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

Lela Arabuli; Lali Murvanidze; Anna Faltynkova; Levan Mumladze

doi:10.3897/BDJ.12.e110201

Biodiversity Data Journal : Taxonomy & Inventories

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Taxonomy & Inventories

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

Lela Arabuli^‡, Lali Murvanidze^‡, Anna Faltynkova^§, Levan Mumladze^‡

‡ Institute of Zoology, Ilia State University, Tbilisi, Georgia

§ Mendel University in Brno, Brno, Czech Republic

Corresponding author: Lela Arabuli (lela.arabuli.1@iliauni.edu.ge)

Academic editor: Simona Georgieva

Received: 28 Jul 2023 | Accepted: 29 Nov 2023 | Published: 08 Jan 2024

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Arabuli L, Murvanidze L, Faltynkova A, Mumladze L (2024) Checklist of digeneans (Platyhelminthes, Trematoda, Digenea) of Georgia. Biodiversity Data Journal 12: e110201. https://doi.org/10.3897/BDJ.12.e110201

Abstract

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.

Keywords

biodiversity, catalogue, Caucasus, helminths, host, parasite

Introduction

Digeneans (Platyhelminthes, Trematoda, Digenea) have complex life-cycles with several hosts involved (Cribb et al. 2003). They are parasites of all vertebrate groups and of a vast variety of invertebrates (Ditrich et al. 1997, Esteban et al. 1997, Dvořák et al. 1999, Faltýnková et al. 2009, Faltýnková et al. 2016, Manga-González and Ferreras 2019). Some groups of trematodes can be detrimental to human health and cause considerable loss to livestock (Ginetsinskaya 1968, Keiser and Utzinger 2009, Beesley et al. 2017, Esteban et al. 2019, Mas-Coma et al. 2019, Rizwan et al. 2022). However, trematodes are mainly important players in the functioning of ecosystems and they are an inherent part of food webs constituting a substantial part of biomass and, because of their sequencial use of intermediate and definitive hosts, they reflect the biodiversity of free-living host animals (Hechinger and Lafferty 2005).

Therefore, knowledge of trematode diversity and distribution, their life-cycles and transmission pathways is crucial not only for understanding the epidemiology and application of preventative measures, but also for biodiversity conservation (Dobson et al. 2008, Han et al. 2016, Selbach et al. 2020). Since Georgia is situated in the centre of the Caucasus biodiversity hotspot and in an area with a high degree of endemism for both vertebrate and invertebrate fauna (Myers et al. 2000, Mumladze et al. 2019), it is of vital importance to assess the species spectrum of the parasites which occur in this region.

In Georgia, the first studies on trematodes were conducted in the 1930s and early 1940s (Kamalov 1935, Burdjanadze 1937a, Burdjanadze 1937b, Chulkova 1939, Gamtsemlidze 1941, Kirshenblat 1941, Kurashvili 1941). Concerning digeneans of medical and veterinary importance, data on the distribution and prevalence of fascioliasis and dicrocoeliasis were published by Kurashvili and Rodonaia (1954). Since the 1950s, authors mainly focused their studies on the helminth fauna of specific host groups, i.e. fish (Kurashvili et al. 1951, Qoiava 1966, Petriashvili 1971, Kurashvili et al. 1973, Kurashvili et al. 1975, Kurashvili et al. 1990, Gogebashvili and Petriashvili 2002), amphibians and reptiles (Petriashvili 1964, Petriashvili 1966, Jankarashvili 1978, Jankarashvili 1985, Petriashvili et al. 1985), birds (Kurashvili 1953a, Kurashvili 1957, Kurashvili 1961b, Kurashvili et al. 1966, Kurashvili et al. 1976) and mammals (Rodonaia 1951, Matsaberidze 1961, Matsaberidze 1966a, Matsaberidze 1966b, Rodonaia 1966a, Rodonaia 1966b, Rodonaia 1971, Matsabaridze 1976). Fragmentary records of trematodes in freshwater and terrestrial molluscs serving as intermediate hosts were provided by several authors (Natsvlishvili 1968, Chiaberashvili and Javelidze 1968, Chiaberashvili 1971a, Chiaberashvili 1971b, Chiaberashvili and Javelidze 1977, Tkachenko 1988, Tkachenko 1990). The occurrence of human fascioliasis was examined by Gigitashvili (1965), Gigitashvili (1969), Zenaishvili et al. (2004a), Zenaishvili et al. (2004b) and Semyenova et al. (2006), who studied the genetic differentiation of European and Asian populations of Fasciola hepatica revealing that the Georgian population was of European origin and providing the only DNA sequences of trematodes from Georgia. In recent years, a few annotated checklists of helminths from different host taxa have been published by Murvanidze et al. (2008a), Murvanidze et al. (2008b) and Murvanidze et al. (2018).

