Vigilance Behaviour in Grazing African Antelopes

Behavioral decisions made under the risk of predation: a review and prospectus

Abstract: Predation has long been implicated as a major selective force in the evolution of several morphological and behavioral characteristics of animals. The importance of predation during evolutionary ti...

Predator vigilance and group size in mammals and birds: a critical review of the empirical evidence

Abstract: One commonly cited benefit to animals that forage in groups is an increase in the probability of detecting a predator, and a decrease in the time spent in predator detection. A mathematical model (Pulliam 1973) predicts a negative relationship between group size and vigilance rates. Over fifty studies of birds and mammals report that the relationship at least partly explains why individuals forage in groups. This review evaluates the strength of these conclusions based on their evidence. Those variables that may confound the relationship between vigilance and group size are outlined, and their control is assessed for each study. The variables I consider to be important include the density and type of food; competition between individuals; the proximity to both a safe place and the observer; the presence of predators; the visibility within the habitat; the composition of the group; the ambient temperature and the time of day. Based on these assessments, most of the studies fail to adequately demonstrate an unambiguous relationship between vigilance behavior and group size. Nevertheless, many studies reveal interesting features of the relationship between vigilance and group size that should provide fruitful avenues for future research.

Wolves, elk, and bison: reestablishing the "landscape of fear" in Yellowstone National Park, U.S.A.

Abstract: The elk or wapiti (Cervus elaphus) and bison (Bison bison) of Yellowstone National Park have lived in an environment free of wolves (Canis lupus) for the last 50 years. In the winter of 1994-1995, wolves were reintroduced into parts of Yellowstone National Park. Foraging theory predicts that elk and bison would respond to this threat by increasing their vigilance levels. We tested this prediction by comparing vigilance levels of elk and bison in areas with wolves with those of elk still in "wolf-free" zones of the Park. Male elk and bison showed no response to the reintroduction of wolves, maintaining the lowest levels of vigilance throughout the study (12 and 7% of the time was spent vigilant, respectively). Female elk and bison showed significantly higher vigilance levels in areas with wolves than in areas without wolves. The highest vigilance level (47.5 ± 4.1%; mean ± SE) was seen by the second year for female elk with calves in the areas with wolves and was maintained during the subsequent 3 years of...

Forage Quality and Aggregation by Large Herbivores

Abstract: Three general hypotheses have been proposed to explain why many large herbivores have highly aggregated patterns of distribution: dilution of predation risk, maintenance of forage in an immature bu...

Foraging-efficiency-predation-risk trade-off in the grey squirrel

Abstract: Many animals must often respond to environmental patterns that simulatneously influence both foraging efficiency and predation risk. We noted that grey squirrels ( Sciurus carolinensis ) sometimes immediately consume food items in areas of relatively great exposure to predators, and at other times carry food items to the safety of a tree prior to consumption. We outlined a hypothesis that the squirrels were somehow trading-off energy intake rate against predation risk. A simple model shows that maximal energetic efficiency is associated with immediate consumption, whereas (under the field conditions studied) carrying items to the safety of trees provides for minimal exposure to predation. Our analysis of the model predicts that the tendency to carry a food item should decrease with distance of food from cover (travel time) and increase with item size (handling time). To test our predictions, we presented free-roaming grey squirrels with patches containing a fixed number of identical food items. We estimated the proportion of items carried to trees before consumption for 12 different combinations of distance to the nearest tree and item size. The results support our hypothesis and indicate that a simple behavioural criterion based solely on foraging rate or time exposed to predators is insufficient to explain the variation in the data.

Observational study of behavior: sampling methods.

Abstract: Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.

Geometry for the selfish herd.

Abstract: This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator. It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others. Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations. The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.

The Social Organisation of Antelope in Relation To Their Ecology

Abstract: The types of social organisation displayed by the African antelope species have been assigned in this paper to five classes, distinguished largely by the strategies used by the reproductively active males in securing mating rights, and the effects of those strategies on other social castes. The paper attempts to show that these strategies are appropriate to each class because of the effects of other, ecological, aspects of their ways of life. The paper describes different feeding styles among antelope, in terms of selection of food items and coverage of home ranges. It argues that these feeding styles bear a relationship to maximum group size of feeding animals through the influence of dispersion of food items upon group cohesion. The feeding styles also bear a relationship to body size and to habitat choice, both of which influence the antelope species' antipredator behaviour. Thus feeding style is related to anti-predator behaviour which, in many species, influences minimum group size. Group size and the pattern of movement over the annual home range affect the likelihood of females being found in a given place at a given time, and it is this likelihood which, to a large extent, determines the kind of strategy a male must employ to achieve mating rights. The effects of the different strategies employed by males can be seen in such aspects of each species' biology as sexual dimorphism, adult sex ratio, and differential distribution of the sexes.

