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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.
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Ecosistema , Agua Subterránea , Biodiversidad , Agua Dulce , Contaminación AmbientalRESUMEN
Bat flies (Diptera: Nycteribiinae) are highly specialized bloodsucking bat ectoparasites. Some of the ectoparasitic bat flies are themselves parasitized with an ectoparasitic fungus of the genus Arthrorhynchus (Laboulbeniales). Ascospores of the fungus attach to the cuticle of a bat fly and develop a haustorium that penetrates the host cuticle. This interaction defines the fungus as a hyperparasite. Both the fly and the fungus are obligate parasites and this peculiar case of hyperparasitism has remained largely unstudied. We studied the prevalence of Laboulbeniales, genus Arthrorhynchus, in natural populations of bat flies infesting the bat species Miniopterus schreibersii, Myotis bechsteinii, My. blythii, My. daubentonii, My. escalerai and My. myotis in Portuguese caves. Laboulbeniales were found infecting 10 of the 428 screened bat flies (2.3%) in natural populations, with fewer infections in winter. Images obtained with transmission electron microscopy show the fungal haustorium within the bat fly host tissue, from where it extracts nutrition.
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Quirópteros/parasitología , Dípteros/parasitología , Micosis/veterinaria , Animales , Ascomicetos , Cuevas , Interacciones Huésped-ParásitosRESUMEN
The genera Diplopodomyces and Troglomyces (Laboulbeniales, Ascomycota) parasitizing millipedes (Diplopoda) are reviewed. We describe four new species: D. lusitanipodos parasitic on Lusitanipus from Portugal and on Cyphocallipus from Spain, D. veneris on Lusitanipus from Portugal, T. botryandrus on Leptoiulus from Italy and T. rossii on Ophyiulus, Leptoiulus and Typhloiulus from Italy. Diplopodomyces veneris is a dimorphic species, with different morphotypes growing on the male copulatory organs and on the surroundings of the female gonopore respectively. The type species of both genera also have been studied, their descriptions are revised and a neotype for T. manfrediae is designated. New records from Italy are given for D. callipodos and T. manfrediae.
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Artrópodos/microbiología , Ascomicetos/clasificación , Animales , Ascomicetos/aislamiento & purificación , Ascomicetos/ultraestructura , Femenino , Italia , Masculino , Microscopía Electrónica de RastreoRESUMEN
A new genus Sireuma is described to accommodate a new species, S. nobile, from a cave in Portugal. The unique features of the male gonopods and the absence of a postvulvar organ support the creation of a new genus. The new species is illustrated with scanning electron micrographs and diagrammatic drawings. Differences vis-à-vis its relatives are discussed, and information on the environment in which the new species was found is also provided.
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Artrópodos/anatomía & histología , Artrópodos/clasificación , Animales , Artrópodos/fisiología , Artrópodos/ultraestructura , Demografía , Femenino , Masculino , Portugal , España , Especificidad de la EspecieRESUMEN
The new hypogean ground beetle, Duvalius abyssimus n. sp., from Krubera-Voronja, world's deepest cave (Arabika massif, Western Caucasus) is described and illustrated. Diagnostic morphological characters of the imago, male and female genitalia are provided. Its relations with other Duvalius Delarouzée species from the Western Caucasus geographic area are discussed. An updated overview of the biocoenosis of its peculiar habitat is made.
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Escarabajos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Animales , Cuevas , Escarabajos/anatomía & histología , Ecosistema , Femenino , MasculinoRESUMEN
Two new species of the genus Cylindroiulus Verhoeff, 1894, C. julesvernei and C. oromii, are described from the subterranean ecosystem of Madeira Island, Portugal. Species are illustrated with photographs and diagrammatic drawings. The new species belong to the Cylindroiulus madeirae-group, an insular species swarm distributed in the archipelagos of Madeira and the Canary Islands. We discuss the differences between the new species and their relatives and present information on the subterranean environment of Madeira. An updated overview of the subterranean biodiversity of millipedes in Macaronesia is also provided.
