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Do Animal Communication Systems Have Phonemes?

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Daniel Bowling at Stanford University
Daniel Bowling
W Tecumseh Fitch
W Tecumseh Fitch
W Tecumseh Fitch
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Abstract

Biologists often ask whether animal communication systems make use of conceptual entities from linguistics, such as semantics or syntax. A new study of an Australian bird species argues that their communication system has phonemes, but we argue that imposing linguistic concepts obscures, rather than clarifyies, communicative function.
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Spotlight
Do Animal
Communication
Systems Have
Phonemes?
Daniel L. Bowling,
1
and
W. Tecumseh Fitch
1,
*
Biologists often ask whether ani-
mal communication systems make
use of conceptual entities from
linguistics, such as semantics or
syntax. A new study of an Austra-
lian bird species argues that their
communication system has pho-
nemes, but we argue that impos-
ing linguistic concepts obscures,
rather than clarifyies, communica-
tive function.
Human spoken language is fundamen-
tally combinatorial. Speech sounds
combine to form words, and words
combine to form phrases and senten-
ces. The flexibility of these combinations,
transforms our limited-precision motor
and perceptual systems into an
unbounded system of thought and com-
munication. Wilhelm von Humboldt char-
acterized this as ‘making infinite use of
finite means’[1].
Linguists tradtionally separate the com-
binatorial nature of language, into two
levels, each operating on a different
time-scale [2]. At the shorter phonologi-
cal level, meaningless sounds are
combined to form meaningful words
(e.g., /c/+/æ/+/t/ = ‘cat’). At the longer
syntactic level, words are combined to
form sentences (‘cats purr’). Research
on animal communication often makes
use of such linguistic terminology, charac-
terizing systems as primarily phonological
or primarily syntactic, sometimes drawing
parallels between specific elements of ani-
mal vocalization and particular linguistic
concepts.
A new paper by Engesser and colleagues
[3] provides an interesting example: they
propose that certain elements of chest-
nut-crowned babbler (Pomatostomus
ruficeps) vocalizations behave like human
phonemes (Box 1). They focus on two
tonal call elements, ‘A’and ‘B’, differenti-
ated by pitch contour. These elements
occur naturally as part of behavior associ-
ated with flight (in the sequence ‘AB’) and
prompting begging during nestling provi-
sioning (in the sequence ‘BAB’). Using
playback experiments with captured wild
birds, the authors demonstrate that the
prompt call sequence BAB stimulates
changes in listener behavior that are not
observed in response to the B element
alone or in response to the flight call
sequence AB. Accordingly, they argue
that the addition of the B element to the
sequence AB changes the meaning of the
call, just as an additional phoneme
changes the meaning of a word (e.g.,
/b/+ ‘it’=‘bit’).
This is an intriguing demonstration of a
novel form of combinatoriality in animal
communication. However, several key dif-
ferences between the elements of babbler
vocalization and human phonemes sug-
gest that a parallel is incomplete. First, the
babbler system is not very generative.
Neither A nor B occurs in the babbler
repertoire outside of flight and prompt
calls, whereas human phonemes are
widely distributed throughout a language.
Second, A and B are acoustically isolated
(separated by silences), whereas most
human phonemes are fused together into
unbroken syllables. Third, the difference
between AB and BAB is a case of pres-
ence or absence (like ‘cat’versus ‘at’),
whereas human phonemic contrasts are
typically discriminative (like ‘cat’vs ‘bat’).
While these superficial issues might be
argued to represent matters of degree
rather than kind, a deeper issue arises
when the term ‘phoneme’is applied out-
side human language. In linguistics, a pho-
neme is defined as the smallest unit in the
sound system of a language that serves to
distinguish meaning [4]. Because this def-
inition relies on ‘meaning’–a subjectively
informed concept that currently defies
neurobiological analysis –it cannot be
transparently applied to other species.
Attempts to shoehorn elements of animal
communication into the phoneme con-
cept quickly lead to questions that do
not allow empirical analysis, such as what
do babbler vocalizations mean. In human
language, the same phoneme string can
be uttered with myriad different intona-
tions and flourishes, conveying subtly dif-
ferent emotional and pragmatic intentions
(e.g., saying ‘no’can convey displeasure,
sarcasm, sincerity, uncertainty, etc.). The
lack of an objective definition for meaning
renders parallels between animals and
humans problematic, if not theoretically
misleading. Similar issues arise when
other linguistic concepts are applied to
animal communication. For example, the
distinction between phonology and syntax
is also dependent on meaning. Given that
our access to meaning in animal commu-
nication is fundamentally limited to what
we can infer from behavioral observation’
–we have no introspective access as for
human language –characterizing certain
levels of organization as phonological and
others as syntactic risks anthropomorphi-
cally obscuring rather than clarifying how
animal communication systems work.
