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Nature's Music: The Science of Birdsong
Nature's Music: The Science of Birdsong
Nature's Music: The Science of Birdsong
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Nature's Music: The Science of Birdsong

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The voices of birds have always been a source of fascination. Nature’s Music brings together some of the world’s experts on birdsong, to review the advances that have taken place in our understanding of how and why birds sing, what their songs and calls mean, and how they have evolved. All contributors have strived to speak, not only to fellow experts, but also to the general reader. The result is a book of readable science, richly illustrated with recordings and pictures of the sounds of birds. Bird song is much more than just one behaviour of a single, particular group of organisms. It is a model for the study of a wide variety of animal behaviour systems, ecological, evolutionary and neurobiological. Bird song sits at the intersection of breeding, social and cognitive behaviour and ecology. As such interest in this book will extend far beyond the purely ornithological - to behavioural ecologists psychologists and neurobiologists of all kinds.

* The scoop on local dialects in birdsong* How birdsongs are used for fighting and flirting* The writers are all international authorities on their subject
LanguageEnglish
Release dateOct 5, 2004
ISBN9780080473550
Nature's Music: The Science of Birdsong

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    Nature's Music - Peter R. Marler

    Table of Contents

    Cover image

    Title page

    Copyright

    Preface: the Editors

    List of contributors

    Foreword: A tribute to the late Luis Felipe Baptista

    Acknowledgements

    Chapter 1: Science and birdsong: the good old days

    INTRODUCTION

    IN THE BEGINNING

    SONG STUDIES IN MID-CENTURY

    DIALECTS

    THE MYSTERY OF SPECIES UNIVERSALS

    SUBSONG: THE START OF A LONG JOURNEY

    INNATENESS AND OTHER TABOOS

    INNATE SONGS, LEARNED SONGS?

    THE VERSATILITY OF BIRDSONG

    CONCLUSIONS

    Chapter 2: Vocal fighting and flirting: the functions of birdsong

    INTRODUCTION

    EVIDENCE THAT SONG IS IMPORTANT

    MALE–MALE COMPETITION: VOCAL FIGHTING

    MATE CHOICE: VOCAL FLIRTING

    ONE SINGER: MULTIPLE FUNCTIONS

    FEMALE SONG AND DUETTING

    CONCLUSIONS

    Chapter 3: Learning to sing

    INTRODUCTION

    CHALLENGES IN A YOUNG BIRD’S WORLD: HOW TO PROCEED FROM HEARING TO MEMORIZING

    PROPERTIES OF SENSORIMOTOR LEARNING: HOW TO ACHIEVE VOCAL EXPERTISE

    FUNCTIONAL ASPECTS OF SONG LEARNING: HOW TO DEVELOP AN APPROPRIATE REPERTOIRE

    COMPARATIVE ASPECTS OF LEARNING TO SING

    CONCLUSIONS

    Chapter 4: The diversity and plasticity of birdsong

    INTRODUCTION

    TO LEARN OR NOT

    DIALECTS, AND OTHER FORMS OF GEOGRAPHIC VARIATION

    WHEN, WHERE, AND FROM WHOM TO LEARN

    REPERTOIRE SIZE

    MIMICRY

    CONCLUSIONS

    Chapter 5: Bird calls: a cornucopia for communication

    INTRODUCTION

    HOW MANY CALLS DOES A BIRD HAVE?

    ALARM CALLS: THE HAZARDS OF PREDATION

    CONTACT AND SEPARATION CALLS: KEEPING IN TOUCH

    ROOSTING CALLS: A PLACE FOR THE NIGHT

    CALLING TO FOOD

    BEGGING CALLS: ASKING FOR FOOD

    AGONISTIC CALLS: SIGNALS FOR AGGRESSION

    CALLING ACROSS SPECIES BOUNDARIES

    VOCAL SEMANTICS: WHAT DO BIRD CALLS MEAN?

    CONCLUSIONS

    Chapter 6: Singing in the wild: the ecology of birdsong

    INTRODUCTION

    SOUND TRANSMISSION: GETTING THE MESSAGE ACROSS

    COMMUNICATION WITH DEGRADATION: INSIGHTS FROM PLAYBACKS

    THE HABITAT AS A SPACE FOR ACOUSTIC PERFORMANCE

    NOISE ANNOYS: COMPETITION FOR ACOUSTIC SIGNAL SPACE

    ENVIRONMENTAL SELECTION AND SIGNAL EVOLUTION

    CONCLUSIONS

    Chapter 7: Audition: can birds hear everything they sing?

    INTRODUCTION

    AUDITORY DETECTION

    AUDITORY DISCRIMINATION

    PERCEPTION OF VOCAL SIGNALS

    CONCLUSIONS

    Chapter 8: Brains and birdsong

    INTRODUCTION

    GENERAL BRAIN ORGANIZATION

    THE SEARCH FOR VOCAL LEARNING BRAIN AREAS

    THE SONGBIRD VOCAL COMMUNICATION BRAIN NETWORK

    FUNCTIONS OF VOCAL PATHWAYS AND NUCLEI

    THE BRAIN AND AUDITION

    SENSITIVE PERIODS

    SPECIES AND SEXUAL DIFFERENCES

    ADULT NEUROGENESIS DISCOVERED

    EVOLUTION OF VOCAL LEARNING: SONGBIRDS, HUMMINGBIRDS, AND PARROTS

    PARALLELS BETWEEN BIRDS AND HUMANS?

    CONCLUSIONS

    Chapter 9: How birds sing and why it matters

    INTRODUCTION

    AN ORGAN FOR SINGING

    CARDINAL SONG: A CASE STUDY

    LEARNING TO SING

    PUSHING THE ACOUSTIC ENVELOPE

    PERFORMANCE CONSTRAINTS: INSIGHTS FROM A VOCAL MIMIC

    VOCAL GYMNASTICS: ARE FEMALES IMPRESSED?

    CONCLUSIONS

    Chapter 10: Birdsong and evolution

    INTRODUCTION

    EVOLUTION AND SONG: A TWO-WAY INTERACTION

    RECONSTRUCTING VOCAL EVOLUTION IN STREPTOPELIA DOVES

    HOW ABOUT OTHER BIRDS?

    CONCLUSIONS

    Chapter 11: Performance limits on birdsong

    INTRODUCTION

    MECHANISMS OF VOCAL PRODUCTION

    VOCAL MECHANICS AND EVOLUTION: CONSTRAINT AND OPPORTUNITY

    SONGS OF DARWIN’S FINCHES

    CONCLUSIONS

    Chapter 12: Birdsong and conservation

    INTRODUCTION

    LEVELS AND ISSUES IN CONSERVATION

    APPLICATIONS OF BIRD COMMUNICATION TO CONSERVATION

    INSTRUMENTS AND METHODS

    CONCLUSIONS

    Chapter 13: Grey parrots: learning and using speech

    INTRODUCTION

    COGNITIVE AND COMMUNICATIVE ABILITIES

    TRAINING PARROTS TO LEARN AND USE REFERENTIAL SPEECH

    CONCLUSIONS

    Chapter 14: Singing, socializing, and the music effect

    INTRODUCTION

    SONG, GESTURE, AND COWBIRD RELATIONSHIPS

    SONG, MUSIC, AND HUMAN DEVELOPMENT

    BIRDSONG AND MUSIC: CROSS-SPECIES INSPIRATIONS

    CONCLUSIONS

    Bibliography

    Glossary

    Species list

    Index

    Introduction to the CD’s

    Color Plates

    Copyright

    This book is printed on acid-free paper

    Copyright © 2004, Elsevier (USA). All rights reserved

    No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means electronic, mechanical, photocopying, recording or otherwise, without the prior written permission of the publisher

    Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (+44) 1865 843830, fax: (+44) 1865 853333, e-mail: permissions@elsevier.co.uk. You may also complete your request on-line via the Elsevier homepage (http://www.elsevier.com), by selecting ‘Customer Support’ and then ‘Obtaining Permissions’

    Elsevier Academic Press

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    http://www.elsevier.com

    Elsevier Academic Press

    84 Theobald’s Road, London WC1X 8RR, UK

    http://www.elsevier.com.

