Freshwater fish biodiversity restoration in floodplain rivers requires connectivity and habitat heterogeneity at multiple spatial scales

With a sixth mass extinction looming and freshwater biodiversity declining at unprecedented rates, evaluating ecological efficacy of river restoration efforts is critical in combatting global biodiversity loss. Here, we present a comprehensive study of the functioning for fishes of 46 river restoration projects in the river Rhine, one of the world's most heavily engineered lowland rivers. Floodplains with permanent, either one- or two-sided lateral connectivity to the main channel, favour total fish abundance, and are essential as nursery areas for riverine fishes. Habitat heterogeneity had a strong positive effect on species richness but was negatively related with fish abundances. However, the effects of environmental variables varied between ecological groups and spatial scales. Surprisingly, richness of critical rheophilic fishes declined with large-scale habitat heterogeneity (~1000 m), while it increased at small scales (~100 m), possibly because of the presence of unfavourable habitats for this ecological group at larger scales. Clearly, there is no one-size-fits-all design for river restoration projects. Whether a river section is free-flowing or impounded dictates the scope and efficacy of restoration projects and, within a river section, multiple complementary restoration projects might be key to mitigate freshwater fish biodiversity loss. An essential element for success is that these projects should retain permanent lateral connection to the main channel