Groundwater is a hidden global keystone ecosystem.

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.

A horizon scan exercise for aquatic invasive alien species in Iberian inland waters.

As the number of introduced species keeps increasing unabatedly, identifying and prioritising current and potential Invasive Alien Species (IAS) has become essential to manage them. Horizon Scanning (HS), defined as an exploration of potential threats, is considered a fundamental component of IAS management. By combining scientific knowledge on taxa with expert opinion, we identified the most relevant aquatic IAS in the Iberian Peninsula, i.e., those with the greatest geographic extent (or probability of introduction), severe ecological, economic and human health impacts, greatest difficulty and acceptability of management. We highlighted the 126 most relevant IAS already present in Iberian inland waters (i.e., Concern list) and 89 with a high probability of being introduced in the near future (i.e., Alert list), of which 24 and 10 IAS, respectively, were considered as a management priority after receiving the highest scores in the expert assessment (i.e., top-ranked IAS). In both lists, aquatic IAS belonging to the four thematic groups (plants, freshwater invertebrates, estuarine invertebrates, and vertebrates) were identified as having been introduced through various pathways from different regions of the world and classified according to their main functional feeding groups. Also, the latest update of the list of IAS of Union concern pursuant to Regulation (EU) No 1143/2014 includes only 12 top-ranked IAS identified for the Iberian Peninsula, while the national lists incorporate the vast majority of them. This fact underlines the great importance of taxa prioritisation exercises at biogeographical scales as a step prior to risk analyses and their inclusion in national lists. This HS provides a robust assessment and a cost-effective strategy for decision-makers and stakeholders to prioritise the use of limited resources for IAS prevention and management. Although applied at a transnational level in a European biodiversity hotspot, this approach is designed for potential application at any geographical or administrative scale, including the continental one.

Using invertebrate functional traits to improve flow variability assessment within European rivers.

Rivers are among the most threatened ecosystems worldwide and are experiencing rapid biodiversity loss. Flow alteration due to climate change, water abstraction and augmentation is a severe stressor on many aquatic communities. Macroinvertebrates are widely used for biomonitoring river ecosystems although current taxonomic approaches used to characterise ecological responses to flow have limitations in terms of generalisation across biogeographical regions. A new macroinvertebrate trait-based index, Flow-T, derived from ecological functional information (flow velocity preferences) currently available for almost 500 invertebrate taxa at the European scale is presented. The index was tested using data from rivers spanning different biogeographic and hydro-climatic regions from the UK, Cyprus and Italy. The performance of Flow-T at different spatial scales and its relationship with an established UK flow assessment tool, the Lotic-invertebrate Index for Flow Evaluation (LIFE), was assessed to determine the transferability of the approach internationally. Flow-T was strongly correlated with the LIFE index using both presence-absence and abundance weighted data from all study areas (r varying from 0.46 to 0.96). When applied at the river reach scale, Flow-T was effective in identifying communities associated with distinct mesohabitats characterised by their hydraulic characteristics (e.g., pools, riffles, glides). Flow-T can be derived using both presence/absence and abundance data and can be easily adapted to varying taxonomic resolutions. The trait-based approach facilitates research using the entire European invertebrate fauna and can potentially be applied in regions where information on taxa-specific flow velocity preferences is not currently available. The inter-regional and continental scale transferability of Flow-T may help water resource managers gauge the effects of changes in flow regime on instream communities at varying spatial scales.

Multiple co-occurrent alien invaders constrain aquatic biodiversity in rivers.