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Artrópodos/clasificación , Animales , Demografía , Ecosistema , Femenino , Masculino , Portugal , Especificidad de la EspecieRESUMEN
Background: Domitiuslusitanicus (Fage, 1931) is a troglobiont spider, endemic from caves in the largest karst massif in Portugal, the Estremenho. It was the first described cave-adapted species from Portugal, but the male of the species was only described in 1988. New information: Over the last two decades, the knowledge on the distribution of D.lusitanicus increased significantly. We assess the conservation status of D.lusitanicus, providing new information on its extent of occurrence and the anthropogenic threats and present a IUCN Red List profile. D.lusitanicus faces various anthropogenic threats, such as habitat loss, agriculture, pollution and tourism impacts. Despite a large part of its distribution is included in a Natural Park, it expands outside of the areas deemed for protection in the Natura 2000 network. This species has the widest spread distribution of all troglobionts in the Estremenho Massif; therefore, it may be used as an umbrella species for the protection of other cave-adapted species of invertebrates of the massif.
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Transition to novel environments, such as groundwater colonization by surface organisms, provides an excellent research ground to study phenotypic evolution. However, interspecific comparative studies on evolution to groundwater life are few because of the challenge in assembling large ecological and molecular resources for species-rich taxa comprised of surface and subterranean species. Here, we make available to the scientific community an operational set of working tools and resources for the Asellidae, a family of freshwater isopods containing hundreds of surface and subterranean species. First, we release the World Asellidae database (WAD) and its web application, a sustainable and FAIR solution to producing and sharing data and biological material. WAD provides access to thousands of species occurrences, specimens, DNA extracts and DNA sequences with rich metadata ensuring full scientific traceability. Second, we perform a large-scale dated phylogenetic reconstruction of Asellidae to support phylogenetic comparative analyses. Of 424 terminal branches, we identify 34 pairs of surface and subterranean species representing independent replicates of the transition from surface water to groundwater. Third, we exemplify the usefulness of WAD for documenting phenotypic shifts associated with colonization of subterranean habitats. We provide the first phylogenetically controlled evidence that body size of males decreases relative to that of females upon groundwater colonization, suggesting competition for rare receptive females selects for smaller, more agile males in groundwater. By making these tools and resources widely accessible, we open up new opportunities for exploring how phenotypic traits evolve in response to changes in selective pressures and trade-offs during groundwater colonization.
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Isópodos , Animales , Filogenia , Isópodos/genética , Ecosistema , ADN , Secuencia de BasesRESUMEN
A new cave-dwelling species of the genus Scutogona Ribuat, 1913, S. minor n. sp., is described from caves of Sicó karst in central Portugal. The classification and delimitation of Scutogona vis-à-vis related genera, in particular Meinerteuma Mauriès, 1982, is discussed.
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Artrópodos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Animales , Artrópodos/anatomía & histología , Cuevas , Ecosistema , Femenino , Masculino , PortugalRESUMEN
Two new blind, cave-dwelling species of the genus Acipes Attems, 1937, are described from the Algarve, southern Portugal: A. machadoi n. sp. and A. bifilum n. sp. Acipes andalusius Enghoff & Mauriès, 1999, is reported from the mesovoid shallow substratum in Alicante (Spain), 250 km from the type locality in Andalusia.
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Artrópodos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Estructuras Animales/crecimiento & desarrollo , Animales , Artrópodos/anatomía & histología , Artrópodos/crecimiento & desarrollo , Tamaño Corporal , Ecosistema , Masculino , Tamaño de los Órganos , PortugalRESUMEN
Three new hypogean species of the Iberian genus Roncocreagris Mahnert, 1974 are described from mainland Portugal: R. borgesi sp. nov. and R. gepesi sp. nov. from caves in the Sicó massif, and R. occidentalis sp. nov. from caves in the Montejunto and Cesaredas karst plateau. This brings to nine the number of known hypogean species of the mostly Iberian genus Roncocreagris: five from Portugal and four from Spain. Ecological comments and new localities for some of the previously known species are also included.
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Arácnidos/clasificación , Distribución Animal , Estructuras Animales/anatomía & histología , Estructuras Animales/crecimiento & desarrollo , Animales , Arácnidos/anatomía & histología , Arácnidos/crecimiento & desarrollo , Tamaño Corporal , Ecosistema , Femenino , Masculino , Tamaño de los Órganos , PortugalRESUMEN
Background: Amongst the cave-dwelling millipedes (Diplopoda), there are several endemic species in Portugal with a very small geographical distribution. These species play an important role in the decomposition of organic matter in subterranean ecosystems and are vulnerable to disturbance from human activities, such as habitat destruction, pollution infiltrating from the surface and cave tourism. New information: We present the IUCN Red List profiles for cave-adapted millipedes (Diplopoda) from Portugal and propose conservation measures to prevent extinction. Overall, cave-adapted millipedes from Portugal represent an endemic part of the country's biodiversity and conservation efforts will help maintain the delicate ecological balance of subterranean ecosystems.
