Migratory Disturbance Thresholds with Mule Deer and Energy Development

Fine-scale movement data has transformed our knowledge of ungulate migration ecology and now provides accurate, spatially explicit maps of migratory routes that can inform planning and management at local, state, and federal levels. Among the most challenging land use planning issues has been developing energy resources on public lands that overlap with important ungulate habitat, including the migratory routes of mule deer (Odocoileus hemionus). We generally know that less development is better for minimizing negative effects and maintaining habitat function, but we lack information on the amount of disturbance that animals can tolerate before reducing use of or abandoning migratory habitat. We used global positioning system data from 56 deer across 15 years to evaluate how surface disturbance from natural gas well pads and access roads in western Wyoming, USA, affected habitat selection of mule deer during migration and whether any disturbance threshold(s) existed beyond which use of migratory habitat declined. We used resource and step selection functions to examine disturbance thresholds at 3 different spatial scales. Overall, migratory use by mule deer declined as surface disturbance increased. Based on the weight of evidence from our 3 independent but complementary metrics, declines in migratory use related to surface disturbance were non-linear, where migratory use sharply declined when surface disturbance from energy development exceeded 3%. Disturbance thresholds may vary across regions, species, or migratory habitats (e.g., stopover sites). Such information can help with management and land use decisions related to mineral leasing and energy development that overlap with the migratory routes of ungulates. © 2020 The Wildlife Society.     

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.     

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.     

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.     

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.     

Mule Deer Migrations and Highway Underpass Usage in California,USA

Roadways may pose barriers to long-distance migrators such as some ungulates. Highway underpasses mitigate wildlife-vehicle collisions and can be an important management tool for protecting migration corridors. In northern California, 3 underpasses were built on United States Route 395 (Route 395) in Hallelujah Junction Wildlife Area (HJWA) in the 1970s for a migratory mule deer (Odocoileus hemionus) herd that had been negatively affected by highway traffic. To determine whether these underpasses were still reducing mule deer mortalities >40 years after construction, we investigated deer use of the underpasses from 2006–2019 using cameras, global positioning system (GPS) collars, and roadkill records. We used occupancy models, approximations of GPS-collared mule deer movement paths, and roadkill locations to estimate the highway crossing patterns of deer. From camera data, there was higher use of the underpasses by deer during migration (spring [Mar–Jun], fall [Oct–Dec]) than in summer (Jul–Sep), when only resident deer were present. Higher underpass usage occurred in the spring compared to fall migrations. Eleven of 21 GPS-collared migrating mule deer crossed Route 395. We estimated 30% of the crossings (by 7 of the 11 deer) occurred south of the underpasses where deer could easily access the highway because of short (1-m high) and deteriorating highway fencing. Roadkill data confirmed that deer-vehicle collisions were occurring south of the underpasses and at the underpasses. This was likely due to deteriorating infrastructure at the underpasses that allows wildlife access to the highway. Overall, our study indicated that although underpasses can provide safe passage for migratory deer decades (>40 yr) after their construction, deteriorating infrastructure such as fencing and gates can lead to wildlife mortalities on highways near underpasses. © 2021 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.     

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.     

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.     

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.     

Evaluating Mule Deer Body Condition Using Serum Thyroid Hormone Concentrations

ABSTRACT Body condition of ungulates is a determinant of fecundity and survival rates. Ultrasonography and body condition scoring techniques allow reliable estimation of body fat but may not be feasible to employ in some circumstances. A reliable blood chemistry index for assessing relative condition of different ungulate populations or groups would be useful in ongoing population monitoring programs. We provided a nutrition supplement (treatment) to a group of free-ranging mule deer (Odocoileus hemionus) during 2 consecutive winters in southwest Colorado. In late February each year, we evaluated whether percent body fat and serum concentrations of total thyroxine (T4), total triiodothyronine (T3), free thyroxine (FT4), and free triiodothyronine (FT3) were higher among treatment deer than an adjacent group of deer that did not receive treatment (control). As a corroborative analysis, we modeled body fat as a function of thyroid hormone concentrations and morphometric variables. Estimated body fat of treatment deer averaged 12.3% (SE = 0.327), whereas estimated body fat of control deer averaged 7.0% (SE = 0.333) during the 2 winters of study. Concentrations of T4 and FT4 averaged 48.07 nanomole/L (SE = 3.80) and 12.61 picomole/L (SE = 1.04) higher, respectively, in treatment deer than control deer. Our optimal model of estimated body fat included T4, T4², FT4, and deer chest girth (%FAT = −4.8015 − 0.0946 × T4 + 0.000603 × T4² + 0.1474 × FT4 + 0.1426 × chest girth, R² = 0.609). Serum thyroid hormones effectively discerned treatment deer from control deer and were related to estimated body fat. Ultrasound and body condition scoring should be used to estimate body fat whenever possible. However, in cases where only a blood sample can be obtained, we documented potential utility of T4 and FT4 during late winter for evaluating relative body condition of mule deer.     

