The transition to selfing in Capsella rubella accompanies its recent divergence from the ancestral outcrossing C. grandiflora species about 100,000 years ago. Whether the change in mating system was accompanied by the evolution of additional reproductive barriers that enforced species divergence remained unknown. Here, we show that C. rubella and C. grandiflora are reproductively separated by an endosperm-based, non-reciprocal postzygotic hybridization barrier. While hybridizations of C. rubella maternal plants with C. grandiflora pollen donors resulted in complete seed abortion caused by endosperm cellularization failure, the reciprocal hybridization resulted in the formation of small seeds with precociously cellularized endosperm. Strikingly, the transcriptomic response of both hybridizations mimicked respectively the response of paternal and maternal excess hybridizations in Arabidopsis thaliana, suggesting unbalanced genome strength causes hybridization failure in both species. These results provide strong support for the theory that crosses between plants of different mating systems will be unbalanced, with the outcrosser behaving like a plant of increased ploidy, evoking a response that resembles an interploidy-type seed failure. Seed incompatilibity of C. rubella pollinated by C. grandiflora followed the Bateson-Dobzhansky-Muller model, involving negative genetic interaction of multiple paternal C. grandiflora loci with at least one maternal C. rubella locus. Given that both species only recently diverged, our data suggest that a fast evolving mechanism underlies the post-zygotic hybridization barrier(s) separating both species.

Based on the biological species concept, two species are considered distinct if reproductive barriers prevent gene flow between them. In Central Europe, the diploid species Arabidopsis lyrata and Arabidopsis arenosa are genetically isolated, thus fitting this concept as "good species." Nonetheless, interspecific gene flow involving their tetraploid forms has been described. The reasons for this ploidy-dependent reproductive isolation remain unknown. Here, we show that hybridization between diploid A. lyrata and A. arenosa causes mainly inviable seed formation, revealing a strong postzygotic reproductive barrier separating these two species. Although viability of hybrid seeds was impaired in both directions of hybridization, the cause for seed arrest differed. Hybridization of A. lyrata seed parents with A. arenosa pollen donors resulted in failure of endosperm cellularization, whereas the endosperm of reciprocal hybrids cellularized precociously. Endosperm cellularization failure in both hybridization directions is likely causal for the embryo arrest. Importantly, natural tetraploid A. lyrata was able to form viable hybrid seeds with diploid and tetraploid A. arenosa, associated with the reestablishment of normal endosperm cellularization. Conversely, the defects of hybrid seeds between tetraploid A. arenosa and diploid A. lyrata were aggravated. According to these results, we hypothesize that a tetraploidization event in A. lyrata allowed the production of viable hybrid seeds with A. arenosa, enabling gene flow between the two species.interspecies hybrid seed failure | Arabidopsis | reproductive isolation | endosperm cellularization | EBN

Pleistocene climatic fluctuations had major impacts on desert biota in southwestern North America. During cooler and wetter periods, drought-adapted species were isolated into refugia, in contrast to expansion of their ranges during the massive aridification in the Holocene. Here, we use Melampodium leucanthum (Asteraceae), a species of the North American desert and semi-desert regions, to investigate the impact of major aridification in southwestern North America on phylogeography and evolution in a widespread and abundant drought-adapted plant species. The evidence for three separate Pleistocene refugia at different time levels suggests that this species responded to the Quaternary climatic oscillations in a cyclic manner. In the Holocene, once differentiated lineages came into secondary contact and intermixed, but these range expansions did not follow the eastwardly progressing aridification, but instead occurred independently out of separate Pleistocene refugia. As found in other desert biota, the Continental Divide has acted as a major migration barrier for M. leucanthum since the Pleistocene. Despite being geographically restricted to the eastern part of the species' distribution, autotetraploids in M. leucanthum originated multiple times and do not form a genetically cohesive group.

Chromosome evolution (including polyploidy, dysploidy, and structural changes) as well as hybridization and introgression are recognized as important aspects in plant speciation. A suitable group for investigating the evolutionary role of chromosome number changes and reticulation is the medium-sized genus Melampodium (Millerieae, Asteraceae), which contains several chromosome base numbers (x=9, 10, 11, 12, 14) and a number of polyploid species, including putative allopolyploids. A molecular phylogenetic analysis employing both nuclear (ITS) and plastid (matK) DNA sequences, and including all species of the genus, suggests that chromosome base numbers are predictive of evolutionary lineages within Melampodium. Dysploidy, therefore, has clearly been important during evolution of the group. Reticulate evolution is evident with allopolyploids, which prevail over autopolyploids and several of which are confirmed here for the first time, and also (but less often) on the diploid level. Within sect. Melampodium, the complex pattern of bifurcating phylogenetic structure among diploid taxa overlain by reticulate relationships from allopolyploids has non-trivial implications for intrasectional classification.

