Earthquakes are among the most destructive natural events. The 6 April 2009, 6.3-Mw 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.

SUMMARY1. 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 Wü rmian 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 Wü rmian 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.

SUMMARY1. 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 aquiferselection 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.

Subterranean ecosystems are among the most widespread environments on Earth, yet we still have poor knowledge of their biodiversity. To raise awareness of subterranean ecosystems, the essential services they provide, and their unique conservation challenges, 2021 and 2022 were designated International Years of Caves and Karst. As these ecosystems have traditionally been overlooked in global conservation agendas and multilateral agreements, a quantitative assessment of solution-based approaches to safeguard subterranean biota and associated habitats is timely. This assessment allows researchers and practitioners to understand the progress made and research needs in subterranean ecology and management. We conducted a systematic review of peer-reviewed and grey literature focused on subterranean ecosystems globally (terrestrial, freshwater, and saltwater systems), to quantify the available evidence-base for the effectiveness of conservation interventions. We selected 708 publications from the years 1964 to 2021 that discussed, recommended, or implemented 1,954 conservation interventions in subterranean ecosystems. We noted a steep increase in the number of studies from the 2000s while, surprisingly, the proportion of studies quantifying the impact of conservation interventions has steadily and significantly decreased in recent years. The effectiveness of 31% of conservation interventions has been tested statistically. We further highlight that 64% of the reported research occurred in the Palearctic and Nearctic biogeographic regions. Assessments of the effectiveness of conservation interventions were heavily biased towards indirect measures (monitoring and risk assessment), a limited sample of organisms (mostly arthropods and bats), and more accessible systems (terrestrial caves). Our results indicate that most conservation science in the field of subterranean biology does not apply a rigorous quantitative approach, resulting in sparse evidence for the effectiveness of interventions. This raises the important question of how to make conservation efforts more feasible to implement, cost-effective, and long-lasting. Although there is no single remedy, we propose a suite of potential solutions to focus our efforts better towards increasing statistical testing and stress the importance of standardising study reporting to facilitate metaanalytical exercises. We also provide a database summarising the available literature, which will help to build quantitative knowledge about interventions likely to yield the greatest impacts depending upon the subterranean species and habitats of interest. We view this as a starting point to shift away from the widespread tendency of recommending conservation interventions based on anecdotal and expert-based information rather than scientific evidence, without quantitatively testing their effectiveness.

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