Groundwater is a hidden global keystone ecosystem.

Groundwater is a vital ecosystem of the global water cycle, hosting unique biodiversity and providing essential services to societies. Despite being the largest unfrozen freshwater resource, in a period of depletion by extraction and pollution, groundwater environments have been repeatedly overlooked in global biodiversity conservation agendas. Disregarding the importance of groundwater as an ecosystem ignores its critical role in preserving surface biomes. To foster timely global conservation of groundwater, we propose elevating the concept of keystone species into the realm of ecosystems, claiming groundwater as a keystone ecosystem that influences the integrity of many dependent ecosystems. Our global analysis shows that over half of land surface areas (52.6%) has a medium-to-high interaction with groundwater, reaching up to 74.9% when deserts and high mountains are excluded. We postulate that the intrinsic transboundary features of groundwater are critical for shifting perspectives towards more holistic approaches in aquatic ecology and beyond. Furthermore, we propose eight key themes to develop a science-policy integrated groundwater conservation agenda. Given ecosystems above and below the ground intersect at many levels, considering groundwater as an essential component of planetary health is pivotal to reduce biodiversity loss and buffer against climate change.

The role of turnover in structuring subterranean aquatic communities in Southeastern Brazil.

Similarly to other animal communities, the diversity of subterranean aquatic fauna is influenced by several factors and processes, such as habitat fragmentation, dispersion, environmental heterogeneity, and physical and chemical water characteristics. Here, we studied cave aquatic communities of the Alto Ribeira hydrographic basin, regarding troglobitic and non-troglobitic species, located in a single karst area to evaluate the influence of sub-basins in fauna differentiation. We investigated how abiotic variables (flow, electrical conductivity, temperature, pH, and substrate) influence the fauna composition and the contribution of beta diversity components (nestedness and turnover) in explaining communities' dissimilarities. Fauna composition differed between sub-basins, as most species did not co-occur in different caves. Caves with higher flow and substrate diversity were the richest. In addition, each cave community was influenced by a unique set of abiotic variables. Dissimilarity among caves was mainly explained by turnover, and our findings suggest the restricted species distribution could be due to ecological (e.g., limited dispersion capacity, tolerance to abiotic variables), hydrogeological (e.g., dispersion barriers, isolation of sub-basins), and historical (e.g., colonization, paleoclimatic events) factors and processes. Therefore, different elements are responsible for determining the composition of cave aquatic communities in different sub-basins, reflecting the variability within a single karst area.

Gross morphology of the brain and some sense organs of subterranean pencil catfishes of the genus Ituglanis Costa and Bockmann, 1993 (Siluriformes, Trichomycteridae), with a discussion on sensory compensation versus preadaptation in subterranean fishes.

Subterranean organisms provide excellent opportunities to investigate morphological evolution, especially of sensory organs and structures and their processing areas in the central nervous system. We describe the gross morphology of the brain and some cephalic sensory organs (olfactory organ, eye, semicircular canals of the inner ear) and the swim bladder (a non-sensory accessory structure) of subterranean species of pencil catfishes of the genus Ituglanis Costa and Bockmann, 1993 (Siluriformes, Trichomycteridae) and compare them with an epigean species of the genus, Ituglanis goya Datovo, Aquino and Langeani, 2016. We compared qualitatively the size of the different brain regions and sense organs of the subterranean species with those of the epigean one, searching for modifications possibly associated with living in the subterranean environment. Our findings suggest that species of Ituglanis exhibit sensory characteristics that are preadaptive for the subterranean life, as only slight modifications were observed in the brains and sense organs of the subterranean species of the genus when compared with the epigean one. Because most subterranean fish species belong to lineages putatively preadapted for subterranean life, our results, discussed in the context of available information on the brain and sense organs of other subterranean species, help identify general trends for the evolution of the brain and sensory organs of subterranean fishes in general.

Influence of substrate on the richness and composition of Neotropical cave fauna.

