Cognitive Ecology II
edited by Reuven Dukas and John M. Ratcliffe
University of Chicago Press, 2009
Cloth: 978-0-226-16935-4 | Paper: 978-0-226-16936-1 | Electronic: 978-0-226-16937-8
DOI: 10.7208/chicago/9780226169378.001.0001


Merging evolutionary ecology and cognitive science, cognitive ecology investigates how animal interactions with natural habitats shape cognitive systems, and how constraints on nervous systems limit or bias animal behavior. Research in cognitive ecology has expanded rapidly in the past decade, and this second volume builds on the foundations laid out in the first, published in 1998.

Cognitive Ecology II integrates numerous scientific disciplines to analyze the ecology and evolution of animal cognition. The contributors cover the mechanisms, ecology, and evolution of learning and memory, including detailed analyses of bee neurobiology, bird song, and spatial learning. They also explore decision making, with mechanistic analyses of reproductive behavior in voles, escape hatching by frog embryos, and predation in the auditory domain of bats and eared insects. Finally, they consider social cognition, focusing on alarm calls and the factors determining social learning strategies of corvids, fish, and mammals.

With cognitive ecology ascending to its rightful place in behavioral and evolutionary research, this volume captures the promise that has been realized in the past decade and looks forward to new research prospects.


Reuven Dukas is associate professor of psychology, neuroscience, and behavior, and a member of the Animal Behaviour Group at McMaster University. John Ratcliffe is assistant professor at the Center for Sound Communication at the Institute of Biology of the University of Southern Denmark.


“A fundamental challenge in modern science is to understand the links from brains to the plastic behaviour of individuals, groups, societies and species assemblages—and to understand how these interactions are fashioned within an evolutionary and ecological context. Dukas and Ratcliffe and their impressive team of contributors accept this challenge in a fascinating and expertly edited volume that begins with the ultimate and proximate mechanisms of learning, then addresses exciting advances in avian cognition, before proceeding to decision making in mate choice and predator-prey interaction and culminating in the role of cognition in sociality. Biologists of all descriptions as well as psychologists and social scientists need to read this book.”

— Stephen J. Simpson, University of Sydney

Cognitive Ecology II is truly impressive. The wide range of questions addressed demonstrates this field’s vitality and broad appeal. It illustrates basic principles using examples from across the animal kingdom and a wide variety of sensory systems and adeptly combines progress reports on well-established topics (e.g. bird song) with syntheses of emerging topics (e.g. social information processing). Ten years ago, Dukas’s first Cognitive Ecology identified a new field; Cognitive Ecology II will define and motivate this exciting and important discipline for the next decade.”

— David Stephens, University of Minnesota

“In the first Cognitive Ecology, Reuven Dukas succeeded in bringing together the disparate threads of research in a then just emerging discipline. The book was certainly not alone in attending to animal cognition, but for behavioural ecologists it served as a reference that made clear that learning, attention, and perception could no longer be ignored. More than ten years later, Dukas’ and Ratcliffe’s Cognitive Ecology II provides vivid evidence that the field has achieved maturity and structure. The book updates some of the major topics covered in the first volume, such as avian song, predator-prey interactions, and mate choice, and offers a treatment of emerging fields such as social information use and innovations. But most importantly, it continues in its role of reference for all those that ask questions about behavioural mechanisms. If you need to assess the current state of animal cognition and want to know which questions will pave the way for the coming decade, then Cognitive Ecology II is an absolute must-read. Buy it.”

— Luc-Alain Giraldeau, L'Université du Québec à Montréal



- Reuven Dukas, John M. Ratcliffe
DOI: 10.7208/chicago/9780226169378.003.0001
[cognitive ecology, ecology, cognition, neuronal processes, antipredatory behavior, social behavior]
This chapter provides an introduction to cognitive ecology that gives attention to ecology and the evolution of “cognition,” defining it as the neuronal processes concerned with the acquisition, retention, and use of information. Cognition can be divided into several interrelated and inseparable components including perception, learning, working memory, attention, long-term memory, and decision making. The chapter provides an overview of research programs relating cognition to avian ecology, as well as a brief discussion on cognitive aspects of decisions made within reproduction and antipredator behavior categories. It also shows a link between antipredatory and social behavior by analyzing the alarm calls of meerkats. (pages 1 - 4)
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Part I: Learning: Ultimate and Proximate Mechanisms

