Terrestrial reserve networks do not adequately represent aquatic ecosystems.

Protected areas are a cornerstone of conservation and have been designed largely around terrestrial features. Freshwater species and ecosystems are highly imperiled, but the effectiveness of existing protected areas in representing freshwater features is poorly known. Using the inland waters of Michigan as a test case, we quantified the coverage of four key freshwater features (wetlands, riparian zones, groundwater recharge, rare species) within conservation lands and compared these with representation of terrestrial features. Wetlands were included within protected areas more often than expected by chance, but riparian zones were underrepresented across all (GAP 1-3) protected lands, particularly for headwater streams and large rivers. Nevertheless, within strictly protected lands (GAP 1-2), riparian zones were highly represented because of the contribution of the national Wild and Scenic Rivers Program. Representation of areas of groundwater recharge was generally proportional to area of the reserve network within watersheds, although a recharge hotspot associated with some of Michigan's most valued rivers is almost entirely unprotected. Species representation in protected areas differed significantly among obligate aquatic, wetland, and terrestrial species, with representation generally highest for terrestrial species and lowest for aquatic species. Our results illustrate the need to further evaluate and address the representation of freshwater features within protected areas and the value of broadening gap analysis and other protected-areas assessments to include key ecosystem processes that are requisite to long-term conservation of species and ecosystems. We conclude that terrestrially oriented protected-area networks provide a weak safety net for aquatic features, which means complementary planning and management for both freshwater and terrestrial conservation targets is needed.