The Potential Impacts of Invasive Quagga and Zebra Mussels on Macroinvertebrate Communities: An Artificial Stone Substrate Based Field Experiment Using Stable Isotopes

Over the past decades, the zebra mussel (Dreissena polymorpha) and quagga mussel (D. rostriformis bugensis) invaded multiple freshwater systems and posed major threats to the overall ecosystem. In Lake Constance where zebra mussels invaded in the 1960s, the quagga mussel invasion progressed at a very high rate since 2016, providing an opportunity to study the ecological impact of both species at an early stage. We conducted a field experiment in the littoral region of the lake and monitored differences in macroinvertebrate community colonization. We used standardized stone substrates, which were blank, glued with empty shells of mussels, with living adult quagga mussels, and with living adult zebra mussels. Empty shells and the shells of both living adult quagga and zebra mussels created more colonization areas for newly settled macroinvertebrates. The abundance of newly settled quagga mussels was higher than zebra mussels, indicating the outcompeting behavior of quagga mussels. We used stable isotopes (delta C-13 and delta N-15) of both dreissenids and their potential competitors, which include two snail species (New Zealand mud snail Potamopyrgus antipodarum and faucet snail Bithynia tentaculate) and additional invasive gammarid species (killer shrimp Dikerogammarus villosus), in order to investigate their feeding ecology and to evaluate their potential impacts on macroinvertebrate community. The delta C-13 and delta N-15 of neither the newly settled quagga mussels nor the well-established zebra mussels differed significantly among various treatments. Newly settled quagga mussels had higher delta C-13 values than newly settled zebra mussels and showed similar differences in all four stone setups. During the experimental period (with quagga and zebra mussels still coexisting in some regions), these two dreissenids exhibited clear dietary (isotopic) niche segregation. The rapid expansion of invasive quagga mussels coupled with the higher mortality rate of zebra mussels might have caused a dominance shift from zebra to quagga mussels. The study offers the first overview of the progressive invasion of quagga mussel and the reaction of zebra mussels and other newly settled macroinvertebrates, and compliments the hypothesis of facilitative associations between invasive dreissenids. Results provide an experimental benchmark by which future changes in trophic ecology and invasion dynamics can be measured across the ecosystem.