Epigean Habitats Research Articles

Phylogenetically independent behavior mediating geographic distributions suggests habitat is a strong driver of phenotype in crangonyctid amphipods

It is unclear if geographic distributions of animals are behaviorally mediated or simply maintained by ecologically-driven deleterious effects on fitness. Furthermore, it is not well known how behaviors that may affect geographic distributions and responses to environmental stressors evolve. To explore this, we examined behavioral and physiological reactions to light in six species of amphipods in the family Crangonyctidae collected from a variety of subterranean and epigean habitats. Stark differences between epigean and subterranean habitats occupied by different crangonyctid species allowed this clade to serve as an appropriate model system for studying the link between habitat and phenotype. We sampled habitats in or adjacent to the Edwards Aquifer in central Texas and collected two epigean and four stygobiontic species. We examined respiratory and behavioral responses to light in all study species. We found that similarities in behavioral and physiological responses to light between species were only weakly correlated with genetic relatedness but were correlated with habitat type. However, the breadth of variation in phenotype was found to be correlated with phylogenetic relationships, suggesting that population level trait evolution likely involves interactions between standing population level variation and strength of selection. Our findings suggest that natural selection via environmental conditions may outweigh history of common ancestry when predicting phenotypic similarities among species, and that behavioral and physiological phenotypes may mediate the evolution of biogeographic distributions.

The systematics of the pseudoscorpion genus Indohya (Pseudoscorpiones: Hyidae) in Australia.

The pseudoscorpion genus Indohya Beier, 1974 is known to occur in three Gondwanan fragments around the Indian Oceansouthern India, Madagascar and north-western Australiasuggesting that the genus had evolved prior to the breakup of Gondwana and was present on each landmass as they rifted apart during the Mesozoic. The Australian fauna is the most diverse, with nine species previously described from Cape Range and the Kimberley region of north-western Australia. The present study documents the genus Indohya in Australia using a combination of morphology and DNA sequence data. We found a total of 36 species, including 27 new species. The majority of the Pilbara fauna consist of blind troglobites collected from subterranean ecosystems, with an additional three eyed species from epigean habitats. The new species consist of one from Cape Range (I. anastomosa Harvey & Burger, n. sp.), 21 from the Pilbara (I. adlardi Harvey & Burger, n. sp., I. alexanderi Harvey & Burger, n. sp., I. aphana Harvey & Burger, n. sp., I. aquila Harvey & Burger, n. sp., I. arcana Harvey & Burger, n. sp., I. arnoldstrongi Harvey & Burger, n. sp., I. boltoni Harvey & Burger, n. sp., I. cardo Harvey & Burger, n. sp., I. catherineae Harvey & Burger, n. sp., I. cockingi Harvey & Burger, n. sp., I. cribbi Harvey & Burger, n. sp., I. draconis Harvey & Burger, n. sp., I. furtiva Harvey & Burger, n. sp., I. incomperta Harvey & Burger, n. sp., I. jessicae Harvey & Burger, n. sp., I. lynbeazlyeae Harvey & Burger, n. sp., I. morganstrongi Harvey & Burger, n. sp., I. rixi Harvey & Burger, n. sp., I. sagmata Harvey & Burger, n. sp., I. scanloni Harvey & Burger, n. sp. and I. silenda Harvey & Burger, n. sp.) and five from the Kimberley (I. currani Harvey & Burger, n. sp., I. finitima Harvey & Burger, n. sp., I. julianneae Harvey & Burger, n. sp., I. karenae Harvey & Burger, n. sp. and I. sachsei Harvey & Burger, n. sp.). The study is augmented with sequence data from 29 species of Indohya, including all of the 24 species recorded from the Pilbara and Cape Range, and five of the 12 known Kimberley species. Seven clades recovered during the molecular analysis are only represented by nymphs, but we used COI sequence data to diagnose these species in the absence of adult morphological data.

Six new epigean species of the genus Falcileptoneta Komatsu, 1970 (Araneae: Leptonetidae) from South Korea.

Six new leptonetid species belonging to Falcileptoneta Komatsu, 1970 are described from South Korea: F. alboclavata sp. nov., F. decemspina sp. nov., F. innuta sp. nov., F. jindoensis sp. nov., F. stilla sp. nov., and F. triformis sp. nov. All newly described species were sampled from epigean habitats (e.g. leaf litter or under the stone). Based on both male and female (except F. jindoensis sp. nov.), diagnostic characteristics, microscopic images of adult specimens and reproductive structures, and distribution records are provided. Additionally, morphological details to diagnose Falcileptoneta is also discussed.

