Acoustic Communication in Birds. Production, Perception & Design Features of Sounds

391 Pages·1983·6.729 MB·English

by  O. Kroodsma (Auth.)

COMMUNICATION AND BEHAVIOR
AN INTERDISCIPLINARY SERIES
Under the Editorship of Duane M. Rumbaugh,
Georgia State University and Yerkes Regional
Primate Research Center off Emory University
DUANE M. RUMBAUGH (ED.), LANGUAGE LEARNING BY A
CHIMPANZEE: THE LANA PROJECT, 1977
ROBERTA L. HALL AND HENRY S. SHARP (EDS.),
WOLF AND MAN: EVOLUTION IN PARALLEL, 1978
HORST D. STEKLIS AND MICHAEL J. RALEIGH (EDS.), NEUROBIOLOGY OF SOCIAL
COMMUNICATION IN PRIMATES: AN EVOLUTIONARY PERSPECTIVE, 1979
P. CHARLES-DOMINIQUE, H. M. COOPER, A. HLADIK, C. M. HLADIK, E. PAGES,
G. F. PARIENTE, A. PETTER-ROUSSEAUX, J. J. PETTER, AND A. SCHILLING (EDS.),
NOCTURNAL MALAGASY PRIMATES: ECOLOGY, PHYSIOLOGY, AND BEHAVIOR, 1980
JAMES L. FOBES AND JAMES E. KING (EDS.), PRIMATE BEHAVIOR, 1982
DONALD E. KROODSMA AND EDWARD H. MILLER (EDS.),
ACOUSTIC COMMUNICATION IN BIRDS, VOLUME 1: PRODUCTION,
PERCEPTION, AND DESIGN FEATURES OF SOUNDS, 1982. VOLUME 2: SONG
LEARNING AND ITS CONSEQUENCES, 1982
Acoustic Communication
in Birds
Volume 1
Production, Perception, and
Design Features of Sounds
Edited by
DONALD E. KROODSMA
Department of Zoology
University of Massachusetts
Amherst, Massachusetts
EDWARD H. MILLER
Vertebrate Zoology Division
British Columbia Provincial Museum
Victoria, British Columbia, Canada
Taxonomic Editor
HENRI OUELLET
Vertebrate Zoology Division
Museum of Natural Sciences
National Museums Canada
Ottawa, Ontario, Canada
1982
ACADEMIC PRESS
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Library of Congress Cataloging in Publication Data
Main entry under title:
Acoustic communication in birds.
(Communication and behavior)
Vol. 1: Edited by Donald E. Kroodsma, Edward H.
Miller, and Henri Ouellet.
Includes bibliographical references and indexes.
1. Bird-song. 2. Animal communication.
I. Kroodsma, Donald E. II. Miller, Edward H.
III. Ouellet, Henri. IV. Series.
QL698.5.A26 598.2'59 82-6730
ISBN 0-12-426801-3 (v. 1)
PRINTED IN THE UNITED STATES OF AMERICA
82 83 84 85 9 8 7 6 5 4 3 2 1
The contributors dedicate these volumes to Peter
Marler. Peter's research and writings have placed the
study of animal communication in the fore of modern
evolutionary and ecological thought in ethology, and
his impact is apparent from the sheer number and
diversity of citations he receives in these two volumes.
Research into almost any facet of bird acoustics puts
one into contact with his work, on topics such as
learning, geographic variation, species-specificity,
"design features," individual variation, and grading.
In addition, Peter has touched many workers indi­
vidually, as graduate students, postdoctoral re­
searchers, and peers, whereby his influence upon re­
search and thinking on animal communication has
carried yet further.
We are sure that this treatise reflects Peter Marler's
influence upon the field of avian acoustics, and hope
that it also reflects our esteem and respect for him as a
scientist, teacher, and colleague.
Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
Arthur P. Arnold (75), Department of Psychology, and Laboratory of Neuroen-
drocrinology, Brain Research Institute, University of California, Los An­
geles, California 90024
Peter H. Becker (213), Institut fur Vogelforschung, Vogelwarte Helgoland, D
2940 Wilhelmshaven 15, West Germany
John H. Brackenbury (53), Department of Biology, University of Salford, Sal-
ford M5 4WT, England
Clive K. Catchpole (297), Department of Zoology, Bedford College, University
of London, London NW1 4NS, England
Robert J. Dooling (95), Department of Psychology, University of Maryland,
College Park, Maryland 20742
Edward H. Miller (253), Vertebrate Zoology Division, British Columbia Provin­
cial Museum, and Biology Department, University of Victoria, Victoria,
British Columbia, Canada
Eugene S. Morton (183), National Zoological Park, Smithsonian Institution,
Washington, D.C. 20008
Douglas G. Richards (131), Kewalo Basin Marine Mammal Laboratory, Univer­
sity of Hawaii, Honolulu, Hawaii 96814
David C. Wickstrom (1), Library of Natural Sounds, Laboratory of Ornithology,
Cornell University, Ithaca, New York 14850
R. Haven Wiley (131), Department of Zoology, University of North Carolina,
Chapel Hill, North Carolina 27514
Foreword
It is entirely appropriate that these volumes of collected essays on aspects of
the study of bird vocalizations should be dedicated to Peter Marler. While the scien­
tific study of bird vocalizations was initiated by W. H. Thorpe, its subsequent
expansion and elaboration was due primarily to Peter Marler.
W. H. Thorpe was initially an entomologist. Finding that some aspects of
insect behavior were more labile than had formerly been supposed, he came to
feel that the most pressing problems in the study of animal behavior concerned
the interface between "instinct," as it was then called, and learning. As an
amateur ornithologist, he realized that birds provided exceptionally suitable ma­
terial for this work. Birds have a repertoire of relatively stereotyped movement
patterns and yet a4t4 the same time exhibit marked learning ability. Accordingly, in
1950 he set up An Ornithological Field Station" (now called the Sub Depart­
ment of Animal Behaviour) at Madingley, Cambridge. Knowing that Chaffinch
{Fringilla coelebs) song was subject to at least some individual variation and
some flexibility (e.g., Poulsen, 1951), he decided to study the ontogeny of
Chaffinch vocalization, and initiated a program of hand-rearing Chaffinches and
studying their song development after varying exposure to the species' song
(Thorpe, 1963).
Peter Marler was the first graduate student at Madingley. When he came, he
already had a Ph.D. in botany and a tenured job. It must have been a decision of
considerable courage to give up that job in order to take a Ph.D. in a subject that
really interested him. His Ph.D. was a field study of the Chaffinch: he wanted to
work in the field and the choice of the Chaffinch as subject meshed well with
Thorpe's own work. At that time I was studying the courtship of captive Chaf­
finches and I am sure that Peter will remember the long discussions, and some­
times disputes, that we had over whether this or that posture should be called the
lopsided wings-drooped posture or something a bit snappier—issues that seemed
terribly important to us then!
Peter stayed on as a postdoctoral worker and took on the song learning work.
His training as an all-round biologist stood him in good stead, and at that time he
became interested not only in the ontogenetic problem that Thorpe was study-
xiii
xiv Foreword
ing—how does Chaffinch song develop—but also in the other three questions
with which ethologists are concerned—those of causation, function, and evolu­
tion. When he subsequently moved to Berkeley, he took the song problem with
him. Changing his British speech for what seemed to some of us a rather extreme
American accent, and the Chaffinch for White-crowned Sparrows (Zonotrichia
leucophrys) and juncos, he started his well-known studies of ontogeny. At
around that time the study of bird song really took off and it can now be regarded
as having produced material of crucial importance for all four major questions of
ethology.
Taking the ontogenetic question first, three fundamentally important princi­
ples have been established largely or entirely through work on bird song. The
principle of sensitive periods in development, though apparent from earlier work
on imprinting and other phenomena, owes a great deal to studies of bird song.
The fact that what an animal learns is constrained in part by its species, though
again an idea coming also from other sources of evidence, was established most
firmly in the 1950s and early 1960s by the work on bird song. And third, the
view that the elaboration and perfection of the song pattern depend on compari­
son between the vocal output and a previously established template casts a new
light on many aspects of ontogeny. In all of these issues Peter Marler and his
colleagues played a leading role. Understanding the processes of song develop­
ment was facilitated by comparisons between studies of different species (e.g.,
Immelmann, 1969; Konishi, 1964; Nicolai, 1956), and Marler himself used a
comparative approach to good effect, relating plasticity to the ecology of the
species (Marler, 1967).
