Table Of ContentS.L. Hopp· M.J. Owren· C.S. Evans (Eds.)
Anima! Acoustic Communication
Springer
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S.1. Hopp • M.}.Owren . C.S. Evans (Eds.)
Animal
Acoustic
Communication
Sound Analysis
and Research Methods
With 115 Figures
Springer
DR. STEVEN 1. Hopp
University of Arizona
Dept. of Ecology and Evolutionary Biology
Tucson, AZ 85721
USA
J.
DR. MICHAEL OWREN
Cornell University
Dept.ofPsychology
Ithaca, NY 14853
USA
DR. CHRISTOPHER S. EvA NS
Macquarie University
Dept of Psychology
Sydney, NSW 2109
Australia
Library of Congress
Animal acoustic communication : sound analysis and research methods /
Steven 1. Hopp, Michael J. Owren, and Christopher S. Evans (eds.).
p. cm.
Includes bibliograhical references and index.
ISBN-13: 978-3-642-76222-2 e-ISBN-13: 978-3-642-76220-8
DOI: 10.1007/978-3-642-76220-8
1. Animal sounds. 2. Sound production by animals. 3. Animal
communication. I. Hopp, Steven 1., 1954-. 11. Owren Michael
J., 1955-. 111. Evans, Christopher Stuart, 1959·
QL765.A.645 1977
591.59#4-dc21 97-10402
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PREFACE
The last decades have brought a significant increase in research on acoustic communi
cation in animals. Publication of scientific papers on both empirical and theoretical
aspects of this topic has greatly increased, and a new journal, Bioacoustics, is entirely
devoted to such articles. Coupled with this proliferation of work is a recognition that
many of the current issues are best approached with an interdisciplinary perspective,
requiring technical and theoretical contributions from a number of areas of inquiry
that have traditionally been separated. With the notable exception of a collection edited
by Lewis (1983), there have been fewvolumes predominatelyfocused on technical issues
in comparative bioacoustics to follow up the earlyworks edited by Lanyon and Tavolga
(1960) and Busnel (1963). It was the tremendous growth of expertise c:()ncerning this
topic in particular that provided the initial impetus to organize this volume, which
attempts to present fundamental information from both theoretical and applied aspects
of current bioacoustics research. While a completely comprehensive review would be
impractical, this volume offers a basic treatment of a wide variety of topics aimed at
providing a conceptual framework within which researchers can address their own
questions. Each presentation is designed to be useful to the broadest possible spectrum
of researchers, including both those currently working in any of the many and diverse
disciplines of bioacoustics, and others that may be new to such studies.
Several previous collections have addressed communication-related topics in par
ticular taxonomic groups, such as fish (Tavolga et al. 1981), birds (Kroodsma et al. 1982;
Catchpole and Slater 1995; Kroodsma andMiller1996), andnonhuman primates (Snow
don et al. 1982; Todt et al. 1988; Zimmermann et al. 1995). Typically these treatments
have provided a mix of technical, empirical, and theoretical contributions. Other com
piled volumes have been devoted to specific subtopics in animal communication in
cluding animal perception (e.g., Berkley and Stebbins 1990; Stebbins and Berkley 1990)
and sound-broadcast, or playback, studies (McGregor 1992). This book differs from
these earlier volumes in that it specifically responds to recent developments in digital
computing, and its applications. In addition, we include several chapters outlining the
current technological and methodological approaches to studies in particular areas of
animal acoustic communication.
A large part of the growth in animal acoustic communication research can be attrib
uted to the development of accessible computer hardware and software that allows
sophisticated digital signal processing (DSP) algorithms to be applied to sound-derived
data. A technological revolution of the last decades has produced a wide variety of new
approaches, newways of thinking about problems, and much greater capacity for proc
essing and manipulating information. However, the extent to which new developments
have been transferred across disciplines has, in part, determined some of the advances
in a given area. For research in acoustics, these advances include the development of
processing hardware, the discovery of efficient methods for the imaging and analysis
of signals, and development of applications for these procedures across disciplines and
even across taxa. Many of these techniques have been used widely in engineering fields
but have only recently been employed routinely by comparative researchers. Theoreti
cal discussions of digital signal processing have been published primarily in the engi
neering literature, and have consequently been relatively inaccessible to comparative
researchers. Similar difficulties also occur within the discipline of comparative bioa-
VI
coustics. Here, important factors indude the great diversity of backgrounds found
among animal-oriented investigators, as weIl as significant variation in both the nature
of the sound signals of interest and the circumstances oft he animal species that produce
them. The result has been that barriers to mutually beneficial sharing ofknowledge have
arisen due to rather arbitrary divergence of terminology and methods. One overriding
purpose in presenting this volume, then, is to introduce common techniques in DSP in
a manner that is comprehensible to both students and professionals conducting bioa
coustics research in diverse subdisciplines. In addition, the various authors here have
tried to provide dear examples ofboth typical and possible applications of given meth
ods or techniques, and outlining practical suggestions wherever possible.
