The role of Anatolia in the origin of the Caucasus biodiversity hotspot illustrated by land snails in the genus Oxychilus.

Several taxa that are distributed in the Caucasus and/or the adjacent Pontic Mountains also have representatives in the East Mediterranean region. These disjunctions could have been caused by long-distance dispersal or be the result of extinctions in Central Anatolia caused by the aridification of the Anatolian Plateau during the Pliocene. We studied the Longiphallus-Hiramia group of Oxychilus as an example showing such distribution patterns. Phylogenetic analyses of the Oxychilus species previously classified in Longiphallus, Hiramia and related subgenera resulted in a new delimitation of these taxa and the recognition of Anatoloxychilus Neiber, Walther & Hausdorf n. subgen. as an additional clade. Based on phylogenetic and population genetic analyses, O. reticulatus from Mingrelia is revalidated and the populations from the Pontic Mountains previously identified with O. mingrelicus koutaisanus are recognised as a distinct species. Three species pairs of the Longiphallus-Hiramia group with deep splits predating the aridification of the Anatolian Plateau during the Pliocene show disjunctions between the Caucasus/Pontic region and the Mediterranean. The majority of taxa with such a distribution pattern probably had more continuous distributions before the aridification started. The relationships between the Hiramia species from the Caucasus, the Pontic Mountains and the East Mediterranean highlight the importance of the Anatolian land as a source area for the colonisation of the Caucasus region. The dating of the divergences of the Caucasian Hiramia species in the middle to late Miocene indicated that they colonised the Caucasus when it was still an island in the Paratethys Sea and that their divergence was triggered by the orogenesis of the Greater Caucasus. A common pattern within the Caucasus region, also found in Hiramia, is the separation of taxa in the north-western Greater Caucasus from taxa inhabiting the southern slopes of the central Greater Caucasus.

Phylogeny, phylogeography and hybridization of Caucasian barbels of the genus Barbus (Actinopterygii, Cyprinidae).

Phylogenetic relationships and phylogeography of six species of Caucasian barbels, the genus Barbus s. str., were studied based on extended geographic coverage and using mtDNA and nDNA markers. Based on 27 species studied, matrilineal phylogeny of the genus Barbus is composed of two clades - (a) West European clade, (b) Central and East European clade. The latter comprises two subclades: (b1) Balkanian subclade, and (b2) Ponto-Caspian one that includes 11 lineages mainly from Black and Caspian Sea drainages. Caucasian barbels are not monophyletic and subdivided for two groups. The Black Sea group encompasses species from tributaries of Black Sea including re-erected B. rionicus and excluding B. kubanicus. The Caspian group includes B. ciscaucasicus, B. cyri (with B. goktschaicus that might be synonymized with B. cyri), B. lacerta from the Tigris-Euphrates basin and B. kubanicus from the Kuban basin. Genetic structure of Black Sea barbels was influenced by glaciation-deglaciation periods accompanying by freshwater phases, periods of migration and colonization of Black Sea tributaries. Intra- and intergeneric hybridization among Caucasian barbines was revealed. In the present study, we report about finding of B. tauricus in the Kuban basin, where only B. kubanicus was thought to inhabit. Hybrids between these species were detected based on both mtDNA and nDNA markers. Remarkably, Kuban population of B. tauricus is distant to closely located conspecific populations and we consider it as relic. We highlight revealing the intergeneric hybridization between evolutionary tetraploid (2n = 100) B. goktschaicus and evolutionary hexaploid (2n = 150) Capoeta sevangi in Lake Sevan.

Patterns of oribatid mite species diversity: testing the effects of elevation, area and sampling effort.

Elevational gradients in species diversity and species area relationships are two well established patterns that are not mutually exclusive in space and time. Elevation and area are both considered as good proxies to detect and characterize the patterns of species diversity distribution. However, such studies are hampered by the incomplete biodiversity data available for ecologists, which may affect the pattern perceptions. Using the large dataset of oribatid mite communities sampled in Georgia, we tested the effects of altitude and area on species distribution using various approaches, while explicitly considering the biases from sampling effort. Our results showed that elevation and area are strongly correlated (with increasing absolute elevation, land area decreases) and both have strong linear effects on species diversity distribution when studied separately. Approaches based on multiple regression and direct removal of co-varied factors, indicated that the effect of area can actually override the effect of elevation in describing the oribatid species diversity distribution along with elevation. On the other hand, the bias of sampling proved significant in perception of elevational species richness pattern with less effect on elevational species area relationship. We suggest that the sampling alone may be responsible for patterns observed and thus should be considered in ecological studies when eligible.

