In animals, scaling relationships between appendages and body size exhibit high interspecific variation but low intraspecific variation. This pattern could result from natural selection for specific allometries or from developmental constraints on patterns of differential growth. We performed artificial selection on the allometry between forewing area and body size in a butterfly to test for developmental constraints, and then used the resultant increased range of phenotypic variation to quantify natural selection on the scaling relationship. Our results show that the shortterm evolution of allometries is not limited by developmental constraints. Instead, scaling relationships are shaped by strong natural selection.

Dividing sister pairs of the butterfly Bicyclus anynana (reared in a common environment) between high and low temperature shows that oviposition temperature induces a plastic response in egg size. Females at a lower temperature laid significantly larger (but fewer) eggs than their sisters kept at a higher temperature, whereas total reproductive investment increased with temperature. Cross‐transfer experiments demonstrated that this plastic response in egg size is reversible. Interestingly, this pattern parallels an almost universal temperature‐induced developmental response in ectotherm body size. In both cases, however, we do not yet understand the underlying mechanisms or the potential adaptive significance. By cross‐transferring the experimentally manipulated eggs between temperatures, we showed that the larger eggs produced at a lower temperature had a higher hatching success, and yielded larger hatchlings with a slightly higher probability of reaching maturity and shorter larval development time (at the lower temperature) compared to the smaller eggs produced at a higher temperature. Overall, the lower temperature was slightly more detrimental for smaller than for larger eggs. The slight differences in survival across temperatures, together with more substantial differences in fecundity, indicate that it could pay off to produce fewer but larger offspring (with increased fitness) at a lower temperature, but more and smaller offspring at a higher temperature where offspring survival was generally high. Our data present for the first time evidence that selection may favor larger eggs at a lower temperature, and thus that a plastic response of egg size to temperature in ectotherms could be adaptive.

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An important issue in the study of the evolution of aging in Drosophila melanogaster is whether decreased early fecundity is inextricably coupled with increased life span in selection experiments on age at reproduction. Here, this problem has been tackled using an experimental design in which selection is applied directly to longevity. Selection appeared successful for short and long life, in females as well as males. Progeny production of females selected for long life was lower than for short-lived females throughout their whole life. No increase of late-life reproduction in long-lived females occurred, as has been found in selection experiments on age at reproduction. This discrepancy is explained in terms of the inadequacy of the latter design to separate selection on life span from selection on late-life fecundity. Moreover, starvation resistance and fat content were lower for adults selected for short life. In general, the data support the negative-pleiotropy-disposable-soma theory of aging, and it is hypothesized that the pleiotropic allocation of resources to maintenance versus to reproduction as implicated in the theory might involve lipid metabolism. It is argued that further research on this suggestion is urgent and should certainly comprise observations on male reproduction because these are for the greater part still lacking. In conclusion, the longevity of D. melanogaster can be genetically altered in a direct-selection design, and such an increase is accompanied by a decreased general reproduction and thus early reproduction.

SummaryEvidence is accumulating that aging is hormonally regulated by an evolutionarily conserved insulin/IGF-1 signalling (IIS) pathway. Mutations in IIS components affect lifespan in Caenorhabditis elegans, Drosophila melanogaster and mice. Most long-lived IIS mutants also show increased resistance to oxidative stress. In D. melanogaster and mice, the longlived phenotype of several IIS mutants is restricted to females. Here, we analysed the relationship between IIS signalling, body height and longevity in humans in a prospective follow-up study. Based on the expected effects (increased or decreased signalling) of the selected variants in IIS pathway components ( GHRHR, GH1, IGF1, INS, IRS1 ), we calculated composite IIS scores to estimate IIS pathway activity. In addition, we analysed the relative impact on lifespan and body size of the separate variants in multivariate models. In women, lower IIS scores are significantly associated with lower body height and improved old age survival. Multivariate analyses showed that these results were most pronounced for the GH1 SNP, IGF1 CA repeat and IRS1 SNP. In females, for variant allele carriers of the GH1 SNP, body height was 2 cm lower ( P = 0.007) and mortality 0.80-fold reduced ( P = 0.019) when compared with wild-type allele carriers. Thus, in females, genetic variation causing reduced IIS activation is beneficial for old age survival. This effect was stronger for the GH1 SNP than for variation in the conserved IIS genes that were found to affect longevity in model organisms.

Polyphenisms-the expression of discrete phenotypic morphs in response to environmental variation-are examples of phenotypic plasticity that may potentially be adaptive in the face of predictable environmental heterogeneity. In the butterfly Bicyclus anynana, we examine the hormonal regulation of phenotypic plasticity that involves divergent developmental trajectories into distinct adult morphs for a suite of traits as an adaptation to contrasting seasonal environments. This polyphenism is induced by temperature during development and mediated by ecdysteroid hormones. We reared larvae at separate temperatures spanning the natural range of seasonal environments and measured reaction norms for ecdysteroids, juvenile hormones (JHs) and adult fitness traits. Timing of peak ecdysteroid, but not JH titres, showed a binary response to the linear temperature gradient. Several adult traits (e.g. relative abdomen mass) responded in a similar, dimorphic manner, while others (e.g. wing pattern) showed a linear response. This study demonstrates that hormone dynamics can translate a linear environmental gradient into a discrete signal and, thus, that polyphenic differences between adult morphs can already be programmed at the stage of hormone signalling during development. The range of phenotypic responses observed within the suite of traits indicates both shared regulation and independent, trait-specific sensitivity to the hormone signal.

Abstract Genetic variation is the fuel of evolution, with standing genetic variation especially important for short-term evolution and local adaptation. To date, studies of spatiotemporal patterns of genetic variation in natural populations have been challenging, as comprehensive sampling is logistically difficult, and sequencing of entire populations costly. Here, we address these issues using a collaborative approach, sequencing 48 pooled population samples from 32 locations, and perform the first continent-wide genomic analysis of genetic variation in European Drosophila melanogaster. Our analyses uncover longitudinal population structure, provide evidence for continent-wide selective sweeps, identify candidate genes for local climate adaptation, and document clines in chromosomal inversion and transposable element frequencies. We also characterize variation among populations in the composition of the fly microbiome, and identify five new DNA viruses in our samples.

Among the animals, the Lepidoptera (moths and butterflies) are second only to beetles in number of described species and are known for their striking intra- and interspecific diversity. Within species, sexual dimorphism is a source of variation in life history (e.g., sexual size dimorphism and protandry), morphology (e.g., wing shape and color pattern), and behavior (e.g., chemical and visual signaling). Sexual selection and mating systems have been considered the primary forces driving the evolution of sexual dimorphism in the Lepidoptera, and alternative hypotheses have been neglected. Here, we examine opportunities for sexual selection, natural selection, and the interplay between the two forces in the evolution of sexual differences in the moths and butterflies. Our primary goal is to identify mechanisms that either facilitate or constrain the evolution of sexual dimorphism, rather than to resolve any perceived controversy between hypotheses that may not be mutually exclusive.