Search

The leopard Panthera pardus is in range-wide decline, and many populations are highly threatened. Prey depletion is a major cause of global carnivore declines, but the response of leopard survival and density to this threat is unclear: by reducing the density of a dominant competitor (the lion Panthera leo) prey depletion could create both costs and benefits for subordinate competitors. We used capture–recapture models fitted to data from a 7-year camera-trap study in Kafue National Park, Zambia, to obtain baseline estimates of leopard population density and sex-specific apparent survival rates. Kafue is affected by prey depletion, and densities of large herbivores preferred by lions have declined more than the densities of smaller herbivores preferred by leopards. Lion density is consequently low. Estimates of leopard density were comparable to ecosystems with more intensive protection and favourable prey densities. However, our study site is located in an area with good ecological conditions and high levels of protection relative to other portions of the ecosystem, so extrapolating our estimates across the Park or into adjacent Game Management Areas would not be valid. Our results show that leopard density and survival within north-central Kafue remain good despite prey depletion, perhaps because (1) prey depletion has had weaker effects on preferred leopard prey compared to larger prey preferred by lions, and (2) the density of dominant competitors is consequently low. Our results show that the effects of prey depletion can be more complex than uniform decline of all large carnivore species, and warrant further investigation.

Many studies have shown that behavioral responses to the risk posed by predators can carry costs for prey by reducing fecundity or survival, with consequent effects on population dynamics. Responses to risk include increased vigilance and reduced foraging, movement to safe habitats, increases or decreases in group size, and changes in patterns of movement. While we know that prey can detect and respond to both long term (LT) and short term (ST) variation in risk, field studies have only recently begun to consider how these responses might differ. Here, we hypothesize that prey movement patterns should respond differently to cues of LT and ST variation in risk. Specifically, cues of elevated LT risk might lead to decreased movement to improve the assessment of ST risk, while elevated ST risk might favor increased movement to reduce the proximity or duration of risks that are already assessed to be acute. We further hypothesize that decreases in movement are likely to be a general response to LT risk, while responses to ST risk are likely to vary in a manner that depends on the type of predator. In Liuwa Plain National Park, we found that wildebeest movements responded to the local intensity of predator use (LT risk), after controlling for other seasonal, diurnal and bottom-up effects. Speed decreased considerably and turning angles increased considerably, combining to markedly decrease linear movements. In contrast, immediate encounters with predators (ST risk) typically provoked fast, linear flight, and this effect was stronger for encounters with coursing predators. The effect of long term risk was to cause wildebeest to move more slowly and less linearly, i.e. to slow down and turn around, as part of a suite of behavioral responses, which also includes increased vigilance, that promote cautious assessment of ST risks when in locations with high levels of LT risk. This result has broad implications of understanding the influence of predation risk on foraging patterns of ungulates as this relationship is much more complex than simple avoidance of areas of ‘high risk’.

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.

Allocating resources to growth and reproduction requires grazers to invest time in foraging, but foraging promotes dental senescence and constrains expression of proactive antipredator behaviors such as vigilance. We explored the relationship between carnivore prey selection and prey foraging effort using incisors collected from the kills of coursing and stalking carnivores. We predicted that prey investing less effort in foraging would be killed more frequently by coursers, predators that often exploit physical deficiencies. However, such prey could expect delayed dental senescence. We predicted that individuals investing more effort in foraging would be killed more frequently by stalkers, predators that often exploit behavioral vulnerabilities. Further these prey could expect earlier dental senescence. We tested these predictions by comparing variation in age‐corrected tooth wear, a proxy of cumulative foraging effort, in adult (3.4–11.9 years) wildebeest killed by coursing and stalking carnivores. Predator type was a strong predictor of age‐corrected tooth wear within each gender. We found greater foraging effort and earlier expected dental senescence, equivalent to 2.6 additional years of foraging, in female wildebeest killed by stalkers than in females killed by coursers. However, male wildebeest showed the opposite pattern with the equivalent of 2.4 years of additional tooth wear in males killed by coursers as compared to those killed by stalkers. Sex‐specific variation in the effects of foraging effort on vulnerability was unexpected and suggests that behavioral and physical aspects of vulnerability may not be subject to the same selective pressures across genders in multipredator landscapes.

Conservation of competitively subordinate carnivores presents a difficult challenge because they are limited by dominant competitors. Prey depletion is one of the leading causes of large carnivore decline worldwide, but little is known about the net effect of prey depletion on subordinate carnivores when their dominant competitors are also reduced. African wild dogs are often limited by high densities of dominant competitors, particularly lions. We measured African wild dog density and survival, using mark-recapture models fit to 8 years of data from 425 known individuals in the Greater Kafue Ecosystem, Zambia. The GKE is affected by prey depletion, particularly of large herbivores, and thus the density of lions is significantly lower than ecologically comparable ecosystems. Counter to expectations from mesopredator release theory, wild dog density in GKE was far lower than comparable ecosystems with higher lion and prey density, though annual survival rates were comparable to large and stable populations. Average pack size was small and home range size was among the largest recorded. Our results show that low lion density did not competitively release the GKE wild dog population and we infer that the low density of wild dogs was a product of low prey density. Our results suggest that there is an optimal ratio of prey and competitors at which wild dogs achieve their highest densities. This finding has immediate implications for the conservation of the endangered African wild dog, and broad implications for the conservation of subordinate species affected by resource depletion and intraguild competition.

Despite some controversy, a wide range of research across multiple taxa have established that carnivores strongly influence prey population dynamics both through direct offtake and indirect risk effects. Because of these powerful top-down effects carnivores can influence ecosystems across multiple trophic levels. Here we discuss research addressing carnivore direct- and indirect effects on prey, and how these effects can influence overall ecosystem structure and function.

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.

Understanding the influence of environmental conditions and people on ungulate density and distribution is of key importance for conservation. We evaluated the effects of ecological and anthropogenic factors on the density of migratory wildebeest and zebra and resident oribi in Zambia's Liuwa Plain National Park where human settlements were present. We conducted transect surveys from 2010 to 2013 using distance sampling methods and then developed a set of 38 candidate models to describe results and predict density. Models included the effects of variables in three classes: environmental (year, season, vegetation, predominant grass height, burn, water presence), predation risk (hyaena density) and anthropogenic (distance to park boundary and settlements). Densities ranged from 6.2 to 60.8 individuals km−2 for wildebeest, 1.1 to 14.5 individuals km−2 for oribi and 1.8 to 8.1 individuals km−2 for zebra. The most complex models were strongly supported for all three species. The magnitude and sign of variable effects differed among species, indicating that local densities of wildebeest, oribi and zebra are affected by a complex set of anthropogenic and ecological factors. Results reveal resource partitioning among ungulate species and indicate that predation risk and proximity to humans affect ungulate distributions with implications for managing migrations in the Greater Liuwa Ecosystem.

