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978-0-521-87242-3 - Bird Song: Biological Themes and Variations, Second Edition

C. K. Catchpole and P. J. B. Slater
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chapter one

THE STUDY OF BIRD SONG

And your bird can sing

John Lennon Popular song

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978-0-521-87242-3 - Bird Song: Biological Themes and Variations, Second Edition
C. K. Catchpole and P. J. B. Slater
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the study of bird so ng

1.1 Introduction
This chapter is a brief introduction to the theory, terminology and tech-
niques used in the scientific study of bird songs. Although everyone may
assume that they do know what a bird song is, how does it differ from the
other sounds that birds make? There are calls, notes, syllables and phrases
to consider – and what are repertoires? Before we start using these words,
it is just as well to define them and become acquainted with a terminology
which can be confusing. Only then can we move on to consider the role of
song in the lives of birds and to review the many studies that have
attempted to shed some light upon it. Animal communication is a rapidly
expanding field, and at this early stage it is also useful to consider some of
the recent theoretical background. For example, what is Ôcommunication’
and how do we know it has occurred? What is Ôinformation’ and who
benefits from sending and receiving it? What are Ôsignals’ and why is
sound transmission particularly effective? Finally, recording, analysing
and experimenting with sounds is a highly technical field, currently being
revolutionised by the use of computers at various points. We will also
attempt to give the reader just a brief introduction to some of the more
important techniques and any recent developments. But first, let us set the
current study of bird song in its historical context.

1.2 History
Clearly, from references to it in music and in literature, particularly
poetry, people have been interested in bird song for a very long
time. But its detailed scientific study is a comparatively recent
phenomenon. One reason for this is that making a permanent record of
it, as we now do routinely on tapes or discs, was not easy until half a
century or so ago. Song can be so rapid and complicated that only with
such a permanent record that could be slowed down, repeated and ana-
lysed in various ways, is it possible to make a serious study of many aspects
of it.
Some interesting work was done before that time. The Hon. Daines
Barrington wrote a letter to the President of the Royal Society in 1773
recounting a variety of observations he had made. He established the
existence of song learning, for example, because he heard the song of

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978-0-521-87242-3 - Bird Song: Biological Themes and Variations, Second Edition
C. K. Catchpole and P. J. B. Slater
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SOME BASIC THEORY

a wren emanating from a house he was passing and, knowing how difficult
such birds were to keep in captivity, knocked on the door out of curiosity,
only to discover that the singer was a captive goldfinch. Presumably this
bird had been exposed to wren song at some stage and had picked it up.
At around the same time, in 1789, the great English parson and naturalist
Gilbert White described how birds previously known as willow wrens
could be separated by their songs into three separate species. These we
now call the willow warbler, the wood warbler and the chiff-chaff. Those
with a good ear were also able to detect that birds had repertoires of songs
and study the way these were strung together into sequences, as Craig
(1943) did with eastern wood pewee song, or that song could vary from
place to place, as found by Marler (1952) for the chaffinches singing in
different glens in the Scottish highlands.
The depth of such studies was severely limited, not just by lack of
the possibility of recording, except latterly on wax drums, but most
importantly by the lack of analytical equipment. The real revolution
came with the invention of the sound spectrograph, first used to provide
a visual representation of song by Thorpe in 1954. Such equipment was
not cheap, and therefore its use was somewhat restricted, but it still led
to a huge growth in studies of song. Today, equivalent visualisations of
song, together with many other forms of analysis, can be carried out
using a variety of computer packages at a fraction of the cost. The detailed
study of bird song is within the scope and budget of many laboratories and
even amateurs: as a result the subject is advancing with great strides.
Thanks to these very powerful techniques, there are now few areas of
animal behaviour research that have not been illuminated by studies of
bird song.

1.3 Some basic theory

1.3.1 Signals and communication
When a bird sings, it produces a sound which serves to communicate with
other members of the same species. Because the song is a special structure
used solely in communication, we call it a signal. But how do we know
whether communication has occurred? It is generally held that if the signal
modifies the behaviour of the receiving animal then we can infer that
communication has taken place (Slater 1983c). For example, if we play

