Condition,Survival, and Cause-Specific Mortality of Adult Female Mule Deer in North-Central New Mexico

Abstract: From December 2001 to December 2004 we monitored 30–44 adult female mule deer (Odocoileus hemionus) annually to assess the factors affecting survival and cause-specific mortality. We found adult female survival of 0.63 (SE = 0.08), 0.90 (SE = 0.05), and 0.91 (SE = 0.04), 2002–2004, respectively. Starvation was the most common cause of mortality, accounting for 11/23 mortalities. Mean ingesta-free body fat (IFBF) levels of adult females in December were low (6–9%), despite few (0–13%) lactating adult females, indicative of extremely nutritionally deficient summer—autumn ranges throughout the study site. A priori levels of IFBF and rump body condition scores (rBCS) were higher in deer that survived the following year regardless of cause of mortality. Logistical analysis indicated that models containing individual body fat, rBCS, mean population body fat, winter precipitation, precipitation during mid- to late gestation, and total annual precipitation were related (x² ≥ 9.1; P ≤ 0.003) to deer survival, with individual IFBF (β =-0.47 [SE = 0.21]; odds ratio = 0.63 [0.42-0.94]) and population mean IFBF (β = -1.94 [SE = 0.68]; odds ratio = 0.14 [0.04-0.54]) the best predictors; with either variable, probability of dying decreased as fat levels increased. Fawn production was low (2–29 fawns/100 ad F) and, combined with adult survival, resulted in estimated population rates of increase of -35%, -5%, and +6% for 2002–2004, respectively. Deer survival and population performance were limited in north-central New Mexico, USA, due to poor condition of deer, likely a result of limited food resulting from both drought and long-term changes in plant communities. Precipitation during mid- to late gestation was also important for adult female survival in north-central New Mexico.     

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.     

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.     

Evaluating precision of cementum annuli analysis for aging mule deer from southern California

We evaluated the precision of age estimates produced by cementum annuli analysis (CAA) of blind-duplicate specimens taken from 994 southern mule deer (Odocoileus hemionus) collected over 15 years. We found that the mean annual proportion of unreliably aged incisor pairs was greater for females (0.48, SD = 0.13) than for males (0.22, SD = 0.07). Most of the 308 unreliably aged tooth pairs disagreed by only 1 year. Sex, precipitation, and certainty codes assigned by Matson's Lab to the age estimates were the best predictors for agreement of estimated ages within incisor pairs. Our estimated overall age error rate of CAA (17%) was >2 times as large as estimated error rates from Montana and South Dakota, but less than half of error rates estimated for Mississippi and south Texas. Knowing the error rate of age estimates from a specific deer population allows wildlife managers to perform tasks requiring specific age class information such as monitoring the harvest rate of older female deer in a hunted population or performing population reconstruction. © 2011 The Wildlife Society.     

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.     

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.     

Evaluating Dependence Among Mule Deer Siblings in Fetal and Neonatal Survival Analyses

Abstract: The assumption of independent sample units is potentially violated in survival analyses where siblings comprise a high proportion of the sample. Violation of the independence assumption causes sample data to be overdispersed relative to a binomial model, which leads to underestimates of sampling variances. A variance inflation factor, c, is therefore required to obtain appropriate estimates of variances. We evaluated overdispersion in fetal and neonatal mule deer (Odocoileus hemionus) datasets where more than half of the sample units were comprised of siblings. We developed a likelihood function for estimating fetal survival when the fates of some fetuses are unknown, and we used several variations of the binomial model to estimate neonatal survival. We compared theoretical variance estimates obtained from these analyses with empirical variance estimates obtained from data-bootstrap analyses to estimate the overdispersion parameter, c. Our estimates of c for fetal survival ranged from 0.678 to 1.118, which indicate little to no evidence of overdispersion. For neonatal survival, 3 different models indicated that ĉ ranged from 1.1 to 1.4 and averaged 1.24–1.26, providing evidence of limited overdispersion (i.e., limited sibling dependence). Our results indicate that fates of sibling mule deer fetuses and neonates may often be independent even though they have the same dam. Predation tends to act independently on sibling neonates because of dam-neonate behavioral adaptations. The effect of maternal characteristics on sibling fate dependence is less straightforward and may vary by circumstance. We recommend that future neonatal survival studies incorporate additional sampling intensity to accommodate modest overdispersion (i.e., ĉ = 1.25), which would facilitate a corresponding ĉ adjustment in a model selection analysis using quasi-likelihood without a reduction in power. Our computational approach could be used to evaluate sample unit dependence in other studies where fates of individually marked siblings are monitored.     