The food base in the subterranean environment consists mainly of allochthonous materials. In this environment the resources are distributed generally in a heterogeneous dispersed way and the distribution of resources and their availability determine where the terrestrial invertebrates will reside, which is important for understanding ecological relationships and to establish conservation strategies. Thus, we tested how the complexity of substrates influences the richness and composition of the subterranean terrestrial invertebrates in the Presidente Olegário karst area, southeastern Brazil. We carried out collections in six caves during both dry and wet seasons, using combined collection methods. We observed different distributions in relation to the substrate, because the environmental heterogeneity increases the amount of available niches for the fauna. Some taxa showed a preference for specific substrates, probably related to the availability of food resources and humidity and to body size restriction, emphasizing the niche differentiation between species. Anthropogenic impacts can cause irreversible alterations in the subterranean fauna because the subterranean environment is dependent on the surface for input of trophic resources. On-going impacts in the Presidente Olegario karst area, like agriculture, pastures, gas extraction, and hydroelectric projects, are therefore a serious threat to subterranean biodiversity and this region should be prioritized for conservation.

Taxonomic distinctness and conservation of a new high biodiversity subterranean area in Brazil.

Subterranean environments, even though they do not possess a primary production (photosynthesis), may present high biodiversity, faunistic originality, endemism, phylogenetic isolations and unique ecological and/or evolution events, in addition to rare taxa. Studies investigating the biological diversity in Neotropical caves are relatively rare and recent, and most of them have been conducted in Brazil. We sampled caves from the state of Bahia, northeastern Brazil, and through sampling sufficiency tests and richness estimators, we demonstrate that the normatization for the Brazilian cave laws is not adequate for its conservation and that only α diversity index is not enough to verify faunistic patterns. We suggest that a phylogenetic diversity index be more robust and accurate for conservation purposes, particularly the Taxonomic Distinctness index. Moreover, we propose that the sandstone complex caves from Chapada Diamantina National Park need to be classified as being of high subterranean biodiversity in a global scope.

First amphibious Crinocheta (Isopoda, Oniscidea) from the Neotropics with a troglobitic status: a relictual distribution.

The first freshwater amphibian representative of Crinocheta (Oniscidea) from the Neotropics is described from caves within the Brazilian Cerrado biome, state of Mato Grosso do Sul. Kadiweuoniscusrebellisgen. et sp. nov. is placed in the family Philosciidae. The present work represents a significant contribution to future studies seeking to understand the ecological and evolutionary processes of Crinocheta within the Neotropical region. Moreover, it highlights the importance of biodiversity surveys in subterranean environments toward effective conservation efforts of these unique habitats and their surroundings.

Tmesiphantes hypogeus sp. nov. (Araneae, Theraphosidae), the first troglobitic tarantula from Brazil.

A new species of Tmesiphantes Simon, 1892, is described from sandstone/quartizitic caves of Chapada Diamantina, Bahia State, Brazil. This is the fifth species of the genus and the first record of a troglobitic mygalomorph in Brazil. A key is presented for all Tmesiphantes species.

A New Species of Spelaeometra Polhemus & Ferreira, 2018 (Hemiptera: Heteroptera: Hydrometridae) from a Hotspot of Troglobites in Brazil, Serra do Ramalho Karst Area.

Spelaeometra Polhemus & Ferreira, 2018 (Hemiptera: Heteroptera: Hydrometridae) is a monotypic and troglobitic genus, recently described based on material collected in the state of Minas Gerais, Brazil. From specimens collected in the Gruna do Enfurnado Cave in the Serra do Ramalho karst area, municipality of Coribe, state of Bahia, Brazil, we describe Spelaeometra hypogea Cordeiro & Moreira, sp. nov. and compare it with Spelaeometra gruta Polhemus & Ferreira, 2018. The new species is diagnosed by the general body color being pale-yellow to light-brown; antennal segments I and II being subequal in length; the reduced gular lobe, not covering any labial segment; the male proctiger without apical projections; and the male paramere with convex dorsal margin after the curvature, with a narrow hook-like apex.

Coarazuphium bambui (Carabidae: Zuphiini), a new cave-dwelling beetle from the threatened region of Serra do Ramalho, Brazil.

