The Moran effect and environmental vetoes: phenological synchrony and drought drive seed production in a Mediterranean oak

Masting is the highly variable production of synchronized seed crops, and is a common reproductive strategy in plants. Weather has long been recognized as centrally involved in driving seed production in masting plants. However, the theory behind mechanisms connecting weather and seeding variation has only recently been developed, and still lacks empirical evaluation. We used 12-year long seed production data for 255 holm oaks (Quercus ilex), as well as airborne pollen and meteorological data, and tested whether masting is driven by environmental constraints: phenological synchrony and associated pollination efficiency, and drought-related acorn abscission. We found that warmsprings resulted in short pollen seasons, and length of the pollen seasons was negatively related to acorn production, supporting the phenological synchrony hypothesis. Furthermore, the relationship between phenological synchrony and acorn production was modulated by spring drought, and effects of environmental vetoes on seed production were dependent on last year's environmental constraint, implying passive resource storage. Both vetoes affectedamong-tree synchrony in seed production. Finally, precipitation preceding acorn maturation was positively related to seed production, mitigating apparent resource depletion following high crop production in the previous year. These results provide new insights into mechanisms beyond widely reported weather and seed production correlations.