Large-scale ecosystem engineering by flamingos and fiddler crabs on West-African intertidal flats promote joint food availability

Although the ecosystem engineering concept is well established in ecology, cases of joint engineering by multiple species at large scales remain rare. Here, we combine observational studies and exclosure experiments to investigate how co-occurring greater flamingos Phoenicopterus roseus and fiddler crabs Uca tangeri promote their own and each other's food availability by creating a spatially complex mosaic of depressions (bowls, gullies) and hummocks (plateaus, mounds) in the intertidal zone. This results in a mosaic of microhabitats with different tidal inundation regimes. These microhabitats are spatially organized with labyrinth-like patterns in the high intertidal zone and spotted patterns in the lower intertidal, both of which likely arise from biophysical interactions between these organisms and hydrodynamic forces. We show that the resulting spatial complexity is vital for biofilm production. The depression microhabitats were wetter and richer in organic matter and biofilms compared with hummocks. Excluding flamingos and crabs resulted in an increase in biofilm biomass over the shorter term (six months), but a decrease over the longer term (after one year). Moreover, our results strongly suggest that these biogeomorphological microhabitats in the mosaics were maintained by the feeding activities of flamingos and to a lesser extent crabs. During a period of flamingo exclusion, all the spotted patterns filled up with sediment, while the exclusion of crabs led to gradual sediment accumulation in the labyrinth-like patterns. Collectively, these findings provide empirical evidence for large-scale joint promotion of food availability by multiple taxa in a marine ecosystem.