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.     

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

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

Winter body condition of moose (Alces alces) in a declining population in northeastern Minnesota

Assessments of the condition of moose (Alces alces) may be particularly informative to understanding the dynamics of populations and other influential factors. During February-March 2003 to 2005, we assessed the nutritional condition of 79 moose (39 females, 40 males) in northeastern Minnesota by body condition scoring (BCS(F), scale of 0-10); 67 of these by were assessed by ultrasonographic measurements of rump fat (Maxfat), which was used to estimate ingesta-free body fat (IFBF) in all but two of the females. Scores of the BCS(F) were related (r(2)=0.34, P<0.0001) to Maxfat. Body condition scores were not affected by sex × capture-year, capture-year, or age-at-capture, but the mean body condition score of males (6.5 ± 0.2 [SE], n=40) was less (P ≤ 0.009) than that of females (7.4 ± 0.2, n=39). Overall, Maxfat ranged from 0.0 to 4.6 and 0.3 to 2.8 cm in females and males, respectively, and was unaffected by age-at-capture. There was a sex×capture-year effect (P=0.021) on Maxfat; mean values were stable for males during the winters of 2003 to 2005 but in females were lowest during 2003, consistent with the lowest pregnancy rates and lowest winter and spring survival compared to 2004 and 2005. Based on estimates of percent IFBF, late winter-early spring survival in 2003 of at least 11% of the collared animals assessed by Maxfat, 21% of the adult females, specifically, may have been seriously challenged directly by poor condition. Data from this study provide reference values and assessments of body condition of moose that will be an essential component of the additional, comprehensive research needed to better understand the influence of extrinsic and intrinsic factors on the performance of this viable, but declining, population. For future research, we will concentrate on developing a more-reliable BCS which would allow IFBF estimation once rump fat is depleted.     

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.     

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.     

Linking White-Tailed Deer Density,Nutrition, and Vegetation in a Stochastic Environment

