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1. Diplura is the sister group to insects and one of the three basal hexapod groups with unique entognathan mouthparts. The order is divided into 10 families, which include 1008 species in 141 genera, with a high proportion of monotypic genera. They are ubiquitous in soils and subsurface terrestrial habitats, as well as have an important role in overall biogeochemical cycles. 2. We present the first comprehensive review of the global biodiversity and ecology of Diplura. We highlight four aspects of this basal hexapod group: diversity in morpho logical body plans and sizes; ecology in terrestrial environments from soil to caves; food preference and trophic levels, and their biogeographical and paleobiogeographical significance. 3. Diplura depends on high humidity and moderate temperatures. They are presum ably very sensitive to anthropogenic pressures and climate change, and therefore are a suitable model for ecophysiological studies and evident priority targets for conservation. 4. We conclude that the future efforts should focus on establishing a molecular phy logeny to clarify the relationships between and within families, as well as to reveal global biogeographical patterns. This will require an increase in sampling effort in several regions of the globe, especially in tropical regions.

Current standardized laboratory test protocols use model species that have limitations to accurately assess native species responses to stressors. We developed and tested a novel acute insitu protocol for testing field-collected organisms. We used Asellus aquaticus and NaCl as a reference toxicant to test for the effects of location (laboratory vs. insitu), medium (synthetic vs. field water), substrate (presence vs. absence), and protocol replicability. We further tested the protocol using groundwater-adapted isopods: Proasellus assaforensis for the effect of location, P. cavaticus of medium and P.lusitanicus of substrate. Our results showed that A.aquaticus’ lethality obtained with the novel acute insitu protocol did not significantly differ from those from laboratory testing. However, laboratory tested P.assaforensis showed a higher sensitivity, suggesting that its acclimation to laboratory conditions might have pernicious effects. A. aquaticus and P. cavaticus showed a higher mortality using synthetic medium insitu and under laboratory conditions, which overestimated the stressor’s effect. Besides, substrate use had no significant effect. The novel acute insitu protocol allows the use of native species under realistic scenarios. It is particularly well adapted for assessing the risk of groundwater ecosystems but it can be applied to a wide range of ecosystems.

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.

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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.

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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Decomposition of organic matter is a process, which includes mostly physical breakdown and biochemical transformation of complex organic molecules into simpler organic and inorganic molecules. The decomposition of organic matter is an important contributor to ecosystem respiration, which together with photosynthesis controls the net carbon emission from ecosystems. Subterranean ecosystems are extended all over the subsurface of our planet, and lack of light and consequently of photosynthetic activity. Understanding the drivers that control the dynamics of the decomposition processes in the deep subterranean spaces is important because they might differ from those at surface, due to factors as low species diversity and abundance, low microbial biomass, nutrient poor conditions, less pronounced variation of temperature, and higher humidity inside cave. Here, we review the existing studies of organic matter decomposition in caves. Decomposition rates are known from only nine caves representing four biogeographic regions, including Europe, North and South America. Most of the studies were performed in the aquatic compartment of caves. The decay of nine different organic substrates have been followed and the incubation time varied from 36 to 439 days. From a cave located in Australia the mass loss of leaf material from three plant species was investigated after 9 days incubation in the terrestrial compartment of the cave. Based on these observations, litter quality seems to be an important driver of decomposition in caves, and invertebrates have a stimulating effect on the decomposition within individual cave zones. The degree of connection to the surface also influences decomposition rate inside the cave. The lack of standard data among the studies is currently the major impediment to evaluate how differently the process proceeds in the underground compared to the surface, and to disentangle the main drivers of decomposition in caves across biomes. Improving our understanding of organic matter decomposition dynamics in caves will require the standardization of protocols and evaluation of the process over space and time, and a better comprehension on how decomposition changes over latitudinal, altitudinal and depth gradients.

