Animal Communication Theory: Information and Influence

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Specificity and multiplicity in the recognition of individuals: implications for the evolution of social behaviour. Biological Reviews Alfonso Olalla and the ornithological exploration of Peru.

American Museum of Natural History Bulletin Trade-offs in the design of experiments. Journal of Comparative Psychology Acoustic and morphological identification of the sympatric cricket frogs Acris crepitans and A. Zootaxa Production and perception of communicatory signals in a noisy environment. Biology Letters 5: Signal transmission in natural environments. In Squire, L. Elsevier, Oxford. Signaller:receiver coordination and the timing of communication in Amazonian birds. Biology Letters 4: Signal detection and animal communication.

Publications: R. H. Wiley

Advances in the Study of Behavior Animal communication: signal detection. In Brown, K. Individuality in songs of Acadian flycatchers and recognition of neighbours. Is there an ideal behavioural experiment? Possibilities for error during communication by Neotropical frogs in a complex acoustic environment. Behavioral Ecology and Sociobiology Background noise from a natural chorus alters female discrimination of male calls in a Neotropical frog. Review: estimating the distance to a source of sound: mechanisms and adaptations for long-range communication.

Sexual selection and mating systems: trade-offs for males and females.

Apollonio, M. Festa-Bianchet, and D. Mainardi editors , Vertebrate Mating Systems. Steadman, L. Chadwick, and L. Social inertia in white-throated sparrows results from recognition of opponents. Acoustic interference limits call detection in a Neotropical frog Hyla ebraccata. Errors, exaggeration, and deception in animal communication.

Real ed. Second, signal traits are a classic system for studying how sexual selection works because of the increased strength and constancy of sexual selection compared to natural selection [ 6 ] and the greater potential for rapid trait divergence if traits and preferences are genetically linked [ 7 ]. The idea that sexual selection drives signal diversity, as first emphasized by West-Eberhard [ 7 ], has been an important yet controversial idea in biology.

For example, Seddon and colleagues [ 8 ] showed that sexual selection promotes trait divergence during speciation, while recent theoretical work [ 9 ] suggests that sexual selection might instead limit signal divergence in some contexts. A third explanation for disagreement over the role of social selection in driving signal diversity might be our lack of understanding of the interrelationships between different aspects of complex signals [ 10 — 12 ]. Early research into animal signals involved the use of standard color swatches or phonetic renderings of bird songs for example, the "whip-poor-will" call of the eastern whip-poor-will.

In the last 30 years, technological advances in, and falling prices of, devices to quantify signals numerically for example, portable spectrophotometers, high-speed digital cameras, or self-powered magnetic recorders [ 13 ] have driven an explosion of research into signal diversity in nature [ 1 ]. Now is an exciting time to be an evolutionary biologist, with vast amounts of life-history data collected through painstaking fieldwork [ 14 ], as well as acoustic for example, xeno-canto.

We are at the precipice of breakthroughs in the integration of these different data streams to answer new questions about the evolution of complex signals. However, despite recent advances in capturing signals in nature and making them readily available, researchers face new challenges with comparing highly divergent signals among species and analyzing them in an evolutionary framework. A new paper in this issue of PLOS Biology [ 16 ] uses novel analytical approaches to quantify diversity and richness for distinct aspects of courtship signals behavioral, acoustic, colorimetric and test hypotheses for how signal complexity itself evolves, both among species and among signal components see Fig 1.

The diverse birds-of-paradise BOPs are an excellent focal group for this work because they provide a classic example of how phenotypic and behavioral diversity is shaped by sexual selection. An integrative framework for studying the causes and consequences of signal diversity. Image credit : Chad M. A challenge with variable datasets like BOP signals is comparing different signals among species.

For example, how can we compare a whistle in one species to a chirp in another species? Ligon and colleagues [ 16 ] tackle this challenge by tabulating all aspects of signals for example, individual color patches or movements across 40 BOP species. Next, they use techniques borrowed from the field of information theory for example, color pixel clustering to calculate complexity for distinct axes of signal diversity acoustic, visual, and behavioral on a species-by-species basis.

