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An Introduction to Animal Behaviour: An Integrative Approach
Animal Behaviour 84 (2012) 1279–1280
Contents lists available at SciVerse ScienceDirect
Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav
Book Review An Introduction to Animal Behaviour: An Integrative Approach, By Michael J. Ryan, Walter Wilczynski. Cold Spring Harbor, New York: Cold Spring Harbor Press (2011). Pp. xiD258. Price $46.00 paperback. Many introductory texts and courses in animal behaviour begin with a treatment of the so-called levels of analysis in behaviour. Sometimes these levels are grouped as proximate versus ultimate causes of behaviour, sometimes as the four levels of analysis (mechanism, ontogeny, function and evolution) enumerated by Niko Tinbergen (1963) in his well-known paper, ‘On aims and methods of ethology’. Often, chapters and lectures in textbooks are organized roughly in sequence according to these levels. Material on the proximate causes of behaviour (mechanism and development) is typically presented first, followed by material on the ultimate causes (function and evolution). In this context, the uniqueness of this 258-page volume by Ryan & Wilczynski is apparent at the outset. In a first chapter that is recommended reading for instructors irrespective of whether they adopt the book for their course, Ryan & Wilczynski take the position that it is time to abandon the levels approach as currently employed in instruction, and to actively endeavour to integrate across these levels, as Tinbergen himself surely intended. The authors make their case in the first chapter using examples of the ways in which one level of analysis informs another level. A case study given of mechanism informing function centres around Hamilton & Brown’s (2001) hypothesis that bright red and yellow autumn foliage functions as a warning signal to aphids of the low nutrient value of these leaves. Studies of aphid colour vision do not support this hypothesis (Chittka & Döring 2007). Red foliage, far from acting as a conspicuous warning, likely appears dark or dull to aphids, which lack a red receptor. Bright yellow autumn foliage, by contrast, should be readily detectable. In fact, the amount of yellow in green foliage is used by insect herbivores such as aphids to distinguish high-quality foliage (high in yellow reflectance) from low-quality foliage (low in yellow reflectance). For that reason, yellow foliage might even be attractive to aphids, rather than repellent, and trapping studies suggest that indeed it is (Döring & Chittka 2007). Not only can mechanism inform function, but function can also inform mechanism, as the authors illustrate using bat navigation as an example. Simply put, the observation that bat flight is functional in the dark suggested to 18th century scientists that bats possessed a sense that we do not. Thus, the authors contend that the study of echolocation was ultimately inspired by an early appreciation of the function of that as-yet-undiscovered mechanism. Examples of development (or acquisition, as the authors call it) informing evolutionary history, and of evolutionary history informing function, are also presented in the first chapter. Chapters 2 and 3 continue the theme of integration, pairing proximate causes (mechanism and 0003-3472/$38.00 http://dx.doi.org/10.1016/j.anbehav.2012.09.004
acquisition) and ultimate causes (function and evolution), respectively. Both chapters serve to ground the reader in basic concepts throughout these levels, ranging widely from kin selection and selfish genes to central pattern generators and hormone function. The fourth chapter is on foraging and provides a good example of the authors’ integrative approach, illustrating both its strengths and weaknesses. The chapter begins with a characterization of physiological state as it affects the appetitive phase of foraging. This is followed by a description of the use of spatial memory by food-caching birds. Both parts give strong attention to underlying mechanism. Treatment of the consummatory phase of foraging extends the treatment of mechanism by reviewing several classic case studies of sensory systems in neuroethology: visually-based prey capture in toads, owl hunting using audition, and infrared sensing by snakes. From there, the chapter introduces the notion of food preferences, first discussing the Brodie group’s work on the coevolution of newt toxicity and snake prey preference (Geffeney et al. 2005), and later introducing foraging theory, including the classic two-prey diet choice model (MacArthur & Pianka 1966). The chapter ends by scrutinizing the perfect information and fitness proxy assumptions of the diet choice model. In short, the coverage of foraging in this volume is as broad-ranging as is found in any animal behaviour textbook, and more so than most, drawing from research and concepts in psychology, neuroethology, behavioural ecology and evolutionary biology. One challenge with ranging so broadly is that the field itself is not well integrated; for example, I know of no literature reviews that cover all these bases in foraging behaviour. The authors are thus navigating in uncharted waters, and the synthesis is consequently somewhat choppy and perhaps best termed a work in progress. The chapter is none the less absorbing and thorough. Chapters on migration and orientation, species recognition and mate choice, social bonding and cooperation, and conflict and aggression round out the book, all using a similar integrative approach. Integrative or not, readers will perceive a stronger emphasis on proximate causes in Ryan & Wilczynski’s effort than in other texts. Whereas behaviour textbooks have, over time, generally devoted less space to proximate causes of behaviour (Alcock 2003), this book is radically different in being rich in description of genetic, neural and hormonal mechanisms of behaviour. However, in contrast to the early textbooks, which focused on mechanism for its own sake, information on mechanism and development in Ryan & Wilczynski’s book is presented in a strongly ecological and evolutionary context. The relatively robust coverage of mechanism and development from a perspective of ultimate causes is what sets this book apart from other textbooks in behaviour. Given that knowledge of mechanisms and development, from genes to neural systems and endocrinological processes, is growing with astonishing rapidity, time would seem nigh for instructors with a functional and evolutionary bent to integrate mechanism and