Oribatid mite communities along an elevational gradient in Sairme gorge (Caucasus).

Many aboveground animals and plant communities have been studied along elevational gradients whereas studies on soil animals are scarce. Here, we studied oribatid mite community distribution along an elevational gradient from 600 to 2200 m in forest ecosystems of the Western Lesser Caucasus Mountains in Georgia. Overall, 86 oribatid mite species were found at the study sites. Oribatid mite densities were generally low (~9800 ind./m(2)), and 74% of all species reproduced sexually indicating that resource conditions at the study sites are generally poor. Oribatids mainly comprised Brachypylina (76%), Mixonomata (13%), Desmonomata (6%) and Enarthronota (5%). Oribatid mite community structure changed along the elevational gradient and the changes correlated with temperature, pH, litter thickness and density of the herb layer. The dominance of sexually reproducing taxa and low overall abundance indicate that the studied elevational gradient is characterized by poor resource conditions for soil microarthropods. Oribatid mite species richness and density declined with elevation suggesting that decreasing temperature in concert with resource limitation is a main driver of oribatid mite communities whereas stochastic factors (such as mid-domain effects) are of minor importance.

Checklist of digeneans (Platyhelminthes, Trematoda, Digenea) of Georgia.

Background: In the present study, we aim to provide an inventory of digenetic trematodes (Platyhelminthes, Trematoda, Digenea) from Georgia including records from the freshwater, marine and terrestrial realms. The checklist is based on a critical review of data from 109 papers, 11 monographs and four Ph.D. theses published between 1935 and 2019 and our new records. The checklist includes information on synonymy, the host species, site of infection, geographical distribution and bibliographical references. The present data will serve as a baseline for further studies on trematodes from Georgia focused on integrative taxonomy, life-cycle elucidation, parasite ecology and epidemiology. New information: We compiled data on the digenean trematode fauna of Georgia, which is represented by 186 species (of these 173 identified to species level) belonging to 108 genera, 47 families and 17 superfamilies. This is the first checklist of the digeneans of Georgia. The majority of digenean species were recorded as adults (160 species), only a small fraction being found as cercariae (33 species) or metacercariae (24 species), in their first or second intermediate hosts, respectively. Predominantly, records of trematodes (62 species) from birds were found, followed by those parasitising fish (50 species, i.e. 32 species as adults and 18 as metacercariae), mammals (33 species) and amphibians (25 species, i.e. 23 species as adults and 2 as metacercariae), with the least number of species reported from reptiles (12 species, i.e. 9 species as adults and 3 as metacercariae). Adult digeneans recorded together with another life-cycle stage (metacercariae and/or cercariae) comprised 28 species, i.e. for 15% of the total trematode species number, a part of their life-cycle is known.

The impact of insect herbivory on biogeochemical cycling in broadleaved forests varies with temperature.

Herbivorous insects alter biogeochemical cycling within forests, but the magnitude of these impacts, their global variation, and drivers of this variation remain poorly understood. To address this knowledge gap and help improve biogeochemical models, we established a global network of 74 plots within 40 mature, undisturbed broadleaved forests. We analyzed freshly senesced and green leaves for carbon, nitrogen, phosphorus and silica concentrations, foliar production and herbivory, and stand-level nutrient fluxes. We show more nutrient release by insect herbivores at non-outbreak levels in tropical forests than temperate and boreal forests, that these fluxes increase strongly with mean annual temperature, and that they exceed atmospheric deposition inputs in some localities. Thus, background levels of insect herbivory are sufficiently large to both alter ecosystem element cycling and influence terrestrial carbon cycling. Further, climate can affect interactions between natural populations of plants and herbivores with important consequences for global biogeochemical cycles across broadleaved forests.

Development and application of a second-generation multilingual tool for invasion risk screening of non-native terrestrial plants.

