recorded injuries and mortalities during capture and ranked the seriousness of injuries on a scale from 0 to 4. We recorded 281 injuries (8.4%) and as a result of capture, at least 206 deer had broken antlers (6.1%), 55 were injured (1.6%), and 20 were direct mortalities (0.6%). The most common antler injury was broken antler tines and the most common body injury was broken legs. Postcapture mortality rates were low (1%) for this capture method. Based on capture-related injuries, mortalities, and postcapture survival, we found the helicopter and net gun to be a safe capture technique compared to other capture techniques, particularly when conditions are favorable.     

Capture method affects survival estimates and subsequent interpretation of ecological covariates for a long‐lived cervid

Understanding what variables affect ungulate neonate survival is imperative to successful conservation and management of the species. Predation is commonly cited as a cause‐specific source of mortality, and ecological covariates often influence neonate survival. However, variation in survival estimates related to capture methodology has been documented with opportunistically captured neonates generally displaying greater survival than those captured via aid of vaginal implant transmitters (VITs), likely because of increased left truncation observed in the opportunistically captured datasets. Our goal was to assess whether 3‐ and 6‐month survival estimates varied by capture method while simultaneously assessing whether capture method affected model selection and interpretation of ecological covariates for white‐tailed deer neonates captured from three study sites from 2014 to 2015 in North Dakota and South Dakota, USA. We found survival varied by capture method for 3‐month neonate survival with opportunistically captured neonates displaying up to 26% greater survival than their counterparts captured via VITs; however, this relationship was not present for 6‐month survival. We also found model selection and subsequent interpretation of ecological covariates varied when analyzing datasets comprised of neonates captured via VITs, neonates captured opportunistically, and all neonates combined regardless of capture method. When interpreting results from our VIT‐only analysis for 3‐month survival, we found survival varied by three time intervals and was lowest in the first two weeks of life. Capture method did not affect 6‐month survival, which was most influenced by total precipitation occurring during 3 – 8 weeks of a neonate''s life and percent canopy cover found at a neonate''s capture site. Our results support previous research that capture method must be accounted for when deriving survival estimates for ungulate neonates as it can impact derived estimates and subsequent interpretation of results.     

Effects of helicopter net-gunning on survival of bighorn sheep

Wildlife capture, and the data collection associated with it, has led to major advancements in ecology that are integral to decision making pertaining to wildlife conservation. Capturing wildlife, however, can cause lethal and non-lethal risks to animals. Understanding the factors that contribute to the level of risk involved in wildlife capture is therefore important for the development and implementation of the safest and most effective methodologies. We used data from 736 animal captures of 389 individuals for 2 subspecies of female bighorn sheep (Rocky Mountain bighorn [Ovis canadensis canadensis], Sierra Nevada bighorn sheep [O. c. sierrae]) in Wyoming and California, USA, in 2002–2020 to evaluate the degree and extent of time that capture via helicopter net-gunning affects survival. We compared pre- and post-capture survival during a 10-week window centered on a capture event, and post-capture survival between captured animals and animals that were monitored but not captured during the 10-week window. Additionally, we evaluated the effects of handling techniques (number of times captured, season of capture event, handling time, chase time, and body temp) and biological factors (age and nutritional condition) on probability of capture mortality. Mean daily survival was 0.9992 during a 5-week pre-capture window, dropped to 0.9864 on the day of capture, and rebounded within 3 days of capture to pre-capture levels and that of sheep that were not captured. Overall, direct mortality resulting from capture was 1.36%, with 0.54% mortality occurring within the 3 days following a capture event for an overall 1.90% capture-related mortality. The only handling and biological metrics that influenced the probability of capture mortality were rectal temperature and nutritional condition; high initial rectal temperatures and poor body condition were associated with increased risk of mortality in the days following capture. Overall, helicopter net-gunning imposed low and short-term risk to survival of female bighorn sheep. To reduce bias in survival estimates, we recommend using a 3-day censorship window for post-capture mortalities as opposed to the common practice of a 2–5-week censor window. Helicopter net-gunning, including annual or seasonal recaptures, remains an effective and comparatively safe technique for capture and associated data collection of bighorn sheep.     

