Zebra mussel effects on benthic invertebrates: physical or biotic?

In soft sediments, Dreissena spp. create firm substrate in the form of aggregates of living mussels (druses) that roll free on the sediments. Druses provide physical structure which increases habitat heterogeneity, and the mussels increase benthic organic matter through the production of pseudofeces and feces. Descriptive and experimental studies were used to determine: 1) whether the density of benthic invertebrates in soft sediments increased in the presence of druses, and 2) whether the invertebrate assemblage responded to the physical structure provided by a druse or to some biotic effect associated with the presence of living mussels. In core samples collected biweekly during summer in Presque Isle Bay, Erie, Pennsylvania, amphipods, chironomids, oligochaetes, turbellarians, and hydrozoans were significantly more abundant in sand with druses than in bare sand. When mesh bags containing either a living druse, non-living druse, or no druse were incubated in the bay for 33 d, we found that chironomids were significantly more abundant in treatments with living druses than with non-living druses, and in treatments with non-living druses than with no druse; turbellarians, amphipods, and hydrozoans were significantly more abundant in treatments with living or non-living druses than with no druse; oligochaetes showed no significant differences among treatments. This study demonstrates that most taxa of benthic invertebrates in soft substrate respond specifically to the physical structure associated with aggregates of mussel shells, but further study is needed to examine chironomid responses to some biotic effect dependent on the presence of living mussels.