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Georgia's rich biodiversity, as a key part of the Caucasus hotspot, makes it an ideal destination for scientific research. Nevertheless, knowledge about the species diversity of different animal groups in this area remains limited. In this article we provide information on the nine centipede species (Henia brevis (Silvestri, 1896), H. hirsuta Verhoeff, 1928, H. taurica (Sseliwanoff, 1884), Strigamia caucasia (Verhoeff, 1938), S. pusilla (Sseliwanoff, 1884), Harpolithobius spinipes Folkmanová, 1958, Lithobius antipai Matic, 1969, L. foviceps Muralevitch, 1926, and L. micropodus (Matic, 1980)) newly recorded in Georgia. Three of these species (H. brevis, H. hirsuta, and L. micropodus) are also new records for the Caucasus region. Data on their distribution, maps of the localities, and photos of the specimens studied are also given.

Abstract 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.

A new species, Psammitis abuliensis sp. nov., from Didi Abuli Mt. (Georgia, Samtskhe-Javakheti region), is described based on two male specimens. Diagnostic drawings, measurements, collecting data, diagnosis, and DNA barcodes are given. A new combination, Psammitis marmoratus (Thorell, 1875), comb. nov. (ex. Xysticus), is proposed based on the COI subunit barcode results and morphological characters.

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.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.

Due to anthropogenic influences, habitat degradation and a continuous loss of biodiversity in freshwater ecosystems are occurring on a large scale, while these ecosystems constitute invaluable natural resources. Therefore, it is essential to study and monitor freshwater ecosystems to guide conservation efforts. Freshwater ecosystems are one of the less-studied fields in Georgia. Studies about the species distribution of many taxa and/or regions carried out during the last century have not been updated for decades. Here, we report the results of an environmental DNA (eDNA) metabarcoding exercise, based on samples collected from the Rioni River, a tributary to the Black Sea and a crucial aquatic ecosystem regionally and globally. The only comprehensive review of the fish of the Rioni River dates back to 1956. We compared the eDNA-based taxonomic composition to the known faunal composition within the Rioni River and found that the eDNA-based taxonomic coverage approached 75% of the expected total fish fauna. A number of new species occurrences were also found, including the first detection of three invasive alien species (Carassius gibelio, Pseudorasbora parva, Rhinogobius lindbergi) in the Rionis River Basin and a new country record of the ninespine stickleback (genus Pungitius) for Georgia. In spite of the usefulness of the eDNA metabarcoding approach, the sparsity of the fish DNA barcode reference library for the region emerged as a limitation to this study. However, our findings still represent a great leap forward in updating fish status on the Rioni River and testing the effectiveness of the eDNA sampling for aquatic species.

Elevational gradients in species richness and species-area relationships are among the most interesting patterns in ecology and biogeography. Both patterns can be characteristic of the same system; however, current knowledge of how these patterns co-exist and how we can disentangle their contributions to biodiversity structure is insufficient. In this article, we tested the effect of elevation and area on the formation of plant species diversity patterns in the forest-free Javakheti Highlands (Georgia). Samples (170 plots) were collected within elevations of 1400-3100 m, and the diversity distribution was examined in relation to altitude, available band area, and sampling. In total, 564 species from 67 families were recorded. Plant species richness was highest at mid-elevations (1900–2200 m), irrespective of area and sampling effort. This was in line with other studies from the Caucasus indicating the generality of plant elevational diversity patterns in the region. Area was not an important predictor of species richness; however, this may be considered a result of insufficient sampling. Our study shows that more research is needed to understand the effect of area on patterns of elevational biodiversity distribution.

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.

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.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.

The fossorial sand cockroach Hemelytroblatta livida (Brunner von Wattenwyl, 1865) is reported from Georgia for the first time, with a commentary on the species composition of the Corydiinae Saussure, 1864 subfamily in the Caucasus. Collection data, pictures of the male and female, and DNA barcodes are also provided. Furthermore, information on Polyphaga aegyptiaca (Linnaeus, 1758) is provided, along with images of the male, female, and juvenile.

The representatives of the genus Ametropus (Insecta, Ephemeroptera) were recorded 80 years ago in Georgia. Due to the taxonomic uncertainty, the recorded specimens were considered potentially new taxa. Since then, no other record of Ametropus has been documented from the whole South Caucasus and its nearby regions further south. In the present contribution, we provide a new record of Ametropus from the Alazani River (Kura River Basin, Georgia). Morphological and DNA barcoding of the COI gene fragment showed that the species belongs to the widespread Holarctic species A. fragilis. Morpho-anatomical characteristics also confirmed the high degree of similarity between the newly recorded specimens and those once recorded 80 years ago. In addition, the COI barcode showed a strikingly closer relationship between Caucasian A. fragilis and North American A. neavei (3.54% divergence) compared to Caucasian and European A. fragilis populations (4.3% divergence). The observed pattern clearly indicates the need for further and more thorough revision of the Ametropus species complex in the Holarctic region.

