Effects of Hydrologic Regime Changes on a Taxonomic and Functional Trait Structure of Earthworm Communities in Mountain Wetlands.

Disturbances, both natural and anthropogenic, influence the patterning of species and species traits. The shift in species composition and distribution pattern of functional traits can demonstrate if the community is resistant, sensitive or resilient to the disturbance. Based on species- and trait-based approaches, we examined the response of the earthworm community to changing hydrologic conditions caused by the artificial drainage of mountain fens, in which cumulative effects of disturbance events over space and time are much less dynamic than in riverine wetlands. We hypothesized that the drainage-related changes of mountain fen peat soils have an effect on the earthworm community composition and its functional structure. We assume that the shift in species composition and value of community-weighted functional traits reflect changes in the resilience or resistance of the earthworm community to environmental change. Our results demonstrate that the total density of earthworms was almost three times lower under drained conditions compared to natural ones. Artificial drainage of fens had a neutral effect on the species-based diversity indices. However, there were species-specific traits that responded to hydrologic changes and which led to the species' replacements and to the co-occurrence of eurytopic, surface-browsing and more drought- and low-pH-resistant earthworm species in the drained fens. Based on these results, we conclude that abiotic-based environmental filtering was the main process responsible for sorting earthworms according to species and traits in the disturbed hydrologic conditions. The greater earthworm functional trait variations in semi-natural hydrologic conditions emphasizes the impact of transient dynamics in an altered disturbance regime on the earthworm assembly. Results also showed that in the changing hydrologic conditions of mountain fens, the functional trait approach revealed only slightly more predictive power than the taxonomic one, but it proved better with processes responsible for earthworm species filtering.

Omnivorous Notoxus trinotatus Pic (Coleoptera: Anthicidae) is a newly recognized vector of northern leaf blight in maize.

The adaptations of omnivorous insects to food are manifested in a multifaceted manner, and the availability of food resources directly determines insect feeding tendencies, which contribute to a complex insect-food relationship and impact insect functionality in the environment. Stable isotope analysis was applied to test the feeding preference and further define the functional role of omnivorous beetles in cropland. Our results confirmed that as an omnivorous beetle, the fungivorous nature of Notoxus trinotatus accounted for a prominent proportion food selection at the adult stage, and more importantly, this dietary feature contributed to the dispersal of the northern corn leaf blight in maize (NLB) during the feeding trials. In addition to the preference for fungi, water supplementation was an essential element extending adult longevity, which directly prolonged the contact time of adults with pathogenic fungi in agricultural fields. Consistent with the herbivorous characteristics of beetles, before the emergence of NLB fungal pathogens, corn tissues served as the main food, which provided the beetles with more opportunities to transmit fungal pathogen propagules. We conclude that the role of N. trinotatus in carrying NLB pathogen is due to its feeding on this plant mycopathogen, and an increased abundance of beetles carrying the pathogen may increase the rate of NLB disease infestation. More focus should be concentrated on the functions of fungivorous beetles, not only as pathogen-transmitting pests, but also as an element among the balanced biotic factors in farmland.

Gut microbiome reflect adaptation of earthworms to cave and surface environments.

BACKGROUND: Caves are special natural laboratories for most biota and the cave communities are unique. Establishing population in cave is accompanied with modifications in adaptability for most animals. To date, little is known about the survival mechanisms of soil animals in cave environments, albeit they play vital roles in most terrestrial ecosystems. Here, we investigated whether and how gut microbes would contribute to the adaptation of earthworms by comparing the gut microbiome of two earthworm species from the surface and caves. RESULTS: Two dominant earthworm species inhabited caves, i.e., Allolobophora chlorotica and Aporrectodea rosea. Compared with the counterparts on the surface, A. rosea significantly decreased population in the cave, while A. chlorotica didn't change. Microbial taxonomic and phylogenetic diversities between the earthworm gut and soil environment were asynchronic with functional diversity, with functional gene diversity been always higher in earthworm gut than in soil, but species richness and phylogenetic diversity lower. In addition, earthworm gut microbiome were characterized by higher rrn operon numbers and lower network complexity than soil microbiota. CONCLUSIONS: Different fitness of the two earthworm species in cave is likely to coincide with gut microbiota, suggesting interactions between host and gut microbiome are essential for soil animals in adapting to new environments. The functional gene diversity provided by gut microbiome is more important than taxonomic or phylogenetic diversity in regulating host adaptability. A stable and high-efficient gut microbiome, including microbiota and metabolism genes, encoded potential functions required by the animal hosts during the processes of adapting to and establishing in the cave environments. Our study also demonstrates how the applications of microbial functional traits analysis may advance our understanding of animal-microbe interactions that may aid animals to survive in extreme ecosystems.

