Lake-stream transition zones support hotspots of freshwater ecosystem services: Evidence from a 35-year study on unionid mussels

Securing adequate supply of high-quality water is of increasing global importance and relies in large part on ecosystem services provided by freshwater biota. Unionid mussels are important keystone species and habitat engineers that shape freshwater ecosystems through water filtration, nutrient cycling and provision of habitats; their rapid global declines result in dramatic losses of ecosystem functions. Maintenance and enhancement of the services they provide depend on the identification of their crucial habitats. Following theoretical assumptions, this study analyses the importance of lake-stream transition zones for unionid mussels, based on data collected in 1984 and 2019 from an undisturbed stream flowing through five consecutive lakes. Mussel distribution matched the distribution of host fish and was strongly influenced by lakes: densities were highest near lake outlets, reaching 290 ind. m^-2 (14.7 kg m^-2) in 2019, and declined with downstream distance following a negative power function. This pattern was spatially consistent and sustained over time. All six unionid species native to north-central Europe were present, but common species (Anodonta anatina, Unio pictorum, U. tumidus) contributed about 80% of individuals and were responsible for most of the ecosystem services provided by unionid mussels. Estimated 1.9 × 10⁶ mussel individuals inhabiting 3.2 km of stream length filtered a water volume equivalent to the total stream discharge approximately 2.5 times daily. Aggregations of spent shells, up to 17 kg m^-2, accumulated downstream of lakes, forming extensive shell and mussel beds, providing habitats and contributing shell hash that improved stream-bed conditions. Globally invasive Dreissena polymorpha was present at low densities and did not spread or increase in abundance, indicating a long-term biotic resistance of the natural native community. Our study underscores the importance of undisturbed lake outlets, longitudinal connectivity of riverine ecosystems, and of common mussel species in maintaining freshwater ecosystem functionality and provision of vital services.