    Library of Congress Catalog Number: 2004108979

    British Library Cataloguing in Publication Data

    A catalogue record for this book is available from the British Library

    ISBN 0 12 473070 1

    Typeset by Replica Press, India

    Printed and bound in Italy

    04 05 06 07 08 09 9 8 7 6 5 4 3 2 1

    Preface: the Editors

    PETER. MARLER, HANS. SLABBEKOORN

    On a sad day in June 2000 Luis Felipe Baptista died unexpectedly near his home in San Francisco, California, at the age of 59. The science of birdsong lost one of its most erudite and charismatic authorities. In a prologue to this book, Baptista’s long time friend and colleague Robert Bowman contributes a brief overview of some of his most illustrious accomplishments. Within months of his death still grieving colleagues decided to celebrate his remarkable career with a symposium, and ultimately with a book, on the remarkable progress that has been made by him and many other scientists around the globe in the study of birdsong.

    In an enterprise spanning more than three years, the debts the editors owe to others are many. Among his many responsibilities Luis had served as Curator of Ornithology and Mammalogy at The California Academy of Sciences. The staff members of the Academy were unstinting in their support of the Baptista symposium, especially Terry Gosliner, Nina Jablonski, and Doug Bell. Sylvia Hope played a key role at many stages, including the planning and execution of the book, and Kathleen Berge was most helpful in organizing and conducting the symposium. A cadre of distinguished experts assembled on November 3 2001 for the daylong meeting. Although the speakers had been encouraged to present up-to-the-minute overviews of their area of expertise, no one was quite prepared for the extent and the richness of the new information presented. It covered all aspects of research on the vocal behavior of birds, never brought together before in the same forum. This book was the next step, after supplementing the list of authors to complete the coverage.

    In an era when everyone specializes, it is all too easy for people working in one corner of the science of birdsong, say the neurobiology of vocal learning, to get out of touch with progress in other aspects of their discipline, such as the impact of the environment on the evolution of birdsong. We hope that this book will help to familiarize everyone working in the field with the remarkable progress that their colleagues have been making. Above all we hope that this book will appeal to general readers, especially to bird watchers and others who already appreciate nature’s music. Luis Baptista was renowned for his skill in communicating to others, both his love of birdsong, and the excitement of scientific ornithology. In that same spirit we have strived to ensure that, wherever possible jargon has been held in check, while still maintaining the highest scientific standards, in both text and figures. The one-page ’boxes’ spaced at intervals throughout the text, written specifically at the invitation of the editors, provide extra detail on some topics, and further explication of technical issues. We have solicited sound recordings of songs and other avian vocal signals wherever these could be found and assembled them in the two compact discs at the back of this book

    In our editorializing it was a pleasure to interact with the sixteen biologists who joined us in writing chapters for the book. Without exception they were cooperative in dealing with our questions and suggestions, and sympathetic to our desire, wherever possible, to strike a balance between technical content and readable prose, an endeavor in which Gregory Ball and William Searcy gave some valuable help. At all stages, from preparing, editing and checking the manuscripts, organizing and formatting figures and color plates and with Judith Marler, scrutinizing proofs, Rebecca Wylie was a tower of strength. We owe special thanks to the 44 authors who made extra efforts with the boxes, creating new text and figures for the purpose and, together with the chapter authors, helping to assemble and donate sound recordings for the compact discs. Tackling the daunting task of putting the C.D.’s together would have been unthinkable without the generous help of Jack Bradbury and his staff at the Macaulay Library of Natural Sounds at the Laboratory of Ornithology, Cornell University, especially Steve Pantle, Bob Grotke and Mary Guthrie. For the color plates that Elsevier/Academic Press encouraged us to include, we are pleased to display the work of photographers who specialize in creating images of birds caught in the very act of vocalizing. They add a further dimension to our efforts to convey something of the saga of Nature’s Music.

    List of contributors

    Bridget Appleby

    BBC Natural History Unit, Bristol, England, UK.