Despite all the above-mentioned, a summarising review providing detailed information on the diversity of trematodes in Georgia has not been published so far. Only Kurashvili (1961a) reviewed the occurrence of some of the trematode species of medical and veterinary importance; and Kurashvili et al. (1980) published a monograph with a survey of parasites of fishes from the River Mtkvari (Kura), with digeneans included; however, there are only a few records which can be regarded as truly from the Georgian territory.

Thus, the aim of the present study was to compose a critically revised checklist of the digenetic trematodes recorded from Georgia. These data will serve as a baseline for future studies on biodiversity, taxonomy, life-cycles, ecological and epidemiological aspects of trematodes in Georgia.

Materials and methods

For records of digeneans from Georgia, we searched the following databases: Google Scholar, Science Direct, Scopus and the Host-parasite database of the Natural History Museum London, using relevant keywords (‘(trematoda or digenea) and Georgia’; excluding records from Georgia, USA). Further, we searched for literary data in relevant papers and monographs. Data from monographs were checked with caution, because, in many cases, the information on geographical distribution was vague providing only general locations, for example, the rivers or mountain ranges expanding over more countries. The present checklist is based on 109 primary papers, 11 monographs and four Ph.D. theses published predominantly in Georgian or Russian language between 1935 and 2019 and our unpublished data.

We provide a parasite-host list with the parasites arranged in alphabetical order by superfamily and family; within each family, species are also listed alphabetically. The host species of each parasite are grouped by families and listed in alphabetical order by family and species. Each parasite taxon contains information on synonyms (listed under "Nomenclature" only when appearing in Georgian literature), host species, site of infection within hosts, geographical distribution and bibliographical references associated with the parasite and its hosts in Georgia. Global geographical distribution is mainly compiled from monographs of Skryabin (1947), Dubois (1968), Dubois (1970), Yamaguti (1971), Gibson et al. (2002), Niewiadomska (2003), Jones et al. (2005), Sitko et al. (2006), Bray et al. (2008) and Fauna Europea (de Jong et al. 2014), GBIF (GBIF.org 2022) and WoRMS (WoRMS 2022). The geographical localities, indicating the distribution of trematodes in Georgia, are listed in alphabetical order. Georgia is geographically divided by the Surami (Likhi) Range into two parts, Western Georgia (WG) and Eastern Georgia (EG); and the abbreviations (WG and EG) are used throughout the text to reflect this division. These regions have different climatic conditions – humid and mild climate in the Western and dry continental in the Eastern part of the country (Maruashvili 1964, Murvanidze and Mumladze 2016).

Life-cycle stages other than the sexual adults are indicated in parentheses following the host category, i.e. intramolluscan stages in first intermediate mollusc hosts comprising sporocysts, rediae and cercariae (the latter are asexual free-living stages emerging from the snail first intermediate hosts); and metacercariae encysted or unencysted in tissue of second intermediate or paratenic hosts.

Species names were checked for their current state and, for the nomenclature of all taxa, we refer to the Fauna Europea (de Jong et al. 2014), GBIF (GBIF.org 2022), WoRMS (WoRMS 2022), Catalogue of Life (Gibson et al. 2023) and Molluscabase (2023); for the classification of trematodes at family level, we follow the “Keys to the Trematoda” (Gibson et al. 2002, Jones et al. 2005, Bray et al. 2008) and, at the superfamily level, we follow WoRMS (2022) and Catalogue of Life (Gibson et al. 2023).