A greater understanding and effective management of biological invasions is a priority for biodiversity conservation globally. Many freshwater ecosystems are experiencing the colonization and spread of multiple co-occurrent alien species. Here the implications of both the relative abundance and richness of alien invaders on aquatic macroinvertebrate taxonomic and functional richness, ecosystem quality, and functional redundancy are assessed using long-term data from rivers in England. Based on the most common aquatic invaders, results indicated that their richness, rather than abundance, was the most important factor negatively affecting aquatic macroinvertebrate biodiversity. However, the response of functional redundancy was negatively affected by invader abundance at the river basin scale. The response of communities varied as the number of invading taxa increased, with the most marked reductions following the colonization of the first few invaders. Results indicate that different facets of multiple biological invasions influence distinct aspects of aquatic biodiversity. Preventing the establishment of new invaders and limiting invader taxa richness within a community should therefore be a conservation priority. These findings will assist river scientists in understanding mechanisms driving changes in biodiversity and facilitate the testing of ecological theories while also ensuring environmental managers and regulators can prioritize conservation / management opportunities.

Exploring longitudinal trends and recovery gradients in macroinvertebrate communities and biomonitoring tools along regulated rivers.

Flow regime alteration by dams has been recognized as a major impact factor for aquatic communities. Spain is currently the member state of the EU with the largest number of large reservoirs. With the broad objective of diminishing the ongoing river degradation trend through the management of environmental flows and the use of biomonitoring tools, we investigated the effects of dams on stream macroinvertebrates in several regulated rivers in Spain with contrasting environmental settings. Specifically, we studied longitudinal trends in macroinvertebrate communities to test: i) if currently used biomonitoring tools and multivariate community analyses can detect hydrological impact responses and biological recovery; ii) if an applicable quantification of the recovery gradient, in terms of distance downstream from dams, can be obtained for Iberian fluvial systems; iii) if macroinvertebrate community structure respond different to flow regulation, depending on the contrasting environmental river typologies; and iv) if the type and intensity of hydrological alteration modulates the observed community responses/recovery. Biotic indices and metrics displayed a decrease in 5 out of 6 systems immediately downstream of infrastructure. Complete recovery could not be clearly detected, but some recovery patterns started at a distance >11km. Multivariate community patterns and biomonitoring metrics showed the most pronounced hydrological alteration impacts and weaker recovery of the downstream macroinvertebrate communities within dammed Mediterranean streams (comparing to other rivers with continental or oceanic climate influence). Finally, both the intensity and type of hydrological alteration (highlighting the alteration of the floods and droughts components) were related to changes in common biomonitoring metrics. Our results could help in recognizing heavily modified water bodies (sensu European Water Framework Directive) downstream of dams or the delineation of fluvial zones or reserves. Furthermore, applied research areas dealing with environmental flows or the bioassessment of hydrological impacts could benefit from our main findings.

Simulating rewetting events in intermittent rivers and ephemeral streams: A global analysis of leached nutrients and organic matter.

Climate change and human pressures are changing the global distribution and the extent of intermittent rivers and ephemeral streams (IRES), which comprise half of the global river network area. IRES are characterized by periods of flow cessation, during which channel substrates accumulate and undergo physico-chemical changes (preconditioning), and periods of flow resumption, when these substrates are rewetted and release pulses of dissolved nutrients and organic matter (OM). However, there are no estimates of the amounts and quality of leached substances, nor is there information on the underlying environmental constraints operating at the global scale. We experimentally simulated, under standard laboratory conditions, rewetting of leaves, riverbed sediments, and epilithic biofilms collected during the dry phase across 205 IRES from five major climate zones. We determined the amounts and qualitative characteristics of the leached nutrients and OM, and estimated their areal fluxes from riverbeds. In addition, we evaluated the variance in leachate characteristics in relation to selected environmental variables and substrate characteristics. We found that sediments, due to their large quantities within riverbeds, contribute most to the overall flux of dissolved substances during rewetting events (56%-98%), and that flux rates distinctly differ among climate zones. Dissolved organic carbon, phenolics, and nitrate contributed most to the areal fluxes. The largest amounts of leached substances were found in the continental climate zone, coinciding with the lowest potential bioavailability of the leached OM. The opposite pattern was found in the arid zone. Environmental variables expected to be modified under climate change (i.e. potential evapotranspiration, aridity, dry period duration, land use) were correlated with the amount of leached substances, with the strongest relationship found for sediments. These results show that the role of IRES should be accounted for in global biogeochemical cycles, especially because prevalence of IRES will increase due to increasing severity of drying events.