1280
Book Review / Animal Behaviour 84 (2012) 1279–1280
development more completely into their teaching. This book can be an aid towards that end. Ryan & Wilczynski acknowledge that their book is not intended to be an exhaustive review of animal behaviour, as is found in the textbooks commonly used in animal behaviour courses. Topics that receive little if any treatment include personality and behavioural syndromes, and (perhaps surprisingly) intersexual conflict. Moreover, the effort at integration may sometimes trade off against adequate detail. One obstacle faced in using this book in a general animal behaviour course is a recurrent tendency for authors to allude to methods or topics without providing enough depth or citations for an inquisitive student to gain a full understanding. For instance, commenting on methods for phylogenetic reconstruction, authors write, ‘Parsimony is a nonparametric method that does not perform optimally under certain conditions, and many phylogeneticists rely on parametric techniques such as maximum likelihood or Bayesian inference’ (page 30). No further explanation is provided, no literature is cited within the text, and, sadly, the book does not have a glossary. Hence, substantial familiarity about statistical methods is presumed here. In my classes, the terms parametric techniques and nonparametric method would have significant name recognition mainly to my upperclassmen, and the terms maximum likelihood and Bayesian inference would probably be familiar to just a few of those. The same would be true of the definition of heritability (page 19), which includes the term additive genetic variation without defining what that is. Most likely, authors sacrificed such detail so as to keep the book of reasonable length; nevertheless, the brevity of explanation could frustrate students, especially motivated students who lack the necessary preparation. Textbooks tend to reflect the interests and expertise of their authors. In terms of coverage and proficiency in material, for example, the book is especially strong in the area of sensory systems; this strength is most palpable in sections on communication, navigation, and mate choice, which are superb. By comparison, treatment of cognitive processes, including learning, is less refined and not as in depth (the lack of depth perhaps exaggerated because material on learning is dispersed throughout the text). Not surprisingly, given Ryan & Wilczynski’s research interests, the chapter entitled ‘Species recognition, mate choice, and sexual selection’ far and away provides the most engaging and compelling application of the integrative approach. The book is predictably heavy in vertebrate case studies; that said, information on invertebrates, especially insects, is thoughtfully selected and expertly treated. In aesthetic terms, this compact book, a tad smaller in area than an iPad and about twice as thick, is a success. Figures are ample, generally simple in composition, and easy on the eyes. Figure legends are comprehensive and clearly written, albeit sometimes so extensive in length that perhaps the ‘box’ approach to
presentation might have been substituted for the figure and legend. Hard as I tried, I could not find typographical errors or errors in content. The writing is certainly more technical than in texts that focus less on mechanism, and is generally free of anecdotes and personal rejoinders. As such, the book does not have the high ‘readability factor’ that Alcock’s (2009) text, for example, possesses. While students may find the presentation drier than that text, the prose overall is nevertheless clear and straightforward enough for the motivated student to follow. The lack of exhaustive coverage, glossary, and teaching aids such as discussion questions may render this book less appealing to the instructor of large undergraduate classes in animal behaviour. As a text, the book seems most appropriate for a small class in animal behaviour composed of advanced undergraduates and first-year graduate students, in which students have ample opportunity to query the instructor for insights directly related to book content. For an instructor in animal behaviour who selects a more conventional textbook for their course, this volume would still be very useful as a reference. The examples are particularly well chosen and explanations are first-rate. A useful and welcome feature as a reference book is inclusion of material not found in many behaviour texts, for example on the evolution of sex, sex determination mechanisms, and meiotic drive as an example of selection at the level of the gene. In closing, I applaud the authors for taking a fundamentally different direction in their textbook by integrating multiple levels of analysis thoroughly in individual chapters. This book may not immediately steal the market away from various established texts in the area of animal behaviour, but it could nevertheless have an important impact on how animal behaviour is taught in the future. Daniel R. Papaj Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, U.S.A.