Trematodes not identified to species level were included in the checklist as well; in ambiguous cases, when it was impossible to discern the identity of the records (Echinochasmus sp., Cephalogonimus sp. and Telorchis sp.), they were listed with 'spp.'. However, we did not include taxa of doubtful identity, but we listed them in the Discussion with an explanation of their status and history; the names of these taxa are provided exactly as they occur in literature. Records of cercarial names, which are regarded as provisional, because they do not conform to the taxonomy of adult digeneans, were not included in the checklist either. However, to have a complete view on the potential biodiversity state of trematodes in Georgia, they are included in the Discussion provided with remarks on their occurrence.

Previously unpublished trematode specimens included in the checklist are deposited in the collections of the Institute of Zoology, Ilia State University.

Annotated checklist of Digenea of Georgia

Phylum Minot, 1876

Class Trematoda Rudolphi, 1808

Subclass Digenea Carus, 1863

Order Diplostomida Olson, Cribb, Tkach, Bray & Littlewood, 2003

Superfamily Brachylaimoidea Joyeux & Foley, 1930

Family Brachylaimidae Joyeux & Foley, 1930

Genus Brachylaima Dujardin, 1843

Brachylaima fulvum Dujardin, 1843

Nomenclature:

Panopistus europaeus Soltys, 1952

Parasite of:

mammals - Soricidae: Sorex araneus, S. raddei.

Site of infection: small intestine.

Distribution:

Occurring in Europe; in Georgia: EG: Borjomi; WG: Batumi, Gagra, Oni, Tskaltubo reported by Matsaberidze (1966a), Matsaberidze (1966b), Matsaberidze (1967), Rodonaia (1971), Matsabaridze (1976) and Kurashvili (1984b).

Brachylaima fuscata (Rudolphi, 1819) Joyeux, Baer & Timon-David, 1932

Nomenclature:

Genus Pseudoleucochloridium Pojmanska, 1959

Pseudoleucochloridium soricis (Soltys, 1952) Pojmanska, 1959

Nomenclature:

Leucochloridium soricis Soltys, 1952

Parasite of:

mammals - Soricidae: Sorex araneus, S. minutus, S. raddei.

Site of infection: small intestine.

Distribution:

Occurring in Europe and Asia; in Georgia: EG: Borjomi; WG: Samtredia, Sokhumi, Qutaisi reported by Matsaberidze (1967), Rodonaia (1971), Matsabaridze (1976) and Kurashvili (1984b).

Superfamily Diplostomoidea Poirier, 1886

Family Cyathocotylidae Mühling, 1898

Genus Mesostephanus Lutz, 1933

Mesostephanus appendiculatus (Ciurea, 1916) Lutz, 1935

Parasite of:

Alaria alata (Goeze, 1782) Krause,1914

Parasite of:

Nomenclature:

Neascus brevicaudatus (von Nordmann, 1832) Hughes, 1928

Parasite of:

fishes (metacercariae) - Esocidae: Esox lucius.

Site of infection: musculature, skin.

Distribution:

With Holarctic distribution; in Georgia: EG: Bazaleti Lake, Rivers: Alazani, Iori, Mtkvari; WG: Paliastomi Lake reported by Chiaberashvili (1968), Chernova (1973) and Murvanidze et al. (2018).

Posthodiplostomum cuticola (von Nordmann, 1832) Dubois, 1936

Parasite of:

fishes (metacercariae) - Cyprinidae: Abramis brama, Alburnus chalcoides, Ballerus sapa, Chondrostoma cyri, Cyprinus carpio, Leuciscus aspius, Luciobarbus capito, Scardinius erythrophthalmus, Tinca tinca, Vimba vimba.

Site of infection: fins, gills, skin.

Distribution:

Occurring in Europe and Asia; in Georgia: EG: River Mtkvari, Sagarejo reported by Kurashvili and Petriashvili (1977), Kurashvili et al. (1980), Kurashvili et al. (1983a), Kurashvili et al. (1990), Gogebashvili and Petriashvili (2002) and Murvanidze et al. (2018).

Posthodiplostomum sp.

