The coloration of species can have multiple functions, such as predator avoidance and sexual signalling, that directly affect fitness. As selection should favour traits that positively affect fitness, the genes underlying the trait should reach fixation, thereby preventing the evolution of polymorphisms. This is particularly true for aposematic species that rely on coloration as a warning signal to advertise their unprofitability to predators. Nonetheless, there are numerous examples of aposematic species showing remarkable colour polymorphisms. We examined whether colour polymorphism in the wood tiger moth is maintained by trade-offs between different functions of coloration. In Finland, males of this species have two distinct colour morphs: white and yellow. The efficacy of the warning signal of these morphs was tested by offering them to blue tits in the laboratory. Birds hesitated significantly longer to attack yellow than white males. In a field experiment, the survival of the yellow males was also higher than white males. However, mating experiments in the laboratory revealed that yellow males had lower mating success than white males. Our results offer an explanation for the maintenance of polymorphism via trade-off between survival selection and mating success.

Melanin production is often considered costly, yet beneficial for thermoregulation. Studies of variation in melanization and the opposing selective forces that underlie its variability contribute greatly to understanding natural selection. We investigated whether melanization benefits are traded off with predation risk to promote observed local and geographical variation in the warning signal of adult male wood tiger moths (Parasemia plantaginis). Warning signal variation is predicted to reduce survival in aposematic species. However, in P. plantaginis, male hindwings are either yellow or white in Europe, and show continuous variation in melanized markings that cover 20 to 90 per cent of the hindwing. We found that the amount of melanization increased from 40 to 59 per cent between Estonia (588 N) and north Finland (678 N), suggesting melanization carries thermoregulatory benefits. Our thermal measurements showed that more melanic individuals warmed up more quickly on average than less melanic individuals, which probably benefits flight in cold temperatures. With extensive field experiments in central Finland and the Alpine region, we found that more melanic individuals suffered increased predation. Together, our data suggest that warning signal efficiency is constrained by thermoregulatory benefits. Differences in relative costs and benefits of melanin probably help to maintain the geographical warning signal differences.

Aim To investigate the phylogeography of the aposematic wood tiger moth (Parasemia plantaginis) across its Holarctic distribution and to explore how its genetic structure relates to geographical differences in hindwing warning coloration of males and females. Males have polymorphic hindwing coloration, while female hindwing coloration varies continuously, but no geographical analyses of coloration or genetic structure exist.Location The Holarctic. MethodsWe sequenced a fragment of the mitochondrial cytochrome c oxidase subunit I gene (COI) from 587 specimens. We also examined more current population structure by genotyping 569 specimens at 10 nuclear microsatellite loci. Species distribution modelling for present conditions and the Last Glacial Maximum (LGM) was performed to help understand genetic structure. Geographical patterns in hindwing warning coloration were described from 1428 specimens and compared to the genetic analyses. ResultsWe found only two instances of genetic divergence that coincided with distinct, yet imperfect, shifts in male hindwing coloration in the Caucasus region and Japan. A shift in female hindwing colour did not appear to be associated with genetic structure. A change from sexual monomorphism to sexual dimorphism was also observed. Mitogenetic (mtDNA) structure does not show the influence of glacial refugia during the LGM. Climate shifts following the LGM appear to have isolated the red Caucasus populations and other southerly populations. Populations at opposite ends of the moth's distribution showed high levels of differentiation in the microsatellite data analysis compared to the shallow mitogenetic structure, supporting a more recent divergence.Main conclusions Parasemia plantaginis populations appeared to have been historically well connected, but current populations are much more differentiated. This raises the possibility that incipient speciation may be occurring in portions of the species' distribution. Some changes in colour align to genetic differences, but others do not, which suggests a role for selective and nonselection based influences on warning signal variation.

Aposematic organisms are not predicted to show high levels of warning signal diversity because they are expected to be under stabilizing selection to decrease costs of 'educating' predators about their unpalatability. However, systematic changes in warning signals (polytypism) can be expected if they represent adaptations to local predators. The aposematic strawberry poison frog (Oophaga pumilio) is red throughout its mainland distribution in Costa Rica and Panamá, but displays high levels of warning signal diversity in the Bocas del Toro Archipelago of Panamá. Both coloration and spot pattern vary in a polytypic sense. Sexual selection contributes to maintaining the polytypism, but little work has investigated the potential influence of predation. We used unspotted models of O. pumilio to determine if predation might help explain the color polytypism on Isla Colón in the Bocas del Toro Archipelago of Panamá. We tested whether attack rates differed among the red mainland morph, green/yellow Isla Colón morph, and the brown control. We found that frog color significantly predicted being attacked. The local green Isla Colón models were attacked more than foreign red or brown models. No difference in attack rate existed between red and brown control models. Our results suggest that the red mainland morph possesses a more effective warning signal, even when it is not the local morph. Honest signaling of unpalatability, neophobia, and the use of search images by local predators are potential explanations. Similarity of the brown model to other local poison frogs might explain the lower attack rate compared to previous work. The attack rate was lower on Isla Colón compared to mainland Costa Rica, which supports the hypothesis that less overall 123Evol Ecol DOI 10.1007/s10682-012-9605-z predation in the Bocas del Toro Archipelago may contribute to the overall warning signal diversity in O. pumilio there by relaxing selection for aposematic traits.