An integrated population model reveals source-sink dynamics for competitively subordinate African wild dogs linked to anthropogenic prey depletion.

Many African large carnivore populations are declining due to decline of the herbivore populations on which they depend. The densities of apex carnivores like the lion and spotted hyena correlate strongly with prey density, but competitively subordinate carnivores like the African wild dog benefit from competitive release when the density of apex carnivores is low, so the expected effect of a simultaneous decrease in resources and dominant competitors is not obvious. Wild dogs in Zambia's South Luangwa Valley Ecosystem occupy four ecologically similar areas with well-described differences in the densities of prey and dominant competitors due to spatial variation in illegal offtake. We used long-term monitoring data to fit a Bayesian integrated population model (IPM) of the demography and dynamics of wild dogs in these four regions. The IPM used Leslie projection to link a Cormack-Jolly-Seber model of area-specific survival (allowing for individual heterogeneity in detection), a zero-inflated Poisson model of area-specific fecundity and a state-space model of population size that used estimates from a closed mark-capture model as the counts from which (latent) population size was estimated. The IPM showed that both survival and reproduction were lowest in the region with the lowest density of preferred prey (puku, Kobus vardonii and impala, Aepyceros melampus), despite little use of this area by lions. Survival and reproduction were highest in the region with the highest prey density and intermediate in the two regions with intermediate prey density. The population growth rate ( λ ) was positive for the population as a whole, strongly positive in the region with the highest prey density and strongly negative in the region with the lowest prey density. It has long been thought that the benefits of competitive release protect African wild dogs from the costs of low prey density. Our results show that the costs of prey depletion overwhelm the benefits of competitive release and cause local population decline where anthropogenic prey depletion is strong. Because competition is important in many guilds and humans are affecting resources of many types, it is likely that similarly fundamental shifts in population limitation are arising in many systems.

A retrospective view of early research on dominance, stress and reproduction in cooperatively breeding carnivores.

Social carnivores have been central in studies of cooperative breeding, and research using noninvasive methods to examine behavioral and endocrine mechanisms of reproductive suppression started in the 1980s with dwarf mongooses in Serengeti National Park. Here, I synthesize the methods, findings and limitations of a research program that examined relationships between social dominance, age, mass, aggression, mating, gonadal steroids, glucocorticoids and reproduction in female and male dwarf mongooses, African wild dogs and wolves. Infanticide is a reliable backstop for reproductive suppression in females, and reproduction is energetically costly in these species. These conditions favor hypothalamic - pituitary - gonadal (HPG) adaptations that reduce the fertility of subordinate females to avoid the cost of producing doomed offspring. Infanticide also favors close synchronization of reproduction when subordinate females do become pregnant. In males, infanticide is a less reliable backstop and reproduction is less costly, so direct effects of subordination on fertility are less pronounced. Age is a strong predictor of social dominance in these species, but the evolutionary reason for this is not clear. In dwarf mongooses and wild dogs, alpha females were never deposed by younger packmates, but alpha males were: this difference is also not understood. Patterns of reproduction supported models predicting that alphas are less likely to share reproduction when the fitness costs of reproduction are high, when the fitness expected for dispersers is low, and with young subordinates to whom they are more closely related. Correlations between dominance and adrenal glucocorticoid concentrations varied between species and sexes, but did not support the hypothesis that chronic stress causes reproductive suppression.

Loss of an apex predator in the wild induces physiological and behavioural changes in prey.

Predators can impact prey via predation or risk effects, which can initiate trophic cascades. Given widespread population declines of apex predators, understanding and predicting the associated ecological consequences is a priority. When predation risk is relatively unpredictable or uncontrollable by prey, the loss of predators is hypothesized to release prey from stress; however, there are few tests of this hypothesis in the wild. A well-studied predator-prey system between white sharks (Carcharodon carcharias) and Cape fur seals (Arctocephalus pusillus pusillus) in False Bay, South Africa, has previously demonstrated elevated faecal glucocorticoid metabolite concentrations (fGCMs) in seals exposed to high levels of predation risk from white sharks. A recent decline and disappearance of white sharks from the system has coincided with a pronounced decrease in seal fGCM concentrations. Seals have concurrently been rafting further from shore and over deeper water, a behaviour that would have previously rendered them vulnerable to attack. These results show rapid physiological and behavioural responses by seals to release from predation stress. To our knowledge, this represents the first demonstration in the wild of physiological changes in prey from predator decline, and such responses are likely to increase given the scale and pace of apex predator declines globally.

