Empty native and invasive bivalve shells as benthic habitat modifiers in a large river

Bivalves are remarkable ecosystem engineers and their long-lasting shells may provide important physical structures for benthic organisms. In the last decades the Danube River has experienced great changes in the bivalve fauna, i.e. several native species have been declining and several invasive species have been introduced. The invasive Corbicula fluminea and Sinanodonta woodiana are now widespread and produce large amounts of shells. In this study, we investigated empty shells of native (Anodonta anatina, Unio tumidus) and invasive (C. fluminea, S. woodiana) bivalves (including their mixtures) as benthic substrates and compared them to clay granules (control), which mimics the natural hard substrates in the Danube River (Hungary). Macroinvertebrate colonization was compared between (i) empty shells and control substrate; (ii) different bivalve species (native and invasive) and (iii) three scenarios (before invasion, and short and long time after invasion) by using a mix of empty shells (native, native plus invasive, and invasive species). In comparison to control treatments the empty shells facilitated the presence of amphipods, caddis larvae and isopods, which contributed to a shift in the trophic structure by decreasing the proportion of gathering collectors while increasing the presence of shredders and predators. Several shell traits such as size, outer-shell surface roughness, hardness, thickness, 3D shape and chemical composition may be important attributes in the habitat modifying effects; however, this study could not disentangle which contribute most for the differences found. Given the capability of invasive C. fluminea and S. woodiana to accumulate large amounts of empty shells on several sites of the Danube, its habitat modifying effects can be particularly important, especially on the macroinvertebrate community structure. Moreover, these effects may increase in near future due to the predicted more frequent and severe extreme climatic conditions, which have been responsible for massive mortalities in both species.