Evaluating Indirect Effects of Hunting on Mule Deer Spatial Behavior

Mule deer (Odocoileus hemionus) are widely hunted throughout western North America and are experiencing population declines across much of their range. Consequently, understanding the direct and indirect effects of hunting is important for management of mule deer populations. Managers can influence deer mortality rates through changes in hunting season length or authorized tag numbers. Little is known, however, about how hunting can affect site fidelity patterns and subsequent habitat use and movement patterns of mule deer. Understanding these patterns is especially important for adult females because changes in behavior may influence their ability to acquire resources and ultimately affect their productivity. Between 2008 and 2013, we obtained global positioning system locations for 42 adult female deer at the Starkey Experimental Forest and Range in northeast Oregon, USA, during 5-day control and treatment periods in which hunters were absent (pre-hunt), present but not actively hunting (scout and post-hunt), and actively hunting male mule deer (hunt) on the landscape. We estimated summer home ranges and 5-day use areas during pre-hunt and hunt periods and calculated overlap metrics across home ranges and use areas to assess site fidelity within and across years. We used step selection functions to evaluate whether female mule deer responded to human hunters by adjusting fine-scale habitat selection and movement patterns during the hunting season compared to the pre-hunt period. Mule deer maintained site fidelity despite disturbance by hunters with 72 ± 4% (SE) within-year overlap between summer home ranges and hunt use areas and 54 ± 7% inter-annual overlap among pre-hunt use areas and 56 ± 7% among hunt use areas. Mule deer diurnal movement rates, when hunters are active on the landscape, were higher during the hunting period versus pre-hunt or scout periods. In contrast, nocturnal movement rates, when hunters are inactive on the landscape, were similar between hunting and non-hunting periods. Additionally, during the hunt, female mule deer hourly movements increased in areas with high greenness values, indicating that mule deer spent less time in areas with more vegetative productivity. Female mule deer maintained consistent habitat selection patterns before and during hunts, selecting areas that offered more forest canopy cover and high levels of vegetative productivity. Our results indicate that deer at Starkey are adopting behavioral strategies in response to hunters by increasing their movement rates and selecting habitat in well-established ranges. Therefore, considering site fidelity behavior in management planning could provide important information about the spatial behavior of animals and potential energetic costs incurred, especially by non-target animals during hunting season. © 2020 The Wildlife Society.     

Survival,Birth Characteristics,and Cause-Specific Mortality of White-Tailed Deer Neonates

ABSTRACT Understanding survival of and factors that may predispose newborn deer (Odocoileus spp.) to mortality contribute to improved understanding of population dynamics. We captured free-ranging white-tailed deer neonates (n = 66) of radiocollared females that survived severe (Winter Severity Index [WSI] = 153) and mild (WSI = 45) winters 2000–2001 and 2001–2002. Mean dates of birth (26 May ± 1.7 [SE] days and 26 May ± 1.3 days) and estimated birth-masses of 2.8 ± 0.1 kg and 3.0 ± 0.1 kg were similar between springs 2001 (n = 31) and 2002 (n = 35), respectively. Neonate survival was similar between years; pooled mortality rates of neonates were 0.14, 0.11, and 0.20 at 0–1 weeks, 2–4 weeks, and 5–12 weeks of age, respectively, and overall survival rate for neonates to 12 weeks of age was 0.47. Predation accounted for 86% of mortality; the remaining 14% of deaths were attributed to unknown causes. Black bears (Ursus americanus) were responsible for 57% and 38% of predation of neonates in springs 2001 and 2002, respectively, whereas bobcats (Felis rufus) accounted for 50% in 2002. Wolves (Canis lupus) accounted for only 5% of predator-related deaths. Low birth-mass, smaller body size, and elevated concentrations of serum urea nitrogen (26.1 ± 2.6 mg/dL vs 19.3 ± 0.8 mg/dL) and tumor necrosis factor-α (82.6 ± 78.6 pg/mL vs. 2.3 ± 0.5 pg/mL) were associated with neonates that died within 1 week of birth. Even though we did not detect a direct relation between winter severity and birth or blood characteristics of neonates, evidence suggests that birth-mass and key serum indices of neonate nutrition were associated with their early mortality. Thus, managers can make more informed predictions regarding survival and cause-specific mortality of fawns and adjust management strategies to better control deer population goals.     