Does size matter? Bat diversity and the use of rock shelters in Brazil

A positive correlation between cave size and bat diversity has been recorded by some authors for different cave areas in South America. We analyzed published and otherwise available data from three Brazilian karst areas and phytogeography domains, including NE and SE Atlantic Forest and Cerrado (savannah-like vegetation), encompassing a wide range of cave sizes. We found no such correlation, as expected in view of previous observations by researchers and speleologists. We discuss the ecological implications of different ways to measure bat diversity, i.e. the accumulative number of species (sequential use of caves) vs. the number of species at a given occasion (species in actual cohabitation) vs. bat abundance (numbers of individuals), as well as speleological topics related to the concept of caves (based on an anthropomorphic definition, not applicable to other species), the problems of estimating topographic variables such as cave area and volume, and the value of including cave temperatures in the analyses. Multiple interacting variables may influence the composition of cave bat communities and bat species richness and abundance. The most important ones would be the density of caves in a given area (related to lithology – calcareous vs. siliciclastic caves), regional climates, and degree of preservation of epigean habitat. Cave morphology, number and size of entrances, and presence of water bodies, as well as interactions with other species, must also be considered. In conclusion, as a general statement for tropical and subtropical bats, cave size (in the human sense) by itself does not matter, except perhaps for one-off situations. Finally, a brief overview of the diversity of cave bats in Brazil is presented.

Algae and cyanobacteria in the aphotic habitats of Veternica Cave (Medvednica Mt., Croatia) and selected caves of the Dinaric karst (South-Eastern Europe)

Microphototrophs (algae and cyanobacteria) in karst environments have been intensively studied in aquatic epigean habitats. In recent decades knowledge about the communities inhabiting cave entrances and lampenflora has grown substantially, but the data about the communities in aphotic cave zone are scarce. This study aimed to investigate spatio-temporal presence of microphototrophs in the aphotic zone of Veternica Cave (Mt. Medvednica karst) and to present additional preliminary data from 22 caves of the Dinaric karst. The data were collected over ten years, in parallel with research on cave phagotrophic protists. In addition to the remains of microphototrophs, living algae and cyanobacteria were found in the cave aphotic zone. Diatoms (Bacillariophyta) were the most frequent group found, followed by green algae (Chlorophyta), golden-brown algae (Chrysophyta) and the filamentous cyanobacteria (Cyanobacteria). The presence of microphototrophs was detected throughout the year but showed spatio-temporal variations. Microphototrophs were absent in the parts of Veternica Cave with seeping and dripping water, while they were occasionally present in the hydrologically active parts of the cave. The presence of diatoms in the aphotic zone of Veternica Cave was related to hydrological conditions, and was not affected by the distance from the cave entrance. The presence of microphototrophs in caves of the Dinaric karst has been associated with caves subject to various types of flooding by endogenous and exogenous water. Despite the fact that microphototrophs are passively transported to the caves from the surface habitats, the presence of live individuals in the cave aphotic zone implies that they should not be neglected in discussions about cave food webs. Future research of microphototrophs should be focused on the species identification, their abundance, survival time, and detail description of conditions that determine their presence in caves.

Activity of stygobionts in spring habitats: behavioral, ecological and evolutionary insights

Cave-dwelling animals might be regarded as dead-end points when it comes to their evolution and habitat exploitation. However, in the past different observations of the olms’ typical troglomorphic populations have been reported for springs of Venetia Giulia, along with a community rich in stygobionts (Bressi et al. 1999; Stoch 2017). The aim of this work is to point out the non-random active use of surface habitats by this community, providing a comparison with the occurrence observed in caves and performing an assessment of factors favouring ecotone habitats exploitation. Since 2020 we started multiple day and night surveys of olms in both 71 springs and 11 caves. Each spring and cave habitat monitored has been characterised by respect to abiotic and biotic features, including planktonic and benthic prey availability. Additionally, cave and spring populations of Troglocaris sp. were tested to determine whether they showed any behavioral difference regarding their response to light stimuli and predatory cues, as potential adaptations to the different conditions found in the two habitats. We used 34 individuals sampled from three springs and 24 from two caves of the Italian karst. We detected the olm at least once in 12 springs, with a maximum of 9 individuals occurring together. Detection probability in springs and caves was similar. Spring habitats provided higher density of potential prey available. Olms seem to prefer springs without predator fish and temporary hydroperiod. We recorded in one spring a larva of 3.5 cm which could be the smallest ever recorded in the field. Preliminary results of the behavioral experiments show no susceptibility to light neither for spring nor cave populations of Troglocaris sp., as for chemical cues we expect a difference in reactivity depending on the habitat of origin of the shrimp, consistently with the top predator present. We suggest that epigean habitats and borders with surface may have an underestimated importance for animals adapted to subterranean environments (Manenti and Piazza 2021), including the olm. Further studies focusing on the ipogean community which actively exploits spring habitats are underway, with the aim of determining the diel occupation of these complex ecotones.

