High Selfing Research Articles

Evolutionary breakdown of distyly to homostyly is accompanied by reductions of floral scent inPrimula oreodoxa

AbstractReproductive traits that function in pollinator attraction may be reduced or lost during evolutionary transitions from outcrossing to selfing. Although floral scent plays an important role in attracting pollinators in outcrossing species, few studies have investigated associations between floral scent variation and intraspecific mating system transitions. The breakdown of distyly to homostyly represents a classic example of a shift from outcrossing to selfing and provides an opportunity to test whether floral fragrances have become reduced and/or changed in composition with increased selfing. Here, we evaluate this hypothesis by quantifying floral volatiles using gas chromatography‐mass spectrometry in two distylous and four homostylous populations ofPrimula oreodoxaFranchet, a perennial herb from SW China. Our analysis revealed significant variation of volatile organic compounds (VOCs) among populations ofP. oreodoxa. Although there was no difference in VOCs between floral morphs in distylous populations as predicted, we detected a substantial reduction in VOC emissions and the average number of scent compounds in homostylous compared with distylous populations. A total of 12 compounds, mainly monoterpenoids and sesquiterpenoids, distinguished homostylous and distylous morphs; of these, (E)‐β‐ocimene was the most important in contributing to the difference in volatiles, with significantly lower emissions in homostyles. Our findings support the hypothesis that the transition from outcrossing to selfing is accompanied by the loss of floral volatiles. The modification to floral fragrances inP. oreodoxaassociated with mating system change might occur because high selfing rates in homostylous populations result in relaxed selection for floral attractiveness.

Why are deleterious mutations maintained in selfing populations? An analysis of the effects of early- and late-acting mutations by a two-locus two-allele model

Frequencies of deleterious mutations are higher than expected in many plants. Here, by developing a two-locus two-allele model, I examine the effects of differential timing of the expression of deleterious mutations (two-stage effects) on the maintenance of mutations. I assume early- and late-acting loci to distinguish whether maintenance of mutations in populations with high selfing rates is explained better by two-stage effects of single mutations, or by separate mutations in both early- and late-acting loci. I found that, when ovules are overproduced, the stable frequency of early-acting mutations is higher if mutations also occur in a late-acting locus than if a late-acting mutation is lacking. The stable frequency of late-acting mutations is higher if mutations also occur in an early-acting locus than if an early-acting mutation is lacking. Selective interference does not account for these results because analyses in which the number of loci subject to mutations is equalized are included. Overproduction of ovules has little effect on maintenance if either early- or late-acting mutations are lacking, whereas when ovules are not overproduced, the two-stage effect does not enhance the maintenance of mutations. Hence, mutations occurring in both loci coupled with overproduction of ovules enhances the maintenance of mutations in populations with high selfing rates. The detailed mechanisms underlying the two-stage effect were also analyzed.

Pollination efficiency and pollen limitation in bistaminate vs tetrastaminate Lamiaceae

AbstractThe evolution of bilabiate flowers in Lamiaceae has several times led to a reduction in stamen number. It is generally assumed that stamen reduction goes along with decreasing pollen production. One should, therefore, expect that bistaminate flowers have either a more efficient pollen transfer mechanism or a higher selfíng rate than tetrastaminate species to adapt to the stamen reduction. We tested this hypothesis in a community of Lamiaceae including three bistaminate and three tetrastaminate species. The results of selfing experiments, extra‐pollination, counting pollen number, and stigmatic pollen load analysis were used to calculate the autonomous selfing index, the pollen limitation index, and the pollination efficiency index. The results indicate a significantly lower pollen number in bistaminate flowers than in tetrastaminate flowers of similar anther size. Bistaminate species shared a higher selfing rate which was related to their structural adaptation to delayed selfing. In Ziziphora clinopodioides, the delayed selfing was accompanied by low pollination efficiency, and plants suffered from pollen limitation at the location studied. However, in Salvia sclarea and Salvia hypoleuca, which benefited from a relatively high pollination efficiency, no pollen limitation was observed. Our results reveal a close association between adaptive mechanisms such as increasing precision of pollen transfer and evolution of delayed selfing with stamen reduction in bistaminate Lamiaceae.

