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Caves are unique environments characterized by spatial limitations, partial or total absence of direct light, and scarcity of organic carbon and nutrients. Caves are shelters for a variety of adapted animals and microorganisms such as fungi, many of which are still unknown. Amphichorda is a fungal genus belonging to the family Bionectriaceae, which includes cave-dwelling and entomopathogenic species with biotechnological applications. In this study, a new fungal species was identified using morphological and multi-locus phylogenetic analyses of the ITS, LSU, and TEF loci, in the Gruta Velha Nova limestone cave located in the Southern Espinhaço Range, Monjolos, Minas Gerais, Brazil. During the exposure of potato dextrose agar plates to the cave environment, an insect from the family Rhaphidophoridae passed by and fed on the culture medium, resulting in three fungal isolates. Phylogenetic analyses showed that these isolates formed a clade distinct from all known species, leading us to introduce a new species, Amphichorda monjolensis, which may be associated with this insect. Here, we also proposed two new combinations for species of acremonium-like fungi in the Bionectriaceae: Bulbithecium globosisporum (synonym: Acremonium globosisporum) and Hapsidospora curva (synonym: Acremonium curvum). The discovery of A. monjolensis highlights the potential of caves as shelters for new species with significant biotechnological importance.

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

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Subterranean environments host a substantial amount of biodiversity, however assessing the distribution of species living underground is still extremely challenging. Environmental DNA (eDNA) metabarcoding is a powerful tool to estimate biodiversity in poorly known environments and has excellent performance for soil organisms. Here, we tested 1) whether eDNA metabarcoding from cave soils/sediments allows to successfully detect springtails (Hexapoda: Collembola) and insects (Hexapoda: Insecta); 2) whether eDNA mostly represents autochthonous (cave-dwelling) organisms or it also incorporates information from species living in surface environments; 3) whether eDNA detection probability changes across taxa with different ecology. Environmental DNA metabarcoding analyses detected a large number of Molecular Operational Taxonomic Units (MOTUs) for both insects and springtails. For springtails, detection probability was high, with a substantial proportion of hypogean species, suggesting that eDNA provides good information on the distribution of these organisms in caves. Conversely, for insects most of MOTUs represented taxa living outside caves, and the majority of them represented taxa/organisms living in freshwater environments (Ephemeroptera, Plecoptera and Trichoptera). The eDNA of freshwater insects was particularly abundant in deep sectors of caves, far from the entrance. Furthermore, average detection probability of insects was significantly lower than the one of springtails. This suggests that cave soils/sediments act as "conveyer belts of biodiversity information", possibly because percolating water lead to the accumulation of eDNA of organisms living in nearby areas. Cave soils hold a complex mix of autochthonous and allochthonous eDNA. eDNA provided unprecedented information on the understudied subterranean cave organisms; analyses of detection probability and occupancy can help teasing apart local eDNA from the eDNA representing spatially-integrated biodiversity for whole landscape.

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A new species of Cryptops, C. (Cryptops) legagussp. nov., occurs in caves in the Koanaka and Gcwihaba Hills in northwestern Botswana. Bayesian molecular phylogenetics using 18S rRNA, 28S rRNA, 16S rRNA and cytochrome c oxidase subunit I corroborates a morphological assignment to the subgenus Cryptops and closest affinities to southern temperate species in South Africa, Australia and New Zealand. The new species is not conspicuously modified as a troglomorph.

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A new genus of terrestrial isopod (Oniscidea) belonging to the family Trichoniscidae, subfamily Haplophthalminae, Baeticoniscus n. gen., is described. We discuss its affinities and differences with the other genera of the same subfamily, which are characterized by presenting a completely smooth third pleonite. Baeticoniscus bullonorum n. sp. is described and designated as a type species of the new genus. The specimens of this new isopod come from the Cueva de la Pileta of Benaoján in the province of Málaga (Andalusia, Spain).

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A new troglobitic species of the subfamily Pselaphinae (Coleoptera: Staphylinidae), Tychobythinus inopinatus sp. nov., is described from Monello Cave (Syracuse, Syracuse Province, Sicily). Major diagnostic features are illustrated based on both male and female specimens. The new species shows some adaptations to cave life, i.e., pale brown colour; setation consisting of long and flattened setae and suberect shorter setae; absence of wings; and anophthalmy and elongate legs and antennae. It can be easily separated from the related taxa by the different shapes of the head, palpi, gular carina of the male, and aedeagus. Tychobythinus inopinatus sp. nov. is known only from Monello Cave, a limestone cave in the south-eastern Sicily.