It’s All about the Zone: Spider Assemblages in Different Ecological Zones of Levantine Caves

Caves possess a continuum of ecological zones that differ in their microhabitat conditions, resulting in a gradient of nutrients, climate, and illumination. These conditions engender relatively rapid speciation and diverse assemblages of highly specialised spider fauna. It is unclear, however, how zonation of these caves affects spider assemblage composition and structure. Surveys of 35 Levantine caves were conducted to compare the assemblages of spiders between their different ecological zones. The diverse spider assemblages of these caves differed between the entrance, twilight, and dark zones, with troglophiles and accidental species occupying the cave entrance, endemic troglobites occupying the dark zones, and hybrid assemblages existing in the twilight zones. The progression of assemblage composition and divergence throughout cave zones is suggestive of processes of ecological specialisation, speciation, and adaptation of cave-endemic troglobites in the deepest zones of caves, while cave entrance assemblages are composed of relatively common species that can also be found in epigean habitats. Moreover, the cave entrance zone assemblages in our study were similar in the different caves, while the cave dark zone assemblages were relatively distinct between caves. Cave entrance assemblages are a subset of the regional species pool filtered by the cave conditions, while dark zone assemblages are likely a result of adaptations leading to local speciation events.

Environmental factors structuring the assemblage of aquatic insects in an epigean and hypogean stretch of a Neotropical karst stream

The aim of the present study was to identify which variables structure the assemblage of aquatic insects in the subterranean environment, because knowledge about how subterranean communities are structured in rivers with hypogean sections is still emerging. In addition to identifying whether the hypogean assemblage is structured in the same way as the epigean assemblage, we sought to identify whether the environmental restrictions caused by the subterranean habitat resulted in reduced species richness and abundance compared with the epigean habitat. We sampled a 345-m stretch of a stream in Brazil and collected aquatic insects from both the epigean and hypogean environments. In the stream stretch investigated, there was a positive relationship between richness and the percentage of immersion, fine substrate and organic matter, and a negative relationship between richness and conductivity. We also verified differences between the epigean and hypogean environments regarding composition. The variables that best explained the overall assemblage structure included the epigean–hypogean environment and the percentage of coarse and fine substrate. Given the differences between these environments, we emphasise the need for management actions in karst regions that consider the discontinuity of these regions’ river communities to conserve all components of diversity.

Phylogeny and biogeography of the Cavernicola (Platyhelminthes: Tricladida): Relicts of an epigean group sheltering in caves?

The planarian suborder Cavernicola Sluys, 1990 was originally created to house five species of triclad flatworms with special morphological features and a surprisingly discontinuous and broad geographic distribution. These five species could not be accommodated with any degree of certainty in any of the three taxonomic groups existing at that moment, viz., Paludicola Hallez, 1892, Terricola Hallez, 1892, and Maricola Hallez, 1892. The scarce representation of the group and the peculiarities of the morphological features of the species, including several described more recently, have complicated new tests of the monophyly of the Cavernicola, the assessment of its taxonomic status, as well as the resolution of its internal relationships. Here we present the first molecular study including all genera currently known for the group, excepting one. We analysed newly generated 18S and 28S rDNA data for these species, together with a broad representation of other triclad flatworms. The resulting phylogenetic trees supported the monophyly of the Cavernicola, as well as its sister-group relationship to the Maricola. The sister-group relationship to the Maricola and affinities within the Cavernicola falsify the morphology-based phylogeny of the latter that was proposed previously. The relatively high diversity of some cavernicolan genera suggests that the presumed rarity of the group actually may in part be due to a collecting artefact. Ancestral state reconstruction analyses suggest that the ancestral habitat of the group concerned epigean freshwater conditions. Our results point to an evolutionary scenario in which the Cavernicola (a) originated in a freshwater habitat, (b) as the sister clade of the marine triclads, and (c) subsequently radiated and colonized both epigean and hypogean environments. Competition with other planarians, notably members of the Continenticola, or changes in epigean habitat conditions are two possible explanations -still to be tested- for the loss of most epigean diversity of the Cavernicola, which is currently reflected in their highly disjunct distributions.