Under the increasing threat to native ecosystems posed by non-native species invasions, there is an urgent need for decision support tools that can more effectively identify non-native species likely to become invasive. As part of the screening (first step) component in non-native species risk analysis, decision support tools have been developed for aquatic and terrestrial organisms. Amongst these tools is the Weed Risk Assessment (WRA) for screening non-native plants. The WRA has provided the foundations for developing the first-generation WRA-type Invasiveness Screening Kit (ISK) tools applicable to a range of aquatic species, and more recently for the second-generation ISK tools applicable to all aquatic organisms (including plants) and terrestrial animals. Given the most extensive usage of the latter toolkits, this study describes the development and application of the Terrestrial Plant Species Invasiveness Screening Kit (TPS-ISK). As a second-generation ISK tool, the TPS-ISK is a multilingual turnkey application that provides several advantages relative to the WRA: (i) compliance with the minimum standards against which a protocol should be evaluated for invasion process and management approaches; (ii) enhanced questionnaire comprehensiveness including a climate change component; (iii) provision of a level of confidence; (iv) error-free computation of risk scores; (v) multilingual support; (vi) possibility for across-study comparisons of screening outcomes; (vii) a powerful graphical user interface; (viii) seamless software deployment and accessibility with improved data exchange. The TPS-ISK successfully risk-ranked five representative sample species for the main taxonomic groups supported by the tool and ten angiosperms previously screened with the WRA for Turkey. The almost 20-year continuous development and evolution of the ISK tools, as opposed to the WRA, closely meet the increasing demand by scientists and decision-makers for a reliable, comprehensive, updatable and easily deployable decision support tool. For terrestrial plant screening, these requirements are therefore met by the newly developed TPS-ISK.

Ponticolaalasanicus sp. n. (Gobiiformes, Gobiidae) from the Alazani River Basin, Georgia.

Background: The South Caucasus Region and Georgia, in particular, is a biodiversity hotspot and characterised by high diversity of landscapes and ecosystems, as well as high levels of endemism. At the same time, diversity of freshwater organisms in the region remains poorly studied, including fishes. The freshwater fish fauna of the South Caucasus Region consists of 119 fish species, of which 13 species belong to the order Gobiiformes. It should be noted that gobies are amongst the poorly studied taxa in Georgia and probably unknown/undescribed species still living in the Georgian freshwater ecosystems which requires further research. New information: Ponticolaalasanicus, a new species is described from the Alazani River, western Caspian Sea Basin, Georgia. It is distinguished from its congeners in the Caspian and Black Sea Basins by having the following features: dorsal fin with VI-VII spines and 15½-16½ branched rays, anal fin with 10½-12½ branched rays; lateral line with 48-55 scales; laterally compressed body with dark brown and black blotches - scales ctenoid; first and second dorsal fins almost touching with dorsal fins bases; head large, depressed, wider than deep, its length approaches almost 3.4th of standard length; nape scaled completely; cycloid scales cover upper part of opercle, cheeks noticeably swollen; snout longer than eye, eye diameter 4.5 times its head length; lower jaw slightly protruding; upper lip is uniform; pelvic disc short, elongated and flat, not reaching the anus; the pectoral fins extends vertically through first branched dorsal fin; caudal fin rounded. Ponticolaalasanicus sp. n. belongs to P.syrman group and it is separated by a minimum Kimura 2-parameter distance of 3.5, 3.6 and 4.8% from P.syrman, P.iranicus and P.patimari, respectively.

Bats of Georgia - an occurrence dataset from 1835 through 2022.

Background: In Georgia, currently, 30 species of bats are recorded from four families and eleven genera. Although the oldest record of bats is from 1835 and continues until today, there are no comprehensive data available for bat diversity and distribution in Georgia. Thus, we aimed to fill that gap and make complete, expertly curated literature and our own published data openly available (through GBIF) for researchers and conservationists. New information: In this publication, out of 1987 records, 1243 (62.4%) are new and unpublished data. Generally, out of all records, 34% are literature and museum data and 66% are data collected by us. Additionally, for the first time in the history of the study of bats in Georgia, we initiated surveys in forested areas of the country.

Annotated checklist of Georgian oribatid mites-II.