A new species of the genus Archiboreoiulus Brolemann, 1921 (Diplopoda, Julida) from Movile Cave (Southern Dobrogea, Romania).

Up to the present, the genus Archiboreoiulus included only two species: A. sollaudi from France and A. pallidus with a wide European distribution range. Here we describe a previously unknown species of the genus Archiboreoiulus, A. serbansarbui, collected from the mesothermal sulfurous Movile Cave, Mangalia, Romania, harboring a rich and diverse troglobitic community.

Subterranean biodiversity and depth distribution of myriapods in forested scree slopes of Central Europe.

The shallow underground of rock debris is a unique animal refuge. Nevertheless, the research of this habitat lags far behind the study of caves and soil, due to technical and time-consuming demands. Data on Myriapoda in scree habitat from eleven localities in seven different geomorphological units of the Czech and Slovak Republics were processed. Based on previous studies, as well as knowledge of cave and soil fauna, it was hypothesised that the occurrence of a varied and peculiar fauna would show a pattern of depth distribution with variations due to local specificities. From 2005-2016 (at least one year on each site), macrofauna was collected via sets of three long-term exposed subterranean traps consisting of 110 cm long perforated tube, with ten cups located in a gradient at 5-95 cm below the soil surface. In total, 14 symphylans (not identified to species level), 271 centipedes (23 spp.) and 572 millipedes (32 spp.) were sampled. The overall depth distribution of centipedes and millipedes appeared to have relatively similar pattern, with both groups being found at all depth levels. Nevertheless, this pattern depends on locations. The depth distribution trend lines are mostly in the form of an asymmetric 'U', with decreased abundance until the middle of the gradient, followed by increase in the deepest levels. Epigeic species were sporadically distributed along the whole depth gradient, but concentrated at the soil surface, while some subterranean species, such as the centipede Lithobius lucifugus and the millipedes Geoglomeris subterranea, Cibiniulus slovacus and Archiboreoiulus pallidus, were recorded in the deepest parts of the gradient. This characterises the debris community as a mixture of soil and subterranean species with an absence of species exclusively found in caves. The use of different fixation methods in traps had a significant and selective impact on samples; millipedes were either attracted by ethylene glycol or repelled by formaldehyde. Centipedes were also captured more frequently in ethylene glycol; however, the species composition varied in each of the fixatives. Depth distribution of myriapods was similar in both fixative solutions. Traps with these fixatives could be recommended for similar ecological studies.

Responses of soil microarthropod taxon (Hexapoda: Protura) to natural disturbances and management practices in forest-dominated subalpine lake catchment areas.

Disturbances are intrinsic drivers of structure and function in ecosystems, hence predicting their effects in forest ecosystems is essential for forest conservation and/or management practices. Yet, knowledge regarding belowground impacts of disturbance events still remains little understood and can greatly vary by taxonomic and functional identity, disturbance type and local environmental conditions. To address this gap in knowledge, we conducted a survey of soil-dwelling Protura, across forests subjected to different disturbance regimes (i.e. windstorms, insect pest outbreaks and clear-cut logging). We expected that the soil proturan assemblages would differ among disturbance regimes. We also hypothesized that these differences would be driven primarily by variation in soil physicochemical properties thus the impacts of forest disturbances would be indirect and related to changes in food resources. To verify that sampling included two geographically distant subalpine glacial lake catchments that differed in underlying geology, each having four different types of forest disturbance, i.e. control, bark beetle outbreak (BB), windthrow + BB (wind + BB) and clear-cut. As expected, forest disturbance had negative effects on proturan diversity and abundance, with multiple disturbances having the greatest impacts. However, differences in edaphic factors constituted a stronger driver of variability in distribution and abundance of proturans assemblages. These results imply that soil biogeochemistry and resource availability can have much stronger effects on proturan assemblages than forest disturbances.

Changes in the Body Size of Carabid Beetles Along Elevational Gradients: A Multispecies Study of Between- and Within-Population Variation.