Top predators can exert strong influences on community structure and function, both via direct, consumptive effects, as well as through non-consumptive, fear-based effects (i.e. predation risk). However, these effects are challenging to quantify, particularly for mobile predators in marine ecosystems. To advance this field of research, here we used baited remote underwater video stations (BRUVs) to assess how the behavior of mobile fish species off Cape Cod, Massachusetts, was affected by exposure to large sharks. We categorized sites into three levels of differential shark predation exposure (white sharks, Carcharodon carcharias) and quantified the relative abundance and arrival times (elapsed time before appearing on screen) for six mobile fish prey groups to the BRUV stations. Increased large shark exposure was associated with a decrease in overall prey abundance, but the overall response was prey group-specific. Foraging of smooth dogfish, a likely important prey item for large sharks in the system, was significantly reduced in areas frequented by white sharks. Specifically, the predicted probabilities of smooth dogfish bait contacts or bite attempts occurring were reduced by factors of 5.7 and 8.4, respectively, in areas of high exposure as compared to low exposure. These modifications were underscored by a decrease in smooth dogfish abundance in areas of high exposure as well. Our results suggest that populations of large, roving sharks may induce food-related costs in prey. We discuss the implications of this work within the context of the control of risk (COR) hypothesis, for the purposes of advancing our understanding of the ecological role and effects of large sharks on coastal marine ecosystems.

If access to food is affected by the risk of predation, then the number of individuals killed by predators is an incomplete measure of the limiting effect of predation. Nonetheless, it is often assumed that the costs of antipredator responses (risk effects) are either small enough to be ignored or positively correlated with direct predation, and thus unlikely to alter inferences based on predation rates. These assumptions are rarely tested. Here we studied five large carnivores and ten prey species in three Zambian ecosystems to test relationships between direct predation, antipredator vigilance and trade-offs with foraging. The presence of a predator caused vigilance to increase by a factor of 2.4, with substantial variation among prey species in the strength of this response. This was associated with a 28% decrease in the proportion of individuals foraging, a trade-off that was consistent across species. We detected no correlation between direct predation and the strength of antipredator responses, which undermines the gambit of ignoring risk effects. The strength of antipredator responses was uncorrelated with broad attributes of predators and environments, but was correlated with attributes of prey. Responses were stronger for small species and for browsers/mixed feeders relative to grazers. It has previously been noted that small ungulates face higher rates of direct predation. Building on this inference, our results suggest that carnivore loss/restoration will also have stronger behaviorally-mediated effects on small ungulates, particularly browsers and mixed feeders. If such species increase their representation where carnivores are depleted, then cascading effects on vegetation would be expected.

Within a large carnivore guild, subordinate competitors (African wild dog, Lycaon pictus, and cheetah, Acinonyx jubatus) might reduce the limiting effects of dominant competitors (lion, Panthera leo, and spotted hyena, Crocuta crocuta) by avoiding them in space, in time, or through patterns of prey selection. Understanding how these competitors cope with one other can inform strategies for their conservation. We tested how mechanisms of niche partitioning promote coexistence by quantifying patterns of prey selection and the use of space and time by all members of the large carnivore guild within Liuwa Plain National Park in western Zambia. Lions and hyenas specialized on wildebeest, whereas wild dogs and cheetahs selected broader diets including smaller and less abundant prey. Spatially, cheetahs showed no detectable avoidance of areas heavily used by dominant competitors, but wild dogs avoided areas heavily used by lions. Temporally, the proportion of kills by lions and hyenas did not detectably differ across four time periods (day, crepuscular, early night, and late night), but wild dogs and especially cheetahs concentrated on time windows that avoided nighttime hunting by lions and hyenas. Our results provide new insight into the conditions under which partitioning may not allow for coexistence for one subordinate species, the African wild dog, while it does for cheetah. Because of differences in responses to dominant competitors, African wild dogs may be more prone to competitive exclusion (local extirpation), particularly in open, uniform ecosystems with simple (often wildebeest dominated) prey communities, where spatial avoidance is difficult.

Interspecific competition between large carnivores is not well understood due to the complexity of carnivore guild dynamics and the array of ecological and human factors affecting them. The loss of a carnivore guild's dominant species can therefore provide considerable insights. We evaluated spotted hyaena density and demography from Zambia's Liuwa Plain National Park, a recovering ecosystem where the hyaena's main competitor, the African lion, was severely reduced to 3–5 animals. Using data from 233 individuals in five clans and mark-resight robust design models we estimated population size, density, age and sex-specific annual survival rates for the hyaena population from 2010 to 2013 and tested for the effect of human settlements, prey density, competition with lions and hyaena clan size on mean hyaena survival. We estimated a minimum density of 0.13 hyaenas/km2 and maximum density of 0.52 hyaenas/km2. Density fluctuated with seasonal migrations of the main prey species, the blue wildebeest. Mean annual survival across all age classes was 0.93 (95%CI: 0.79–0.99). With high survival rates across all sex and age classes, we found no detectable effects of ecological or anthropogenic factors on survival. These results suggest that the combination of abundant prey, weak human-carnivore conflict, and weak competition from lions provide very favourable conditions for hyaena survival. Additional investigations of hyaena demography concurrent with lion population recovery efforts can provide considerable insights into the dynamics of carnivore competition and predation in African savannahs and the human and ecological factors affecting them.