    Arthur P. Arnold

    University of California, Los Angeles, USA

    Myron C. Baker

    Colorado State University, Fort Collins, USA

    Gregory F. Ball

    John Hopkins University, Baltimore, Maryland, USA

    Robert Bowman

    California Academy of Sciences, San Francisco, USA

    Deborah Buitron

    North Dakota State University, Fargo, USA

    Clive K. Catchpole

    University of London, Egham, England, UK

    MarthaLeah Chaiken

    Hofstra University, Hempstead, New York, USA

    Claude Chappuis

    Museum of Natural History, Paris, France

    Sarah Collins

    University of Nottingham, England, UK

    Jeffrey Cynx

    Vassar College, Poughkeepsie, New York, USA

    Robert Dooling

    University of Maryland, College Park, USA

    Françoise Dowsett-Lemaire

    Ganges, France

    Chris Evans

    MacQuarie University, Sydney, Australia

    Adriana R. Ferreira

    Duke University, Durham, North Carolina, USA

    W. Tecumseh Fitch

    University of St. Andrews, Scotland, UK

    Sandra L.L. Gaunt

    Ohio State University, Columbus, USA

    Jean-Pierre Gautier

    Universite de Rennes, Paimpont, France

    Michael H. Goldstein

    Franklin & Marshall College, Lancaster, Pennsylvania, USA

    Franz Goller

    University of Utah, Salt Lake City, USA

    Thomas Hahn

    University of California, Davis, USA

    Paul Handford

    University of Western Ontario, London, Canada

    Henrike Hultsch

    Freie Universität, Berlin, Germany

    Darren E. Irwin

    University of California, Los Angeles, USA

    Jessica H. Irwin

    University of California, Los Angeles, USA

    Erich D. Jarvis

    Duke University, Durham, North Carolina, USA

    Andrew P. King

    Indiana University, Bloomington, USA

    Kohta I. Kobayasi

    Chiba University, Chiba-city, Japan

    Michel Kreutzer

    Université Paris X-Nanterre, France

    Donald Kroodsma

    University of Massachusetts, Amherst, USA

    Anthony Leonardo

    Harvard University, Cambridge, Massachusetts, USA

    Daniel Margoliash

    University of Chicago, Illinois, USA

    Peter Marler

    University of California, Davis, USA

    Archibald McCallum

    Applied Bioacoustics, Eugene, Oregon, USA

    Douglas A. Nelson

    Ohio State University, Columbus, USA

    Stephen Nowicki

    Duke University, Durham, North Carolina, USA

    Gary L. Nuechterlein

    North Dakota State University, Fargo, USA

    Kazuo Okanoya

    Chiba University, Chiba-city, Japan

    Robert B. Payne

    University of Michigan, Ann Arbor, USA

    Irene M. Pepperberg

    Massachusetts Institute of Technology, Cambridge, USA

    Jeffrey Podos

    University of Massachusetts, Amherst, USA

    Marilyn Ramenofsky

    University of Washington, Seattle, USA

    Steve Redpath

    Center for Ecology and Hydrology, Banchory, Scotland, UK

    Katharina Riebel

    Leiden University, The Netherlands

    William A. Searcy

    University of Miami, Coral Gables, Florida, USA

    Hans Slabbekoorn

    Leiden University, The Netherlands

    Jill Soha

    Ohio State University, Columbus, USA

    Roderick A. Suthers

    Indiana University, Bloomington, USA

    Carel ten Cate

    Leiden University, The Netherlands

    Dietmar Todt

    Freie Universität, Berlin, Germany

    Mari A. Tokuda

    Tokyo Institute of Technology, Japan

    Meredith J. West

    Indiana University, Bloomington, USA

    Jay Withgott

    Portland, Oregon, USA

    Ayako Yamaguchi

    Boston University, Massachusetts, USA

    Richard A. Zann

    La Trobe University, Melbourne, Australia

    Foreword: A tribute to the late Luis Felipe Baptista

    ROBERT I. BOWMAN

    This volume is a celebration of the life of Luis Felipe Baptista, and it is appropriate and timely to review some of the highlights of Dr Baptista’s scientific career. Tales of his productivity are legendary, and many of his achievements were widely publicized in tributes by his friends and professional associates after his death, in speeches, by radio and television, in scientific journals, in newspapers and magazines. I knew Luis as a friend and colleague since his graduate student days at the University of California, Berkeley, commencing in the fall of 1967. He had just completed his master’s degree studies at the University of San Francisco under the tutelage of the renowned ornithologist Dr Robert T. Orr, who was at that time also the Curator of Ornithology and Mammalogy at the California Academy of Sciences. It was through this first close relationship with a professional biologist that Luis was amazed to learn that some people actually made a living doing what he had been doing for pleasure since boyhood – watching birds!

    As a youth raised by Portuguese-Chinese parents in Hong Kong and Macau, it was painless for him to learn to speak Portuguese, English, and Chinese. Later he picked up Spanish and German with little difficulty. He also became very familiar with the habits and voices of native birds, as well as the widely prevalent Chinese art of aviculture. Early in his youth he realized that some birds learned their songs by imitation, a major focus of his later research. Indeed his graduate research on the song of the white-crowned sparrow – an abundant bird on the Berkeley campus and at other San Francisco Bay Area localities – provided him with a plethora of thematic vocal variations. He later broadened his documentation of variations in song from California to British Columbia. He demonstrated that boundaries between dialects are not always clear-cut, and that birds living at the interface may be bilingual.

    A great source of amusement and amazement for those who were privileged to hear Luis report on his white-crowned sparrow songs were his precise imitations of their dialects. He had a remarkable memory and an acute sensitivity to the many nuances of pitch and timing that individual birds displayed.

    After the completion of his doctoral dissertation at Berkeley, Luis received a grant from NATO to visit Germany where he spent more than a year at the Max Planck Institüt für Verhaltensphysiologie. While stationed at Vögelwarte Radolfzell, he recorded the learned ’rain call’ of the chaffinch around Lake Constance, trying to define dialect boundaries. The project was later expanded to include a transect across Germany and resulted in a remarkable documentation of geographical dialects in a bird call.

    Upon his return to the United States in 1973 Luis assumed an academic appointment at Occidental College in Los Angeles where he became Professor of Biology and Curator of the Moore Laboratory collection of birdsongs. In 1980, Luis was appointed Chairman and Associate Curator of Ornithology and Mammalogy at the California Academy of Sciences in San Francisco, where he remained until his untimely death on June 10, 2000.

    Without enumerating details of his more than 120 scientific publications, the majority of which concern avian vocalizations, Luis’ broad and detailed knowledge of birds of the world is evident in some of the other topics he wrote about, often with co-authors, including cowbird parasitism; leaf bathing in emberizine sparrows; the use of courtship displays in taxonomy; behavioral interactions within and among animal species; handedness and its possible taxonomic significance in grassquits; taxonomic revision of the Mexican Piculus woodpecker complex; hybridization in Calypte hummingbirds; behavior and taxonomic status of Grayson’s dove; photoperiodically induced ovarian growth in white-crowned sparrows; testosterone, aggression, and dominance in Gambel’s white-crowned sparrows; the origin of Darwin’s finches; the fine structural basis of the cuteaneous water barrier in nesting zebra finches; production and control of birdsong; the behavior, status and relationships of the endemic St Lucia black finch; inheritance and loss of the straw display in estrildid finches; field observations of some New Guinea mannikins; the role of song in the evolution of passerine diversity; seasonal changes in song behavior and song nuclei in the brain of Gambel’s white-crowned sparrow; nature and its nurturing in avian vocal development; the program of reproduction and reintroduction of the dove on Socorro Island; bioacoustics as a tool in conservation studies; guidelines for the use of wild birds in research; relationships of some mannikins and waxbills in the Estrildidae; linguistics and anthropological study of Chinese birds; cognitive processes in avian vocal development; what the white-crowned sparrow can teach us about language.

    Luis’ breadth of knowledge of birds of the world qualified him superbly for the task of revising one of the most famous modern ornithology texts, The Life of Birds by J.C. Welty. At the time of his death, Luis was involved with a National Academy of Sciences project on the biology of music and its relevance to birdsong.

    Much of Luis’ avian research was conducted and co-authored cooperatively with colleagues and junior scientists. He shared his knowledge of birds generously, with students and the general public alike. He was a great communicator. He possessed a brilliant mind, an encyclopedic memory of facts and faces, and a warmth of personality that charmed all who visited his quarters at the California Academy of Sciences. His presence served as a magnet for visiting scientists and the lay public.

    Through the years Luis’ presence greatly enhanced the scientific stature of the Academy through his public lectures, foreign travel on research projects, his prolific publication record, his generous coaching of graduate students, and his concern for the survival of endangered bird species. The latter brought him close to the problem of environmental degradation and the sorrowful fate of the endemic dove of Socorro Island off Baja California. Fortunately, a few surviving individuals were to be found in the collections of mainland aviculturists. Working with Mexican biologists, Luis initiated an intensive breeding program in the hope of reintroducing the species to its ancestral homeland, once the island environment was rehabilitated. The good news is that plans were launched in collaboration now under way with Mexican authorities to reintroduce the Socorro dove to its ancestral home. The year 2003 is significant because it commemorates the 100th anniversary of the first expedition to the Revillagigedo Archipelago by the California Academy of Sciences. The sad news is that Luis will never see the fruits of his conservation efforts. But he took pride in his award from the Rolex Corporation for initiating this pioneering enterprise in the field of conservation.

    The departure of Luis Baptista from the halls of science at universities, and at the California Academy of Sciences is deeply felt by his many friends and admirers around the world. This symposium volume is a deserving tribute to the legacy of his creative spirit, his voracious curiosity, and his many insights into the minds of birds. His unabashed love for scientific investigation was nourished by his faithful and supportive staff, and especially by his beloved long-term companion, Helen Horblit. I know that he would be deeply pleased and appreciative of the dramatic progress in many aspects of the science of birdsong reviewed in this volume.