Occurring in the Palaearctic Region, Africa; in Georgia: EG: Bazaleti Lake, Jinvali, Khazbegi, surroundings of Tbilisi; WG: Khobi, surroundings of Batumi, Zugdidi reported by Petriashvili (1966) and Jankarashvili (1985).

Superfamily Schistosomatoidea Stiles & Hassall, 1898

Family Clinostomidae Lühe, 1901

Distribution:

In the Holarctic Region, India, Southeast Asia, Africa; in Georgia: EG: Dusheti – Bazaleti Lake, Tbilisi – Digomi, Tsalka reported by Kurashvili (1948), Kurashvili (1950), Kurashvili (1953a), Kurashvili (1957) and Kurashvili (1961a).

Family Schistosomatidae Stiles & Hassall, 1898

Genus Bilharziella Looss, 1899

Bilharziella polonica (Kowalewski, 1895) Looss, 1899

Parasite of:

birds - Anatidae: Anas crecca, A. penelope, A. plathyrynchos f. domestica.

Site of infection: abdominal cavity, blood vessels of liver.

Distribution:

Holarctic distribution; in Georgia: EG: Dusheti – Bazaleti Lake, Gardabani – Tbilisi Sea, Kumisi Lake, Tsalka; WG: Mestia – Lenjeri, Zugdidi reported by Kurashvili et al. (1976) and Kurashvili (1984b).

Genus Dendritobilharzia Skrjabin & Zakharov, 1920

Dendritobilharzia pulverulenta (Braun, 1901) Skrjabin, 1924

Parasite of:

birds - Anatidae: Anas crecca, A. penelope, A. plathyrynchos f. domestica, Anser anser f. domestica.

Site of infection: blood vessels – large veins.

Distribution:

Holarctic distribution; in Georgia: EG: Gardabani – Tbilisi Sea, Lagodekhi, Lisi Lake; WG: Surroundings of Poti reported by Kurashvili et al. (1976).

Order Plagiorchiida La Rue, 1957

Superfamily Azygioidea Lühe, 1909

Distribution:

Occurring in Europe, Asia; in Georgia: River Mtkvari; WG: Lakes: Didi Narionali, Paliastomi reported by Chernova (1973), Kurashvili et al. (1980) and Murvanidze et al. (2018).

Superfamily Echinostomatoidea Looss, 1902

Family Cyclocoelidae Stossich, 1903

Genus Harrahium Witenberg, 1926

Harrahium halli (Harrah, 1922) Witenberg, 1926

Nomenclature:

Cyclocoelum halli Harrah, 1922

Parasite of:

birds - Anatidae: Anas querquedula.

Site of infection: abdominal cavity, air sacs of lungs.

Distribution:

Occurring in North America; in Georgia: EG: Mtkvari River, Mtskheta reported by Kurashvili (1957) and Kurashvili (1961a).

Genus Hyptiasmus Kossack, 1911

Hyptiasmus magniproles Witenberg, 1928

Parasite of:

birds - Anatidae: Mergellus albellus.

Site of infection: infraorbital cavity, nasal cavity.

Distribution:

Occurring in Asia; in Georgia: EG: Marneuli reported by Kurashvili (1957).

Genus Morishitium Witenberg, 1928

Morishitium bivesiculatum (Prudhoe, 1944) Yamaguti, 1958

Nomenclature:

Distribution:

Occurring in the Holarctic region; in Georgia: EG: Tbilisi; WG: Batumi reported by Burdjanadze (1937a), Gamtsemlidze (1941), Burjanadze (1943), Kurashvili (1961a), Kurashvili et al. (1980) and Kurashvili (1984b).

Echinochasmus spp.

Parasite of:

fishes (metacercariae) - Cyprinidae: Luciobarbus escherichii, Squalius cephalus.

mollusc (intramolluscan stages) - Melanopsidae: Melanopsis praemorsa.

Site of infection: gills.

Distribution:

Cosmopolitan distribution; in Georgia: WG: Rivers: Pichori, Rioni reported by Chiaberashvili (1962b), Olenev and Dobrovolsky (1975), Kurashvili (1984a) and Manafov (2011).

