Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

There is growing recognition of the important roles played by predators in regulating ecosystems and sustaining biodiversity. Much attention has focused on the consequences of predator-regulation of herbivore populations, and associated trophic cascades. However apex predators may also control smaller 'mesopredators' through intraguild interactions. Removal of apex predators can result in changes to intraguild interactions and outbreaks of mesopredators ('mesopredator release'), leading in turn to increased predation on smaller prey. Here we provide a review and synthesis of studies of predator interactions, mesopredator release and their impacts on biodiversity. Mesopredator suppression by apex predators is widespread geographically and taxonomically. Apex predators suppress mesopredators both by killing them, or instilling fear, which motivates changes in behaviour and habitat use that limit mesopredator distribution and abundance. Changes in the abundance of apex predators may have disproportionate (up to fourfold) effects on mesopredator abundance. Outcomes of interactions between predators may however vary with resource availability, habitat complexity and the complexity of predator communities. There is potential for the restoration of apex predators to have benefits for biodiversity conservation through moderation of the impacts of mesopredators on their prey, but this requires a whole-ecosystem view to avoid unforeseen negative effects. 'Nothing has changed since I began. My eye has permitted no change. I am going to keep things like this.' From 'Hawk Roosting', by Ted Hughes.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1461-0248.2009.01347.x

Predator interactions, mesopredator release and biodiversity conservation

We review the concepts and research associated with measuring fear and its consequences for foraging. When foraging, animals should and do demand hazardous duty pay. They assess a foraging cost of predation to compensate for the risk of predation or the risk of catastrophic injury. Similarly, in weighing foraging options, animals tradeoff food and safety. The foraging cost of predation can be modelled, and it can be quantitatively and qualitatively measured using risk titrations. Giving-up densities (GUDs) in depletable food patches and the distribution of foragers across safe and risky feeding opportunities are two frequent experimental tools for titrating food and safety. A growing body of literature shows that: (i) the cost of predation can be big and comprise the forager’s largest foraging cost, (ii) seemingly small changes in habitat or microhabitat characteristics can lead to large changes in the cost of predation, and (iii) a forager’s cost of predation rises with risk of mortality, the forager’s energy state and a decrease in its marginal value of energy. In titrating for the cost of predation, researchers have investigated spatial and temporal variation in risk, scale-dependent variation in risk, and the role of predation risk in a forager’s ecology. A risk titration from a feeding animal often provides a more accurate behavioural indicator of predation risk than direct observations of predator-inflicted mortality. Titrating for fear responses in foragers has some well-established applications and holds promise for novel methodologies, concepts and applications. Future directions for expanding conceptual and empirical tools include: what are the consequences of foraging costs arising from interference behaviours and other sources of catastrophic loss? Are there alternative routes by which organisms can respond to tradeoffs of food and safety? What does an animal’s landscape of fear look like as a spatially explicit map, and how do various environmental factors affect it? Behavioural titrations will help to illuminate these issues and more.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1461-0248.2004.00661.x

Hazardous duty pay and the foraging cost of predation

Landscape Ecology in Theory and Practice

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.

https://www.jstor.org/stable/pdfplus/3450747.pdf

Elk alter habitat selection as an antipredator response to wolves

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...

/pdf/wolves-elk-and-bison-reestablishing-the-landscape-of-fear-in-2mfl8dchlb.pdf

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

Predation risk\" and \"fear\" are concepts well established in animal behavior literature. We expand these concepts to develop the model of the \"landscape of fear\". The landscape of fear represents relative levels of predation risk as peaks and valleys that reflect the level of fear of predation a prey experiences in different parts of its area of use. We provide observations in support of this model regarding changes in predation risk with respect to habitat types, and terrain characteristics. We postulate that animals have the ability to learn and can respond to differing levels of predation risk. We propose that the landscape of fear can be quantified with the use of well documented existing methods such as giving- up densities, vigilance observations, and foraging surveys of plants. We conclude that the landscape of fear is a useful visual model and has the potential to become a unifying ecological concept.

