Molecular phylogeny confirms the subtribal classification of Fumarieae based on morphology. However it provides different results regarding the relationships among genera within each subtribe, which affects the inference of the evolutionary pathway followed by the four selected characters. The disjunct distribution of the tribe is explained by different vicariance scenarios.

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Currently, three species of Muscari subg. Botryanthus are recognized in the Iberian Peninsula: two diploids (2n = 18), M. atlanticum and M. cazorlanum, and one morphologically variable species with three different ploidy levels, M. neglectum (2n = 36, 45, 54). We have made a comparative study of numerous Iberian populations to clarify the taxonomy and evolution of this group. To this end we carried out morphological and cytogenetic analyses, and phylogenetic analysis of the internal transcribed spacers of nuclear ribosomal DNA. Comparative and UPGMA analyses of the morphological characteristics show that the different ploidy levels of M. neglectum represent different species. We describe the pentaploid and hexaploid levels as two new species, M. olivetorum (2n = 45) and M. baeticum (2n = 54), each with an exclusive combination of morphological characters and a characteristic ecological behavior pattern. Phylogenetic study of ITS shows that the two new species are not autopolyploids from M. neglectum but allopolyploids. These findings are supported by the additivity of the three ITS variants found in M. olivetorum with the ITS of M. neglectum and M. baeticum, and also by morphology. Possible parents for both new species are proposed. Absence of homogenization between homeologous M. olivetorum nrDNA loci is explained by the absence of sexual reproduction and by nucleolar dominance, indicating that this is a recent species.

Pollen grains show an enormous variety of aperture systems. What genes are involved in the aperture formation pathway and how conserved this pathway is in angiosperms remains largely unknown. INAPERTURATE POLLEN1 (INP1) encodes a protein of unknown function, essential for aperture formation in Arabidopsis, rice and maize. Yet, because INP1 sequences are quite divergent, it is unclear if their function is conserved across angiosperms. Here, we conducted a functional study of the INP1 ortholog from the basal eudicot Eschscholzia californica (EcINP1) using expression analyses, virus-induced gene silencing, pollen germination assay, and transcriptomics. We found that EcINP1 expression peaks at the tetrad stage of pollen development, consistent with its role in aperture formation, which occurs at that stage, and showed, via gene silencing, that the role of INP1 as an important aperture factor extends to basal eudicots. Using germination assays, we demonstrated that, in Eschscholzia, apertures are dispensable for pollen germination. Our comparative transcriptome analysis of wild-type and silenced plants identified over 900 differentially expressed genes, many of them potential candidates for the aperture pathway. Our study substantiates the importance of INP1 homologs for aperture formation across angiosperms and opens up new avenues for functional studies of other aperture candidate genes.

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