    Acknowledgements

    CHAPTER 1

    Peter Marler: I dedicate this brief historical review to my late mentor, William Homan Thorpe, the pioneer who launched the new discipline. For help in preparing the chapter I am grateful to Hans Slabbekoorn, my co-editor. In preparing figures and obtaining sound recordings, I was aided generously by Allan Baker, Jack Bradbury, Doug Nelson, Jürgen Nicolai, Steve Nowicki, Peter Slater, Bill Searcy, Jill Soha, and Rebecca Wylie. Thanks are due to the Macaulay Library of Natural Sounds at Cornell University for allowing use of sound recordings from their CD on the Diversity of Animal Sounds. For research assistance over the years I am grateful to Miwako Tamura, Mary Sue Waser, Roberta Pickert, Virginia Sherman, and especially to Susan Peters. Don Kroodsma gave generous access to material on the swamp sparrows of the Great Vly. Research was supported by the National Science Foundation and the National Institute of Mental Health.

    CHAPTER 2

    Sarah Collins: Thanks for comments on earlier versions of the chapter to Bill Searcy, Hans Slabbekoorn, and Peter Marler and to Selvino de Kort for discussions on the nature of birdsong. Thanks also to the students of the University of Nottingham on the Behavioural Ecology Field Course for their enthusiasm in recording birdsong and conducting playback experiments. My work was supported by a grant from the Nuffield Foundation.

    CHAPTER 3

    Henrike Hultsch & Dietmar Todt: We thank the editors for their help in improving the quality of this chapter, and Nicole Geberzahn for preparing most of the figures and the sound samples. The studies on nightingales reported here were supported by a fund of the German Science Foundation DFG (Az: To 13/30-1).

    CHAPTER 4

    Donald Kroodsma: I’m grateful that I’ve spent much of my adult life thinking about the diversity of bird sounds, and have key people to thank for those opportunities: my parents, Sewall Pettingill, John Wiens, Peter Marler, and my wife Melissa, who has tolerated and encouraged all of this frivolity. The National Science Foundation has provided substantial support over the years. Thanks also to Peter Marler and Hans Slabbekoorn, two gracious and patient editors, and to Luis Baptista, with whom I shared more than a quarter century of birdsong adventures.

    CHAPTER 5

    Peter Marler: I thank Sylvia Hope for much help in preparing and commenting on this review. Many people contributed generously to the background research, including Mike Baker, Mike Beecher, Hans-Heiner Bergmann, Jack Bradbury, Thomas Bugnyar, Nick Davies, Chris Evans, Bruce Falls, Millicent Ficken, Tom Hahn, Bill Hamilton, Berndt Heinrich, Arla Hile, Georg Klump, Walt Koenig, Indrikis Krams, Paul Mundinger, Marilyn Ramenofsky, Steve Rothstein, Hans Slabbekoorn, Georg Striedter, and Sandy Vehrencamp. I am grateful to Don Kroodsma and the Laboratory of Ornithology at Cornell University for permission to reproduce figures and sound recordings from the chapter on vocal behavior in their Home Study Course in Bird Biology.

    CHAPTER 6

    Hans Slabbekoorn: I thank Tom Smith of the Center for Tropical Research for moral and financial support during my studies in Cameroon, California and Colorado. Luis Baptista was a great pleasure to meet and I am grateful to him and the California Academy of Sciences for allowing me to use their acoustic laboratory. Jacintha Ellers, Peter Marler, and Katharina Riebel made helpful comments that improved my chapter. Many thanks to Thierry Aubin, Theo Beerenfenger, Frank Dorritie, Lang Elliott, Albertina Leitao, and Gary Ritchison for help in getting sound recordings. I thank Andrea Jesse for her work and pleasant collaboration on the white-crowned sparrows of the San Francisco Bay Area (Figure 1.6). R. Manin is the photographer of both pictures in Box 39. I thank Merijn de Bakker, John van Dort, Bart Houx, Caroline van Heijningen, Rob Lachlan, Helder Perreira, Martin Poot, Pien Verburg, and Paige Warren for suggestions and material about musicians inspired by birdsong. My work was supported by the Netherlands Organization for Scientific Research. NWO and the US National Science Foundation.

    CHAPTER 7

    Robert Dooling: I thank all of the students, postdoctoral fellows, and colleagues who have collaborated on the work contained herein over the years, especially Beth Brittan-Powell, Bernard Lohr, and Marjorie Leek who also read and commented on previous drafts of this manuscript. This work was supported by grants from the National Institute of Deafness and Communication Disorders.

    CHAPTER 8

    Erich D. Jarvis: For assistance with figures and/or text I thank: Art Arnold, Gregory Ball, Eliot Brenowitz, Catherine Carr, Timothy DeVoogd, Fred Gage, Adriana Ferreira, John Kirn, Lubica Kubikova, Dan Margoliash, Claudio Mello, Richard Mooney, Paul Nealen, David Perkel, Constance Scharff, Kazuhiro Wada, Martin Wild. For sound files I thank: Michael Brainard, Fernando Nottebohm, Kazuo Okanoya, Constance Scharff. In Figure 1: I am grateful to John W. Sundsten for permission to reproduce the human brain image from the Digital Anatomist Project, Dept. of Biological Structure, University of Washington at http://www9.biostr.washington.edu/da.html, and to John Kirn for illustration material.

    CHAPTER 9

    Roderick A. Suthers: I am deeply indebted to my students and colleagues whose names appear in the authorship of papers cited from my laboratory. Without their talents, enthusiasm and hard work many of these studies would not have been possible. I thank Masakazu Konishi for encouraging me to apply to songbirds the techniques developed for studying echolocating bats and birds. Special thanks are also due to Michel Kreutzer and Eric Vallet at the University of Paris for the invitation to collaborate in their research on canary song and to Carel ten Cate at Leiden University for the opportunity to participate in studies on dove vocalizations. The author’s research was supported by grants from the U.S. National Science Foundation and the National Institutes of Health.

    CHAPTER 10

    Carel ten Cate: I am grateful to Mechteld Ballintijn, Gabriël Beckers, Selvino de Kort and Hans Slabbekoorn, all of whom contributed significantly to the Streptopelia-project. They and other members of the Leiden Behavioural Biology group were excellent discussion partners for the ideas presented in the chapter. Selvino and Hans collected the recordings gathered on the CD. I also thank J. Jordan Price for providing the sonograms in Figure 10.1 as well as additional unpublished material. Gabriël, Selvino and, in particular, the editors of this book provided constructive and useful criticisms on earlier drafts.

    CHAPTER 11

    Jeffrey Podos & Stephen Nowicki: We thank the editors for the opportunity to contribute to this volume, and for their helpful suggestions on our chapter. Our laboratory groups at Duke and UMass Amherst have provided valuable feedback over the years on the ideas presented here. J.P.’s research in the Galapagos has been made possible through the kind support of the Charles Darwin Research Station and the Galapagos National Park Service. We are both grateful to the National Science Foundation for funding our research (IBN 0077891 and IBN 0347291 to JP, and IBN 9974743 and IBN 0315377to SN).

