Phylogeny of Lithobiidae Newport, 1844, with emphasis on the megadiverse genus Lithobius Leach, 1814 (Myriapoda, Chilopoda).

Phylogenetic analyses based on molecular and morphological data were conducted to shed light on relationships within the mostly Palaearctic/Oriental centipede family Lithobiidae, with a particular focus on the Palaearctic genus Lithobius Leach, 1814 (Lithobiidae, Lithobiomorpha), which contains >500 species and subspecies. Previous studies based on morphological data resolved Lithobius as nonmonophyletic, but molecular-based phylogenetic analyses have until now sampled few species. To elucidate species inter-relationships of the genus, test the validity of its classification into subgenera, and infer its relationships with other Lithobiidae, we obtained molecular data (nuclear markers: 18S rRNA, 28S rRNA; mitochondrial markers: 16S rRNA, COI) and 61 morphological characters for 44 species of Lithobius representing four of its eight subgenera and nine other representatives of Lithobiidae. The data were analyzed phylogenetically using maximum-likelihood, parsimony and Bayesian inference. This study suggests that (i) a close relationship between L. giganteus and the pterygotergine Disphaerobius loricatus highlighted in recent morphological analyses is also strongly supported by molecular data, and Pterygoterginae is formally synonymized with Lithobiinae; (ii) the Oriental/Australian genus Australobius is consistently resolved as sister group to all other sampled Lithobiidae by the molecular and combined data; (iii) the subfamily Ethopolyinae may be paraphyletic; (iv) the genus Lithobius is nonmonophyletic; (v) the subgenera Lithobius, Sigibius and Monotarsobius are nonmonophyletic and should not be used in future taxonomic studies; and (vi) there are instances of cryptic species and cases in which subspecies should be elevated to full species status, as identified for some European taxa within Lithobius.

Local- versus broad-scale environmental drivers of continental ß-diversity patterns in subterranean spider communities across Europe.

Macroecologists seek to identify drivers of community turnover (ß-diversity) through broad spatial scales. However, the influence of local habitat features in driving broad-scale ß-diversity patterns remains largely untested, owing to the objective challenges of associating local-scale variables to continental-framed datasets. We examined the relative contribution of local- versus broad-scale drivers of continental ß-diversity patterns, using a uniquely suited dataset of cave-dwelling spider communities across Europe (35-70° latitude). Generalized dissimilarity modelling showed that geographical distance, mean annual temperature and size of the karst area in which caves occurred drove most of ß-diversity, with differential contributions of each factor according to the level of subterranean specialization. Highly specialized communities were mostly influenced by geographical distance, while less specialized communities were mostly driven by mean annual temperature. Conversely, local-scale habitat features turned out to be meaningless predictors of community change, which emphasizes the idea of caves as the human accessible fraction of the extended network of fissures that more properly represents the elective habitat of the subterranean fauna. To the extent that the effect of local features turned to be inconspicuous, caves emerge as experimental model systems in which to study broad biological patterns without the confounding effect of local habitat features.

Investigation on the true identity of Entomobryanigriventris Stach, 1929 (Collembola, Entomobryidae) with the description of a new species.

The present paper gives a detailed and illustrated redescription of Entomobryanigriventris Stach, 1929, and the description of a new species collected from open sand steppe habitat in Hungary. Based on the colour pattern, E.arenaria Winkler, Flórián & Dányi, sp. nov. is close to E.violaceolineata Stach, 1963 but differs from it by the morphology of the labral papillae and the dorsal macrochaetotaxy of the head, Th II, and Abd II-IV. The new species is also characterised by dark ventral body colouration in adult specimens. In this regard, an overview of European Entomobrya species in which the dark ventral side may occur is also provided.

Editorial.

Editorial.

Taxonomic status and behavioural documentation of the troglobiont Lithobiusmatulici (Myriapoda, Chilopoda) from the Dinaric Alps: Are there semiaquatic centipedes in caves?

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.

