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.

Biological invasions cause biodiversity erosion on a global scale. Invasive species spreading beyond their natural range compete with native fauna for food and space, push native species to suboptimal habitats, impairing their behaviour and thus limiting their occurrence. Freshwater ecosystems are especially vulnerable to biological invasions and their ecological and economic impacts. The invasive Asian clams (Corbicula spp.), due to their opportunistic life style, can occur at densities of thousands ind. m-2. They act as ecosystem engineers transforming bottom substrata through accumulation of shells. Our goal was to determine the effect of substratum modification by living Corbicula and their shells on substratum choice and behaviour of Unio tumidus and Anodonta anatina, two European freshwater mussel species of the highly imperilled Unionidae family. We assessed their substratum selection in pairwise choice tests (pure sand vs. sand modified by living Corbicula or their shells, sand modified by shells vs. living Corbicula). Next, we tested locomotion and burrowing of unionids on pure substratum and substrata modified by Corbicula. Unionids avoided sand modified by living Corbicula and their empty shells, not distinguishing between these two types of substratum modification. In the presence of Corbicula, their burrowing was shallower or it took them longer to obtain the same depth as in the pure sand. Additionally, on sand modified by Corbicula shells, we observed a locomotion increase (U. tumidus) or slowing down (A. anatina). Our research showed a novel mechanism of negative impact of Corbicula on unionids, consisting in pushing them away from their optimal habitats. This may contribute to their habitat loss and future declines in invaded ecosystems.

1. Ecosystem engineering freshwater bivalves, burrowing in the substratum and accumulating shell deposits, transform bottom habitats. Especially the invasive Asian bivalve Sinanodonta woodiana (SW), due to its rapid growth, large size, and high fecundity, can affect benthic communities. Here, we determined its effect on habitat selection and behaviour of endangered native bivalves, Anodonta cygnea and Unio tumidus. 2. We conducted laboratory preference assays (Experiment 1: choice between two substrata) exposing the native bivalves to pure sand (control), shells (several densities on the sand surface or burrowed), or living SW. Then, we tested their locomotion and burrowing (Experiment 2) on pure sand and substrata contaminated with shells or living SW. 3. In Experiment 1, native bivalves avoided shells, but not living SW. Burrowed and larger shells were avoided compared with those on the surface and smaller ones, respectively. 4. In Experiment 2, U. tumidus exposed to SW delayed activity initiation (in response to living bivalves), increased locomotion (living bivalves, surface shells), and reduced burrowing depth (living bivalves, all shells). Anodonta cygnea exposed to SW reduced locomotion speed (living bivalves, shells), and reduced burrowing duration (burrowed shells) and depth (living bivalves, burrowed shells). 5. SW (especially shell beds) constitutes another emerging threat to native bivalves, impairing their burrowing and inducting active avoidance. As SW expands its distribution with climate warming, the range and strength of its impact is likely to increase, reducing the area available to native bivalves, exposing them to environmental dangers (due to burrowing limitation) and deteriorating physical condition (energetic resources used for excessive locomotion).

Clams from the Corbicula species complex of Asian origin are invasive all over the world, currently extending their distribution to waters of the natural thermal regime in Central European areas. Features linking the clams to their habitats in newly invaded areas have not been fully examined. These include substratum preferences and mobility, which may be key to determining potential niche overlap with native bivalves and gaining a better knowledge of their potential active dispersal and impact. Therefore, our aim was to determine the substratum preferences, horizontal locomotion and burrowing behaviour of three Corbicula morphospecies from Poland, Central Europe: two populations of undetermined Corbicula sp. from an artificially heated habitat and water of the natural thermal regime (cold water), as well as Corbicula leana, and Corbicula fluminea (from artificially heated and cold water, respectively). To determine the preferred substratum range, we used pairwise choice tests with a range of muddy, sandy, and gravelly substrata, as well as zebra mussel shells. Then, we conducted behavioural observations (locomotion and burrowing) by CCTV cameras on substrata found as preferred and non-preferred by clams. All morphospecies preferred fine-grained over coarser substrata. C. fluminea from cold water had the widest range of substratum preferences. By contrast, C. leana from artificially heated water and Corbicula sp. from cold water showed the narrowest substratum preferences and spent the longest time deeply burrowed. The longest duration of burrowing activity was noted on non-preferred coarser substrata, which may indicate difficulties in burrowing. On non-preferred coarse substrata, C. fluminea (cold water) crawled more than the other morphospecies. Our results show that sediment preferences and behaviour (burrowing and locomotion) differ among various Corbicula morphospecies in Europe, probably also varying their invasive potential. This knowledge is important for determining the overlap of habitat preferences of non-native Corbicula clams with native bivalves, which is critical for determining potential competition strength, impact, and/or displacement. Our results also indicate that habitats with fine sediments could be more threatened by Corbicula invasions.

Infaunal freshwater mussels are highly threatened and declining worldwide. One of the potential threats to mussels consists of biological invasions. We intended to investigate the habitat overlap and behavioural differences between native (Unio pictorum, Unio tumidus, Anodonta anatina, Anodonta cygnea) and invasive (Asian Sinanodonta woodiana) unionid bivalves to determine potential sources of competition. Furthermore, we investigated differences between S. woodiana from the established population in artificially heated waters and from the recent population in a natural thermal regime. We used pairwise choice tests on mud, medium, coarse and very coarse sand, mixture of medium and coarse sand, fine, medium and coarse gravel, and observed mussel locomotion and burrowing in preferred and non-preferred substrata. All species generally preferred fine-grained materials. The widest preference range was exhibited by S. woodiana (both populations), whereas A. cygnea was the most selective. The preferences of the cold-water population of S. woodiana were shifted towards coarser materials compared to conspecifics from the heated waters, and highly overlapped with the preferences of the native species. Anodonta cygnea most often moved horizontally and spent the shortest time deeply burrowed. Both Unio species were deeply burrowed for the largest amount of time and the horizontal locomotion of U. tumidus was the lowest among the test species. Sinanodonta woodiana, especially from the heated water population, exhibited relatively weak locomotion (compared to A. cygnea) and burrowing (compared to Unio spp. and A. anatina). Deep burrowing was more common on fine-grained materials. Our results suggest that the native mussels can be threatened by S. woodiana due to their overlapping habitat preferences, potentially hindering habitat separation. However, mobile native mussels may be capable of migrating and avoiding competition. Accumulating knowledge of the biology and ecology of freshwater mussels could contribute to the creation and improvement of conservation plans to protect these threatened animals.

Abstract Invasive species are those that have been transferred by humans out of their natural range. Native crayfish species in Polish waters include: Astacus astacus and Pontastacus leptodactylus, whereas invasive species are: Pacifastacus leniusculus, Faxonius limosus, Procambarus clarkii and Procambarus virginalis. The objective of this study was to determine how abiotic and biotic environmental factors contribute to interspecific competition of Polish crayfish based on the available literature. Abiotic factors affecting the interspecific competition include tolerance to extreme pH values, calcium ion content, temperature, oxygenation, water salinity, preferred substrate and the type of water bodies. Biotic factors are, inter alia, pathogens, food base, plant cover and interactions in the prey–predator system, as well as interactions between crayfish species. The most important abiotic factors are water temperature and oxygenation, while the most important biotic factor is the crayfish plague – a deadly disease for native species. Each invasive species has a different set of traits and adaptations that enable a successful invasion. However, a successful invasion of a given species is not determined by one, but many adaptations that coexist.