The checklist of oribatid mites from Georgia is updated using sampling data from more than 230 locations during field works conducted between 2016 and 2022. We report 32 species of oribatid mites, sixteen of them were collected between 2017 and 2019, six were missing in Checklist I, while eleven species are new records for the country. The genus Ameronothrus was found in Georgia for the first time, but could not be identified to species level. Additional information about the distribution of Epilohmannia styriaca Schuster, 1960, Sphaerozetes orbicularis (C.L. Koch, 1835) and Fuscozetes fuscipes (C.L. Koch, 1844) is given. For each species we provide notes about regional and global distribution as well as information about their ecology. This list brings the number of identified species known for Georgia up to 563.

The genus Acicula W. Hartmann, 1821 in the Caucasus (Gastropoda: Caenogastropoda: Aciculidae).

The Aciculidae is a family of terrestrial operculate snails with a western Palaearctic distribution. Historically three Acicula species were reported from the Caucasus Mountains, which represents the eastern edge of the family's distribution: A. limbata, A. moussoni and A. parcelineata, all from a few localities. We examined extensive material from the entire range of the Caucasus Mountains by means of classic morphology-based taxonomy and geometric morphometries. We arrived at the following conclusions: (1) At least three species inhabit the Caucasus Mountains, two of which are problematic taxonomically and regarding their nomenclature. (2) The two syntypes of A. moussoni have been lost, and both lectotype designations by the former authors were invalid. Therefore, we designate a neotype for A. moussoni. (3) Acicula limbata was described from Lower Miocene deposits of the Czech Republic. Although the fossil Central European and extant Caucasian shells differ only slightly from one another, it is highly unlikely that they would belong to the same species. Therefore, we suggest using A. limbata only for the Czech fossil species and A. moussoni for the Caucasian extant species. (4) In multiple samples, the generally smaller Acicula parcelineata was found syntopic with the generally larger A. moussoni. Therefore, the two are possibly distinct species. However, it is very difficult to distinguish, and the problem of A. moussoni and A. parcelineata needs further attention. (5) A species with a bullet (or torpedo)-shaped shell and an opisthocline apertural profile from the northern Caucasus region has been treated as A. moussoni by previous authors, but it is distinct from the other two Acicula species by consistent conchological characters and was also found with them in syntopy, and is described here as Acicula telum Páll-Gergely, n. sp.

Diatom endemism and taxonomic turnover: Assessment in high-altitude alpine lakes covering a large geographical range.

Diatoms are widely used as ecological indicators and show various degrees of endemism. Many studies that support the idea of endemic species integrate several climate zones, a variety of ecosystem types, and often focus on a global scale. Here, we investigated whether endemism could be detected when considering a homogeneous type of ecosystem in a single climate zone. We sampled stone biofilms at 40-50 cm depth in high-altitude lakes in the Alpine climate zone. A total of 149 samples were obtained from the French and Georgian mountains, two areas separated by ∼3000 km. Using Amplicon Sequence Variants derived from DNA metabarcoding, we assessed taxonomic turnover and Zeta-diversity (a measure of endemism). We ran haplotype networks and phylogenetic tests to measure geographical signal in the phylogenies of dominant taxa. The French and Georgian communities shared 51 % of species. Species that were not shared across both regions were mostly rare, and often not characteristic of lakes but of neighboring habitats instead. In contrast, at the sub-species level, 87 % of the genotypes showed restricted distributions. Whereas endemism was the rule at sub-species level, most species were shared across both French and Georgian lakes, suggesting that geographic barriers strongly limited dispersal at the sub-species level but not species level. Dominant species hosted higher levels of sub-specific diversity than rare species. In contrast to global-scale studies, we did not find any significant geographical structuring in the phylogeny of the investigated species. This could indicate ongoing dispersal at a frequency fast enough to prevent allopatric divergence, yet slow enough to prevent sharing most haplotypes between France and Georgia. These results have implications for biomonitoring: depending on the taxonomic level chosen, robust generic tools (species level) or tools dedicated to a region able to discriminate fine pressures differences (sub-species level) may be developed.

Into the unknown: the first barcode-assisted checklist of Psocoptera (Insecta, Psocodea) of Georgia with a census on country species richness.

This checklist reports 47 species of Psocoptera from 15 families and three suborders from Georgia, of which 31 species are recorded for the first time, increasing the known fauna of the country by more than 65%. Of these, 37 species have been barcoded, representing 210 Barcode Identification Numbers (BINs). An additional 14 species are expected to occur in Georgia but remain undiscovered, meaning that only ≈ 77% of the fauna is currently documented. Barcodes, comments on distributions, and images of voucher specimens are given followed by a map of the sampling sites.

