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The evolutionary histories of many polyploid plant species are difficult to resolve due to a complex interplay of hybridization, incomplete lineage sorting, and missing diploid progenitors. In the case of octoploid strawberry with four subgenomes designated ABCD, the identities of the diploid progenitors for subgenomes C and D have been subject to much debate. By integrating new sequencing data from North American diploids with reticulate phylogeny and admixture analyses, we uncovered introgression from an extinct or unsampled species in the clade ofFragaria viridis,Fragaria nipponica, andFragaria nilgerrensisinto the donor of subgenome A of octoploidFragariaprior to its divergence fromF. vescasubsp. bracteata. We also detected an introgression event fromF. iinumaeinto an ancestor ofF. nipponicaandF. nilgerrensis.Using an LTR-age-distribution-based approach, we estimate that the octoploid and its intermediate hexaploid and tetraploid ancestors emerged approximately 0.8, 2, and 3 million years ago, respectively. These results provide an explanation for previous reports ofF. viridisandF. nipponicaas donors of the C and D subgenomes and suggest a greater role than previously thought for homoploid hybridization in the diploid progenitors of octoploid strawberry. The integrated set of approaches used here can help advance polyploid genome analysis in other species where hybridization and incomplete lineage sorting obscure evolutionary relationships. 

Abstract The identity of the diploid progenitors of octoploid cultivated strawberry (Fragaria × ananassa) has been subject to much debate. Past work identified four subgenomes and consistent evidence forF. californica(previously namedF. vescasubsp.bracteata) andF. iinumaeas donors for subgenomes A and B, respectively, with conflicting results for the origins of subgenomes C and D. Here, reticulate phylogeny and admixture analysis support hybridization betweenF. viridisandF. vescain the ancestry of subgenome A, and betweenF. nipponicaandF. iinumaein the ancestry of subgenome B. Using an LTR-age-distribution-based approach, we estimate that the octoploid and its intermediate hexaploid and tetraploid ancestors emerged approximately 0.8, 2, and 3 million years ago, respectively. These results provide an explanation for previous reports ofF. viridisandF. nipponicaas donors of the C and D subgenomes and unify conflicting hypotheses about the evolutionary origin of octoploidFragaria. 

Abstract Over the decades, evolutionists and ecologists have shown intense interest in the role of polyploidization in plant evolution. Without clear knowledge of the diploid ancestor(s) of polyploids, we would not be able to answer fundamental ecological questions such as the evolution of niche differences between them or its underlying genetic basis. Here, we explored the evolutionary history of two Fragaria tetraploids, Fragaria corymbosa and Fragaria moupinensis. We de novo assembled five genomes including these two tetraploids and three diploid relatives. Based on multiple lines of evidence, we found no evidence of subgenomes in either of the two tetraploids, suggesting autopolyploid origins. We determined that Fragaria chinensis was the diploid ancestor of F. corymbosa while either an extinct species affinitive to F. chinensis or an unsampled population of F. chinensis could be the progenitor of F. moupinensis. Meanwhile, we found introgression signals between F. chinensis and Fragaria pentaphylla, leading to the genomic similarity between these two diploids. Compared to F. chinensis, gene families related to high ultraviolet (UV)-B and DNA repair were expanded, while those that responded towards abiotic and biotic stresses (such as salt stress, wounding, and various pathogens) were contracted in both tetraploids. Furthermore, the two tetraploids tended to down-regulate defense response genes but up-regulate UV-B response, DNA repairing, and cell division gene expression compared to F. chinensis. These findings may reflect adaptions toward high-altitude habitats. In summary, our work provides insights into the genome evolution of wild Fragaria tetraploids and opens up an avenue for future works to answer deeper evolutionary and ecological questions regarding the strawberry genus. 