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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.

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A new species of Cryptops Leach, 1814, C. speleorex sp. nov., is described from Movile Cave, Dobrogea, Romania. The cave is remarkable for its unique ecosystem entirely dependent on methane- and sulfur-oxidising bacteria. Until now, the cave was thought to be inhabited by the epigean species C. anomalans, which is widespread in Europe. Despite its resemblance to C. anomalans, the new species is well-defined morphologically and molecularly based on two mitochondrial (cytochrome c oxidase subunit I COI and 16S rDNA) and one nuclear (28S rDNA) markers. Cryptops speleorex sp. nov. shows a number of troglomorphic traits such as a generally large body and elongated appendages and spiracles, higher number of coxal pores and saw teeth on the tibia of the ultimate leg. With this record, the number of endemic species known from the Movile Cave reaches 35, which ranks it as one of the most species-rich caves in the world.

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A new troglobitic Pselaphinae, Tychobythinus villasmundi sp. nov., is described from Villasmundo Cave (Melilli, Syracuse province, Sicily). Major diagnostic features are illustrated based on both male and female specimens. The new species shows some adaptations to cave life, such as pale brown colour, setation consisting of long and flattened setae and suberect shorter setae, absence of wings, anophthalmy, and elongated legs and antennae. It can be easily separated from the related taxa by the different structure of the aedeagus. Tychobythinus villasmundi sp. nov. is known only from Villasmundo Cave, a limestone cave in the south-eastern Sicily.

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Lithobiusmatulici Verhoeff, 1899 is redescribed based on type material and newly collected specimens. Strandiolusjugoslavicus Hoffer, 1937, described from another cave in the same region in Bosnia and Hercegovina, is presented as a junior subjective synonym of L.matulici (syn. nov.). L.matulici is shown to be most closely related to Lithobiusremyi Jawlowski, 1933, type species of the subgenus Thracolithobius Matic, 1962. The completeness of the chitin-lines on the forcipular coxosternite is discussed as a promising character for interspecific differentiation within Lithobiomorpha. Documentation of hitherto unknown semiaquatic behaviour in L.matulici and other cave-dwelling centipede species from Herzegovinian-, Montenegrin- and Pyrenean caves is presented.

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This paper is the first in a series describing the previously unstudied cave spiders from Cyprus. Two new species, Dysderocrateskibrisensis sp. n. and Harpacteakalavachiana sp. n., are described. Detailed morphological descriptions and diagnostic characteristics are presented. This is the first report of the genus Dysderocrates Deeleman-Reinhold & Deeleman, 1988 from Cyprus.

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Seven new species of the cavernicolous and anophthalmous genus Thaumastocephalus Poggi, Nonveiller, Colla, Pavicevic T. Rada, 2001 are described: T. bilandzijae sp. n., T. kirini sp. n., T. marsici sp. n., T. rujnicensis sp. n., T. slavkoi sp. n. and T. troglavi sp. n. from Croatia and T. dahnae sp. n. from Bosnia and Herzegovina. Aedeagi of all species are illustrated. A key to all species is provided. The records of all specimens of the genus treated here are given, and their distributions are discussed and shown on maps. The distribution of all genera of cavernicolous Pselaphinae in the Dinarides is discussed.

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The family Microstigmatidae is composed of two subfamilies, Microstigmatinae and Micromygalinae, seven genera and 16 species. Micromygalinae is monotypic, comprising the species Micromygalediblemma Platnick & Forster, 1982 from Panama. A new genus, Tonton is described as a new member of the Micromygalinae. Masteriaemboaba Pedroso, Baptista & Bertani, 2015, is transferred to the new genus and six new species from Brazil are described and attributed to Tonton gen. n.: the type species, T.itabirito sp. n., T.queca sp. n., T.matodentro sp. n. and T.sapalo sp. n., all from the state of Minas Gerais; T.ipiau sp. n. from the state of Bahia and T.quiteria sp. n. from the state of Maranhão. Among the cavernicolous species, only T.itabirito sp. n. is considered troglobitic by the total absence of eyes.