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C. K. Catchpole and P. J. B. Slater
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the study of bird so ng

back a tape recording of a male great tit song to another male, we may
cause the second male to respond by approaching the speaker and dis-
playing aggressively. As the song appears to have modified his behaviour,
we are entitled to conclude that communication has occurred. This is a
somewhat restricted definition of communication, as it relies upon a
behavioural response and thus excludes passive signal detection by the
receiver. For example, if we repeated the experiment on another great tit
and obtained no response, it may be that the great tit had heard the song
but decided for some reason not to respond. Such behaviour may often
occur, so we must temper our definition with caution and also be sure to
carry out a number of experiments before we draw any firm conclusions.
Although it may be relatively easy to demonstrate that communication
takes place, it is much more difficult to suggest why it has evolved. Early
theories emphasised the benefits that might accrue to both the sender and
the receiver (e.g. Smith 1977) and saw communication as a sharing of
information between individuals to their mutual advantage. Modern ethol-
ogists are much more inclined to view communication as the outcome of
conflict rather than cooperation between sender and receiver.
Krebs & Dawkins (1984) examined how different kinds of signals may
result from the benefits that accrue to senders and receivers. Sometimes,
cooperation rather than conflict is involved, and they suggest that a system
which benefits both receiver and sender would give rise to the evolution of
relatively quiet, inconspicuous signals. For example, a great tit may give
an alarm call to warn its fledglings that a sparrowhawk is approaching. The
call should be loud enough to reach the fledglings but not loud enough to
reach the hawk and give away the position of the caller. To be able to hear
the call, the fledglings should develop sensitive hearing, and so a coevolu-
tionary process will lead to the production of calls that are Ôcost-minimiz-
ing conspiratorial whispers’. The fascinating story of the evolution of
alarm calls, and their possible detection by predators, will be discussed
in detail later in the book.
But where the interests of sender and receiver conflict, as in a territorial
dispute, there will be a different kind of coevolution. Here, Krebs &
Dawkins (1984) suggest that there is an evolutionary arms race between
Ômanipulation’ by the sender and Ôsales resistance’ by the receiver. The
male great tit this time sings as long and loud as he can manage, simply to
force his message across. As we will see in later chapters, loudness and
repetition are a particular feature of the songs of males when defending

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C. K. Catchpole and P. J. B. Slater
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SOME BASIC THEORY

their territorial boundaries. Although sales resistance may occur among

rival males, there are even more compelling reasons why females should
be wary of male signals. If a listening male makes a mistake, he may just
waste energy in a display or a fight, but if a listening female chooses a male
of the wrong species, or one of inferior quality, she may pay a severe
penalty in reduced breeding success. We will also see in later chapters that
there is now considerable evidence that females have been selected for fine
discrimination of both quantity and quality of male songs.
So far, we have assumed that the signals transmitted give reliable
information from sender to receiver. At this stage, we should mention that
the word Ôinformation’ is used in two different ways. Technically, informa-
tion theory considers it to be Ôa reduction in uncertainty’ about the senderÕs
future behaviour on the part of the receiver. In other words, information is
said to have passed from sender to receiver when the senderÕs behaviour
becomes more predictable to the receiver (Halliday 1983). When informa-
tion is transmitted between birds, it is generally about something quite
precise, such as species, sex, identity, likely next action, and so on. There are
some grounds for expecting that receivers will be selected to detect unre-
liable or false information. Zahavi (1979, 1987) has suggested that only
signals which are honest indicators of size, strength or motivation should
evolve. One reason for this is that many signals are costly to produce, and so
it is difficult, for example, for a smaller, weaker animal to cheat or bluff the
receiver into accepting it as a larger, stronger rival or mate.
The view that, because of costs incurred by the sender, evolution has
generally favoured Ôhonest advertising’ in communication has now
become widely accepted. However, Dawkins & Guilford (1991) have
pointed out that receivers also pay costs when assessing signals. If the
costs of long, detailed assessment are high in relation to the value of the
extra information gained, then receivers might settle for cheaper, less
reliable signals. If so, the receiver may be open to being bluffed, cheated
and manipulated to the senderÕs advantage. In their review, Krebs &
Davies (1993) suggest that such coevolutionary arms races between sender
and receiver may have two different end-points. In one, the outcome is the
evolution of honest signalling, but in the other we may find unreliable
signals have evolved as one animal attempts to manipulate the behaviour
of the other. Recent developments in this field have also been reviewed by
Maynard Smith & Harper (2003), and by Searcy & Nowicki (2005), who
focus particularly on the evolution of reliability and deception, using bird

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C. K. Catchpole and P. J. B. Slater
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the study of bird so ng

Table 1.1. A comparison of difference sensory channels of communication

Acoustic Visual Chemical Tactile

Nocturnal use Good Poor Good Good

Around objects Good Poor Good Poor
Range Long Medium Long Short
Rate of change Fast Fast Slow Fast
Locatability Medium Good Poor Good
Energetic cost Low Low Low Low
Modified from Alcock 1989.