- Reuven Dukas
DOI: 10.7208/chicago/9780226169378.003.0002
[acquisition, retention, learning, memory, ecology, behavior, speciation]
This chapter discusses the ecological and evolutionary significance of acquisition and retention. The mechanistic research on animal learning and memory is typically conducted under the necessary controlled laboratory conditions using a few animal species whose ecology and behavior in the wild are not well known. The author finds that the increased understanding of the neurogenetic mechanisms underlying learning and memory has led to the realization that there is great similarity in these mechanisms across all animals. Though learning is a key factor in the life history of most animals, it has not been well integrated into the life history literature, which has focused on physical traits such as growth, effort, and senescence. Also, recent work on mechanisms of speciation has made it clear that learning can play an important role in population divergence. (pages 7 - 26)
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- Susan E. Fahrbach, Scott Dobrin
DOI: 10.7208/chicago/9780226169378.003.0003
[plasticity, honeybee, brain, Apis mellifera, genes, experience-dependent changes]
The mechanistic research on animal learning and memory is typically conducted under the necessary controlled laboratory conditions using a few animal species whose ecology and behavior in the wild are not well known. A notable exception is the honeybee, Apis mellifera, which has been studied extensively in this chapter. The authors illustrate the honeybee as an ideal model system for integrating mechanistic knowledge on genes, neurons, and hormones with whole-animal information on behavior and ecology. Though the chapter focuses on studies of the honeybee, experience-dependent brain plasticity is not a rare phenomenon. A behavioral neuroscientist can be confident that it is happening in the animal model. Association of the experience-dependent changes in brain structure to their functional consequences is important because doing so will provide an insight into a powerful source of individual (experience-dependent) differences in animal behavior. (pages 27 - 46)
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Part II: Avian Cognition: Memory, Song, and Innovation

- Michael D. Beecher, John M. Burt
DOI: 10.7208/chicago/9780226169378.003.0004
[song sparrow, song learning, song communication, social ecology, cognitive ecology, songbirds]
In later life This chapter presents an update on a long-term research program examining the rules that young song sparrows (Melospiza melodia) use to decide which songs to learn, retain, and later use, and the role that the social ecology of song sparrows has played in shaping these rules. The strategy of song learning in song sparrows and the mechanisms of song communication between neighboring adult males are shaped by cognitive factors at the proximate level and variables in the species' social ecology at the ultimate level. The chapter presents an argument around the fact that a cognitive ecology perspective captures the key features of song learning and song communication in this species, and will probably do so as well in other species of songbirds. The authors find that understanding the song-learning process may ultimately be the key to understanding why it is that song learning in one life stage parallels song communication stages. (pages 49 - 70)
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- William A. Searcy, Stephen Nowicki
DOI: 10.7208/chicago/9780226169378.003.0005
[nutritional stress, male song, male quality, stress, brood size, song system]
This chapter reviews the bird song learning by examining the recent nutritional stress hypothesis. It suggests that conditions during early development have long-term effects on features of the male song, which females can readily perceive, and on other aspects of male quality, which females cannot easily assess. It is therefore found that females rely on male song as a reliable indicator of overall male quality. Also, a variety of forms of stress has been shown to affect either adult song or the song system or both, including nutritional limitation, orticosterone treatment, and parasite infection. The authors observe that the one exception has been brood size manipulation, which had no effect on either song or the song system in zebra finches. Hence, further studies of the effects of early stress on adult social dominance, immunocompetence, and cognitive abilities would be particularly interesting. (pages 71 - 87)
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- Vladimir V. Pravosudov
DOI: 10.7208/chicago/9780226169378.003.0006
[spatial memory, hippocampus, nutritional stress, developmental constraint, birds, brain]
This chapter explains the effects of nutritional stress during development, with a focus on the hippocampus and spatial memory. Research work done on spatial memory in birds has provided a clear link between the ecological need to store food, a relatively enhanced spatial memory used to retrieve the cached food, and the relative volume of the hippocampus, the brain region processing spatial memory. With the importance of spatial memory for the survival of certain bird species, the author predicted that hippocampal development, and hence spatial memory, would remain intact even under nutritional stress. The data in the chapter refute that prediction and suggest that constraints during development preclude the insulation of certain brain regions from nutritional stress. Also, favoring animals with better spatial learning might provide an easier path for natural selection to select for better parents to produce high-quality offspring, rather than resolving the potential developmental constraint issue. (pages 88 - 110)
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- Daniel Sol
DOI: 10.7208/chicago/9780226169378.003.0007
[cognitive-buffer hypothesis, large brain, brain evolution, positive-feedback, body size, brain]
This chapter relies on recent data to address the old question of why some animals have large brains relative to body size even though such brains incur substantial costs in terms of delayed maturation and high maintenance. It reviews recent studies providing support for the cognitive-buffer hypothesis, which states that a relatively large brain is associated with an enhanced ability to handle novel situations and hence with increased probability of survival in novel or altered environments. The cognitive-buffer hypothesis is the most general explanation for the benefits of the evolution and development of large brains, proposing that a major advantage of a large brain is to produce behavioral responses that protect the animal from the vagaries of the environment. The buffer function of the brain has the potential to generate “autocatalytic” and positive-feedback processes that, although still not well understood, could accelerate brain evolution. (pages 111 - 134)
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Part III: Decision Making: Mate Choice and Predator-Prey Interactions

