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.

Research that integrates animal behavior theory with mechanics-including biomechanics, physiology, and functional morphology-can reveal how organisms accomplish tasks crucial to their fitness. Despite the insights that can be gained from this interdisciplinary approach, biomechanics commonly neglects a behavioral context and behavioral research generally does not consider mechanics. Here, we aim to encourage the study of "mechanoethology," an area of investigation intended to encompass integrative studies of mechanics and behavior. Using examples from the literature, including papers in this issue, we show how these fields can influence each other in three ways: 1) the energy required to execute behaviors is driven by the kinematics of movement, and mechanistic studies of movement can benefit from consideration of its behavioral context; 2) mechanics sets physical limits on what behaviors organisms execute, while behavior influences ecological and evolutionary limits on mechanical systems; and 3) sensory behavior is underlain by the mechanics of sensory structures, and sensory systems guide whole-organism movement. These core concepts offer a foundation for mehanoethology research. However, future studies focused on merging behavior and mechanics may reveal other ways by which these fields are linked, leading to further insights in integrative organismal biology.

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.

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.

Abstract
In the context of mate choice, males may vary continuously in their expression of assessment signals, typically reflecting information about variation in mate quality. Similarly, females may exhibit variation in mate preference, which could be due to differences in how individual females perceive signals. The extent to which perception varies across individuals, however, and whether differences in sensory physiology underlie perceptual differences is poorly understood. Carotenoid pigments create the orange-red coloration of many assessment signals, and they also play a role in color discrimination in many vertebrates via their presence in retinal oil droplets. Here, we link variation in oil droplet carotenoid concentration with the ability of female zebra finches (Taeniopygia guttata) to discriminate an orange-red color continuum that parallels variation in male beak color, a mate assessment signal. We have shown previously that zebra finch females perceive this color range categorically, meaning they label color stimuli from this continuum as belonging to two categories and exhibit better discrimination between colors from different categories as compared with equally different colors from within a category. We quantified behavioral color discrimination and R-type (red) cone oil droplet spectral absorption, a proxy for carotenoid concentration. Oil droplet absorption was strongly predictive of variation in behavioral color discrimination ability. In particular, higher carotenoid concentration in oil droplets correlated with increased discrimination of colors from different sides of the previously identified category boundary. These data show that differences in the sensory periphery can correlate with individual variation in perception of a signal-relevant color range.

Significance statement
Signal receivers vary in their preferences for signaling traits, but whether this is due to variation in how different receivers perceive signals is not well-understood. We show that variation between individual zebra finch females in perception of an orange-red continuum range correlates with the carotenoid concentration of retinal oil droplets. These data provide the first direct evidence that individual variation in oil droplet carotenoid concentration can lead to variation in color discrimination ability. Linking variation in signal-relevant color discrimination ability with variation in retinal physiology suggests a potential mechanism contributing to individual variation in signal assessment.

. 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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Measurements of energy use, and its scaling with size, are critical to understanding how organisms accomplish myriad tasks. For example, energy budgets are central to game theory models of assessment during contests and underlie patterns of feeding behavior. Clear tests connecting energy to behavioral theory require measurements of the energy use of single individuals for particular behaviors. Many species of mantis shrimp (Stomatopoda: Crustacea) use elastic energy storage to power high-speed strikes that they deliver to opponents during territorial contests and to hard-shelled prey while feeding. We compared the scaling of strike kinematics and energetics between feeding and contests in the mantis shrimp Neogonodactylus bredini. We filmed strikes with high-speed video, measured strike velocity, and used a mathematical model to calculate strike energy. During contests, strike velocity did not scale with body size but strike energy scaled positively with size. Conversely, while feeding, strike velocity decreased with increasing size and strike energy did not vary according to body size. Individuals most likely achieved this strike variation through differential compression of their exoskeletal spring prior to the strike. Post-hoc analyses found that N. bredini used greater velocity and energy when striking larger opponents, yet variation in prey size was not accompanied by varying strike velocity or energetics. Our estimates of energetics inform prior tests of contest and feeding behavior in this species. More broadly, our findings elucidate the role behavioral context plays in measurements of animal performance.

. 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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