Density-dependent behavior underpins white-tailed deer (Odocoileus virginianus) theory and management application in North America, but strength or frequency of the phenomenon has varied across the geographic range of the species. The modifying effect of stochastic environments and poor-quality habitats on density-dependent behavior has been recognized for ungulate populations around the world, including white-tailed deer populations in South Texas, USA. Despite the importance of understanding mechanisms influencing density dependence, researchers have concentrated on demographic and morphological implications of deer density. Researchers have not focused on linking vegetation dynamics, nutrition, and deer dynamics. We conducted a series of designed experiments during 2004–2012 to determine how strongly white-tailed deer density, vegetation composition, and deer nutrition (natural and supplemented) are linked in a semi-arid environment where the coefficient of variation of annual precipitation exceeds 30%. We replicated our study on 2 sites with thornshrub vegetation in Dimmit County, Texas. During late 2003, we constructed 6 81-ha enclosures surrounded by 2.4-m-tall woven wire fence on each study site. The experimental design included 2 nutrition treatments and 3 deer densities in a factorial array, with study sites as blocks. Abundance targets for low, medium, and high deer densities in enclosures were 10 deer (equivalent to 13 deer/km²), 25 deer (31 deer/km²), and 40 deer (50 deer/km²), respectively. Each study site had 2 enclosures with each deer density. We provided deer in 1 enclosure at each density with a high-quality pelleted supplement ad libitum, which we termed enhanced nutrition; deer in the other enclosure at each density had access to natural nutrition from the vegetation. We conducted camera surveys of deer in each enclosure twice per year and added or removed deer as needed to approximate the target densities. We maintained >50% of deer ear-tagged for individual recognition. We maintained adult sex ratios of 1:1–1:1.5 (males:females) and a mix of young and older deer in enclosures. We used reconstruction, validated by comparison to known number of adult males, to make annual estimates of density for each enclosure in analysis of treatment effects. We explored the effect of deer density on diet composition, diet quality, and intake rate of tractable female deer released into low- and high-density enclosures with natural nutrition on both study sites (4 total enclosures) between June 2009 and May 2011, 5 years after we established density treatments in enclosures. We used the bite count technique and followed 2–3 tractable deer/enclosure during foraging bouts across 4 seasons. Proportion of shrubs, forbs, mast, cacti, and subshrubs in deer diets did not differ (P > 0.57) between deer density treatments. Percent grass in deer diets was higher (P = 0.05) at high deer density but composed only 1.3 ± 0.3% (SE) of the diet. Digestible protein and metabolizable energy of diets were similar (P > 0.45) between deer density treatments. Likewise, bite rate, bite size, and dry matter intake did not vary (P > 0.45) with deer density. Unlike deer density, drought had dramatic (P ≤ 0.10) effects on foraging of tractable deer. During drought conditions, the proportion of shrubs and flowers increased in deer diets, whereas forbs declined. Digestible protein was 31%, 53%, and 54% greater (P = 0.06) during non-drought than drought during autumn, winter, and spring, respectively. We studied the effects of enhanced nutrition on the composition and quality of tractable female deer diets between April 2007 and February 2009, 3 years after we established density treatments in enclosures. We also estimated the proportion of supplemental feed in deer diets. We used the 2 low-density enclosures on each study site, 1 with enhanced nutrition and 1 with natural nutrition (4 total enclosures). We again used the bite count technique and 2–3 tractable deer living in each enclosure. We estimated proportion of pelleted feed in diets of tractable deer and non-tractable deer using ratios of stable isotopes of carbon. Averaged across seasons and nutrition treatments, shrubs composed a majority of the vegetation portion of deer diets (44%), followed by mast (26%) and forbs (15%). Enhanced nutrition influenced the proportion of mast, cacti, and flowers in the diet, but the nature and magnitude of the effect varied by season and year. The trend was for deer in natural-nutrition enclosures to eat more mast. We did not detect a statistical difference (P = 0.15) in the proportion of shrubs in diets between natural and enhanced nutrition, but deer with enhanced nutrition consumed 7–24% more shrubs in 5 of 8 seasons. Deer in enhanced-nutrition enclosures had greater (P = 0.03) digestible protein in their overall diet than deer in natural-nutrition enclosures. The effect of enhanced nutrition on metabolizable energy in overall diets varied by season and was greater (P < 0.04) for enhanced-nutrition deer during summer and autumn 2007 and winter 2008. In the enhanced-nutrition treatment, supplemental feed averaged 47–80% of the diet of tractable deer. Of non-tractable deer in all density treatments with enhanced nutrition, 97% (n = 128 deer) ate supplemental feed. For non-tractable deer averaged across density treatments, study sites, and years, percent supplemental feed in deer diets exceeded 70% for all sex and age groups. We determined if increasing deer density and enhanced nutrition resulted in a decline in preferred forbs and shrubs and an increase in plants less preferred by deer. We sampled all 12 enclosures via 20, 50-m permanent transects in each enclosure. Percent canopy cover of preferred forbs was similar (P = 0.13) among deer densities averaged across nutrition treatments and sampling years (low density: = 8%, SE range 6–10; medium density: 5%, 4–6; high density: 4%, 3–5; SE ranges are presented because SEs associated with backtransformed means are asymetrical). Averaged across deer densities, preferred forb canopy cover was similar between nutrition treatments in 2004; but by 2012 averaged 20 ± 17–23% in enhanced-nutrition enclosures compared to 10 ± 8–13% in natural-nutrition enclosures (P = 0.107). Percent canopy cover of other forbs, preferred shrubs, other shrubs, and grasses, as well as Shannon's index, evenness, and species richness were similar (P > 0.10) among deer densities, averaged across nutrition treatments and sampling years. We analyzed fawn:adult female ratios, growth rates of fawns and yearlings, and survival from 6 to 14 months of age and for adults >14 months of age. We assessed adult body mass and population growth rates (lambda apparent, λ_APP) to determine density and nutrition effects on deer populations in the research enclosures during 2004–2012. Fawn:adult female ratios declined (P = 0.04) from low-medium density to high density in natural-nutrition enclosures but were not affected (P = 0.48) by density in enhanced nutrition enclosures although, compared to natural nutrition, enhanced nutrition increased fawn:adult female ratios by 0.15 ± 0.12 fawns:adult female at low-medium density and 0.44 ± 0.17 fawns:adult female at high density. Growth rate of fawns was not affected by deer density under natural or enhanced nutrition (P > 0.17) but increased 0.03 ± 0.01 kg/day in enhanced-nutrition enclosures compared to natural nutrition (P < 0.01). Growth rate of yearlings was unaffected (P > 0.71) by deer density, but growth rate increased for males in some years at some density levels in enhanced-nutrition enclosures. Adult body mass declined in response to increasing deer density in natural-nutrition enclosures for both adult males (P < 0.01) and females (P = 0.10). Enhanced nutrition increased male body mass, but female mass did not increase compared to natural nutrition. Survival of adult males was unaffected by deer density in natural- (P = 0.59) or enhanced- (P = 0.94) nutrition enclosures. Survival of adult females was greatest in medium-density enclosures with natural nutrition but similar at low and high density (P = 0.04). Enhanced nutrition increased survival of females (P < 0.01) and marginally for males (P = 0.11). Survival of fawns 6–14 months old was unaffected (P > 0.35) by density in either natural- or enhanced-nutrition treatments but was greater (P = 0.04) under enhanced nutrition. Population growth rate declined (P = 0.06) with increasing density in natural-nutrition enclosures but not (P = 0.55) in enhanced nutrition. Enhanced nutrition increased λ_APP by 0.32. Under natural nutrition, we found only minor effects of deer density treatments on deer diet composition, nutritional intake, and plant communities. However, we found density-dependent effects on fawn:adult female ratios, adult body mass, and population growth rate. In a follow-up study, deer home ranges in our research enclosures declined with increasing deer density. We hypothesized that habitat quality varied among home ranges and contributed to density-dependent responses. Variable precipitation had a greater influence on deer diets, vegetation composition, and population parameters than did deer density. Also, resistance to herbivory and low forage quality of the thornshrub vegetation of our study sites likely constrained density-dependent behavior by deer. We posit that it is unlikely that, at our high-density (50 deer/km²) and perhaps even medium-density (31 deer/km²) levels, negative density dependence would occur without several wet years in close association. In the past century, this phenomenon has only happened once (1970s). Thus, density dependence would likely be difficult to detect in most years under natural nutrition in this region. Foraging by deer with enhanced nutrition did not result in a reduction in preferred plants in the vegetation community and had a protective effect on preferred forbs because ≤53% of deer diets consisted of vegetation. However, enhanced nutrition improved fitness of individual deer and deer populations, clearly demonstrating that nutrition is limiting for deer populations under natural conditions in western South Texas. © 2019 The Authors. Wildlife Monographs published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Survival,cause-specific mortality,and population growth of white-tailed deer in western Virginia