Geographic variation in body size has fascinated biologists since the 19th century as it can provide insight into the evolution of the body size of various organisms. In this study, we investigated body size variation in eight carabid species/subspecies (Coleoptera: Carabidae) along elevational gradients in six Central European mountain ranges. First, we examined elevational variation in body size and whether female and male body sizes differed in their responses to elevation. Second, we examined intrapopulation variation in body size along an elevational gradient, and we compared the degrees of intrapopulation variation between males and females. The investigated species either followed a converse Bergmann's cline (Carabus auronitens auronitens Fabricius 1792; Carabus linnei Panzer 1810; Pterostichus melanarius (Illiger, 1798); Pterostichus pilosus (Host, 1789)) or their size was unaffected by elevation (Carabus auronitens escheri Palliardi, 1825; Carabus sylvestris sylvestris Panzer, 1796; Carabus sylvestris transsylvanicus Dejean, 1826; Pterostichus burmeisteri Heer, 1838). Females were the larger sex in all the investigated species, but the degree of sexual size dimorphism differed between species. In general, the degree of sexual size dimorphism showed no change with elevation. The degree of intrapopulation variation in body size slightly increased with elevation in C. sylvestris sylvestris and P. pilosus. Overall, the intrapopulation variation in body size significantly differed among the investigated carabid species. The existing literature on intrapopulation variation in the body size of insects is limited, but further investigation of this issue could provide a better understanding of the mechanisms that generate geographical clines.

The Siberian centipede species Lithobiusproximus Sseliwanoff, 1878 (Chilopoda, Lithobiomorpha): a new member of the Polish fauna.

The centipede Lithobiusproximus Sseliwanoff, 1878 is presented for the first time as a new member of the Polish fauna. This species, originally characterized as a widespread Siberian boreal species, seems to possess high plasticity with regards to environmental requirements. Its actual distribution range covers several geographical zones where local conditions have allowed it to survive. The present research in the Wigry National Park, northeast Poland, shows that its distribution extends to the ends of the East European Plain embracing the East Suwalki Lake District, where it occurs almost exclusively in the oak-hornbeam forests: in summer it is one of the three dominant lithobiomorph centipedes inhabiting litter layers.

Terrestrial isopods (Oniscidea) of the White Carpathians (Czech Republic and Slovakia).

This paper summarizes data regarding the terrestrial isopods of the White Carpathians range in the Western Outer Carpathians based on field research undertaken during the past several decades in natural meadow pasture and forest localities. Using a combination of four collection methods 19 species belonging to nine families were recorded. The most common representatives were Protracheoniscuspolitus Trachelipusrathkii and Ligidiumhypnorum. In general the biodiversity of isopod communities in the studied area was considerable with half of the localities explored inhabited by six to ten species. The composition of the isopod assemblages was determined by the character of the biotope and its geographical location. Forest habitats were considerably richer in species than the meadow and pasture ecosystems. Some xerotermic localities in the Slovak part of the area were inhabited by Trachelipusnodulosus and Orthometoponplanum thermophilic species typically associated with warmer parts of Europe. Two relic species (Hyloniscusmariae and Ligidiumgermanicum) were confirmed for this area. Except for only one finding of Porcellioscaber no other evidently introduced or synanthropic species were recorded. Based on the data analyzed the high nature conservancy value of the given area is emphasised.

Influence of surface flattening on biodiversity of terrestrial arthropods during early stages of brown coal spoil heap restoration.

Heterogeneity of environmental conditions is the crucial factor supporting biodiversity in various habitats, including post-mining sites. The effects of micro-topographic heterogeneity on biodiversity and conservation potential of arthropod communities in post-industrial habitats had not been studied before now. At one of the largest European brown coal spoil heaps, we sampled eight groups of terrestrial arthropods with different life strategies (moths, spiders, ground beetles, ants, orthopteroids, centipedes, millipedes, and woodlice), in successionally young plots (5-18 y), with a heterogeneous wavy surface after heaping, and compared the communities with plots flattened by dozing. A combination of the standardized quantitative sampling, using two different methods, and a paired design of the plot selection enabled a robust analysis. Altogether, we recorded 380 species of the focal arthropods, 15 of them nationally threatened. We revealed the importance of the micro-topographic heterogeneity for the formation of the biodiversity of arthropods in their secondary refuges. The communities with higher biodiversity and conservation value were detected in the plots with heterogeneous surfaces; exceptions were ground beetles and millipedes. The surface flattening, often the first step of technical reclamation projects, thus suppress biodiversity of most terrestrial arthropods during the restoration of post-mining sites. Since the communities of both surface types differed, the proportional presence on both surfaces could be more efficient in supporting the local biodiversity. We suggest reducing the surface dozing for the cases with other concerns only, to achieve a proportional representation of both surface types. Such a combination of different restoration approaches would, thus, efficiently support high biodiversity of groups with various needs.