For example, a species such as the paradise-crow Lycocorax pyrrhopterus with a uniformly glossy-black plumage would have a lower visual complexity score than the red bird-of-paradise Paradisaea rubra , a species with an iridescent green throat, red tail, yellow head, and brown back and wings.

Insights from statistical decision theory

This composite approach gets around the problem of incomparability in complex signal traits and lets the authors ask the question: How does signal complexity itself change over evolutionary timescales? Evolutionary tradeoffs involving reduced expression of one trait in response to elaboration of another trait are commonplace in nature. Darwin [ 17 ] first discussed the possibility of such tradeoffs in the context of signaling in birds.

Recent studies have provided mixed support for this prediction. For example, in cardueline finches, brighter species have simpler songs [ 18 ], while in Asian barbets, more colorful species have more complex longer songs [ 11 ]. These differences could reflect real biological differences among different clades or differences in methodology. By analyzing signal complexity in unprecedented detail, Ligon and colleagues [ 16 ] uncover concerted evolutionary increases in color and acoustic diversity, as well as between behavior and acoustic diversity, within BOPs.

That is, species with diverse acoustic signals also have diverse color patterns. These positive relationships in complexity among signal types provide evidence of sexual selection acting on an interrelated set of signal traits, which the authors describe as the "courtship phenotype" [ 16 ]. This work highlights a potential role for phenotypic integration that is, concerted evolutionary changes among traits in explaining the diverse signals we see in BOPs. Further behavioral and comparative work is needed to determine whether this pattern is unique to BOPs and their distinct evolutionary history or if positive correlations in complexity among signal types are more common in birds than previously recognized.

Form—function relationships are well studied in ecological traits [ 19 ] but less so in ornamental traits [ 20 , 21 ]. Whether signals evolve by natural or sexual selection, evolutionary changes in signals will depend on underlying physicochemical bases for signal production Fig 1 , such as the evolution of new pigment types [ 22 ], variation in hormones that affect signals [ 23 ], or anatomical changes in structures or transmission environments used for signaling [ 20 ].

Further studies on the proximate bases of signal traits will therefore be crucial to our understanding of how complex signals ultimately evolve and help answer longstanding questions about how signal diversity relates to diversity in signal-generating mechanisms.

Animal Communication Theory : Information and Influence

Exciting discoveries about how genetic mutations translate to brilliant signals, like red feather ornaments in birds, are now being made with next-generation sequencing technology and genome-wide association studies [ 24 ]. Comparative genomic approaches can further be used to identify target genes underpinning novel signals [ 3 ]. As access to large datasets continues to grow, researchers will face new challenges with identifying shared aspects of complex signals that is, signal homology that are comparable across species.

Perhaps individual aspects of complex songs or plumages are not comparable in different species, but certain patterns are for example, black and white bars, or repeated combinations of notes in a song , akin to how a few developmental precursors might give rise to diverse patterns through reaction-diffusion mechanisms in the context of developmental biology for example, fur patterns in mammals [ 25 ].

Answers to how signal variation reflects regulatory, genic, or even epigenetic changes within the genome will rely on novel approaches for rapid phenotyping along with new analytical tools for linking genomic changes with complex signal variation. Studying the evolution of single traits shared by a group of closely related species is relatively straightforward.

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Increases in the number of traits necessary to capture variation in complex signals bring new analytical challenges associated with multivariate datasets. For example, as a general rule of thumb, the number of traits should not be greater than the number of species used in a comparative analysis [ 26 ].

Recent methods for studying evolutionary trends in multivariate data that use distance-based methods [ 26 ] may get around this limitation. However, we are still left with the challenge of identifying comparable subunits shared by species, which is particularly difficult for complex animal signals.

Assessing complex signals on a species-by-species basis and deriving composite scores—for example, describing signal complexity [ 16 ] or plumage appearance [ 27 ]—is a good starting point for asking questions about how complex signals evolve.