Bighorn sheep (Ovis canadensis) can live in extremely harsh environments and subsist on submaintenance diets for much of the year. Under these conditions, energy stored as body fat serves as an essential reserve for supplementing dietary intake to meet metabolic demands of survival and reproduction. We developed equations to predict ingesta-free body fat in bighorn sheep using ultrasonography and condition scores in vivo and carcass measurements postmortem. We then used in vivo equations to investigate the relationships between body fat, pregnancy, overwinter survival, and population growth in free-ranging bighorn sheep in California and Nevada. Among 11 subpopulations that included alpine winter residents and migrants, mean ingesta-free body fat of lactating adult females during autumn ranged between 8.8% and 15.0%; mean body fat for nonlactating females ranged from 16.4% to 20.9%. In adult females, ingesta-free body fat > 7.7% during January (early in the second trimester) corresponded with a > 90% probability of pregnancy and ingesta-free body fat > 13.5% during autumn yielded a probability of overwinter survival > 90%. Mean ingesta-free body fat of lactating females in autumn was positively associated with finite rate of population increase (λ) over the subsequent year in bighorn sheep subpopulations that wintered in alpine landscapes. Bighorn sheep with ingesta-free body fat of 26% in autumn and living in alpine environments possess energy reserves sufficient to meet resting metabolism for 83 days on fat reserves alone. We demonstrated that nutritional condition can be a pervasive mechanism underlying demography in bighorn sheep and characterizes the nutritional value of their occupied ranges. Mountain sheep are capital survivors in addition to being capital breeders, and because they inhabit landscapes with extreme seasonal forage scarcity, they also can be fat reserve obligates. Quantifying nutritional condition is essential for understanding the quality of habitats, how it underpins demography, and the proximity of a population to a nutritional threshold.

Migratory species face well-documented global declines, but the causes of these declines remain unclear. One obstacle to better understanding these declines is uncertainty surrounding how migratory behavior is maintained. Most migratory populations are partially migratory, displaying both migrant and resident behaviors. Theory only provides two possible explanations for this coexistence of migration and residency: either these behaviors are fixed at the individual level or both behaviors are part of a single conditional strategy in which an individual’s migratory status (adoption of migrant or resident behavior) is plastic. Here we test for plasticity in migratory status and tactics (timing, distance, and duration of migration) in a federally endangered mountain caprid, the Sierra Nevada bighorn sheep (Ovis canadensis sierrae Grinnell, 1912). We used nonlinear modeling to quantitatively describe migratory behavior, analyzing 262 animal-years of GPS location data collected between 2005 and 2016 from 161 females across 14 subpopulations. Migratory tactics and prevalence varied by subpopulation. On average, individuals from partially migratory subpopulations switched migratory status every 4 years. Our results support the hypothesis that partial migration is maintained through a single conditional strategy. Understanding plasticity in migratory behavior will improve monitoring efforts and provide a rigorous basis for evaluating threats, particularly those associated with changing climate.

Predation can disproportionately affect endangered prey populations when generalist predators are numerically linked to more abundant primary prey. Apparent competition, the term for this phenomenon, has been increasingly implicated in the declines of endangered prey populations. We examined the potential for apparent competition to limit the recovery of Sierra Nevada bighorn sheep (Ovis canadensis sierrae), an endangered subspecies under the US Endangered Species Act. Using a combination of location, demographic, and habitat data, we assessed whether cougar (Puma concolor) predation on endangered bighorn sheep was a consequence of their winter range overlap with abundant mule deer (Odocoileus hemionus). Consistent with the apparent competition hypothesis, bighorn sheep populations with higher spatial overlap with deer exhibited higher rates of cougar predation which had additive effects on adult survival. Bighorn sheep killed by cougars were primarily located within deer winter ranges, even though those areas constituted only a portion of the bighorn sheep winter ranges. We suspect that variation in sympatry between bighorn sheep and deer populations was largely driven by differences in habitat selection among bighorn sheep herds. Indeed, bighorn sheep herds that experienced the highest rates of predation and the greatest spatial overlap with deer also exhibited the strongest selection for low elevation habitat. Although predator-mediated apparent competition may limit some populations of bighorn sheep, it is not the primary factor limiting all populations, suggesting that the dynamics of different herds are highly idiosyncratic. Management plans for endangered species should consider the spatial distributions of key competitors and predators to reduce the potential for apparent competition to hijack conservation success.

Survival in small populations (e.g., Sierra Nevada bighorn sheep or Sierra bighorn [Ovis canadensis sierrae]) is often highly variable. External selective pressures vary in the degree to which they regulate survival by sex and age class. Understanding the important factors and risks for different demographic classes helps managers design strategies that enhance the recovery of endangered species, including Sierra bighorn. Our goal was to determine what population‐level factors (e.g., climate, habitat, population size, predation) affect survival and whether there are interactions between these factors by age and sex, and then apply our findings to recovery strategies. To this end, we conducted a known‐fate survival analysis for female and male Sierra bighorn with data collected over 12 years, and used model selection to evaluate models with spatial, environmental, and other population‐level factors hypothesized to be related to survival. Survival of adult Sierra bighorn declined continuously with age for both sexes; survival was generally higher for females than males, and there were no interactions between age and any environmental or population‐level factors. The top model for both sexes included the date of peak value of normalized difference vegetation index (NDVI) from the previous summer; NDVI had a similar positive relationship with survival for both sexes, which indicates that the later the growing season persists into the summer, the better survival the subsequent year. For females, survival also was negatively related to an index of abundance for mountain lions (Puma concolor), whereas the relationship was less apparent for males. Instead, top models for males indicated elevated survival during warm wet years, but years with late peaks in NDVI the previous year ameliorated the effect of a cold, dry winter. Finally, competitive models for males and females included a variable representing avalanche risk, indicating reduced survival in areas with increased avalanche risk. From a recovery management perspective, the lack of any interaction between age and other covariates suggests that although we may still select younger female Sierra bighorn for translocations (an essential recovery action) because they have higher reproductive value than old females, there were no additional negative synergies between age and other factors to consider. All variables are of value in guiding expectations for newly established populations and established source populations and some may help fine tune the selection of translocation areas. In addition, including predation, weather covariates, and catastrophic effects, such as avalanche risk, in projection models is important for realistic estimation of the time required to meet recovery goals and predicting population trajectories under likely climate change scenarios. Our approach is generalizable to other systems; we demonstrated how survival analyses can inform endangered species recovery management by indicating ideal areas for translocations and provided realist...