Summary African rangelands support diverse ungulate communities whose member species exhibit unique combinations of body morphology and behaviour that have evolved over millions of years to limit the effects of competition and predation on fitness, and more recently, to cope with people and livestock. The mechanisms by which native ungulates cope with the combined effects of competition, predation and human disturbance are poorly understood. Addressing this knowledge gap will help guide management and conservation plans for large mammal communities outside of strictly protected areas. We conducted animal counts on line transects and used a spatially explicit distance sampling model to test the influence of bottom–up effects (vegetation), top–down effects from African lion Panthera leo and spotted hyena Crocuta crocuta and human disturbance (livestock density, occupied human settlements) on native ungulate densities and distributions across a multiple-use rangeland in Kenya. We examined five species that varied in body morphology and foraging strategy, including two large-bodied grazers (zebra Equus quagga; wildebeest Connochaetes taurinus), two medium-bodied, mixed-feeders (Grant's gazelle Nanger granti; impala Aepyceros melampus) and one very large-bodied browser (giraffe Giraffa camelopardalis tippelskirchi). Densities and distributions of all species varied across available land-use types, with bottom–up effects (vegetation) and human disturbance having the strongest overall effects. Responses varied by ungulate foraging strategy. Distributions of grazers and the browser (giraffe) changed seasonally, while distributions of mixed-feeders and a grazer (zebra) changed diurnally. At broad spatial scales, ungulates did not respond to long-term variation in predation risk though they have been shown to respond behaviourally to short-term variation in risk. Synthesis and applications. Our results revealed that native ungulates in this multiple-use rangeland occur at densities comparable to many flagship protected area populations. We attribute observed densities to spatial heterogeneity across this landscape that included variation in human land use (community conservation area, seasonal settlements and livestock grazing, permanent settlements) and vegetation (grassland, bushland, woodland) that changed through time (seasonally, diurnally). This variation allowed ungulates to carve out ecological niches while reducing exposure to the potential limiting effects of competition and human disturbance. For some species, securing access to forage resources and avoiding human disturbance may be associated with costs through increased risk of predation. Our research reveals the trade-offs native ungulates make to cope with changes in forage availability, human disturbance and predation risk, providing important insights to help guide native ungulate conservation efforts in multiple-use rangelands.

Factors that limit African lion populations are manifold and well-recognized, but their relative demographic effects remain poorly understood, particularly trophy hunting near protected areas. We identified and monitored 386 individual lions within and around South Luangwa National Park, Zambia, for five years (2008-2012) with trophy hunting and for three additional years (2013-2015) during a hunting moratorium. We used these data with mark-resight models to estimate the effects of hunting on lion survival, recruitment, and abundance. The best survival models, accounting for imperfect detection, revealed strong positive effects of the moratorium, with survival increasing by 17.1 and 14.0 percentage points in subadult and adult males, respectively. Smaller effects on adult female survival and positive effects on cub survival were also detected. The sex-ratio of cubs shifted from unbiased during trophy-hunting to female-biased during the moratorium. Closed mark-recapture models revealed a large increase in lion abundance during the hunting moratorium, from 116 lions in 2012 immediately preceding the moratorium to 209 lions in the last year of the moratorium. More cubs were produced each year of the moratorium than in any year with trophy hunting. Lion demographics shifted from a male-depleted population consisting mostly of adult (≥4 years) females to a younger population with more (>29%) adult males. These data show that the three-year moratorium was effective at growing the Luangwa lion population and increasing the number of adult males. The results suggest that moratoria may be an effective tool for improving the sustainability of lion trophy hunting, particularly where systematic monitoring, conservative quotas, and age-based harvesting are difficult to enforce.

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.

Several elk herds in the Greater Yellowstone Ecosystem are fed during winter to alleviate interactions with livestock, reduce damage to stored crops, and to manage for high elk numbers. The effects of supplemental feeding on ungulate population dynamics has rarely been examined, despite the fact that supplemental feeding is partially justified as necessary for maintaining or enhancing population growth rates. We used linear regression to assess how the presence of feedgrounds, snowpack, summer rainfall, indices of grizzly bear density and wolves per elk, elk population trend counts, brucellosis seroprevalence, and survey date were correlated with midwinter calf:cow ratios, a metric correlated with population growth, from 1983–2010 from 12 ecologically similar elk herd units (7 fed and 5 unfed) in Wyoming, USA. Our statistical approach allowed for rigorous tests of the hypotheses that supplemental feeding had positive effects on calf:cow ratios and reduced sensitivity of calf:cow ratios to bottom-up limitation relative to top-down limitation from native predators. Calf:cow ratios generally declined across all herd units over the study period and varied widely among units with feedgrounds. We found no evidence that the presence of feedgrounds had positive effects on midwinter calf:cow ratios in Wyoming. Further, fed elk showed stronger correlations with environmental factors, whereas calf:cow ratios for unfed elk showed stronger correlations with predator indices. Although we found no consistent association between winter feeding and higher calf:cow ratios, we did not assess late winter mortality and differences in human offtake between fed and unfed regions, which remain a priority for future research. Published 2015. This article is a U.S. Government work and is in the public domain in the USA.

Animals that are exposed to environmental stressors may experience chronically elevated glucocorticoid (GC) levels, which can lead to deleterious effects such as immune and reproductive system suppression. Such effects are of special concern in rare species. We measured fecal GC concentrations in endangered San Joaquin kit foxes (Vulpes mocrotis mutica) to assess stress responses in natural and urban habitats. Basal GC levels were significantly higher (P < 0.01) among urban foxes (5.3 ± 2.2 ng cortisol/g dry feces; n = 32) than among foxes in natural habitats (4.7 ± 0.5 ng cortisol/g dry feces; n = 179). In the natural habitat, potential stressors included predation risk, particularly in areas with shrubs where coyote (Canis latrans) abundance was higher. In the urban habitat, the primary stressors were anthropogenic disturbances (e.g., human activity, vehicles). Foxes in natural habitats exhibited acute stress responses when trapped, but exhibited similar mean GC levels between areas of high an...

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.

Both short-term and long-term variation in predation risk can affect the behaviour of prey, thus affecting growth, reproduction, survival and population dynamics. Inferences about the strength of such ‘risk effects’ in the wild have been limited by a lack of studies that relate antipredator responses to the magnitude of direct predation, measure responses of prey to risk from complete predator guilds, and quantify risk in more than one way. Here, we quantify behavioural responses of a complete ungulate prey guild to long-term and short-term variation in risks from all of the large predators in Liuwa Plain National Park, with known patterns of direct predation. Our analysis allows the first direct test for interaction between responses to long-term and short-term risk in the wild, and reveals that prey vigilance responds strongly to locations with high long-term risk when short-term risk is high, but not when short-term risk is low. This result has broad ramifications for the design and interpretation of field studies of antipredator behaviour, its costs and its consequences for population dynamics. Predator–prey interactions in a Zambian national park reveal the interaction between types of risk: strong prey vigilance responses to locations with high long-term risk when short-term risk was high, but not when short-term risk was low.