Understanding the role of recruitment in population dynamics of white-tailed deer (Odocoileus virginianus) is important for management. In the central Appalachian Mountains, deer are part of a largely forested ecosystem that supports 3 carnivore species thought to be capable of influencing white-tailed deer recruitment: black bears (Urus americanus), coyotes (Canis latrans), and bobcats (Lynx rufus). Yet little is known about predation, how other environmental factors influence recruitment, or the importance of neonate survival to white-tailed deer population performance in the region. Our objectives were to identify causes of mortality for neonates, analyze effects of landscape attributes on survival of neonates, estimate survival rates for neonates and adult female white-tailed deer, and to model population growth trends based on current vital rates and hypothetical harvest and neonate survival scenarios. During 2019–2020, we captured 57 neonate deer in Bath County, Virginia, USA, by monitoring 38 pregnant females equipped with global positioning system collars and vaginal implant transmitters and by conducting transect searches for recently born neonates. We observed 37 neonate mortalities and identified cause of death using field and genetic evidence. Mortalities included 28 predation events and 9 deaths from other causes (e.g., abandonment, malnutrition, disease). Black bears accounted for 48.6% of neonate mortalities, and 64.2% of predation events (n = 18), followed by bobcats (n = 5) and coyotes (n = 3). Annual survival for adult female deer was 0.871 and neonate survival to 12 weeks old was 0.310. Elevation was a significant predictor of neonate survival; mortality risk increased 20% for every 100-m increase in elevation. Models of annual population growth using observed vital rates predicted an increasing population (λ = 1.10). A 10% increase in female harvest would still result in a potential population increase of 2% (λ = 1.02), but a 20% increase in harvest rate would result in a potential 7% decline (λ = 0.93). Neonate survival was higher near fertile valley bottoms and lower along forested ridges characterized by shallow, infertile soils and limited edge or early successional forests. While predation, largely influenced by black bears, was the leading cause of neonate mortality and contributed to low neonate survival, we observed little evidence of population decline, and suggest there is opportunity for a modest increase in harvest of female deer.     