Genus Stephanoprora Odhner, 1902

Stephanoprora pseudoechinata (Olsson, 1876) Yamaguti, 1958

Nomenclature:

Mesorchis pseudoechinatus (Olsson, 1876) Dietz, 1909

Parasite of:

birds - Anatidae: Aythya nyroca; Laridae: Hydrocoloeus minutus, Larus canus.

Site of infection: small intestine.

Distribution:

Occurring in the Holarctic, Africa; in Georgia: EG: Kumisi Lake, Lagodekhi, Marneuli; WG: River Kaparchina, Rioni Valley – Poti, Sokhumi reported by Kurashvili (1950), Kurashvili (1953a), Kurashvili (1957), Kurashvili (1961a) and Kurashvili (1984b).

Family Echinostomatidae Looss, 1899

Genus Chaunocephalus Dietz, 1909

Chaunocephalus ferox subsp. orientalis Baschkirova, 1941

Parasite of:

Parasite of:

birds - Anatidae: Anas acuta, A. crecca, A. platyrhynchos, A. platyrhynchos f. domestica, Anser anser, Marmaronetta angustirostris; Ardeidae: Botaurus stellaris; Phasianidae: Gallus gallus f. domestica.

molluscs (intramolluscan stage) - Planorbidae: Ancylus fluviatilis, Planorbis planorbis.

Site of infection: caecum, small intestine.

Distribution:

Cosmopolitan distribution; in Georgia: EG: Dusheti – Bazaleti Lake, Gardabani, Marneuli, Sartichala, Tsalka; WG: Abasha, Khobi, Lanchkhuti – Mamati, Poti – River Kaparchina, Samtredia, Senaki, Tsageri, Tsalenjikha reported by Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967), Chiaberashvili (1971a), Kurashvili et al. (1976) and Kurashvili (1984b).

Genus Echinostoma Rudolphi, 1809

Echinostoma miyagawai Ishii, 1932

Nomenclature:

Echinostoma robustum Yamaguti, 1935

Parasite of:

birds - Anatidae: Anas platyrhynchos, A. platyrhynchos f. domestica, Anser anser f. domestica, Netta rufina; Columbidae: Spilopelia chinensis, Streptopelia turtur; Phasianidae: Gallus gallus f. domestica, Meleagris gallopavo.

molluscs (intramolluscan stage) - Lymnaeidae: Ampullaceana lagotis.

Site of infection: hepatopancreas, intestine.

Distribution:

Occurring in Europe, Asia; in Georgia: EG: Dedoflistskaro – Qvemo qedi, Dusheti – Bazaleti Lake, Gardabani – Jandari Lake, Marneuli, Mtskheta, Sartichala, Tbilisi – Samgori, Lisi Lake, Shuakhevi, Tetriwkaro – Koda, Tsalka; WG: Paliastomi Lake, Qareli, Rioni Valley, Samtredia, Sokhumi, Terjola, Zugdidi reported by Kurashvili (1941), Kurashvili (1953a), Chiaberashvili (1954), Chiaberashvili (1957), Kurashvili (1961a), Japaridze (1962), Japaridze and Savateeva (1967), Chiaberashvili (1971a), Kurashvili et al. (1976) and Kurashvili (1984b).

Echinostoma paraulum Dietz, 1909

Parasite of:

birds - Anatidae: Anas clypeata, A. platyrhynchos, A. platyrhynchos f. domestica, Anser anser f. domestica, Aythya ferina, Cygnus cygnus, Netta rufina; Columbidae: Columba livia.

molluscs (intramolluscan stage) - Lymnaeidae: Ampullaceana balthica.

Site of infection: hepatopancreas, intestine.

Distribution:

Occurring in Europe, Asia; in Georgia: EG: Gardabani, Sartichala, Tskhinvali; WG: Lanchkhuti, lowland of Rioni, Samtredia, surroundings of Poti, Tetritskaro, Zugdidi reported by Kurashvili (1950), Kurashvili (1953a), Chiaberashvili (1954), Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967), Chiaberashvili (1971a), Kurashvili et al. (1976) and Kurashvili (1984b).