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.     

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.     

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.     

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.     

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.     

Postpartum Group Cohesion of Sympatric Deer in Texas

ABSTRACT Postpartum behavior of maternal deer may be specific to species of deer and predators. We captured sympatric white-tailed deer (Odocoileus virginianus) and mule deer (O. hemionus eremicus) fawns from radiocollared adult females in 2004–2006 on rangelands of west central Texas, USA, where predators larger than bobcats (Lynx rufus) were absent. Our objective was to determine whether differences in postpartum antipredator behavior existed between deer species, and if so, examine efficacy of those strategies. We collected postpartum group cohesion data in 2004 and 2005 by using radiotelemetry and examined dead fawns for cause of mortality. During fawns' hider phase, <3 weeks postpartum, mule deer females kept fawns closer to themselves (95% CI = 39−66 m) and twins closer to each other (95% CI = 25–49 m) than did white-tailed deer females (95% CIs = 152–234 m and 163–255 m, respectively). After 30 days postpartum, familial group cohesion was similarly tight for both species. During hider phases from 2004 to 2006, predated carcasses of white-tailed deer fawns (11 of 11) were dismembered or consumed more than mule deer fawns (7 of 13, P = 0.016), which was one line of evidence for maternal defense by mule deer adults. During hider phases in 2004 and 2005, predation rate of mule deer fawns was lower than that for white-tailed deer fawns. In 2006, predation rate increased for mule deer but was similar for white-tailed deer fawns compared with previous years. The tight cohesion strategy of mule deer exhibited in 2004 and 2005 seemed successful at thwarting small predators. Without large predators, the loose cohesion strategy of white-tailed deer females was maladaptive. When meso-predators are abundant due to extermination of larger predators, predation on fawns could increase if a deer species has relatively fixed postpartum maternal antipredator behavior.     

Interannual Variability in Survival Rates for Adult Female White-Tailed Deer

Adult female survival is an important component to population models and management programs for white-tailed deer (Odocoileus virginianus), but short-term survival studies (1–3 yrs) may not accurately reflect the variation in interannual survival, which could alter management decisions. We monitored annual survival and cause-specific mortality rates of adult female white-tailed deer (n = 158) for 6 years (2010–2012, 2016–2018) in southern Delaware, USA. Annual survival rate differed among years. Survival rates (±SE) and mortality causes were similar in 3 years (2011 = 0.72 ± 0.08, 2017 = 0.68 ± 0.08, 2018 = 0.74 ± 0.09) and comparable to previous research from mixed forest-agricultural landscapes. A relatively low survival rate in 2010 (0.48 ± 0.11) was influenced by hunter harvest and potentially compounded by abnormally severe winter conditions in the prior year. A peracute outbreak of hemorrhagic disease occurred during summer 2012, resulting in an annual survival rate of 0.38 ± 0.11, and to our knowledge is the first reported case of a hemorrhagic disease outbreak in a monitored wild population with known fates. In 2016, we did not observe any harvest mortality, resulting in high annual survival (0.96 ± 0.04). Our results demonstrate the degree of variability in annual survival and cause-specific mortality rates within a population. We caution against the use of short-term survival studies to inform management decisions, particularly when incorporating survival data into population models or when setting harvest objectives. © 2020 The Wildlife Society.     

How Size and Condition Influence Survival and Cause-Specific Mortality of Female Elk