Species diversity and DNA barcode library of freshwater Molluscs of South Caucasus.

This study provides the first attempt to investigate the molecular diversity of South Caucasian freshwater molluscs (Mollusca, Gastropoda) and lay down the first bricks to build up a DNA-barcode library. In total, 289 COI barcode sequences were obtained from 33 morpho-species belonging to 24 molluscan genera and 10 families that represent nearly 30% of known freshwater molluscan diversity of the South Caucasus region. DNA barcodes were analysed by means of the Barcode Index Number (BIN) and the other tools available in BOLD Systems. Results showed that the knowledge of freshwater molluscs diversity in the South Caucasus is far from comprehensive. For the studied 33 morpho-species, 289 barcodes were clustered into 40 BINs, from which unique BINs were defined for 12 species and five species were characterised with more than a single BIN. From the studied taxa, 60% were characterised larger than 2.2% sequence divergence indicating high genetic variation or cryptic diversity. Within our limited taxonomic coverage, we found one new species for the Republic of Georgia (Galbaschirazensis) and at least three undescribed species belonging to the genera Stagnicola, Segmentina and Anisus. Uniqueness and high molecular diversity of the studied species emphasise the need for further intensive morphological and molecular investigations of the South Caucasian freshwater molluscan fauna.

Species diversity and distribution of freshwater molluscs of Javakheti Highlands (Republic of Georgia).

The diversity and distribution of freshwater molluscs is poorly studied in the Republic of Georgia, due to the scarcity of field studies during the last 50 years. Here, we present the results of the first concerted investigation of freshwater mollusc biodiversity in the Javakheti Highlands, in the southern, mountainous region of Georgia. In total, we were able to collect 22 species from 42 sampling localities, including different kinds of freshwater habitats. Amongst the 22 collected species, 12 were recorded for the first time from Javakheti. From the newly-recorded species, Bathyomphalus contortus is a new country record, whose identity is supported by 16S rRNA sequence data.

Genetic diversity, phylogenetic position and morphometric analysis of Astacus colchicus (Decapoda, Astacidae): a new insight into Eastern European crayfish fauna.

The phylogeny of European crayfish fauna, especially with respect to Eastern European species, is still far from being completely resolved. To fill this gap, we analyzed most of the European crayfish species focusing on the phylogenetic position of the endemic crayfish Astacus colchicus, inhabiting Georgia. Three mitochondrial and one nuclear marker were used to study evolutionary relationships among European crayfish species, resulting in the unique phylogenetic position of A. colchicus indicating independent species status to A. astacus. Phylogenetic analyses revealed a deep molecular divergence of A. colchicus in comparison to A. astacus (6.5-10.9% in mtDNA and 1.1% in nDNA) as well as to Pontastacus leptodactylus and P. pachypus (5.5-10.0% in mtDNA and 1.4-2.4% in nDNA). Absent ventral process on second male pleopod and abdominal somites II and III with pleura rounded lacking prominent spines clearly indicate taxonomic assignment to the genus Astacus; however, the species is distributed almost in the middle of Ponto-Caspian area typical by occurrence of the genus Pontastacus. Several morphological indices linked to head length, carapace, and total body length and width were found to demonstrate apparent differences between A. colchicus and A. astacus. Although this study provides a novel insight into European crayfish phylogeography, we also point out the gaps in comprehensive study of the P. leptodactylus species complex, which could reveal details about the potential species status of particular species and subspecies within this genus.

Towards retrieving the Promethean treasure: a first molecular assessment of the freshwater fish diversity of Georgia.