Response of lion demography and dynamics to the loss of preferred larger prey.

Large carnivores are experiencing range contraction and population declines globally. Prey depletion due to illegal offtake is considered a major contributor, but the effects of prey depletion on large carnivore demography are rarely tested. We measured African lion density and tested the factors that affect survival using mark-recapture models fit to six years of data from known individuals in Kafue National Park (KNP), Zambia. KNP is affected by prey depletion, particularly for large herbivores that were preferred prey for KNP lions a half-century ago. This provides a unique opportunity to test whether variables that explain local prey density also affect lion survival. Average lion density within our study area was 3.43 individuals/100 km2 (95% CI, 2.79-4.23), which was much lower than lion density reported for another miombo ecosystem with similar vegetation structure and rainfall that was less affected by prey depletion. Despite this, comparison to other lion populations showed that age- and sex-specific survival rates for KNP lions were generally good, and factors known to correlate with local prey density had small effects on lion survival. In contrast, recruitment of cubs was poor and average pride size was small. In particular, the proportion of the population comprised of second-year cubs was low, indicating that few cubs are recruited into the subadult age class. Our findings suggest that low recruitment might be a better signal of low prey density than survival. Thus, describing a lion population's age structure in addition to average pride size may be a simple and effective method of initially evaluating whether a lion population is affected by prey depletion. These dynamics should be evaluated for other lion populations and other large carnivore species. Increased resource protection and reducing the underlying drivers of prey depletion are urgent conservation needs for lions and other large carnivores as their conservation is increasingly threatened by range contraction and population declines.

The control of risk hypothesis: reactive vs. proactive antipredator responses and stress-mediated vs. food-mediated costs of response.

Inducible defences against predators evolve because they reduce the rate of direct predation, but this benefit is offset by the cost (if any) of defence. If antipredator responses carry costs, the effect of predators on their prey is partitioned into two components, direct killing and risk effects. There is considerable uncertainty about the strength of risk effects, the factors that affect their strength, and the mechanisms that underlie them. In some cases, antipredator responses are associated with a glucocorticoid stress response, and in other cases they are associated with trade-offs between food and safety, but there is no general theory to explain this variation. Here, I develop the control of risk (COR) hypothesis, predicting that proactive responses to predictable and controllable aspects of risk will generally have food-mediated costs, while reactive responses to unpredictable or uncontrollable aspects of predation risk will generally have stress-mediated costs. The hypothesis is grounded in laboratory studies of neuroendocrine stressors and field studies of food-safety trade-offs. Strong tests of the COR hypothesis will require more studies of responses to natural variation in predation risk and the physiological consequences of these responses, but its explanatory power can be illustrated with existing case studies.

Physiological stress responses to natural variation in predation risk: evidence from white sharks and seals.

Predators can impact ecosystems through consumptive or risk effects on prey. Physiologically, risk effects can be mediated by energetic mechanisms or stress responses. The predation-stress hypothesis predicts that risk induces stress in prey, which can affect survival and reproduction. However, empirical support for this hypothesis is both mixed and limited, and the conditions that cause predation risk to induce stress responses in some cases, but not others, remain unclear. Unusually clear-cut variation in exposure of Cape fur seals (Arctocephalus pusillus pusillus) to predation risk from white sharks (Carcharodon carcharias) in the waters of Southwestern Africa provides an opportunity to test the predation-stress hypothesis in the wild. Here, we measured fecal glucocorticoid concentrations (fGCM) from Cape fur seals at six discrete islands colonies exposed to spatiotemporal variation in predation risk from white sharks over a period of three years. We found highly elevated fGCM concentrations in seals at colonies exposed to high levels of unpredictable and relatively uncontrollable risk of shark attack, but not at colonies where seals were either not exposed to shark predation or could proactively mitigate their risk through antipredatory behavior. Differences in measured fGCM levels were consistent with patterns of risk at the site and seasonal level, for both seal adults and juveniles. Seal fGCM levels were not correlated with colony population size, density, and geographic location. Investigation at a high risk site (False Bay) also revealed strong correlations between fGCM levels and temporal variation in shark attack rates, but not with shark relative abundance. Our results suggest that predation risk will induce a stress response when risk cannot be predicted and/or proactively mitigated by behavioral responses.

The relationship between direct predation and antipredator responses: a test with multiple predators and multiple prey.