Predators alter prey dynamics by direct killing and through the costs of antipredator responses or risk effects. Antipredator behavior includes proactive responses to long-term variation in risk (e.g., grouping patterns) and reactive responses to short-term variation in risk (e.g., intense vigilance). In a 3-year field study, we measured variation in antipredator responses and the foraging costs of these responses for 5 ungulates (zebra, wildebeest, Grant’s gazelle, impala, and giraffe) that comprised more than 90% of the prey community available to the 2 locally dominant predators, lions and spotted hyenas. Using a model-selection approach, we examined how vigilance and group size responded to attributes of the predator, prey, and environment. We found that 1) the strength of antipredator responses was affected by attributes of the predator, prey, and environment in which they met; 2) grouping and vigilance were complementary responses; 3) grouping was a proactive response to the use of dangerous habitats, whereas vigilance was a reactive response to finer cues about predation risk; 4) increased vigilance caused a large reduction in foraging for some species (but not all); and 5) there was no clear relationship between direct predation rates and the foraging costs of antipredator responses. Broadly, our results show that antipredator responses and their costs vary in a complex manner among prey species, the predators they face, and the environment in which they meet.

Terrestrial large carnivores have great ecological, economic and cultural importance, but are in global decline due to habitat loss, prey depletion, poaching, retributive killing and regulated hunting. While regulated carnivore hunting potentially reduces conflict with humans and livestock, increases social tolerance and provides revenue for conservation, it can also drive population declines. Some policies regulating carnivore hunting acknowledge and address negative effects on demography and population dynamics, but others do not. Using wolves as an example, we identify four aspects of hunting policy that do not align well with ecological theory and data, and suggest resolutions for these problems that have broad relevance to exploited carnivore populations.

Field studies that rely on fixes from GPS-collared predators to identify encounters with prey will often underestimate the frequency and strength of antipredator responses. These underestimation biases have several mechanistic causes. (1) Step bias: The distance between successive GPS fixes can be large, and encounters that occur during these intervals go undetected. This bias will generally be strongest for cursorial hunters that can rapidly cover large distances (e.g., wolves and African wild dogs) and when the interval between GPS fixes is long relative to the duration of a hunt. Step bias is amplified as the path travelled between successive GPS fixes deviates from a straight line. (2) Scatter bias: Only a small fraction of the predators in a population typically carry GPS collars, and prey encounters with uncollared predators go undetected unless a collared group-mate is present. This bias will generally be stronger for fission-fusion hunters (e.g., spotted hyenas, wolves, and lions) than for highly cohesive hunters (e.g., African wild dogs), particularly when their group sizes are large. Step bias and scatter bias both cause underestimation of the frequency of antipredator responses. (3) Strength bias: Observations of prey in the absence of GPS fix from a collared predator will generally include a mixture of cases in which predators were truly absent and cases in which predators were present but not detected, which causes underestimation of the strength of antipredator responses. We quantified these biases with data from wolves and African wild dogs and found that fixes from GPS collars at 3-h intervals underestimated the frequency and strength of antipredator responses by a factor >10. We reexamined the results of a recent study of the nonconsumptive effects of wolves on elk in light of these results and confirmed that predation risk has strong effects on elk dynamics by reducing the pregnancy rate.

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.

We have previously found that the behavioral responses of elk to the risk of predation by wolves are correlated with changes in habitat selection, altered diets, reduced food intake, decreased fecal progesterone concentrations, and decreased calf recruitment. P. J. White et al. recently questioned these results, concluding that “multiple lines of evidence and/or strong validation should be brought to bear before relying on indirect measures of how predators affect pregnancy rates.” Here, we systematically surveyed available data and found that five studies (with data from 10 widely distributed populations) have directly detected decreases of 24–43% in elk pregnancy rates in response to increased predation risk. This survey includes data not discussed by White et al. from their own research, which reveal a 32% decrease in pregnancy rates following wolf recolonization in central Yellowstone. Following the survey of available data, we discuss several methodological and statistical problems in White et al.'s study that would be expected to mask the effects of predation risk. While other factors also clearly affect elk recruitment, multiple lines of evidence using a broad array of methods have detected an association between predation risk and reduced reproduction in elk.

Summary 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 )a nd 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.

Recent increases in the height and growth ring width of willow (Salix spp.) and other woody plants in the Greater Yellowstone Ecosystem (GYE) have been attributed to a behaviorally mediated trophic cascade from wolves (Canis lupus) to elk (Cervus elaphus) to willows. This hypothesis predicts that individual elk consume less willow in response to the presence of wolves, but this prediction has not been directly tested with data from elk. We collected 727 fecal samples from elk in the Gallatin Canyon portion of the GYE over three winters and used microhistological methods to quantify the proportion of willow in each sample. We then tested the effect of wolf presence on willow consumption by elk, controlling for the effects of snow conditions, sex, and habitat type. During the period of study, 8-17 wolves occupied the study area, and wolves were locally present on 49% of 260 sampling days, stratified at two-week intervals across three drainages. Over the three years combined, willow consumption was related to snow conditions, wolf presence, and a wolf X sex interaction. As expected, willow consumption increased with deeper and less penetrable snow, and this effect was strong. Contrary to expectation, willow consumption increased in the presence of wolves. As with other aspects of antipredator behavior, wolves had different effects on willow consumption by males and females. Finally, we aggregated the data to estimate winter-long mean willow consumption within each drainage; at this broader scale, willow consumption again increased as predation risk increased. In summary, willow consumption was more strongly affected by snow conditions than by the presence of wolves. Interactions between elk and willow were affected by wolves, but not as predicted by the hypothesis that wolf presence favors willow release through a reduction in the selection of willow by individual elk. If a trophic cascade is operating, our results suggest that a decline in the size of the elk population (to roughly one-half its size immediately prior to wolf recovery) may be more important than changes in the willow consumption of individual elk. Finally, reduced grazing of herbaceous vegetation may be equally important for vegetation dynamics.

Considerable research has tested two axioms about antipredator behaviour: prey may respond to temporal variation in risk (the risky times hypothesis) or to spatial variation in risk (the risky places hypothesis). The risk allocation hypothesis extends the risky times hypothesis, noting that the context into which a period of risk or safety is imbedded should affect the level of response, and predicting that vigilance should be greatest during a brief pulse of risk and lowest during a brief pulse of safety. We tested these models using natural variation in true predation risk, by observing elk, Cervus elaphus, in the presence or absence of wolves, Canis lupus, on three sites in the Greater Yellowstone Ecosystem, over 4 winters. Elk were exposed to wolves frequently in the Northern Range, intermittently in the Gallatin Canyon, and never in the Elkhorn Mountains. We used GLM to control the effects of herd size and composition while testing for responses of vigilance to daily wolf presence (temporal variation) and to site-specific, background levels of risk (spatial variation). The data gave strong support to the risk allocation hypothesis and equivocal support to the risky times hypothesis, and did not support the risky places hypothesis. Contrary to the risky places hypothesis but as predicted by risk allocation, Gallatin elk were more vigilant than Northern Range elk despite lower background risk in the Gallatin Canyon. The contradictory predictions of the risk allocation and risky places hypotheses have important ramifications for research on the ecological consequences of antipredator behaviour.