The size of animal populations fluctuates with number of births, rate of immigration, rate of emigration, and number of deaths. For many ungulate populations, adult female survival is the most important factor influencing population growth. Therefore, increased understanding of survival and causes of mortality for adult females is fundamental for conservation and management. The objectives of our study were to quantify survival rates of female elk (Cervus canadensis) and determine cause-specific mortality. We predicted that hunter harvest would be the leading cause of mortality. Further, we predicted that hunters would harvest animals that were in prime age (2–9 yr) and in better condition than elk predated by mountain lions (Puma concolor). From 2015 to 2017, we captured 376 female elk in central Utah, USA. We assessed body size and condition of captured elk, fitted each animal with a global positioning system-collar, and determined cause of death when we received mortality signals. We estimated survival using Kaplan-Meier estimates and Cox proportional hazard models within an Akaike's Information Criterion model selection framework to identify covariates that influenced survival. We analyzed differences in size and condition measurements between harvested elk and predated elk using analysis of variance tests. Our best model indicated consistent survival across years; mean survival was 78.3 ± 3.5% (SE) including hunter harvest and 95.5 ± 1.7% without hunter harvest. In decreasing order of importance, elk mortality occurred from hunter harvest (21.2%), mountain lion predation (3.7%), depredation removal (0.5%), automobile collision (0.3%), disease (0.3%), complications during calving (0.3%), and those characterized as undetermined (1.3%). Neck circumference and body length were negatively associated with survival, suggesting that larger animals in good condition had lower survival as a result of hunter harvest. Individuals that died because of cougar predation were smaller and had less loin muscle than the average animal. Hunters removed large, healthy, prime-aged females, individuals that likely have a greater effect on population growth than elk lost to other predators. If the proportion of larger, healthy females in the population begins to decline, hunting practices may require adjustment because hunters may be removing individuals with the greatest reproductive value. © 2021 The Wildlife Society.     

Effects of Chemical Immobilization on Survival of African Buffalo in the Kruger National Park

ABSTRACT Capturing, immobilizing, and fitting radiocollars are common practices in studies of large mammals, but success is based on the assumptions that captured animals are representative of the rest of the population and that the capture procedure has negligible effects. We estimated effects of chemical immobilization on mortality rates of African buffalo (Syncerus caffer) in the Kruger National Park, South Africa. We used a Cox proportional hazards approach to test for differences in mortality among age, sex, and capture classes of repeatedly captured radiocollared buffalo. Capture variables did not improve model fit and the Cox regression did not indicate increased risk of death for captured individuals up to 90 days postcapture [exp (β) = 1.07]. Estimated confidence intervals, however, span from a halving to a doubling of the mortality rate (95% CI = 0.56–2.02). Therefore, capture did not influence survival of captured individuals using data on 875 captures over a 5-year period. Consequently, long-term research projects on African buffalo involving immobilization, such as associated with research on bovine tuberculosis, should result in minimal capture mortality, but monitoring of possible effects should continue.     

Comparative evaluation of effort, capture and handling effects of drive nets to capture roe deer (Capreolus capreolus), Southern chamois (Rupicapra pyrenaica) and Spanish ibex (Capra pyrenaica)

The objective of this study is to assess the usefulness of drive nets to capture roe deer (Capreolus capreolus), Southern chamois (Rupicapra pyrenaica) and Spanish ibex (Capra pyrenaica), comparing the results obtained with other capture methods and amongst the three species. Sixty-five drive net capture operations using beaters were conducted from January 1998 to September 2004. A total of 161 wild ungulates (31 roe deer, 95 Southern chamois and 35 Spanish ibexes) were captured. The average number of animals captured per operation was 1.07 for roe deer, 3.96 for Southern chamois and 2.92 for Spanish ibex. The average number of person–days per captured animal was 21.5, 7.1 and 10.6 for roe deer, Southern chamois and Spanish ibex, respectively. Specificity was 100% for Southern chamois and Spanish ibex (only the target species captured) and 77.5% for roe deer. Risk for the animals (mortality plus injuries) was 3.23% for roe deer, 5.27% for Southern chamois and 0% for Spanish ibex, whereas injuries to the operators occurred with 3.1% of the handled animals. Sex ratio was skewed towards females in roe deer, towards males in Southern chamois and balanced in Southern chamois. Drive nets showed good performance, although many operators were required. Safety for the animals and specificity were higher than traditionally attributed to this capture method. It is concluded that drive nets are an efficient and safe method to capture many ungulate species.     

