Interactive effects between physical forces and ecosystem engineers on seed burial: A case study using Spartina anglica

Seed burial (i.e. vertical seed dispersal) has become increasingly valued for its relevance for seed fate and plant recruitment. While ecosystem engineers have been generally considered as the most important drivers of seed burial, the role of physical forces, such as wind or water flow, has been largely overlooked. Using tidal habitats as a model system, and a combination of flume and mesocosm experiments, we investigated the effects of 1) currents, 2) benthic animals with different engineering activities and 3) their interplay on seed burial of a common salt marsh pioneer plant, Spartina anglica. Our results reveal that in such systems, water flow can be of equal or higher importance than ecosystem engineers for seed burial. For passive seed-burying engineers (PSE), coupling their actions with currents produced synergistic seed burial effects, whereas the interactive effects were only additive for active seed-burying engineers (ASE). This paper extends current understanding of seed burial and seed bank formation by revealing the need to incorporate physical forces into seed burial mechanisms. We provide the first empirical evidence that physical forces influence seed burial by synergistically interacting with ecosystem engineers, thus highlighting the role of biophysical interactions as important drivers for vertical seed movement.