Risk effects arise when prey alter their behavior in response to predators, and these responses carry costs. Empirical studies have found that risk effects can be large. Nonetheless, studies of predation in vertebrate conservation and management usually consider only direct predation. Given the ubiquity and strength of behavioral responses to predators by vertebrate prey, it is not safe to assume that risk effects on dynamics can be ignored. Risk effects can be larger than direct effects. Risk effects can exist even when the direct rate of predation is zero. Risk effects and direct effects do not necessarily change in parallel. When risk effects reduce reproduction rather than survival, they are easily mistaken for limitation by food supply.

Risk effects in ungulates are poorly understood but have recently been implicated as an important driver of elk (Cervus elaphus) population dynamics since wolves were reintroduced into the Greater Yellowstone Ecosystem (GYE) of Montana and Wyoming, USA. From December to May in 2004, 2005, and 2006, we recorded the daily presence of wolves on 3 sites occupied by the Upper Gallatin elk population in the northwest corner of the GYE. We estimated the proportion of grasses, conifers, evergreen shrubs, and woody stems in 980 elk fecal samples collected from those 3 sites and tested whether wolf presence affected elk diets. The winter of 2005 was extremely mild allowing us the opportunity to investigate how elk‐wolf interactions might change if winter snowpack continues to decline in western North America due to global warming. Snow accumulation consistently favored browsing, and diets during the mild winter were dominated by grass, very similar to the spring diet. In normal winters, adult males grazed less than adult females except when wolves were near because females decreased grazing in response to wolves. Adult males decreased browsing on conifers by half whereas adult females doubled conifer browsing on days when wolves were near. Overall, the sexes had different diets when wolves were absent but showed strong overlap when wolves were present. Diet shifts due to wolves may be causing trophic cascades that have gone unrecognized and probably carry nutritional consequences for wintering elk. Key words: behaviorally mediate trophic cascade, browsing, diet, elk, grazing, predation, risk effects, sexual segregation, wolves. [Behav Ecol]

We quantified the vigilance levels of elk (Cervus elaphus) preyed on by wolves (Canis lupus) in Yellowstone National Park, between January and May in 2005 and 2006, and used Akaike’s information criterion to compare a set of 38 regression models for vigilance levels. These models combined up to 9 predictor variables of 3 types: characteristics of the prey group (herd size and composition), characteristics of the predator (wolf pack size, distance away, and the presence/absence of a kill), and characteristics of the local environment (distance to woodland edges, snow depth, and snow cover). The set of models spanned a range of complexity from simple univariate models to complex combinations with up to 3 variables of each type. Complex models incorporating the characteristics of the wolf pack, the structure of the elk herd, and the environmental conditions had higher information content than simple models. Although univariate models of vigilance detect significant relationships, they have low information content relative to multivariate models. These results show that elk assesses factors of several types when assessing risk and deciding how much time to allocate to vigilance. In particular, we found that all well-supported models of vigilance included several ‘‘prey’’ variables and several ‘‘predator’’ variables. This result highlights the need to consider information about predators when trying to explain variation in vigilance levels in prey. Key words: antipredator behavior, Canis lupus, Cervus elaphus, elk, predation risk, vigilance, wolf. [Behav Ecol]

San Joaquin kit foxes (Vulpes macrotis mutica) are an endangered species with a narrow geographic range whose natural populations are limited by predation by coyotes (Canis latrans). In the warm, arid grassland and shrubland habitats where kit foxes occur, coyotes are more cover dependent than kit foxes, creating the possibility of habitat segregation. Effects of habitat variation on coyote and kit fox competition are unknown. We assessed exploitation and interference competition between coyotes and kit foxes in grassland and shrubland habitats to determine if such competition varies among habitats. With respect to exploitation competition, we evaluated habitat and spatial partitioning, diet, prey abundance, and survival for kit foxes and coyotes at the Lokern Natural Area in central California, USA, from January 2003 through June 2004. Kit foxes partitioned habitat, space, and diet with coyotes. Coyotes primarily used shrubland habitats whereas kit foxes selectively used burned grasslands. Kit foxes and coyotes had high dietary overlap with regards to items used, but proportional use of items differed between the 2 species. Kit foxes selected for Heermann's kangaroo rats (Dipodomys heermanni), which were closely tied to shrub habitats. With respect to interference competition, predation was the primary source of mortality for kit foxes, and survival of individual kit foxes was inversely related to proportion of shrub habitat within their home ranges. Our results suggest that a heterogeneous landscape may benefit kit foxes by providing habitat patches where predation risk may be lower.

We studied individual and herd level behavioural responses of elk to spatial and temporal variation in the risk of predation by wolves over three winters in the Upper Gallatin drainage, Montana, U.S.A. Within a given drainage, elk of both sexes moved into or closer to protective cover (timber) in response to wolf presence. Cow elk responded to elevated risk by increasing vigilance in exchange for foraging, and large mixed (cow, calf, spike) herds substantially decreased in size. In contrast, when wolves were present, bulls did not increase vigilance levels, or decrease feeding, and small bull-only groups slightly increased in size. As a consequence, small bull-only herds and large mixed-sex herds converged on a similar size when wolves were present. We think this response is a balancing of the benefits of risk dilution with increased detectability or attractiveness of larger herds to wolves. Based on proportions in the population, wolves overselected bulls and underselected cows as prey. Thus, bulls showed weaker antipredator responses than cows, despite facing a greater risk of predation. Using marrow fat content from elk killed by wolves as an indicator of body condition, bulls were in significantly worse body condition than cows throughout the winter, and condition deteriorated for both sexes as winter progressed. Overall, we conclude that antipredator behaviours carry substantial foraging costs, that bulls, because of their poorer body condition, are less able to pay these costs than cows, and that differences in ability to pay foraging costs probably explain sex-specific differences in antipredator behaviours.