Species conservation profiles of cave-dwelling arthropods from Azores, Portugal.

BackgroundAzorean volcanic cave biodiversity is under considerable pressure due to ongoing threats of pollution, land use change, touristic activities or climate change. In this contribution, we present the IUCN Red List profiles of 15 cave-adapted arthropod species, endemic to the Azorean archipelago, including species belonging to the speciose genus Trechus (Carabidae), which is represented in Azores by seven species. The objective of this paper is to assess all endemic Azorean cave-adapted species and advise on possible future research and conservation actions critical for the long-term survival of the most endangered species.New informationMost species have a restricted distribution (i.e. occur in one or two caves), very small extent of occurrence (EOO) and a small area of occupancy (AOO). A continuing decline in the number of mature individuals is inferred from the ongoing cave habitat degradation. The two troglobitic species of the homopteran genus Cixius are in great danger of extinction due to major land-use changes in epigean habitats above their known localities. We suggest, as future measures of conservation, the regular monitoring of the species (every five years), the creation of additional protected caves, the limitation of several aggressive activities around the caves (e.g. decreasing pasture intensification) and in some cases the creation of fences in the entrance of the most important caves.

Behavioural adaptations of cave orb spiders

Given the low abundance of flying insects in subterranean habitats, it is surprising that troglophile orb spiders in the genus Meta (family Tetragnathidae) are relatively abundant in subterranean habitats, especially in the twilight zone of European caves. As is the case for most subterranean invertebrates, we know very little about their behaviour. However, they have been reported to possess a number of specific behavioural adaptations to the cave environment. These include modifications of the traditional orb web such that webs do not have frame and anchor threads, but instead have the radii (the spokes in the web) attaching directly to the cave walls, and that cave spiders, unlike epigean orb spiders, allegedly engage in off-web foraging as well as on-web foraging. Here I review the evidence for these claims and compare these to the behaviour of epigean spiders in the related Metellina genus including preliminary comparative data on both morphology and web characteristics. In addition, I will discuss the potential of using the cave Meta spiders, especially M. menardi and M. bourneti, as model organisms for studying search behaviour, mate finding behaviour and sexual selection, as these spiders show an interesting life cycle with young instars leaving the cave environment to spend an instar in the epigean environment before returning. It is currently not known how spiderlings locate subterranean habitats, how males find female webs or whether the level of sexual selection is lower in the resource-limited subterranean habitat compared to the epigean habitat.

Terrace springs: habitat haven for macrobenthic fauna in the lower plain of the River Ticino (Lombardy, Northern Italy)

Springs are important environments between hypogean and epigean habitats; the interaction between aquatic and terrestrial ecosystems is an important factor for their biotic communities. We investigated the ecology of the macrobenthic community of two lowland springs in the River Ticino valley, focusing on the autoecology of some relevant species and on the role of springs as hotspots of biodiversity in an area threatened by anthropogenic pressure. We collected 26 taxa in total: diptera (8), trichoptera (6), gastropods (5), coleoptera (2), crustacea (2), lumbricidae (1), odonata (1), plathelminthes (1). Some of them are stenothermal and oligotrophic species thus quite unusual for a flood plain area characterized by intensive agricultural activities. In conclusion, our study highlights the importance of lowland springs in conserving some habitat-selective macrobenthic species.

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.

