Does scale-dependent feedback explain spatial complexity in salt-marsh ecosystems

Complexity theory highlights scale-dependent feedback mechanisms as an explanation for regular spatial patterning in ecosystems. To what extent scale-dependent feedback clarifies spatial structure in more complex, non-regular systems remains unexplored so far. We report on a scale-dependent feedback process generating patchy landscapes at the interface of intertidal flats and salt marshes. Here, vegetation was characterized by Spartina anglica tussocks, surrounded by erosion gullies. To demonstrate the presence of a scale-dependent feedback, we determined if vegetation induced habitat modification resulted in local facilitation and large scale-inhibition of plant growth. Field surveys revealed that larger tussocks have deeper gullies, suggesting that gully erosion is caused by increased water flow around tussocks. This was confirmed by flume experiments, showing that feedback effects vary with current velocity and water depth. Transplantation of small Spartina units inside and just outside present tussocks revealed that the growth of Spartina transplants compared to transplant growth on bare sediment was higher within the raised Spartina tussocks, but lower in the gully just outside Spartina tussocks, providing clear evidence of scale-dependent feedback. Our results emphasize that scale-dependent feedback is a more general explanation for spatial complexity in ecosystems than previously considered.