Abstract Background and AimsPollination failure occurs from insufficient pollen quantity or quality. However, the relative contributions of pollen quantity vs. quality to overall pollen limitation, and how this is affected by the co-flowering context, remain unknown for most plant populations. Here, we studied patterns of pollen deposition and pollen tube formation across populations of four predominately outcrossing species in the genus Clarkia to evaluate how the richness of co-flowering congeners affects the contribution of pollen quantity and quality to pollen limitation. MethodsWe partition variation in pollen deposition and pollen tube production across individuals, populations and species to identify the main sources of variation in components of reproductive success. We further quantify the relative contribution of pollen quantity and quality limitation to the reproductive success of the four Clarkia species using piecewise regression analyses. Finally, we evaluate how variation in the number of co-flowering Clarkia species in the community affects the strength of pollen quality and quality limitation. ResultsAcross all contexts, pollen deposition and the proportion of pollen tubes produced varied greatly among individuals, populations and species, and these were not always correlated. For instance, C. xantiana received the smallest pollen loads yet produced the highest proportion of pollen tubes, while C. speciosa exhibited the opposite pattern. Yet, co-flowering richness had variable effects on the strength of pollen quantity and quality limitation among populations. Specifically, breakpoint values, which are an indicator of overall pollen limitation, were two-fold higher in the four-species community compared with one- and two-species communities for two Clarkia species, suggesting that pollen limitation can increase with increasing richness of co-flowering congeners. ConclusionsOur results reveal a complex interplay between the quantity and quality of pollen limitation and co-flowering context that may have different evolutionary outcomes across species and populations. 

Abstract PremisePolyploidy is a widespread mutational process in angiosperms that may alter population performance of not only plants but also their interacting species. Yet, knowledge of whether polyploidy affects plant–herbivore dynamics is scarce. Here, we tested whether aphid herbivores exhibit preference for diploid or neopolyploid plants, whether polyploidy impacts plant and herbivore performance, and whether these interactions depend on the plant genetic background. MethodsUsing independently synthesized neotetraploid strains paired with their diploid progenitors of greater duckweed (Spirodela polyrhiza), we evaluated the effect of neopolyploidy on duckweed's interaction with the water‐lily aphid (Rhopalosiphum nymphaeae). Using paired‐choice experiments, we evaluated feeding preference of the herbivore. We then evaluated the consequences of polyploidy on aphid and plant performance by measuring population growth over multiple generations. ResultsAphids preferred neopolyploids when plants were provided at equal abundances but not at equal surface areas, suggesting the role of plant population surface area in driving this preference. Additionally, neopolyploidy increased aphid population performance, but this result was dependent on the plant's genetic lineage. Lastly, the impact of herbivory on neopolyploid vs. diploid duckweed varied greatly with genetic lineage, where neopolyploids appeared to be variably tolerant compared to diploids, sometimes mirroring the effect on herbivore performance. ConclusionsBy experimentally testing the impacts of polyploidy on trophic species interactions, we showed that polyploidization can impact the preference and performance of herbivores on their plant hosts. These results have significant implications for the establishment and persistence of plants and herbivores in the face of plant polyploidy. 

Whole-genome duplication is a common macromutation with extensive impacts on gene expression, cellular function, and whole-organism phenotype. As a result, it has been proposed that polyploids have “general-purpose” genotypes that perform better than their diploid progenitors under stressful conditions. Here, we test this hypothesis in the context of stresses presented by anthropogenic pollutants. Specifically, we tested how multiple neotetraploid genetic lineages of the mostly asexually reproducing greater duckweed (Spirodela polyrhiza) perform across a favorable control environment and 5 urban pollutants (iron, salt, manganese, copper, and aluminum). By quantifying the population growth rate of asexually reproducing duckweed over multiple generations, we found that across most pollutants, but not all, polyploidy decreased the growth rate of actively growing propagules but increased that of dormant ones. Yet, when considering total propagule production, polyploidy increased tolerance to most pollutants, and polyploids maintained population-level fitness across pollutants better than diploids. Furthermore, broad-sense genetic correlations in growth rate among pollutants were all positive in neopolyploids but not so for diploids. Our results provide a rare test and support for the hypothesis that polyploids are more tolerant of stressful conditions and can maintain fitness better than diploids across heterogeneous stresses. These results may help predict that polyploids may be likely to persist in stressful environments, such as those caused by urbanization and other human activities. 