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The family Microstigmatidae is composed of two subfamilies, Microstigmatinae and Micromygalinae, seven genera and 16 species. Micromygalinae is monotypic, comprising the species Micromygale diblemma Platnick & Forster, 1982 from Panama. A new genus, Tonton is described as a new member of the Micromygalinae. Masteria emboaba Pedroso, Baptista & Bertani, 2015, is transferred to the new genus and six new species from Brazil are described and attributed to Tonton gen. n.: the type species, T. itabirito sp. n., T. queca sp. n., T. matodentro sp. n. and T. sapalo sp. n., all from the state of Minas Gerais; T. ipiau sp. n. from the state of Bahia and T. quiteria sp. n. from the state of Maranhao. Among the cavernicolous species, only T. itabirito sp. n. is considered troglobitic by the total absence of eyes.

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The orb-weaver spider Meta bourneti Simon 1922 (Araneae: Tetragnathidae) is one of the most common cave predators occurring in the Mediterranean basin. Although the congeneric M. menardi represented the model species in several studies, our knowledge of M. bourneti is only founded on observations performed on a handful of populations. In this study M. bourneti spiders were studied in caves of Monte Albo (Sardinia, Italy) over a year. Generalized Linear Mixed Models were used to analyze spider occupancy inside cave environments, as well as spider abundance. Analyses on M. bourneti occupancy and abundance were also repeated for adults and juveniles separately. Generalized Linear Models, were used to weight species absence based on its detection probability. Linear Mixed Models were used to detect possible divergences in subterranean spatial use between adult and juvenile spiders. Although widespread on the mountain, M. bourneti generally showed low density and low detection probability. Most of the individuals observed were juveniles. The spiders generally occupied cave sectors with high ceilings that were deep enough to show particular microclimatic features. Adults tended to occupy less illuminated areas than juveniles, while the latter were more frequently found in sectors showing high humidity. The abundance of M. bourneti was strongly related to high humidity and the presence of two troglophile species, Hydromantes flavus Wake, Salvador & Alonso-Zarazaga, 2005 (Amphibia: Caudata) and Oxychilus oppressus (Shuttleworth, 1877) (Gastropoda: Panpulmonata). The abundance of juveniles was related to sector temperature and humidity, the presence of H. flavus and O. oppressus and to morphological sector features. However, when only adults were considered, no significant relationships were found. Adult and juvenile spiders did not differ in their spatial distribution inside the caves studied, but a seasonal distribution of the species along cave walls was observed. Microclimate was one of the most important features affecting both the presence and abundance of M. bourneti in subterranean environments. Individuals tended to occupy lower heights during hot seasons.

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The processes of vicariance and dispersal are central to our understanding of diversification, yet determining the factors that influence these processes remains a significant challenge in evolutionary biology. Caves offer ideal systems for examining the mechanisms underlying isolation, divergence, and speciation. Intrinsic ecological differences among cavernicolous organisms, such as the degree of cave dependence, are thought to be major factors influencing patterns of genetic isolation in caves. Using a comparative phylogeographic approach, we employed mitochondrial and nuclear markers to assess the evolutionary history of two ecologically distinct groups of terrestrial cave-dwelling springtails (Collembola) in the genera Pygmarrhopalites (Arrhopalitidae) and Pogonognathellus (Tomoceridae) that are codistributed in caves throughout the Salem Plateau-a once continuous karst region, now bisected by the Mississippi River Valley in Illinois and Missouri. Contrasting phylogeographic patterns recovered for troglobiotic Pygmarrhopalites sp. and eutroglophilic Pogonognathellus sp. suggests that obligate associations with cave habitats can restrict dispersal across major geographic barriers such as rivers and valleys, but may also facilitate subterranean dispersal between neighboring cave systems. Pygmarrhopalites sp. populations spanning the Mississippi River Valley were estimated to have diverged 2.9-4.8 Ma, which we attribute to vicariance resulting from climatic and geological processes involved in Mississippi River Valley formation beginning during the late Pliocene/early Pleistocene. Lastly, we conclude that the detection of many deeply divergent, morphologically cryptic, and microendemic lineages highlights our poor understanding of microarthropod diversity in caves and exposes potential conservation concerns.