Anthropogenic impacts in groundwater ecosystems remain poorly known. Climate change is omnipresent, while groundwater salinization poses serious long-term environmental problems in arid and semi-arid regions, and is exacerbated by global warming. Both are present threats to the conservation of groundwater ecosystems, which harbour highly specialized species, with peculiar traits and limited geographic distributions. We tested the temperature and salinity tolerance of groundwater-adapted invertebrates to understand the effect of global warming and salinization in groundwater ecosystems. We used species representative of groundwater-adapted crustaceans: two copepods (harpacticoid and cyclopoid) and one syncarid, endemic to Australia. Our results show that 50% of the populations died at salt concentrations between 2.84 to 7.35 g NaCl/L after 96 h, and at 6.9 °C above the ambient aquifer temperature for copepods and more than 10 °C for syncarids. Both copepods were more sensitive to temperature and NaCl than the syncarid. We calculated a salinity risk quotient of 9.7 and predicted the risk of loss of 10% of syncarid and 20% of copepod population abundances under a worst-case scenario of global warming predictions for 2070. These results highlight that both salinity and temperature increases pose a risk to the ecological integrity of groundwater ecosystems.

The first eyeless species of the suborder Spirostreptidea, Odontostreptus fadriquei sp. nov., is described from a cave in Morocco. The new species, which exhibits some troglomorphic traits, is compared with the highly variable O. maroccanus (Attems, 1914) of which new material is presented.

Laboulbeniales are highly specialized arthropod-associated fungi. The majority of the almost 2200 known species live on insects, although they also occur on other arthropod hosts. Recently, the number of Laboulbeniales associated with millipedes has increased considerably. Here we describe the first species of a Laboulbeniales fungus, Troglomyces twitterisp. nov., from an American millipede. The new species was initially discovered on a photo of Cambala annulata (Say, 1821) from Ohio, USA, which had been shared on Twitter. A subsequent microscopic study of Cambala millipedes in museum collections in Denmark and France confirmed the discovery.

Este artículo contiene 138 páginas, 213 figuras, 14 tablas., Diplura is a group of entognathous hexapods, often considered a sister group to insects. They play an important role in recycling organic matter in soil and subterranean terrestrial ecosystems. The Campodeidae is the most diverse family, divided into four subfamilies. The subfamily Plusiocampinae has a subterranean life-style with many species distributed in the Euro-Mediterranean area. The incertae sedis tachycampoids (“lignée Tachycampoïde”) is a group within the family Campodeidae that share with the Plusiocampinae a strong preference for subterranean habitats and several morphological characters, such as slender body shape, elongated appendages, considerable increment in the number of antennomeres and cercal articles, and complexity of sensorial structures. The present monograph provides a taxonomic revision of the subfamily Plusiocampinae and the genera belonging to the tachycampoid lineage from Europe and the Mediterranean region. It comprises detailed morphological descriptions and illustrations together with data on the habitats and distributions of 87 species, 10 subspecies and 11 affinis forms. Seven new species are described among those, namely: Plusiocampa (Plusiocampa) apollo Sendra, Giachino & Vailati sp. nov., P. (P.) chiosensis Sendra & Gasparo sp. nov., P. (P.) dublanskii Sendra & Turbanov sp. nov., P. (P.) hoffmanni Sendra & Paragamian sp. nov., P. (P.) rhea Sendra sp. nov., P. (P.) ternovensis Sendra & Borko sp. nov. and P. (Venetocampa) ferrani Sendra & Delić sp. nov., DS’s field trips were funded by the Serbian Ministry of Education, Science and Technology (Grant 173038); KP was partially funded by the HISR project “Conservation of the Cave Fauna of Greece” funded by the MAVA Foundation and WWF Greece; PMG and DV carried out sampling in Greece using permits from the Ministry of Environment no. 124085/1362/2015, 135366/373/2016 and 166238/248/2018; IT was funded within the framework of the state assignment of FASO Russia (themes no. АААА-А18-118012690106-7 and АААА-А18-118012690105), supported in part by RFBR (project no. 17-54-40017Абх_а); sampling efforts by AF and colleagues resulted from various projects on the evolution of cave fauna diversity, some of them funded by the German Research Foundation, DFG (DFG FA 1042/1-1 and DFG BA 2152/14-1); PMG and DV were partly supported by the program “Research Missions in the Mediterranean Basin” sponsored by the World Biodiversity Association onlus XLI contribution; ASPSR was supported by a research grant (15471) from Villum Fonden.