Estimating chronic wasting disease effects on mule deer recruitment and population growth

Chronic wasting disease (CWD), a prion disease of mule deer (Odocoileus hemionus), accelerates mortality and in so doing has the potential to influence population dynamics. Although effects on mule deer survival are clear, how CWD affects recruitment is less certain. We studied how prion infection influenced the number of offspring raised to weaning per adult (≥2 yr old) female mule deer and subsequently the estimated growth rate (λ) of an infected deer herd. Infected and presumably uninfected radio-collared female deer were observed with their fawns in late summer (August-September) during three consecutive years (2006-2008) in the Table Mesa area of Boulder, Colorado, USA. We counted the number of fawns accompanying each female, then used a fully Bayesian model to estimate recruitment by infected and uninfected females and the effect of the disease on λ. On average, infected females weaned 0.95 fawns (95% credible interval=0.56-1.43) whereas uninfected females weaned 1.34 fawns (95% credible interval=1.09-1.61); the probability that uninfected females weaned more fawns than infected females was 0.93). We used estimates of prevalence to weight recruitment and survival parameters in the transition matrix of a three-age, single-sex matrix model and then used the matrix to calculate effects of CWD on λ. When effects of CWD on both survival and recruitment were included, the modeled λ was 0.97 (95% credible interval = 0.82-1.09). Effects of disease on λ were mediated almost entirely by elevated mortality of infected animals. We conclude that although CWD may affect mule deer recruitment, these effects seem to be sufficiently small that they can be omitted in estimating the influences of CWD on population growth rate.     

How has the remnant population of the threatened frog Leiopelma pakeka (Anura: Leiopelmatidae) fared on Maud Island,New Zealand,over the past 25 years?

Despite widespread global reports of declining amphibian populations, supporting long‐term census data are few, limiting opportunities to study changes in numbers and survival over time. However, in New Zealand, for the past 25 years (1983–2008), we studied Leiopelma pakeka, a threatened, terrestrial frog that inhabits rocky boulder banks under forest on Maud Island. Using night sampling at least annually on two 12 × 12 m plots, we had 5390 captures of 1000+ individuals, 327 on one plot (grid 1), 751 on the other (grid 2). The mean (±SE) number of frogs found per night was 11.3 (±0.6) on grid 1 and 25.6 (±1.4) on grid 2. We used capture‐recapture models to estimate population size, proportion of animals remaining beneath the surface and survival rate. The mean (±SE) population estimate was 131 (±14.7) frogs on grid 1 and 367 (±38.7) on grid 2. Over 25 years the estimated population increased on grid 1 and fluctuated on grid 2. Some frogs were captured on most sampling visits, others less often, evidently failing to emerge from cover each visit. Using a combination of open and closed population models, we estimated the mean (±SE) proportion remaining underground was 0.63 (±0.12) on grid 1 and 0.53 (±0.07) on grid 2. Our research represents one of the longest‐run population studies of any frog, and we recorded significant longevity, two males reaching 35+ and 37+ years, a female 34+ years. No significant differences occurred between mean annual survival rates of apparent females and males, or between the two sites. The number of toes clipped for individual identification had little influence on the return rate, once the effect of time of first capture was removed.     

Netguns: a technique for capturing Black‐backed Woodpeckers

ABSTRACT Effective capture techniques are essential for studying bird populations, but commonly used techniques have proven ineffective for capturing Black‐backed Woodpeckers (Picoides arcticus) during the nonbreeding period. As a result, little is known about the winter ecology of Black‐backed Woodpeckers. We used two netguns, one powered with a 0.308 cartridge and another with CO₂ propellant, to capture 101 Black‐backed Woodpeckers (N= 75 initial captures and 26 recaptures) in the Black Hills of South Dakota from 2008 to 2011. Captures with the 0.308 netgun resulted in an impact mortality probability of 0.061 ± 0.034 (SE), whereas no impact mortalities were associated with the CO₂ netgun. We also tracked birds for 72 h post‐release, and determined a capture‐related mortality rate of 0.102 ± 0.04 with the 0.308 netgun and 0.038 ± 0.027 with the CO₂ netgun. With the CO₂ netgun, we captured woodpeckers in 31 of 43 net deployments (72%), with an average of 7.2 ± 0.4 h of capture effort for each bird. Many unsuccessful attempts were caused by tree branches that prevented net deployment. Netguns powered by CO₂ provide an effective capture technique that we recommend for studies of Black‐backed Woodpeckers and possibly other species of birds that forage low on trees.