Groundwater is a vital ecosystem of the global water cycle, hosting unique biodiversity and providing essential services to societies. Despite being the largest unfrozen freshwater resource, in a period of depletion by extraction and pollution, groundwater environments have been repeatedly overlooked in global biodiversity conservation agendas. Disregarding the importance of groundwater as an ecosystem ignores its critical role in preserving surface biomes. To foster timely global conservation of groundwater, we propose elevating the concept of keystone species into the realm of ecosystems, claiming groundwater as a keystone ecosystem that influences the integrity of many dependent ecosystems. Our global analysis shows that over half of land surface areas (52.6%) has a medium‐to‐high interaction with groundwater, reaching up to 74.9% when deserts and high mountains are excluded. We postulate that the intrinsic transboundary features of groundwater are critical for shifting perspectives towards more holistic approaches in aquatic ecology and beyond. Furthermore, we propose eight key themes to develop a science‐policy integrated groundwater conservation agenda. Given ecosystems above and below the ground intersect at many levels, considering groundwater as an essential component of planetary health is pivotal to reduce biodiversity loss and buffer against climate change.

Summary: 1. The distribution patterns of stygobiotic invertebrates were examined with a stratified sampling design at 197 sites selected among four hydrogeographic basins in the Lessinian Massif (northern Italy). The sites were approximately evenly distributed among four hydrogeological zones: unsaturated and saturated zone of karstic aquifers, and hyporheic and saturated zone of porous aquifers.2. Outlying Mean Index (OMI) analysis which assesses deviation of habitat conditions from reference conditions, was used to evaluate the importance of 14 selected environmental variables in shaping groundwater biodiversity patterns in the region (total of 89 stygobiotic species). The measured variables explained 80% of the variability in the data set.3. Sampling sites were distributed along the environmental gradients defined by OMI analysis. Significant differences were detected between karstic and porous site, as well as among sites located in the four hydrogeological zones. Differences among the four hydrogeographic basins were not observed.4. Ordination of stygobiotic species along the environmental gradients was best explained by historical variables (mainly Würmian glaciation and age of the underlying geological formation), while variables related to hydrogeology (mainly pH, calcium concentration and habitat fragmentation) influenced species distributions in the hydrogeological zones. An Environmental Integrity Index and nitrate concentration were significantly correlated with altitude, but appeared not to play a significant role in determining stygobiotic biodiversity patterns at the regional scale.5. Results of the OMI analysis were highly significant for all taxa, suggesting that stygobiotic species are sensitive to the environmental factors studied. Thirty‐five species showed high habitat specialisation (OMI index > 10). These species were usually rare and endemic to the Lessinian Massif. Most of them were found in a single hydrogeological zone.6. Quaternary glaciations appear not to have lowered stygobiotic species richness in the Lessinian Massif. This may be because of the marginal location of the region with respect to the Würmian glacier limit and because of extensive networks of fractures in the vadose zone of the karst, which may have allowed stygobionts to move deep down in the aquifers to seek refuge during surface freezing and to recolonise ancestral habitats after the glaciers retreated.

Groundwater comprises the largest freshwater ecosystem on the planet. It has a distinct regime of extreme, yet stable environmental conditions that have favoured the development of similar morphological and functional traits in the resident invertebrate fauna (stygofauna). The analysis of community traits is increasingly used as an alternative to taxonomy‐based assessments of biodiversity, especially for monitoring ecosystem status and linking the functions of organisms to ecological processes, yet it has been rarely applied to stygofauna and groundwater ecosystems. In this paper, we review the variation in functional traits among the invertebrate fauna of this important ecosystem. We focus on the stygofauna and processes of alluvium and fractured rock aquifers that are typified by small voids and fissures that constrain the habitats and environmental conditions. As a first step, we compare trait variability between groundwater and surface water invertebrate communities and then examine the significance of the ranges of these traits to the vulnerability of the ecosystem to change. Fifteen potentially useful functional traits are recognised. Eight of these have narrower ranges (i.e. exhibit fewer states, or attributes, of a particular trait) in groundwater than they do in surface water. Two traits have wider ranges. Our synthesis suggests that the relative stability of groundwater environments has led to low trait variability. The low biomass and low reproductive rate of stygofauna suggest that recovery potential following disturbance is likely to be low. For the purposes of both improved understanding and effective management, further work is needed to document additional functional traits and their states in groundwater fauna, enabling a better understanding of the relationship between response and effect traits in these ecosystems. Read the free Plain Language Summary for this article on the Journal blog.

Earthquakes are among the most destructive natural events. The 6 April 2009, 6.3-M_w earthquake in L'Aquila (Italy) markedly altered the karstic Gran Sasso Aquifer (GSA) hydrogeology and geochemistry. The GSA groundwater invertebrate community is mainly comprised of small-bodied, colourless, blind microcrustaceans. We compared abiotic and biotic data from two pre-earthquake and one post-earthquake complete but non-contiguous hydrological years to investigate the effects of the 2009 earthquake on the dominant copepod component of the obligate groundwater fauna. Our results suggest that the massive earthquake-induced aquifer strain biotriggered a flushing of groundwater fauna, with a dramatic decrease in subterranean species abundance. Population turnover rates appeared to have crashed, no longer replenishing the long-standing communities from aquifer fractures, and the aquifer became almost totally deprived of animal life. Groundwater communities are notorious for their low resilience. Therefore, any major disturbance that negatively impacts survival or reproduction may lead to local extinction of species, most of them being the only survivors of phylogenetic lineages extinct at the Earth surface. Given the ecological key role played by the subterranean fauna as decomposers of organic matter and “ecosystem engineers”, we urge more detailed, long-term studies on the effect of major disturbances to groundwater ecosystems.

Summary: 1. A possible conservation strategy to minimise the risk of groundwater biodiversity loss due to human activities consists in designing a network of reserve areas at the continental scale that collectively include most groundwater species. To this end, we compared the efficiency of three area selection methods (species richness hotspots, endemism hotspots and complementarity) and examined the influence of spatial constraints (reduced extent and increased aggregation of reserve areas) on the representation of 1059 groundwater species in six European regions.2. Presence data from a data base elaborated as part of a European initiative on groundwater biodiversity, the PASCALIS project, were referenced onto 4675 grid cells (0.2 by 0.2°). Complementary performed much better than traditional selection methods for maximising species representation in a reserve network arbitrarily limited to 10% of all the cells containing groundwater fauna. It captured 155 more species than areas selected on richness and 77 more species than areas selected on endemism hotspots.3. Representing species in a specified proportion of their area of occupancy (i.e. 100%, 50% and 10% of the area of occupancy of species occurring in 1, ≤10, and >10 cells, respectively) required inclusion of 46% of the cells containing groundwater fauna. The reserve network needed to achieve this level of coverage may be too large and fragmented to be implemented and managed in practice.4. Reduction of the reserve areas to 10% of the landscape containing groundwater fauna and their aggregation into a smaller number of cell clusters resulted in a more realistic reserve network that represented 73.8% (782 species) and 59.1% (274 endemics) of the total number of species and endemics, respectively.5. We propose several research priorities to improve the design of effective groundwater reserve networks in Europe: (i) devising sampling strategies that reduce uncertainties in the placement of reserves and increase the number of alternative reserve networks and (ii) shifting from a grid‐cell selection approach to an aquifer‐selection approach that incorporates species representation targets, minimum space requirement and also socio‐economic costs related to the vulnerability of aquifers and degree of human activity in the catchment.