The role of isolation on contrasting phylogeographic patterns in two cave crustaceans

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

A new stem bythinine in Cretaceous Burmese amber and early evolution of specialized predatory behaviour in pselaphine rove beetles (Coleoptera: Staphylinidae)

Comprising more than 10,000 valid species, the staphylinid subfamily Pselaphinae is a major element of epigean habitats, and among the most diverse groups of rove beetles. Pselaphinae is split basally into two principal clades: the small supertribe Faronitae, and its sister group, the hyper-diverse ‘higher Pselaphinae’ containing the remaining five supertribes. Deducing the origins and divergence times of major higher Pselaphinae clades requires direct fossil evidence. Here we describe a new pselaphine rove beetle, Cretobythus excavatus Yin, Parker & Cai gen. et sp. nov., based on a well-preserved individual embedded in mid-Cretaceous amber from Myanmar (Cenomanian, c. 99 Ma). Cretobythus does not obviously belong to any Recent tribe, but Bayesian phylogenetic placement using morphological characters supports a position within the stem-group of the tribe Bythinini, sister to Boreotethys Parker, a genus also recently described from Burmese amber. Together, Cretobythus + Boreotethys comprise the sister group of modern Bythinini. Despite some external similarities to Recent Bythinini, Cretobythus exhibits several plesiomorphic traits, including a generally flattened body plan, and metacoxae that are positioned close to the ventral midline. The resemblance in form of the enlarged maxillary palpi of Cretobythus to extant bythinines implies a similar function in prey capture, indicating that the unusual employment of the maxillary palps to trap moving prey in Bythinini had probably evolved by the mid-Cretaceous, at the latest.http://zoobank.org/urn:lsid:zoobank.org:pub:9FEB523F-76A7-4502-AF02-FCC0FAAB262A

Revision of the Remaneicaris argentina-group (Copepoda, Harpacticoida, Parastenocarididae): supplementary description of species, and description of the first semi-terrestrial Remaneicaris from the tropical forest of Southeast Mexico.

Remaneicaris is a species-rich Neotropical monophyletic group, easily recognized by the synapomorphic position of the outer seta of the third exopodite of leg 4, localized at 2/3 of the outer margin. The genus, comprising 35 species in five monophyletic groups, plus R. ignotus and R. meyerabichi, retains an unusual set of plesiomorphic characters. Herein we supplement the descriptions of the species belonging to the Remaneicaris argentina-group, and describe a new species from the tropical forest of Southeast Mexico. The present study extends the geographic distribution of the genus, with the northernmost record until now being from El Salvador. The genus having hitherto been known from interstitial groundwater habitats, this is its first record in epigean semi-terrestrial habitats. Remaneicaris siankaan sp. nov. was found in phytotelmata (bromeliads), leaf litter, moist soil, permanent ponds (known locally as "aguadas"), and temporal and permanent wetlands (savannahs). The new species can be easily characterized by its finely pitted cuticle, the ε (epsilon)-shaped thumb of the male P3 and the bifid accessory spine with distal hyaline inner tip, which precedes the thumb. A new method for the 3D reconstruction of microcrustaceans is described.

A new species of Moraria (Copepoda, Harpacticoida, Canthocamptidae) from groundwaters of Germany, including a key for the identification of the species of the Western Palaearctic Region

A new species of Moraria collected from interstitial waters of southern Germany is described. The new species Moraria glitzae n. sp. can be differentiated from other members of the genus by a combination of morphological traits related to the shape of the anal operculum, the armature of leg 4 and the ornamentation of the urosome in both sexes. Males of M. glitzae n. sp. have a typical number of armature elements on the endopods of legs 2 to 4, some of which with unique morphology. Females can be distinguished from those of other species of the genus by the number of antennular segments, the number of setae on the exopod of the antenna and the mandibular palp, the armature of legs 1, 3 and 4, and the size and armature of leg 5. The relationship of the new species to closely related species, viz., M. catalana, M. jana, M. stankovitchi and M. varica is discussed. The morphological differences to other species inhabiting groundwaters and epigean habitats of the region, viz., M. alpina, M. brevipes, M. fontinalis, M. mrazeki mrazeki, M. mrazeki macedonica, M. pectinata, M. poppei and M. radovnae are presented. A distribution map of the Moraria species known from Germany and a dichotomous identification key of the 20 species from the Western Palaearctic Region are provided.

A Minimal Invasive Method to Forecast the Effects of Anthropogenic Disturbance on Tropical Cave Beetle Communities.

