Abstract
. The information an animal gathers from its environment, including that associated with signals, often varies continuously. Animals may respond to this continuous variation in a physical stimulus as lying in discrete categories rather than along a continuum, a phenomenon known as categorical perception. Categorical perception was first described in the context of speech and thought to be uniquely associated with human language. Subsequent work has since discovered that categorical perception functions in communication and decision-making across animal taxa, behavioral contexts, and sensory modalities. We begin with an overview of how categorical perception functions in speech perception and, then, describe subsequent work illustrating its role in nonhuman animal communication and decision-making. We synthesize this work to suggest that categorical perception may be favored where there is a benefit to 1) setting consistent behavioral response rules in the face of variation and potential overlap in the physical structure of signals, 2) especially rapid decision-making, or 3) reducing the costs associated with processing and/or comparing signals. We conclude by suggesting other systems in which categorical perception may play a role as a next step toward understanding how this phenomenon may influence our thinking about the function and evolution of animal communication and decision-making.

© 2020 by the University of Chicago. 0003-0147/2021/19702-60002$15.00. All rights reserved. Sensory systems are predicted to be adapted to the perception of important stimuli, such as signals used in communication. Prior work has shown that female zebra finches perceive the carotenoid-based orange-red coloration of male beaks—a mate choice signal—categorically. Specifically, females exhibited an increased ability to discriminate between colors from opposite sides of a perceptual category boundary than equally different colors from the same side of the boundary. The Bengalese finch, an estrildid finch related to the zebra finch, is black, brown, and white, lacking carotenoid coloration. To explore the relationship between categorical color perception and signal use, we tested Bengalese finches using the same orange-red continuum as in zebra finches, and we also tested how both species discriminated among colors differing systematically in hue and brightness. Unlike in zebra finches, we found no evidence of categorical perception of an orange-red continuum in Bengalese finches. Instead, we found that the combination of chromatic distance (hue difference) and Michelson contrast (difference in brightness) strongly correlated with color discrimination ability on all tested color pairs in Bengalese finches. The pattern was different in zebra finches: this strong correlation held when discriminating between colors from different categories but not when discriminating between colors from within the same category. These experiments suggest that categorical perception is not a universal feature of avian—or even estrildid finch—vision. Our findings also provide further insights into the mechanism underlying categorical perception and are consistent with the hypothesis that categorical perception is adapted for signal perception.

Animals often use assessment signals to communicate information about their quality to a variety of receivers, including potential mates, competitors, and predators. But what maintains reliable signaling and prevents signalers from signaling a better quality than they actually have? Previous work has shown that reliable signaling can be maintained if signalers pay fitness costs for signaling at different intensities and these costs are greater for lower quality individuals than higher quality ones. Models supporting this idea typically assume that continuous variation in signal intensity is perceived as such by receivers. In many organisms, however, receivers have threshold responses to signals, in which they respond to a signal if it is above a threshold value and do not respond if the signal is below the threshold value. Here, we use both analytical and individual-based models to investigate how such threshold responses affect the reliability of assessment signals. We show that reliable signaling systems can break down when receivers have an invariant threshold response, but reliable signaling can be rescued if there is variation among receivers in the location of their threshold boundary. Our models provide an important step toward understanding signal evolution when receivers have threshold responses to continuous signal variation.