    CHAPTER 12

    Sandra L. L. Gaunt & D. Archibald McCallum: For discussion and comment on the manuscript for this chapter we thank Dr. Douglas A. Nelson, Director and Dr. Jill Soha, Curator of The Borror Laboratory of Bioacoustics, Museum of Biological Diversity, The Ohio State University. Tom Scott provided fruitful discussions on applying bioacoustic technology to monitoring.

    CHAPTER 13

    Irene V. Pepperberg: This chapter was written with support from the MIT School of Architecture and Planning and donors to The Alex Foundation. Research reported in this chapter was supported by donors to The Alex Foundation and several grants from the National Science Foundation.

    CHAPTER 14

    Meredith J. West, Andrew P. King, & Michael H. Goldstein: The authors’ work was supported by grants from the National Science Foundation and the National Institute of Health.

    Science and birdsong: the good old days

    PETER. MARLER

    INTRODUCTION

    The invention of a novel analytical technique often helps to launch a new science. What microscopes were for the emergence of cell biology as a discipline, or the cathode-ray oscilloscope for neurophysiology, it was the sound spectrograph that, immediately after the Second World War, enabled the birth of the science of birdsong. There had been no lack of interest in birdsong previously, and fascinating and important discoveries were made, especially about the functions of song. But never before had researchers come together to form a coherent discipline. Until about 1950, everyone interested in birdsong had no choice but to work by ear. Only when the sound spectrograph became available was it possible, for the first time, to grapple objectively with the daunting variability of birdsong, and to specify its structure precisely. Almost immediately a multitude of new issues became accessible for scientific scrutiny and experimentation.

    IN THE BEGINNING

    Visible Speech, Visible Birdsongs

    With remarkable rapidity sonograms became the standard method for birdsong study. Previously the only machine available for the empirical visualization of animal sounds was the same cathode-ray oscillograph that revolutionized neurophysiology. Oscillograms were useful for the study of insect sounds, but not very helpful for analyzing sounds with a more complex frequency structure, like birdsongs or, for that matter, human speech (Fig. 1.1). Around 1940, a group of researchers at the Bell Telephone Laboratories decided that it was time to develop methods for making the details of speech more visible and intelligible, in part because they thought it might help in teaching deaf people to learn to speak and use the telephone (Potter et al. 1947). Also it was wartime and, as with other acoustic technologies, another driving force was the need to monitor movements of ships and submarines, analyzing their far-traveling ocean-born sounds, each with its own particular signature. As a consequence, many of the details remained classified until the War was over. Soon after hostilities ended the Kay Electric Company was created as an offshoot of the Bell Laboratories with the assignment to build and market a machine for visible speech (Fig. 1.1). The success of the Vibralyser, as it was first called, was mixed. There were some dramatic achievements. Sound spectrograms of speech became a basic tool in linguistics, and remained so for years. As far as speech therapy was concerned, sonograms of speech proved to be difficult even for experts to read in real time, greatly limiting their value in working with the deaf. But they were perfect for the study of birdsongs.

    Figure 1.1 Some historical memorabilia. An oscillogram of Visible Speech (Potter et al. 1947) (A); followed by a much more informative spectrogram of the same words (B). The Vibralyser diagram by Borror comes from his 1953 paper with Reese (C). (D) represents a sonogram of an obviously two-voiced song of a wood thrush, from Borror and Reese (1956).

    The potential of the sound spectrograph for the study of birdsong was appreciated almost immediately. During the War ornithologist and entomologist Donald Borror worked in Naval intelligence and learned of the visible speech project. When he arrived back in Ohio State, he discovered a sound spectrograph sent there for a declassified project that did not work out. He was able to arrange access, and sonograms of songs of sparrows, thrushes, and wrens soon followed (Borror 1956; Borror & Reese 1953, 1956). For the first time the extraordinary virtuosity of the avian voice was revealed in all its glorious detail. Although Borror was primarily interested in field identification and taxonomy, he proved to be a perceptive bioacoustician (Borror 1960). In some of the first sonograms of birdsong published, he showed that birds are capable of singing with two independent voices (Fig. 1.1), something already hinted at by Potter (1945) in comments on brown thrasher song, sonograms of which were reproduced in the original book Visible Speech (Potter et al. 1947).

    At about the same time, Nicholas Collias, a student of Chicago ecologist W.C. Allee, and a pioneer in the study of animal behavior, collaborated with linguist Martin Joos who had worked on the sound spectrograph project, to publish a sonogram-based functional analysis of the vocal signals of chickens, describing their structure and exploring their function (Collias & Joos 1953; Collias 1960). The stage was set to launch the science of birdsong in earnest.

    Studying by Ear

    In an era dominated by computers, with so many elegant methods available for the analysis of complex sounds, it is hard for us now to imagine the extraordinary impact of the sound spectrograph. Until that time, the only useful descriptions we had, aside from oscillograms, were either musical transcriptions, or verbal renditions, along the lines of the ‘drink-your-tea’ of towhees. Evocative as they were, they hardly did justice to all the acoustic intricacies, as was immediately obvious with William Homan Thorpe’s (1954) first sonograms of chaffinch song (Fig. 1.2).

    Figure 1.2 Three renditions of chaffinch song, from Garstang (1922), Messiaen (Johnson 1975) and Thorpe (1954). The musical transcriptions are from Messiaen’s ‘Vingt Regards sur l’Enfant-Jesus’ (1944) and ‘Chronochromie’ (1960). Thorpe’s chaffinches are three different birds.

    Musical notations are another possibility. The late French composer Olivier Messiaen created some nice renditions of chaffinch song (Fig. 1.2), and birdsongs have engaged many composers over the centuries. The relationship between birdsong and music is an interesting one. Despite his many song-inspired compositions, Messiaen was heard to remark that birds sing in extremely quick tempi which are absolutely impossible for our instruments (Johnson 1975). So he often found it necessary to transcribe them to a slower tempo. The excessively high registers of most birds often required him to transcribe them several octaves lower, suppressing very small pitch intervals. Long before, Barrington (1773) had made the same point. As a bird’s pitch, therefore, is higher than that of any instrument, we are consequently at a still greater loss when we attempt to mark their notes in musical characters, which we can so readily apply to such as we can distinguish with precision. Similarly, the intervals used by birds are commonly so minute, that we cannot judge at all of them from the more gross intervals into which we divide our musical octave (Barrington 1773, p. 266). As a consequence, even though in pieces like ‘Oiseux exotiques,’ and ‘Quatuor pour la Fin du Temps,’ Messiaen captured the essential rhythms of birds singing quite wonderfully, he often rendered particular songsters all but unrecognizable to an ornithologist. Musical transcriptions are in fact not very helpful from a scientific point of view. Sonograms do a much better job (Boxes 1 & 2, pp. 5–6).

    BOX 1 GRAPHIC REPRESENTATION OF SOUNDS

    Three types of graphs are commonly used to visualize sounds; those below were all made from one recording of a white-crowned sparrow song. The first graph illustrates intensity fluctuations over time: the amplitude wave form (A). The x-axis represents the passing of time while the sound volume is reflected in the height of the spikes above and below this axis. Usually the y-axis indicates the relative amplitude: no sound results in no spikes and the loudest sound reaches the maximum extension possible in the graph.