Effects of single and repeated drought on soil microarthropods in a semi-arid ecosystem depend more on timing and duration than drought severity.

Soil moisture is one of the most important factors affecting soil biota. In arid and semi-arid ecosystems, soil mesofauna is adapted to temporary drought events, but, until now, we have had a limited understanding of the impacts of the different magnitudes and frequencies of drought predicted to occur according to future climate change scenarios. The present study focuses on how springtails and mites respond to simulated repeated drought events of different magnitudes in a field experiment in a Hungarian semi-arid sand steppe. Changes in soil arthropod activities were monitored with soil trapping over two years in a sandy soil. In the first year (2014), we applied an extreme drought pretreatment, and in the consecutive year, we applied less devastating treatments (severe drought, moderate drought, water addition) to these sites. In the first year, the extreme drought pretreatment tended to have a negative effect (either significantly or not significantly) on the capture of all Collembola groups, whereas all mite groups increased in activity density. However, in the consecutive year, between the extreme drought and control treatments, we only detected differences in soil microbial biomass. In the cases of severe drought, moderate drought and water addition, we did not find considerable changes across the microarthropods, except in the case of epedaphic Collembola. In the cases of the water addition and drought treatments, the duration and timing of the manipulation seemed to be more important for soil mesofauna than their severity (i.e., the level of soil moisture decrease). We suggest that in these extreme habitats, soil mesofauna are able to survive extreme conditions, and their populations recover rapidly, but they may not be able to cope with very long drought periods.

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.

Catalogue of the type material of pseudoscorpions (Arachnida: Pseudoscorpiones) deposited in the Hungarian Natural History Museum, Budapest.

A catalogue of the pseudoscorpion types deposited in the Hungarian Natural History Museum (Budapest) is presented. The Collection of Soil Invertebrates at the Museum harbours type material of 60 described (49 valid) pseudoscorpion species or subspecies, belonging to 13 families and 33 genera. Most of these are types of species described by Ödön Tömösváry, Jeno Daday, Max Beier and Volker Mahnert, along with a handful described by Lodovico di Caporiacco, Mark Harvey and János Novák. Lists are also given of types of species that were, or might be expected to have been, deposited in HNHM but cannot now be located there. A lectotype is designated for Chelifer rutilans Tömösváry, 1883, which is considered a junior subjective synonym of Chelifer cancroides (Linnaeus, 1758) (syn. nov.).

Life history characteristics of a cave isopod (Mesoniscusgraniger Friv.).

The special environmental conditions of caves provide habitat for several endemic and relict species, among them terrestrial isopods. The Baradla Cave system (north-eastern Hungary) hosts Mesoniscusgraniger (Frivaldszky, 1865) (Oniscidea, Microcheta, Mesoniscidae), a pygmy, blind, fragile troglophile woodlice species. Its stable environment can be characterised by the lack of light, high relative humidity (96%), low and constant temperature (about 10 °C). We explored the population characteristics (sex ratio, size distribution) and life history traits of the species (e.g. longevity, reproductive strategy, offspring number, and size). Sex ratio and size distribution of the individuals (head-width measurements) were estimated based on a yearly pooled pitfall-trap data set (N = 677). We studied the species' reproductive strategy under natural conditions (Baradla Cave, Aggtelek National Park). Model populations were set up in the cave and checked monthly between March and October, 2016 (15 replicates, each with 12 randomly chosen adult individuals; ΣN = 180). Digital photos were taken of the live animals and their length was estimated based on the photos by using ImageJ software (average body length: 6.56 ± 0.79 mm). The results showed female dominance in the population [(male:female = 0.43:0.57); p < 0.001 (GLM)]. Female head width (0.87 ± 0.18 mm) was significantly greater than that of males [0.79 ± 0.08 mm; p < 0.001 (t-test)]. Based on our present data we assume that the offspring number per single female is low (3-5), and new-borns have a relatively large size (body length: 4.22 ± 0.53 mm) compared to the adults. The probability of reproduction was continuous by monthly intervals (binomial test) and longevity exceeds one year. Our results suggest that the species follows a stenodynamic life history.

