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Environmental influences on acoustic and electric animal communication.

Animal communication in any sensory modality is influenced by environmental constraints on signal transmission and detection. Acoustic signals are subject to frequency-dependent attenuation, temporal degradation, and masking by environmental noise; these factors constrain the spectral and temporal structures of long-range sound signals. Signals produced by electric fish are largely free of such physical environmental constraints on their spectral and temporal structures, but experience severe attenuation due to geometrical spreading. This effectively limits electric communication to short ranges, whereas acoustic signals are more flexible in the range over which they are effective.

Acoustics↗

Signalers and receivers in animal communication.

In animal communication natural selection favors callers who vocalize to affect the behavior of listeners and listeners who acquire information from vocalizations, using this information to represent their environment. The acquisition of information in the wild is similar to the learning that occurs in laboratory conditioning experiments. It also has some parallels with language. The dichotomous view that animal signals must be either referential or emotional is false, because they can easily be both: The mechanisms that cause a signaler to vocalize do not limit a listener's ability to extract information from the call. The inability of most animals to recognize the mental states of others distinguishes animal communication most clearly from human language. Whereas signalers may vocalize to change a listener's behavior, they do not call to inform others. Listeners acquire information from signalers who do not, in the human sense, intend to provide it.

Affect↗

Some general comments on the evolution and design of animal communication systems.

Animal communication systems have evolved so that individuals can make decisions based upon the behaviour, physiology or morphology of others. Receiving mechanisms probably evolve to increase the efficiency and reliability of information reception whereas signals probably evolve to increase the efficiency of communication and reliability of manipulation of the receiving individual to the benefit of the emitter. The minimum requirement for clear reception suggests that any study of the evolution and design of communication systems must consider the factors that affect the quality of the received and processed signal. Critical information is needed about how the signal is generated and emitted, how it fares during transmission through air, water or substrate, how it is received and processed by the receiver's sensory and cognitive systems, and the factors which affect the fitness consequences of alternative ways of reacting to the information contained in the signal. These should allow predictions about the kinds and forms of signals used by animals signalling under known conditions. Phylogenetic history, and the geological time a clade spends in different signalling environments, will also affect signal evolution, and hence the success of predictions about signal design. We need to use methods of many different biological fields to understand the design and evolution of signals and signalling systems.

Animal Communication↗

Is there any intermediate stage between animal communication and language?

Animal communication and human language have fundamental differences in their structures and functions. Furthermore, there is no living species demonstrating an intermediate stage of language evolution. Thus, we have difficulty in finding characteristics attributable to a communication system which can already be considered as a starting point for linguistic evolution. However, some findings coming from neurolinguistic research give us the opportunity to suppose that varying and arranging linguistic elements can be detached from other grammatical functions. Further information in this direction comes from apes' language-teaching experiments; namely bonobos (Pan paniscus) are able to understand and produce differences in meaning by varying word arrangements. Based on these results one can suppose that an acoustic signal system, which possesses discrete units for variable use, might be very ancient and might exist independent and prior to a more advanced language state. In the natural setting, acoustic territorial marking behaviour is exposed to selection pressure to elaborate sign systems built up from discrete, variable units. In addition to the well-known territorial bird songs, some monkey species and all species of lesser apes have territorial songs fitting these criteria. The analyses of the so-called long calls in chimpanzees and bonobos make it likely that the group-living great apes preserved the ability to create syntactically different calls, which would be developed by requirements of social life. A call repertoire emerged in these species, which contained a large number of call variants at group level available for each group member via social learning. This type of animal call is different from ordinary animal communication; it shows some features of human language. It can represent an intermediate stage between animal communication and language, and communication systems similar to this one can be considered as a starting point or first stage of language evolution.

Animal Communication↗

A simple frequency-scaling rule for animal communication.

Different animals use widely different frequencies for sound communication, and it is reasonable to assume that evolution has adapted these frequencies to give greatest conspecific communication distance for a given vocal effort. Acoustic analysis shows that the optimal communication frequency is inversely proportional to about the 0.4 power of the animal's body mass. Comparison with observational data indicates that this prediction is well supported in practice. For animals of a given class, for example mammals, the maximum communication distance varies about as the 0.6 power of the animal's mass. There is, however, a wide spread of observed results because of the different emphasis placed upon vocal effort in the evolution of different animal species.

Animal Communication↗

Economic models of animal communication.

