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The role of the tolerance-fecundity trade-off in maintaining intraspecific seed trait variation in a widespread dimorphic herb.

Coexistence of species with different seed sizes is a long-standing issue in community ecology, and a trade-off between fecundity and stress tolerance has been proposed to explain co-occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade-off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade-off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life-cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance-fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size-number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.