Warning colouration functions to deter predators from attacking unprofitable (e.g., unpalatable or toxic) prey items. While warning colours are often characterized by long-wavelength components (e.g., orange, red or yellow), many distasteful animals across the animal kingdom display orange or red colour patches adjacent to or within a field of short-wavelength colouration such as blue (e.g., strawberry poison dart frogs, pipevine swallowtail and pyjama nudibranch), which yields a multicomponent visual warning signal. Here we show that, in such signals, blue and orange patches can function as redundant signal components; avian predators trained not to attack the intact blue and orange colouration of the pipevine swallowtail (Battus philenor) recognised the butterflies as distasteful even when the blue and orange were presented individually. Our results demonstrate that blue colouration and potential multiple, unimodal, signal components should be considered in research on visual warning signals, including in well-studied animals, such as dendrobatid frogs and swallowtail butterflies.

Predation on distasteful animals should favor warning coloration that is relatively conspicuous and phenotypically invariable. However, even among similarly colored individuals there can be variation in their warning signals. In butterflies, individual differences in larval feeding history could cause this variation. The warning signal of the pipevine swallowtail butterfly, Battus philenor L. (Lepidoptera: Papilionidae) consists of both a blue iridescent patch and pigmentbased orange spots on the ventral hindwing. B. philenor males also display a dorsal surface iridescent patch that functions as a sexual indicator signal. A previous study of iridescence in B. philenor found that the iridescent blue on both the dorsal and ventral hind wings is variable and significantly different between lab-reared and field-caught individuals. These differences could be the result of larval food deprivation in the field. Through experimental manipulation of larval diet, larval food deprivation was evaluated as a potential cause of the differences observed between lab and field individuals, and if food deprivation is a source of inter-individual variation in warning signals. B. philenor larvae were food restricted starting at two points in the last larval instar, and one group was fed through pupation. Adult coloration was then compared. Food deprivation led to poorer adult condition, as indicated by lower adult body mass, forewing length, and fat content of stressed individuals. As the level of food deprivation increased, the hue of the iridescent patches on both the dorsal and ventral hind wing shifted to shorter wavelengths, and the chroma of the orange spots decreased. The shifts in iridescent color did not match the differences previously found between lab and field individuals. However, the treatment differences indicate that food deprivation may be a significant source of warning color variation. The differences between the treatment groups are likely detectable by predators, but the effect of the variation on signal effectiveness and function remains to be empirically explored.

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Warning coloration deters predators from attacking distasteful or toxic prey. Signal features that influence warning color effectiveness are not well understood, and in particular, we know very little about how effective short-wavelength and iridescent colors are as warning color elements in nature and how warning signal effectiveness changes throughout the day. We tested the effect of these factors on predation risk in nature using specimens of the distasteful pipevine swallowtail butterfly, Battus philenor. B. philenor adults display both iridescent blue and diffusely reflecting orange components in their warning signal. We painted B. philenor wings to create five different model types: all-black, only-iridescent-blue, onlyorange, iridescent-blue-and-orange (intact signal), and matte-blue-andorange. We placed 25 models in each of 14 replicate field sites for 72 h and checked for attacks three times each day. Model type affected the likelihood of attack; only-orange models were, the only model attacked significantly less than the all-black model. Iridescence did not enhance or decrease warning signal effectiveness in our experiment because matteblue-and-orange models were attacked at the same rate as iridescentblue-and-orange models. Time of day did not differentially affect model type. Video recordings of attacks revealed that insectivorous birds were responsible. The results of this experiment, when taken with previous work, indicate that the response to blue warning coloration is likely dependent on predator experience and context, but that iridescence per se does not affect warning signals in a natural context.

In many animals, males bear bright ornamental color patches that may signal both the 6 direct and indirect benefits that a female might accrue from mating with him. Here we test 7 whether male coloration in the Pipevine Swallowtail butterfly, Battus philenor, predicts two 8 potential direct benefits for females, copulation duration and the quantity of materials the male 9 passes to the female during mating. In this species, males have a bright iridescent blue field on 10 the dorsal hindwing surface while females have little or no dorsal iridescence. Females 11 preferentially mate with males who display a bright and highly chromatic blue on their dorsal 12 hindwing. In this study, we show that the chroma of the blue on the male dorsal hindwing and 13 male body size (forewing length) significantly predict the mass of material or spermatophore that 14 a male forms within the female's copulatory sac during mating. We also found that 15 spermatophore mass correlated negatively with copulation duration, but that color variables did 16 not significantly predict this potential direct benefit. These results suggest that females may 17 enhance the material benefits they receive during mating by mating with males based on the 18 coloration of their dorsal hindwing.