    Second is the sonogram (B) which includes information on the pitch or more precisely the frequency of the sound. The x-axis again represents time and the y-axis pitch, with low-frequency sound near the baseline and high-frequency sound higher up. The frequency of birdsongs usually falls between 500 Hz and 10,000 Hz or, as often indicated, 0.5 and 10 kHz. In this frequency–time graph, information on amplitude is depicted by the darkness of the gray-scale with black reflecting frequencies of the highest amplitude. This gray-scale often disappears in print, replaced by a black and white version. The gray-scale is replaced by a color-scale in many computer software programs.

    Third is the power spectrogram (C), which displays frequency versus amplitude, summated for a segment of sound, or an entire song. It shows the distribution of power through the sound spectrum for a certain song, call, or note, depicting the sound energy present at each frequency.

    The sonogram is the most widely used, often to measure temporal and spectral characteristics of songs. It is also used more and more in bird guides to describe songs, instead of onomatopoeic renditions or musical script. The sonographic representation of sound is objective, but the form it takes depends on the temporal and spectral resolution chosen; the same sound can appear very different if generated with different analysis bandwidths. A narrow-band analysis, with high-frequency resolution yields short, wide graphs, compressed in time (D); a wideband analysis, with high temporal resolution leads to tall, narrow graphs, compressed on the frequency axis (E). It is technically important to state the bandwidth employed in an analysis, but generally people use similar, wideband settings for birdsong, making sonograms more or less comparable, as in the illustrations for this book.

    Hans Slabbekoorn

    BOX 2 A ROUGH GUIDE TO READING SONOGRAMS

    Birds produce an enormous variety of sounds, which can be evaluated by ear, but better by eye via sonographic representations. As a sound is played back, you can read along with a sonogram from left to right (CD2 #3–15). A ‘note’ is the usual term for the smallest sound unit in birdsong appearing in a sonogram as a continuous sound trace. A set of two or more notes repeated coherently in a ‘trill’ is a ‘syllable,’ an unrepeated cluster is a ‘note complex,’ and a series of note-complexes and trills is a ‘song.’

    Pure tone notes at a constant frequency are the simplest components of birdsongs (A). There are examples in the two notes of the African cuckoo (B), the introductory note of a white-crowned sparrow song (C), and the alarm ‘seet’ of the great tit (D).

    The fox sparrow has a few simple notes that change gradually in frequency (E) mingled with other more complex notes in the territorial song (F). Frequency changes in such pure notes can be heard by the human ear, but the details only become clear when a recording is played at a reduced speed, bearing in mind that at a slower speed sounds are lower in pitch. We can simulate birdsong complexity by creating artificial frequency modulations at various rates (G) – at a slow pace, frequency upsweeps and downslurs are easily heard, but become more difficult to discern at the fast pace of the frequency modulations in yellow warbler song (H).

    Tonal sounds often change gradually in frequency, but more discrete changes also occur, as if the bird’s voice is breaking. This can lead to stereotypic note variants, with and without abrupt frequency jumps; four of the seven notes of the second song of a diederik cuckoo display a sudden increase in frequency (I).

    Harmonics or overtones are represented in a sonogram by a typical ladder pattern. One frequency, the fundamental, appears with one or more other sound traces at frequencies that are multiples of the fundamental; harmonics of a fundamental frequency of 500 Hz occur at 1.0 kHz, 1.5 kHz, and so on (J). The relative amplitude of these harmonics may vary, with a strong effect on the quality of the sound, and some can even be completely missing, including the fundamental itself. Overtones are sometimes not harmonics but ‘side-bands.’ If a tone is frequency-modulated it can be described in two ways, as a warble or as a complex tone. Both are valid and what you see on a sonogram depends on the analyzing bandwidth. A warble appears on a sonogram with high-time resolution as a tone fluctuating in frequency. As a complex tone, on a sonogram with high-frequency resolution, you see side-bands spaced at the modulation frequency, above and below the tone. The more sidebands there are, the more emphatic the warble sounds. Amplitude modulation can also create side-bands, just one on each side of the carrier frequency. Side-bands are characteristic of some birds, such as red-winged and yellow-headed blackbirds giving their songs a buzzy, nasal tone. Harmonics are found in many bird calls and songs, such as in the zebra finch (K) and in the human voice.

    Not all songs and calls are tonal and smooth; some sound like hissing, others are noisy and croaky. Noisy sounds lead to messy sonograms with sound diffused across the spectrum, as in some zebra finch syllables and in calls of the Eurasian jay (L). When birds use a double voice they produce two independent sounds at the same time, which can vary independently. The songs of the black-bellied seedcracker contain harmonically related sounds together with another independently varying note (M).

    Hans Slabbekoorn

    As a kind of compromise, several people, especially Saunders (1935), developed schematized frequency/time diagrams that captured some of the dominant features of birdsongs (Fig. 1.3), and this is the method I adopted in my youthful studies of birdsong. Beginning in 1949, playing hooky from my graduate studies in botany and my duties as a plant ecologist for the Nature Conservancy, I hiked around in England, Scotland, France, and the Azores chasing chaffinches. Altogether I transcribed by ear more than five hundred chaffinch songs, learning much about their behavior and ecology in the process.

    Figure 1.3 Transcriptions by Saunders (1935) of song and swamp sparrow songs.

    Looking back, my conclusions about the mosaic structure of chaffinch song dialects were not far off target. But with my primitive methods no one else could tell whether my results were believable or not, including the person who mattered most, destined to be my new boss, W.H. Thorpe at Cambridge University. Thorpe had invited me to join him in 1951 as a research assistant. I already had a PhD in plant ecology, so in one sense I went to Cambridge as a postdoctoral fellow. But I was a neophyte as far as animal behavior was concerned, so I became a graduate student again, this time in zoology, with Thorpe as my major professor.

    SONG STUDIES IN MID-CENTURY

    Thorpe’s laboratory was on the top floor of the zoology department, next door to the legendary insect physiologist, Victor Wigglesworth; this was an appropriate location for Thorpe as an expert on insect behavior, and a wartime innovator in developing new biological methods for pest control. But now his mind was filled with questions about birdsong and the role of learning in its development. The breadth of his erudition was an inspiration to me, as was that of Robert Hinde, the new director of the just-founded Madingley Ornithological_Field Station: both were sources of endless revelations. Above all, Thorpe had just acquired in 1950 a sound spectrograph, only the second to be imported into Great Britain; the first went to the Admiralty Research Laboratory at Teddington, presumably for use by the Royal Navy. With free access to this new machine, and the library of long playing records of bird sounds donated from the archives of the British Broadcasting Corporation (BBC), I was in heaven. We were set for the great leap forward, and never looked back.

    At that time, Thorpe, like everyone else, had to make sound recordings by cutting wax discs, calling for much care and cumbersome equipment; tape recorders, including some that were at least semi-portable, arrived somewhat later. The mysteries of wow and flutter loomed large, and fluctuations in tape speed were a major problem. Among the more imaginative solutions was the heavy cast iron flywheel you screwed on to the tape transport of the famous Magnemite portable tape recorder, before lugging it off into the field. It was a while before truly portable tape recorders came on the market, driven in part by the burgeoning needs of filmmakers, a market force without which the magnificent Swiss-made Nagra recorders would never have seen the light of day.

    Quality playback equipment that was portable enough for field use had to be custom made, one of a host of problems we now solve comfortably with personal computers. The elaborate interactive playback experiments of recent years (McGregor 1992; Dabelsteen & McGregor 1996; McGregor & Dabelsteen 1996) would have been unthinkable even a short time ago.