Hungarosoma bokori Verhoeff, 1928 (Diplopoda: Chordeumatida): new insights into its taxonomy, systematics, molecular genetics, biogeography and ecology.

Hungarosoma bokori Verhoeff, 1928 is a millipede species which was originally classified solely on the basis of a female specimen. Subsequently, a long history of field searching for and surmising about the systematic position of this small, enigmatic species followed. In April 2013, 85 years after its first description, a series of nine specimens were sampled in the type locality, the Abaliget Cave, in southern Hungary. An adult male was collected for the first time, along with females and juveniles. Descriptions of the gonopods and the female vulvae, both important for considerations of the systematic position of the species, are presented for the first time. Revision and re-designation of the type material was made.The cryptic life of the species is connected with its activity in winter, and its known fragmented distribution corresponds with its presence in undisturbed microhabitats having a specific microclimate, often in the soil at cave entrances.Molecular methods showed a positive detection of the intracellular prokaryotic parasite Wolbachia in H. bokori, reflecting its highly probable parthenogenetic character in the main part of its known area of occurrence. This is the first demonstration of Wolbachia in a millipede.The legitimacy of the family Hungarosomatidae Ceuca, 1974, as a separate taxon was analysed using morphological and molecular approaches. Results of both methods confirmed the existence of a distinct phyletic line. DNA barcoding has shown its closest position to Attemsiidae Verhoeff, 1899, or Neoatractosomatidae Verhoeff, 1901. Based on records from Austria, the Czech Republic, Hungary and Slovakia, the residual circum-pannonic distribution that the whole genus (family) probably represents is proposed.

Spectroscopic parameters of the cuticle and ethanol extracts of the fluorescent cave isopod Mesoniscusgraniger (Isopoda, Oniscidea).

The body surface of the terrestrial isopod Mesoniscusgraniger (Frivaldsky, 1863) showed blue autofluorescence under UV light (330-385 nm), using epifluorescence microscopy and also in living individuals under a UV lamp with excitation light of 365 nm. Some morphological cuticular structures expressed a more intense autofluorescence than other body parts. For this reason, only the cuticle was analyzed. The parameters of autofluorescence were investigated using spectroscopic methods (molecular spectroscopy in infrared, ultraviolet-visible, fluorescence, and X-ray fluorescence spectroscopy) in samples of two subspecies of Mesoniscusgraniger preserved in ethanol. Samples excited by UV light (from 350 to 380 nm) emitted blue light of wavelengths 419, 420, 441, 470 and 505 nm (solid phase) and 420, 435 and 463 (ethanol extract). The results showed that the autofluorescence observed from living individuals may be due to some ß-carboline or coumarin derivatives, some crosslinking structures, dityrosine, or due to other compounds showing similar excitation-emission characteristics.

Methane production and methanogenic Archaea in the digestive tracts of millipedes (Diplopoda).

Methane production by intestinal methanogenic Archaea and their community structure were compared among phylogenetic lineages of millipedes. Tropical and temperate millipedes of 35 species and 17 families were investigated. Species that emitted methane were mostly in the juliform orders Julida, Spirobolida, and Spirostreptida. The irregular phylogenetic distribution of methane production correlated with the presence of the methanogen-specific mcrA gene. The study brings the first detailed survey of methanogens' diversity in the digestive tract of millipedes. Sequences related to Methanosarcinales, Methanobacteriales, Methanomicrobiales and some unclassified Archaea were detected using molecular profiling (DGGE). The differences in substrate preferences of the main lineages of methanogenic Archaea found in different millipede orders indicate that the composition of methanogen communities may reflect the differences in available substrates for methanogenesis or the presence of symbiotic protozoa in the digestive tract. We conclude that differences in methane production in the millipede gut reflect differences in the activity and proliferation of intestinal methanogens rather than an absolute inability of some millipede taxa to host methanogens. This inference was supported by the general presence of methanogenic activity in millipede faecal pellets and the presence of the 16S rRNA gene of methanogens in all tested taxa in the two main groups of millipedes, the Helminthophora and the Pentazonia.

The genus Hylebainosoma Verhoeff, 1899 (Diplopoda, Chordeumatida, Haaseidae): redescription of Hylebainosoma tatranum, description of a new troglobiont species and notes to the Hylebainosoma-Romanosoma species group.

