Nobel laureate Niko Tinbergen laid the foundations for the scientific study of animal behaviour with his work on causation, development, function and evolution. In this book, an international cast of leading animal biologists reflect on the enduring significance of Tinbergen's groundbreaking proposals for modern behavioural biology. It includes a reprint of Tinbergen's original article on the famous 'four whys' and a contemporary introduction, after which each of the four questions are discussed in the light of contemporary evidence. There is also a discussion of the wider significance of recent trends in evolutionary psychology and neuroecology to integrate the 'four whys'. With a foreword by one of Tinbergen's most prominent pupils, Aubrey Manning, this wide-ranging book demonstrates that Tinbergen's views on animal behaviour are crucial for modern behavioural biology. It will appeal to graduate students and researchers in animal behaviour, behavioural ecology and evolutionary biology.

About the Authors

Simon Verhulst is assistant professor of Behavioural Biology at the University of Groningen, The Netherlands. His research interests have included ecological immunology, avian life history evolution and avian energetics, all with the aim to study the interplay between function and mechanism of behavior. In 2004 he was elected as member of 'The Young Academy', the junior section of the Dutch Royal Society. He has been an Associate Editor for the Journal of Animal Ecology since 2007.

Johan J. Bolhuis is professor of Behavioural Biology at the University of Utrecht, The Netherlands. His main research interests are in the behavioral, cognitive and neural mechanisms of learning, memory and development, in particular song learning in birds. He has been an editor of the leading journal Animal Behaviour and President of the Royal Dutch Zoological Society.

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