BACKGROUND: Processes driving ploidal diversity at the population level are virtually unknown. Their identification should use a combination of large-scale screening of ploidy levels in the field, pairwise crossing experiments and mathematical modelling linking these two types of data. We applied this approach to determine the drivers of frequencies of coexisting cytotypes in mixed-ploidy field populations of the fully sexual plant species Pilosella echioides. We examined fecundity and ploidal diversity in seeds from all possible pairwise crosses among 2x, 3x and 4x plants. Using these data, we simulated the dynamics of theoretical panmictic populations of individuals whose progeny structure is identical to that determined by the hybridization experiment. RESULTS: The seed set differed significantly between the crossing treatments, being highest in crosses between diploids and tetraploids and lowest in triploid-triploid crosses. The number of progeny classes (with respect to embryo and endosperm ploidy) ranged from three in the 2x-2x cross to eleven in the 3x-3x cross. Our simulations demonstrate that, provided there is no difference in clonal growth and/or survival between cytotypes, it is a clear case of minority cytotype exclusion depending on the initial conditions with two stable states, neither of which corresponds to the ploidal structure in the field: (i) with prevalent diploids and lower proportions of other ploidies, and (ii) with prevalent tetraploids and 9% of hexaploids. By contrast, if clonal growth differs between cytotypes, minority cytotype exclusion occurs only if the role of sexual reproduction is high; otherwise differences in clonal growth are sufficient to maintain triploid prevalence (as observed in the field) independently of initial conditions. CONCLUSIONS: The projections of our model suggest that the ploidal structure observed in the field can only be reached via a relatively high capacity for clonal growth (and proportionally lower sexual reproduction) in all cytotypes combined with higher clonal growth in the prevailing cytotype (3x).

Department of Botany Natural History Museum National Museum CZ 193 00 Prague Horní Počernice Czech Republic

Institute of Botany The Czech Academy of Sciences CZ 252 43 Průhonice Czech Republic

Institute of Botany The Czech Academy of Sciences CZ 252 43 Průhonice Czech Republic Department of Botany Faculty of Science Charles University Prague CZ 128 01 Prague Czech Republic

Institute of Botany The Czech Academy of Sciences CZ 252 43 Průhonice Czech Republic Department of Botany Faculty of Science Charles University Prague CZ 128 01 Prague Czech Republic Faculty of Agriculture University of South Bohemia CZ 370 05 České Budějovice Czech Republic

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