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.     

White-Tailed Deer Population Dynamics Following Louisiana Black Bear Recovery

Changing predator communities have been implicated in reduced survival of white-tailed deer (Odocoileus virginianus) fawns. Few studies, however, have used field-based age-specific estimates for survival and fecundity to assess the relative importance of low fawn survival on population growth and harvest potential. We studied white-tailed deer population dynamics on Tensas River National Wildlife Refuge (TRNWR) in Louisiana, USA, where the predator community included bobcats (Lynx rufus), coyotes (Canis latrans), and a restored population of Louisiana black bear (Ursus americanus luteolus). During 2013–2015, we radio-collared and monitored 70 adult (≥2.5 yrs) and 21 yearling (1.5-yr-old) female deer. Annual survival averaged 0.815 (95% CI = 0.734–0.904) for adults and 0.857 (95% CI = 0.720–1.00) for yearlings. We combined these estimates with concurrently collected fawn survival estimates (0.27; 95% CI = 0.185–0.398) to model population trajectories and elasticities. We used estimates of nonhunting survival (annual survival estimated excluding harvest mortality) to project population growth (λ) relative to 4 levels of harvest (0, 10%, 20%, 30%). Finally, we investigated effects of reduced fawn survival on population growth under current management and with elimination of female harvest. Despite substantial fawn predation, the deer population on TRNWR was increasing (λ = 1.06) and could sustain additional female harvest; however, the population was expected to decline at 20% (λ = 0.98) and 30% (λ = 0.94) female harvest. With no female harvest, the population was projected to increase with observed (λ = 1.15) and reduced fawn survival (λ = 1.02), but the population could not sustain current female harvest (10%) if fawn survival declined (λ = 0.90). For all scenarios, adult female survival was the most elastic parameter. Given the importance of adult female survival, the relative predictability in response of adult survival to harvest management, and the difficulty in altering fawn survival, reducing female harvest is likely the most efficient approach to compensate for low fawn survival. On highly productive sites such as ours, reduction, but not necessarily elimination, of harvest can mitigate effects of low fawn survival on population growth. © 2020 The Wildlife Society.     