Five decades ago, a landmark paper in Science titled The Cave Environment heralded caves as ideal natural experimental laboratories in which to develop and address general questions in geology, ecology, biogeography, and evolutionary biology. Although the 'caves as laboratory' paradigm has since been advocated by subterranean biologists, there are few examples of studies that successfully translated their results into general principles. The contemporary era of big data, modelling tools, and revolutionary advances in genetics and (meta)genomics provides an opportunity to revisit unresolved questions and challenges, as well as examine promising new avenues of research in subterranean biology. Accordingly, we have developed a roadmap to guide future research endeavours in subterranean biology by adapting a well-established methodology of 'horizon scanning' to identify the highest priority research questions across six subject areas. Based on the expert opinion of 30 scientists from around the globe with complementary expertise and of different academic ages, we assembled an initial list of 258 fundamental questions concentrating on macroecology and microbial ecology, adaptation, evolution, and conservation. Subsequently, through online surveys, 130 subterranean biologists with various backgrounds assisted us in reducing our list to 50 top-priority questions. These research questions are broad in scope and ready to be addressed in the next decade. We believe this exercise will stimulate research towards a deeper understanding of subterranean biology and foster hypothesis-driven studies likely to resonate broadly from the traditional boundaries of this field. S.M acknowledge support by the CAWEB project “Testing macroecological theory using simplified systems”, funded by the European Commission through Horizon 2020 Marie Skłodowska-Curie Actions (MSCA) individual fellowships (Grant no. 882221). I.R.A. was supported by Portuguese funds through FCT – Fundação para a Ciência e a Tecnologia, I.P., under the Norma Transitória – DL57/2016/ CP1375/CT0003. D.E. was supported by IFREMER and by the CERES “Climate change and European Aquatic Resources” project funded by European Commission through Horizon 2020 research and innovation programme (Grant no. 678193). C.F., Z.F., and M.Z. were supported by the Slovenian Research Agency (program P1-0184, project N1-0069). J.E.K. was funded by NSF awards DEB1754231 and IOS1933428, and EDGE award 1923372. F.M. was supported by the French National Research Agency projects CONVERGENOMICS (ANR-15-CE32-0005) and EUR H2O’Lyon (ANR-17-EURE-0018). A.M. was supported by the ANCAVE project “Anchialine caves to understand evolutionary processes”, funded by the European Commission through Horizon 2020 Marie Skłodowska-Curie Actions (MSCA) individual fellowships (Grant no. 745530). P.A.V.B. was supported by the project AZORESBIOPORTAL – PORBIOTA (ACORES-01-0145-FEDER-000072), financed by FEDER in 85% and by Azorean Public funds by 15% through Operational Program Azores 2020. A.S.P.S.R. was supported by VILLUM FONDEN (Grant no. 15471) and by a Carlsbergfondet grant (CF19-0609). T.G.P. was supported by Vale S.A. and Fundação de Amparo à Pesquisa de Minas Gerais (FAPEMIG) for individual fellowship (RDP 00092-18). T.P. was supported by the Karst Research Programme P6-0119, LifeWatch ERIC, RI-SI LifeWatch, and EU H2020 project eLTER. info:eu-repo/semantics/publishedVersion

Diplurans (Hexapoda) are considered the ‘ideal cavernicolous animal’ having one of the highest ratios of cave-adapted vs. non-cave-adapted species. They are successful colonizers of subterranean habitats, thriving in all cryptic, dark, terrestrial environments. Diplurans play an important role in the decomposition of organic matter below the ground and are sensitive to anthropogenic pressures. We present the first comprehensive review about cave Diplura diversity, ecology, evolution, distribution and biogeography. We provide a roadmap for research questions regarding the ecology, aimed at stimulating the pursuit of new studies on this fascinating group. Filling these current knowledge gaps will contribute to conservation efforts for cave ecosystems.