song as a key example. The latter point out that reliability requires there to
be a correlation between some aspect of the signal and some attribute of
the signaler that the receiver benefits from knowing about. Hence the
receiver benefits from assessing the signal rather than ignoring it. Deceit
requires not only that the correlation between signal and attribute be
broken, but that the signaler benefits from that breakdown. Searcy &
Nowicki (2005) present a detailed review of this complex area and discuss
how opinions about the prevalence of reliability and deception in animal
communication have changed over the years.
The apparent coevolutionary arms race between senders and receivers
involves many different aspects of communication, which we will be
considering throughout this book. It is important to emphasise that we
will not just consider the signal (song) itself, but how it is transmitted
through the environment, how it is perceived by receivers and, in partic-
ular, how males and females react to both natural and experimental signals.

1.3.2 Why sound?

Sound is only one of several channels of communication that are open to
birds, and the advantages and disadvantages of the different channels have
been summarised in Table 1.1. In general, birds have rather a poorly
developed olfactory system, and so this method is less important than
the main channels of sound or vision. This contrasts with mammals, where
olfaction is a very important method of communication. Olfaction is rather
less important to humans, as their tiny noses indicate, and, like birds,
humans rely particularly upon sound and vision. There is no doubt that

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C. K. Catchpole and P. J. B. Slater
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SOME BASIC THEORY

visual signalling is of great importance to birds, as indicated by their

elaborate plumage and coloration and as seen in their eye-catching visual
displays. What then are the particular advantages of sounds, especially
when compared to visual signals?
Visual signals have several disadvantages, for example in darkness or
poor light. But bad conditions for visual signalling can occur at any time in
dense habitats such as forest or reeds and when animals move out of view
behind objects. Try looking for a small bird as it moves through the
canopy. Now you see it – now you donÕt! But if it calls or sings you
can always hear it, long after it moves out of sight. Sound travels in all
directions, it can penetrate Ôthrough’ or Ôround’ objects, and it travels over
long distances. Sound is an ideal method for communicating over long
distances, and although birds also call softly to each other, their songs are
often loud and can carry for several kilometres. How natural selection may
have acted to Ôshape’ song structures for optimal transmission through
different habitats is one of many topics we will discuss later in this book.
Other advantages stem mainly from the rapid and transient nature of
sound communication. A song or call is only produced when needed, and
large amounts of information can be transmitted rapidly and efficiently
through the sound channel. There might perhaps be one disadvantage if,
as has sometimes been suggested, singing is very costly in terms of ener-
getics. A song has to be Ômade’ each time it is produced, and some birds
sing thousands of songs per day. However, recent evidence suggests that
this is not the case and that song is comparatively cheap to produce
compared, for example, with the cost to a bird of hopping or flying around
its cage (e.g. Oberweger & Goller 2001, Ward et al. 2004). It does there-
fore seem that the many advantages of sound communication rather easily
outweigh its costs. Birds, like humans, are intensely vocal creatures, and
communication by sound has come to play a central role in their lives.

1.3.3 Songs, calls and terminology

Bird vocalisations can be divided into songs and calls. The distinction is
both traditional and arbitrary, but as these terms are still retained in the
literature we must attempt some clarification. There is also a taxonomic
reason for the distinction. One particular group, the oscines, were origi-
nally separated from the rest of the order Passeriformes, primarily on the
number and complexity of their syringeal muscles. As these birds

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C. K. Catchpole and P. J. B. Slater
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generally produced more complicated sounds or Ôsongs’, the oscines