Migration without interbreeding: Evolutionary history of a highly selfing Mediterranean grass inferred from whole genomes.

Wild plant populations show extensive genetic subdivision and are far from the ideal of panmixia which permeates population genetic theory. Understanding the spatial and temporal scale of population structure is therefore fundamental for empirical population genetics – and of interest in itself, as it yields insights into the history and biology of a species. In this study we extend the genomic resources for the wild Mediterranean grass Brachypodium distachyon to investigate the scale of population structure and its underlying history at whole‐genome resolution. A total of 86 accessions were sampled at local and regional scales in Italy and France, which closes a conspicuous gap in the collection for this model organism. The analysis of 196 accessions, spanning the Mediterranean from Spain to Iraq, suggests that the interplay of high selfing and seed dispersal rates has shaped genetic structure in B. distachyon. At the continental scale, the evolution in B. distachyon is characterized by the independent expansion of three lineages during the Upper Pleistocene. Today, these lineages may occur on the same meadow yet do not interbreed. At the regional scale, dispersal and selfing interact and maintain high genotypic diversity, thus challenging the textbook notion that selfing in finite populations implies reduced diversity. Our study extends the population genomic resources for B. distachyon and suggests that an important use of this wild plant model is to investigate how selfing and dispersal, two processes typically studied separately, interact in colonizing plant species.

So close, so far: spatial genetic structure and mating system inPetunia exserta, an endemic from a peculiar landscape in the Brazilian Pampa grasslands

AbstractGene flow via seed or pollen dispersal is fundamental for establishing population diversity and structure of plants, especially in naturally fragmented environments. Petunia exserta (Solanaceae) is endemic to small shelters in rocky towers in the Brazilian Pampa grassland, an ancient and isolated region. The landscape is a long-term fragmented habitat, and ecological conditions inside the shelters constitute an inhospitable environment for other Petunia spp., which usually inhabit open and sunny grasslands. We aimed to evaluate the mating system and gene flow impact on genetic diversity and population structure in P. exserta throughout its geographical range. We used eight microsatellite markers to employ fine-scale genetic structure and paternity analyses in 15 populations, including 361 adults and 244 progeny. Our results showed that P. exserta has low genetic diversity and a homozygous excess compared with its congeners. We identified four genetic clusters that did not reflect the spatial population distribution and a strong genetic structure at the first spatial distance. Pollen and seed dispersal mainly occurred at short distances, and the species has a mixed mating system with high selfing levels. We did not observe recent population reduction, and most population clusters showed a small effective population size. The landscape micro-habitat features contribute to pollen flow that occurs mainly inside shelters through geitonogamy or biparental inbreeding. The self-compatible status of P. exserta and related lineages could be important in the colonization of a new environment for the genus.

Faster Rates of Molecular Sequence Evolution in Reproduction-Related Genes and in Species with Hypodermic Sperm Morphologies.

Sexual selection drives the evolution of many striking behaviors and morphologies and should leave signatures of selection at loci underlying these phenotypes. However, although loci thought to be under sexual selection often evolve rapidly, few studies have contrasted rates of molecular sequence evolution at such loci across lineages with different sexual selection contexts. Furthermore, work has focused on separate sexed animals, neglecting alternative sexual systems. We investigate rates of molecular sequence evolution in hermaphroditic flatworms of the genus Macrostomum. Specifically, we compare species that exhibit contrasting sperm morphologies, strongly associated with multiple convergent shifts in the mating strategy, reflecting different sexual selection contexts. Species donating and receiving sperm in every mating have sperm with bristles, likely to prevent sperm removal. Meanwhile, species that hypodermically inject sperm lack bristles, potentially as an adaptation to the environment experienced by hypodermic sperm. Combining functional annotations from the model, Macrostomum lignano, with transcriptomes from 93 congeners, we find genus-wide faster sequence evolution in reproduction-related versus ubiquitously expressed genes, consistent with stronger sexual selection on the former. Additionally, species with hypodermic sperm morphologies had elevated molecular sequence evolution, regardless of a gene’s functional annotation. These genome-wide patterns suggest reduced selection efficiency following shifts to hypodermic mating, possibly due to higher selfing rates in these species. Moreover, we find little evidence for convergent amino acid changes across species. Our work not only shows that reproduction-related genes evolve rapidly also in hermaphroditic animals, but also that well-replicated contrasts of different sexual selection contexts can reveal underappreciated genome-wide effects.