Contribution to the knowledge of the earthworm fauna of Turkey with description of three new species (Clitellata: Megadrili).

During a recent survey, 29 earthworm species were collected from different parts of Turkey, 27 of which belong to the family Lumbricidae, one to Acanthodrilidae and one to Megascolecidae. Dendrobaena proved to be the most speciose genus with 12 recorded species including two species new to science, Dendrobaena pavliceki and Dendrobaena taurica spp. nov. In addition a third new species was also found belonging to the East Mediterranean genus Healyella: Healyella zicsii sp. nov. From the species recorded earlier for Turkey, D. fridericae uludagi Omodeo Rota, 1991 and He. boluana Omodeo Rota, 1989 were found for the first time since the original description. With these new records the number of earthworm species recorded for Turkey is raised to 84.

On the identity of Photophilus Folkmanová, 1928: A new generic synonymy (Chilopoda: Geophilomorpha: Geophilidae).

The enigmatic monotypic genus Photophilus Folkmanová, 1928 and its type species, Photophilus griseus Folkmanová, 1928, were described almost 90 years ago from the Czech Republic. The new genus was erected not only based on some morphological characters, but also on behavioural features rather peculiar among Central European geophilomorphs. Although the patria of these taxa is among the best studied regions regarding Chilopoda, and focussed effort has been made to collect further representatives of these taxa, none have been found since the original description As the type material is lost and we failed to collect new specimens from the type locality, we tried to elucidate the true identity of these genus and species by analysing the characters given in the original description in comparison to all other geophilomorphs known from Europe. The structure of the mouth-parts, the shape of the ventral pore-fields and the arrangement of the coxal pores indicate Photophilus to be in the closest relation with the genus Geophilus Leach, 1814 from which it is separated by characters which are shown here as artefacts or misinterpretations according to the judgement of the original descriptions illustrations. Considering the remaining characters of Photophilus and its type species-including the behavioural features-they actually fit only to the widespread European species Geophilus carpophagus Leach, 1815, which is formerly proposed here to be the subjective senior synonym of Photophilus griseus Folkmanová, 1928 (syn. nov.). Photophilus is formerly proposed here as subjective junior synonym of Geophilus (syn. nov.).

An annotated checklist of earthworms of Greece (Clitellata: Megadrili).

The earthworm fauna of Greece is reviewed. According to the up-to-date checklist, the earthworm fauna of Greece consists of 67 species and subspecies, of which 59 taxa belong to the family Lumbricidae, three to Megascolecidae, two to Acanthodrilidae and to Ocnerodrilidae and one taxon to the family Criodrilidae. Three species are recorded for the first time from the country: Allolobophora kosowensis kosowensis Karaman, 1968, Amynthas gracilis (Kinberg, 1867) and Eukerria saltensis (Beddard, 1895). Eisenia spelaea var. athenica Cernosvitov, 1938 is proposed to be a synonym of Aporrectodea rosea (Savigny, 1826). The earthworm fauna of Greece is characterized by a large number of strict endemic species belonging to the family Lumbricidae (9 taxa), together with the occurrence of another 10 Balkanic endemic species.

Out of Africa: The first introduced African geophilomorph centipede record from a European greenhouse (Chilopoda: Geophilidae).

In Europe, 184 species of Geophilomorpha are recognised (Bonato & Minelli 2014) of which eight are evaluated as alien to the region (Stoev et al. 2010, Decker et al. 2014). Four of these have been reported from greenhouses exclusively: Mecistocephalus guildingii Newport, 1843, Mecistocephalus maxillaris (Gervais, 1837), Tygarrup javanicus Attems, 1929 and Pectiniunguis pauperatus Silvestri, 1907. In this paper, we report another species, Polygonarea silvicola Lawrence, 1955, which is the first Geophilomorpha species of unambiguous African origin in Europe. Description of the specimen found in Olomouc (Czech Republic) is provided. Co-occurrence of another African species, the lithobiomorph Lamyctes africanus (Porath, 1871) (also new for Czech Republic) is also reported here.