The first portion of the book is dedicated to sound analysis and computer-applica
tions. In the first chapter, Gerhardt outlines the basic techniques of recording and ana
lyzing animal sounds. The methods discussed therein provide a solid background for
applications in diverse areas of research. This presentation is followed by a pair of com
plementary chapters discussing the applications of digital technology to sound signal
representation and processing in the time-and frequency-domains. The first, by ele
ments, outlines the general principles of DSP in both mathematical and verbal form.
The second, by Beeman, explicitly explains and demonstrates these principles in the
context of the computing equipment and research topics that might be typical of sci
entists working in bioacoustics. The next chapter, by Stoddard, offers a practical dis
cussion of the crucial role of electronic filters in bioacoustics research, along with ex
planations of related terminology, design, and application. The final chapter in this
section, by Owren and Bernacki, addresses the conceptually narrower topic of how to
characterize energy patterning in the frequency spectrum of sound. The approach de
scribed, based on linear prediction, has been used in analogous applications in other
disciplines, particularly in the analysis and synthesis ofhuman speech.
The second section addresses four general areas of study in which recent technologi
cal advances have allowed researchers both to address long-standing questions of in
terest with greatly increased power and to pioneer entirely new lines of inquiry. Each
of these chapters combines the presentation of important theoretical and methodologi
cal topics with both an overview of the associated empiricalliterature and examples of
studies in which new approaches have been used. The first two chapters discuss three
realms of acoustics that fall outside the typical human auditory experience. The chapter
by Pye and Langbauer on infrasound and ultrasound, airborne acoustic signals that fall
belowand above the range ofhuman sensitivity, respectively, is complemented by the
discussion by Tyack, on underwater sound communication. Each of these signaling
domains presents unique technical challenges and have given rise to quite unexpected
research findings. In the third chapter, Rubin and Va tikiotis-Bateson discuss the acous
tics of species-typical communication in human speech, an extensively studied com
munication system. The techniques and theoretical models outlined therein provide a
rich base for studies with a comparative focus. This area of research is typically not
considered in a comparative perspective, although it has historically been immensely
important to the investigation ofs ound signals in nonhuman animals. The development
of much of the hardware and software currently in use in comparative bioacoustics can
be directly linked to studies of speech production or perception. For example, ideas
about the source-filter theory of voice production in humans have provided models for
vn
understanding some aspects of sound production by both primates and birds. In the
final chapter in this section, Gaunt and Nowicki specifically examine the latter topic,
i.e. sound production by birds, and in doing so demonstrate the power of combining
technological advances with interdisciplinary thinking.
The three chapters in the final section focus on different aspects of the question of
how communicative signals affect the receiver. The methods are diverse, ranging from
how animals overtly or covertly respond to signals, to basic questions such as whether
organisms can discriminate among similar signals. Hopp and Morton discuss sound
playback studies, which are used to assess the re action of animals to the broadcast of
particular sounds. Such studies are typically conducted in the field with free-ranging
animals. Where naturally occurring responses to sounds are lirnited, training animals
to respond differentially to signals allows researchers to ask questions about sound
perception and categorization of signals. These techniques are discussed by Cynx and
Clark. The last chapter is a contrast to the other contributions in that its focus is not
primarily on issues related to measurement and analysis of acoustic signals. In this
chapter, Ball and Dufty provide detailed, practical methodological information about
the interplay between acoustic signals and the neuroendocrine systems of animals. As
the chapter demonstrates, hormonal responses to auditory stimulation play an impor
tant and immediate role in many aspects of acoustic communication.
As we prepare to enter a new century, it is nearly impossible to anticipate the inevi
table advances in technology, or the profound effects that these advances will have on
studies of animal acoustic communication. We hope this volume will provide readers
with a framework for understanding the dramatic changes that have taken place in
animal communication research. It is meant as a stepping stone to our shared future
in comparative bioacoustics.