Self-Fertilization, Inbreeding, and Yield in Alfalfa Seed Production.

Selfing (self-pollination) is the ultimate form of inbreeding, or mating among close relatives. Selfing can create yield loss when inbreeding depression, defined as a lower survival and reproduction of inbred relative to outbred progeny, is present. To determine the impact of selfing in alfalfa (Medicago sativa L.), we quantified the selfing rate of 32 alfalfa seed production fields located in three regions, namely, the Pacific Northwest (PNW), the Central Valley of California (CEV), and the Imperial Valley of California (IMP). Selfing rates (the proportion of selfed seeds) varied between 5.3 and 30% with an average of 12.2% over the 32 seed production fields. In both the parents and their progeny, we observed an excess of heterozygotes relative to Hardy–Weinberg expectations. We detected notable levels of inbreeding in parents (0.231 ± 0.007 parental inbreeding coefficient) and progeny (0.229 ± 0.005). There were a 15% decrease in the number of seeds per stem (seed set) and a 13% decline in the number of seeds per pod in selfed relative to outcrossed stems, but negligible inbreeding depression for pods per raceme and seed weight. The number of racemes on selfed stems increased significantly in fields with greater selfing rates, supporting the presence of geitonogamous or among flower selfing. Despite the significant level of inbreeding depression, seed set did not decrease in fields with higher selfing rates, where the greater number of racemes on the selfed stems increased the seed set. The effects of the field selfing rate on the seed yield metrics were mostly indirect with direct effects of the number of racemes per stem. Available data indicate that the majority of selfing in alfalfa is pollinator-mediated, and thus, eliminating selfing in alfalfa seed production would require the selection of self-incompatible varieties, which, by eliminating inbreeding depression, would provide a 15% potential increase in seed yield and an increase in future hay yield.

New genomic resources and comparative analyses reveal differences in floral gene expression in selfing and outcrossing Collinsia sister species.

The evolutionary transition from outcross- to self-fertilization is one of the most common in angiosperms and is often associated with a parallel shift in floral morphological and developmental traits, such as reduced flower size and pollen to ovule ratios, known as the “selfing syndrome.” How these convergent phenotypes arise, the extent to which they are shaped by selection, and the nature of their underlying genetic basis are unsettled questions in evolutionary biology. The genus Collinsia (Plantaginaceae) includes seven independent transitions from outcrossing or mixed mating to high selfing rates accompanied by selfing syndrome traits. Accordingly, Collinsia represents an ideal system for investigating this parallelism, but requires genomic resource development. We present a high quality de novo genome assembly for the highly selfing species Collinsia rattanii. To begin addressing the basis of selfing syndrome developmental shifts, we evaluate and contrast patterns of gene expression from floral transcriptomes across three stages of bud development for C. rattanii and its outcrossing sister species Collinsia linearis. Relative to C. linearis, total gene expression is less variable among individuals and bud stages in C. rattanii. In addition, there is a common pattern among differentially expressed genes: lower expression levels that are more constant across bud development in C. rattanii relative to C. linearis. Transcriptional regulation of enzymes involved in pollen formation specifically in early bud development may influence floral traits that distinguish selfing and outcrossing Collinsia species through pleiotropic functions. Future work will include additional Collinsia outcrossing-selfing species pairs to identify genomic signatures of parallel evolution.