Marler's field studies of the Chaffinch had inevitably alerted him to problems
concerned with the causation of bird song, and his thesis contains a great deal of
observational material on the factors determining when a Chaffinch sings. He
soon became interested also in the patterning of bird song, and was one of the
first people to study the detailed sequencing of different song types in an indi­
vidual's vocal output (e.g., Isaac and Marler, 1963). However, a more detailed
study of the neural mechanisms underlying bird song arose from one of the
findings of the studies of song ontogeny. Domestic Fowl (Gallus domes-
ticus) and Ringed Turtle-Doves (Streptopelia risoria) were found to be capable of
developing all the normal species' vocal signals, even though deafening took
place soon after hatching. On the other hand, Konishi and Nottebohm (e.g.,
1969) found that, in a number of species of songbirds, deafening before full song
had developed resulted in consistent abnormalities in their song, and often in a
regression to a rather amorphous type of vocal output. However, birds that had
already learned to sing could continue singing after deafening. Since it was
known that the control of human speech is influenced by the speaker's perception
of himself speaking, the finding that deaf birds could continue to sing normally
was surprising. One possibility was that the deafened bird was using feedback
Foreword XV
from the muscles. Nottebohm therefore investigated the effects of severing the
hypoglossal nerves in the syrinx. This has led to the discovery of laterality in the
motor control of bird vocalizations, and what is more of a partially reversible
laterality. Furthermore, it has led to a detailed investigation of the brain mecha­
nisms underlying bird song, and has provided us with quite new data on the role
of hormones in the development of neural mechanisms (e.g., Nottebohm, 1980).
Marler's training as an all-round naturalist led him early on to ask functional
questions. He was concerned with the functions of different calls in the Chaf­
finch's repertoire. What information did each carry? This led him into an attempt
to categorize the nature of the information carried in animal signals (Marler,
1961). While not everyone will agree with the view that natural selection always
acts to enhance the effectiveness of signals in transmitting information about the
signaler, Marler's formulations greatly facilitated the study of communication.
He was also concerned with the diversity of avian vocalizations. Noticing that
the alarm calls of different species tended to resemble each other, while the songs
were very different, he speculated about the selective factors controlling the
form of avian vocalizations—selective factors that included the optimal degree of
audibility in the habitat in question as well as the particular response to be
elicited in the responding individual (e.g., Marler, 1955). In addition, in com­
paring the ontogeny of different avian species he was forced to ask why learning
plays a much greater part in some species than others, why the sensitive period
occurs at different ages in different species, why learning is constrained in one
way in some species and in another way in others, and so on. We do not yet
know the answers to all these questions, but it was Marler's pioneering work in
the early 1950s that posed them.
Finally, and inevitably, functional questions led to questions about the course
of evolution of avian vocalizations. Peter Marler was concerned with this prob­
lem at a very early stage: as a student, he went on an expedition to the Canary
Islands and became concerned with the differences between the songs of the
species found there and their mainland counterparts (Marler and Boatman,
1951). His work on bird ontogeny led him into questions of the relations between
song and call notes, and the functional questions he raised were inevitably linked
with evolutionary questions about how song evolved.
While Peter Marler's work on avian vocalizations has led to progress in
answering all four of the major questions in which ethologists are interested, that
is not all. Perhaps influenced by Thorpe, who was a pioneer in emphasizing the
importance of perceptual processes at a time when most ethologists were think­
ing in relatively mechanistic terms and were not yet sensitive to his suggestions,
Peter Marler became interested at an early stage in the relations between the
physical structure of bird vocalization and their quality as perceived by the
recipient (e.g., Marler, 1969). And while outside the scope of this volume,
Marler's more recent work on primate vocalizations, and his emphasis on com-
xvi Foreword
mon features between avian communication and human language (Marler, 1970)
has done much to stimulate research.
I would suggest that the study of bird song is the example par excellence of the
ethological approach. It involves the study of a naturally occurring pattern of
behavior against a background of the natural history of the species concerned,
but employs an experimental methodology. It involves questions about the
causation, ontogeny, function, and evolution of the pattern in question, questions
that are at the same time independent and interfertile. It is probably true to say
that the study of bird song has done as much for the advancement of ethology as
the study of any other specific aspect of behavior. In this, Peter Marler has
played a major role.
