The ecological role of permanent ponds in Europe: a review of dietary linkages to terrestrial ecosystems via emerging insects

Permanent ponds are valuable freshwater systems and biodiversity hotspots. They provide diverse ecosystem services (ESs), including water quality improvement and supply, food provisioning, and biodiversity support, despite significant pressure from multiple anthropogenic stressors and the impacts of ongoing global change. However, ponds are largely overlooked in management plans and legislation, and ecological research has focused on large freshwater ecosystems, such as rivers or lakes. Protection of ponds is often insufficient or indirectly provided via associated habitats such as wetlands. This situation is likely exacerbated by the lack of a full-scale understanding of the importance of ponds. In this review, we provide a detailed overview of permanent ponds across Europe, including their usages and the biodiversity they support. By discussing the concepts of pondscape and metacommunity theory, we highlight the importance of connectivity among and between ponds and identified fluxes of emerging insects as another ES of ponds. Those insects are rich in essential nutrients such as polyunsaturated fatty acids (PUFAs), delivered through them to the terrestrial environment; however, the extent and impact of this ES remains largely unexplored. Several potential stressors, especially related to ongoing global change, that influence pond diversity and integrity are discussed. We provide our insights on future pond management. Adaptive measures, taking into account the pond system per se within the pondscape, are the most promising to mitigate the loss of natural ponds and restore and conserve natural small waterbodies as refuges and diversity hotspots in increasingly urbanized landscapes.