Mating system of Datura inoxia: association between selfing rates and herkogamy within populations.

Plant mating system determines, to a great extent, the demographic and genetic properties of populations, hence their potential for adaptive evolution. Variation in plant mating system has been documented between phylogenetically related species as well between populations of a species. A common evolutionary transition, from outcrossing to selfing, is likely to occur under environmental spatial variation in the service of pollinators. Here, we studied two phenotypically (in floral traits) and genetically (in neutral molecular markers) differentiated populations of the annual, insect-pollinated, plant Datura inoxia in Mexico, that differ in the service of pollinators (Mapimí and Cañada Moreno). First, we determined the populations’ parameters of phenotypic in herkogamy, outcrossing and selfing rates with microsatellite loci, and assessed between generation (adults and seedlings) inbreeding, and inbreeding depression. Second, we compared the relationships between parameters in each population. Results point strong differences between populations: plants in Mapimí have, on average, approach herkogamy, higher outcrossing rate (tm = 0.68), lower primary selfing rate (r = 0.35), and lower inbreeding at equilibrium (Fe = 0.24) and higher inbreeding depression (δ = 0.25), than the populations of Cañada. Outcrossing seems to be favored in Mapimí while selfing in Cañada. The relationship between r and Fe were negatively associated with herkogamy in Mapimí; here, progenies derived from plants with no herkogamy or reverse herkogamy had higher selfing rate and inbreeding coefficient than plants with approach herkogamy. The difference Fe–F is positively related to primary selfing rate (r) only in Cañada Moreno which suggests inbreeding depression in selfing individuals and then genetic purging. In conclusion, mating system evolution may occur differentially among maternal lineages within populations of Datura inoxia, in which approach herkogamy favors higher outcrossing rates and low levels of inbreeding and inbreeding depression, while no herkogamy or reverse herkogamy lead to the evolution of the “selfing syndrome” following the purge of deleterious alleles despite high inbreeding among individuals.

Similar genetic diversity but increased differentiation revealed among a 58-year-old Pinus massoniana seed-tree stand and its progenies generated at different ages

The genetic diversity and mating status of seed production stands are crucial factors impacting the genetic quality and ecological stability of progeny plantations. The genetic homogeneity of seeds collected from a seed production population in different years has not yet been clearly described. In this study, the genetic diversity and differentiation of a seed-tree stand of 58-year-old Pinus massoniana (P1959) and four-open pollinated plantations of its progeny (OS1996, OS2000, OS2006 and OS2015) were determined by using microsatellite markers. The results showed that similar high genetic diversity was preserved in the seed-tree stand and its four progenies. This implied that current artificial seed harvesting from the seed-tree stand for reforestation does not lead to a significant reduction in genetic diversity; the F index across most loci was significantly greater than zero, indicating population homozygote excess. The pairwise genetic differentiation of the five populations ranged from 0.035 to 0.075, among which OS2015 was the most differentiated from the parent. The differentiation between the parental population and its progeny seemed to increase with increasing age of the seed-trees. This implied that a limited pollination coupled with variable within-stand pollen availability has created opportunities for both self- and foreign pollination to be successful in various periods, resulting in a seemingly paradoxical scenario of both high selfing and gene flow, ultimately resulting in gradually increasing differences in the gene frequency composition between the parental population and its progeny. Management considerations in addressing these problems in similar seed production stands were discussed.

How rapidly do self-compatible populations evolve selfing? Mating system estimation within recently evolved self-compatible populations of Azorean Tolpis succulenta (Asteraceae).