Most species adjust their behavior to reduce the likelihood of predation. Many experiments have shown that antipredator responses carry energetic costs that can affect growth, survival, and reproduction, so that the total cost of predation depends on a trade-off between direct predation and risk effects. Despite these patterns, few field studies have examined the relationship between direct predation and the strength of antipredator responses, particularly for complete guilds of predators and prey. We used scan sampling in 344 observation periods over a four-year field study to examine behavioral responses to the immediate presence of predators for a complete antelope guild (dominated by wildebeest, zebra, and oribi) in Liuwa Plains National Park, Zambia, testing for differences in response to all large carnivores in the ecosystem (lions, spotted hyenas, cheetahs, and African wild dogs). We quantified the proportion that each prey species contributed to the kills made by each predator (516 total kills), used distance sampling on systematic line transects to determine the abundance of each prey species, and combined these data to quantify the per-capita risk of direct predation for each predator-prey pair. On average, antelopes increased their vigilance by a factor of 2.4 when predators were present. Vigilance varied strongly among prey species, but weakly in response to different predators. Increased vigilance was correlated with reduced foraging in a similar manner for all prey species. The strength of antipredator response was not detectably related to patterns of direct predation (n = 15 predator-prey combinations with sufficient data). This lack of correlation has implications for our understanding of the role of risk effects as part of the limiting effect of predators on prey.

Inclusive fitness theory and eusociality.

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Nowak et al. argue that inclusive fitness theory has been of little value in explaining the natural world, and that it has led to negligible progress in explaining the evolution of eusociality. However, we believe that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature. We will focus our comments on three general issues.

Hunting on a hot day: effects of temperature on interactions between African wild dogs and their prey.

As global temperatures increase, interactions between species are affected by changes in distribution, abundance and phenology, but also by changes in behavior. The heat dissipation limitation hypothesis suggests that the ability to dissipate heat commonly limits the activity of endotherms, a problem that should be particularly acute for cursorial predators and their prey in equatorial ecosystems. Allometric relationships suggest that heat dissipation should be a stronger constraint for larger species, so that (smaller) predators should be less affected than (larger) prey. We used data from 266 complete days of direct observation of African wild dogs (Lycaon pictus) in five packs over a period of 2 yr to test how deviations of temperature from that expected for the time of day affected eight measures of hunting effort and success. We found that higher temperatures disadvantaged the prey of wild dogs more than the dogs themselves, with increased hunting success and shorter pursuits on warmer days. Broadly, our results demonstrate that effects of temperature on behavior can alter interactions between species, exacerbating or offsetting the direct effects of climate change.

Assessing the sustainability of African lion trophy hunting, with recommendations for policy.

While trophy hunting provides revenue for conservation, it must be carefully managed to avoid negative population impacts, particularly for long-lived species with low natural mortality rates. Trophy hunting has had negative effects on lion populations throughout Africa, and the species serves as an important case study to consider the balance of costs and benefits, and to consider the effectiveness of alternative strategies to conserve exploited species. Age-restricted harvesting is widely recommended to mitigate negative effects of lion hunting, but this recommendation was based on a population model parameterized with data from a well-protected and growing lion population. Here, we used demographic data from lions subject to more typical conditions, including source-sink dynamics between a protected National Park and adjacent hunting areas in Zambia's Luangwa Valley, to develop a stochastic population projection model and evaluate alternative harvest scenarios. Hunting resulted in population declines over a 25-yr period for all continuous harvest strategies, with large declines for quotas >1 lion/concession (~0.5 lion/1,000 km2 ) and hunting of males younger than seven years. A strategy that combined periods of recovery, an age limit of ≥7 yr, and a maximum quota of ~0.5 lions shot/1,000 km2 yielded a risk of extirpation <10%. Our analysis incorporated the effects of human encroachment, poaching, and prey depletion on survival, but assumed that these problems will not increase, which is unlikely. These results suggest conservative management of lion trophy hunting with a combination of regulations. To implement sustainable trophy hunting while maintaining revenue for conservation of hunting areas, our results suggest that hunting fees must increase as a consequence of diminished supply. These findings are broadly applicable to hunted lion populations throughout Africa and to inform global efforts to conserve exploited carnivore populations.

Long-term data reveal fitness costs of anthropogenic prey depletion for a subordinate competitor, the African wild dog (Lycaon pictus).