The new species of millipedeCylindroiulusvillumiis described from a cave in the Estremenho karst massif in central Portugal. It is the first cave-adapted species of its genus with a strict subterranean life-style in continental Europe, and is the fifth blind species of the genus. The new species is illustrated with photographs and diagrammatic drawings. It is tentatively placed in the purely IberianCylindroiulusperforatus-group. The differences between the new species and its relatives are discussed, as well as its adaptations to a subterranean life-style.

Five new species of the recently created genus Occidenchthonius Zaragoza, 2017 are described from caves of Portugal: Occidenchthonius alandroalensis sp. nov., O. algharbicus sp. nov., O. duecensis sp. nov., O. goncalvesi sp. nov. and O. vachoni sp. nov. The species Occidenchthonius cardosoi (Zaragoza, 2012) and Chthonius ischnocheles (Hermann, 1804) are reported from new localities in different karst units of Portugal. New morphological characters are proposed for use in the Chthoniidae taxonomy. An updated key to the genus Occidenchthonius is given.

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. 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.

The mesovoid shallow substratum (MSS) can act as a climatic refuge for invertebrates, as a biogeographic corridor to deeper substrates or as a permanent habitat for some species. This study characterizes the seasonal invertebrate diversity and abundance of MSS ecosystems in central Portugal focusing on Diplopoda, Diplura, Orthoptera and Coleoptera during one year. Sampling was performed with standard MSS pitfalls in scree slopes (colluvial MSS) of karst areas and environmental parameters (temperature, pH, conductivity, water content, organic carbon, nitrate, phosphate and ammonium) were quantified. Our results show that winter was the season with the highest arthropod abundance and that the MSS acts as a permanent habitat for chordeumatidan millipedes and as a climatic refuge for orthopterans and most beetles. All Diplura collected belong to a single species known previously from surface habitats in the Iberian Peninsula, which does not seem to use the Portuguese MSS as a refuge. MSS habitats in central Portugal, classified as western Mediterranean and thermophile deposits protected by the Natura 2000 network based on plant communities and geology, revealed an abundant and diverse invertebrate community that urges characterization and protection.

The aim of this study was to unravel the relative role played by speleogenesis (i.e., the process in which a cave is formed), landscape‐scale variables, and geophysical factors in the determination of species richness in caves. Biological inventories from 21 caves located in the southeastern Iberian Peninsula along with partial least square (PLS) regression analysis were used to assess the relative importance of the different explanatory variables. The caves were grouped according to the similarity in their species composition; the effect that spatial distance could have on similarity was also studied using correlation between matrices. The energy and speleogenesis of caves accounted for 44.3% of the variation in species richness. The trophic level of each cave was the most significant factor in PLS regression analysis, and epigenic caves (i.e., those formed by the action of percolating water) had significantly more species than hypogenic ones (i.e., those formed by the action of upward flows in confined aquifers). Dissimilarity among the caves was very high (multiple‐site βsim = 0.92). Two main groups of caves were revealed through the cluster analysis, one formed by the western caves and the other by the eastern ones. The significant—but low—correlation found between faunistic dissimilarity and geographical distance (r = .16) disappeared once the caves were split into the two groups. The extreme beta‐diversity suggests a very low connection among the caves and/or a very low dispersal capacity of the species. In the region under study, two main factors are intimately related to the richness of terrestrial subterranean species in caves: the amount of organic material (trophic level) and the formation process (genesis). This is the first time that the history of a cave genesis has been quantitatively considered to assess its importance in explaining richness patterns in comparison with other factors more widely recognized.