Genome‐wide genotyping and Bayesian inference method (BORICE) were employed to estimate outcrossing rates and paternity in two small plant populations of Tolpis succulenta (Asteraceae) on Graciosa island in the Azores. These two known extant populations of T. succulenta on Graciosa have recently evolved self‐compatibility. Despite the expectation that selfing would occur at an appreciable rate (self‐incompatible populations of the same species show low but nonzero selfing), high outcrossing was found in progeny arrays from maternal plants in both populations. This is inconsistent with an immediate transition to high selfing following the breakdown of a genetic incompatibility system. This finding is surprising given the small population sizes and the recent colonization of an island from self‐incompatible colonists of T. succulenta from another island in the Azores, and a potential paucity of pollinators, all factors selecting for selfing through reproductive assurance. The self‐compatible lineage(s) likely have high inbreeding depression (ID) that effectively halts the evolution of increased selfing, but this remains to be determined. Like their progeny, all maternal plants in both populations are fully outbred, which is consistent with but not proof of high ID. High multiple paternity was found in both populations, which may be due in part to the abundant pollinators observed during the flowering season.

Seasonal change of flower sex ratio and pollinator dynamics in three reproductive ecotypes of protandrous plant

AbstractCimicifuga simplex has three genetically and ecologically distinct pollination morphs with different flowering phenology, flower sex expressions, and selfing rates. A previous study showed that strong protandry in hermaphroditic flowers of C. simplex causes there to be seasonal minority sexes; for example, unisexual female ramets are advantageous in the first half of the flowering season and bloom then (minority sex hypothesis). That study, however, did not distinguish among the three pollination morphs of C. simplex. We investigated seasonal sex ratio changes and pollinator environments of the three morphs to verify and expand the minority sex hypothesis. We investigated flowering phenology, pollinator quantity and quality, seasonal population sex ratio dynamics, and stamen/ovule ratios in hermaphroditic flowers. We also examined the seasonal female reproductive success of hermaphroditic flowers, and the effect of male flower excision on fruit set by morph II hermaphroditic flowers. Morph I (mainly hermaphroditic and female ramets) had high pollinator quality and quantity. Fruit set of hermaphroditic ramets was high throughout the flowering season. Morph II (hermaphroditic and andromonoecious ramets) had low pollinator quality, and few pollinators visited near the beginning and end of the season. Removal of male flowers led to a pollen limitation. Morph III (hermaphroditic ramets and a high selfing rate) had very low pollinator abundance throughout the flowering season, and male and female phases largely overlapped. The scarcity of male flowers in morph I is likely a result of the high pollinator quantity and quality. The high fruit set suggests that no pollen limitation existed. The low pollinator quality and quantity of the morph II population caused it to suffer from pollen limitation and may explain the absence of unisexual female ramets in this morph. The high selfing rate of morph III may be due to the extremely low pollinator abundance. Theoretical studies have indicated that the optimal investment allocation to male and female functions depends on whether a pollen limitation exists. In C. simplex, the observed relationships between pollinator environment and reproductive systems are consistent with these theoretical models.

Hidden genetic variance contributes to increase the short-term adaptive potential of selfing populations.

Standing genetic variation is considered a major contributor to the adaptive potential of species. The low heritable genetic variation observed in self-fertilizing populations has led to the hypothesis that species with this mating system would be less likely to adapt. However, a non-negligible amount of cryptic genetic variation for polygenic traits, accumulated through negative linkage disequilibrium, could prove to be an important source of standing variation in self-fertilizing species. To test this hypothesis, we simulated populations under stabilizing selection subjected to an environmental change. We demonstrate that, when the mutation rate is high (but realistic), selfing populations are better able to store genetic variance than outcrossing populations through genetic associations, notably due to the reduced effective recombination rate associated with predominant selfing. Following an environmental shift, this diversity can be partially remobilized, which increases the additive variance and adaptive potential of predominantly (but not completely) selfing populations. In such conditions, despite initially lower observed genetic variance, selfing populations adapt as readily as outcrossing ones within a few generations. For low mutation rates, purifying selection impedes the storage of diversity through genetic associations, in which case, as previously predicted, the lower genetic variance of selfing populations results in lower adaptability compared to their outcrossing counterparts. The population size and the mutation rate are the main parameters to consider, as they are the best predictors of the amount of stored diversity in selfing populations. Our results and their impact on our knowledge of adaptation under high selfing rates are discussed.

