Facilitating your replacement? Ecosystem engineer legacy affects establishment success of an expanding competitor

Interactions with resident species can affect the rate that expanding species invade novel areas. These interactions can be antagonistic (biotic resistance), where resident species hinder invasive establishment, or facilitative (biotic assistance), where residents promote invasive establishment. The predominance of resistance or assistance could vary with the abiotic context. We examined how the effects of a resident ecosystem engineer interact with abiotic stress to resist or assist the establishment of an expanding competitor. In Florida salt marshes, native cordgrass, Spartina alterniflora, is an influential ecosystem engineer that, when dead, exerts a legacy effect by forming persistent wrack patches. We examined how the legacy effect of Spartina wrack varies with spatial context and abiotic conditions to influence establishment of the northward-expanding black mangrove, Avicennia germinans. Field surveys documented that Spartina wrack and Avicennia propagules co-occur in the high intertidal zone, and we conducted two outdoor mesocosm experiments to investigate this association. Wrack positively affected propagule establishment when propagules were beneath wrack, but negatively affected establishment when propagules were above wrack. The abiotic tidal regime influences the magnitude of wrack effects by controlling ambient moisture, and the positive and negative effects of wrack were stronger in low moisture conditions that simulated desiccation stress during harsh neap tides. Thus, the same resident engineer can either resist or assist an expanding competitor and the magnitude of these effects depends on abiotic conditions. We propose that under harsh conditions, there is greater scope for an engineer's mediating influence to affect associated species, both positively and negatively.