- Michael J. Ryan, Karin L. Akre, Mark Kirkpatrick
DOI: 10.7208/chicago/9780226169378.003.0008
[sexual selection, mate choice, mating signals, cognitive biology, social groups, phenotype]
Female choice of mates has been studied for a long time, and as this chapter explains, it is well established that females choose mates based on their perceived quality. It is observed that such female choice has generated sexual selection, which is responsible for the evolution of many features of male mating signals. Social groups can provide females with an enormous amount of information to consider in mate choice, as demonstrated by mate choice copying; thus, a male's utility is defined not only by his phenotype but also by how others in the social group, besides the choosing female, react to his phenotype. The authors analyze how cognitive mechanisms could affect female mate choice and identify exciting directions in this relatively unexplored avenue of research. (pages 137 - 155)
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- Steven M. Phelps, Alexander G. Ophir
DOI: 10.7208/chicago/9780226169378.003.0009
[sexual behavior, mammalian model, mating systems, pair-bonding, V1aR, tactic]
This chapter focuses on male sexual behavior within a unique mammalian model that relies on natural variation in the mating systems of voles of the genus Microtus. The authors use their thorough knowledge of voles' natural history and behavior, and discuss the neuroanatomy, neurochemistry, and genetics underlying males' pair-bonding and their corresponding use of space. It is observed that by making pair-bonding and residency contingent on repeated and prolonged mating, the mechanism ensures that a male is likely to be a successful resident before committing to the tactic. The association of cingulate V1aR variation with intrapair and extrapair paternity highlights how behavioral specializations can make conflicting demands of cognitive substrates. Because new methods permit the manipulation of V1aR and other genes in natural environments, such studies promise to clarify both the mechanisms of natural behavior and the origins of behavioral diversity. (pages 156 - 176)
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- Karen M. Warkentin
DOI: 10.7208/chicago/9780226169378.003.0010
[embryonic stage, predation, information, detection, decision theory]
This chapter reviews recent experiments related to cognition at the embryonic stage. It is considered that that embryo possesses sophisticated abilities to assess and respond to cues of predation. Information, detection, and decision theory have played key roles in studies of animal communication. Students of predator–prey interactions have considered how prey should behave under the typical context of imperfect information about risk. The authors note that embryonic cognition is an essential yet relatively neglected aspect of cognitive research even though it is relevant for many animals. The association of empirical studies of risk assessment mechanisms to the theoretical framework of information, detection, and decision theory provides both testable hypotheses for empirical work and new contexts in which to assess the generality of these bodies of theory. (pages 177 - 200)
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- John M. Ratcliffe
DOI: 10.7208/chicago/9780226169378.003.0011
[bat, moth, auditory, predation, reproduction, call emission rate, predator–prey interaction, auditory world]
This chapter reviews the historically well-studied system of bats and moths. This classic model system focuses on the auditory domain used by bats to detect insects and by many moths to attempt to avoid impending predation. While long-lived bats have been underestimated with respect to behavioral flexibility and learning, moths are short-lived vehicles for reproduction and are not expected to be terribly plastic in their sensoribehavioral responses. The decisions noctuid moths make when faced with female pheromones and batlike ultrasound, and the ability of arctiids to assess the relative risk of hunting bats based on call emission rate alone, imply that they too employ evolutionary strategies. Because bats produce echolocation calls to perceive their environment, and because some moths generate sounds to deter attacking bats, researchers eavesdropping on these signals have a unique opportunity to examine how the available auditory information translates into decisions made by both predator and prey. (pages 201 - 226)
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Part IV: Cognition and Sociality