    It was not until we resettled in Berkeley in 1957, that I was able to pause and gather my thoughts, and reap the full benefit of the open and generous opportunities that Thorpe had given me to sift through the treasures in the BBC birdsong recordings with the sound spectrograph. Many of the sonograms I made then only saw the light of day in 1959, in an invited chapter on animal communication I wrote for a book on Darwin, edited by a colleague from my botanical past, Peter Bell. A few are reproduced in Figures 1.4 and 1.5

    Figure 1.4 Song thrush and mistle thrush songs from BBC archival recordings that I analyzed on Thorpe’s sound spectrograph, in the mid-1950s. At that time we did not know whether birds like the song thrush have a repertoire of discrete song types, or how repertoires are constructed..

    From Marler 1959

    Figure 1.5 Sonograms of various finch calls I recorded in aviaries at Thorpe’s Madingley Ornithological Field Station outside Cambridge in the mid-1950s. Below are the original figure legends edited from Marler (1959). (A) The ‘social’ calls of various finches, all showing affinities with the same basic pattern. Those of the chaffinch and hawfinch, also used as mobbing calls, are the most divergent, but the chaffinch ‘chink’ can be traced back to the same type, as shown in (B). On the left are developmental stages of the chaffinch ‘chink’ call, relating it to the basic finch type. On the right are suggested evolutionary relationships between the ‘primitive’ alarm call (a), used only by young chaffinches but widespread among other finches, and certain adult calls. Different harmonics may have been selected for the ‘huit’ call (c), the ‘seee’ hawk alarm (d); an unusual form of the hawk alarm call, and the ‘chink’ call (e). Call b shows an unusual form of the ‘huit’ call, which may correspond to an intermediate stage. (C) Hawk alarm calls of various finches, showing their essential similarity. The chaffinch only uses this call when young (below); in the adult male it is replaced by the type of call difficult to locate (above), which none of the other finches has evolved. (D) The ‘flight’ calls of some finches, showing how they tend to conform to similar patterns, with the exception of the hawfinch.

    The Debut of the Chaffinch

    This period in the 1950s, when Thorpe presided over the birth of the science of birdsong, was also a time of ferment in the behavioral sciences. Thorpe assembled a massive review of the learning abilities of birds in 1951, setting aside once and for all the shibboleth inherited from the era of Maier and Schneirla (1935) that birds are reflexive machines; this view went back at least to 1924, when Herrick had said that it is everywhere recognized that birds possess highly complex instinctive endowments and that their intelligence is very limited.

    Although he did not actually discover song learning, the blossoming of research on avian vocal learning, in all its dimensions, would almost certainly never have happened without Thorpe’s seminal contributions. His historic papers on song learning in chaffinches, in 1954 and 1958, the first in-depth studies based firmly on sound spectrographic analyses, were crucial in launching the new discipline and they evoked enormous interest (CD1, #3). It was already clear that the chaffinch was an ideal subject for learning studies. Holger Poulsen in Denmark published a paper on inheritance and learning in chaffinch song which I saw in proof while I was completing my pre-Cambridge field project on song dialects. Writing in 1951, just before I arrived in Cambridge I had already concluded, somewhat prematurely, that the geographical variation and development of dialects in the song of the chaffinch are phenotypic variations, arising and persisting because of the two processes of learning to vocalize from associates and of retaining a preference to breed in certain localities (Poulsen 1951; Marler 1952).

    Ethology on Stage

    Meanwhile, the newly emerging discipline of animal behavior was in a state of high excitement. In 1950–51 Thorpe served as the Prather lecturer at Harvard, and he seized on the opportunity to spread the gospel according to European ethologists Konrad Lorenz and Niko Tinbergen; he was preparing the way for his own 1953 book. Learning and Instinct in Animals was a source of inspiration for the next generation. Many of the ideas had already been set forth in his paper on The concepts of learning and their relation to those of instinct; this was presented at a remarkable conference for the Society for Experimental Biology in 1949 in Cambridge on Physiological mechanisms in animal behavior that he helped to organize. Prophets gathered from various disciplines, anticipating future prospects of the behavioral and neuroethological sciences. Lorenz and Tinbergen spoke at length on the principles of ethology, aided and abetted by the Dutch pioneer Gerard Baerends and by Thorpe himself, bringing this material together for a general biological audience for the first time.

    The conference proceedings amounted to a manifesto for research on proximate mechanisms in the behavioral sciences; there were contributions from such famous figures as Weiss on development, Gray and von Holst on the coordination of motor patterns, Boycott and Young on octopus brains and behavior, Lashley and his search for the engram, and many others, culminating with Thorpe’s magisterial overview of learning and instinct. He mentioned song learning only in passing, along with imprinting, pleading for more study. In one sentence, Thorpe anticipated a significant contribution that ethology in general and birdsong research in particular was destined to make in behavioral science. Where the innate powers of recognition can only carry the animal a part of the way towards its goal, the process is completed and adjusted by a tendency to learn in certain restricted times and directions (as in the tendency of a bird to learn and copy the song of its own species in preference to the song of another) so that experience completes for the individual the process commenced for him by his inherited constitution (Thorpe 1950).

    As the sound spectrograph became more widely available, its use in bioacoustics spread like wildfire. Studies of geographic variation in birdsong began to appear, first as a trickle, then as a flood, in both Canada and the USA. The role of song in territorial defense and the impact of sexual selection now became tractable research targets. The significance of local dialects came under renewed scrutiny. Sonograms provided the bedrock for the initiative that Thorpe had launched on the elusive problem of vocal learning, paving the way for study of the underlying neural mechanisms, culminating in the discovery of the song system in the avian brain more than twenty years later (Nottebohm et al. 1976).

    DIALECTS

    In one important step, the reality of local dialects in birdsong became clear; what had been suspected by ear became fact. Dialects exist in birdsong, on a scale so local that, for creatures as vagile as birds, it hardly seemed plausible to attribute them to genetic differences between populations. All of the early pioneers, including the leading evolutionary biologists of the day, like Julian Huxley (1942) and Ernst Mayr (1942), as well as Thorpe (1951), recognized the scientific challenge that dialects represent. The many insights that flow from immersion in the study of song dialects is evident from the career of Luis Baptista (Baptista 1975, 1990), to whom this book is dedicated. To this day dialects remain an important focus in birdsong studies; witness Kroodsma’s unexpected and provocative discovery of song dialects in a sub-oscine, the three-wattled bellbird (see Chapter 4, p. 108).

    The conspicuousness of the dialects in white-crowned sparrow songs in western North America (Fig. 1.6) was instrumental in the emergence of this species as a subject of choice in research on song variation and development, in both laboratory and field studies (Marler & Tamura 1962, 1964;Konishi 1965a; Baptista 1975; Baker & Cunningham 1985). Sonographic analyses elevated studies of song dialects of such European birds as chaffinches, corn buntings, yellowhammers, and redwings, from the birdwatcher level (Huxley 1947; Marler 1952) to the status of scientific research (Slater et al. 1984; McGregor 1983; Baker et al. 1987a; Espmark et al. 1989).