Summary Animal group size distributions are often right‐skewed, whereby most groups are small, but most individuals occur in larger groups that may also disproportionately affect ecology and policy. In this case, examining covariates associated with upper quantiles of the group size distribution could facilitate better understanding and management of large animal groups.We studied wintering elk groups in Wyoming, where group sizes span several orders of magnitude, and issues of disease, predation and property damage are affected by larger group sizes. We used quantile regression to evaluate relationships between the group size distribution and variables of land use, habitat, elk density and wolf abundance to identify conditions important to larger elk groups.We recorded 1263 groups ranging from 1 to 1952 elk and found that across all quantiles of group size, group sizes were larger in open habitat and on private land, but the largest effect occurred between irrigated and non‐irrigated land [e.g. the 90th quantile group size increased by 135 elk (95% CI = 42, 227) on irrigation].Only upper quantile group sizes were positively related to broad‐scale measures of elk density and wolf abundance. For wolf abundance, this effect was greater on elk groups found in open habitats and private land than those in closed habitats or public land. If we had limited our analysis to mean or median group sizes, we would not have detected these effects. Synthesis and applications. Our analysis of elk group size distributions using quantile regression suggests that private land, irrigation, open habitat, elk density and wolf abundance can affect large elk group sizes. Thus, to manage larger groups by removal or dispersal of individuals, we recommend incentivizing hunting on private land (particularly if irrigated) during the regular and late hunting seasons, promoting tolerance of wolves on private land (if elk aggregate in these areas to avoid wolves) and creating more winter range and varied habitats. Relationships to the variables of interest also differed by quantile, highlighting the importance of using quantile regression to examine response variables more completely to uncover relationships important to conservation and management., Our analysis of elk group size distributions using quantile regression suggests that private land, irrigation, open habitat, elk density and wolf abundance can affect large elk group sizes. Thus, to manage larger groups by removal or dispersal of individuals, we recommend incentivizing hunting on private land (particularly if irrigated) during the regular and late hunting seasons, promoting tolerance of wolves on private land (if elk aggregate in these areas to avoid wolves) and creating more winter range and varied habitats. Relationships to the variables of interest also differed by quantile, highlighting the importance of using quantile regression to examine response variables more completely to uncover relationships important to conservation and management.

For elk (Cervus elaphus) in the Gallatin drainage of the Greater Yellowstone Ecosystem, Montana, USA, wolf movements caused local predation risk to vary substan- tially on a time scale of days. Spatially and temporally fine-scaled data from GPS radio collars show that elk moved into the protective cover of wooded areas when wolves were present, reducing their use of preferred grassland foraging habitats that had high predation risk. By constraining habitat selection, wolves may have greater effects on elk dynamics than would be predicted on the basis of direct predation alone. Based on changes in the woody vegetation following the reintroduction of wolves, it has been suggested that an- tipredator responses by elk may be driving a trophic cascade in the Yellowstone ecosystem. However, studies to date have been hampered by a lack of direct data on spatial variation in predation risk, and the ways in which elk respond to variation in risk. Our data support a central portion of the hypothesis that elk antipredator behavior could drive a trophic cascade, but changes in elk numbers are also likely to have affected elk-plant interactions.

Many studies have examined grouping as a form of antipredator behaviour, but relatively few studies have examined how group size responds to natural variation in predation risk across space and through time. We studied the responses of elk, Cervus elaphus, herd size and composition to natural variation in the risk of predation by wolves, Canis lupus, in the Gallatin Canyon of Montana. We found that elk herd size increased as distance to protective cover increased. A positive association between group size and distance to cover is often interpreted as evidence that grouping is an antipredator response. However, we found that herd size increased only on days that wolves were absent. When wolves were present, herd sizes remained small at all distances from cover. This suggests that aggregation far from cover on days that wolves were absent was a foraging response, rather than an antipredator response. These data highlight interaction between temporal and spatial variation in predation risk, and suggest caution in conclusions about the antipredator benefits of grouping in the absence of direct data on risk or predator presence.

The reintroduction of wolves (Canis lupus) to Yellowstone provided the unusual opportunity for a quasi-experimental test of the effects of wolf predation on their primary prey (elk – Cervus elaphus) in a system where top-down, bottom-up, and abiotic forces on prey population dynamics were closely and consistently monitored before and after reintroduction. Here, we examined data from 33 years for 12 elk population segments spread across southwestern Montana and northwestern Wyoming in a large scale before-after-control-impact analysis of the effects of wolves on elk recruitment and population dynamics. Recruitment, as measured by the midwinter juvenile∶female ratio, was a strong determinant of elk dynamics, and declined by 35% in elk herds colonized by wolves as annual population growth shifted from increasing to decreasing. Negative effects of population density and winter severity on recruitment, long recognized as important for elk dynamics, were detected in uncolonized elk herds and in wolf-colonized elk herds prior to wolf colonization, but not after wolf colonization. Growing season precipitation and harvest had no detectable effect on recruitment in either wolf treatment or colonization period, although harvest rates of juveniles∶females declined by 37% in wolf-colonized herds. Even if it is assumed that mortality due to predation is completely additive, liberal estimates of wolf predation rates on juvenile elk could explain no more than 52% of the total decline in juvenile∶female ratios in wolf-colonized herds, after accounting for the effects of other limiting factors. Collectively, these long-term, large-scale patterns align well with prior studies that have reported substantial decrease in elk numbers immediately after wolf recolonization, relatively weak additive effects of direct wolf predation on elk survival, and decreased reproduction and recruitment with exposure to predation risk from wolves.

Limitation of predator populations by prey availability and the effects of predators on prey populations are widely recognized as important ecological processes that affect carnivore conservation. Interspecific competition can also be a strong limiting factor for carnivore populations, and the effects of competition help explain why some carnivore species are prone to extinction. Competition among carnivores is unusual in some ways, so some predictions from traditional models of competition do not hold. For example, an increase in the density of prey can increase the effect of competition among carnivores, rather than weakening it. I used published data from African wild dogs ( Lycaon pictus) to highlight four complexities that can modify the effects of competition on the population dynamics of carnivores: habitat fragmentation, counterintuitive effects of prey density, predator-prey size ratios, and habitat type. Resumen: La restriccion sobre poblaciones de depredadores ocasionada por la disponibilidad de presas y los efectos de depredadores en poblaciones de presas son ampliamente reconocidos como procesos ecologicos importantes que afectan la conservacion de carnivoros. A pesar de ello, la competencia interespecifica tambien puede ser un fuerte factor limitante para las poblaciones de carnivoros y los efectos de la competencia ayudan a explicar el porque algunas especies carnivoras son propensas a extinguirse. La competencia entre carnivoros es poco comun de cierta forma, asi que algunas predicciones de modelos tradicionales de competencia no se cumplen. Por ejemplo, un incremento en la densidad de presas puede incrementar el efecto de la competencia entre carnivoros, en lugar de disminuirlo. Utilice datos de trabajos publicados de perros silvestres africanos ( Lycaon pictus) para resaltar cuatro complicaciones que pueden modificar los efectos de la competencia en la dinamica poblacional de carnivoros: fragmentacion del habitat, efectos contraintuitivos de la densidad de presas, tamano de las proporciones depredador/presa y tipo de habitat.