Within carnivore guilds, dominant competitors (e.g., lions, Panthera leo) are limited primarily by the density of prey, while subordinate competitors (e.g., African wild dogs, Lycaon pictus) have been limited by the density of dominant competitors. Historically, the fitness and population density of subordinate competitors have not been tightly linked to prey density. However, populations of large herbivores have declined substantially across sub-Saharan Africa due to human impacts, and where prey depletion is severe, fitness costs for competitive subordinates may begin to outweigh the benefits of competitive release. Using long-term intensive monitoring of African wild dogs in Zambia's Luangwa Valley Ecosystem (LVE), we tested the effects of prey depletion on survival and reproduction. We hypothesized that African wild dog fitness would be lower in prey-depleted areas, despite lower lion densities. Our study area included four contiguous regions that varied in protection level, prey density, and lion density. We fit Bayesian Cormack-Jolly-Seber and closed-capture models to estimate effects on survival and population density, and generalized linear models to estimate effects on reproductive success. We found that the LVE is a stronghold for African wild dogs, with an estimated median density of 4.0 individuals/100 km2. Despite this high density, survival and reproduction differed among regions, and both components of fitness were substantially reduced in the region with the lowest prey density. Anthropogenic prey depletion is becoming an important limiting factor for African wild dogs. If prey depletion (or any other form of habitat degradation) becomes severe enough that its fitness costs outweigh the benefits of competitive release, such changes can fundamentally alter the balance between limiting factors for competitively subordinate species.

Inferential consequences of modeling rather than measuring snow accumulation in studies of animal ecology.

It is increasingly common for studies of animal ecology to use model-based predictions of environmental variables as explanatory or predictor variables, even though model prediction uncertainty is typically unknown. To demonstrate the potential for misleading inferences when model predictions with error are used in place of direct measurements, we compared snow water equivalent (SWE) and snow depth as predicted by the Snow Data Assimilation System (SNODAS) to field measurements of SWE and snow depth. We examined locations on elk (Cervus canadensis) winter ranges in western Wyoming, because modeled data such as SNODAS output are often used for inferences on elk ecology. Overall, SNODAS predictions tended to overestimate field measurements, prediction uncertainty was high, and the difference between SNODAS predictions and field measurements was greater in snow shadows for both snow variables compared to non-snow shadow areas. We used a simple simulation of snow effects on the probability of an elk being killed by a predator to show that, if SNODAS prediction uncertainty was ignored, we might have mistakenly concluded that SWE was not an important factor in where elk were killed in predatory attacks during the winter. In this simulation, we were interested in the effects of snow at finer scales (< 1 km2) than the resolution of SNODAS. If bias were to decrease when SNODAS predictions are averaged over coarser scales, SNODAS would be applicable to population-level ecology studies. In our study, however, averaging predictions over moderate to broad spatial scales (9-2200 km2) did not reduce the differences between SNODAS predictions and field measurements. This study highlights the need to carefully evaluate two issues when using model output as an explanatory variable in subsequent analysis: (1) the model's resolution relative to the scale of the ecological question of interest and (2) the implications of prediction uncertainty on inferences when using model predictions as explanatory or predictor variables.

Advances in wildlife abundance estimation using pedigree reconstruction.

The conservation and management of wildlife populations, particularly for threatened and endangered species are greatly aided with abundance, growth rate, and density measures. Traditional methods of estimating abundance and related metrics represent trade-offs in effort and precision of estimates. Pedigree reconstruction is an emerging, attractive alternate approach because its use of one-time, noninvasive sampling of individuals to infer the existence of unsampled individuals. However, advances in pedigree reconstruction could improve its utility, including forming a measure of precision for the method, establishing required spatial sampling effort for accurate estimates, ascertaining the spatial extent of abundance estimates derived from pedigree reconstruction, and assessing how population density affects the estimator's performance. Using established relationships for a stochastic, spatially explicit simulated moose (Alces americanus) population, pedigree reconstruction provided accurate estimates of the adult moose population size and trend. Novel bootstrapped confidence intervals performed as expected with intensive sampling but underperformed with moderate sampling efforts that could produce abundance estimates with low bias. Adult population estimates more closely reflected the total number of adults in the extant population, rather than number of adults inhabiting the area where sampling occurred. Increasing sampling effort, measured as the proportion of individuals sampled and as the proportion of a hypothetical study area, yielded similar asymptotic patterns over time. Simulations indicated a positive relationship between animal density and sampling effort required for unbiased estimates. These results indicate that pedigree reconstruction can produce accurate abundance estimates and may be particularly valuable for surveying smaller areas and low-density populations.

Spatial and temporal patterns of neutral and adaptive genetic variation in the endangered African wild dog (Lycaon pictus).