Maintenance of high inbreeding depression in selfing populations: Two-stage effect of early- and late-acting mutations

High estimates of inbreeding depression have been obtained in many plant populations with high selfing rates. However, deleterious mutations might be purged from such populations as a result of selfing. I developed a simulation model assuming the presence of mutations at two sets of loci, namely, early- and late-acting loci, and the selective abortion of embryos coupled with ovule overproduction. In the model, early-acting loci are expressed during embryo initiation, and less vigorous embryos are aborted. Late-acting loci are expressed after selective abortion ends; the surviving embryos (seeds) compete, and some of them form the next generation. If mutations are allowed to occur in both early- and late-acting loci, both types increase in frequency in populations with high selfing rates. However, this phenomenon does not occur if mutations occur only in the early- or only in the late-acting loci. Consistent results are observed even if the total number of loci in which mutations are allowed to occur is the same for simulations with both early- and late-acting loci, only early-acting loci, or only late-acting loci, indicating that the presence of both types of loci is the causal factor. Thus, the two-stage effect, or occurrence of both early- and late-acting mutations, promotes the maintenance of these mutations in populations with high selfing rates.

Management Practices and Breeding History of Varieties Strongly Determine the Fine Genetic Structure of Crop Populations: A Case Study Based on European Wheat Populations

As the effects of climate change begin to be felt on yield stability, it is becoming essential to promote the use of genetic diversity in farmers’ fields. The presence of genetic variability in variety could fulfil this purpose. Indeed, the level of intra-varietal genetic diversity influences the spatio-temporal stability of yields and the disease susceptibility of crop species. Breeding history of varieties and their management practices are two factors that should influence intra-varietal genetic diversity. This paper describes the genetic diversity of eight wheat samples covering a gradient from modern single varieties to on-farm mixtures of landraces. This gradient discriminates between landrace, historical and modern varieties, considering the breeding history of varieties, between single-varieties and mixtures of varieties, and between ex situ and in situ de facto strategy in terms of management practices. Genetic diversity of these samples was analyzed with the help of 41 single nucleotide polymorphism markers located in neutral regions, through computing genetic indices at three different levels: Allelic, haplotypic and genetic group level. Population structure and kinship were depicted using discriminant analysis and kinship network analysis. Results revealed an increase in the complexity of the genetic structure as we move on the gradient of variety types (from modern single variety to in situ on-farm mixtures of landraces). For the landraces, the highest levels of genetic diversity have been observed for a landrace (Solina d’Abruzzo) continuously grown on-farm in the region of Abruzzo, in Italy, for many decades. This landrace showed an excess of haplotypic diversity compared to landraces or the historical variety that were stored in genebanks (ex situ conservation). Genetic analyses of the mixtures revealed that, despite a very high selfing rate in wheat, growing in evolutionary mixtures promotes recombination between different genetic components of the mixture, a second way to increase the level of haplotype diversity. When management practices such as growing in mixture and on-farm management are combined, they substantially increase the different levels of genetic diversity of the populations (allelic, haplotypic, genetic group diversity), and consequently promote their adaptability. Our results confirm the need to develop and manage evolving diversified large populations on-farm. These results invite crop diversity managers such as genebank curators, community seed bank managers and farmers’ organizations to adapt their management strategies to the type of variety they wish to manage, because we have shown that their choices have a strong influence on the genetic composition of the crop populations.

Selfing and Drought-Stress Strategies Under Water Deficit for Two Herbaceous Species in the South American Andes.

