Living on the edge – An overview of invertebrates from groundwater habitats prone to extreme environmental conditions

Groundwater ecosystems from cold polar and circumpolar regions, hot springs, as well as those developed in salt, gypsum or in volcanic rocks are one of the environments considered to exhibit extreme environmental conditions such as low (below 0°C) or high (over 45°C) temperatures, hypersaline waters, or with elevated content of toxic gases like hydrogen sulfide or methane. They represent the “unseen ecosystem beneath our feet” and are inhabited by a large diversity of organisms, persisting and flourishing under severe environmental conditions that are usually hostile to the majority of organisms. These types of groundwater ecosystems are remarkable “evolutionary hotspots” that witnessed the adaptive radiation of morphologically and ecologically diverse species, whereas the organisms living here are good models to understand the evolutionary processes and historical factors involved in speciation and adaptation to severe environmental conditions. Here, we provide an overview of the groundwater invertebrates living in continental groundwater habitats prone to extreme environmental conditions in one or more physico-chemical parameters. Invertebrates are represented by a wide variety of taxonomic groups, however dominated by crustaceans that show specific adaptations mostly metabolic, physiologic, and behavioral. Symbiotic associations among bacteria and invertebrates are also discussed enlightening this biological interaction as a potential adaptation of different groundwater invertebrates to cope with severe environmental conditions. Given the high pressures that anthropogenic activities pose on groundwater habitats worldwide, we predict that several of these highly specialized organisms will be prone to extinction in the near future. Finally, we highlight the knowledge gaps and future research approaches in these particular groundwater ecosystems by using integrative-omic studies besides the molecular approach to shed light on genetic variation and phenotypic plasticity at species and populational levels.GRAPHICAL ABSTRAC

Thermal acclimation and metabolic scaling of a groundwater asellid in the climate change scenario

Metabolic rate has long been used in animal adaptation and performance studies, and individual oxygen consumption is used as proxy of metabolic rate. Stygofauna are organisms adapted to groundwater with presumably lower metabolic rates than their surface relatives. How stygofauna will cope with global temperature increase remains unpredictable. We studied the thermal acclimation and metabolic scaling with body mass of a stygobitic crustacean, Proasellus lusitanicus, in the climate change scenario. We measured oxygen consumption rates in a thermal ramp-up experiment over four assay temperatures and tested two hypotheses: (i) P. lusitanicus exhibits narrow thermal plasticity, inadequate for coping with a fast-increasing thermal regime; and (ii) oxygen consumption rates scale with the body mass by a factor close to 0.75, as commonly observed in other animals. Our results show that P. lusitanicus has low thermal plasticity in a fast-increasing thermal regime. Our data also suggest that oxygen consumption rates of this species do not follow mass-dependent scaling, potentially representing a new trait of metabolic optimization in groundwater habitats, which are often limited in food and oxygen. Species with limited dispersal capacities and rigid metabolic guilds face extinction risk due to climate change and omitting groundwater ecosystems from climate change agendas emphasizes the unprotected status of stygofauna.info:eu-repo/semantics/publishedVersio

Sensitivity of hypogean and epigean freshwater copepods (Crustacea Copepoda) to agricultural pollutants: single toxicants and mixtures

Widespread pollution from agriculture is one of the major causes of poor freshwater quality currently observed across Europeand worldwide. Nutrient loads (nitrogen and phosphorous) from fertilizers and pesticides are known to adversely impact freshwater ecosystems, both surface- and ground water. The Crustacea Copepoda are by far the most abundant and species-rich taxon in ground water and they are consistently represented in ecotonal environments also, as in the hyporheic zone. The direct impact of agricultural pollutants on freshwater biota has been addressed in several studies by means of laboratory bioassays. However, the ecotoxicological research concerning freshwater copepods is scarce for epigean species and almost non-existent for the hypogean ones. Moreover, when available, ecotoxicological studies have been performed considering the effect of toxicants taken individually. Actually, this approach does not reflect the conditions in the field high concentrations of both N-fertilizers and pesticides co-occur. In this study we assessed the acute (at 96h) sensitivity of adults of a hypogean and an epigean species, both belonging to the Crustacea, Copepoda, Cyclopoida, Cyclopidae, to two agricultural toxicants: the ammonium nitrate and the herbicides Imazamox. Both chemicals are widely used for cereal agriculture inEurope. We tested the sensitivity considering the LC50 with mortality endpoints for individual and combined (a mixture solution of ammonium nitrate and Imazamox) toxicant concentrations. The hypogean species was more sensitive than the epigean one to both chemicals and their mixture. Ionized ammonia proved to be more toxic than the herbicide Imazamox for both species. However, the LC50 of both chemicals were lower than the actual standard law limits for good freshwater quality status defined by the Water Framework Directive (2000/60/CE). The effect of the mixture, of the two toxicants was fairly synergic. Concerning ionized ammonia, the LC50-96h in the mixture was higher than the law limits for both species. According to these results, the actual law limits for the good quality of freshwater bodies should be revised accordingly by authorities in charge of water management

Exploring copepod distribution patterns at three nested spatial scales in a spring system. Habitat partitioning and potential for hydrological bioindication