The Ethology of Predation

Abstract: 1 Internal Factors.- A. Hunger: Expression through Overt behavior.- I. Predatory Schedules.- 1. Patterns of Satiation.- 2. Feast and Famine.- II. Hunger and Diel Rhythms.- III. The Ramification of Hunger Effects.- 1. Capture-eliciting Prey Stimuli.- 2. Search behavior.- IV. The Motivation Underlying Feeding Responses.- 1. Hunger Thresholds of Feeding Response Components.- 2. The Complexity of Predatory Motivation.- V. The Diversity of Foraging Tactics.- VI. Feeding Components Affected and not Affected by Hunger.- B. The Control of Feeding Responses by Factors Other than Hunger.- I. The Readiness to Hunt.- II. Prey Storing.- III. Providing Food for Dependent Family Members.- C. The Problem of Specific Hungers.- I. Switching of Prey.- II. The Prey-density Predation Curve.- III. Swamping the Appetite of Predators.- D. Daily and Annual Rhythms in Predator-Prey Interactions.- I. Daily Rhythm of Predation.- II. Daily Activity Patterns of the Prey.- III. Annual Rhythm of Predation.- 2 Searching for Prey.- A. Path of Searching and Scanning Movements.- B. Area-concentrated Search.- I. Short-term Area Concentration.- 1. Living Scattered and Area-concentrated Search.- 2. The Nature of the Path Changes.- 3. Search Behavior after the Disappearance of Prey.- II. Long-term Area Concentration.- III. One-prey : One-place Association.- C. Object-concentrated Search.- I. Existence and Properties of "Searching Image".- 1. Ecological Evidence.- 2. Experimental Evidence.- II. Social Facilitation of Searching Image Formation.- III. Searching Image and "Training Bias".- IV. Searching Image and Profitability of Hunting.- 1. Ecological Evidence for Profitability of Hunting.- 2. Experimental Evidence for Profitability of Hunting.- V. Prey-specific Expectation.- VI. Ecological Implications of Searching Image.- 3 Prey Recognition.- A. The Stimulus-specificity of Prey Capture.- I. Capture-eliciting Prey Stimuli.- II. Capture-inhibiting Prey Stimuli.- B. One-prey : One-response Relationships.- C. The Assessment of the Circumstances of a Hunt.- D. Prey Recognition by Prey-related Signals.- E. Prey Stimulus Summation.- F. Novelty Versus Familiarity.- I. The Rejection of Novel Prey.- II. Familiarization with Prey and Its Consequences.- G. The Multi-channel Hypothesis of Prey Recognition.- 4 Prey Selection.- A. Preying upon the Weak and the Sick.- B. Preying upon the Odd and the Conspicuous.- C. The Mechanics of Prey Selection.- D. Evolutionary Implications.- 5 Hunting for Prey.- A. Modes of Hunting.- I. Hunting by Speculation.- II. Stalking and Ambushing.- 1. Stalking.- 2. Ambushing.- III. Prey Attack under Disguise.- IV. Pursuit of the Prey.- 1. Changes of Velocity of Attack (Pursuit).- 2. Interception of the Flight Path.- 3. Counteradaptations of the Prey.- V. Exhausting Dangerous Prey.- VI. Insinuation.- VII. Scavenging and Cleptoparasitism.- 1. Modes and Extent.- 2. Cleptoparasitism and Competition.- 3. Counter-measures of the Robbed.- VIII. Tool-use.- IX. Mutilation.- B. The Diversity of Hunting Methods.- I. Prey-specific Methods.- II. Situation-specific Methods.- III. Mechanisms and Causes of Predatory Versatility.- 1. General.- 2. Individual Predatory Repertories.- 3. The Persistence of Individual Traits.- 4. Predatory Specialization and Structural Modification.- 5. Predatory Versatility in Relation to Prey Availability.- C. Behavioral Aspects of Hunting Success.- I. A Comparison of Hunting Success across Predator Species.- II. Variables Influencing Hunting Success within Predator Species.- III. Aspects of Communal Hunting.- 1. Modes and Properties of Communal Hunting.- 2. Factors Conducive to Communal Hunting.- 3. Benefits of Communal Hunting.- References.- Scientific Names of Animals and Plants.