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.     

Validating Predictive Models of Nutritional Condition for Mule Deer

Abstract: We developed new, and validated existing, indices of nutritional condition for live and dead mule deer (Odocoileus hemionus). Live animal indices included a body condition score (BCS), thickness of subcutaneous fat and selected muscles using ultrasonography, and body mass. Dead animal indices included femur, metatarsal, and mandible marrow fat, 3 kidney fat indices, and 2 carcass scoring methods. We used 21 female deer and 4 castrates (1-11 yr old) varying widely in nutritional condition (2-28% ingesta-free body fat). Deer were euthanized and homogenized for chemical analysis of fat, protein, water, and ash content. Estimates of fat and gross energy (GE) were regressed against each condition indicator using regression. Subcutaneous fat thickness, a rump BCS, and rLIVINDEX (an arithmetic combination of subcutaneous fat thickness and the rump BCS) were most related to condition for live animals (r² ≥ 0.87, P < 0.001) whereas the Kistner score and kidney fat were most related to fat and GE for dead animals (r² ≥ 0.77, P < 0.001). We also evaluated range of usefulness and sensitivity to small changes in body condition for all models. In general, indices with moderate or highly curvilinear statistical relations to body fat or those based on only one fat depot or a small number of ranking scores will have limitations in their use. Our results identify robust tools for a variety of research and monitoring designs useful for evaluating nutrition's effect on mule deer populations.     

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.     

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.     

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

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

Consequences of migratory strategy on habitat selection by mule deer

Ungulate behavior is often characterized as balancing selection for forage and avoidance of predation risk. Within partially migratory ungulate populations, this balancing occurs across multiple spatial scales, potentially resulting in different exposure to costs and benefits between migrants and residents. We assessed how availability and selection of forage and risk from predators varied between summer ranges of migrant and resident mule deer (Odocoileus hemionus; a species in which individual migratory strategies are generally fixed for life) in 3 study areas in western Montana, USA, during summers 2017–2019. We hypothesized that mule deer would face a tradeoff between selecting forage and avoiding predation risk, and that migration and residency would pose contrasting availability of forage and risk at a broad (summer range) spatial scale. We hypothesized deer exposed to lower forage at a given spatial scale would compensate for reduced availability by increasing selection of forage at the cost of reduced avoidance of predators, a mechanism whereby migrants and residents could potentially achieve similar exposure to forage despite disparate availability. We compared the availability of forage (kcal/m²) and predation risk from wolves (Canis lupus) and mountain lions (Puma concolor) between summer ranges of each migratory strategy, then assessed how selection for those factors at the home range (second order) and within-home range (third order) scales varied using resource selection functions (RSFs). As forage availability increased among mule deer summer ranges and individual home ranges, selection for forage decreased at the second-order (P = 0.052) and third-order (P = 0.081) scales, respectively, but avoidance of predators varied weakly. In 1 study area, summer range of residents contained lower forage and higher risk than summer range of migrants, but residents compensated for this disadvantage through stronger selection of forage and avoidance of risk at finer spatial scales. In the other 2 study areas, summer range of migrants contained lower forage and higher risk than residents, but migrants did not compensate through stronger selection for beneficial resources. The majority of mule deer in our study system were migratory, though the benefits of migration were unclear, suggesting partial migration may persist in populations even when exposure to forage and predation risk appears unequal between strategies.     

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

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