References Alcock, J. 2003. A textbook history of animal behaviour. Animal Behaviour, 65, 3–10. Alcock, J. 2009. Animal Behaviour: an Evolutionary Approach. 9th edn. Sunderland, Massachusetts: Sinauer. Chittka, L. & Döring, T. F. 2007. Are autumn foliage colors red signals to aphids? PLoS Biology, 5, e187. Döring, T. F. & Chittka, L. 2007. Visual ecology of aphids: a critical review on the role of colours in host finding. Arthropod–Plant Interactions, 1, 3–16. Geffeney, S. L., Fujimoto, E., Brodie, E. D., III, Brodie, E. D., Jr. & Ruben, P. C. 2005. Evolutionary diversification of TTX-resistant sodium channels in a predator– prey interaction. Nature, 434, 759–763. Hamilton, W. D. & Brown, S. P. 2001. Autumn tree colours as a handicap signal. Proceedings of the Royal Society B, 268, 1489–1493. MacArthur, R. H. & Pianka, E. R. 1966. On optimal use of a patchy environment. American Naturalist, 100, 603–609. Tinbergen, N. 1963. On aims and methods of ethology. Zeitschrift für Tierpsychologie, 20, 410–433.
Contents lists available at SciVerse ScienceDirect
Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav
Book Review An Introduction to Animal Behaviour: An Integrative Approach, By Michael J. Ryan, Walter Wilczynski. Cold Spring Harbor, New York: Cold Spring Harbor Press (2011). Pp. xiD258. Price $46.00 paperback. Many introductory texts and courses in animal behaviour begin with a treatment of the so-called levels of analysis in behaviour. Sometimes these levels are grouped as proximate versus ultimate causes of behaviour, sometimes as the four levels of analysis (mechanism, ontogeny, function and evolution) enumerated by Niko Tinbergen (1963) in his well-known paper, ‘On aims and methods of ethology’. Often, chapters and lectures in textbooks are organized roughly in sequence according to these levels. Material on the proximate causes of behaviour (mechanism and development) is typically presented first, followed by material on the ultimate causes (function and evolution). In this context, the uniqueness of this 258-page volume by Ryan & Wilczynski is apparent at the outset. In a first chapter that is recommended reading for instructors irrespective of whether they adopt the book for their course, Ryan & Wilczynski take the position that it is time to abandon the levels approach as currently employed in instruction, and to actively endeavour to integrate across these levels, as Tinbergen himself surely intended. The authors make their case in the first chapter using examples of the ways in which one level of analysis informs another level. A case study given of mechanism informing function centres around Hamilton & Brown’s (2001) hypothesis that bright red and yellow autumn foliage functions as a warning signal to aphids of the low nutrient value of these leaves. Studies of aphid colour vision do not support this hypothesis (Chittka & Döring 2007). Red foliage, far from acting as a conspicuous warning, likely appears dark or dull to aphids, which lack a red receptor. Bright yellow autumn foliage, by contrast, should be readily detectable. In fact, the amount of yellow in green foliage is used by insect herbivores such as aphids to distinguish high-quality foliage (high in yellow reflectance) from low-quality foliage (low in yellow reflectance). For that reason, yellow foliage might even be attractive to aphids, rather than repellent, and trapping studies suggest that indeed it is (Döring & Chittka 2007). Not only can mechanism inform function, but function can also inform mechanism, as the authors illustrate using bat navigation as an example. Simply put, the observation that bat flight is functional in the dark suggested to 18th century scientists that bats possessed a sense that we do not. Thus, the authors contend that the study of echolocation was ultimately inspired by an early appreciation of the function of that as-yet-undiscovered mechanism. Examples of development (or acquisition, as the authors call it) informing evolutionary history, and of evolutionary history informing function, are also presented in the first chapter. Chapters 2 and 3 continue the theme of integration, pairing proximate causes (mechanism and 0003-3472/$38.00 http://dx.doi.org/10.1016/j.anbehav.2012.09.004