became known as Ôthe true songbirds’. But, as we will see later, the differ-
ences between oscines and sub-oscines may have more to do with how
they learn their songs (and the underlying brain structure) rather than with
the actual complexity of their vocalisations.
As this book is largely devoted to the study of songs, it is only fair that
we should attempt a definition. In general, Ôsongs’ tend to be long, com-
plex, vocalisations produced by males in the breeding season. Song also
appears to occur spontaneously and is often produced in long spells with a
characteristic diurnal rhythm. But to these features there are innumerable
exceptions. Especially in the tropics, it is common for females to sing as
well as males, and both sexes may do so throughout the year even though
breeding only occurs during a restricted period (e.g. Langmore 1998).
Even in temperate regions, song may occur well before egg-laying and
there is also often a bit of a resurgence in the autumn. In the European
robin, for example, song may be heard in every month of the year and in
the winter it is produced by both males and females singing on separate
territories (Lack 1946). However as far as complexity is concerned, it is not
easy to generalise and, as we shall see in Chapter 8, species differ enor-
mously in how varied their songs are. There are even songbirds that
appear not to ÔsingÕ at all, but the simple ÔcheepingÕ of a male house
sparrow on a rooftop may fulfil the same function so that it is, in effect,
a very simple song.
What then are calls? ÔCalls’ tend to be shorter, simpler and produced by
both sexes throughout the year. Unlike songs, calls are less spontaneous
and usually occur in particular contexts which can be related to specific
functions such as flight, threat, alarm and so on. As with the house sparrow
example, there are obviously areas of overlap between simple song and
complex calls, and plenty of exceptions to the criteria we have presented.
But in general, ornithologists and ethologists recognise these distinctions
and continue to find them useful. Why the oscines have evolved such
complex songs, and a special brain pathway to learn them, is one of the
central themes of this book.
Having stated that one of the main characteristics of most songs is their
complexity, we have a number of other categories and units to define. Most
birds have more than one version of their species song, and some have
many. For example, most male chaffinches have more than one version of
their song and will repeat one several times in a bout of singing, and then

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C. K. Catchpole and P. J. B. Slater
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Fig. 1.1. Sonagrams of

two different song types
from the repertoire of a
male chaffinch, also
illustrating divisions into
phrases, syllables and
elements (from Slater &
Ince 1979). As explained
in Section 1.3.3,
sonagrams are plots of
frequency against time
with the trace being dark
where there is energy at
that particular point, so
giving a visual
representation of the
pattern of sound.

switch to another. Each version is called a song type, and the male chaf-
finch is said to have a repertoire of song types (see Fig. 1.1, and the more
detailed discussion of repertoires in Chapter 8).
Moving our analysis to a more detailed level, we can also see that each
chaffinch song consists of a number of distinct sections. These are called
phrases, and each phrase consists of a series of units which occur together
in a particular pattern. Sometimes, the units in a phrase are all different, as
in the end phrase shown in Fig. 1.1. The units themselves are usually
referred to as syllables. Syllables can be very simple or quite complex in
their structure. When complex, they are constructed from several of the
smallest building blocks of all, called elements or notes (but the latter is
usually avoided because of its musical connotations). One definition of an
element is simply a continuous line on a sonagram, as illustrated in Fig. 1.1.
Songs, syllables and elements can also be defined by the time intervals
which separate them, intersong intervals are the longest, and so on down-
wards. Because of the great variety of form and structure in songs, indi-
vidual workers often use and define their own terms, which may be slightly
different from those given above. These can only serve as a general guide
for this book, for there can be linguistic as well as technical problems with

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C. K. Catchpole and P. J. B. Slater
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definitions. For example, in German there are two different words for
Ôsong’: Ôgesang’ means the song of a particular species, whereas Ôstrophe’
means a particular delivery of a song. This last word is now often adopted
in English to refer to a single rendition.

1.4 Some basic techniques

1.4.1 Observing
The very idea of Ôobserving sounds’ seems like a contradiction in terms.
However, if the main objective is to determine what possible functions
a sound has, then this is where to start. Currently, it has become fashion-
able in many branches of modern biology to construct a hypothesis,
perhaps even a model, and then test selected predictions by experiment.
Naturally, this book is full of such examples, as experiments have played a
leading role in the scientific study of bird sounds. But to formulate an
appropriate hypothesis or model, a period of observation should first be
undertaken. This should preferably be a thorough field study which relates
the singing bird to its habitat, to its other behaviour and to its general life
history.
The experimenter may have rather less enthusiasm for this phase,
regarding the necessary field work as difficult, dull and somewhat old-
fashioned. However, it is vitally important for several reasons. For exam-
ple, it will provide an accurate source of basic information from which a
proper hypothesis can be constructed. It should reveal such essential
information as when, where and to whom the bird sings. The first clues
as to the probable functions of song invariably come from simple, con-
textual observations in the field. Does a male sing only in his territory?
Does he countersing against rival males? Does he stop singing when he
has paired with a female? These are very basic questions, and their answers
will help to give initial clues to function and will allow appropriate hypoth-
eses to be formulated. Does the male sing only at dawn? Does he stop
when his mate appears? Does he sing more in her fertile period? More
precise questions such as these can also be answered by careful observa-
tions and may lead to the eventual design of suitable playback experiments
to test more detailed functional hypotheses. Nor need the modern field
worker feel too old-fashioned. The traditional note-book can be replaced
by an electronic one, and a number of software packages will allow a full,

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