African wild dogs (Lycaon pictus) always live at low population densities relative to sympatric large carnivores. This suggests that there are basic ecological reasons for the wild dog's endangered status. We examined the effects of intraspecific and interspecific competition, infectious diseases, foraging success, genetics and human activities on wild dogs. We present data from wild dogs in the Selous Game Reserve, and make comparisons with other populations to identify the limiting factors that are broadly important. The high density of wild dogs in Selous (40 adults/1000 km2) is associated with weak competition from lions and spotted hyenas. Across ecosystems, population density is negatively related to the intensity of interference competition with larger carnivores. Predation by lions and hyenas accounted for 13% of known-cause deaths in Selous and 33–50% in other populations. Intraspecific competition caused 69% of known-cause deaths in Selous, through infanticide and fights between packs, although most of the victims were juveniles with low reproductive value. Infectious diseases had little apparent impact in Selous (4% of deaths) or Kruger National Park (5%), but did play a role in the extinction of a small population in Serengeti. Infectious diseases and competition will generally interact because competitors harbor and transmit the diseases that affect wild dogs. Human activities caused 12% of deaths in Selous, even though it is large (43 000 km2) and does not border large human or livestock populations. Humans were the major agent of mortality in some populations. Foraging success varied little across ecosystems and was not apparently limiting. Mitochondrial DNA (mtDNA) genotypes revealed clinal variation between Selous and distant populations (rather than geographically isolated subspecies, as previously suggested). All wild dog populations have a genetically effective size (Ne) less than 500, so gene flow is necessary to maintain genetic diversity within populations.

African wild dogs (Lycaon pictus) are endangered largely because their population-density is low under all conditions. Interspecific competition with larger carnivores may be a factor limiting wild dog density. The density of wild dogs on a 2600-km2 area of the Selous Game Reserve (Tanzania) was 0.04 adults/km2. Spotted hyaena (Crocuta crocuta) density for the same area was estimated by audio playbacks as 0.32 hyaenas/km2. Lion (Panthera leo) density, determined from the ratio of hyaenas to lions, was 0.11 lions/km2. Across six ecosystems including Selous, there were strong negative correlations between wild dog and hyaena densities (r = −0.92; p = 0.01) and between wild dog and lion densities (r = −0.91; p = 0.03). Hyaenas out-numbered wild dogs by ratios ranging from 8:1 to 122:1. Ratios of lions to wild dogs ranged from 3:1 to 21:1. The diets of hyaenas and wild dogs overlap extensively; those of wild dogs and lions show less overlap. Where hyaenas are common and visibility is good, interference competition from hyaenas at wild dog kills is common and reduces wild dogs’ feeding time. Where hyaena density is lower and visibility is poor, interference competition at wild dog kills is rare. Wild dogs are commonly killed by lions and occasionally by hyaenas. These data suggest that competition with spotted hyaenas may limit or exclude wild dogs when hyaena density is high. Competition with lions appears less intense, but direct predation by lions on wild dogs is important. Competition and predation by larger carnivores may be of broad importance to the conservation of wild dogs and other medium-sized carnivores. Los perros salvajes Africanos (Lycaon pictus), se encuentran en peligro debido principalmente a que su densidad poblacional es baja. La competencia interespecifica con carnivoros mayores, podria ser un factor que limita la densidad de los perros salvajes. La densidad de los perros salvajes en un area de 2600 km2 de la Reserva de Caza de Selous (Tanzania) fue de 0.04 adultos/km2. La densidad de la hiena moteada (Crocuta crocuta) en la misma zona, fue estimada en 0.32 hienas/km2. La densidad de los leones (Panthera leo) fue estimada a partir de la proporcion entre hienas y leones y fue de 0.11 leones/km2. Existieron fuertes correlaciones negativas entre la densidad de perros salvajes y la de las hienas (r = −0.92, P = 0.01) y entre aquella de los perros salvajes y la de los leones (r = −0.91, P = 0.03) a traves de seis ecosistemas, incluyendo al de Selous. Las hienas fueron mas numerosas que los perros salvajes con proporciones que variaron entre 8: 1 y 122:1. La proporcion entre leones y perros salvajes vario entre 3:1 y 21:1. Las dietas de las hienas y de los perros salvajes se superponen en forma extenso, mientras que las dietas de los perros salvajes y de los leones muestran una menor superposicion. En las areas donde las hienas son comunes y la visibilidad es buena, la competencia de interferencia de las hienas con las presas cazadas por los perros salvajes, es comun y reduce la longitud del periodo de alimentacion de los perros salvajes. En las areas donde la densidad de las hienas es baja y la visibilidad es pobre, la competencia por interferencia con las presas cazadas por los perros salvajes es baja. Los perros salvajes son comunmente cazados por los leones y ocasionalmente por las hienas. Estos datos sugieren que la competencia con las hienas moteadas podria limitar o excluir a los perros salvajes cuando la densidad de las hienas es alta. La competencia con los leones parece ser menos intensa, pero la de predacion directa de los leones sobre los perros salvajes es importante. La competencia y la de predacion por parte de carnivoros mayores podria ser de gran importancia para la conservacion de los perros salvajes y otros carnivoros de tamano intermedio.

African wild dogs are 20–25 kg social carnivores whose major prey are ungulates ranging from 15 to 200 kg. In the Selous Game Reserve, Tanzania, wild dog pack size ranged from three to 20 adults (3–44 including yearlings and pups). Data from 905 hunts and 404 kills showed that hunting success, prey mass and the probability of multiple kills increased with number of adults. Chase distance decreased with number of adults. None the less, the distribution of per capita food intake across adult pack size was U-shaped, with a minimum close to the modal pack size. A similar result has been used to conclude that cooperative hunting does not favour sociality in lions (Packer et al. 1990, Am. Nat., 136, 1–19), and to argue that cooperative hunting is not responsible for group living in any carnivore (Caro 1994, Cheetahs of the Serengeti Plains: Group Living in an Asocial Species). Daily per capita food intake only accounts for variation in the benefits to cooperative hunting, ignoring variation in costs. For Selous wild dogs, per capita food intake per km chased peaked close to the modal adult pack size (where per capita food intake per day was near its minimum). Thus, the energetics of cooperative hunting favour sociality in Selous wild dogs. Analyses that incorporate variation in both costs and benefits of hunting may show that cooperative hunting favours sociality in other species where its influence has previously been rejected.

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.