Postfledging Survival of Laysan Ducks

ABSTRACT Precise and unbiased estimates of demographic parameters are necessary for effective population monitoring and to parameterize population models (e.g., population viability analyses). This is especially important for endangered species, where recovery planning and managers' decisions can influence species persistence. In this study, we used mark—recapture methods to estimate survival of fledged juveniles (hatch-yr [HY]) and adult (after-hatch-yr [AHY]) Laysan ducks (Anas laysanensis), an endangered anatid restricted to Laysan Island in the northwestern Hawaiian Islands. To better understand population dynamics, we examined how survival varied as a function of Laysan duck density during 1998–2004. Using random effects models, we also quantified process variation in survival, thereby quantifying the appropriate source of variation for future population models. The dataset supported variation in survival that was time (yr), age (AHY vs. HY), and sex specific. Due to small sample sizes, we did not examine time specificity in the survival of HY ducks. Survival of HY ducks was 0.832 (SE = 0.087) for females (n = 21) and 0.999 (SE < 0.001) for males (n = 15) during 1998–2001. Trends in time and density lacked support as sources of variation in the survival of AHY ducks during 1998–2004. After-hatch-year survival ranged from 0.792 (SE = 0.033) to 0.999 (SE < 0.001). Where we modeled survival as a random effect, annual survival for AHY females was 0.881 (SE = 0.017) and process variation (σ_s) was 0.034. For AHY males, annual survival (μ_s) was 0.906 (SE = 0.019) and process variation (σ_s) was 0.040. This information will improve existing population viability analysis models for Laysan ducks. We believe that monitoring the source and translocation populations will be paramount for increasing our understanding of Laysan duck dynamics, recovery planning, and population management.     

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.     

Habitat and weather influence body condition in white-tailed deer,Wisconsin, USA

In temperate and northern ecosystems where there are pronounced seasonal patterns in weather and available energy, there are corresponding patterns of body condition among white-tailed deer (Odocoileus virginianus). Body condition of white-tailed deer can affect survival and reproduction, which has large repercussions for state-level natural resource agencies that allocate hunting permits. In this study, we investigated how variation in winter weather, spring phenology, habitat composition, and browse quantity affected white-tailed deer body condition across a large spatial scale. Several body condition indicators (e.g., carcass mass, heart fat, antler size) were measured by hunters for 795 deer during September–December 2016–2018 in Wisconsin, USA. Winter severity in the previous year was an unreliable predictor of fall body condition of deer when winters were considered mild or moderate. The timing of spring green-up had a consistent effect on the body condition of all age and sex classes of deer. Earlier spring green-up resulted in heavier fawns and larger antlers among adult males. Region and spring green-up interacted to affect the heart fat of adult females. Earlier springs resulted in adult females in northern and central Wisconsin having a higher probability of heavy heart fat, whereas spring green-up had no effect on adult female heart fat in southern Wisconsin. Effects of habitat differed by age and sex class of deer, and by the body condition metric being evaluated, indicating that there are important physiological differences among age and sex classes of deer that are affected by the environment. Our study demonstrates that the hunting public can contribute large-scale, cost-effective, and quality data to deer monitoring and research projects. It is important that natural resource agencies be able to identify and recruit highly engaged members of the hunting public to ensure project success. The timing of spring green-up can have lasting effects on deer health that can be consistently observed the following fall, which is in contrast to the effects of winter severity that did not appear to persist when previous winters were mild or moderate. We encourage managers in northern or temperate regions to consider measures of spring green-up timing in conjunction with traditional winter severity when making deer population management decisions, such as antlerless tag allocation.     

Survival of white-tailed deer fawns in the grasslands of the northern Great Plains