. Although perception begins when a stimulus is transduced by a sensory neuron, numerous perceptual mechanisms can modify sensory information as it is processed by an animal's nervous system. One such mechanism is categorical perception, in which (1) continuously varying stimuli are labelled as belonging to a discrete number of categories and (2) there is enhanced discrimination between stimuli from different categories as compared with equally different stimuli from within the same category. We have shown previously that female zebra finches (
. Taeniopygia guttata
. ) categorically perceive colours along an orange–red continuum that aligns with the carotenoid-based coloration of male beaks, a trait that serves as an assessment signal in female mate choice. Here, we demonstrate that categorical perception occurs along a blue–green continuum as well, suggesting that categorical colour perception may be a general feature of zebra finch vision. Although we identified two categories in both the blue–green and the orange–red ranges, we also found that individuals could better differentiate colours from within the same category in the blue–green as compared with the orange–red range, indicative of less clear categorization in the blue–green range. We discuss reasons why categorical perception may vary across the visible spectrum, including the possibility that such differences are linked to the behavioural or ecological function of different colour ranges.
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. Safe and effective conflict resolution is critical for survival and reproduction. Theoretical models describe how animals resolve conflict by assessing their own and/or their opponent's ability (resource holding potential, RHP), yet experimental tests of these models are often inconclusive. Recent reviews have suggested this uncertainty could be alleviated by using multiple approaches to test assessment models. The mantis shrimp
. Neogonodactylus bredini
. presents visual displays and ritualistically exchanges high-force strikes during territorial contests. We tested how
. N. bredini
. contest dynamics were explained by any of three assessment models—pure self-assessment, cumulative assessment and mutual assessment—using correlations and a novel, network analysis-based sequential behavioural analysis. We staged dyadic contests over burrow access between competitors matched either randomly or based on body size. In both randomly and size-matched contests, the best metric of RHP was body mass. Burrow residency interacted with mass to predict outcome. Correlations between contest costs and RHP rejected pure self-assessment, but could not fully differentiate between cumulative and mutual assessment. The sequential behavioural analysis ruled out cumulative assessment and supported mutual assessment. Our results demonstrate how multiple analyses provide strong inference to tests of assessment models and illuminate how individual behaviours constitute an assessment strategy.
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. Cleaner shrimp and their reef fish clients are an interspecific mutualistic interaction that is thought to be mediated by signals, and a useful system for studying the dynamics of interspecific signalling. To demonstrate signalling, one must show that purported signals at minimum (a) result in a consistent state change in the receiver and (b) contain reliable information about the sender's intrinsic state or future behaviour. Additionally, signals must be perceptible by receivers. Here, we document fundamental attributes of the signalling system between the cleaner shrimp
. Ancylomenes pedersoni
. and its clients. First, we use sequential analysis of
. in situ
. behavioural interactions to show that cleaner antenna whipping reliably predicts subsequent cleaning. If shrimp do not signal via antenna whipping, clients triple their likelihood of being cleaned by adopting darker coloration over a matter of seconds, consistent with dark colour change signalling that clients want cleaning. Using experimental manipulations, we found that visual stimuli are sufficient to elicit antenna whipping, and that shrimp are more likely to ‘clean' dark than light visual stimuli. Lastly, we show that antenna whipping and colour change are perceptible when accounting for the intended receiver's visual acuity and spectral sensitivity, which differ markedly between cleaners and clients. Our results show that signalling by both cleaners and clients can initiate and mediate their mutualistic interaction.
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. Mantis shrimp strike with extreme impact forces that are deadly to prey. They also strike conspecifics during territorial contests, yet theoretical and empirical findings in aggressive behaviour research suggest competitors should resolve conflicts using signals before escalating to dangerous combat. We tested how
. Neogonodactylus bredini
. uses two ritualized behaviours to resolve size-matched contests: meral spread visual displays and telson (tailplate) strikes. We predicted that (i) most contests would be resolved by meral spreads, (ii) meral spreads would reliably signal strike force and (iii) strike force would predict contest success. The results were unexpected for each prediction. Contests were not resolved by meral spreads, instead escalating to striking in 33 of 34 experiments. The size of meral spread components did not strongly correlate with strike force. Strike force did not predict contest success; instead, winners delivered more strikes. Size-matched
. N. bredini
. avoid deadly combat not by visual displays, but by ritualistically and repeatedly striking each other's telsons until the loser retreats. We term this behaviour ‘telson sparring', analogous to sparring in other weapon systems. We present an alternative framework for mantis shrimp contests in which the fight itself is the signal, serving as a non-lethal indicator of aggressive persistence or endurance.
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