Integrative taxonomic revision of the genera Nesticella and Howaia in Japan with the description of five new species (Araneae, Nesticidae, Nesticellini).

The Japanese species of the genera Nesticella Lehtinen & Saaristo, 1980 and Howaia Lehtinen & Saaristo, 1980 (Araneae, Nesticidae, Nesticellini) are revised using an integrative taxonomic approach. Their morphology, phylogenetic position within the genera, assignment to species groups, and distribution in mainland Japan and the Ryukyu islands are discussed herein. A phylogenetic and species delimitation analysis was conducted to confirm the boundaries between the putative species that were initially discriminated based on external and genital morphology. As a result of the present study, six species of Nesticella and three species of Howaia are proposed for the fauna of Japan based on the combined results of morphology and molecular analyses. Male and female of the previously known species H.mogera (Yaginuma, 1972), N.brevipes (Yaginuma, 1970), and N.okinawaensis (Yaginuma, 1979) are redescribed and illustrated using type specimens or specimens collected at the type locality. Nesticellaterrestris (Yaginuma, 1970) is resurrected as a valid species and distinguished from the closely related N.brevipes based on morphological and molecular evidence. The male of N.terrestris is described for the first time. We herein describe three new species of Nesticella and two new species of Howaia from different islands in the Ryukyu Archipelago, southwest Japan: N.insulanasp. nov. (♂, ♀) from Yonaguni-jima Island, N.occultasp. nov. (♀) from Ishigaki-jima Island, N.silvicolasp. nov. (♂, ♀) from Yakushima Island, H.albasp. nov. (♂, ♀) from Miyako-jima Island and H.subterraneasp. nov. (♀) from Okinoerabu-jima Island. Nesticellaoccultasp. nov., H.albasp. nov. and H.subterraneasp. nov. dwell exclusively in caves and show extensive morphological adaptation to subterranean life.

Cave-adapted millipedes from Portugal: species conservation profiles.

Background: Amongst the cave-dwelling millipedes (Diplopoda), there are several endemic species in Portugal with a very small geographical distribution. These species play an important role in the decomposition of organic matter in subterranean ecosystems and are vulnerable to disturbance from human activities, such as habitat destruction, pollution infiltrating from the surface and cave tourism. New information: We present the IUCN Red List profiles for cave-adapted millipedes (Diplopoda) from Portugal and propose conservation measures to prevent extinction. Overall, cave-adapted millipedes from Portugal represent an endemic part of the country's biodiversity and conservation efforts will help maintain the delicate ecological balance of subterranean ecosystems.

Species conservation profiles of cave-adapted terrestrial isopods from Portugal.

Background: Terrestrial isopods (Oniscidea) are the most diverse group of troglobionts in caves of continental Portugal. They occur in all karst regions of Portugal, play a major role in decomposition of organic matter in caves and may act as umbrella species for the conservation of all other cave-adapted invertebrates. New information: We present the IUCN Red List profiles for the cave-adapted terrestrial isopods from continental Portugal, based on recent distribution data from caves.

Two new cave Hyleoglomeris species (Glomerida, Glomeridae) from northern Vietnam.

Two new glomerid species from caves in Cao Bang Province, Northern Vietnam, namely, Hyleoglomerishalang Kuroda, Nguyen & Eguchi, sp. nov. and Hyleoglomerisalba Nguyen, Kuroda & Eguchi, sp. nov., are described. The former is characterized by a distinct body color pattern; telopods with a large, quadrate, medially concave, sparsely setose, central syncoxital lobe; and syncoxital horns approximately 1.5-2.0 times as long as the lobe. The latter is distinguished by its completely troglobiotic form without eyes, an unpigmented body, and a roundly triangular syncoxital lobe of telopods. An identification key is also provided for the cave glomerids of Vietnam.

Species conservation profiles of the endemic spiders Troglohyphantes (Araneae, Linyphiidae) from the Alps and the north-western Dinarides.

