Variation in seed packaging of a fleshy-fruited conifer provides insights into the ecology and evolution of multi-seeded fruits.

The study of intraspecific seed packaging (i.e. seed size/number strategy) variation across different populations may allow better understanding of the ecological forces that drive seed evolution in plants. Juniperus thurifera (Cupressaceae) provides a good model to study this due to the existence of two subspecies differentiated by phenotypic traits, such as seed size and cone seediness (number of seeds inside a cone), across its range. The aim of this study was to analyse seed packaging (seed mass and cone seediness) variation at different scales (subspecies, populations and individuals) and the relationship between cone and seed traits in European and African J. thurifera populations. After opening more than 5300 cones and measuring 3600 seeds, we found that seed packaging traits followed different patterns of variation. Large-scale effects (region and population) significantly contributed to cone seediness variance, while most of the seed mass variance occurred within individuals. Seed packaging differed between the two sides of the Mediterranean Sea, with African cones bearing fewer but larger seeds than the European ones. However, no differences in seed mass were found between populations when taking into account cone seediness. Larger cones contained more pulp and seeds and displayed a larger variation in individual seed mass. We validated previous reports on the intraspecific differences in J. thurifera seed packaging, although both subspecies followed the same seed size/number trade-off. The higher seediness and variation in seed mass found in larger cones reveals that the positive relationship between seed and cone sizes may not be straightforward.We hypothesise that the large variation of seed size found within cones and individuals in J. thurifera, but also in other fleshy-fruited species, could represent a bet-hedging strategy for dispersal.