The species Paraphaenops breuilianus (Jeannel, 1916) is an Iberian iconic cave beetle with “aphaenopsian” facies, monospecific until now. An extensive field work over the last years allowed the revision of this peculiar genus using the combinations of morphological, molecular and ecological approaches and led to the description of new taxa Paraphaenops fadriquei Ortuno and Faille sp. nov., Paraphaenops breuilianus espanoli Ortuno and Faille ssp. nov. and the full larval diagnosis of the third larval instar of the type species. The sequencing of two mitochondrial (cox1, rrnl + tRNA-Leu + nad1) and one nuclear (LSU) gene fragments evidenced a strong divergence for all markers considered between P. breuilianus and the new species P. fadriquei sp. nov. The strong genetic differences between the two taxa contrast singularly with the extreme morphological homogeneity of the genus. The ecological data recorded for Paraphaenops Jeannel, 1916 species show that they are stenotherms and stenohygrobionts, perfectly adapted to the subterranean environment, living at temperatures between 4.4 and 10 °C with high relative humidity. The new localities and taxa enlarge the former distribution area from 72 to 5091 square kilometres sprinkled between twelve caves: six in the Mola de Cati (P. breuilianus breuilianus); three in the aforementioned karst region towards the south, crossing the geographical barrier “Barranc de Regatxol” (P. breuilianus espanoli ssp. nov.); and three, 60 km towards the west, in a different karst region: the “Sierra de la Dehesas” (P. fadriquei sp. nov.) This geographical range overlaps a geostructural area quoted as “junction area” between the Iberian System and the Catalan coastal Mountain range, an area up to 1000 meters above sea level.

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.

Jeekelosoma Mauriès, 1985, is upgraded from subgenus status under Eviulisoma Silvestri, 1910 to full genus status. The type species, Jeekelosoma abadi (Mauriès, 1985) is redescribed based on topotypical material from a cave in Morocco. Jeekelosoma heptarachne sp. nov. and J. viginti sp. nov. are described from two further Moroccan caves.

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.

As a consequence of the growing global dependence on groundwater resources, environmental risk assessments (ERA) for groundwater are increasingly required and, with that, ecotoxicological studies with groundwater fauna (stygofauna). However, the literature on the ecotoxicological studies with stygobiotic species (i.e. species that complete their life cycle exclusively in groundwater) has not expanded significantly since the first paper published in this field. The limitations regarding the use of stygobiotic species for ecotoxicological testing are clear and consistent across the globe; stygobiotic species are often 1) naturally present in low numbers, 2) difficult to collect, and 3) difficult to culture under laboratory conditions. This paper reviews the methods used in ecotoxicological studies performed with stygobiotic species, and provides ten recommendations for Good Laboratory Practice (GLP) for such tests. The recommendations focused on the following 10 points: 1) the taxonomic identification, the life stage/size and gender of the test organisms; 2) collection methodology of the organisms, including collection location, conditions and methods; 3) holding and acclimation conditions in the laboratory; 4) exposure conditions such as test set up and exposure time, number of replicates and densities of organisms in tests and in test vessels; 5) range-finding test set up and schedule; 6) final test design, including details of controls and treatments, and replication options; 7) incubation conditions, specifying temperature, pH and oxygenation levels throughout the test; 8) test duration; 9) observations and endpoints; 10) test validity criteria and compliance. The recommendations were developed for the purpose of supporting future short-term ecotoxicological testing with stygofauna through providing consistency in the protocols. A discussion of the potential implications for groundwater managers and decision-makers committed to ERA for groundwater is included.

The whip spider genus Sarax Simon, 1892 is widely distributed throughout Southeast Asia and part of the Indo-Malayan region. The genus is recorded from several Indonesian islands, but no species are known from inside the area that comprises the biogeographical region of Wallacea, despite being recorded from both sides of the area. An expedition to survey the biology of caves in Timor-Leste (formerly East-Timor) discovered populations of amblypygids living underground and including a remarkable new species of Sarax, S.timorensissp. n., the first Amblypygi known from the island of Timor. The new species is here described bears the unique character state of two pairs of lateral eyes, instead of three or none as in all other living species of Amblypygi, and expands the biogeographic range of the genus. New records of amblypygids are given for two caves in Timor-Leste. A detailed description and a discussion of its distribution and the species characters are also provided.