Deciphering patterns of genetic variation within a species is essential for understanding population structure, local adaptation and differences in diversity between populations. Whilst neutrally evolving genetic markers can be used to elucidate demographic processes and genetic structure, they are not subject to selection and therefore are not informative about patterns of adaptive variation. As such, assessments of pertinent adaptive loci, such as the immunity genes of the major histocompatibility complex (MHC), are increasingly being incorporated into genetic studies. In this study, we combined neutral (microsatellite, mtDNA) and adaptive (MHC class II DLA-DRB1 locus) markers to elucidate the factors influencing patterns of genetic variation in the African wild dog (Lycaon pictus); an endangered canid that has suffered extensive declines in distribution and abundance. Our genetic analyses found all extant wild dog populations to be relatively small (N(e) < 30). Furthermore, through coalescent modelling, we detected a genetic signature of a recent and substantial demographic decline, which correlates with human expansion, but contrasts with findings in some other African mammals. We found strong structuring of wild dog populations, indicating the negative influence of extensive habitat fragmentation and loss of gene flow between habitat patches. Across populations, we found that the spatial and temporal structure of microsatellite diversity and MHC diversity were correlated and strongly influenced by demographic stability and population size, indicating the effects of genetic drift in these small populations. Despite this correlation, we detected signatures of selection at the MHC, implying that selection has not been completely overwhelmed by genetic drift.

Glucocorticoid stress hormones and the effect of predation risk on elk reproduction.

Predators affect prey demography through direct predation and through the costs of antipredator behavioral responses, or risk effects. Experiments have shown that risk effects can comprise a substantial proportion of a predator's total effect on prey dynamics, but we know little about their strength in wild populations, or the physiological mechanisms that mediate them. When wolves are present, elk alter their grouping patterns, vigilance, foraging behavior, habitat selection, and diet. These responses are associated with decreased progesterone levels, decreased calf production, and reduced population size [Creel S, Christianson D, Liley S, Winnie JA (2007) Science 315:960]. Two general mechanisms for the effect of predation risk on reproduction have been proposed: the predation stress hypothesis and the predator-sensitive-food hypothesis. Here, we used enzyme immunoassay to measure fecal glucocorticoid metabolite concentrations for 1,205 samples collected from 4 elk populations over 4 winters to test the hypothesis that the effect of predation risk on elk reproduction is mediated by chronic stress. Across populations and years, fecal glucocorticoid concentrations were not related to predator-prey ratios, progesterone concentrations or calf-cow ratios. Overall, the effect of wolf presence on elk reproduction is better explained by changes in foraging patterns that carry nutritional costs than by changes in glucocorticoid concentrations.

African wild dog movements show contrasting responses to long and short term risk of encountering lions: analysis using dynamic Brownian bridge movement models.

BACKGROUND: Prey depletion is a threat to the world's large carnivores, and is likely to affect subordinate competitors within the large carnivore guild disproportionately. African lions limit African wild dog populations through interference competition and intraguild predation. When lion density is reduced as a result of prey depletion, wild dogs are not competitively released, and their population density remains low. Research examining distributions has demonstrated spatial avoidance of lions by wild dogs, but the effects of lions on patterns of movement have not been tested. Movement is one of the most energetically costly activities for many species and is particularly costly for cursorial hunters like wild dogs. Therefore, testing how top-down, bottom-up, and anthropogenic variables affect movement patterns can provide insight into mechanisms that limit wild dogs (and other subordinate competitors) in resource-depleted ecosystems. METHODS: We measured movement rates using the motion variance from dynamic Brownian Bridge Movement Models (dBBMMs) fit to data from GPS-collared wild dogs, then used a generalized linear model to test for effects on movement of predation risk from lions, predictors of prey density, and anthropogenic and seasonal variables. RESULTS: Wild dogs proactively reduced movement in areas with high lion density, but reactively increased movement when lions were immediately nearby. Predictors of prey density had consistently weaker effects on movement than lions did, but movements were reduced in the wet season and when dependent offspring were present. CONCLUSION: Wild dogs alter their patterns of movement in response to lions in ways that are likely to have important energetic consequences. Our results support the recent suggestion that competitive limitation of wild dogs by lions remains strong in ecosystems where lion and wild dog densities are both low as a result of anthropogenic prey depletion. Our results reinforce an emerging pattern that movements often show contrasting responses to long-term and short-term variation in predation risk.

Predation strongly limits demography of a keystone migratory herbivore in a recovering transfrontier ecosystem.