Environmental factors, such as forest characteristics, have been linked to fawn survival in eastern and southern white-tailed deer (Odocoileus virginianus) populations. In the Great Plains, less is known about how intrinsic and habitat factors influence fawn survival. During 2007–2009, we captured and radiocollared 81 fawns in north-central South Dakota and recorded 23 mortalities, of which 18 died before 1 September. Predation accounted for 52.2% of mortality; remaining mortality included human (hunting, vehicle, and farm accident; 26.1%) and hypothermia (21.7%). Coyotes (Canis latrans) accounted for 83.3% of predation on fawns. We used known-fate analysis in Program MARK to estimate summer (15 May–31 Aug) survival rates and investigated the influence of intrinsic and habitat variables on survival. We developed 2 a priori model sets, including intrinsic variables and a test of annual variation in survival (model set 1) and habitat variables (model set 2). Model set 1 indicated that summer survival varied among years (2007–2009); annual survival rates were 0.94 (SE = 0.06, n = 22), 0.78 (SE = 0.09, n = 27), and 0.54 (SE = 0.10, n = 32), respectively. Model set 2 indicated that survival was further influenced by patch density of cover habitats (Conservation Reserve Program [CRP]-grasslands, forested cover, and wetlands). Mean CRP-grassland and wetland patch density (no. patches/100 ha) were greater (P < 0.001) in home-range areas of surviving fawns ( = 1.81, SE = 0.10, n = 63; = 1.75, SE = 0.14, n = 63, respectively) than in home-range areas of fawns that died ( = 0.16, SE = 0.04, n = 18; = 1.28, SE = 0.10, n = 18, respectively). Mean forested cover patch density was less (P < 0.001) in home-range areas of surviving fawns ( = 0.77, SE = 0.10, n = 63) than in home-range areas of fawns that died ( = 1.49, SE = 0.21, n = 18). Our results indicate that management activities should focus on CRP-grassland and wetland habitats in order to maintain or improve fawn survival in the northern Great Plains, rather than forested cover composed primarily of tree plantings and shelterbelts. © 2012 The Wildlife Society.     

Mortality of Rocky Mountain elk in Michigan due to meningeal worm

Mortality from cerebrospinal parelaphostrongylosis caused by the meningeal worm (Parelaphostrongylus tenuis) has been hypothesized to limit elk (Cervus elaphus nelsoni) populations in areas where elk are conspecific with white-tailed deer (Odocoileus virginianus). Elk were reintroduced into Michigan (USA) in the early 1900s and subsequently greatly increased population size and distribution despite sympatric high-density (>or=12/km2) white-tailed deer populations. We monitored 100 radio-collared elk of all age and sex classes from 1981-94, during which time we documented 76 mortalities. Meningeal worm was a minor mortality factor for elk in Michigan and accounted for only 3% of mortalities, fewer than legal harvest (58%), illegal kills (22%), other diseases (7%), and malnutrition (4%). Across years, annual cause-specific mortality rates due to cerebrospinal parelaphostrongylosis were 0.033 (SE=0.006), 0.029 (SE=0.005), 0.000 (SE=0.000), and 0.000 (SE=0.000) for calves, 1-yr-old, 2-yr-old, and >or=3-yr-old, respectively. The overall population-level mortality rate due to cerebrospinal parelaphostrongylosis was 0.009 (SE=0.001). Thus, meningeal worm had little impact on elk in Michigan during our study despite greater than normal precipitation (favoring gastropods) and record (>or=14 km2) deer densities. Further, elk in Michigan have shown sustained population rates-of-increase of >or=18%/yr and among the highest levels of juvenile production and survival recorded for elk in North America, indicating that elk can persist in areas with meningeal worm at high levels of population productivity. It is likely that local ecologic characteristics among elk, white-tailed deer, and gastropods, and degree of exposure, age of elk, individual and population experience with meningeal worm, overall population vigor, and moisture determine the effects of meningeal worm on elk populations.     

Survival and cause-specific mortality rates of female sika deer in eastern Hokkaido, Japan

Survival and cause-specific mortality rates of female sika deer (Cervus nippon) were studied using radio telemetry in eastern Hokkaido, Japan. We captured and radio-collared 18 female deer, and monitored their survival from April 1993 to May 1996. Estimated annual survival rate for adult females was 0.779 (95% confidence interval was 0.609–0.997). The harvest mortality rate of adult females was higher than the natural mortality rate. Experimental female hunting during 1994–1996 contributed to an increase in the mortality rate for females and was useful in the control of the sika deer population.     

