Native and non-native unionids respond differently to the presence of fouling dreissenid mussels

Unionid mussels are globally threatened by several human disturbances, including the introduction of non-native species. Among these, biofouling zebra and quagga mussels of Ponto-Caspian origin are considered to be especially detrimental to unionid locomotion, filtration and physical condition. The aim of our study was to determine and compare the impact of dreissenid fouling and/or presence on locomotion and burrowing of the native Unio tumidus and invasive Sinanodonta woodiana , a novel invader expanding its range in Europe in recent decades. We tested unionids collected from Lake Balaton (central Europe) that were fouled by dreissenids (zebra and quagga mussels mixed), cleaned of fouling or non-fouled (collected without any signs of dreissenid fouling). Moreover, unionids were tested in the presence or absence of other fouled individuals and dreissenids isolated in mesh bags to determine the influence of direct fouling and presence of dreissenids in the environment on unionid behaviour. Movement initiation time, locomotion distance and burrowing level were retrieved from videos recorded for 24 hours. Direct fouling affected only the behaviour of U. tumidus , limiting their burrowing and delaying movements. After removal of fouling, movement timing returned back to normal, but mussels still burrowed less than the control non-fouled individuals, indicating persisting effects of fouling on physical condition. Moreover, U. tumidus reduced their locomotion in the presence of fouled unionids. Sinanodonta woodiana responded to the presence of dreissenids (especially quagga mussels) with increased burrowing. These different responses of the two unionid species to Dreissena spp. indicate that biofoulers may influence biotic interactions between the unionids by promoting the invasive species (less susceptible to negative effects of fouling). Moreover, S. woodiana may indirectly affect U. tumidus through apparent competition, constituting an environmental reservoir of biofoulers exerting a stronger impact on the native species.