    Figure 1.6 Local dialects around the San Francisco Bay in white-crowned sparrow song. The figure on the left, based on Marler & Tamura (1962) was reproduced in many textbooks. On the right side are the dialects thirty years later, from Slabbekoorn et al. 2003.

    Problems of micro- and macro-geographic variation now began to come into focus, eventually providing new windows on cultural evolution and speciation (Mundinger 1982; Lynch 1996; Martens 1996; Payne 1996; Slabbekoorn & Smith 2002b). For the first time we began to gain some sense of what the entire song repertoire of a species encompasses, and of the challenges confronting a bird in recognizing its own species song and distinguishing it from the many others with which it might be confused. Given the enormous variability of birdsong this might seem to be a daunting task, and yet birdwatchers usually have no trouble telling one from another. How in fact do birds do it?

    THE MYSTERY OF SPECIES UNIVERSALS

    Song Recognition

    There is some evidence of innate song recognition, and Thorpe was the first to suggest that it must occur. He was preoccupied with questions about song recognition from the earliest days, especially since some of his laboratory subjects seemed to resist being taught alien songs when first learning to sing (Thorpe 1961). He was greatly impressed by the instant recognizability of the songs of chaffinches that had lived for generations in completely new environments, after introduction by homesick colonials in New Zealand and South Africa (CD1, #4). It seemed to him that they must be able to recognize the songs they should be attending to while learning to sing. In Cambridge, his young chaffinches found his artificially created songs quite unacceptable, but they would learn re-articulated chaffinch songs. They also learned some elements of a tree-pipit song selected by Thorpe because he thought the tonal quality was reminiscent of the chaffinch (Fig. 1.7).

    Figure 1.7 Six chaffinch songs, four of them from W.H. Thorpe’s original recordings. A & B are Cambridge birds from the same local dialect, recorded by Thorpe in 1954. C is a bird raised in social isolation by Thorpe as a Kaspar Hauser. D is a bird trained by Thorpe with a recording of a tree pipit song. E is a song from Sussex, England, the source of the chaffinches introduced to New Zealand between 1840 and 1860. F is a chaffinch song from New Zealand (CD1 #3–4).

    Thorpe tentatively concluded that there must be something distinctive about the tonal quality of songs, providing the basis for any innate ‘blueprint’ they might possess. He thought that young birds might be innately cued to these tonal qualities, hence their tendency to learn conspecific song. He seemed reluctant to admit the possibility that innate recognition of song might be based not on its tonal quality, but on a much more complete mental image of many or even all of the basic components of its species not necessarily all sharing the same tonal quality. As the sound spectrograph made much more comprehensive analysis possible, we could begin conducting large-scale investigations of the entire song repertoire of a species, to identify features present in all songs, and thus candidates as possible cues for species recognition. The results were fascinating, and suggested that we needed to reopen the whole question of song recognition.

    We have long known that song plays a crucial role, not only in mate choice, but also in territorial defense, where same-species, same-sex birds are usually the primary targets (Howard 1920; Hinde 1956a). As methods for song playback to territorial males were perfected (Bremond 1968, 1976; Krebs 1977a), and the importance of singer location and individual identity and the ability to match songs with rivals became clear (Box 3, p. 49; see Chapter 4) (Brooks & Falls 1975 a, b; Falls & Brooks 1975; Falls 1978, 1982), it was inevitable that individual and species differences in song characteristics would eventually come under renewed scrutiny (Becker 1982; Falls 1992). Almost without exception, playback of songs of other species proved to be much less potent in evoking territorial response from resident males than own-species songs. But the task of establishing which particular song features were critical in own-species/other species discrimination was a daunting one, and not many people had the tenacity to grapple with it. Again, the sound spectrograph came to the rescue. Amidst the close scrutiny of all aspects of song variation, comprehensive analyses of song structure now began to emerge for a few species.

    The obvious first step in using song playback to study the responses of territorial males was the selection of particular songs as stimuli. Because learned birdsongs are so incredibly variable the choice is harder than it sounds, if the choice is to be representative. Attempting to characterize the song of an entire species is a daunting undertaking, and compromises are inevitable. Archived sound recordings like those in the Borror Laboratory of Bioacoustics at Ohio State, and the Macaulay Library of Natural Sounds at Cornell University (see Chapter 12) become a treasured resource, and much time and devotion to duty is required of the researcher to gather the necessarily large volume of new material. Luis Baptista devoted much of his life to recording songs of the white-crowned sparrow. He often worked in suburbia, stopping his car in mid-traffic to record a new bird, oblivious to the resulting traffic jam. Other early enthusiasts for comprehensive sonographic song coverage included Gerhardt Thielcke and his European tree-creepers, Fernando Nottebohm’s chingolos in Argentina, Peter Slater and chaffinch song, the nightingales of Henrike Hultsch and Dietmar Todt, Michel Kreutzer’s cirl buntings, Peter Becker’s goldcrests and firecrests, Bob Lemon’s cardinals, not forgetting Don Kroodsma’s lifelong devotion to the voices of the wrens of the world. These long-term studies of species’ song repertoires have all been invaluable in providing a solid basis for subsequent work on recognition and song ontogeny. They provided an intriguing opportunity to follow up in a new way on Thorpe’s early reflections on how we should interpret the apparent contradiction between the great variability associated with the learnability of song, the clear evidence of song universals shared by all the singers of a species, and problems of song recognition.

    The Indigo Bunting

    Before launching his path-breaking studies of the social life of bee-eaters and birds navigating by the stars, Steven Emlen at Cornell did one of the first studies of song recognition; he quickly found the selection of playback stimuli to be a problem even with the relatively simple song of the indigo bunting (CD1 #5). This bird became the subject of what is still one of the most comprehensive analyses of the mechanisms underlying species and individual song recognition. Emlen addressed the problem in an interesting way, by first looking at sonograms of indigo bunting song from several parts of its range, in and around Ithaca, New York, trying to figure out what they had in common (Emlen 1971, 1972). Thompson (1970) had already conducted a similar exercise in a different part of the species range, and the comparison turned out to be fascinating and unexpected (Shiovitz & Thompson 1970; Shiovitz 1975; Thompson 1976; Margoliash et al. 1991; Baker & Boylan 1995; Payne 1996). Shiovitz and Thompson had assembled a comprehensive sonogram catalog of less than a hundred distinct syllable types, from which every one of the hundreds of songs they analyzed from Kentucky and Michigan could be assembled. Amazingly, miles away in New York, the identical catalog proved to be equally serviceable in classifying almost all of Emlen’s songs (Fig. 1.8). He only had to add a few more types for the coverage to be complete. Since then the Thompson catalog has been the basis for every study of indigo bunting song, only updated 20 years later to 127 syllable types by Baker & Boylan (1995). The bottomline is that, over thousands of square miles, indigo bunting songs are constructed from essentially the same set of building blocks.

    Figure 1.8 The catalog of the universal repertoire of indigo bunting song syllables, from Baker and Boylan (1996). Also shown are nine syllable types recurring in similar form in three states.

    From Shiovitz 1975.

    Although its significance was not fully appreciated at the time, this was a very surprising discovery. Even though these are learned birdsongs, species universals are present as a dominant feature of indigo bunting song. Despite the variability that you would expect to be an inevitable consequence of the cultural transmission of song, and the copying errors that must ensue, despite the existence of dialects, and the modification of song by improvization and invention, and the great individuality of these and all other learned birdsongs,

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