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For cooperative breeders, we hypothesize that the effects of group size on reproduction and survival might run in opposition if the benefits of grouping cannot be shared without cost. We tested this hypothesis by examining relationships between group size, survival, and reproduction in African wild dogs (Lycaon pictus), cooperative hunters with highly cohesive packs within which reproduction is monopolized by the dominant male and female. The production and survival of pups are known to increase with increasing pack size, but the effect of pack size on adult survival has not been examined previously. Data from 366 individuals over a period of 6 years showed that the survival of adults decreased with increasing pack size, with a 25% difference between the largest and smallest packs after controlling for the effects of age, sex, social status, year of study, and pack identity. Several tests confirmed that undetected dispersal is unlikely to have produced this pattern. These results suggest that cooperative breeding in wild dogs cannot be fully explained by mutual direct benefit, thus reinforcing the prior inference that kin selection plays an important role in the evolution of their cooperation. The results also weaken support for the hypothesis that wild dogs are extinction prone due to group-level Allee effects. More broadly, the relationship of effects of group size on survival and reproduction might be predicted by considering whether cooperation yields benefits that accrue to all group members (e.g., through cooperative vigilance) or benefits that must be apportioned to individuals (e.g., through cooperative hunting).

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.

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]

Models dealing with the relationship of inbreeding to dispersal, the evolution of cooperative breeding, and the adaptive significance of reproductive suppression all involve a parameter describing die probability that emigration ends in death. Surprisingly, estimates of this parameter (or equivalently, of the probability of dispersing successfully) are virtually absent from the literature. There are formidable difficulties in estimating dispersal risk by direct observation of emigrants, but we contend that many investigators have already collected demographic information that can be used to estimate it indirectly. The root problem is that the proportion of the population diat emigrates from its natal group or home range, e, is not generally observable. Nevertheless, for each value of e diere is only one possible survival rate of emigrants (st) and of philopatric animals (sf) diat can explain the overall survival rate. For study sites Uiat are neither sources nor sinks, we describe a graphical framework within which census data and behavioral observations can be used to place bounds on e, s,, and sp. We use data from a Serengeti population of the dwarf mongoose (HelogaU parvula) to illustrate how different types of data and assumptions allow us to construct better estimates of these parameters. For dwarf mongooses, die approach reveals unexpected differences in dispersal risk among sex and age classes.

Dwarf mongooses (Helogale parvula) are small, communally breeding carnivores found in woodland and tree-savanna throughout Africa. Within a pack, socially subordinate mongooses do not normally breed, yet they invariably participate in all aspects of parental care. The primary alternative to tolerating reproductive suppression is dispersal, which shortens the wait for dominance and breeding. Here, we calculate the annual inclusive fitness payoffs to the dispersing and nondispersing strategies for males and females of ages one to seven, using data from a 14-year study in Serengeti National Park, Tanzania. Factors with effects on inclusive fitness included relatedness to pack mates before and after dispersal, the effect of help on recipients' reproductive success, the probability of dispersing successfully, the probability of attaining dominance, and reproductive success after attaining dominance. All of these factors differed between male and female dwarf mongooses. We compared the contributions of direct and indirect fitness to the total fitness of mongooses pursuing each of the strategies, across a range of ages. In our population, dispersal and nondispersal both yielded direct and indirect payoffs at most ages. For dispersers of both sexes, direct fitness was the primary component of total fitness but indirect fitness was substantial for young (< 2 years old) dispersers. For nondispersers of both sexes, indirect fitness was the major component of total fitness among young mongooses (up to 2 or 3 years), but direct fitness was the major component among older mongooses. By comparing the inclusive fitness payoffs for the two strategies, we determined the range of ages at which dispersal should be favored for each sex. These comparisons correctly predicted that males should be more dispersive than females at all ages, and that males should disperse over a broader range of ages.