Maternal carryover, winter severity, and brown bear abundance relate to elk demographics.

Ungulates are key components of ecosystems due to their effects on lower trophic levels, role as prey, and value for recreational and subsistence harvests. Understanding factors that drive ungulate population dynamics can inform protection of important habitat and successful management of populations. To ascertain correlates of ungulate population dynamics, we evaluated the effects of five non-exclusive hypotheses on ungulate abundance and recruitment: winter severity, spring nutritional limitation (spring bottleneck), summer-autumn maternal condition carryover, predation, and timber harvest. We used weather, reconstructed brown bear (Ursus arctos) abundance, and timber harvest data to estimate support for these hypotheses on early calf recruitment (calves per 100 adult females in July-August) and population counts of Roosevelt elk (Cervus canadensis roosevelti) on Afognak and Raspberry islands, Alaska, USA, 1958-2020. Increasing winter temperatures positively affected elk abundance, supporting the winter severity hypothesis, while a later first fall freeze had a positive effect on elk recruitment, supporting the maternal carry-over hypothesis. Increased brown bear abundance was negatively associated with elk recruitment, supporting the predation hypothesis. Recruitment was unaffected by spring climate conditions or timber harvest. Severe winter weather likely increased elk energy deficits, reducing elk survival and subsequent abundance in the following year. Colder and shorter falls likely reduced late-season forage, resulting in poor maternal condition which limited elk recruitment more than winter severity or late-winter nutritional bottlenecks. Our results additionally demonstrated potential negative effects of brown bears on elk recruitment. The apparent long-term decline in elk recruitment did not result in a decline of abundance, which suggests that less severe winters may increase elk survival and counteract the potential effects of predation on elk abundance.

Variable effects of wolves on niche breadth and density of intraguild competitors.

The parallel niche release hypothesis (PNR) indicates that reduced competition with dominant competitors results in greater density and niche breadth of subordinate competitors and which may support an adaptive advantage.We assessed support for the PNR by evaluating relationships between variation in niche breadth and intra- and interspecific density (an index of competition) of wolves (Canis lupus) coyotes (C. latrans), and bobcats (Lynx rufus).We estimated population density (wolf track surveys, coyote howl surveys, and bobcat hair snare surveys) and variability in space use (50% core autocorrelated kernel density home range estimators), temporal activity (hourly and overnight speed), and dietary (isotopic δ13C and δ15N) niche breadth of each species across three areas of varying wolf density in the Upper Peninsula of Michigan, USA, 2010-2019.Densities of wolves and coyotes were inversely related, and increased variability in space use, temporal activity, and dietary niche breadth of coyotes was associated with increased coyote density and decreased wolf density supporting the PNR. Variability in space use and temporal activity of wolves and dietary niche breadth of bobcats also increased with increased intraspecific density supporting the PNR.Through demonstrating decreased competition between wolves and coyotes and increased coyote niche breadth and density, our study provides multidimensional support for the PNR. Knowledge of the relationship between niche breadth and population density can inform our understanding of the role of competition in shaping the realized niche of species.

Subsidies from anthropogenic resources alter diet, activity, and ranging behavior of an apex predator (Canis lupus).

Acquisition of resources can be costly and individuals are predicted to optimize foraging strategies to maximize net energy gain. Wolves (Canis lupus) would be expected to scavenge on subsidies from anthropogenic resources when these resources provide an energetic benefit over the capture of wild prey. We examined the effects of subsidies from anthropogenic resources in the form of livestock carcass dumps (LCDs) on wolf space use, activity, tortuosity, and diet in portions of North America's northern hardwood/boreal ecosystem. We fitted 19 wolves with global positioning system collars during May-August of 2009-2011 and 2013-2015. Wolves with LCDs within their home ranges used areas adjacent to LCDs greater than non-LCD sites and had decreased home ranges and activity as compared to wolves without LCDs in their home ranges. Additionally, cattle comprised at least 22% of wolf diet from scavenging in areas with LCDs present as compared to no cattle in the diet of wolves without access to LCDs. Subsidies from anthropogenic resources in the form of LCDs can serve as attractants for wolves and alter wolf diet, activity, and ranging behavior. Apex predators may alter their behavior where subsidies from anthropogenic resources occur and management of these subsidies should be considered when attempting to reduce the impacts of humans on wolf behavior.

Female American black bears do not alter space use or movements to reduce infanticide risk.

Infanticide occurs in a variety of animal species and infanticide risk has large implications for the evolution of behavior. Further, the sex hypothesis of sexual segregation predicts that for species in which infanticide occurs, females with dependent young will avoid males to reduce risk of sexually-selected infanticide. Infanticide risk-avoidance behavior has been studied primarily in social species, but also occurs in some solitary species. We used generalized linear mixed models to determine if space use and movements of female American black bears (Ursus americanus) during the breeding season were consistent with the sex hypothesis of sexual segregation in the Upper Peninsula of Michigan, USA. Space use and movements of female black bears (n = 16) were not consistent with avoidance behavior to reduce sexually-selected infanticide risk. Females with cubs occupied core areas (mean = 4.64 km2, standard error [SE] = 1.28) and home ranges (mean = 19.46 km2, SE = 5.10) of similar size to females without cubs (core area [mean = 4.11 km2, SE = 0.59]; home range [mean = 16.07 km2, SE = 2.26]), and those core areas and home ranges were not in areas with lesser relative probability of male use. Additionally, females with cubs did not reduce movements during times of day when male movements were greatest. As female bears do avoid potentially infanticidal males in populations with greater levels of infanticide, female black bears may exhibit variation in avoidance behavior based on the occurrence of infanticide.

