Colonization dynamics and facilitative impacts of a nitrogen-fixing shrub in primary succession

Whether nitrogen-fixing plants facilitate or inhibit species change in primary succession is best resolved by examining their impacts throughout the plant's entire life cycle from arrival to senescence. We experimentally examined two aspects of the successional impacts of a nitrogen-fixing shrub, Coriaria arborea, on Mt. Tarawera, a volcano in New Zealand: factors limiting Coriaria colonization and impacts of Coriaria-induced soil changes on a later successional tree, Griselinia littoralis. Coriaria germination was promoted by artificial wind protection and by the presence of heath shrubs. Transplanted Coriaria seedlings survived only if nodulated with Frankia, and the addition of Coriaria-enriched soils slowed Coriaria seedling growth and did not improve seedling survival. This explained why Coriaria seedlings were found mostly in protected habitats away from adult Coriaria, and suggested that Coriaria thickets are not self-replacing. Coriaria increased soil fertility by developing a 4 cm thick organic soil horizon that was richer in nitrogen (tenfold) and phosphorus (threefold) than pre-Coriaria stages. These soil changes resulted in three- to sixfold increases in growth of Griselinia when it was grown in Coriaria-enriched soils in a glasshouse. Coriaria's net effect on primary succession is facilitative, but the establishment of Coriaria is itself facilitated by the amelioration of the physical habitat by earlier colonists, suggesting facilitation is important throughout the life cycle of Coriaria. Sequential facilitative events determine the order of species replacements in this study but inhibition, linked to the developmental stages of Cariaria, may determine the rate of species change.