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The genus Melampodium consists of 40 species distributed throughout Mexico and Central America with extensions into the southwestern United States and Colombia and Brasil of South America. The genus reflects broad chromosomal evolution involving dysploidy and polyploidy withn = 9, 10, 11, 12, 14, 18, 20, 23, 24, 27, 28, 30, and 33 having been documented (x = 9, 10, 11, 12, and 14). The most recent classification of the genus, based primarily on morphology and chromosome num bers, recognized six taxonomic sections: Alcina, Bibractiaria, Melampodium, Rhizomaria, Serratura, and Zarabellia. Section Melampodium, which contains 22 species, was divided into five series: Cupulata, Leucantha, Longipila, Melampodium, and Sericea. This hypothesis had been tested by previous morphological phenetic and cladistic analyses, and several problem areas had been highlighted, especially involving section Alcina, but no modifications to the classification have been made. Recent molecular studies utilizing nrITS, 5S rDNA spacer and low‐copy nuclear PgiC gene, plus plastid matK and psbA‐trnH regions allow the classificatory hypothesis to be tested more precisely. Congruence between ITS and matK phylogenies for all species reveals sections Bibractiaria, Rhizomaria, and Serratura to be holophyletic. The largest section Melampodium is holophyletic with both plastid markers, but M. longipilum is shown as an outlier (tying to sect. Rhizomaria) in all nuclear marker phylogenies (ITS, PgiC and 5S rDNA spacer). Section Zarabellia appears holophyletic in matK but biphyletic with ITS, suggesting recogni tion of two distinct series. Section Alcina is the most problematic, being triphyletic in ITS and matK, and with the species not connecting to the same relatives. These insights recommend recognition of three sections from within section Alcina, one hous ing M. nutans (Nutantia Stuessy, sect. nov.) another M. glabrum (Glabrata Stuessy, sect. nov.) and a third M. perfoliatum (sect. Alcina (Cav.) DC.). These may represent ancient independent lines that have diverged from original x = 11 ancestors. Within section Melampodium, 5S rDNA NTS, PgiC and psbA‐trnH, in addition to ITS and matK, provide insights on relationships among taxonomic series. Series Leucantha and Longipila are holophyletic, and series Cupulata nearly so, with M. glabribracteatum deserving treatment in a series of its own. The most complex relationships are between series Melampodium, Cupulata, and Sericea, the latter comprising five exclusively polyploid taxa (4x and 6x). Molecular and cytogenetic data reveal allopolyploid origins for all of these polyploids, in some cases involving hybridization between the two series Cupulata and Melampodium / Sericea, such as with M. mayfieldii and M. longicorne, which exacerbates taxonomic circumscription.

The biennial plant Gentianella bohemica is a subendemic of the Bohemian Massif, where it occurs in seminatural grasslands. It has become rare in recent decades as a result of profound changes in land use. Using amplified fragment length polymorphisms (AFLP) fingerprint data, we investigated the genetic structure within and among populations of G. bohemica in Bavaria, the Czech Republic, and the Austrian border region. The aim of our study was (1) to analyze the genetic structure among populations and to discuss these findings in the context of present and historical patterns of connectivity and isolation of populations, (2) to analyze genetic structure among consecutive generations (cohorts of two consecutive years), and (3) to investigate relationships between intrapopulational diversity and effective population size (Ne) as well as plant traits. (1) The German populations were strongly isolated from each other (pairwise FST= 0.29–0.60) and from all other populations (FST= 0.24–0.49). We found a pattern of near panmixis among the latter (FST= 0.15–0.35) with geographical distance explaining only 8% of the genetic variance. These results were congruent with a principal coordinate analysis (PCoA) and analysis using STRUCTURE to identify genetically coherent groups. These findings are in line with the strong physical barrier and historical constraints, resulting in separation of the German populations from the others. (2) We found pronounced genetic differences between consecutive cohorts of the German populations (pairwise FST= 0.23 and 0.31), which can be explained by local population history (land use, disturbance). (3) Genetic diversity within populations (Shannon index, HSh) was significantly correlated with Ne (RS= 0.733) and reflected a loss of diversity due to several demographic bottlenecks. Overall, we found that the genetic structure in G. bohemica is strongly influenced by historical periods of high connectivity and isolation as well as by marked demographic fluctuations in declining populations.

18The formation of an allopolyploid species involves the merger of two genomes with separate 19 evolutionary histories. In allopolyploids, genes derived from one progenitor species are often 20 expressed at higher levels than those from the other progenitor. It has been suggested that this 21 could be due to differences in transposable element (TE) content among progenitors, as 22 silencing of TEs can affect expression of nearby genes. Here, we examine the role of TEs for 23 expression biases in the widespread allotetraploid Capsella bursa-pastoris and in diploid F1 24 hybrids generated by crossing Capsella orientalis and Capsella rubella, two close relatives of 25 the progenitors of C. bursa-pastoris. As C. rubella harbors more TEs than C. orientalis, we 26 expect C. orientalis alleles to be expressed at higher levels if TE content is key for expression 27 biases. To test this hypothesis, we quantified expression biases at approximately 5800 genes 28 in flower buds and leaves, while correcting for read mapping biases using genomic data. 29While three of four C. bursa-pastoris accessions exhibited a shift toward higher relative 30 expression of C. orientalis alleles, the fourth C. bursa-pastoris accession had the opposite 31 direction of expression bias, as did diploid F1 hybrids. Associations between TE 32 polymorphism and expression bias were weak, and the effect of TEs on expression bias was 33 small. These results suggest that differences in TE content alone cannot fully explain 34 expression biases in these species. Future studies should investigate the role of differences in 35 TE silencing efficacy, as well as a broader set of other factors. Our results are important for a 36 more general understanding of the role of TEs for cis-regulatory evolution in plants. 37 subgenome (e.g. Gossypium; Flagel and Wendel 2010, Brassica; Woodhouse et al 2014, 52