Background: The genus Troglohyphantes Joseph, 1882 (Araneae, Linyphiidae) includes 131 species, mainly distributed across the main European mountain ranges. The Alps and the north-western Dinarides account for 66 species, most of them showing narrow or even point-like distributions. The majority of Troglohyphantes spiders dwell in subterranean habitats including caves, mines, soil litter, rocky debris and other moist and shaded retreats. Despite being intensively studied from taxonomic, ecological and biogeographic standpoints, knowledge on the status of conservation and on the potential risk of extinction of these spiders is lagging. To date, only three species have been included in the global IUCN Red List, but their status has not been updated ever since their last assessment in 1996. The aim of this contribution is to assess the Alpine and north-western Dinaric species of the genus Troglohyphantes and to re-assess the species previously evaluated, according to the last version of the IUCN Red List Categories and Criteria. New information: Amongst the 66 species here considered, 62 had sufficient data to allow the quantification of their Extent Of Occurrence (EOO) and Area Of Occupancy (AOO). Most of the species have a narrow distribution range, with an estimated EOO < 20,000 km2 and AOO < 2,000 km2, meeting the thresholds for the inclusion in the threatened categories. Five species have a more widespread distribution (EOO > 20,000 km2), extending across multiple countries. The quality of the data on distribution of four species was not sufficient to provide a reliable estimation of the distribution range.A continuing decline in EOO, AOO and habitat quality was inferred for 30 species. The majority of them were subterranean specialised species, with a reduced thermal tolerance and a low dispersal ability. Accordingly, changes in subterranean microclimatic conditions due to climate change represent a major threat for these species. Land-use change and habitat alteration were identified as additional relevant threats for several species.A considerable proportion of the species here assessed was found in protected areas and in sites of the Natura 2000 network. In addition, 14 species are formally protected by national and sub-national legislation. At present, 25 species are listed in the regional Red Lists.Long-term monitoring programmes, management plans for both the species and their habitats, expansion of the extant protected areas and designation of new ones, should be considered as the most effective approaches to species conservation.

Cave-inhabiting Cheliferidae (Arachnida, Pseudoscorpiones) from Thailand, with description of four new species of Metachelifer Redikorzev.

Four new species of the genus Metachelifer Redikorzev, 1938 are described from caves in the provinces of Tak (M.takensis sp. nov. and M.thailandicus sp. nov.), Chiangmai (M.mahnerti sp. nov.), and Nakhon Ratchasima (M.cheni sp. nov.). An identification key is provided to all known world representatives of the genus Metachelifer.

Cave-adapted beetles from continental Portugal.

BACKGROUND: The cave biodiversity of continental Portugal faces tremendous conservation challenges, mostly linked to their direct destruction and contamination infiltrating from the surface. Beetles are the most diverse insects and one of the most diverse arthropod groups in caves of Portugal. NEW INFORMATION: We present the IUCN Red List profiles for the cave-adapted beetles from continental Portugal, all endemic to their respective geological units and massifs. Ground beetles (Carabidae) are the most diverse family of cave-adapted beetles in continental Portugal, followed by rove beetles (Staphylinidae). Beetles in caves of Portugal are mostly terrestrial and only one species is known to have evolved to live in groundwater. Trechus is the most diverse genus with four species, followed by Domene with three species and by Speonemadus and Iberoporus, both with one cave-adapted species. The aim of this contribution is to assess all endemic cave-adapted species of beetles from continental Portugal and to support their specific protection, to promote adequate management of surface habitats and the establishment of priority areas for conservation. The main biodiversity erosion drivers that are impacting the conservation of the studied species are pollution infiltrating from the surface, urbaniation, modifications of the natural habitat for touristic purposes and mining, quarrying and energy production infrastructures.This document can be used in spatial planning and territory management in karst, based on the current scientific knowledge.

A rich fauna of subterranean short-range endemic Anillini (Coleoptera, Carabidae, Trechinae) from semi-arid regions of Western Australia.

Globally, the great majority of Anillini species are endogean, adapted to live in the interstices of soil and leaf litter, while the extremely low vagility of these minute ground beetles gives rise to numerous shortrange endemic species. Until recently the Australian Anillini fauna was known only from leaf litter in rain forests and eucalypt forests in the wetter, forested regions of eastern and south eastern Australia, as well as Lord Howe and Norfolk islands. The first hypogean Anillini in Australia (17 species in six genera) were described in 2016 from mineral exploration drill holes in iron-ore bearing rocks of the Pilbara region in Western Australia, representing the first finding of the tribe deep underground in a semi-arid climate region. A further eight new genera and 20 new species are described herein, mostly from the Pilbara region as well as the semi-arid Kimberley and Goldfields regions; all were collected in mineral exploration drill holes. The following new genera are described: Erwinanillus gen. nov., Gregorydytes gen. nov., Pilbaraphanus gen. nov., Neoillaphanus gen. nov., Kimberleytyphlus gen. nov., Gilesdytes gen. nov., Pilbaradytes gen. nov., and Bylibaraphanus gen. nov. The following new species are described: Erwinanillus baehri sp. nov.; Gracilanillus hirsutus sp. nov., G. pannawonicanus sp. nov.; Gregorydytes ophthalmianus sp. nov.; Pilbaraphanus chichesterianus sp. nov., P. bilybarianus sp. nov.; Magnanillus firetalianus sp. nov., M. sabae sp. nov., M. salomonis sp. nov., M. regalis sp. nov., M. serenitatis sp. nov.; Neoillaphanus callawanus sp. nov.; Kimberleytyphlus carrboydianus sp. nov.; Austranillus jinayrianus sp. nov.; Gilesdytes pardooanus sp. nov., G. ethelianus sp. nov.; Pilbaradytes abydosianus sp. nov., P. webberianus sp. nov.; Bylibaraphanus cundalinianus sp. nov.; and Angustanillus armatus sp. nov. Identification keys are provided for all Australian anilline genera, and Western Australian species. All the described species are known from a single locality and qualify as short-range endemics. The Anillini are recognised as a significant and diverse element making up part of Western Australia's remarkable subterranean fauna, and whose conservation may potentially be impacted by mining developments.