A revision of the taxonomy, distribution and ecology of the millipedes of the genus Hylebainosoma Verhoeff, 1899 in the Carpathians is presented. Based on the study of numerous specimens, we consider the subspecies of the species                 Hylebainosoma tatranum Verhoeff, 1899 described in the past as overrated, reflecting simply intrapopulational variation attributable to the rather extended area of species distribution, different habitats of its occurrence (surface and cave habitats, soil, litter and stony debris) and wide altitudinal range from the forest zone in foothills to the alpine zone above timberline. Besides the redescription of males, the morphology of female vulvae is presented for the first time. Hylebainosoma tatranum is endemic to the extensive area of the Western and Eastern Carpathians, involving Slovakia, Hungary, the Czech Republic, Poland and highly probably Ukraine. A new troglobiont species from the karst area near the Tisovec Town (Muránska planina Mts., Central Slovakia), Hylebainosoma gulickai n. sp. is described, and is considered as stenoendemic for this small karstic region, with occurrence in few nearby caves. Hylebainosoma gulickai represents the first troglobiont chordeumatid millipede found in the northern territories of the Carpathians and the northernmost troglobiont in Central Europe in general. Morphological characteristics of both males and females are presented.Taxonomic relationships between the closely related genera Hylebainosoma and Romanosoma Ceuca, 1967 are discussed and replacement of the species Hylebainosoma cavernicola (Ceuca, 1967) n. comb., Hylebainosoma oltenica (Ceuca, 1967) n. comb. and Hylebainosoma odici (Ceuca, 1979) n. comb. into the genus Hylebainosoma is proposed. The fourth species described as Romanosoma (?) birtei Ceuca, 1967 we consider as invalid taxon. Romanosoma becomes a junior subjective synonym of the genus Hylebainosoma. 

Soil food web changes during spontaneous succession at post mining sites: a possible ecosystem engineering effect on food web organization?

Parameters characterizing the structure of the decomposer food web, biomass of the soil microflora (bacteria and fungi) and soil micro-, meso- and macrofauna were studied at 14 non-reclaimed 1- 41-year-old post-mining sites near the town of Sokolov (Czech Republic). These observations on the decomposer food webs were compared with knowledge of vegetation and soil microstructure development from previous studies. The amount of carbon entering the food web increased with succession age in a similar way as the total amount of C in food web biomass and the number of functional groups in the food web. Connectance did not show any significant changes with succession age, however. In early stages of the succession, the bacterial channel dominated the food web. Later on, in shrub-dominated stands, the fungal channel took over. Even later, in the forest stage, the bacterial channel prevailed again. The best predictor of fungal bacterial ratio is thickness of fermentation layer. We argue that these changes correspond with changes in topsoil microstructure driven by a combination of plant organic matter input and engineering effects of earthworms. In early stages, soil is alkaline, and a discontinuous litter layer on the soil surface promotes bacterial biomass growth, so the bacterial food web channel can dominate. Litter accumulation on the soil surface supports the development of the fungal channel. In older stages, earthworms arrive, mix litter into the mineral soil and form an organo-mineral topsoil, which is beneficial for bacteria and enhances the bacterial food web channel.

Assemblages of terrestrial isopods (Isopoda, Oniscidea) in a fragmented forest landscape in Central Europe.

Terrestrial isopods were collected in 13 forest fragments differing in area (within the range of 0.1 and 254.5 ha), shape and composition of forest vegetation (thermophilous oak, mesophilous oak-hornbeam, thermophilous oak-hornbeam, acidophilous oak, basiphilous oak, beech oak-hornbeam, moist mixed deciduous forest, plantations of deciduous and coniferous trees), all situated in the Ceský kras Protected Landscape Area, Czech Republic, Central Europe. Number of sites sampled in each fragment of forest depended on its size and ranged from 1 to 7. Altogether 30 sites were sampled. Soil samples (5 per site collected twice a year) and pitfall trapping (5 traps per site in continuous operation throughout a year) during 2008-2009 yielded a total of 14 species of terrestrial isopods. The highest densities and highest epigeic activities of terrestrial isopods were recorded in the smallest fragments of woodland. Although a wider range of habitats were sampled in the larger fragments of woodland there was not a greater diversity of species there and the population densities and epigeic activities recorded there were lower. Porcellium collicola was most abundant in small fragments of woodland regardless the vegetation there. Armadillidium vulgare and Protracheoniscus politus were statistically more abundant in the larger fragments of woodland. The results indicate that forest fragmentation does not necessarily result in a decrease in the species richness of the isopod assemblages in such habitats.