Intrinsic postzygotic barriers can play an important and multifaceted role in speciation, but their contribution is often thought to be reserved to the final stages of the speciation process. Here, we review how intrinsic postzygotic barriers can contribute to speciation, and how this role may change through time. We outline three major contributions of intrinsic postzygotic barriers to speciation. (i) reduction of gene flow : intrinsic postzygotic barriers can effectively reduce gene exchange between sympatric species pairs. We discuss the factors that influence how effective incompatibilities are in limiting gene flow. (ii) early onset of species boundaries via rapid evolution : intrinsic postzygotic barriers can evolve between recently diverged populations or incipient species, thereby influencing speciation relatively early in the process. We discuss why the early origination of incompatibilities is expected under some biological models, and detail how other (and often less obvious) incompatibilities may also serve as important barriers early on in speciation. (iii) reinforcement : intrinsic postzygotic barriers can promote the evolution of subsequent reproductive isolation through processes such as reinforcement, even between relatively recently diverged species pairs. We incorporate classic and recent empirical and theoretical work to explore these three facets of intrinsic postzygotic barriers, and provide our thoughts on recent challenges and areas in the field in which progress can be made. This article is part of the theme issue ‘Towards the completion of speciation: the evolution of reproductive isolation beyond the first barriers’.

Highlights d Mimulus decorus is a cryptic species complex within the M. guttatus species complex d M. decorus is reproductively isolated from M. guttatus by hybrid seed inviability d Patterns of hybrid seed inviability conform to the predictions of parental conflict d Differences in conflict between species scale with genomewide diversity

Seed dormancy can prevent germination under unfavourable conditions that reduce the chances of seedling survival. Freshly harvested seeds often have strong primary dormancy that depends on the temperature experienced by the maternal plant and which is gradually released through afterripening. However, seeds can be induced into secondary dormancy if they experience conditions or cues of future unfavourable conditions. Whether this secondary dormancy induction is influenced by seed-maturation conditions and primary dormancy has not been explored in depth. In this study, we examined secondary dormancy induction in seeds of Arabidopsis thaliana matured under different temperatures and with different levels of afterripening. We found that low water potential and a range of temperatures, from 8°C to 35°C, induced secondary dormancy. Secondary dormancy induction was affected by the state of primary dormancy of the seeds. Specifically, afterripening had a non-monotonic effect on the ability to be induced into secondary dormancy by stratification; first increasing in sensitivity as afterripening proceeded, then declining in sensitivity after 5 months of afterripening, finally increasing again by 18 months of afterripening. Seed-maturation temperature sometimes had effects that were independent of expressed primary dormancy, such that seeds that had matured at low temperature, but which had comparable germination proportions as seeds matured at warmer temperatures, were more easily induced into secondary dormancy. Because seed-maturation temperature is a cue of when seeds were matured and dispersed, these results suggest that the interaction of seed-maturation temperature, afterripening and post-dispersal conditions all combine to regulate the time of year of seed germination.

Range and niche expansion are commonly associated with transitions to asexuality, polyploidy and hybridity (allopolyploidy) in plants. The ability of asexual polyploids to colonize novel habitats may be due to widespread generalist clones, multiple ecologically specialized clones, or may be a neutral by-product of multiple, independent origins of asexual polyploids throughout the range. We have quantified niche size and divergence for hawthorns of the Pacific Northwest using data from herbarium vouchers with known cytotypes. We find that all polyploid niches diverge from that of the diploid range, and allopolyploids have the broadest niches. Allotetraploids have the largest niche and the widest geographic distribution. We then assessed the genetic mechanism of range expansion by surveying the ecological and geographic distribution of genotypes within each cytotype from sites in which fine-scale habitat assessments were completed. We find no isolation by either geographic or ecological distance in allopolyploids, suggesting high dispersal and colonization ability. In contrast, autotriploids and diploids show patterns of isolation by geographic distance. We also compared the geographic and ecological distributions of clonal genotypes with those of randomly drawn sites of the most widespread cytotype. We found that most clones are geographically widespread and occur in a variety of habitats. We interpret these findings to suggest that patterns of range and niche expansion in Pacific Northwest Hawthorns may stem from these widespread, ecologically generalist clones of hybrid origin.