acquisition) and ultimate causes (function and evolution), respectively. Both chapters serve to ground the reader in basic concepts throughout these levels, ranging widely from kin selection and selfish genes to central pattern generators and hormone function. The fourth chapter is on foraging and provides a good example of the authors’ integrative approach, illustrating both its strengths and weaknesses. The chapter begins with a characterization of physiological state as it affects the appetitive phase of foraging. This is followed by a description of the use of spatial memory by food-caching birds. Both parts give strong attention to underlying mechanism. Treatment of the consummatory phase of foraging extends the treatment of mechanism by reviewing several classic case studies of sensory systems in neuroethology: visually-based prey capture in toads, owl hunting using audition, and infrared sensing by snakes. From there, the chapter introduces the notion of food preferences, first discussing the Brodie group’s work on the coevolution of newt toxicity and snake prey preference (Geffeney et al. 2005), and later introducing foraging theory, including the classic two-prey diet choice model (MacArthur & Pianka 1966). The chapter ends by scrutinizing the perfect information and fitness proxy assumptions of the diet choice model. In short, the coverage of foraging in this volume is as broad-ranging as is found in any animal behaviour textbook, and more so than most, drawing from research and concepts in psychology, neuroethology, behavioural ecology and evolutionary biology. One challenge with ranging so broadly is that the field itself is not well integrated; for example, I know of no literature reviews that cover all these bases in foraging behaviour. The authors are thus navigating in uncharted waters, and the synthesis is consequently somewhat choppy and perhaps best termed a work in progress. The chapter is none the less absorbing and thorough. Chapters on migration and orientation, species recognition and mate choice, social bonding and cooperation, and conflict and aggression round out the book, all using a similar integrative approach. Integrative or not, readers will perceive a stronger emphasis on proximate causes in Ryan & Wilczynski’s effort than in other texts. Whereas behaviour textbooks have, over time, generally devoted less space to proximate causes of behaviour (Alcock 2003), this book is radically different in being rich in description of genetic, neural and hormonal mechanisms of behaviour. However, in contrast to the early textbooks, which focused on mechanism for its own sake, information on mechanism and development in Ryan & Wilczynski’s book is presented in a strongly ecological and evolutionary context. The relatively robust coverage of mechanism and development from a perspective of ultimate causes is what sets this book apart from other textbooks in behaviour. Given that knowledge of mechanisms and development, from genes to neural systems and endocrinological processes, is growing with astonishing rapidity, time would seem nigh for instructors with a functional and evolutionary bent to integrate mechanism and
1280
Book Review / Animal Behaviour 84 (2012) 1279–1280
development more completely into their teaching. This book can be an aid towards that end. Ryan & Wilczynski acknowledge that their book is not intended to be an exhaustive review of animal behaviour, as is found in the textbooks commonly used in animal behaviour courses. Topics that receive little if any treatment include personality and behavioural syndromes, and (perhaps surprisingly) intersexual conflict. Moreover, the effort at integration may sometimes trade off against adequate detail. One obstacle faced in using this book in a general animal behaviour course is a recurrent tendency for authors to allude to methods or topics without providing enough depth or citations for an inquisitive student to gain a full understanding. For instance, commenting on methods for phylogenetic reconstruction, authors write, ‘Parsimony is a nonparametric method that does not perform optimally under certain conditions, and many phylogeneticists rely on parametric techniques such as maximum likelihood or Bayesian inference’ (page 30). No further explanation is provided, no literature is cited within the text, and, sadly, the book does not have a glossary. Hence, substantial familiarity about statistical methods is presumed here. In my classes, the terms parametric techniques and nonparametric method would have significant name recognition mainly to my upperclassmen, and the terms maximum likelihood and Bayesian inference would probably be familiar to just a few of those. The same would be true of the definition of heritability (page 