Fungal species identities are often based on morphological features, but current molecular phylogenetic and other approaches almost always lead to the discovery of multiple species in single morpho-species. According to the morphological species concept, the ant-parasitic fungusRickia wasmannii(Ascomycota, Laboulbeniales) is a single species with pan-European distribution and a wide host range. Since its description, it has been reported from ten species ofMyrmica(Hymenoptera, Formicidae), of which two belong to therubra-group and the other eight to the phylogenetically distinctscabrinodis-group. We found evidence forR. wasmanniibeing a single phylogenetic species using sequence data from two loci. Apparently, the original morphological description (dating back to 1899) represents a single phylogenetic species. Furthermore, the biology and host-parasite interactions ofR. wasmanniiare not likely to be affected by genetic divergence among different populations of the fungus, implying comparability among studies conducted on members of different ant populations. We found no differences in total thallus number on workers betweenMyrmicaspecies, but we did observe differences in the pattern of thallus distribution over the body. The locus of infection is the frontal side of the head inMyrmica rubraandM. sabuletiwhereas inM. scabrinodisthe locus of infection differs between worker ants from Hungary (gaster tergites) and the Netherlands (frontal head). Possible explanations for these observations are differences among host species and among populations of the same species in (i) how ant workers come into contact with the fungus, (ii) grooming efficacy, and (iii) cuticle surface characteristics.

In light of recent alarming trends in human population growth, climate change, and other environmental modifications, a “Warning to humanity” manifesto was published in BioScience in 2017. This call reiterated most of the ideas originally expressed by the Union of Concerned Scientists in 1992, including the fear that we are “pushing Earth's ecosystems beyond their capacities to support the web of life.” As subterranean biologists, we take this opportunity to emphasize the global importance and the conservation challenges associated with subterranean ecosystems. They likely represent the most widespread nonmarine environments on Earth, but specialized subterranean organisms remain among the least documented and studied. Largely overlooked in conservation policies, subterranean habitats play a critical role in the function of the web of life and provide important ecosystem services. We highlight the main threats to subterranean ecosystems and propose a set of effective actions to protect this globally important natural heritage.

Iberoporus pluto sp. n., the first stygobiont beetle from Portugal (Dytiscidae, Hydroporinae), is described from a single female from the cave Soprador do Carvalho (Coimbra). The species is highly troglomorphic, depigmented, blind, and with elongated appendages not adapted for swimming. A molecular phylogeny based on a combination of three mitochondrial and two nuclear genes showed the new species to be sister to I. cermenius Castro & Delgado, 2001 from Córdoba (south of Spain), within the subtribe Siettitiina of the tribe Hydroporini. Both species are included in a clade with Siettitia avenionensis Guignot, 1925 (south of France) and Rhithrodytes agnus Foster, 1992 and R. argaensis Fery & Bilton, 1996 (north of Portugal), in turn sister to the rest of species of genus Rhithrodytes Bameul, 1989, in what is here considered the Siettitia group of genera. We resolve the paraphyly of Rhithrodytes by transferring the two Portuguese species to Iberoporus Castro & Delgado, 2001, I. agnus (Foster, 1992), comb. n. and I. argaensis (Fery & Bilton, 1996), comb. n., Molecular work was partly funded by projects CGL2013-48950-C2-1-P and CGL2016-76705-P (AEI/FEDER, UE) to IR. ASR is supported by a research grant (15471) from VILLUM FONDEN.

The postembryonic development of millipedes includes a series of stadia separated by moults, a process known as anamorphosis. The moulting process and especially the moulting into maturity, i.e., with fully developed copulatory organs, remains unknown for most millipede species. We have kept specimens of Lusitanipus alternans (Verhoeff, 1893) in the laboratory for one year and studied its moulting process, including the first study of the maturation moult in the order Callipodida. Unlike the typical silk cocoon reported for other callipodidans, this species builds a new type of solid moulting chamber, using the available substrate reinforced by a silken web. We present the detailed ultrastructure of the moulting chamber and silk. It takes five days to build the moulting chamber and between 29 (female) and 35 (male) days to shed the exuviae. The male maturation moult is preceded by an evagination of a gonopodal sac between the 6th and 7th body rings, in which the gonopods are developed. Females evaginated completely their vulval sacs, retracting them after shedding the exuviae. Vulval sac size seems to increase with the progressive reduction of the second pair of legs.