- Marta B. Manser
DOI: 10.7208/chicago/9780226169378.003.0012
[alarm calls, predators, escape response, adaptive response, functionally referential calls]
This chapter links antipredatory and social behavior by analyzing the alarm calls of meerkats (Suricata suricatta). These cooperatively breeding animals employ referential alarm calls, which indicate the approach of specific predators and cause receivers to show an appropriate escape response to these predators. Functionally, referential calls refer to specific stimuli in the external environment of the caller and cause receivers to show an adaptive response to them. In some species, the predator-specific calls appear to refer to a predator species or category, whereas in others they appear to refer to the spatial area from which the predator is approaching, or its behavior. Neurobiological studies will help to identify whether the same regions of the brain are activated during encounters that elicit functionally referential calls and encounters that cause animals to change their call rate or other acoustic characteristics along a continuum. (pages 229 - 248)
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- Rachel L. Kendal, Isabelle Coolen, Kevin N. Laland
DOI: 10.7208/chicago/9780226169378.003.0013
[social learning, acquisition, trade-offs, asocial learning, behavioral traditions]
This chapter reviews the current knowledge on social learning. From humans' biased perspective, social learning is a basic way of life. Most animals do not rely on social learning, whereas some species use it only conditionally. The authors organize their discussion around the two key questions of when individuals should rely on social learning and whom they should learn from. Individual characteristics of observers, favoring the overriding of social learning strategies and the continued acquisition of personal information, may be influential in determining the innovatory capacities of individuals. It is hoped that consideration of the trade-offs inherent in the adaptive use of social and asocial learning will contribute to an increased understanding of the observed pattern of social learning and behavioral traditions in the animal kingdom, especially as the use of social information may lead to cultural evolution, which may in turn affect biological evolution. (pages 249 - 271)
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- Ira G. Federspiel, Nicola S. Clayton, Nathan J. Emery
DOI: 10.7208/chicago/9780226169378.003.0014
[3Es, ecology, evolutionary history, social information, natural behavior]
This chapter illustrates how integrating knowledge about animals' natural behavior, ecology, and evolutionary history with the powerful empirical techniques of experimental psychology has helped us to understand the use of social information by different bird species. To understand these mechanisms and to define the ways in which they differ between species, an integrative approach must be adopted combining the 3Es with knowledge of experimental psychology, to obtain rigorous experimental validity. More comparative work is clearly needed to determine the influence of ecology and evolutionary history on social information use, and experiments using different tasks should shed light on whether the capacity to use social information is domain specific. The knowledge of the 3Es can thus be used to influence a particular species' life history, not only to drive the development of appropriate research questions and methodology but also to provide post hoc explanations of successes and failures in psychological experiments. (pages 272 - 297)
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- Reuven Dukas, John M. Ratcliffe
DOI: 10.7208/chicago/9780226169378.003.0015
[learning, memory, behavioral flexibility, cognitive mechanisms, cognitive ecology]
This final chapter summarizes the arguments and discussions of the book. The chapters in this book have dealt with a huge range of topics in a vast assortment of animals. Learning, memory, and overall behavioral flexibility have been analyzed from a variety of angles. A clear thread has linked the chapters together, and this is the underlying approach of relating cognitive mechanisms to animal ecology and evolution. This final chapter also talks about what is missing from this discussion and looks towards future directions. Overall, it can stated that the major challenge facing future work in cognitive ecology is the further integration of knowledge on the genetic, neurobiological, and endocrinological mechanisms underlying cognition with information about animal ecology and evolution. (pages 298 - 300)
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