Appearance before performance? Nutritional constraints on life-history traits, but not warning signal expression in aposematic moths.

Trade-offs have been shown to play an important role in the divergence of mating strategies and sexual ornamentation, but their importance in explaining warning signal diversity has received less attention. In aposematic organisms, allocation costs of producing the conspicuous warning signal pigmentation under nutritional stress could potentially trade-off with life-history traits and maintain variation in warning coloration. We studied this with an aposematic herbivore Arctia plantaginis (Arctiidae), whose larvae and adults show extensive variation in aposematic coloration. In larvae, less melanic coloration (i.e. larger orange patterns) produces a more efficient warning signal against predators, whereas high amounts of melanism (smaller orange pattern) enhance thermoregulation, correlate with better immunity and make individuals harder to detect for naïve predators. We conducted a factorial rearing experiment with larvae originating from lines selected for either small or large orange signal size, which were reared on an artificial diet that had either low or high protein content. Protein content of the diet is critical for melanin production. We measured the effects of diet on individual coloration, life-history traits, immune defence and reproductive output. We also compared the responses to dietary conditions between the small and large larval signal genotypes. Protein content of the diet did not affect warning coloration in the larval stage, but larval signal sizes differed significantly among selection lines, confirming that its variation is mainly genetically determined. In adults, signal line or diet did not affect coloration in hindwings, but males' forewings had more melanin on the high than on low protein diet. Contrary to coloration, diet quality had a stronger impact on life-history traits: individuals developed for longer had smaller hindwing sizes in females and lower immune defence on the low protein content diet compared with the high. These costs were higher for more melanic larval signal genotypes in terms of development time and female hindwing size. We conclude that low plasticity in warning signal characteristics makes signal expression robust under varying dietary conditions. Therefore, variation in diet quality is not likely to constrain signal expression, but can have a bigger impact on performance.