Abstract Climatic and soil features influence resources and mate availability for plants. Because of different resource/mating demands of the male and female reproductive pathways, environmental variation can drive geographic patterns of sex‐specific factors in sexually polymorphic species. Yet, the relationship between environment and sex, sexual dimorphism or sex chromosomes at the range‐wide scale is underexamined.Using ~7000 herbarium and iNaturalist specimens we generate a landscape‐scale understanding of how sex ratio and sexual dimorphism vary with geographic, climatic and soil gradients in the sexually polymorphic wild strawberry (Fragaria virginiana) and test whether these conform to predictions from theory. Then, for ~300 specimens we use genotyping of the sex‐determining region (SDR haplotypes) to reveal geographic and phenotypic patterns in sex chromosome types.Across North America, the sex ratio was hermaphrodite/male‐biased and was associated more with soil attributes than climate. Sex ratio‐environment associations matched predictions for subdioecy in the West but for gynodioecy in the East. Climatic factors correlated with sexual dimorphism in traits related to carbon acquisition (leaf size and runnering while flowering) but not mate access (petal size, flowering time). Variation in sexual dimorphism was due to one sex being more responsive to the environmental variation than the other. Specifically, leaf length in females was more responsive to variation in precipitation than in hermaphrodite/males, but the probability of runnering while flowering in hermaphrodite/males was more responsive to variation in temperature than in females. The ancestral sex chromosome type was most common overall. But the frequency of the more derived sex chromosomes varied with environmental factors that differed between East–West regions.Synthesis. A landscape‐level perspective revealed that variation in soil and climate factors can explain geospatial variation in sex ratio and sexual dimorphism in a wild strawberry. Variation in sex ratio was associated more with soil resources than climate, while variation in sexual dimorphism was the result of sex‐differential responses to climate for vegetative traits but a similar response to abiotic factors in mate access traits. Finally, sex chromosome types were associated with soil moisture and precipitation in ways that could contribute to the evolution of sex determination. 

Premise of research. Polyploidy, a major evolutionary process in flowering plants, is expected to 19 impact floral traits which can have cascading effects on pollination interactions, but this may 20 depend on selfing propensity. In a novel use of herbarium specimens, we assessed the effects of 21 polyploidy and mating system on floral traits and the pollination niche of 40 Brassicaceae 22 species. 23 Methodology. We combined data on mating system (self-compatible or self-incompatible) with 24 inferred ploidy level (polyploid or diploid) and use phylogenetically controlled analyses to 25 investigate their influence on floral traits (size and shape) and the degree of pollination 26 generalism based on the frequency and the richness of heterospecific pollen morphospecies 27 captured by stigmas. 28 Pivotal Results. Flower size (but not shape) depended on the interaction between ploidy and 29 mating system. Self-incompatible polyploid species had larger flowers than self-incompatible 30 diploids but there was no difference for self-compatible species. The breadth of pollination niche 31 (degree of generalism) was not affected by ploidy but rather strongly by mating system only. 32 Self-incompatible species had more stigmas with heterospecific pollen and higher heterospecific 33 pollen morphospecies richness per stigma than self-compatible species, regardless of their 34 ploidy. 35 Conclusions. Our results demonstrate that mating system moderated the influence of ploidy on 36 morphological features associated with pollination generalism but that response in terms of 37 heterospecific pollen captured as a proxy of pollination generalism was more variable. 

Abstract Background and Aims When plant communities are exposed to herbicide ‘drift’, wherein particles containing the active ingredient travel off-target, interspecific variation in resistance or tolerance may scale up to affect community dynamics. In turn, these alterations could threaten the diversity and stability of agro-ecosystems. We investigated the effects of herbicide drift on the growth and reproduction of 25 wild plant species to make predictions about the consequences of drift exposure on plant-plant interactions and the broader ecological community. Methods We exposed potted plants from species that commonly occur in agricultural areas to a drift-level dose of the widely used herbicide dicamba or a control solution in the glasshouse. We evaluated species-level variation in resistance and tolerance for vegetative and floral traits. We assessed community-level impacts of drift by comparing species evenness and flowering networks of glasshouse synthetic communities comprised of drift-exposed and control plants. Key Results Species varied significantly in resistance and tolerance to dicamba drift: some were negatively impacted while others showed overcompensatory responses. Species also differed in the way they deployed flowers over time following drift exposure. While drift had negligeable effects on community evenness based on vegetative biomass, it caused salient differences in the structure of coflowering networks within communities. Drift reduced the degree and intensity of flowering overlap among species, altered the composition of groups of species that were more likely to coflower with each other than with others, and shifted species roles (e.g., from dominant to inferior floral producers and vice versa). Conclusions These results demonstrate that even low levels of herbicide exposure can significantly alter plant growth and reproduction, particularly flowering phenology. If field-grown plants respond similarly, then these changes would likely impact plant-plant competitive dynamics and potentially plant-pollinator interactions occurring within plant communities at the agro-ecological interface.