Large herbivore migrations are imperiled globally; however the factors limiting a population across its migratory range are typically poorly understood. Zambia's Greater Liuwa Ecosystem (GLE) contains one of the largest remaining blue wildebeest (Connochaetes taurinus taurinus) migrations, yet the population structure, vital rates, and limiting factors are virtually unknown. We conducted a long-term demographic study of GLE wildebeest from 2012 to 2019 of 107 collared adult females and their calves, 7352 herd observations, 12 aerial population surveys, and concurrent carnivore studies. We applied methods of vital rate estimation and survival analysis within a Bayesian estimation framework. From herd composition observations, we estimated rates of fecundity, first-year survival, and recruitment as 68%, 56%, and 38% respectively, with pronounced interannual variation. Similar rates were estimated from calf-detections with collared cows. Adult survival rates declined steadily from 91% at age 2 years to 61% at age 10 years thereafter dropping more sharply to 2% at age 16 years. Predation, particularly by spotted hyena, was the predominant cause of death for all wildebeest ages and focused on older animals. Starvation only accounted for 0.8% of all unbiased known natural causes of death. Mortality risk differed substantially between wet and dry season ranges, reflecting strong spatio-temporal differences in habitat and predator densities. There was substantial evidence that mortality risk to adults was 27% higher in the wet season, and strong evidence that it was 45% higher in the migratory range where predator density was highest. The estimated vital rates were internally consistent, predicting a stable population trajectory consistent with aerial estimates. From essentially zero knowledge of GLE wildebeest dynamics, this work provides vital rates, age structure, limiting factors, and a plausible mechanism for the migratory tendency, and a robust model-based foundation to evaluate the effects of potential restrictions in migratory range, climate change, predator-prey dynamics, and poaching.

Toward a predictive theory of risk effects: hypotheses for prey attributes and compensatory mortality.

Risk effects, or the costs of antipredator behavior, can comprise a large proportion of the total effect of predators on their prey. While empirical studies are accumulating to demonstrate the importance of risk effects, there is no general theory that predicts the relative importance of risk effects and direct predation. Working toward this general theory, it has been shown that functional traits of predators (e.g., hunting modes) help to predict the importance of risk effects for ecosystem function. Here, I note that attributes of the predator, the prey, and the environment are all important in determining the strength of antipredator responses, and I develop hypotheses for the ways that prey functional traits might influence the magnitude of risk effects. In particular, I consider the following attributes of prey: group size and dilution of direct predation risk, the degree of foraging specialization, body mass, and the degree to which direct predation is additive vs. compensatory. Strong tests of these hypotheses will require continued development of methods to identify and quantify the fitness costs of antipredator responses in wild populations.

Neonatal mortality of elk driven by climate, predator phenology and predator community composition.

1. Understanding the interaction among predators and between predation and climate is critical to understanding the mechanisms for compensatory mortality. We used data from 1999 radio-marked neonatal elk (Cervus elaphus) calves from 12 populations in the north-western United States to test for effects of predation on neonatal survival, and whether predation interacted with climate to render mortality compensatory. 2. Weibull survival models with a random effect for each population were fit as a function of the number of predator species in a community (3-5), seven indices of climatic variability, sex, birth date, birth weight, and all interactions between climate and predators. Cumulative incidence functions (CIF) were used to test whether the effects of individual species of predators were additive or compensatory. 3. Neonatal elk survival to 3 months declined following hotter previous summers and increased with higher May precipitation, especially in areas with wolves and/or grizzly bears. Mortality hazards were significantly lower in systems with only coyotes (Canis latrans), cougars (Puma concolor) and black bears (Ursus americanus) compared to higher mortality hazards experienced with gray wolves (Canis lupus) and grizzly bears (Ursus horribilis). 4. In systems with wolves and grizzly bears, mortality by cougars decreased, and predation by bears was the dominant cause of neonatal mortality. Only bear predation appeared additive and occurred earlier than other predators, which may render later mortality by other predators compensatory as calves age. Wolf predation was low and most likely a compensatory source of mortality for neonatal elk calves. 5. Functional redundancy and interspecific competition among predators may combine with the effects of climate on vulnerability to predation to drive compensatory mortality of neonatal elk calves. The exception was the evidence for additive bear predation. These results suggest that effects of predation by recovering wolves on neonatal elk survival, a contentious issue for management of elk populations, may be less important than the composition of the predator community. Future studies would benefit by synthesizing overwinter calf and adult-survival data sets, ideally from experimental studies, to test the roles of predation in annual compensatory and additive mortality of elk.