Steven L. Hopp Tueson, Arizona, USA
Michael J. Owren Ithaca, NY, USA
Christopher S. Evans Sydney, Australia
References
Berkley MA, Stebbins WC (eds) (1990) Comparative perception, voll. Basic mechanisms. Wiley, New
York
Busnel RG (ed) (1963) Acoustic bel1avior of animals. Elsevier, Amsterdam
Catchpole CK, Slater PIB (1995) Bird song. Cambridge New York
Kroodsma OE, Miller EH, Ouellet H (eds) (1982) Acoustic communication in birds, vols I and 11. Academic
Press, NewYork
Kroodsma OE, Miller EH (eds)(1996) Ecology and evolution of acoustic communication in birds. Cornell
University Press, Ithaca
Lanyon WE, Tavolga WN (1960) Symposium on animal sounds and communication. Am Inst Biol Sei,
Washington
Lewis B (ed) (1983) Bioacoustics: a comparative approach. Academic Press, NewYork
McGregor PK (ed) (1992) Playback and studies of animal communication. Plenum, New York
Snowdon CT, Brown CH, Petersen MR (1982) Primate communication. Cambridge Universitv Press, New
York
Stebbins WC, Berkley MA (eds) (1990) Comparative perception, volII. Complex signals. Wiley, New York
Tavolga WN, Popper AN, Pay RR (eds) (1981) Hearing and sound communication in fishes. Springer
Berlin Heidelberg New York
VIII
Todt D, Goedeking P, Symrnes D (eds) (1988) Primate voca1 communication. Springer Berlin Heidelberg
NewYork
Zinlmerman E, Newrnan JD, Jürgens, U (eds) (1995) Current topics in primate vocal communication.
Plenum, NewYork
Acknowledgments
Over the years there were many people who offered suggestions, support, discussion,
inspiration, time, and chapter reviews. We thank the following people for their contri
butions: W. Au, C.A. Boone, B. Burgess, E. Carterette, F. Cheever, D. Czeschlik, V.J.
DeGhett, U. Gramm, P.S. Kaplan, B.E. Kingsolver, A. Kirby, T. Krammer, D.E. Krood
sma, C.A. Logan, A. Popper, C. Seelinger, J. Spiesberger, R.A. Suthers. In addition we
thank the authors of this volume for their dedication and endurance.
CONTRIBUTORS
GREGORY F. BALL
Department ofP sychology, lohns Hopkins University, 3400 N. Charles Street, Baltimore,
Maryland 21218-2686 USA
KIMBEEMAN
Engineering Design, 43 Newton Street, Belmont, Massachusetts 02178, USA
ROBERT BERNACKI
P.O. Box 3188, Bloomington Scientific, Bloomington, Indiana 47402 USA
STEPHEN CLARK
Psychology Department, 124 Raymond Ave, Vassar College, Poughkeepsie, New York
12601, USA
MARK CLEMENTS
School of Electrical Engineering, Georgia Institute of Technology, Atlanta, Georgia,
30332-0250, USA
JEFFERY CYNX
Psychology Department, 124 Raymond Ave, Vassar College, Poughkeepsie, New York,
12601, USA
ALFRED M. DUFTY, JR.
Department ofBiology, Boise State University, 1910 University Drive, Boise, Idaho, 83725,
USA
CHRISTOPHER S. EvA NS
School ofBehavioural Sciences, Macquarie University, Sydney NSW 2109, Australia
ABBOT S. (TOBY) GAUNT
Department ofZoology, Ohio State University, 1735 Neil A v., Columbus, Ohio, 43210, USA
H. CARL GERHARDT
Division of Biological Sciences, 105 Tucker Hall, University of Missouri at Columbia,
Columbia, Missouri, 65211, USA
Steven L. Hopp
Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Ari
zona, 85721, USA
WILLIAM R. LANGBAUER, JR.
Pittsburgh Zoo, P.O. Box 5250, Pittsburgh, Pennsylvania, 15206, USA
x
EUGBNB S. MORTON
Conservation and Research Center, Smithonian Institution, National Zoological Park,
1500 Remount Road, Front Royat Virginia 22630, USA
STBPHBN NOWICKI
Department ofZoology, Duke University, Durham, North Carolina, 27706, USA
J.
MICHABL OWRBN
Department of Psychology, Cornell University, 224 Uris Halt Ithaca, New York 1485.3
USA
J.DAVIDPYB
Department ofZoology and Cpmparartive Physiology, Queen Mary College, University
ofL ondon, Mile End Road, London E1 4NS, UK
PHILIP RUBIN
Haskins Laboratories and Yale University School ofM edicine, Department of Surgery,
Otolaryngology, 270 Crown Street, New Haven, Conneticut, 06511, USA
PHILIP K. STODDARD
Department of Biological Sciences, Florida International University, University Park,
Miam~ Florida, 33199, USA
PBTBR L. TYACK
Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, 02543, USA
ERIC VATIKIOTIS-BATBSON
ATR Human Information Processing Research Laboratories, 2-2 HikaridailSeika-cho,
Sorkau-gun,Kyoto 619-02, Japan
