W. Talloen

The cost of melanization: Butterfly wing coloration under environmental stress

Abstract Evolutionary studies typically focus on adaptations to particular environmental conditions, thereby often ignoring the role of possible constraints. Here we focus on the case of variation in dorsal wing melanization in a satyrine butterfly Pararge aegeria. Because melanin is a complex polymer, its synthesis may be constrained if ambient conditions limit the resource budget. This hypothesis was tested by comparing melanization among butterflies that fed as larvae on host grasses experiencing different drought‐stress treatments. Treatment differences were validated both at the level of the host plant (nitrogen, carbonate, and water content) and of the butterfly (life‐history traits: survival, development time, and size at maturity). Melanization rate was measured as average gray value of the basal dorsal wing area. This area, close to the thorax, is known to be functionally significant for basking in order to thermoregulate. Individuals reared on drought‐stressed host plants developed paler wings, and development of darker individuals was slower and less stable as estimated by their level of fluctuating asymmetry. These results provide evidence that melanin is indeed costly to synthesize, and that differences in environmental quality can induce phenotypic variation in wing melanization. Therefore, studies dealing with spatial and/or temporal patterns of variation in wing melanization should not focus on adaptive explanations alone, but rather on a cost‐benefit balance under particular sets of environmental conditions.

Does the presence of ant nests matter for oviposition to a specialized myrmecophilous Maculinea butterfly

More than 50% of the lycaenid butterflies have an ant–associated lifestyle (myrmecophily) which may vary from coexistence to specific mutualistic or even parasitic interactions. Ant–related host–plant selection and oviposition has been observed in some myrmecophilous lycaenids. Therefore, it is remarkable that there is no evidence for this behaviour in the highly specialized, obligate myrmecophilous butterflies of the genus Maculinea. In contrast with previous findings, our results provide evidence for ant–related oviposition patterns in Maculinea alcon in relation to the distribution of specific host–ant nests (i.e.Myrmica ruginodis) based on repeated egg counts during the flight period in two populations. We also show that ant–related oviposition can be counterbalanced by intraspecific competition and oviposition deterrency when host plants already carry several eggs. Therefore, the absence of a correlation between egg load and the presence of host–ant nests at the end of the flight period should be interpreted carefully. Whether ovipositional cues are obtained either directly (from ants or their nests) or indirectly (from vegetation structure), and whether alternative explanations based on the phenology and growth form of host plants are possible, is discussed.

THE COST OF MELANIZATION: BUTTERFLY WING COLORATION UNDER ENVIRONMENTAL STRESS

Abstract Evolutionary studies typically focus on adaptations to particular environmental conditions, thereby often ignoring the role of possible constraints. Here we focus on the case of variation in dorsal wing melanization in a satyrine butterfly Pararge aegeria. Because melanin is a complex polymer, its synthesis may be constrained if ambient conditions limit the resource budget. This hypothesis was tested by comparing melanization among butterflies that fed as larvae on host grasses experiencing different drought-stress treatments. Treatment differences were validated both at the level of the host plant (nitrogen, carbonate, and water content) and of the butterfly (life-history traits: survival, development time, and size at maturity). Melanization rate was measured as average gray value of the basal dorsal wing area. This area, close to the thorax, is known to be functionally significant for basking in order to thermoregulate. Individuals reared on drought-stressed host plants developed paler wings, and development of darker individuals was slower and less stable as estimated by their level of fluctuating asymmetry. These results provide evidence that melanin is indeed costly to synthesize, and that differences in environmental quality can induce phenotypic variation in wing melanization. Therefore, studies dealing with spatial and/or temporal patterns of variation in wing melanization should not focus on adaptive explanations alone, but rather on a cost-benefit balance under particular sets of environmental conditions.

Feather development under environmental stress: lead exposure effects on growth patterns in great tits **Parus major**: capsule regrowth rate of tail feathers is more strongly affected compared to feather length and symmetry

Capsule Regrowth rate of tail feathers is more strongly affected compared to feather length and symmetry. Aims To assess the value of avian feathers as bioindicators. Methods The origin and persistence of fluctuating asymmetry (FA) in homologous pairs of regrowing feathers was studied in captive birds under different levels of environmental stress, respresented by exposure to lead (Pb). Homologous feathers of individually housed birds were plucked synchronously or with a delay of seven days. We measured growth rate, regeneration time, final size and FA of regrown feathers and related them to Pb stress. Results Asymmetry decreased as feathers reached their final length. This was not due to compensatory growth but rather a consequence of the programmed growth trajectory of single feathers. Tail feathers grown under higher Pb pollution showed increased regeneration times, decreased growth rates and shorter lengths, but no changes in development times nor in FA. For differences between both (i) original and induced feathers and (ii) control and Pb treatment, effect sizes of parameters related to feather development (growth bar width, growth rate, regeneration and development time) were consistently larger than those related to the resulting phenotype (feather length and FA). Conclusions Growth bar widths in particular provide an applicable, sensitive and reliable indicator of adverse conditions such as Pb pollution and experimental conditions. In general, phenotypic characteristics that retrospectively allow estimation of growth rates may be more suitable for monitoring environmental stress than sizes or asymmetries of full-grown traits.

Environmental stress and quantitative genetic variation in butterfly wing characteristics

Butterfly wing characteristics are extensively used as model system in studies of development, quantitative genetics and phenotypic plasticity. In spite of its evolutionary relevance, however, the effect of stress on the expression of genetic variation itself has only rarely been studied. In this paper, we explore genetic variation of wing characteristics of the Speckled wood Pararge aegeria along a host plant drought stress gradient. Forewing area, basal and distal degree of melanization and the area of five yellow wing spots were measured. We found an increase in (additive) genetic variation in degree of melanization at higher drought stress, and a similar, yet non-significant, relationship for forewing size. As a result, both the upper limits of the narrow-sense heritability and the coefficient of additive genetic variation of wing size and melanization increased with drought stress. Patterns for the different yellow wing spots were less consistent, suggesting trait-specificity in the relationships between genetic variation and environmental stress.