Many tropical landscapes are changing rapidly, with uncertain outcomes for biodiversity, landscape function, and the corresponding landscape services. Therefore, monitoring and adaptively managing the drivers and consequences of landscape change while sustaining the production of essential resources have become research and policy priorities. In this perspective, we have applied a recent framework, the stochastic dynamic methodology (StDM), with the purpose of understanding the effects of natural and anthropogenic disturbances on caves' integrity using cave beetle communities (Coleoptera) as ecological indicators. The proposed method was preceeded by a generalized linear model for discriminating significant relationships between the selected indicators, the structural changes in the caves, and the epigean habitats associated. The obtained results showed different ecological trends in response to the environmental changes. Overall, the simulation results seem to demonstrate the StDM reliability in determining the effects of habitat dynamics, that is, the expansion of agricultural activities, in areas near the caves in the structure of cave beetle communities. The applied method, based on universal information-theoretic principles, can be easily implemented and interpreted by environmental managers and decision makers, enabling anticipating impacts and supporting the development of measures aimed at minimizing the identified problems.

Two new high-altitude subspecies of Tapinopterus (Tapinopterus) cognatus (Dejean, 1831) (Coleoptera: Carabidae: Pterostichinae) from Serbia.

The genus Tapinopterus Schaum, 1858 currently comprises around 60 species belonging to eight subgenera, inhabiting mostly the Balkan Peninsula and Asia Minor (Schatzmayr 1942, 1943; Bousquet 2003; Ćurčić et al. 2008; Arndt et al. 2011; Lohaj et al. 2012). The taxa live in limited geographic areas and are mostly montane (Trautner & Geigenmüller 1987; Guéorguiev & Guéorguiev 1998). The territory of Serbia is inhabited by two species of Tapinopterus, T. (T.) cognatus (Dejean, 1831) (recorded in Bulgaria and Romania as well) and T. (T.) miridita (Apfelbeck, 1904) (recorded in Albania and the Republic of Macedonia as well), as well as by the three subspecies, T. (T.) cognatus cognatus (Dejean, 1831) (known from Mt. Suva Planina in Serbia and from Bulgaria), T. (T.) cognatus winkleri Mandl, 1936 (known from the Svrljiške Planine Mts. and Mt. Stara Planina in both Serbia and Bulgaria), and T. (T.) miridita miridita (Apfelbeck, 1904) (known from Mt. Šar Planina in both Serbia and the Republic of Macedonia), respectively (Mandl 1936; Schatzmayr 1942, 1943; Guéorguiev & Guéorguiev 1995; Drovenik & Peks 1999; Bousquet 2003; Ćurčić et al. 2007; Guéorguiev & Giachino 2008). In our opinion, T. (T.) cognatus cognatus does not live in Bulgaria (as was previously reported) (Bousquet 2003), while the older Schatzmayr's (1943) record "Klisura" should refer to the Iron Gate Gorge in SW Romania and NE Serbia, not to any Bulgarian place. The other mentioned locality, i.e., Mt. Suva Planina in Serbia, is situated far from the type locality (Iron Gate). Since nearly all Tapinopterus taxa are regional or local endemics, this southernmost record in SE Serbia for T. cognatus cognatus seems unlikely. In view of the high diversity of Tapinopterus on the Balkan Peninsula, it is expected that new taxa exist in border areas of the distribution in Serbia. During the last years, the first (SĆ), second (NI) and fourth author (DA) of the paper investigated a number of epigean montane habitats in SE and C Serbia in order to confirm such an assumption, by collecting a series of ground beetles at high altitude sites. After careful examination of the samples, we identified two Tapinopterus subspecies new to science.

DNA Barcoding Cleans House Through the Carychiidae (Eupulmonata, Ellobioidea)*

Abstract: The terrestrial, ellobioid taxon, Carychiidae provides an excellent case study for testing integrative taxonomy and addressing the plethora of historical species designations based on vague morphological characters. Since the Carychiidae are ephemeral, hermaphroditic microgastropods (< 2mm) inhabiting permanently moist aphotic zones of epigean (Carychium O. F. Muller, 1773) and subterranean habitats (eutroglobiont Zospeum Bourguignat, 1856), studies investigating their reproductive system for species delimitation have been futile. Many carychiid species designations were established during the mid-18th to 20th Century heydays of species discovery. Naturalists either split (“splitters”) species by recognizing them according to trivial differences in shell morphology or grouped them (“lumpers”) based upon common morphological traits. The concept of phenotypic variability was not considered or greatly underestimated in these species hypotheses. Although integration of DNA barcodes has since enabled...