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A variety of veterinary and human medicinal products (VHMPs) are found in groundwater, an often-neglected habitat inhabited by species with unique traits, stygobitic species. It is crucial to understand the effect of VHMPs on stygobitic species because they may respond differently to stressors than surface species. Our hypothesis is that groundwater species may be more susceptible to environmental contaminants due to less plasticity in their detoxification response and acquisition of energy because subterranean habitats are more stable and isolated from anthropogenic activities. We performed a battery of biomarkers associated with important physiological functions on the stygobitic asellid crustacean Proasellus lusitanicus, after a 14-day exposure to acetaminophen, a commonly used pharmaceutical and pollutant of groundwaters. Our results show a decrease in total glutathione levels and an increase in glutathione S-transferase activity, suggesting a successful detoxification response. This helps explaining why acetaminophen did not cause oxidative damage, as well as had no effect on cholinesterase activity nor in aerobic production of energy. This study shows the remarkable capacity of P. lusitanicus to tolerate sublethal concentrations of VHMP acetaminophen. Most ecotoxicological studies on stygobitic species focused on the lethal effects of these compounds. The present study focuses on consequences at sublethal concentrations. Future studies should assess the stress levels induced to better predict and estimate the impacts of contaminants on groundwater ecosystems.
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Acetaminofén , Antioxidantes , Humanos , Acetaminofén/toxicidad , Antioxidantes/metabolismo , Ecosistema , Estrés Oxidativo , Glutatión/metabolismoRESUMEN
Climate change affects all ecosystems, but subterranean ecosystems are repeatedly neglected from political and public agendas. Cave habitats are home to unknown and endangered species, with low trait variability and intrinsic vulnerability to recover from human-induced disturbances. We studied the annual variability and cyclicity of temperatures in caves vis-à-vis surface in different climatic areas. We hypothesize that cave temperatures follow the average temperature pattern at the surface for each location with a slight delay in the signal, but we found three different thermal patterns occurring in caves: (1) high positive correlation and a similar thermal pattern to the surface, (2) low correlation and a slight thermal delay of the signal from the surface, and (3) high negative correlation with an extreme delay from the surface. We found daily thermal cycles in some caves, which may potentially control the circadian rhythms of cave organisms. Our results show that caves had lower thermal amplitude than the surface, and that thermal averages within caves approximately correspond to the to the annual average of surface temperature. Caves buffer external temperature and act as refugia for biota in extreme climatic events. Likewise, temperature increases at surface will lead to increment in caves, threatening subterranean biota and ecosystem services.
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Cuevas , Ecosistema , Animales , Humanos , Temperatura , Biota , Especies en Peligro de ExtinciónRESUMEN
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.
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Cambio Climático , Agua Subterránea , Animales , Ecosistema , Temperatura , Biodiversidad , Aclimatación , OxígenoRESUMEN
Background: Terrestrial isopods (Oniscidea) are the most diverse group of troglobionts in caves of continental Portugal. They occur in all karst regions of Portugal, play a major role in decomposition of organic matter in caves and may act as umbrella species for the conservation of all other cave-adapted invertebrates. New information: We present the IUCN Red List profiles for the cave-adapted terrestrial isopods from continental Portugal, based on recent distribution data from caves.
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BACKGROUND: The cave biodiversity of continental Portugal faces tremendous conservation challenges, mostly linked to their direct destruction and contamination infiltrating from the surface. Beetles are the most diverse insects and one of the most diverse arthropod groups in caves of Portugal. NEW INFORMATION: We present the IUCN Red List profiles for the cave-adapted beetles from continental Portugal, all endemic to their respective geological units and massifs. Ground beetles (Carabidae) are the most diverse family of cave-adapted beetles in continental Portugal, followed by rove beetles (Staphylinidae). Beetles in caves of Portugal are mostly terrestrial and only one species is known to have evolved to live in groundwater. Trechus is the most diverse genus with four species, followed by Domene with three species and by Speonemadus and Iberoporus, both with one cave-adapted species. The aim of this contribution is to assess all endemic cave-adapted species of beetles from continental Portugal and to support their specific protection, to promote adequate management of surface habitats and the establishment of priority areas for conservation. The main biodiversity erosion drivers that are impacting the conservation of the studied species are pollution infiltrating from the surface, urbaniation, modifications of the natural habitat for touristic purposes and mining, quarrying and energy production infrastructures.This document can be used in spatial planning and territory management in karst, based on the current scientific knowledge.