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.Ponticola alasanicus, 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. Ponticola alasanicus 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.

The Caucasus ecoregion is one of the most important biodiversity hotspots in western Eurasia. Georgia is situated in the middle of this area, between the ranges of the Greater and Lesser Caucasus, a position providing exceptionally balanced environmental conditions for its flora and fauna. Animal groups of low mobility, such as land snails, are important indicators of the past and present geographic and climatic events shaping these unique ecosystems. We give an overview of the land snail family Clausiliidae in Georgia that includes 30 species, among which eight are endemic to this country. Their distribution areas are reassessed in the light of novel georeferenced occurrence data resulting from recent field work. Aspects of the origin, diversity, habitat requirements and conservation status of the Georgian Clausiliidae fauna are discussed. Additionally, Mucronaria (Mucronaria) kartvelica sp. n. is described from the Racha-Lechkhumi and Kvemo Svaneti Region.

Aquatic invasions are one of the major threats for freshwater ecosystems. However, in developing countries, knowledge of biological invasions, essential for the implementation of appropriate legislation, is often limited if not entirely lacking. In this regard, the identification of potentially invasive non-native species by risk screening, followed by a full risk assessment of the species ranked as higher risk, enables decision-makers to be informed about the extent of the threats posed to the recipient (risk assessment) area. In this study, 32 non-native extant and horizon fish species were screened for their risk of invasiveness under current and predicted climate conditions for the South Caucasus – a biodiversity and geopolitical hotspot that includes the countries of Armenia, Azerbaijan and Georgia. Overall, the number of very high-risk species increased from four (12.5%) under current climate conditions to 12 (37.5%) under predicted climate conditions. The highest-risk species under both conditions included the already established gibel carp Carassius gibelio and topmouth gudgeon Pseudorasbora parva, the locally translocated pikeperch Sander lucioperca and the horizon North African catfish Clarias gariepinus. Under predicted climate conditions, a very high risk of invasiveness was predicted also for the translocated three-spined stickleback Gasterosteus aculeatus and Eurasian perch Perca fluviatilis, for the already established eastern mosquitofish Gambusia holbrooki, ruffe Gymnocephalus cernua, sharpbelly Hemiculter leucisculus and Nile tilapia Orechromis niloticus, and for the horizon pumpkinseed Lepomis gibbosus and largemouth bass Micropterus salmoides. Future research on the non-native species in the South Caucasus should be conducted both country- and region-wide and should account not only for the high biodiversity, but also for the critical geopolitical situation affecting the study area.

Electronic decision-support tools are becoming an essential component of government strategies to tackle non-native species invasions. This study describes the development and application of a multilingual electronic decision-support tool for screening terrestrial animals under current and future climate conditions: the Terrestrial Animal Species Invasiveness Screening Kit (TAS-ISK). As an adaptation of the widely employed Aquatic Species Invasiveness Screening Kit (AS-ISK), the TAS-ISK question template inherits from the original Weed Risk Assessment (WRA) and related WRA-type toolkits and complies with the ‘minimum requirements’ for use with the recent European Regulation on invasive alien species of concern. The TAS-ISK consists of 49 basic questions on the species’ biogeographical/historical traits and its biological/ecological interactions, and of 6 additional questions to predict how climate change is likely to influence the risks of introduction, establishment, dispersal and impact of the screened species. Following a description of the main features of this decision-support tool as a turnkey software application and of its graphical user interface with support for 32 languages, sample screenings are provided in different risk assessment areas for one representative species of each of the main taxonomic groups of terrestrial animals supported by the toolkit: mammals, birds, reptiles, amphibians, annelids, insects, molluscs, nematodes, and platyhelminths. The highest-scoring species were the red earthworm Lumbricus rubellus for the Aegean region of Turkey and the New Zealand flatworm Arthurdendyus triangulatus for Croatia. It is anticipated that adoption of this toolkit will mirror that of the worldwide employed AS-ISK, hence allowing to share information and inform decisions for the prevention of entry and/or dispersal of (high-risk) non-native terrestrial animal species – a crucial step to implement early-stage control and eradication measures as part of rapid-response strategies to counteract biological invasions.

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 (Galba schirazensis) 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.

