Plant survival in relation to seed size along environmental gradients: a long-term study from semi-arid and Mediterranean annual plant communities

P>1. A positive relationship between seed size and subsequent offspring survival is a key assumption in ecological theory concerning life-history strategies. Yet, this relationship is uncertain in respect to lifetime survival because sound evidence only exists for early seedling stages. Furthermore, the effect of environmental variation in space and time, and of contrasting plant functional groups, on this relationship has been scarcely examined. 2. Here, we investigated survival and between-year variation therein. We tested for both whether their relationship to seed size (i) is maintained up until reproduction, (ii) changes along environmental gradients and (iii) differs between functional groups (grasses, legumes, forbs). 3. Survival was monitored from established seedlings to reproductive plants in 49 annual species under natural conditions during 7 years in three sites along a steep rainfall gradient. We then related average survival per species and between-year variation in survival to seed size, site along the gradient and functional group. 4. Larger seed size was associated with higher survival and lower between-year variation. Across the rainfall gradient, we detected no difference in the seed size-survival relationships; however, variation between years was lowest in the most mesic site where no relationship for between-year variation with seed size was observed. Legumes showed lower survival and higher between-year variation than grasses. 5. Our findings indicate that larger seed size provides survival advantages beyond seedling establishment up until reproduction among annual species. Larger seed size also provides a bet-hedging strategy in temporally unpredictable environments. Increased abiotic favourability along environmental gradients may have little effect on survival rates but reduces survival variation between years and thus reduces the bet-hedging benefit of larger seed size. We suggest that the contrasting response of legumes and grasses may partly result from their disparity in seed dormancy. 6. Synthesis. Current plant life-history theory can be refined by accounting for both benefits of larger seed size, higher survival rates and bet-hedging. Studies along environmental gradients are needed to generalize findings across ecosystems and to predict patterns of plant traits and plant performance under changing environmental conditions.