A new species of Janusius Bretfeld, 2010 (Collembola, Sminthuridae) from the Korean Peninsula, with a key to the species of the genus.

Janusius geumgangensis sp. nov. is described and illustrated based on specimens from North Korea. The new species is characterised by having 3 spatulate chaetae on all tibiotarsus, 4 chaetae on retinaculum, smooth anal appendage, anterior chaetotaxy of dens 3,3,3,2,2,1,1, colouration with one pair brownish lateral spot and presence of mucronal chaeta. The genus Janusius Bretfeld, 2010 is first mentioned from the Korean Peninsula, although another species described from there as Sminthurus annulatus Kang & Lee, 2005 is recognised here as belonging to this genus (Janusius annulatus (Kang & Lee, 2005) comb. nov.), too. A key including all known species of the genus is presented as well.

Description of the male and the larva of Ixodes ariadnae Hornok, 2014.

Ixodes ariadnae is a tick species of bats so far reported only in Central Europe, with its description based on the female and nymph. This study describes the male and larva in order to complete the description of the species. Male ticks collected from cave walls in Hungary showed a different morphology from those of I. vespertilionis and I. simplex. Molecular analysis of the cytochrome oxidase subunit I (COI) gene of these ticks verified them as conspecific to I. ariadnae. In addition, a larva of I. ariadnae was removed from a Daubenton's bat (Myotis daubentonii Kuhl, 1817). The male of I. ariadnae is characterized by long legs (7-8mm; I. vespertilionis: 8-10mm; I. simplex: 2-2.2mm), relatively short palpal setae (30-100µm; I. vespertilionis: 100-200µm; I. simplex: 20-50µm) and straight lateral edge of palps, the genital aperture (enclosed by a line of sclerotization) situated anteriorly to second intercoxal space and rounded coxae. The larva of I. ariadnae has long legs (2-2.2mm; I. vespertilionis: 1.6-1.8mm; I. simplex: 1mm), broad palps (length×width: 200×90µm; I. vespertilionis: 200×70µm; I. simplex: 140×60µm), pentagonal and posteriorly reverse bell-shaped scutum. These features allow to distinguish the male and the larva of I. ariadnae from those of I. vespertilionis (of which the male has longer palpal setae and curved lateral edge of palps, the genital aperture is situated posterior to the second intercoxal space, and the second coxae are squared; the larva of I. vespertilionis has narrower palps and posteriorly triangular scutum) and I. simplex (of which the male and the larva have considerably shorter legs, palps).

True identity of Folkmanovius paralellus Dobroruka (Chilopoda: Geophilomorpha: Geophilidae).

The mysterious monotypic genus Folkmanovius Dobroruka, 1957, and its type species, Folkmanovius paralellus Dobroruka, 1957, have never been found again since their description, although the patria of these taxa-the Czech Republic-is among the best studied regions regarding Chilopoda. Analysing the characters given in the original description, we found Folkmanovius to be in the closest relation with another European genus, Clinopodes C.L. Koch, 1847. Some characters could fit with the only Central European representative of this latter genus (Clinopodes flavidus) if, and only if, we consider that the description was based on juvenile specimens. Studying C. flavidus specimens of the first adolescens stadia, we found morphological changes during postembryonic development for several characters, documented first time for the genus. These results explain and elucidate all the tentative differential characters of Dobroruka's taxa and prove that Folkmanovius Dobroruka, 1957, is a junior subjective synonym of Clinopodes C.L. Koch, 1847, (syn. nov.), as well as Folkmanovius paralellus Dobroruka, 1957, being a junior subjective synonym of Clinopodes flavidus C.L. Koch, 1847 (syn. nov.).