Following the growth and geographic expansion of wolf (Canis lupus) populations reintroduced to Yellowstone National Park and central Idaho in 1995–1996, Rocky Mountain wolves were removed from the endangered species list in May 2009. Idaho and Montana immediately established hunting seasons with quotas equaling 20% of the regional wolf population. Combining hunting with predator control, 37.1% of Montana and Idaho wolves were killed in the year of delisting. Hunting and predator control are well-established methods to broaden societal acceptance of large carnivores, but it is unprecedented for a species to move so rapidly from protection under the Endangered Species Act to heavy direct harvest, and it is important to use all available data to assess the likely consequences of these changes in policy. For wolves, it is widely argued that human offtake has little effect on total mortality rates, so that a harvest of 28–50% per year can be sustained. Using previously published data from 21 North American wolf populations, we related total annual mortality and population growth to annual human offtake. Contrary to current conventional wisdom, there was a strong association between human offtake and total mortality rates across North American wolf populations. Human offtake was associated with a strongly additive or super-additive increase in total mortality. Population growth declined as human offtake increased, even at low rates of offtake. Finally, wolf populations declined with harvests substantially lower than the thresholds identified in current state and federal policies. These results should help to inform management of Rocky Mountain wolves.

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.

Elk (Cervus elaphus) in the Greater Yellowstone Ecosystem alter patterns of aggregation, habitat selection, vigilance, and foraging in the presence of wolves (Canis lupus). Antipredator behaviors like these can reduce predation risk but are also likely to carry costs. Data from five elk populations studied for 16 site years showed that progesterone concentrations (from 1489 fecal samples) declined with the ratio of elk to wolves. In turn, progesterone concentrations were a good predictor of calf recruitment in the subsequent year. Together, these data suggest that wolves indirectly affect the reproductive physiology and the demography of elk through the costs of antipredator behavior.

The risk of predation drives many behavioral responses in prey. However, few studies have directly tested whether predation risk alters the way other variables influence prey behavior. Here we use information theory (Akaike’s Information Criterion, AICc) in a novel way to test the hypothesis that the decision-making rules governing elk behavior are simplified by the presence of wolves. With elk habitat use as the dependent variable, we test whether the number of independent variables (i.e., the size of the models) that best predict this behavior differ when wolves are present vs absent. Thus, we use AICc scores simply to determine the number of variables to which elk respond when making decisions. We measured habitat use using 2,288 locations from GPS collars on 14 elk, over two winters (14 elk winters), in the Gallatin Canyon portion of the Greater Yellowstone Ecosystem. We found that the use of three major habitat components (grass, conifer, sage) was sensitive to many variables on days that wolves were locally absent, with the best models (ΔAICc≤2) averaging 7.4 parameters. In contrast, habitat use was sensitive to few variables on days when wolves were present: the best models averaged only 2.5 parameters. Because fewer variables affect elk behavior in the presence of wolves, we conclude that elk use simpler decision-making rules in the presence of wolves. This simplification of decision-making rules implies that predation risk imposes selection pressures that do not allow prey to respond to other pressures in ways that they otherwise would. If the affected processes are important, then this indirect effect of predation is likely to be important.

Within predator–prey systems behavior can heavily influence spatial dynamics, and accordingly, the theoretical study of how spatial dynamics relate to stability within these systems has a rich history. However, our understanding of these behaviors in large mammalian systems is poorly developed. To address the relationship between predator selection patterns, prey density, and prey vulnerability, we quantified selection patterns for two fine-scale behaviors of a recovering wolf (Canis lupus) population in Yellowstone National Park, Wyoming, USA. Wolf spatial data were collected between November and May from 1998–1999 until 2001–2002. Over four winters, 244 aerial locations, 522 ground-based telemetry locations, 1287 km of movement data from snow tracking, and the locations of 279 wolf kill sites were recorded. There was evidence that elk (Cervus elaphus) and bison (Bison bison) densities had a weak effect on the sites where wolves traveled and made kills. Wolf movements showed a strong selection for geothermal areas, meadows, and areas near various types of habitat edges. Proximity to edge and habitat class also had a strong influence on the locations where elk were most vulnerable to predation. There was little evidence that wolf kill sites differed from the places where wolves traveled, indicating that elk vulnerability influenced where wolves selected to travel. Our results indicate that elk are more vulnerable to wolves under certain conditions and that wolves are capable of selecting for these conditions. As such, vulnerability plays a central role in predator–prey behavioral games and can potentially impact the systems to which they relate.

Large carnivores are in global decline, particularly outside of government protected areas such as national parks. Much is known about the ecology of lions in protected areas, but little is known about situations in which lions coexist with people and livestock. In the Olkiramatian and Shompole area of Kenya's South Rift, lions move among areas with different land uses to avoid direct interactions with people and livestock. When the separation between lions and people is low, lions mount strong glucocorticoid stress responses. These results confirm that access to areas with low disturbance and little interaction with people is important for the conservation of lions outside of national parks., Large carnivore populations are in global decline, and conflicts between large carnivores and humans or their livestock contribute to low tolerance of large carnivores outside of protected areas. African lions (Panthera leo) are a conflict-prone species, and their continental range has declined by 75% in the face of human pressures. Nonetheless, large carnivore populations persist (or even grow) in some areas that are occupied by humans. Lions attain locally high density in the Olkiramatian and Shompole Group Ranches of Kenya's South Rift region, despite residence by pastoralist Maasai people and their sheep, goats, and cattle. We have previously found that these lions respond to seasonal movements of people by moving away from occupied settlements, shifting into denser habitats when people are nearby, and moving into a protected conservation area when people move into the adjacent buffer zone. Here, we examined lion stress responses to anthropogenic activities, using enzyme-linked immunoassay to measure the concentration of faecal glucocorticoid metabolites in 136 samples collected from five lion groups over 2 years. Faecal glucocorticoid metabolite concentrations were significantly lower for lions in the conservation area than for lions in the human-settled buffer zone, and decreased significantly with increasing distance to the nearest occupied human settlement. Faecal glucocorticoid metabolite concentrations were not detectably related to fine-scaled variation in prey or livestock density, and surprisingly, faecal glucocorticoid metabolite concentrations were higher in the wet season, when regional prey abundance was high. Lions coexist with people and livestock on this landscape by adjusting their movements, but they nonetheless mount an appreciable stress response when conditions do not allow them to maintain adequate separation. Thus, physiological data confirm inferences from prior data on lion movements and habitat use, showing that access to undisturbed and protected areas facilitates human–lion coexistence in a broader landscape that is used by people and livestock.