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Three new species of the genus Plato from caves in the states of Pará and Minas Gerais, Brazil, are described. P. novalimasp. n., from Minas Gerais, is the first record of the genus in the southeastern region of Brazil. P. ferriferussp. n. and P. striatussp. n., from Carajás, Pará, north of Brazil, are also described. The former is an extremely abundant species, whereas the latter has only one known male specimen. Cuacubagen. n. is proposed and represented by two new species, C. marianasp. n. (type species) and C. morrodopilarsp. n., both from the state of Minas Gerais. Morphology of genitalia in Cuacubagen. n. is similar to other Theridiosomatidae genera and is herein discussed. None of the proposed species presents troglomorphic adaptations. They are widespread, abundant inside caves in different and large karst areas, and each genus prefers different lithologies.

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This paper documents the collembolan fauna of the Campanet cave, in the province of Mallorca (Balearic Islands). Sampling was carried out using pitfall traps baited with beer and checked every 15 days. The cave fauna included the following species: Coecobrya tenebricosa, Disparrhopalites patrizi, Entomobrya pazaristei, Neelus murinus and Dicyrtomina ornata. In addition, two new species were found and are described here: Pseudosinella grauae sp. nov. and Oncopodura siquierae sp. nov. The former is characterised by an unpigmented body, eyes absent, dorsal chaetotaxy R0R1R2001/31/0101+2, with accessory chaeta 's' on Abd IV; all claws with three teeth; posterior chaetae on labial triangle as M1, M2, r, E, L1, L2, all ciliated except for r, a smooth microchaetae and smooth remaining anterior labial chaetae. O. siquierae sp. nov is typified by an unpigmented body; eyes absent; PAO as a single small vesicle; Ant IV with four distal leaf­shaped sensilla and a basal pointed cylindrical chaeta; dens with one proximal inner spine, one distal inner tooth and a large disto-external hook­like spine; and mucro with four teeth, the basal one distinctly pointed.

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Three new species of the genus Plato from caves in the states of Para and Minas Gerais, Brazil, are described. P. novalima sp. n., from Minas Gerais, is the first record of the genus in the southeastern region of Brazil. P. ferriferus sp. n. and P. striatus sp. n., from Carajas, Para, north of Brazil, are also described. The former is an extremely abundant species, whereas the latter has only one known male specimen. Cuacuba gen. n. is proposed and represented by two new species, C. mariana sp. n. (type species) and C. morrodopilar sp. n., both from the state of Minas Gerais. Morphology of genitalia in Cuacuba gen. n. is similar to other Theridiosomatidae genera and is herein discussed. None of the proposed species presents troglomorphic adaptations. They are widespread, abundant inside caves in different and large karst areas, and each genus prefers different lithologies.

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BACKGROUND: The underlying mechanisms and processes that prompt the colonisation of extreme environments, such as caves, constitute major research themes of evolutionary biology and biospeleology. The special adaptations required to survive in subterranean environments (low food availability, hypoxic waters, permanent darkness), and the geographical isolation of caves, nominate cave biodiversity as ideal subjects to answer long-standing questions concerning the interplay amongst adaptation, biogeography, and evolution. The present project aims to examine the phylogeographic patterns exhibited by two sympatric species of surface and cave-dwelling peracarid crustaceans (Asellus aquaticus and Niphargus hrabei), and in doing so elucidate the possible roles of isolation and exaptation in the colonisation and successful adaptation to the cave environment. RESULTS: Specimens of both species were sampled from freshwater hypogean (cave) and epigean (surface) habitats in Hungary, and additional data from neighbouring countries were sourced from Genbank. Sequencing of mitochondrial and nuclear loci revealed, through haplotype network reconstruction (TCS) and phylogenetic inference, the genetic structure, phylogeographic patterns, and divergence-time estimates of A. aquaticus and N. hrabei surface and cave populations. Contrasting phylogeographic patterns were found between species, with A. aquaticus showing strong genetic differentiation between cave and surface populations and N. hrabei lacking any evidence of genetic structure mediated by the cave environment. Furthermore, N. hrabei populations show very low levels of genetic differentiation throughout their range, which suggests the possibility of recent expansion events over the last few thousand years. CONCLUSIONS: Isolation by cave environment, rather than distance, is likely to drive the genetic structuring observed between immediately adjacent cave and surface populations of A. aquaticus, a predominantly surface species with only moderate exaptations to subterranean life. For N. hrabei, in which populations exhibit a fully 'cave-adapted' (troglomorphic) phenotype, the lack of genetic structure suggests that subterranean environments do not pose a dispersal barrier for this surface-cave species.

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