In this study, we provide a first estimation of the molecular diversity of the freshwater fishes of Georgia. In addition to field collections, we integrated DNA barcode data obtained from recent works and public databases (BOLD and NCBI GenBank). Currently, the DNA barcode reference library for freshwater fishes of Georgia comprises 352 DNA barcodes for 50 species, 36 genera and 15 families (52% of total Georgian freshwater fish diversity), from which 162 DNA barcodes belonging to 41 species were newly generated as part of this study. A total of 22 species are reported from the Caspian Sea basin and 31 from the Black Sea basin. Amongst the studied taxa, seven species were found with large interspecific divergences (> 2%) while 11 species were found to share DNA barcodes within our dataset. In the course of the study, we found the first evidence of the existence of Gymnocephalus cernua (Linnaeus, 1758) and also confirm the second occurrence of invasive Rhinogobius lindbergi (Berg, 1933) in Georgia. Based on the evaluation of currently-available barcode data for Georgian fishes, we highlighted major gaps and research needs to further progress DNA-based biodiversity studies in Georgia. Though this study lays a solid base for DNA, based biodiversity assessment and monitoring approaches, further efforts within the recently started CaBOL (Caucasus Barcode Of Life) project are needed to obtain reference data for the species still lacking DNA barcodes.

Revealing the stygobiotic and crenobiotic molluscan biodiversity hotspot in Caucasus: Part I. The phylogeny of stygobiotic Sadlerianinae Szarowska, 2006 (Mollusca, Gastropoda, Hydrobiidae) from Georgia with descriptions of five new genera and twenty-one new species.

The position of the southwestern Caucasus as a stygobiotic Mollusca hotspot is confirmed. Molecular data of stygobiotic gastropods revealed the diversity of subfamily Sadlerianinae Szarowska, 2006, inhabiting the subterranean environment of Georgia. In addition to the well-known endemic genera Pontohoratia Vinarski, Palatov & Glöer, 2014 and Motsametia Vinarski, Palatov & Glöer, 2014, five more genera were identified in northwestern Georgia as new to the science: Kartvelobia gen. nov., Imeretiopsis gen. nov., Caucasopsis gen. nov., Caucasogeyeria gen. nov., and Hausdorfenia gen. nov. Additionally, 21 new species were found to inhabit the studied area (Samegrelo, Imereti, Racha regions in Georgia).

A new pseudoscorpion genus from western Georgia (Pseudoscorpiones: Neobisiidae: Cornuroncus n. gen.), with a key to all Neobisiinae genera.

A new genus and species of epigean pseudoscorpions belonging to the subfamily Neobisiinae Chamberlin, 1930, is described from western Georgia and its diagnostic characters are illustrated. Cornuroncus n. gen. resembles the genus Roncus L. Koch, 1873 in several characters but differs by having a short dorso-distal spine on tarsus IV, and the presence of a ventral tooth on one of the claws of tarsus IV. An identification key for all valid Neobisiinae genera is provided.

Parthenogenetic vs. sexual reproduction in oribatid mite communities.

The dominance of sex in Metazoa is enigmatic. Sexual species allocate resources to the production of males, while potentially facing negative effects such as the loss of well-adapted genotypes due to recombination, and exposure to diseases and predators during mating. Two major hypotheses have been put forward to explain the advantages of parthenogenetic versus sexual reproduction in animals, that is, the Red Queen hypothesis and the Tangled Bank/Structured Resource Theory of Sex. The Red Queen hypothesis assumes that antagonistic predator-prey/ parasite-host interactions favor sex. The Structured Resource Theory of Sex predicts sexual reproduction to be favored if resources are in short supply and aggregated in space. In soil, a remarkable number of invertebrates reproduce by parthenogenesis, and this pattern is most pronounced in oribatid mites (Oribatida, Acari). Oribatid mites are abundant in virtually any soil across very different habitats, and include many sexual and parthenogenetic (thelytokous) species. Thereby, they represent an ideal model group to investigate the role of sexual versus parthenogenetic reproduction across different ecosystems and habitats. Here, we compiled data on oribatid mite communities from different ecosystems and habitats across biomes, including tropical rainforests, temperate forests, grasslands, arable fields, salt marshes, bogs, caves, and deadwood. Based on the compiled dataset, we analyzed if the percentage of parthenogenetic species and the percentage of individuals of parthenogenetic species are related to total oribatid mite density, species number, and other potential driving factors of the reproductive mode including altitude and latitude. We then interpret the results in support of either the Red Queen hypothesis or the Structured Resource Theory of Sex. Overall, the data showed that low density of oribatid mites due to harsh environmental conditions is associated with high frequency of parthenogenesis supporting predictions of the Structured Resource Theory of Sex rather than the Red Queen hypothesis.