Angiosperms are highly diverse in their reproductive systems, including predominantly selfing, exclusive outcrossing, and mixed mating systems. Even though selfing can have negative consequences on natural populations, it has been proposed that plants having a predominantly selfing strategy are also associated with fast development strategies through time limitation mechanisms that allow them to complete their life cycle before the onset of severe drought. This relationship might be affected by the challenges imposed by global change, such as a decrease in pollinator availability and the earlier and more severe onset of droughts. In this work, our aim was to investigate whether selfing is correlated with a dehydration avoidance strategy, and how this could affect drought resistance and survival in two species with different types of selfing: pollinator-independent delayed selfing (Schizanthus grahamii) and pollinator-dependent selfing (Schizanthus hookeri), representing a gradient in selfing rates. We hypothesize that delayed selfing species and highly selfing populations will show “fast” plant traits whereas we will find no pattern in more outcrossed populations of the pollinator-dependent species. However, we predicted that high selfing populations would have lower survival rates when exposed to chronic drought early in their development since fast traits imply physiological compromises that will affect their drought survival. To evaluate these hypotheses, we characterized different physiological and morphological traits in response to two contrasting treatments (moist and dry) in a total of six populations of the two species. We found a relationship between the delayed selfing species and a dehydration avoidance strategy and also with low drought survival. Our work offers evidence to support the importance of abiotic factors, such as drought, on the possible variation in selfing rates on natural populations, and the effect that this mating system could have in their ability to face new environmental conditions such as those imposed by climate change.

Multiple founder events explain the genetic diversity and structure of the model allopolyploid grass Brachypodium hybridum in the Iberian Peninsula hotspot.

It is accepted that contemporary allopolyploid species have originated recurrently, but very few cases have been documented using multiple natural formations of the same species. To extend our knowledge, we have investigated the multiple origins, genetic variation and structure of the allotetraploid grass Brachypodium hybridum with respect to its progenitor diploid species B. distachyon (D genome) and B. stacei (S genome). For this, our primary focus is the Iberian Peninsula, an evolutionary hotspot for the genus Brachypodium. We analysed 342 B. hybridum individuals from 36 populations using ten nuclear SSR loci and two plastid loci. The B. hybridum genetic profiles were compared with those previously reported for B. stacei and B. distachyon. In addition, phylogenetic analysis of the plastid data was performed for a reduced subset of individuals. The nuclear simple sequence repeat (SSR) genetic analysis detected medium to high genetic diversity, with a strong south-to-north genetic structure cline, and a high selfing rate in B. hybridum. Comparative genetic analysis showed a close relatedness of current B. hybridum D allelic profiles with those of B. distachyon, but a lack of similarity with those of B. stacei, suggesting another B. stacei source for the B. hybridum S alleles. Plastid analysis detected three different bidirectional allopolyploidization events: two involved distinct B. distachyon-like ancestors and one involved a B. stacei-like ancestor. The south-eastern Iberian Peninsula B. hybridum populations were more genetically diverse and could have originated from at least two hybridization events whereas north-eastern/north-western Iberian Peninsula B. hybridum populations were less diverse and may have derived from at least one hybridization event. The genetic and evolutionary evidence supports the plausible in situ origin of the south-eastern and northern Iberian Peninsula B. hybridum allopolyploids from their respective local B. distachyon and unknown B. stacei ancestors. The untapped multiple origins and genetic variation detected in these B. hybridum populations opens the way to future evolutionary analysis of allopolyploid formation and genomic dominance and expression in the B. hybridum-B. distachyon-B. stacei grass model complex.

Marker‐assisted parentage analysis reveals high individual selfing rates in tetraploid red clover genotypes selected for seed yield

AbstractSeed yield is a major breeding target in tetraploid red clover. We investigated if marker‐assisted parentage analysis can identify progeny plants with two high seed‐yielding parents in tetraploid red clover and if this technique is more advantageous than traditional half‐sib selection. Parentage analysis was successfully performed on the progeny from the 10% highest seed‐yielding genotypes from a second‐cycle family selection trial: 16.0% of progeny were identified with a high seed‐yielding father. However, progeny plants with two high seed‐yielding parents did not produce more seeds than traditionally selected progeny (27.3 g vs. 30.7 g/plant, respectively). The 10% highest seed‐yielding genotypes displayed on average 2% self‐fertilization. Four genotypes were self‐fertile with individual selfing rates up to 20%. Our results discourage the use of marker‐assisted parentage analysis to improve seed yield in tetraploid red clover when the material has been preselected for seed yield. Breeders should be aware that intensive selection for seed yield in tetraploid red clover may inadvertently lead to selection for increased self‐fertility, which may increase inbreeding in the long term.