Influence of body mass and environmental conditions on winter mortality risk of a northern ungulate: Evidence for a late‐winter survival bottleneck

1. A relationship between winter weather and survival of northern ungulates has long been established, yet the possible roles of biological (e.g., nutritional status) and environmental (e.g., weather) conditions make it important to determine which potential limiting factors are most influential.
2. Our objective was to examine the potential effects of individual (body mass and age) and extrinsic (winter severity and snowmelt conditions) factors on the magnitude and timing of mortality for adult (>2.5 years old) female white‐tailed deer (Odocoileus virginianus [Zimmerman, 1780]) during February–May in the Upper Peninsula of Michigan, USA.
3. One hundred and fifty deer were captured and monitored during 2009–2015 in two areas with varying snowfall. February–May survival ranged from 0.24 to 0.89 (mean = 0.69) across years. Mortality risk increased 1.9% with each unit increase in cumulative winter severity index, decreased 8.2% with each cumulative snow‐free day, and decreased 4.3% with each kg increase in body mass. Age and weekly snow depth did not influence weekly deer survival. Predation, primarily from coyote (Canis latrans [Say, 1823]) and wolves (Canis lupus [L., 1758]), accounted for 78% of known‐cause mortalities.
4. Our results suggest that cumulative winter severity, and possibly to a lesser degree deer condition entering winter, impacted deer winter survival. However, the timing of spring snowmelt appeared to be the most influential factor determining late‐winter mortality of deer in our study. This supports the hypothesis that nutrition and energetic demands from weather conditions are both important to northern ungulate winter ecology. Under this model, a delay of several weeks in the timing of spring snowmelt could exert a large influence on deer survival, resulting in a survival bottleneck.

     

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

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

Estimating survival in the Apennine brown bear accounting for uncertainty in age classification

For most rare and elusive species, estimating age-specific survival is a challenging task, although it is an important requirement to understand the drivers of population dynamics, and to inform conservation actions. Apennine brown bears Ursus arctos marsicanus are a small, isolated population under a severe risk of extinction, for which the main demographic mechanisms underlying population dynamics are still unknown, and population trends have not been formally assessed. We present a 12-year analysis of their survival rates using non-invasive genetic sampling data collected through four different sampling techniques. By using multi-event capture–recapture models, we estimated survival probabilities for two broadly defined age classes (cubs and older individuals), even though the age of the majority of sampled bears was unknown. We also applied the Pradel model to provide a preliminary assessment of population trend during the study period. Survival was different between cubs [ϕ = 0.51, 95% CI (0.22, 0.79)], adult males [ϕ = 0.85, 95% CI (0.76, 0.91)] and adult females [ϕ = 0.92, 95% CI (0.87, 0.95)], no temporal variation in survival emerged, suggesting that bear survival remained substantially stable throughout the study period. The Pradel analysis of population trend yielded an estimate of λ = 1.009 [SE = 0.018; 95% CI (0.974, 1.046)]. Our results indicate that, despite the status of full legal protection, the basically stable demography of this relict population is compatible with the observed lack of range expansion, and that a relatively high cub mortality could be among the main factors depressing recruitment and hence population growth.     

A review of vital rates and cause‐specific mortality of elk Cervus elaphus populations in eastern North America

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Bighorn Sheep Abundance Following a Suspected Pneumonia Epidemic in Rocky Mountain National Park

Abstract: Anecdotal evidence of a pneumonia epizootic among bighorn sheep (Ovis canadensis canadensis) in Rocky Mountain National Park (RMNP), Colorado, USA, during the mid-1990s prompted park officials to examine the current condition of the herds. Here we present a mark—resight study design to estimate population abundance that, in many circumstances, is a reliable and cost-effective alternative to traditional mark—recapture or to indices of population abundance. We captured 59 adult females and radiocollared them via helicopter net-gunning during winter 2002–2003. From ground resighting surveys conducted May—September, we estimated the total RMNP bighorn population at 389.9 (SE = 34.9, CI = 327.2–464.6) in 2003 and 366.4 (SE = 34.7, CI = 304.4–441.0) in 2004. Previous abundance estimates suggest a park-wide decline has occurred between the late 1980s and the suspected pneumonia epidemic of the mid-1990s. Although the 2 years of data from our study are not enough to predict whether the herds are capable of recovering to previous levels, they provide park officials the tools necessary to make the most informed decisions for future monitoring and management of this fragile species.