Abstract. While the study of colour patterns is a traditional subject of evolutionary ecology, there are various hypotheses which suffer from a lack of experimental evidence. One intriguing possibility is a trade-off between warning efficiency and detectability. After a certain size threshold, the detrimental effect of increased detectability can outweigh the benefits of warning colouration. One may thus expect corresponding patterns at the level of ontogenetic development: as juveniles grow, they should first acquire warning colouration, and then lose it again. We analysed this possibility in Orgyia antiqua, a moth species with hairy larvae which are polyphenic with respect to the intensity of warning colouration. We detected a regular change in colour patterns through larval life. Indeed, the larvae tend to display warning colouration at intermediate sizes while dull colours dominate in fully grown larvae. In aviary experiments, we confirmed that the colourful phenotype is the one that causes the strongest aversion in birds. Nevertheless, the effect was rather weak and most of the larvae were still eventually consumed when found. Unexpectedly, for human subjects, the warningly coloured larvae were harder, and not easier to find among natural vegetation, most likely due to the disruptive effect of the aposematic colour pattern. Importantly, the trend was reversed in the largest size class, suggesting that the disruptive colouration loses its advantage as the larva grows. This is consistent with the actual patterns of size-dependence of colouration. We present evidence against an alternative explanation which relates size-related change in colouration to behavioural changes prior to pupation. We conclude that even if the efficiency of the warning effect plays a role in determining the size-dependence of colouration, the pattern may be largely explained by the effects of size-dependent detectability alone.
Carotenoid‐based integumental coloration is often associated with individual performance in various animals. This is because the limited amount of the pigment has to be allocated to different vital functions. However, most of the evidence for the carotenoid‐based trade‐off comes from vertebrate studies, and it is unclear if this principle can be applied to insects. This possibility was investigated in Orgyia antiqua L. (Lepidoptera: Lymantriidae). The larvae of this species are polyphenic in their coloration, varying from a highly conspicuous combination of yellow hair tufts on black background to cryptic appearance with brown hair tufts. The conspicuous larvae are aposematic, advertising their aversive hairiness. The maintenance of different colour morphs in O. antiqua requires explanation, as an aposematic signal is expected to evolve towards monomorphism. Chromatographic analysis showed that the yellow coloration of the hair is based on the carotenoid pigment lutein (α‐carotene‐3,3’‐diol). The colour of hair tufts was dependent on their carotenoid content. This justifies an expectation of carotenoid‐based physiological trade‐offs between aposematic coloration and individual performance. To test this hypothesis, we monitored life histories of differently coloured larvae reared on various host plants, recording their body sizes, growth rates, and mortalities in each instar. There was a significant but relatively low heritability of tuft coloration, which allowed us to expect environmental effects. We found no phenotypic associations between hair tuft colour and performance indices in O. antiqua larvae, neither did the quality of host plant affect the frequency of colour morphs. However, the frequency of colour morphs differed between larval instars. Our results suggest that carotenoid‐mediated physiological trade‐offs are not involved in the maintenance of colour morphs in O. antiqua larvae, and factors other than individual condition should be responsible for the observed variability.
Aposematic prey advertise their defence to visually hunting predators using conspicuous warning colouration. Established theory predicts that aposematic signals should evolve towards increased conspicuousness and similarity to enhance predator education. Contrary to theoretical expectations, there is often considerable within- and between-species variation in aposematic signals of animals sharing the same ecological niche, phylogeny and predators. This may be explained by varying responses of predators that weaken the selection pressure for a consistent signal. By presenting painted mealworm larvae as prey to great tits as predators we tested if different aposematic colour patterns have different values as a means of initial protection and learnt avoidance from predators, and how widely birds generalise their learnt avoidance to other colour patterns. We also investigated how the colour and luminance of the pattern elements affect predator attack decisions. Finally, we studied if hunger affects the predators' reaction to differently coloured prey. We found that similarity in colour was not crucial to the survival of aposematic prey, since learnt avoidance was not influenced by colour, and predators remembered and generalised widely in their learnt avoidance to other colours. We found that initial avoidance was, however, apparently influenced by luminance contrast. Interestingly, the predators' level of hunger was more important than the colour of the aposematic signal in determining birds' decisions to attack chemically-defended insect larvae. We discuss the implications of visual properties of prey colour pattern and predator appetite for the evolution of insect defences and warning signals. In addition we propose a methodological approach to effectively control for predator appetite in laboratory experiments.
Ultimate causes of phenotypic plasticity in visual appearance are frequently related to increasing the degree of crypsis in a way specific to the environment. The cues used to elicit such plastic responses may be both direct (i.e. straightforward background matching) as well as indirect. In the latter case, cues other than the visual signals from of the environment are used to predict the phenotype best corresponding to the particular situation. On the basis of a series of laboratory experiments we show that the remarkable variability in the visual appearance of the larvae of the geometrid moth Ematurga atomaria, though genetically based in part, involves a substantial environmental component. Using multiple correspondence analysis, we transformed the multidimensional variation in colour and pattern into two dimensions interpretable as patterning and darkness. Plastic changes in the darkness of the larvae were elicited by direct cues: the larvae were darker when reared on dark host‐plants. Host‐specific degree of patterning was also induced in absolute darkness which indicates the use of an indirect cue. This was unexpected because the study species is broadly polyphagous, and thus not likely to have evolved adaptations specific to particular host‐plant species. Indeed, the larvae of E. atomaria originating from geographic populations using different host‐plants showed analogous plastic responses which indicates that the link between the indirect cue and visual appearance of the host needs not to be specific to plant species. In an additional experiment, we showed that surface roughness is a likely candidate to serve as the proximate cue for determination of some pattern elements, a case not reported for insect larvae earlier.
Abstract. Although the effects of host plant quality on the performance of polyphagous herbivores are largely uniform across insect taxa, there are various exceptions to this rule. In particular, there are scattered reports of cases in which the relative quality of different hosts differs among larval instars of a single insect species. Such cases are explained either in terms of differences in the susceptibility of different aged larvae to plant defences or, alternatively, age-specific nutritional demands. Here we report the results of experiments that show that young larvae of the polyphagous common heath moth Ematurga atomaria (Lepidoptera: Geometridae) consistently attain higher weights on common heather Calluna vulgaris than bilberry Vaccinium myrtillus, whereas the rank order of these host plants is reversed in the final larval instar. Phytochemical analyses showed that differences in nutrient content of these plants are not likely to explain the observed pattern. Instead, the results are more consistent with the idea that the greater chemical defence of bilberry has a relatively stronger influence on young than old larvae.
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