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.

30 years of large river restoration: How long do restored floodplain channels remain suitable for targeted rheophilic fishes in the lower river Rhine?

The ecological efficacy of river restoration projects may change over time, resulting in the loss of their ecological function for targeted species. The goal of this study was to evaluate the rheophilic nursery function of restored floodplain channels over time, by analysing 30 years of monitoring data from 12 restoration projects in the lower river Rhine. We hypothesised that the nursery function would change over time, caused by the combined effects of decreasing flow conditions and succession processes affecting habitat heterogeneity. We found that nursery area suitability for rheophilic fish was almost 4 times higher in two-sided connected channels than in one-sided connected channels, although the response trends of rheophilic fish were similar for both water body types. These response curves showed clear optima with channel age, for rheophilic fish abundance at 13 to 14 years post-restoration, indicating optimal nursery conditions. On the other hand, rheophilic species richness showed a steadily decreasing trend with channel age, suggesting aging channels became less suitable as nursery areas for most rheophilic fish species. The presence of permanent flow was found to be the main driving factor in explaining both rheophilic fish community trends and habitat succession in individual restored channels. We did not observe an effect of habitat heterogeneity on nursery function for rheophilic fish. To create and maintain optimal nursery conditions in restored floodplain channels of strongly anthropogenically influenced rivers such as the river Rhine, we propose a management strategy involving cyclic rejuvenation through human intervention, focusing on restoring permanent flow, with a frequency of on average every 15 years, depending on the rate of aggradation and targeted rheophilic species. We also propose a thorough investigation of the relationship between habitat heterogeneity and nursery success in floodplain channels, as a next step in the identification of suitable nursery areas for rheophilic fishes.