To date, 20 species of obligate cave-dwelling beetles are known from Brazil, and 12 belong to the Coarazuphium genus. In this paper, we describe Coarazuphium bambui sp. n., from two specimens, one male and one female, collected in the Gruna Boca da Lapa cave located in the municipality of Feira da Mata (Bahia state, Brazil). The most striking difference between C. bambui and other species of the genus is the presence of three pairs of fixed setae posteriorly on the head, combined with the slightly elytral sinuosity. We also provide an updated key for species identification of the Coarazuphium genus. Finally, we highlight the importance of the Serra do Ramalho karst area for the conservation of troglobite diversity in the Brazilian context.

Expanding the knowledge on the diversity of the cavernicolous Styloniscidae Vandel, 1952 (Oniscidea, Synocheta) from Brazil, with descriptions of two new species from the semiarid karst regions.

Two new species of Pectenoniscus from two caves in karst areas of the Brazilian semiarid region are described. Pectenoniscuspankaru Campos-Filho, Torres & Bichuette, sp. nov. from Gruna do Govi cave, Serra do Ramalho karst area, state of Bahia, and Pectenoniscusfervens Campos-Filho, Taiti & Bichuette, sp. nov. from Toca Coroa do Frade cave, Barra Bonita karst area, state of Piauí. In addition, specimens of Cylindroniscusflaviae from Gruta da Tapagem (= Caverna do Diabo), Açungui karst area were also recorded. An updated diagnosis of Pectenoniscus and a distribution map of the species examined herein are given.

Nutrient availability in tropical caves influences the dynamics of microbial biomass.

Few studies have evaluated the trophic level in tropical caves, and none related the microbial biomass dynamics in the immobilization of carbon and nitrogen. Here, four tropical caves of Terra Ronca State Park, Brazil, were studied: Angélica, São Bernardo, Terra Ronca I, and Terra Ronca II caves. Physical, chemical, and microbiological parameters (microbial biomass and respiration) were estimated in the dry and wet seasons. São Bernardo, Terra Ronca I, and Terra Ronca II caves presented higher nitrogen and microbial biomass nitrogen (MBN) values in the wet season than in the dry season. On the other hand, the Angélica cave showed larger amounts of nitrogen and lower MBN values in the dry season. These results indicate that caves can be adjusted in two ecological theories known as "stoichiometric decomposition" and "microbial nitrogen mining"-to the effects of nutrient availability on organic matter decomposition. The caves studied showed different environmental dynamics in relation to organic matter decomposition, which allows them to be considered unique and possess specific characteristics. Microbial biomass dynamics can be an important parameter to evaluate the availability of nutrients and ecological dynamics of the trophic network in subterranean environments.

Spooky Interaction at a Distance in Cave and Surface Dwelling Electric Fishes.

Glass knifefish (Eigenmannia) are a group of weakly electric fishes found throughout the Amazon basin. Their electric organ discharges (EODs) are energetically costly adaptations used in social communication and for localizing conspecifics and other objects including prey at night and in turbid water. Interestingly, a troglobitic population of blind cavefish Eigenmannia vicentespelea survives in complete darkness in a cave system in central Brazil. We examined the effects of troglobitic conditions, which includes a complete loss of visual cues and potentially reduced food sources, by comparing the behavior and movement of freely behaving cavefish to a nearby epigean (surface) population (Eigenmannia trilineata). We found that the strengths of electric discharges in cavefish were greater than in surface fish, which may result from increased reliance on electrosensory perception, larger size, and sufficient food resources. Surface fish were recorded while feeding at night and did not show evidence of territoriality, whereas cavefish appeared to maintain territories. Surprisingly, we routinely found both surface and cavefish with sustained differences in EOD frequencies that were below 10 Hz despite being within close proximity of about 50 cm. A half century of analysis of electrosocial interactions in laboratory tanks suggest that these small differences in EOD frequencies should have triggered the "jamming avoidance response," a behavior in which fish change their EOD frequencies to increase the difference between individuals. Pairs of fish also showed significant interactions between EOD frequencies and relative movements at large distances, over 1.5 m, and at high differences in frequencies, often >50 Hz. These interactions are likely "envelope" responses in which fish alter their EOD frequency in relation to higher order features, specifically changes in the depth of modulation, of electrosocial signals.