No abstract

Chromosomal inversions can play an important role in adaptation, but the mechanism of their action in many natural populations remains unclear. An inversion could suppress recombination between locally beneficial alleles, thereby preventing maladaptive reshuffling with less-fit, migrant alleles. The recombination suppression hypothesis has gained much theoretical support but empirical tests are lacking. Here, we evaluated the evolutionary history and phenotypic effects of a chromosomal inversion which differentiates annual and perennial forms of Mimulus guttatus. We found that perennials likely possess the derived orientation of the inversion. In addition, this perennial orientation occurs in a second perennial species, M. decorus, where it is strongly associated with life history differences between co-occurring M. decorus and annual M. guttatus. One prediction of the recombination suppression hypothesis is that loci contributing to local adaptation will predate the inversion. To test whether the loci influencing perenniality pre-date this inversion, we mapped QTLs for life history traits that differ between annual M. guttatus and a more distantly related, collinear perennial species, M. tilingii. Consistent with the recombination suppression hypothesis, we found that this region is associated with life history in the absence of the inversion, and this association can be broken into at least two QTLs. However, the absolute phenotypic effect of the LG8 inversion region on life history is weaker in M. tilingii than in perennials which possess the inversion. Thus, while we find support for the recombination suppression hypothesis, the contribution of this inversion to life history divergence in this group is likely complex.

Aim Asexual organisms frequently have larger ranges than their sexual progenitors, a phenomenon referred to as geographical parthenogenesis (GP). In plants, GP is associated not only with asexuality (apomixis), but also with polyploidization and hybridity (allopolyploidy). Dispersal is thought to play a role in range‐size differences in other taxa, but has not been directly related to GP. Here, we compare resource allocation to dispersal‐related traits in sexual diploids, asexual autopolyploids and asexual allopolyploids, and relate these differences to patterns of GP. Location The Pacific Northwest, North America. Methods We created distribution maps for all cytotypes known in Crataegus series Douglasianae using herbarium records. To quantify dispersal ability, we collected fruit samples from sexual diploids, apomictic allopolyploids and apomictic autopolyploids across their ranges, and used the masses of each fruit component as a proportion of the total fruit mass to gauge relative investment in dispersal and competitive ability. Results The largest ranges belong to apomictic allopolyploids, whereas apomictic autotriploids and sexual diploids have the smallest ranges. Compared to sexual diploids and apomictic autotriploids, the allotetraploids exhibit a more dispersal‐orientated strategy, with proportionally heavier pyrenes and more fruit pulp, but proportionally lighter seeds. Allotriploid taxa, which arose via back‐crosses between sexual diploids and asexual allotetraploids, exhibit an intermediate range size as well as intermediate investment in dispersal. Main conclusions In Crataegus series Douglasianae, GP is associated only with allopolyploids, highlighting the potential role of hybridization in range expansion. The data suggest that allopolyploids are associated with increased resource allocation to dispersal‐related traits, whereas sexual diploid C. suksdorfii and asexual autotriploid C. gaylussacia exhibit a more competition‐orientated strategy. These findings are consistent with the hypothesis that hybridization contributes to patterns of GP in asexual allopolyploids, potentially by increasing their dispersal ability.

Understanding the factors that produce and maintain genetic variation is a central goal of evolutionary biology. Despite a century of genetic analysis, the evolutionary history underlying patterns of exceptional genetic and phenotypic variation in the model organism Drosophila melanogaster remains poorly understood. In particular, how genetic and phenotypic variation is partitioned across global D. melanogaster populations, and specifically in its putative ancestral range in Subtropical Africa, remains unresolved. Here, we integrate genomic and behavioral analyses to assess patterns of population genetic structure, admixture, mate preference, and genetic incompatibility throughout the range of this model organism. Our analysis includes 174 new accessions from novel and under-sampled regions within Subtropical Africa. We find that while almost all Out of Africa genomes correspond to a single genetic ancestry, different geographic regions within Africa contain multiple distinct ancestries, including the presence of substantial cryptic diversity within Subtropical Africa. We find evidence for significant admixture- and variation in admixture rates-between geographic regions within Africa, as well as between African and non-African lineages. By combining behavioral analysis with population genomics, we demonstrate that female mate choice is highly polymorphic, behavioral types are not monophyletic, and that genomic differences between behavioral types correspond to many regions across the genome. These include regions associated with neurological development, behavior, olfactory perception, and learning. Finally, we discovered that many individual pairs of putative incompatibility loci likely evolved during or after the expansion of D. melanogaster out of Africa. This work contributes to our understanding of the evolutionary history of a key model system, and provides insight into the distribution of reproductive barriers that are polymorphic within species.