These issues aside, Engesser et al.’s[3]
experimental results provide new evi-
dence that nonhuman animals, can com-
bine acoustically differentiable elements
productively, and that different sequences
can affect listener behavior in different
ways. Similar abilities have been reported
in mammals and other birds. For example,
field studies with Campbell's monkeys
(Cercopithecus campbelli) have shown
that boom calls are associated with group
cohesion and movement, and krak-oo
calls with nonspecific predator activity,
but sequences of booms followed by
sequences of krak-oos occur almost
exclusively with falling trees or branches
[5,6]. Among birds, black-capped chick-
adees (Poecile atricapillus) have been
TICS 1486 No. of Pages 2
Trends in Cognitive Sciences, September 2015, Vol xx. No. x 1
TICS 1486 No. of Pages 2
shown to alter the number of trailing dee
elements in their eponymous chick-a-dee
alarm calls as a function of predator size,
responding with more dees to more dan-
gerous predators [7]. Thus, call sequen-
ces appear to be a level of organization
worthy of further attention in animal com-
munication and playback studies, which
have typically focused mostly on single
calls: This is true regardless of whether
we label elements as phonemes, or cate-
gorize systems as phonology or syntax.
Finally, this new work on babblers marks
what we hope will be a promising resur-
gence of interest in ‘phonological’aspects
of animal communication which, after a
flurry of interest in the 1970s, has largely
languished (cf. [8]). Research in animal
communication focusing on issues of syn-
tax or meaning often neglects equally fas-
cinating questions about combinatoric
power at the level of acoustically differen-
tiable units. Some of the most complex
and generative animal communication
systems known to science are traditionally
termed ‘song’(e.g., birdsong or hump-
back whale song), suggesting that paral-
lels with music (which lacks propositional
meaning) may be more profitable than
parallels with language. Accordingly,
these systems may provide a richer
source of insight for understanding how
combinatoric communication systems
can evolve than do such ‘meaningful’calls
as alarm or food calls. Thus, parallels
between animal communication and
music or phonology may ultimately prove
more useful for understanding the evolu-
tion of communication than parallels with
syntax or semantics.
1
Department of Cognitive Biology, University of Vienna,
Vienna, Austria
*Correspondence: tecumseh.fitch@univie.ac.at (W.T. Fitch).
http://dx.doi.org/10.1016/j.tics.2015.08.011
References
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Box 1. Definition of Terms
Phones and Phonemes: Linguists define a phone as the smallest perceptually distinguishable segment of
sound in a stream of speech (e.g., the /c/ in ‘the cat meowed’). The concept of a phoneme was initially
motivated by a desire to understand patterns of organization among the tremendous diversity of phones
heard in languages. The traditional method of demonstrating phonemes is to find word pairs (e.g., ‘pat/bat’),
where a minimal change in the phone results in a different meaning. If it does, the sounds (/p/ versus /b/) are
considered to be phonemes. If it doesn’t, the new sound is considered an allophonic variant or allophone
[4,9].
Combinatoriality and Compositionality: Combinatoriality is a property by which new compounds are built
out of smaller parts. This definition is independent of meaning. Many animal communication systems are
clearly combinatorial. Compositionality is a property of human language whereby semantic units (like words)
are combined into larger compounds (phrases and sentences) whose composite meanings are functions of,
but not wholly determined by, the independent units. By present knowledge, no animal communication
system has this property. The eminent birdsong biologist Peter Marler addressed this issue with his
distinction between phonological syntax, which is combinatorial, and lexical syntax, which is combinatorial
and compositional [10].
2Trends in Cognitive Sciences, September 2015, Vol xx. No. x
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... Numerous bird and mammal species demonstrate calling and the integration of vocal cultures into key behaviours [270][271][272][273][274][275][276][277][278][279][280][281]. Increasingly, the complexity of animal call repertoires is being realized. ...
... The vocal production of primates and terrestrial mammals is now known to be greater and more flexible than had been previously appreciated [274]. Moreover, the emotional underpinning of vocal mechanisms is increasingly being recognized [270][271][272][273][274]. The abilities to project affective states as vocal signals and threat-associated alarm calls are only two examples of the flexible use of acoustics to portray a caller's state. ...
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Bibliogr. s. 448-452
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