A new centipede Schendylops Cook from eastern Brazil: the first troglobitic geophilomorph for South America (Geophilomorpha, Schendylidae).

Schendylops janelao sp. n. is described from a limestone cave located in eastern Brazil. There are several reports of geophilomorphs in Brazilian caves, however, the new species represents the first troglobite species for South America and the first for Gruta do Janelão Cave, northern Minas Gerais State. Schendylops janelao sp. n. is described based on a male specimen and differs from its congeners by a combination of 10 characters. We discuss the character-states related to subterranean life and the conservation status of the species.

Two new species and new records of terrestrial isopods (Crustacea, Isopoda, Oniscidea) from Brazilian caves.

To date, approximately 190 species of terrestrial isopods are known from Brazil and only 14 are considered troglobiotic. After the examination of a large collection from caves in Bambuí and Una geomorphological areas, along the states of Bahia, Minas Gerais and Goiás, two new troglobiotic species were recognized. Pectenoniscus liliae Campos-Filho, Bichuette Taiti sp. n. (Styloniscidae) is described from Serra do Ramalho karst area, and Benthana xiquinhoi Campos-Filho, Bichuette Taiti sp. n. (Philosciidae) from sandstone caves of Chapada Diamantina region. The latter constitutes the second troglomorphic species of the genus. Xangoniscus aganju (Styloniscidae) is also recorded from two caves in the Serra do Ramalho karst area. The systematic position of Iuiuniscus iuiuensis is briefly discussed and Venezillo congener (Armadillidae) is redescribed from São Domingos karst area.

A synopsis of centipedes in Brazilian caves: hidden species diversity that needs conservation (Myriapoda, Chilopoda).

This study revises centipede fauna found in Brazilian caves, focusing on troglomorphic taxa and emphasizing conservation status. We present 563 centipede specimens from 274 caves across eleven Brazilian states. Of these, 22 records were derived from existing literature and 252 are newly collected. Specimens represent four orders, ten families, 18 genera, and 47 morphospecies. Together, the cave records represent 21 % of Brazil's centipede fauna. Scolopendromorpha was the most representative order (41 %), followed by Geophilomorpha (26 %), Scutigeromorpha (23 %), and Lithobiomorpha (10 %). Six species were found only in caves, with four considered troglobitic. The distribution of Cryptops iporangensis, the first Brazilian troglobitic centipede species to be discovered, was expanded to other three caves. Cryptops spelaeoraptor and Cryptops iporangensis are two troglobitic species considered Vulnerable and Endangered, respectively, according to the IUCN Red List. Main threats to Brazilian caves are mining, hydroelectric projects, water pollution, and unregulated tourism.

Brazilian obligatory subterranean fauna and threats to the hypogean environment.

The subterranean environment harbors species that are not capable of establishing populations in the epigean environment, i.e., the obligatory subterranean species. These organisms live in a unique selective regime in permanent darkness and usually low food availability, high air humidity in terrestrial habitats, and low temperature range allied to other unique conditions related to lithologies and past climatic influences. The pressure to increase Brazil's economic growth relies on agricultural/pastoral industries and exporting of raw materials such as iron, limestone, ethanol, soybean, cotton, and meat, as well as huge reservoir constructions to generate electricity. Mining (even on a small scale), agricultural expansion, and hydroelectric projects are extremely harmful to subterranean biodiversity, via the modification and even destruction of hypogean habitats. The Brazilian subterranean species were analyzed with respect to their distributions, presence on the IUCN Red List, and current and potential threats to hypogean habitats. A map and three lists are presented, one with the described obligatory subterranean species, one with undescribed taxa, and one with the current and potential threats to the hypogean environment. To date, 150 obligatory subterranean species have been recorded in Brazil, plus at least 156 undescribed troglomorphic taxa, totaling 306 Brazilian troglobites/obligatory cave fauna. We also analyzed the current and potential cave threats and the conservation actions that are underway to attempt to compensate for loss of these habitats. In according to the Brazilian legislation (Decree 6640) only caves of maximum relevance are fully protected. One strategy to protect the subterranean fauna of Brazil is the inclusion of these species in the IUCN Red List (one of attributes that determines maximum relevance for caves); however, one of the IUCN assumptions is that the taxa must be formally described. It is clear that the description and proposed protection of Brazilian subterranean biodiversity depends on more systematics studies.