Taxonomic changes and non-native species: An overview of constraints and new challenges for macroinvertebrate-based indices calculation in river ecosystems.

Freshwater ecosystems face many threats in the form of reduced water quantity, poor water quality and the loss of biodiversity. As a result, aquatic biomonitoring tools are required to enable the evaluation of these critical changes. Currently, macroinvertebrate-based indices are globally the most widely used biomonitoring tools in fluvial ecosystems. However, very little is known about the potential effects of changes in taxonomic understanding (updating of classification and nomenclature) or the presence of new non-native species for biotic indices calculation. This is especially relevant given that errors, incorrect classification or exclusion of new/updated nomenclature may affect ecological status evaluations and have direct consequences for the management and conservation of freshwater systems. In this discussion paper the main constraints, challenges and implications of these issues are outlined and case studies from a range of European countries are discussed. However, similar challenges affect rivers and managers globally and will potentially be amplified further in the future. Bioassessment science needs to be open to improvements, and current tools and protocols need to be flexible so that they can be updated and revised rapidly to allow new scientific developments to be integrated. This discussion highlights specific examples and new ideas that may contribute to the future development of aquatic biomonitoring using macroinvertebrates and other faunal and floral groups in riverine ecosystems.

Freshwater ecosystems and aquatic insects: a paradox in biological invasions.

Biological invasions have increased significantly in response to global change and constitute one of the major causes of biodiversity loss. Insects make up a large fraction of invasive species, in general, and freshwaters are among the most invaded ecosystems on our planet. However, even though aquatic insects dominate most inland waters, have unparalleled taxonomic diversity and occupy nearly all trophic niches, there are almost no invasive insects in freshwaters. We present some hypotheses regarding why aquatic insects are not common among aquatic invasive organisms, suggesting that it may be the result of a suite of biological, ecological and anthropogenic factors. Such specific knowledge introduces a paradox in the current scientific discussion on invasive species; therefore, a more in-depth understanding could be an invaluable aid to disentangling how and why biological invasions occur.

How far could the alien boatman Trichocorixa verticalis verticalis spread? Worldwide estimation of its current and future potential distribution.

Invasions of alien species are considered among the least reversible human impacts, with diversified effects on aquatic ecosystems. Since prevention is the most cost-effective way to avoid biodiversity loss and ecosystem problems, one challenge in ecological research is to understand the limits of the fundamental niche of the species in order to estimate how far invasive species could spread. Trichocorixa verticalis verticalis (Tvv) is a corixid (Hemiptera) originally distributed in North America, but cited as an alien species in three continents. Its impact on native communities is under study, but it is already the dominant species in several saline wetlands and represents a rare example of an aquatic alien insect. This study aims: i) to estimate areas with suitable environmental conditions for Tvv at a global scale, thus identifying potential new zones of invasion; and ii) to test possible changes in this global potential distribution under a climate change scenario. Potential distributions were estimated by applying a multidimensional envelope procedure based on both climatic data, obtained from observed occurrences, and thermal physiological data. Our results suggest Tvv may expand well beyond its current range and find inhabitable conditions in temperate areas along a wide range of latitudes, with an emphasis on coastal areas of Europe, Northern Africa, Argentina, Uruguay, Australia, New Zealand, Myanmar, India, the western boundary between USA and Canada, and areas of the Arabian Peninsula. When considering a future climatic scenario, the suitability area of Tvv showed only limited changes compared with the current potential distribution. These results allow detection of potential contact zones among currently colonized areas and potential areas of invasion. We also identified zones with a high level of suitability that overlap with areas recognized as global hotspots of biodiversity. Finally, we present hypotheses about possible means of spread, focusing on different geographical scales.