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Animal Signals and

Volume 5

Series editors Vincent M. Janik School of Biology University of St Andrews Fife, UK

Peter McGregor Centre for Applied Zoology Cornwall College Newquay, UK More information about this series at http://www.springer.com/series/8824 Mark A. Bee • Cory T. Miller Editors

Psychological Mechanisms in Communication Editors Mark A. Bee Cory T. Miller Department of Ecology, , Department of Psychology, Neurosciences and Behavior Graduate Program University of Minnesota - Twin Cities University of California Saint Paul, Minnesota La Jolla, California USA USA

ISSN 2197-7305 ISSN 2197-7313 (electronic) Animal Signals and Communication ISBN 978-3-319-48688-8 ISBN 978-3-319-48690-1 (eBook) DOI 10.1007/978-3-319-48690-1

Library of Congress Control Number: 2017931073

© Springer International Publishing AG 2016 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic , computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made.

Printed on acid-free paper

This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Preface

Each of the two editors of this volume on Psychological Mechanisms in Animal Communication often sees the world of animal signals and communication very differently from the other editor. This is because each of us tends to view these topics through the eyes of our respective study organisms and the lenses of our past training. One of us (MAB), with a background in , , and , primarily studies acoustic communication in . The other (CTM), with a background in evolutionary biology, cognitive psychol- ogy, and systems neuroscience, studies acoustic communication in monkeys. A number of years ago, however, at the First International Conference on Acoustic Communication by , held at the University of Maryland in July 2003, we discovered that we share in common a deeply held opinion about research on animal communication: psychological mechanisms for signaling and receiving often get eclipsed by a fascination with signals. Animal behaviorists, the media, and the public are frequently captivated by the elaborate and showy signals animals produce to communicate with each other. How can one fail to marvel at the music-like vocalizations of and humpback whales, the cacophony generated by choruses of calling frogs and insects, the dynamic and mesmerizing visual displays of a cuttlefish, the use of chemical signals by foraging , or the multisensory displays of peacock and of paradise? Likewise, biologists and laypersons alike are often amazed by the diver- sity of functions that communication signals serve in the lives of animals, from courting and choosing mates to defending resources, strengthening social bonds, warning of danger, and begging for food or leading others to it. By comparison, the no-less elaborate and complex psychological mechanisms that have evolved for the purposes of signaling and receiving seem to inspire less wonder. One notable exception, of course, is the hunt for parallels and precursors of in other animals, which has been extensive and is ongoing. But many of the sensory, perceptual, and cognitive mechanisms that enable human communication are not language specific and can be studied in many other communicating animals. We

v vi Preface hope this volume will stimulate wonderment regarding the diversity of psycholog- ical mechanisms for signaling and receiving found across a diversity of species. After first crossing paths as graduate students at that 2003 conference, we set for ourselves the task of putting to pen our shared view on the state of research on psychological mechanisms in animal communication. It was only after several false starts, and after navigating the academic job market, that we finally accomplished our task in the form of an essay published in Animal Behaviour (Miller and Bee 2012). In that paper, we questioned whether the receiver psychology paradigm (Guilford and Dawkins 1991, 1993; Rowe 2013), which has been influential in shaping the field’s thinking about the evolution of signal design and signaling behavior, has led to any useful, new insights into the psychology of receivers. Our general conclusion was that it had not. We argued that the receiver psychology paradigm could have a much greater impact on the study of animal signals and communication if it also promoted research on the actual psychological mecha- nisms involved in communication and their potential evolutionary diversity. Spe- cifically, we advocated for seeking a broader and deeper understanding of receivers’ “psychological landscapes” through the comparative study of the sen- sory, perceptual, and cognitive mechanisms that enable receivers to acquire, pro- cess, and act upon signals. This volume represents an extension of the basic thesis of our 2012 essay: understanding animal signals and communication depends fundamentally on understanding the psychological mechanisms operating in sig- nalers and receivers. In putting this book together, we have had the fortunate pleasure to engage as contributing authors a number of our colleagues whose research programs on animal communication have focused on various aspects of sensation, perception, and cognition. Collectively, their chapters span an impressive range of topics, taxa, and tools. We are deeply grateful to each author for their contribution to this volume and, more broadly, for their contribution to making new discoveries about signaler and receiver psychology. Their research inspires our own. We are also grateful to the series editors, Vincent Janik and Peter McGregor, for inviting us to edit this volume and for their input and guidance both on the large, big-picture issues involved with putting together an edited volume and on the small, mundane issues that are an inevitable part of doing so. We thank Andrea Schlitzberger, from Springer, for her help and dedication to coordinating all aspects of this project. The National Institutes of Health and the National Science Foundation have generously supported our research on animal communication over the years. Finally, we thank our families—Meggan, Karis, Abigail, River, and Sierra—for their enduring support.

Mark A. Bee St. Paul, MN, USA Cory T. Miller La Jolla, CA, USA Preface vii

References

Guilford T, Dawkins MS (1991) Receiver psychology and the evolution of animal signals. Anim Behav 42:1–14 Guilford T, Dawkins MS (1993) Receiver psychology and the design of animal signals. Trends Neurosci 16(11):430–436 Miller CT, Bee MA (2012) Receiver psychology turns 20: is it time for a broader approach? Anim Behav 83(2):331–343 Rowe C (2013) Receiver psychology: a receiver’s perspective. Anim Behav 85(3):517–523 Contents

1 Signaler and Receiver Psychology ...... 1 Mark A. Bee and Cory T. Miller 2 Avian Auditory Processing at Four Different Scales: Variation Among Species, Seasons, Sexes, and Individuals ...... 17 Kenneth S. Henry, Megan D. Gall, Alejandro Ve´lez, and Jeffrey R. Lucas 3 Perceptual and Neural Mechanisms of Auditory Scene Analysis in the European Starling ...... 57 Georg M. Klump 4 Mate Searching Animals as Model Systems for Understanding Perceptual Grouping ...... 89 Hamilton E. Farris and Ryan C. Taylor 5 Why Complex Signals Matter, Sometimes ...... 119 Tricia L. Rubi and David W. Stephens 6 Communication Through a Window of Error: Proportional Processing and Signal Categorization ...... 137 Karin L. Akre and Sonke€ Johnsen 7 Social Recognition in Anurans ...... 169 Mark A. Bee 8 Referents and Semantics in Animal Vocalizations ...... 223 Marta B. Manser 9 Social Concepts and Communication in Nonhuman ..... 251 Klaus Zuberbühler

ix x Contents

10 Decisions to Communicate in Ecological and Social Landscapes ...... 271 Camille R. Toarmino, Vladimir Jovanovic, and Cory T. Miller 11 Overcoming Sensory Uncertainty: Factors Affecting Foraging Decisions in -Eating ...... 285 Rachel A. Page and Patricia L. Jones

Index ...... 313