/pdf/the-landscape-of-fear-ecological-implications-of-being-3av2mhurxm.pdf

The Landscape of Fear: Ecological Implications of Being Afraid~!2009-09-09~!2009-11-16~!2010-02-02~!

Predation risk" and "fear" are concepts well established in animal behavior literature. We expand these concepts to develop the model of the "landscape of fear". The landscape of fear represents relative levels of predation risk as peaks and valleys that reflect the level of fear of predation a prey experiences in different parts of its area of use. We provide observations in support of this model regarding changes in predation risk with respect to habitat types, and terrain characteristics. We postulate that animals have the ability to learn and can respond to differing levels of predation risk. We propose that the landscape of fear can be quantified with the use of well documented existing methods such as giving- up densities, vigilance observations, and foraging surveys of plants. We conclude that the landscape of fear is a useful visual model and has the potential to become a unifying ecological concept.

The Landscape of Fear: Ecological Implications of Being Afraid

In 1995, wolves Canis lupus were reintroduced into Yellowstone National Park, USA, where they began to prey on ungulate species. In response to this new predation risk by wolves, we predicted that the two main ungulate species, elk Cervus elaphus and bison Bison bison, should compensate by reducing their use of riskier open meadows and increasing their use of safer forest. Additionally we predicted that this shift in habitat use would result in reduced diet quality. We tested the first prediction by regressing the number of faecal groups in 10-m2 sampling plots against distance from forest edges. To test the second prediction, we compared percent faecal nitrogen in elk and bison faeces between areas with and without wolves. We found a significant negative relationship between number of elk faecal groups and distance from forest edge in areas with wolves (r2 = 0.65, P = 0.001), but we did not find a relationship between these two factors in areas without wolves. Mean percent faecal nitrogen in elk...

/pdf/foraging-in-the-landscape-of-fear-and-its-implications-for-7lh2lioluj.pdf

Foraging in the ‘landscape of fear’ and its implications for habitat use and diet quality of elk Cervus elaphus and bison Bison bison

Identifying factors that may be responsible for regulating the size of animal populations is a cornerstone in understanding population ecology. The main factors that are thought to influence population size are either resources (bottom-up), or predation (top-down), or interspecific competition (parallel). However, there are highly variable and often contradictory results regarding their relative strengths and influence. These varied results are often interpreted as indicating "shifting control" among the three main factors, or a complex, nonlinear relationship among environmental variables, resource availability, predation, and competition. We argue here that there is a "missing link" in our understanding of predator-prey dynamics. We explore whether the landscape-of-fear model can help us clarify the inconsistencies and increase our understanding of the roles, extent, and possible interactions of top-down, bottom-up, and parallel factors on prey population abundance. We propose two main predictions derived from the landscape-of-fear model: (1) for a single species, we suggest that as the makeup of the landscape of fear changes from relatively safe to relatively risky, bottom-up impacts switch from strong to weak as top-down impacts go from weak to strong; (2) for two or more species, interspecific competitive interactions produce various combinations of bottom-up, top-down, and parallel impacts depending on the dominant competing species and whether the landscapes of fear are shared or distinctive among competing species. We contend that these predictions could successfully explain many of the complex and contradictory results of current research. We test some of these predictions based on long-term data for small mammals from the Chihuahuan Desert in the United States, and Mexico. We conclude that the landscape-of-fear model does provide reasonable explanations for many of the reported studies and should be tested further to better understand the effects of bottom-up, top-down, and parallel factors on population dynamics.

Lucina Hernández

Papers

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

The Landscape of Fear: Ecological Implications of Being Afraid~!2009-09-09~!2009-11-16~!2010-02-02~!

The Landscape of Fear: Ecological Implications of Being Afraid

Foraging in the ‘landscape of fear’ and its implications for habitat use and diet quality of elk Cervus elaphus and bison Bison bison

The landscape of fear: the missing link to understand top-down and bottom-up controls of prey abundance?