Speciation is facilitated if ecological adaptation directly causes assortative mating, but few natural examples are known. Here we show that a shift in colour pattern mimicry was crucial in the origin of two butterfly species. The sister species Heliconius melpomene and Heliconius cydno recently diverged to mimic different model taxa, and our experiments show that their mimetic coloration is also important in choosing mates. Assortative mating between the sister species means that hybridization is rare in nature, and the few hybrids that are produced are non-mimetic, poorly adapted intermediates. Thus, the mimetic shift has caused both pre-mating and post-mating isolation. In addition, individuals from a population of H. melpomene allopatric to H. cydno court and mate with H. cydno more readily than those from a sympatric population. This suggests that assortative mating has been enhanced in sympatry.

Abstract. 1. Sown wildflower strips are increasingly being established in European countries within agri-environmental schemes to enhance biodiversity, especially in intensively used agricultural areas.2. The regulations vary between countries regarding the seed mixture, intensity of management and period of time over which subsidies are given. Insects in particular are intended to benefit from these schemes.3. This review treats studies of insect diversity and abundance in sown wildflower strips. Schemes on wildflower strips in several countries in Central and Northern Europe are compared.4. In a significant majority of studies, sown wildflower strips support higher insect abundances and diversity than cropped habitats. In general, numbers and diversity also tend to be higher than in other margin types such as sown grass margins and natural regeneration, but pollen-and nectar-rich flower mixtures may outperform them.5. Common species are the main beneficiaries of the establishment of wildflower strips, although some studies point out the presence of rare or declining insect species.6. Insect groups respond differently to particular characteristics of the strips. Flower abundance, seed mixture, vegetation structure, management, age and landscape have been identified as factors influencing insect abundance and diversity.7. Future work should address under-represented comparisons, such as with pollen-and nectar-rich seed mixes, and neglected groups, in particular parasitoids. Nevertheless, sown wildflower strips can already be seen as a beneficial measure to enhance insect diversity. This is especially the case, where schemes for sown strips vary within a region to favour different species groups.

Understanding the fate of hybrids in wild populations is fundamental to understanding speciation. Here we provide evidence for disruptive sexual selection against hybrids between Heliconius cydno and Heliconius melpomene. The two species are sympatric across most of Central and Andean South America, and coexist despite a low level of hybridization. No-choice mating experiments show strong assortative mating between the species. Hybrids mate readily with one another, but both sexes show a reduction in mating success of over 50% with the parental species. Mating preference is associated with a shift in the adult colour pattern, which is involved in predator defence through Müllerian mimicry, but also strongly affects male courtship probability. The hybrids, which lie outside the curve of protection afforded by mimetic resemblance to the parental species, are also largely outside the curves of parental mating preference. Disruptive sexual selection against F(1) hybrids therefore forms an additional post-mating barrier to gene flow, blurring the distinction between pre-mating and post-mating isolation, and helping to maintain the distinctness of these hybridizing species.

SUMMARY Despite renewed interest in the role of natural selection as a catalyst for the origin of species, the developmental and genetic basis of speciation remains poorly understood. Here we describe the genetics of Müllerian mimicry in Heliconius cydno and H. melpomene (Lepidoptera: Nymphalidae), sister species that recently diverged to mimic other Heliconius. This mimetic shift was a key step in their speciation, leading to pre-and postmating isolation. We identify 10 autosomal loci, half of which have major effects. At least eight appear to be homologous with genes known to control pattern differences within each species. Dominance has evolved under the influence of identifiable "modifier" loci rather than being a fixed characteristic of each locus. Epistasis is found at many levels: phenotypic interaction between specific pairs of genes, developmental canalization due to polygenic modifiers so that patterns are less sharply defined in hybrids, and overall fitness through ecological selection against nonmimetic hybrid genotypes. Most of the loci are clustered into two genomic regions or "supergenes," suggesting color pattern evolution is constrained by preexisting linked elements that may have arisen via tandem duplication rather than having been assembled by natural selection. Linkage, modifiers, and epistasis affect the strength of mimicry as a barrier to gene flow between these naturally hybridizing species and may permit introgression in genomic regions unlinked to those under disruptive selection. Müllerian mimics in Heliconius use different genetic architectures to achieve the same mimetic patterns, implying few developmental constraints. Therefore, although developmental and genomic constraints undoubtedly influence the evolutionary process, their effects are probably not strong in comparison with natural selection.