Temporal relationship between genetic and warning signal variation in the aposematic wood tiger moth (Parasemia plantaginis).

Many plants and animals advertise unpalatability through warning signals in the form of colour and shape. Variation in warning signals within local populations is not expected because they are subject to directional selection. However, mounting evidence of warning signal variation within local populations suggests that other selective forces may be acting. Moreover, different selective pressures may act on the individual components of a warning signal. At present, we have a limited understanding about how multiple selection processes operate simultaneously on warning signal components, and even less about their temporal and spatial dynamics. Here, we examined temporal variation of several wing warning signal components (colour, UV-reflectance, signal size and pattern) of two co-occurring colour morphs of the aposematic wood tiger moth (Parasemia plantaginis). Sampling was carried out in four geographical regions over three consecutive years. We also evaluated each morph's temporal genetic structure by analysing mitochondrial sequence data and nuclear microsatellite markers. Our results revealed temporal differences between the morphs for most signal components measured. Moreover, variation occurred differently in the fore- and hindwings. We found no differences in the genetic structure between the morphs within years and regions, suggesting single local populations. However, local genetic structure fluctuated temporally. Negative correlations were found between variation produced by neutrally evolving genetic markers and those of the different signal components, indicating a non-neutral evolution for most warning signal components. Taken together, our results suggest that differential selection on warning signal components and fluctuating population structure can be one explanation for the maintenance of warning signal variation in this aposematic species.

Galarza JA ，Nokelainen O ，Ashrafi R ，Hegna RH ，Mappes J ... -  《-》

Changes in predator community structure shifts the efficacy of two warning signals in Arctiid moths.

Polymorphism in warning coloration is puzzling because positive frequency-dependent selection by predators is expected to promote monomorphic warning signals in defended prey. We studied predation on the warning-coloured wood tiger moth (Parasemia plantaginis) by using artificial prey resembling white and yellow male colour morphs in five separate populations with different naturally occurring morph frequencies. We tested whether predation favours one of the colour morphs over the other and whether that is influenced either by local, natural colour morph frequencies or predator community composition. We found that yellow specimens were attacked less than white ones regardless of the local frequency of the morphs indicating frequency-independent selection, but predation did depend on predator community composition: yellows suffered less attacks when Paridae were abundant, whereas whites suffered less attacks when Prunellidae were abundant. Our results suggest that spatial heterogeneity in predator community composition can generate a geographical mosaic of selection facilitating the evolution of polymorphic warning signals. This is the first time this mechanism gains experimental support. Altogether, this study sheds light on the evolution of adaptive coloration in heterogeneous environments.

Nokelainen O ，Valkonen J ，Lindstedt C ，Mappes J ... -  《-》

Environment-mediated morph-linked immune and life-history responses in the aposematic wood tiger moth.

1. Warning signals are expected to evolve towards conspicuousness and monomorphism, and thereby hamper the evolution of multiple colour morphs. Here, we test fitness responses to different rearing densities to explain colour polymorphism in aposematic wood tiger moth (Parasemia plantaginis) males. 2. We used larval lines sired by white or yellow adult males selected for small or large melanization patterns of coloration. We reared these selected lines either solitarily (favourable conditions) or in aggregations (challenged conditions), and followed their performance to adult stage. We tested whether differences in larval density affected life-history traits, adult melanin expression, adult morph (white or yellow) survival and immunological responses. 3. We found that the aggregated environment increased mortality of larvae, but decreased larval developmental time and pupa weight. Adult wing melanin pigmentation was dependent on larval melanin expression but not rearing density. We also confirmed that adult wing coloration had a genetic basis (h(2) = 0.42) and was not influenced by larval growth density. Adult yellow males survived better from aggregations in comparison with white males, which may be related to differences in immune defence. White males had better encapsulation ability, whereas yellow males had increased lytic activity of haemolymph in the aggregations. 4. Our main results highlight, that morph-linked immune responses mediated by differential growth density may facilitate the maintenance of colour polymorphism in aposematic species. In nature, risk of diseases and parasites vary spatially and temporally. Therefore, both yellow and white adult morphs can be maintained due to their differential investment in immune defence in heterogeneous environments.

Nokelainen O ，Lindstedt C ，Mappes J 《-》

To quiver or to shiver: increased melanization benefits thermoregulation, but reduces warning signal efficacy in the wood tiger moth.

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 (58° N) and north Finland (67° 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.

Hegna RH ，Nokelainen O ，Hegna JR ，Mappes J ... -  《-》

Geographic mosaic of selection by avian predators on hindwing warning colour in a polymorphic aposematic moth.

Warning signals are predicted to develop signal monomorphism via positive frequency-dependent selection (+FDS) albeit many aposematic systems exhibit signal polymorphism. To understand this mismatch, we conducted a large-scale predation experiment in four countries, among which the frequencies of hindwing warning coloration of the aposematic moth, Arctia plantaginis, differ. Here we show that selection by avian predators on warning colour is predicted by local morph frequency and predator community composition. We found +FDS to be the strongest in monomorphic Scotland and lowest in polymorphic Finland, where the attack risk of moth morphs depended on the local avian community. +FDS was also found where the predator community was the least diverse (Georgia), whereas in the most diverse avian community (Estonia), hardly any models were attacked. Our results support the idea that spatial variation in predator communities alters the strength or direction of selection on warning signals, thus facilitating a geographic mosaic of selection.

Rönkä K ，Valkonen JK ，Nokelainen O ，Rojas B ，Gordon S ，Burdfield-Steel E ，Mappes J ... -  《-》