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Laboulbeniales are a highly specialized group of fungi living only on arthropods. They have high host specificity and spend their entire life-cycle on an arthropod host. Taxonomic characters of Laboulbeniales are based on the architecture of the cells of the parenchymal thallus, i.e. the visible part of the fungus outside the host. The extent of the fungus spreading inside the host-the haustorium-remains largely unknown. The attachment to the arthropod host is fundamental to understand the fungus-animal interaction, but how this truly occurs is unclear. Recent evidences question the strictly parasitic life-style of Laboulbeniales. We used micro-computed tomography (µCT) and 3D reconstructions to visualize, for the first time, the complete structure of Laboulbeniales species in situ on their hosts. We compared the haustoriate species, Arthrorhynchus nycteribiae on an insect host to the non-haustoriate species, Rickia gigas on a millipede host. Our results confirm that some Laboulbeniales species are ectoparasitic and have a haustorial structure that penetrates the host's cuticle, while others are ectobionts and are only firmly attached to the host's cuticle without penetrating it. The presence and the morphology of the haustorium are important traits for Laboulbeniales evolution, and key factors for future understanding of host dependence and specificity.
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Artrópodos/microbiología , Ascomicetos/fisiología , Interacciones Microbiota-Huesped , Animales , Artrópodos/ultraestructura , Especificidad del Huésped , Interacciones Huésped-ParásitosRESUMEN
Groundwater is an indispensable resource for humankind and sustainable biomes functioning. Anthropogenic disturbance threatens groundwater ecosystems globally, but to which extent groundwater organisms respond to stressors remains poorly understood. Groundwater animals are rare, with small populations, difficult to find and to breed in the lab, which poses a main challenge to the assessment of their responses to pollutants. Despite the difficulties, assessing the toxicity of a large spectrum of stressors to groundwater organisms is a priority to inform towards appropriate environmental protection of these ecosystems. We tested the sensitivity to CuSO4, diclofenac, and NaCl of a groundwater population of the copepod Diacyclops crassicaudis crassicaudis and compared its sensitivity with the model organism Daphnia magna. We ranked its sensitivity using a species sensitivity distribution (SSD) approach using the feasible data available for groundwater and surface crustaceans. Our results show that the most toxic compound was CuSO4 for which higher amount of data was recorded and wider variability in response was observed. It was followed by diclofenac, largely lacking data for groundwater-adapted organisms, and the least toxic compound was NaCl. The differential sensitivity between D. crassicaudis and D. magna was contaminant-dependent. As a general trend D. crassicaudis was always distributed in the upper part of the SSD curves together with other groundwater-adapted organisms. Our results highlight that the widespread groundwater populations of the D. crassicaudis species complex, which can be successfully breed in the lab, may provide a reasonable approach to assess the ecological effects of anthropogenic stressors in groundwater ecosystems.
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Copépodos , Agua Subterránea , Contaminantes Químicos del Agua , Animales , Daphnia , Ecosistema , Contaminantes Químicos del Agua/análisis , Contaminantes Químicos del Agua/toxicidadRESUMEN
How anthropogenic stressors affect biodiversity is a central question in a changing world. Subterranean ecosystems and their biodiversity are particularly vulnerable to change, yet, these species are frequently neglected in analyses of global biodiversity and assessments of ecological status and risk. Are these hidden species affected by anthropogenic stressors? Do they survive outside of the current thermal limits of their ecosystems? These and other important questions can be addressed with ecotoxicological testing, relating contaminants and temperature resistance of species with measured environmental concentrations and climatic data. Ecotoxicological knowledge specific to subterranean ecosystems is crucial for establishing thresholds for their protection, but such data are both scarce and scattered. Here, we review the existing ecotoxicological studies of these impacts to subterranean-adapted species. An effort that includes 167 measured endpoints and presents a database containing experimentally derived species' tolerance data for 28 contaminants and temperature, for 46 terrestrial and groundwater species, including fungi and animals. The lack of standard data among the studies is currently the major impediment to evaluate how stressors affect subterranean-adapted species and how differently they respond from their relatives at surface. Improving understanding of ecotoxicological effects on subterranean-adapted species will require extensive analysis of physiological responses to a wide range of untested stressors, standardization of testing protocols and evaluation of exposures under realistic scenarios.