Antibiotics have been developed for more than 80 years but presently there is an evident decline on their effectiveness due to the development of resistance by pathogens. The arising of multi drug resistant pathogens, has become a serious threat to public health. Microorganisms play a central role in nature as well as in drug discovery with a long track record as important sources of bioactive natural products revealing a wide array of molecules. Although many approaches have been applied to solve this crisis, in the last decades the focus on extreme habitats as gain attention due to peculiar features and due to the presence of unique microbial populations inhabiting these ecosystems. Caves are considered extreme environments, usually oligotrophic, being in general devoid of light, and presenting stable conditions (temperature, humidity) throughout the year. We present the first analysis of antimicrobial activity of bacteria isolated from Cerâmica Cave, located in the Sicó karst massif, central Portugal. Microorganism acquisition was performed through a variety of physical and chemical treatments: dry-heat, wet-heat, dry-heat plus phenol (1,5%), phenol (1,5%), microwave irradiation, rehydration centrifugation, and no-treatment, along with three media: R2A agar, Actinomycetes isolation agar and Yeast-malt agar. A primary antimicrobial activity screening, through cross streak method, was performed in 28 isolates, using two different incubation periods and different target agents (Staphylococcus aureus, Bacillus cereus, Klebsiella pneumoniae, Escherichia coli, Pseudomonas aeruginosa, Aeromonas salmonicida, Salmonella typhimurium and Enterococcus faecalis). A total of 7 isolates presented antagonistic activity against B. cereus, P. aeruginosa, E. coli, A. salmonicida S. typhimurium and S. aureus. This study proves the ability of cave microorganisms to inhibit both Gram positive and Gram negative pathogenic strains, highlighting the potential of caves microbiome to be one of the answers to solve the current global crisis of multi drug resistant pathogens.

The whip spider genus Sarax Simon, 1892 is widely distributed throughout Southeast Asia and part of the Indo-Malayan region. The genus is recorded from several Indonesian islands, but no species are known from inside the area that comprises the biogeographical region of Wallacea, despite being recorded from both sides of the area. An expedition to survey the biology of caves in Timor-Leste (formerly East-Timor) discovered populations of amblypygids living underground and including a remarkable new species of Sarax, S.timorensissp. n., the first Amblypygi known from the island of Timor. The new species is here described bears the unique character state of two pairs of lateral eyes, instead of three or none as in all other living species of Amblypygi, and expands the biogeographic range of the genus. New records of amblypygids are given for two caves in Timor-Leste. A detailed description and a discussion of its distribution and the species characters are also provided.

Pharmaceutical compounds have been extensively used in medicine and currently detected in trace concentrations (ranging from ng/L to mg/L) in both surface and groundwater bodies worldwide in the last decades. Despite being mostly found in low concentrations, these are persistent compounds that are continuously discharged into aquatic ecosystems. Pharmaceutical compounds are also present in complex mixtures and the actual absence of appropriate wastewater treatment specifically targeting their elimination, renders them emerging contaminants with high risk for surface and groundwater ecosystems. None of these compounds is included in the European Water Framework Directive as a priority substance and comprehensive data are still being built regarding pharmaceuticals toxicity on standard surface aquatic taxa and almost none concerning groundwater-dweller species. However, the current Directive 2008/105/EC has included some antibiotics, steroids, phenolic-antioxidants and the anti-inflammatory diclofenac in the established watch list for monitoring substances regarding water quality policy. Assessing the toxicity of pharmaceutical compounds in a large spectrum of aquatic environments is needed to assist a realistic environmental risk assessment. Hence, including subterranean species is a priority for establishing conservation measures in groundwater ecosystems. Copepod crustaceans are widely represented in freshwater bodies and dominant in groundwater ecosystems. We have selected the cosmopolitan (freshwater and groundwater) crustacean species Diacyclops crassicaudis crassicaudis Sars, 1863 and the freshwater model species Daphnia magna Müller, 1785 to estimate their response to acute exposures to the pharmaceutical compound diclofenac. Our preliminary results indicate that diclofenac was more toxic to the copepodites C1-C2 than to the neonates of D. magna (half maximal effective concentration: 63 mg/L and 111.3 mg/L at 48 h respectively) (See Suppl. material 1). However, further long-term toxicity testing at more environmentally relevant concentrations and the use of species with similar ecological features and more closely related are required for a better understanding of the potential of pharmaceutical compounds to harm the groundwater biota.

