Dormancy and dispersal in dimorphic achenes of tansy ragwort, Senecio jacobaea L. (Compositae)

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Abstract
Marginal and central florets of the capitula of tansy ragwort Senecio jacobaea yield different kinds of fruit. The central (“disk”) achenes are lighter (x±SE=199±5μg), more numerous (x±SE=58±0.6 achenes per head), and are equipped with a pappus aiding wind transport and rows of trichomes aiding animal transport. The marginal (“ray”) achenes are heavier (x±SE=286±7μg), less numerous (virtually invariant at 13 achenes per head), and lack dispersal structures. Whereas disk achenes are relased shortly after they mature, ray achenes are retained by the parent for a period of months following maturity. Germination at constant temperature (20°C) and with alternating light (12 h light: 12 h dark) demonstrated that disk and ray achenes exhibit different germination syndromes. Germination percentage increases linearly with achene fresh weight in both types; for a given weight, disk achenes have a higher germination percentage than ray achenes. Germination time decreases with increasing achene weight in disk achenes, but increases with achene weight in ray achenes. The germination percentages and germination times for disk and ray achenes diverge progressively with increasing achene weight. The divergence in behavior is a result of diverging patterns of dry matter allocation in the two achene types. Increase in the size of disk achenes favors the embryo fraction, thereby speeding germination while reducing protection. Increase in the size of the ray achenes favors the pericarp fraction, thereby increasing protection while delaying germination. Reduced germination percentage and germination speed of the ray achenes were shown by experimental manipulation to be caused by physical inhibition by their thicker pericarps. Dimorphism in ragwort likely speards germination out in space and time, thereby increasing the number of safe sites an individual parent can exploit in disseminating offspring. The syndrome in other heteromorphic composites resembles that of ragwort, generally combining reduced dispersal-delayed germination in the outer achenes and distance dispersal-quick germination in the central achenes. The outer achenes are generally less numerous and larger. Dispersal traits (large numbers, early release and light wieght) are the direct opposite of dormancy traits (small numbers, delayed release and heavier weight). Thus conflicts between the properties determining dormancy and dispersal appear to require separate dormancy and dispersal phenotypes.