Survival and Cause-Specific Mortality of Neonatal Mule Deer Fawns,North-Central New Mexico

Abstract: Because of significant declines in mule deer (Odocoileus hemionus) populations across New Mexico, USA, we investigated survival of fawns in north-central New Mexico, USA. We captured 19 fawns, 34 fawns, and 47 fawns in 2002, 2003, and 2004, respectively, and used fawn morphological measurements, habitat characteristics, and adult female (hereafter “female”) condition to model preweaning fawn survival. Survival was 0.0, 0.12, 0.52 for 2002, 2003, and 2004, respectively, and was related to birth mass (χ₁² = 9.5, P = 0.002), birth date (χ₁²= 8.4, P = 0.004), litter size (χ₂² = 9.4, P = 0.009), female body fat (χ₁² = 40.9, P < 0.001), annual precipitation (χ₁² = 35.0, P < 0.001), summer precipitation (χ₁²= 37.5, P < 0.001), and winter precipitation (χ₁² = 32.0, P < 0.001). Total ingesta-free body fat of females (β = 3.01, SE = 0.75; odds ratio = 20.19, 95% CI = 4.64-87.91) and birth mass of fawns (β = 1.188, SE = 0.428; odds ratio = 3.38, 95% CI = 1.42-7.59) were the best predictors of survival of individual fawns, although few of the logistic models differed in model selection criteria. Fawn survival in north-central New Mexico was driven by an interaction of total and seasonal precipitation and its effect on plant production, consequential effects on female nutrition, and ultimately, fawn birth attributes. Habitat conditions were so poor throughout north-central New Mexico during 2002 and 2003 (and likely during other drought yr) that, based upon birth attributes, few fawns could have survived regardless of proximate causes of mortality. In 2004, precipitation enhanced security cover, maternal body condition, birth attributes and, thus, survival of fawns. However, more habitat enhancements are needed to improve the nutritional quality of mule deer habitats in north-central New Mexico and further enhance maternal and fawn condition to recover mule deer populations in this region.     

Survival and Cause-Specific Mortality of Male Mule Deer Under Different Hunting Regulations in the Bridger Mountains,Montana

Abstract: We selected 2 adjacent populations of mule deer (Odocoileus hemionus hemionus) in the Bridger Mountains, Montana, USA, to measure effects on survival rates and causes of mortality of 2 hunting regulations designed to enhance representation of mature males. We compared male survival between the West Slope and South 16 Mile populations considering both hunting and nonhunting sources of mortality with respect to age (fawn, yearling, and mature), month (Jun-May), and year (1990-1995). Harvest rates of mature males were greater than for yearlings, demonstrating hunter preferences. We found no differences in yearling monthly survival rates between October and November or between areas or years. In contrast, we found survival of mature males differed between October and November and across years and study areas. During these months, survival rates of mature males averaged 0.602 on the West Slope under the 2-point regulation and 0.762 on South 16 Mile under the outfitted hunt. Monthly survival during summer also differed by age class, but not area, with estimates of 0.963 for yearling males, and corresponding mature male survival estimates of 0.991, demonstrating greater yearling summer mortality. Winter survival rates of yearlings and matures were similar for both areas with a monthly estimate of 0.986. We found differences in spring monthly survival estimates for the 2 areas, mainly for matures. Yearling male monthly survival estimates were 0.959 and 0.958 for the 2 areas, whereas corresponding mature male estimates during spring were 0.991 and 0.936 on the West Slope and South 16 Mile, respectively. Fawn survival rates varied from 0.101 to 0.770 among years and between areas overwinter. Cumulative effects of nonhunting mortality among all age classes reduced the effectiveness of 2 hunting regulations designed to enhance survival of males to age classes ≥4 years associated with maximum antler development despite accomplishing reductions in harvest rates. Low and variable fawn survival and relatively high nonhunting-related losses of yearling and mature males might be typical of many populations in the Northern Rocky Mountains. Deer managers should avoid populations coexisting with a diversity of large predators in environments with strong year effects when considering opportunities for implementing harvest regulations to improve representation of mature males.     

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.     

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.     

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.     

Effects of Winter-Feeding on Mule Deer in Northern Utah

Abstract: During severe winters, mule deer (Odocoileus hemionus) concentrated on ranges in poor condition can experience high mortality. Winter-feeding programs have been implemented to mitigate this mortality. We studied effects on body condition, mortality, fawn production, and migration of mule deer following winter-feeding in the Cache-Wasatch Mountains of northern Utah, USA. Fed deer exhibited 12% higher live body-condition indices both years (main effect feed: F_1,7.32 = 5.39, P = 0.052), lower mortality (33% vs. 55%: χ²₁= 4.58, P < 0.05), and produced more fawns (19 fawns:18 fed F vs. 11 fawns:12 nonfed F; t_27.2 = 2.20, P < 0.036) than nonfed deer. Fed deer migrated later in spring 2004 (x̄ = 13 Apr) than nonfed deer (x̄ = 24 Mar; t₃₄= 3.25, P = 0.003). Fed deer spent more time on winter range in 2003-2004 (x̄ = 157 d) than nonfed deer (x̄ = 121 d; t20 = 3.63, P = 0.002), and more time on winter range for both winters combined (fed deer x̄ = 321 d, nonfed deer x̄ = 257 d; t₂₇ = 3.29, P = 0.003). Concomitantly, wildlife managers need to recognize that any possible benefits accrued to mule deer populations in terms of increased nutritional status as a result of winter-feeding programs may be mitigated by altered timing of migration and increased duration of use of seasonal ranges by fed deer.     

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.