Many models of animal signal evolution fail to incorporate an explicit strategy for receivers prior to the evolution of signals. When reasonable assumptions are made for such strategies, we have shown that there is a minimal accuracy of signal coding that is required before receivers should attend to signals (Bradbury & Vehrencamp 1998, Principles of Animal Communication). Depending upon the relative payoffs of correct and incorrect decisions by receivers, this minimal accuracy can be quite high. Here we use this result to explain why so many signals appear to be traits that provided useful information to receivers before becoming ritualized into signals. Our model also supports one prediction of sensory drive models: that latent preferences may selectively favour some signal precursors over others. However, it imposes a serious constraint on sensory drive by requiring that there be sufficient benefits to a receiver to compensate for the costs of disrupting the optimal receiver strategy used before exploitation. Finally, we discuss the overlap between signal honesty and accuracy and show how senders that completely disagree with receivers about appropriate receiver decisions may still benefit by providing moderately honest and accurate signals. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Quantitative tools for comparing animal communication systems: information theory applied to bottlenose dolphin whistle repertoires.

Comparative analysis of nonhuman animal communication systems and their complexity, particularly in comparison to human language, has been generally hampered by both a lack of sufficiently extensive data sets and appropriate analytic tools. Information theory measures provide an important quantitative tool for examining and comparing communication systems across species. In this paper we use the original application of information theory, that of statistical examination of a communication system's structure and organization. As an example of the utility of information theory to the analysis of animal communication systems, we applied a series of information theory statistics to a statistically categorized set of bottlenose dolphin Tursiops truncatus, whistle vocalizations. First, we use the first-order entropic relation in a Zipf-type diagram (Zipf 1949 Human Behavior and the Principle of Least Effort) to illustrate the application of temporal statistics as comparative indicators of repertoire complexity, and as possible predictive indicators of acquisition/learning in animal vocal repertoires. Second, we illustrate the need for more extensive temporal data sets when examining the higher entropic orders, indicative of higher levels of internal informational structure, of such vocalizations, which could begin to allow the statistical reconstruction of repertoire organization. Third, we propose using 'communication capacity' as a measure of the degree of temporal structure and complexity of statistical correlation, represented by the values of entropic order, as an objective tool for interspecies comparison of communication complexity. In doing so, we introduce a new comparative measure, the slope of Shannon entropies, and illustrate how it potentially can be used to compare the organizational complexity of vocal repertoires across a diversity of species. Finally, we illustrate the nature and predictive application of these higher-order entropies using a preliminary sample of dolphin whistle vocalizations. The purpose of this preliminary report is to re-examine the original application of information theory to the field of animal communication, illustrate its potential utility as a comparative tool for examining the internal informational structure of animal vocal repertoires and their development, and discuss its relationship to behavioural ecology and evolutionary theory. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

The Umwelt and its relevance to animal communication: introduction to special issue.

This introduction applies J. von Uexküll's (1934/1957) concept of the Umwelt to the study of animal communication, particularly as it pertains to studies presented at a recent workshop on animal communication in the context of the environment. The environment is conceived broadly in the articles that follow, including the many physical and social environments in which an animal may find itself. The Umwelt concept is briefly expanded here to include also the personal microenvironment of the signaler in which the signal is embedded into the suite of concurrent nonsignaling behaviors of the individual. Other animals may even infer aspects of the signaler's own immediate Umwelt by noticing accompanying attentional cues such as the direction of eye gaze. In this way, part of the Umwelt can be accessible to companions, facilitating the communication process.

Animal Communication↗

ANIMAL COMMUNICATION.

Semiotics and ethology have converged in a new behavioral science, zoosemiotics. Those who are interested in the theoretical analysis of the complex problems of non-verbal behavior that arise where these two disciplines interact aim to treat comprehensively animal communication systems by the aid of representations that have proved illuminating in the study of sentences of human language. Students of zoosemiotics are concerned with codes and messages much as linguists are concerned with competence, or language, and performance, or speech. They thus face the twin tasks of constructing a model for the addresser to specify how a message is encoded and transformed into a signal carried by a variety of channels to the addressee; and of constructing a model for the addressee to specify the ways in which animals utilize their knowledge of their code to recognize the messages they receive. Finally, they assess the context of the communicative event in the hope of dissecting that which is relevant to the selection process from the rest of the background, a program for which there is as yet neither a procedural eliciting technique nor a satisfactory theoretical solution in sight.

Animal Communication↗