A catalogue of type specimens of basal hexapods (Collembola, Protura, Diplura, Archaeognatha, Zygentoma) in the collection of the Natural History Museum of Denmark, University of Copenhagen is presented. The NHMD collection harbours type material of 175 species, and it is composed of 132 holotypes, 4 neotypes, 3 syntypes, 32 lectotypes, and 39 paratypes. For Diplura one neotype is newly designated. The order Protura is represented by 5 families, Collembola by 7 families, Diplura by 6 families, Archaeognatha by 1 family and Zygentoma by 2 families.

The new species of millipede Cylindroiulusvillumi is described from a cave in the Estremenho karst massif in central Portugal. It is the first cave-adapted species of its genus with a strict subterranean life-style in continental Europe, and is the fifth blind species of the genus. The new species is illustrated with photographs and diagrammatic drawings. It is tentatively placed in the purely Iberian Cylindroiulusperforatus-group. The differences between the new species and its relatives are discussed, as well as its adaptations to a subterranean life-style.

We describe a new anthroleucosomatid millipede, Heterocaucaseuma deprofundum sp. nov., found in the second and third deepest caves of our planet, Krubera-Voronja and Sarma, in Abkhazia, western Caucasus. A brief discussion on its taxonomy and relationships within the Caucaseuma complex and genus Heterocaucaseuma is provided, together with new faunistic records on other three species of Heterocaucaseuma. Notes on the distribution and troglomorphy of the genus Heterocaucaseuma are added. Ecological information and data on the subterranean community of the Krubera-Voronja Cave are updated. We also discuss the vertical distribution of the new species in caves and the presence of an ectoparasitic fungi of the order Laboulbeniales. This new species is currently the world's deepest-occurring millipede, also representing one of the largest known chordeumatidans.

Recently, there has been an increasing demand for new molecules with bioactivity as a result of the increasing number of multi drug resistant bacteria and also of the need to find treatment for various diseases. Thus, to fill this urging need there has been a great investment in developing new techniques to speed up and increase the possibility of finding new molecules with therapeutic potential. Typically, the first stage of drug discovery process relies on in vitro assays and conventional high-throughput LC-MS-MS approaches. This combined with unique extreme environments, as a source of new molecules, has proved to be a good approach to enlarge natural product drug discovery. Considering the present demand for new bioactive molecules, together with the fact that cave microorganisms are already considered an important, yet largely unexplored source, our study aimed at assessing the bioactive potential of cave bacterial extracts, namely its antimicrobial and antitumoral potential. From six soil samples collected in the Cerâmica cave, at the Sicó karst massif, centre of Portugal, a total of 409 isolates were obtained. After a primary screening with the cross-streak method, 91 isolates showed antagonistic activity against four pathogenic agents (Escherichia coli, Pseudomonas aeruginosa, Aeromonas salmonicida and Bacillus aquimaris). Afterwards the secondary screening was carried out with the three isolates that presented higher antimicrobial activity. For the secondary screening, cell-free supernatants (CFS) were tested against eight pathogenic agents (Escherichia coli, Pseudomonas aeruginosa, Aeromonas salmonicida, Bacillus cereus, Enterococcus faecalis, Klebesiella pneumoniae, Salmonella typhimurium and Staphylococcus aureus). The chemical analysis of the CFS is discussed, as well as the results from the in vitro tests with human cell lines to evaluate its effects on cell morphology, viability and proliferation (Suppl. material 1).