19), which includes the term additive genetic variation without defining what that is. Most likely, authors sacrificed such detail so as to keep the book of reasonable length; nevertheless, the brevity of explanation could frustrate students, especially motivated students who lack the necessary preparation. Textbooks tend to reflect the interests and expertise of their authors. In terms of coverage and proficiency in material, for example, the book is especially strong in the area of sensory systems; this strength is most palpable in sections on communication, navigation, and mate choice, which are superb. By comparison, treatment of cognitive processes, including learning, is less refined and not as in depth (the lack of depth perhaps exaggerated because material on learning is dispersed throughout the text). Not surprisingly, given Ryan & Wilczynski’s research interests, the chapter entitled ‘Species recognition, mate choice, and sexual selection’ far and away provides the most engaging and compelling application of the integrative approach. The book is predictably heavy in vertebrate case studies; that said, information on invertebrates, especially insects, is thoughtfully selected and expertly treated. In aesthetic terms, this compact book, a tad smaller in area than an iPad and about twice as thick, is a success. Figures are ample, generally simple in composition, and easy on the eyes. Figure legends are comprehensive and clearly written, albeit sometimes so extensive in length that perhaps the ‘box’ approach to
presentation might have been substituted for the figure and legend. Hard as I tried, I could not find typographical errors or errors in content. The writing is certainly more technical than in texts that focus less on mechanism, and is generally free of anecdotes and personal rejoinders. As such, the book does not have the high ‘readability factor’ that Alcock’s (2009) text, for example, possesses. While students may find the presentation drier than that text, the prose overall is nevertheless clear and straightforward enough for the motivated student to follow. The lack of exhaustive coverage, glossary, and teaching aids such as discussion questions may render this book less appealing to the instructor of large undergraduate classes in animal behaviour. As a text, the book seems most appropriate for a small class in animal behaviour composed of advanced undergraduates and first-year graduate students, in which students have ample opportunity to query the instructor for insights directly related to book content. For an instructor in animal behaviour who selects a more conventional textbook for their course, this volume would still be very useful as a reference. The examples are particularly well chosen and explanations are first-rate. A useful and welcome feature as a reference book is inclusion of material not found in many behaviour texts, for example on the evolution of sex, sex determination mechanisms, and meiotic drive as an example of selection at the level of the gene. In closing, I applaud the authors for taking a fundamentally different direction in their textbook by integrating multiple levels of analysis thoroughly in individual chapters. This book may not immediately steal the market away from various established texts in the area of animal behaviour, but it could nevertheless have an important impact on how animal behaviour is taught in the future. Daniel R. Papaj Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, U.S.A.
References Alcock, J. 2003. A textbook history of animal behaviour. Animal Behaviour, 65, 3–10. Alcock, J. 2009. Animal Behaviour: an Evolutionary Approach. 9th edn. Sunderland, Massachusetts: Sinauer. Chittka, L. & Döring, T. F. 2007. Are autumn foliage colors red signals to aphids? PLoS Biology, 5, e187. Döring, T. F. & Chittka, L. 2007. Visual ecology of aphids: a critical review on the role of colours in host finding. Arthropod–Plant Interactions, 1, 3–16. Geffeney, S. L., Fujimoto, E., Brodie, E. D., III, Brodie, E. D., Jr. & Ruben, P. C. 2005. Evolutionary diversification of TTX-resistant sodium channels in a predator– prey interaction. Nature, 434, 759–763. Hamilton, W. D. & Brown, S. P. 2001. Autumn tree colours as a handicap signal. Proceedings of the Royal Society B, 268, 1489–1493. MacArthur, R. H. & Pianka, E. R. 1966. On optimal use of a patchy environment. American Naturalist, 100, 603–609. Tinbergen, N. 1963. On aims and methods of ethology. Zeitschrift für Tierpsychologie, 20, 410–433.