Spelaeobochica mahnerti, a new cave-dwelling pseudoscorpion from Brazil (Arachnida: Pseudoscorpiones: Bochicidae), with comments on the troglomorphism of the Brazilian bochicid species.

Spelaeobochica mahnerti sp. n., found in a limestone cave located in the municipality of Monjolos (Minas Gerais state, Southeastern Brazil), is described. It differs from congeneric species mainly by the rallum with 3 apically dentate blades, the position of the trichobothrium ib in the far distal half of the hand dorsum (near the base of fixed finger), the smooth subterminal setae, and by the arolia that are a little longer than the claws. A key to the species of Spelaeobochica is provided, as well as information on the habitat and threats for this new species. We further discuss troglomorphic traits in Brazilian species of Bochicidae.

Continental data on cave-dwelling spider communities across Europe (Arachnida: Araneae).

BACKGROUND: Spiders (Arachnida: Araneae) are widespread in subterranean ecosystems worldwide and represent an important component of subterranean trophic webs. Yet, global-scale diversity patterns of subterranean spiders are still mostly unknown. In the frame of the CAWEB project, a European joint network of cave arachnologists, we collected data on cave-dwelling spider communities across Europe in order to explore their continental diversity patterns. Two main datasets were compiled: one listing all subterranean spider species recorded in numerous subterranean localities across Europe and another with high resolution data about the subterranean habitat in which they were collected. From these two datasets, we further generated a third dataset with individual geo-referenced occurrence records for all these species. NEW INFORMATION: Data from 475 geo-referenced subterranean localities (caves, mines and other artificial subterranean sites, interstitial habitats) are herein made available. For each subterranean locality, information about the composition of the spider community is provided, along with local geomorphological and habitat features. Altogether, these communities account for > 300 unique taxonomic entities and 2,091 unique geo-referenced occurrence records, that are made available via the Global Biodiversity Information Facility (GBIF) (Mammola and Cardoso 2019). This dataset is unique in that it covers both a large geographic extent (from 35° south to 67° north) and contains high-resolution local data on geomorphological and habitat features. Given that this kind of high-resolution data are rarely associated with broad-scale datasets used in macroecology, this dataset has high potential for helping researchers in tackling a range of biogeographical and macroecological questions, not necessarily uniquely related to arachnology or subterranean biology.

New data of spiders (Arachnida, Araneae) of Cyprus. 1. Dysderidae found in caves.

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.

Amblypygids of Timor-Leste: first records of the order from the country with the description of a remarkable new species of Sarax (Arachnida, Amblypygi, Charinidae).

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.timorensis sp. 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.

Heat tolerance and acclimation capacity in subterranean arthropods living under common and stable thermal conditions.

Cave-dwelling ectotherms, which have evolved for millions of years under stable thermal conditions, could be expected to have adjusted their physiological limits to the narrow range of temperatures they experience and to be highly vulnerable to global warming. However, most of the few existing studies on thermal tolerance in subterranean invertebrates highlight that despite the fact that they show lower heat tolerance than most surface-dwelling species, their upper thermal limits are generally not adjusted to ambient temperature. The question remains to what extent this pattern is common across subterranean invertebrates. We studied basal heat tolerance and its plasticity in four species of distant arthropod groups (Coleoptera, Diplopoda, and Collembola) with different evolutionary histories but under similar selection pressures, as they have been exposed to the same constant environmental conditions for a long time. Adults were exposed at different temperatures for 1 week to determine upper lethal temperatures. Then, individuals from previous sublethal treatments were transferred to a higher temperature to determine acclimation capacity. Upper lethal temperatures of three of the studied species were similar to those reported for other subterranean species (between 20 and 25°C) and widely exceeded the cave temperature (13-14°C). The diplopod species showed the highest long-term heat tolerance detected so far for a troglobiont (i.e., obligate subterranean) species (median lethal temperature after 7 days exposure: 28°C) and a positive acclimation response. Our results agree with previous studies showing that heat tolerance in subterranean species is not determined by environmental conditions. Thus, subterranean species, even those living under similar climatic conditions, might be differently affected by global warming.

Testing the climatic variability hypothesis in edaphic and subterranean Collembola (Hexapoda).