Cylindroniscus platoi (Isopoda: Oniscidea: Styloniscidae), a new cave-dwelling species from Lagoa Santa Karst, Southeastern Brazil.

The new species Cylindroniscus platoi n. sp. is herein described. As far as known, the species occurs only in three caves developed in a continuous limestone outcrop at Pedro Leopoldo municipality, Minas Gerais State. Cylindroniscus platoi n. sp. is regarded as an endemic troglobite based on its limited distribution and on non-obvious troglomorphisms found on its body surface.

A new species of cave dwelling Neocarus (Acari: Opilioacaridae) from Bahia state, Brazil, with remarks on taxonomic characters.

Neocarus coronatus n. sp., is described from caves and the surrounding epigean habitats from a karst area of São Desidério county, Bahia state, Brazil. A dissection procedure is presented for the better use of scarce Opilioacaridae material, especially to analyze gut contents. Sexual dimorphism is presented, mainly differences in the density of setae on the dorsal shield in the prominent triangular anterior region. Females have an ovipositor with 3 pairs of smooth digitiform projections and 6-8 eugenital setae. Adults have a palpal tibiotarsus with 4 d setae. Using Low-Temperature Scanning Electron Microscopy (LT-SEM), evaluation of the chaetotaxy (especially setal shape) and morphology are greatly improved over standard light microscopy techniques enhancing the interpretation of characters for species differentiation. Also, we discuss the fragility of microinvertebrates in cave habitats.

Everything is similar, everything is different! Trichorhina (Oniscidea, Platyarthridae) from Brazilian caves, with descriptions of 11 new species / Tudo é parecido, tudo é diferente! Trichorhina (Oniscidea, Platyarthridae) de cavernas brasileiras, com descrições de 11 novas espécies

Abstract The present work describes 11 new species of the genus Trichorhina from Brazilian caves in the states of Bahia, Mato Grosso do Sul, Minas Gerais, Pará, and São Paulo. Moreover, the distribution of T. tomentosa has been expanded to include the states of Mato Grosso and Minas Gerais. Additionally, a distribution map and a key for all Brazilian species of Trichorhina, including epigean ones, are provided.

Resumo O presente trabalho descreve 11 novas espécies do gênero Trichorhina de cavernas brasileiras nos estados da Bahia, Mato Grosso do Sul, Minas Gerais, Pará e São Paulo. Além disso, a distribuição de T. tomentosa foi ampliada para incluir os estados de Mato Grosso e Minas Gerais. Adicionalmente, são fornecidos um mapa de distribuição e uma chave para todas as espécies brasileiras de Trichorhina, incluindo as epígeas.

The Laterosensory Canal System in Epigean and Subterranean Ituglanis (Siluriformes: Trichomycteridae), With Comments About Troglomorphism and the Phylogeny of the Genus.

The laterosensory system is a mechanosensory modality involved in many aspects of fish biology and behavior. Laterosensory perception may be crucial for individual survival, especially in habitats where other sensory modalities are generally useless, such as the permanently aphotic subterranean environment. In the present study, we describe the laterosensory canal system of epigean and subterranean species of the genus Ituglanis (Siluriformes: Trichomycteridae). With seven independent colonizations of the subterranean environment in a limited geographical range coupled with a high diversity of epigean forms, the genus is an excellent model for the study of morphological specialization to hypogean life. The comparison between epigean and subterranean species reveals a trend toward reduction of the laterosensory canal system in the subterranean species, coupled with higher intraspecific variability and asymmetry. This trend is mirrored in other subterranean fishes and in species living in different confined spaces, like the interstitial environment. Therefore, we propose that the reduction of the laterosensory canal system should be regarded as a troglomorphic (= cave-related) character for subterranean fishes. We also comment about the patterns of the laterosensory canal system in trichomycterids and use the diversity of this system among species of Ituglanis to infer phylogenetic relationships within the genus. J. Morphol. 278:4-28, 2017. ©© 2016 Wiley Periodicals,Inc.