In groundwater-fed springs, habitat characteristics are primarily determined by a complex combination of geomorphic features and physico-chemical parameters, while species assemblages are even more intricate. Springs host species either inhabiting the spring mouth, or colonizing spring habitats from the surface or from the aquifers which feed the springs. Groundwater species living in springs have been claimed as good candidates for identifying dual aquifer flowpaths or changes in groundwater pathways before reaching the spring outlets. However, the reliability of spring species as hydrological biotracers has not been widely investigated so far. Our study was aimed at analysing a large karstic spring system at three nested spatial scales in order: i) to assess, at whole spring system scale, the presence of a groundwater divide separating two aquifers feeding two spring units within a single spring system, by combining isotope analyses, physico-chemistry, and copepod distribution patterns; ii) to test, at vertical spring system scale, the effectiveness of copepods in discriminating surface and subsurface habitat patches within the complex mosaic spring environment; iii) to explore, at local spring unit level, the relative role of hydrochemistry and sediment texture as describers of copepod distribution among microhabitats. The results obtained demonstrated the presence of a hierarchical spatial structure, interestingly reflected in significant differences in assemblage compositions. Copepod assemblages differed between the two contiguous spring units, which were clearly characterized by their hydrochemistry and by significant differences in the groundwater flowpaths and recharge areas, as derived by the isotope analyses. The biological results suggested that stygobiotic species seem to be related to the origin of groundwater, suggesting their potential role as hydrological biotracers. At vertical scale, assemblage composition in surface and subsurface habitats was significantly different, both between spring units and among microhabitats, supporting strong habitat preferences of copepod species. At the smaller local scale, the response to habitat patchiness of subsurface copepod assemblages resulted in distribution patterns primarily defined by sediment texture, while the sensitivity to differences in hydrochemistry was negligible

Anthropometric, physiological and performance aspects that differentiate male athletes from females and practical consequences

The article provides a concise analysis of the meaningful and comprehensive aspects of the incidence of gender differences in sport performance, in particular analyzes some aspects (physiological, quantitative and qualitative) that determine the performance of the sport and influence the organization of training in elite sports. The results of different scientific papers will allow to better understand and motivate more accurately the methodological choices and the procedural attentions, in the respect physical, physiological, technical, psychic and gender characteristics in the various sports activities. The objective of the study is theoretical argument for the part relating to the revision of the scientific literature of the theory of training and interpretative for the theoretical and documentary results

Earthquakes trigger the loss of groundwater biodiversity

Earthquakes are among the most destructive natural events. The 6 April 2009, 6.3-Mw earthquake in L\u27Aquila (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

Some like it hot: Thermal preference of the groundwater amphipod Niphargus longicaudatus (Costa, 1851) and climate change implications

Groundwater is a crucial resource for humans and the environment, but its global human demand currently exceeds available volumes by 3.5 times. Climate change is expected to exacerbate this situation by increasing the frequency of droughts along with human impacts on groundwater ecosystems. Despite prior research on the quantitative effects of climate change on groundwater, the direct impacts on groundwater biodiversity, especially obligate groundwater species, remain largely unexplored. Therefore, investigating the potential impacts of climate change, including groundwater temperature changes, is crucial for the survival of obligate groundwater species. This study aimed to determine the thermal niche breadth of the crustacean amphipod species Niphargus longicaudatus by using the chronic method. We found that N. longicaudatus has a wide thermal niche with a natural performance range of 7–9 °C, which corresponds to the thermal regime this species experiences within its distribution range in Italy. The observed range of preferred temperature (PT) was different from the mean annual temperature of the sites from which the species has been collected, challenging the idea that groundwater species are only adapted to narrow temperature ranges. Considering the significant threats of climate change to groundwater ecosystems, these findings provide crucial information for the conservation of obligate groundwater species, suggesting that some of them may be more resilient to temperature changes than previously thought. Understanding the fundamental thermal niche of these species can inform conservation efforts and management strategies to protect groundwater ecosystems and their communities.info:eu-repo/semantics/publishedVersio

Temperature effect on the sensitivity of the copepod Eucyclops serrulatus (Crustacea, Copepoda, Cyclopoida) to agricultural pollutants in the hyporheic zone

Abstract The sensitivity of freshwater invertebrates to agricultural pollutants is supposed to increase with rising temperature due to global warming. The aim of this study was to measure the effect of temperature on the lethal toxicity of ammonia-N, the herbicide Imazamox and the mixture of the two chemicals, in the adults and the juveniles of a population of the copepod Eucyclops serrulatus. This is a widely distributed species found in surface waters, in transitional habitats between surface water and groundwater, and in genuine groundwater environments. We tested the sensitivity by short-term bioassays (96 h) at 15°C and 18°C, respectively. Our results highlighted the following: (1) increasing temperature affected the sensitivity of the adults to ammonia-N and of the juveniles to the mixture, all of which were more sensitive to its detrimental effects at 18°C; (2) the juvenile stages were more sensitive than the adults to all toxicants, and (3) for all combinations of chemicals and temperatures, the effects were synergistic and approximately one order of magnitude greater than those expected according to a concentration addition model when comparing the LC50 for each chemical in the mixture with the LC50s of chemicals individually assayed. Overall, in a context of global change, ammonia-N and mixtures of agricultural pollutants may affect the survival rate of species that spend a part or the whole life-cycle in the hyporheic habitat, with detrimental effects to biodiversity and ecosystem services provided by the hyporheic biota