Climate outweighs native vs. nonnative range‐effects for genetics and common garden performance of a cosmopolitan weed

AbstractComparing genetic diversity, genetic differentiation, and performance between native and nonnative populations has advanced our knowledge of contemporary evolution and its ecological consequences. However, such between‐range comparisons can be complicated by high among‐population variation within native and nonnative ranges. For example, native vs. nonnative comparisons between small and non‐representative subsets of populations for species with very large distributions have the potential to mislead because they may not sufficiently account for within‐range adaptation to climatic conditions, and demographic history that may lead to non‐adaptive evolution. We used the cosmopolitan weed Conyza canadensis to study the interplay of adaptive and demographic processes across, to our knowledge, the broadest climatic gradient yet investigated in this context. To examine the distribution of genetic diversity, we genotyped 26 native and 26 nonnative populations at 12 microsatellite loci. Furthermore, we recorded performance traits for 12 native and 13 nonnative populations in the field and in the common garden. To analyze how performance was related to range and/or climate, we fit pedigree mixed‐effects models. These models weighed the population random effect for co‐ancestry to account for the influence of demographic history on phenotypic among‐population differentiation. Genetic diversity was very low, selfing rates were very high, and both were comparable between native and nonnative ranges. Nonnative populations out‐performed native populations in the field. However, our most salient result was that both neutral genetic differentiation and common garden performance were far more correlated with the climatic conditions from which populations originated than native vs. nonnative range affiliation. Including co‐ancestry of our populations in our models greatly increased explained variance and our ability to detect significant main effects for among‐population variation in performance. High propagule pressure and high selfing rates, in concert with the ability to adapt rapidly to climatic gradients, may have facilitated the global success of this weed. Neither native nor nonnative populations were homogeneous groups but responded comparably to similar environments in each range. We suggest that studies of contemporary evolution should consider widely distributed and genotyped populations to disentangle native vs. nonnative range effects from varying adaptive processes within ranges and from potentially confounding effects of demographic history.

Dispersal and local persistence shape the genetic structure of a widespread Neotropical plant species with a patchy distribution.

Isolated populations constitute an ideal laboratory to study the consequences of intraspecific divergence, because intrinsic incompatibilities are more likely to accumulate under reduced gene flow. Here, we use a widespread bromeliad with a patchy distribution, Pitcairnia lanuginosa, as a model to infer processes driving Neotropical diversification and, thus, to improve our understanding of the origin and evolutionary dynamics of biodiversity in this highly speciose region. We assessed the timing of lineage divergence, genetic structural patterns and historical demography of P. lanuginosa, based on microsatellites, and plastid and nuclear sequence data sets using coalescent analyses and an Approximate Bayesian Computation framework. Additionally, we used species distribution models (SDMs) to independently estimate potential changes in habitat suitability. Despite morphological uniformity, plastid and nuclear DNA data revealed two distinct P. lanuginosa lineages that probably diverged through dispersal from the Cerrado to the Central Andean Yungas, following the final uplift of the Andes, and passed through long-term isolation with no evidence of migration. Microsatellite data indicate low genetic diversity and high levels of inbreeding within populations, and restricted gene flow among populations, which are likely to be a consequence of bottlenecks (or founder events), and high selfing rates promoting population persistence in isolation. SDMs showed a slight expansion of the suitable range for P. lanuginosa lineages during the Last Glacial Maximum, although molecular data revealed a signature of older divergence. Pleistocene climatic oscillations thus seem to have played only a minor role in the diversification of P. lanuginosa, which probably persisted through adverse conditions in riparian forests. Our results imply drift as a major force shaping the evolution of P. lanuginosa, and suggest that dispersal events have a prominent role in connecting Neotropical open and forest biomes.