The climatic variability hypothesis was applied to the thermal tolerance of edaphic and cave Collembola occupying contrasting environments. Collembola belonged to four categories - trogloxene, subtroglophile, eutroglophile and troglobiont - with a different degree of affinity to subterranean habitats. Altogether, specimens of 17 species were exposed to a one-hour laboratory survival test. The impact of temperature, species and species-temperature interaction on cold and heat survival was statistically significant. There was a decrease trend in cold and heat tolerance from trogloxenes, over subtroglophiles and eutroglophiles to troglobionts. It was shown that obligate cave species, restricted to climatic-stable cave conditions, retain a functional thermal resistance, i.e. the genetically determined ability to tolerate relatively broader temperature ranges. Our results outlined the direct relationship between the thermal tolerances of species and the size of their geographic distributions. It was also observed that cold resistance of Collembola decreased significantly with increasing species body length, indicating that body size plays an important role in temperature tolerances of arthropods inhabiting soil and subterranean habitats.

On the genus Systaria (Araneae: Clubionidae) in Southeast Asia: new species from caves and forests.

The genus Systaria Simon 1897 is reviewed in Southeast Asia and eight new species are described: S. lannops spec. nov. from Thailand (female), S. longinqua spec. nov. (male, female) and S. luangprabang spec. nov. (female) from Laos, S. procera spec. nov. (male, female) and S. bregibec spec. nov. (male) from Cambodia, S. bifidops spec. nov. from Malaysia (male), S. panay spec. nov. (female) and S. princesa spec. nov. (male, female) from Philippines. S. elberti (Strand 1913) is illustrated and partly re-described. Informal groups of species are proposed for the first time. Distribution ranges and habitat preferences are mapped.

First continental troglobiont Cylindroiulus millipede (Diplopoda, Julida, Julidae).

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.

Taxonomic additions of Embolemidae and Sclerogibbidae (Hymenoptera: Chrysidoidea) from Japan, with description of a new species of Trogloembolemus.

Ampulicomorpha thauma Rasnitsyn Matveev (Hymenoptera: Embolemidae) and Sclerogibba rossi Olmi (Hymenoptera: Sclerogibbidae) were newly recorded from Japan. Females of Embolemus walkeri Westwood were found in the soil together with nymphs of Cixiidae. Trogloembolemus okanoi sp. nov. is described and illustrated. The holotype ♀ was collected in the soil at a depth of one meter. A key to species of Japanese Embolemidae was updated.

Aphaenogaster gamagumayaa sp. nov.: the first troglobiotic ant from Japan (Hymenoptera: Formicidae: Myrmicinae).

Aphaenogaster gamagumayaa sp. nov., a new troglobiotic (true cave-dwelling) ant species, from a limestone cave on the island of Okinawa (Okinawa-jima), Ryukyu Archipelago, Japan is described. This is the first discovery of a troglobiotic ant in Japan and the second verified record worldwide. This species has only been found in a cave area with heavy guano deposits, and some worker ants were observed carrying guano. The evidence for categorizing this new species as troglobiont is discussed.

Opportunistic fungal pathogens isolated from a captive individual of the European blind cave salamander Proteus anguinus.

Proteus anguinus is a neotenic cave amphibian endemic to the Dinaric Karst and represents a symbol of Slovenian natural heritage. It is classified as 'Vulnerable' by the International Union for Conservation of Nature (IUCN) and is one of the EU priority species in need of strict protection. Due to inaccessibility of its natural underground habitat, scientific studies have been primarily conducted on Proteus in captivity where amphibians may be particularly susceptible to opportunistic microbial infections. In this case report, we present the results of an analysis of an individual that had been kept in captivity for 6 yr and then developed clinical symptoms, including ulcers, suggesting opportunistic microbial infection. Pigmented fungal hyphae and yeast-like cells were present in the dermis and in almost all other sampled tissues. Sampling of the ulcer allowed the isolation of a diverse array of bacterial and fungal species. We identified the water-borne, polymorphic black yeast Exophiala salmonis, an opportunistic pathogen of fish, as the cause of the primary infection. This is the first report on a fungal infection of Proteus and on cave salamanders in general.

Cherax acherontis (Decapoda: Parastacidae), the first cave crayfish from the Southern Hemisphere (Papua Province, Indonesia).

Cherax acherontis n. sp., is a crayfish endemic to the submerged river Yumugima in Hagepma/Jugurama cave in the New Guinea Highlands, Jayawijaya Regency, Papua Province, Indonesia. This species is the first cave crayfish from the Southern Hemisphere. The new species is most similar to Cherax monticola. Both species can be easily distinguished by certain morphological characteristics, which easily demonstrate C. acherontis n. sp. is a valid species.