The mechanisms and functions of reversible colour change in arthropods are highly diverse despite, or perhaps due to, the presence of an exoskeleton. Physiological colour changes, which have been recorded in 90 arthropod species, are rapid and are the result of changes in the positioning of microstructures or pigments, or in the refractive index of layers in the integument. By contrast, morphological colour changes, documented in 31 species, involve the anabolism or catabolism of components (e.g. pigments) directly related to the observable colour. In this review we highlight the diversity of mechanisms by which reversible colour change occurs and the evolutionary context and diversity of arthropod taxa in which it has been observed. Further, we discuss the functions of reversible colour change so far proposed, review the limited behavioural and ecological data, and argue that the field requires phylogenetically controlled approaches to understanding the evolution of reversible colour change. Finally, we encourage biologists to explore new model systems for colour change and to engage scientists from other disciplines; continued cross-disciplinary collaboration is the most promising approach to this nexus of biology, physics, and chemistry.

Summary1. Cryptic coloration may evolve in response to selective pressure imposed by predators, yet effective intraspecific communication may require some level of detectability. This creates a tension between the benefits of sexually selected visual traits and the predatory costs imposed by greater conspicuousness, and little is known about how this tension may be ameliorated in highly cryptic species. 2. We explore these competing demands in the false garden mantid Pseudomantis albofimbriata, a colour-blind and seemingly cryptic insect. We use reflectance spectrometry and receptor-noise modelling to characterize the conspicuousness of mantid body regions in the visual systems of mates (mantids), as well as potential predators (birds) and prey (bees). We then use condition manipulation and conspecific choice tests to further explore the colour traits of interest. 3. Based on visual modelling, we find that male mantids are inconspicuous to conspecifics, prey and predators -that is, they are chromatically and achromatically cryptic. In contrast, female mantids are chromatically cryptic to all potential receivers, but their abdomens are achromatically conspicuous. Our food manipulation experiment shows that females in good condition (and therefore with more eggs) have brighter abdomens than females in poor condition. Choice assays show male mantids are consistently attracted to females bearing brighter abdomens. 4. Our results reveal brightness-mediated sexual signalling in a colour-blind and classically cryptic insect. By communicating in the only visual channel available to them, female mantids are conspicuously signalling their quality to mates, while potentially minimizing their conspicuousness to predators and prey. Furthermore, by signalling with only a single body region, female mantids are apparently using coincident disruptive coloration to further decrease detectability to potential eavesdroppers. 5. Our data reveal a novel example of the way in which the trade-off between sexual selection for conspicuousness and natural selection for crypsis may be mediated in a visual signalling system. Such signals may be common in apparently cryptic species, and this study once again demonstrates the importance of analysing visual signals beyond the capacity of human vision.

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Aposematism involves predators learning conspicuous signals of defended prey. However, prey species utilize a wide range of chemical (or physical) defenses, which are not likely to be equally aversive to all predators. Aposematism may therefore only be effective against a physiologically sensitive subset of potential predators, and this can only be identified through behavioral testing. We studied the emerging model organism Tectocoris diophthalmus (Heteroptera: Scutelleridae), an aposematically colored but weakly defended shieldback stinkbug, to test the efficacy of its defenses against a suite of predator types. We predicted the bugs' defenses would be ineffectual against both experienced and naïve birds but aversive to predaceous insects. Surprisingly, the opposite pattern was found. Both habituated wild passerines and naïve chickens avoided the bugs, the chickens after only one or two encounters. To avian predators, T. diophthalmus is aposematic. However, praying mantids showed no repellency, aversion, or toxicity associated with adult or juvenile bugs after multiple trials. Comparison with prior studies on mantids using bugs with chemically similar but more concentrated defenses underscores the importance of dose in addition to chemical identity in the efficacy of chemical defenses. Our results also emphasize the importance of behavioral testing with multiple ecologically relevant predators to understand selective pressures shaping aposematic signals and chemical defenses.

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Theory suggests that aposematism, specifically the learned avoidance of unprofitable prey via memorable color patterns, should result in selection for pattern uniformity. However, many examples to the contrary are seen in nature. Conversely, honest sexual signals are likely to exhibit greater variation because they reflect underlying variation in mate quality. Here we aim to characterize and quantify the mechanistic causes of color in Tectocoris diopthalmus to shed light on the costs of color production, and thus the potential information content of its color signals. We use Tectocoris diopthalmus because it is a weakly-defended stinkbug, and presents elements that have classically been studied in the context of aposematism (red coloring), and sexual selection (sexual dichromatism and iridescent coloring). Pigment analysis reveals that variation in orange coloration is due to the amount of erythropterin pigment, stored in intracellular granules. This pigment is common in Heteroptera, and as an endogenously produced excretory byproduct is unlikely to reflect mate quality or variation in unprofitability of the bug. Electron microscopy reveals the iridescent patches are caused by an epicuticular multilayer reflector, and the hue and patch size are directly related to the layer widths and extent of coverage of this layering. Furthermore, we identified melanin as an essential component of the multilayer reflector system; therefore, the quality of the iridescent patches may be affected by aspects of rearing environment and immunocompetence. We posit that T. diopthalmus has co-opted the melanic patches of a ‘typical’ red and black aposematic signal, transforming it into a complex and variable iridescent signal that may enhance its capacity to display individual quality.

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