When top predators become prey: Black bears alter movement behaviour in response to hunting pressure.

The trade-off between predator avoidance and foraging is a key decision making factor that shapes an organism's adaptive behaviour and movement patterns. Human hunters act as top predators to influence the behaviour of free-ranging mammals, including large carnivorous species such as black bears (Ursus americanus). Analysing the effects of hunting on animal behavioural patterns is essential for understanding the extent to which animals detect and respond to human-induced disturbances. To this end, we assessed whether black bear movement behaviour changed with varying risk from spatially and temporally heterogeneous human predation. Levels of risk were categorized as either low (disturbance from dog training; n=19 bears) or high (disturbance from hunting activities; n=11 bears). Road types were either paved (risk due to vehicles) or non-paved (risk due to hunters) and were used as proxies for hunting effort and amount of disturbance. We began by testing the null hypothesis that bears' distribution before the onset of human disturbance is spatially random. Next, to test temporal movement adjustment between the low and high risk levels, we measured the distance to the nearest road and the road crossing frequency using mixed effects models with risk level, time of day and sex as predictor variables. As disturbance near non-paved roads increased due to the start of the hunting activity, the mean distances of bears to non-paved roads increased while the mean distances of bears to paved roads decreased, despite the continual risk of vehicle collision. These behavioural responses were observed during day and night, with the frequency of crossing paved roads at night five times greater than in daytime during the hunting season. Our findings demonstrate that black bears are able to detect risky places and adjust their spatial movements accordingly. More specifically, bears can perceive changes in the level of risk from human hunting activities on a fine temporal scale.

Scale Dependence of Female Ungulate Reproductive Success in Relation to Nutritional Condition, Resource Selection and Multi-Predator Avoidance.

Female ungulate reproductive success is dependent on the survival of their young, and affected by maternal resource selection, predator avoidance, and nutritional condition. However, potential hierarchical effects of these factors on reproductive success are largely unknown, especially in multi-predator landscapes. We expanded on previous research of neonatal white-tailed deer (Odocoileus virginianus) daily survival within home ranges to assess if resource use, integrated risk of 4 mammalian predators, maternal nutrition, winter severity, hiding cover, or interactions among these variables best explained landscape scale variation in daily or seasonal survival during the post-partum period. We hypothesized that reproductive success would be limited greater by predation risk at coarser spatiotemporal scales, but habitat use at finer scales. An additive model of daily non-ideal resource use and maternal nutrition explained the most (69%) variation in survival; though 65% of this variation was related to maternal nutrition. Strong support of maternal nutrition across spatiotemporal scales did not fully support our hypothesis, but suggested reproductive success was related to dam behaviors directed at increasing nutritional condition. These behaviors were especially important following severe winters, when dams produced smaller fawns with less probability of survival. To increase nutritional condition and decrease wolf (Canis lupus) predation risk, dams appeared to place fawns in isolated deciduous forest patches near roads. However, this resource selection represented non-ideal resources for fawns, which had greater predation risk that led to additive mortalities beyond those related to resources alone. Although the reproductive strategy of dams resulted in greater predation of fawns from alternative predators, it likely improved the life-long reproductive success of dams, as many were late-aged (>10 years old) and could have produced multiple litters of fawns. Our study emphasizes understanding the scale-dependent hierarchy of factors limiting reproductive success is essential to providing reliable knowledge for ungulate management.

Effects of maternal nutrition, resource use and multi-predator risk on neonatal white-tailed deer survival.

Growth of ungulate populations is typically most sensitive to survival of neonates, which in turn is influenced by maternal nutritional condition and trade-offs in resource selection and avoidance of predators. We assessed whether resource use, multi-predator risk, maternal nutritional effects, hiding cover, or interactions among these variables best explained variation in daily survival of free-ranging neonatal white-tailed deer (Odocoileus virginianus) during their post-partum period (14 May-31 Aug) in Michigan, USA. We used Cox proportional hazards mixed-effects models to assess survival related to covariates of resource use, composite predation risk of 4 mammalian predators, fawn body mass at birth, winter weather, and vegetation growth phenology. Predation, particularly from coyotes (Canis latrans), was the leading cause of mortality; however, an additive model of non-ideal resource use and maternal nutritional effects explained 71% of the variation in survival. This relationship suggested that dams selected areas where fawns had poor resources, while greater predation in these areas led to additive mortalities beyond those related to resource use alone. Also, maternal nutritional effects suggested that severe winters resulted in dams producing smaller fawns, which decreased their likelihood of survival. Fawn resource use appeared to reflect dam avoidance of lowland forests with poor forage and greater use by wolves (C. lupus), their primary predator. While this strategy led to greater fawn mortality, particularly by coyotes, it likely promoted the life-long reproductive success of dams because many reached late-age (>10 years old) and could have produced multiple generations of fawns. Studies often link resource selection and survival of ungulates, but our results suggested that multiple factors can mediate that relationship, including multi-predator risk. We emphasize the importance of identifying interactions among biological and environmental factors when assessing survival of ungulates.