Robert A. Hinde
REFERENCES
Immelmann, K. (1969). Song development in the Zebra Finch and other estrilidid finches. In "Bird
Vocalizations. Their Relation to Current Problems in Biology and Psychology" (R. A. Hinde,
ed.), pp. 61-74. Cambridge Univ. Press, London and New York.
Isaac, D., and Marler, P. (1963). Ordering of sequences of singing behaviour of Mistle Thrushes in
relation to timing. Anim. Behav. 11, 179-188.
Konishi, M. (1963). The role of auditory feedback in the vocal behavior of the Domestic Fowl. Z.
Tierpsychol. 20, 349-367.
Konishi, M. (1964). Effects of deafening on song development in two species of juncos. Condor 66,
85-102.
Konishi, M., and Nottebohm, F. (1969). Experimental studies in the ontogeny of avian vocaliza­
tions. In "Bird Vocalizations. Their Relation to Current Problems in Biology and Psychol­
ogy" (R. A. Hinde, ed.), pp. 29-48. Cambridge Univ. Press, London and New York.
Marler, P. (1955). Characteristics of some animal calls. Nature (London) 176, 6.
Marler, P. (1961). The logical analysis of animal communication. J. Theoret. Biol. 1, 295-317.
Marler, P. (1967). Comparative study of song development in sparrows. Proc. 14th Int. Ornith.
Cong. Oxford, pp. 213-244, Blackwell, Oxford.
Marler, P. (1969). Tonal quality of bird sounds. In "Bird Vocalizations. Their Relation to Current
Problems in Biology and Psychology" (R. A. Hinde, ed.), pp. 5-18. Cambridge Univ. Press,
London and New York.
Marler, P. (1970). Birdsong and speech development: could there be parallels? Am. Sci. 58 (6),
669-673.
Marler, P., and Boatman, D. J. (1951). Observations on the birds of Pico, Azores. Ibis 93, 90-99.
Nicolai, J. (1956). Zur Biologie und Ethologie des Gimpels (Pyrrhula pyrrhula L.). Z. Tierpsychol.
13, 93-132.
Nottebohm, F. (1980). Brain pathways for vocal learning in birds. A review of the first 10 years.
Prog, in Psychobiol. and Physiol. Psychol. 9, 85-124.
Poulsen, H. (1951). Inheritance and learning in the song of the Chaffinch (Fringilla coelebs).
Behaviour 3, 216-228.
Thorpe, W. H. (1963). "Learning and Instinct in Animals." Methuen, London.
Preface
We began corresponding about co-editing a book on bird sounds in 1978,
excited by the enormous increase in evolutionary understanding and interpreta­
tion of communication systems since Robert Hinde's edited volume, Bird Vocal­
izations, appeared in 1969. The rapid mushrooming and splintering of ideas and
observations on animal communication were daunting, but we nevertheless
shared the belief that a representative and useful collection of writings on evolu­
tion and ecology of bird acoustics could be assembled. Our original intent was to
compile both taxonomic and conceptual reviews, but the meager knowledge of
acoustic signals of many important avian taxa made the first of these impossible.
Consequently, we solicited contributions from active researchers in bird acous­
tics, for chosen areas of evolutionary and behavioral ecology, and sought to
complement them with reviews of sound recording techniques, and sound pro­
duction, reception, and processing.
Volume 1 begins with several of these background chapters. The first dis­
cusses sound recording, makes certain recommendations, and points out com­
mon errors. The others outline some of the complex events and processes be­
tween sound production and behavioral response to sound. The remaining
chapters stand apart from the first ones, a gap that accurately reflects our poor
understanding of the processes which ultimately link individual physiological
responses to population-genetical changes, and to larger-scale evolutionary
trends. Bridging this gap will require diverse research endeavors and theorizing,
some of which are touched on in Chapters 5-9 of Volume 1 and Chapters 1-9 of
Volume 2.
The chapters are varied. They range from lengthy, well balanced, detailed
syntheses of subjects such as coding of species-specificity (Becker), indi­
viduality (Falls), and environmental acoustics (Wiley and Richards), etc., to
briefer more partisan explications of ideas and observations on dialects (Baker),
subsong (Marler and Peters), sexual selection (Catchpole), "moti­
vation-structural rules" (Morton), etc. Others present novel views and reviews
on doggedly troublesome concepts such as duetting (Farabaugh), vocal mimicry
(Baylis), and geographic variation (Mundinger). We have contributed chapters
xvii