Wildlife capture, and the data collection associated with it, has led to major advancements in ecology that are integral to decision making pertaining to wildlife conservation. Capturing wildlife, however, can cause lethal and non-lethal risks to animals. Understanding the factors that contribute to the level of risk involved in wildlife capture is therefore important for the development and implementation of the safest and most effective methodologies. We used data from 736 animal captures of 389 individuals for 2 subspecies of female bighorn sheep (Rocky Mountain bighorn [Ovis canadensis canadensis], Sierra Nevada bighorn sheep [O. c. sierrae]) in Wyoming and California, USA, in 2002-2020 to evaluate the degree and extent of time that capture via helicopter net-gunning affects survival. We compared pre-and post-capture survival during a 10-week window centered on a capture event, and post-capture survival between captured animals and animals that were monitored but not captured during the 10-week window. Additionally, we evaluated the effects of handling techniques (number of times captured, season of capture event, handling time, chase time, and body temp) and biological factors (age and nutritional condition)

1. Energy stores and migration are important adaptations for animals in seasonal environments, but their roles may vary relative to an animal's endogenous and exogenous environment. In partially migratory populations, migrants and residents experience different seasonal environments; thus, the influence of energy stores on survival may differ relative to migratory tactic, with potential consequences to survival and fitness.2. Using data from Sierra Nevada bighorn sheep (Ovis canadensis sierrae; hereafter, Sierra bighorn), we tested the hypothesis that body fat (energy stores) buffers animals against their environment, but that buffering capacity differs across environments experienced by high-elevation residents (using a single range yearround), traditional migrants (making 1 round-trip movement between high-and low-elevation ranges during winter) and vacillating migrants (making ≥2 round trips between high-and low-elevation ranges during winter). We predicted that: for animals with high levels of body fat, survival would be high regardless of migratory tactic; residents would require larger stores of body fat to survive than migrants; energy stores would be least influential to survival for vacillating migrants.3. High levels of body fat in autumn (≥14% for females and ≥19% for males) largely buffered animals against harsh environments (survival >0.90) regardless of migratory tactic. At lower levels of body fat, traditional migrants had higher survival than residents. Vacillating migrants exhibited nearly 100% survival with no detectable effect of body fat on survival. 4. Collectively, these results support the hypothesis that body fat buffers animals against harsh environments but that the buffering capacity differed relative to the environment and highly flexible behaviours (i.e. vacillating migration) can allow animals to decouple survival from body fat. 5. Our work reveals that synergies between physiological and behavioural adaptations of animals in highly seasonal environments carry potential fitness consequences for individuals and demographic consequences for populations.

Translocation of animals into formerly occupied habitat is a key element of the recovery plan for Sierra Nevada bighorn sheep (Ovis canadensis sierrae), which are state (California) and federally listed as endangered. However, implementing Sierra bighorn translocations is a significant conservation challenge because of the small size of the extant population and the limited number of herds available to donate translocation stock. One such herd, the Mt. Langley herd, recently became unusable as a translocation source following a substantial population decline. At the time of listing in 1999, predation by mountain lions (Puma concolor; hereafter lion) was considered a primary threat to Sierra bighorn, and since then lion predation may have continued to limit the ability of source herds to provide translocation stock. We evaluated the relationship between lion predation and ewe survival rates within three source herds of the Southern Recovery Unit, compared lion abundance and ewe survival among years of varying predation levels, provided a range of estimated times for the Mt. Langley herd to recover to its former status as a translocation source, and determined if the rates lions have been removed to mitigate Sierra bighorn predation exceeded sustainable harvest guidelines. We found compelling evidence that lion predation has impeded the recovery of Sierra bighorn by reducing survival rates of adult ewes (and consequently, population growth) and by preying upon individuals that could have otherwise been translocated. Ewe survival was poor during years of extreme predation but even during years of typical predation, survival rates were below a level needed to ensure population growth, indicating that years with little or no lion predation may be necessary for the population to grow and meet recovery goals. Because the intensity of predation was related to lion abundance, monitoring lion populations could provide managers with advance warning of periods of extreme predation. We found that following a period of particularly extreme predation, the Mt. Langley herd decreased in abundance far below the threshold needed to be considered a source of translocation stock, resulting in the loss of approximately 25% of the recovery program’s capacity for translocations. It is unclear how many years it will take for this herd to recover, but management actions to reduce lion predation are likely needed for this herd to grow to a size that can afford to donate individuals to translocation efforts in the near future, even when optimistic growth rates are assumed. We found that lion removal may also be needed to prevent predation from leading to Sierra bighorn population decline. Lion removal rates that have been implemented thus far are well below what would be needed to reduce the abundance the eastern Sierra lion population itself. We recommend continued monitoring of Sierra bighorn and sympatric lions and note that lion removal may be required to facilitate bighorn recovery for the foreseeable future.

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