Chromosome number data on the Hygromiidae (Gastropoda: Stylommatophora) are summarized and reviewed briefly in the context of the phylogeny of the family. In hygromiids, the haploid chromosome numbers range from 21 to 26. It is supposed that n = 21 is the ancestral chromosome number in the family. The modal haploid number for Hygromiidae is 23. Description of karyotype in terms of chromosome number and morphology of hygromiid land snail Circassina frutis is provided for the first time. The diploid chromosome number of this species is 2n = 46. The karyotype is symmetric and consists of 21 pairs of metacentric and 2 pairs of submetacentric chromosomes. The karyotype formula is as follows: 2n = 42m + 4sm (n = 21m + 2sm). The fundamental number (FN) is 92. Chromosomes range in length from 2.53 μm for the smallest pair to 6.00 μm for the largest pair. The total length of chromosomes in diploid complement (TCL) is 170.40 ± 3.22 μm.

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.

Since 2020, the south-western Caucasus has been recognized as a hotspot of stygobiotic Mollusca diversity after revealing a large number of new, range-restricted species within the spring snail family Hydrobiidae, subfamily Sadlerianinae sensu Szarowska. Meantime, based on extensive material collected in the south-western Caucasus during the last decades, we studied members of another spring snail subfamily Belgrandiellinae Radoman, 1983. Modern integrative taxonomic work revealed hitherto unknown diversity within this subfamily in the region and further proved the importance of the south-western Caucasus as a hotspot of stygobiotic life. In particular, the subterranean environment and springs of Georgia were known to be inhabited by the genus Tschernomorica Vinarski and Palatov, 2019 with four nominal species. Our research, based on a morpho-anatomical study and genetic investigation of COI/H3 mitochondrial/nuclear markers, revealed additionally seven species and three genera new to science—Colchiella lugella gen. et sp. nov., C. nazodelavo gen. et sp. nov., C. shiksa gen. et sp. nov., C. dadiani gen. et sp. nov., Sataplia cavernicola gen. et sp. nov., Aetis starobogatovi gen. et sp. nov., and Tschernomorica kopidophora sp. nov. —to inhabit the Georgian part of south-western Caucasus. The full taxonomic description of each new taxa, along with the review of habitat characteristics and conservation status, is provided. Molecular genetics suggests that the ancestors of Caucasian Belgrandiellinae have migrated from south-western Europe, probably more than once during the late Messinian and early Pliocene periods. Later, Plio-Pleistocene climate oscillations, particularly the repeated rise and fall of the Black Sea water level, resulted in the isolation and radiation of various lineages within the Caucasus and Crimea.

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.

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).

Abstract 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 high density of oribatid mites, indicating abundance of resources, is associated with high frequency of parthenogenesis supporting predictions of the Structured Resource Theory of Sex rather than the Red Queen hypothesis.

Madatapa Lake is an oligotrophic lake located in Javakheti plateau (Southern Georgia) at 2108 m a.s.l. and is the part of the Javakheti Protected Areas since 2011. The lake have a natural volcanic origin with a surface area of 8.8 km2 and with a maximum depth of 1.7 m. The lake is historically known to be free of fish species which is believed to be a result of severe winter conditions. After the middle 90th the gibel carp [Carassius gibelio (Bloch, 1782)] was firstly appeared there. However the exact date and the source of invasion of gibel carp in the lake is unknown and currently untraceable. While gibel carp is a very successful invasive fish species in Georgian inland waters, very limited data is available on its biology, ecology and even distribution as well as its ecological impacts and economic importance in the region. For the first time, we studied the population characteristics of gibel carp in Lake Madatapa (South Georgia) with the aim to reveal the extent of its morphological variability, sex ratio, length-weight relationship and its parasitic community. One hundred and forty one individuals presented a mean weight 37.5 g (SD=53, range [13.3-471.7]) and average length of 12.5 cm (SD=2.9, range[9.4-30]). The number of lateral line scales and gill rakers varied between 29-31, and 36-47, respectively. Sex was determined in 105 specimens (the gonads of remaining specimens were either strongly damaged by the parasites or were not developed) and the sex ratio was significantly female biased (1/1.8, Chi-square test p0.05) between the regression coefficients for males and females and the slope (b) do not differ significantly from 3. A sub-sample of thirty four individuals were examined for parasitic infections and 91% (31) were infected with one or two parasites. The plerocercoids of cestoda (Ligula intestinalis L., 1758) had highest prevalence (75%), whereas only 38% of studied fishes were infected with metacercariae of digenean trematode (Diplostomum spathaceum Rud., 1819). Both parasites were detected in only 9 specimens (27%). Cestodes were found in 5 female whereas sex was not determined for other 8 specimens. In 29 specimens infected with L. intestinalis, 24 (83%) had completely degenerate gonads. From other remaining five specimens for which the sex were determined, only one was the male. In the three specimens without parasites two were females. Since we are not able to infer the difference in parasitism intensity between sexes directly, the healthy specimens have indistinguishable sex ratio as an overall population (Chi-square test p>0.05).