Echinostoma revolutum (Fröhlich, 1802) Looss, 1899

Parasite of:

birds - Anatidae: Anas clypeata, A. platyrhynchos, A. platyrhynchos f. domestica, Anser anser f. domestica, Mergellus albellus, Mergus merganser, M. serrator; Ardeidae: Ardea cinerea; Phasianidae: Gallus gallus f. domestica.

molluscs (intramolluscan stage) - Lymnaeidae: Ampullaceana balthica, A. lagotis, Peregriana peregra, Galba truncatula; Planorbidae: Planorbis planorbis.

Site of infection: intestine.

Distribution:

Occurring in the Holarctic Region; in Georgia: EG: Bazaleti Lake, Sarthichala, Tbilisi – Samgori, River Iori; WG: Batumi, Lentekhi, Mestia, Paliastomi Lake, Samtredia reported by Kirshenblat (1941), Burjanadze (1943), Chiaberashvili (1954), Kurashvili (1957), Kurashvili (1961a), Japaridze (1962), Japaridze and Savateeva (1967), Chiaberashvili (1971a), Kurashvili et al. (1973), Kurashvili et al. (1983b) and Kurashvili (1984b).

Echinostoma stantschinskii Semenov, 1927

Parasite of:

Moliniella anceps (Molin, 1859) Hübner, 1939

Parasite of:

molluscs (intramolluscan stage) - Lymnaeidae: Lymnaea stagnalis.

Site of infection: hepatopancreas.

Distribution:

Occurring in Europe; in Georgia: WG: Madatapa Lake reported by Arabuli et al. (2015).

Genus Neopetasiger Baschkirova, 1941

Neopetasiger megacanthus (Kotlán, 1922) Tkach, Kudlai & Kostadinova, 2016

Nomenclature:

Petasiger megacanthus (Kotlán, 1922) Pande, 1939

Parasite of:

Pegosomum petrowi Kurashvili, 1949

Parasite of:

birds - Ardeidae: Egretta alba.

Site of infection: gallbladder.

Distribution:

Records exist only from Georgia; in Georgia: EG: Kumisi Lake, Lagodekhi, Marneuli reported by Kurashvili (1949), Kurashvili (1950), Kurashvili (1953a), Kurashvili (1957) and Kurashvili (1961a).

Pegosomum skrjabini Shakhtakhtinskaya, 1949

Recorded in Georgia only; in Georgia: WG: River Rioni reported by Tichomirov (1976), Ataev (1991) and Ataev and Dobrovolskiy (1992).

Family Psilostomidae Looss, 1900

Genus Psilochasmus Lühe, 1909

Psilochasmus longicirratus Skrjabin, 1913

Parasite of:

birds - Anatidae: Anas acuta, A. clypeata, Anas platyrhynchos f. domestica, Anser anser f. domestica, Aythya nyroca.

Site of infection: caecum, intestine.

Distribution:

Europe, North America; in Georgia: EG: Dusheti – Bazaleti Lake, Tsalka – Khrami Reservoir. WG: Gurianta, Khobi, Ozurgeti – Gomi, Samtredia, surroundings of Poti reported by Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967), Kurashvili et al. (1976) and Kurashvili (1984b).

Psilochasmus oxyurus (Creplin, 1825) Lühe, 1909

Parasite of:

birds - Anatidae: Anas acuta, Anas plathyrynchos, Tadorna tadorna; Phasianidae: Gallus gallus f. domestica.

Site of infection: intestine.

Distribution:

North America, Europe, Asia (Iraq); in Georgia: EG: Lagodekhi – Alazani Valley; WG: Khobi, Rioni Valley, Samtredia, Senaki, surroundings of Poti reported by Kurashvili (1953a), Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967), Kurashvili et al. (1976) and Kurashvili (1984b).

Family Typhlocoelidae Harrah, 1922

Genus Tracheophilus Skrjabin, 1913

Tracheophilus sisowi Skrjabin, 1913

Parasite of:

birds - Anatidae: Anas acuta, A. clypeata, A. platyrhynchos, A. platyrhynchos f. domestica, A. querquedula, Anser anser f. domestica, Aythya fuligula.

Site of infection: abdominal cavity, lungs, trachea.

Distribution:

Holarctic distribution; in Georgia: EG: Bolnisi, Dusheti – Bazaleti Lake, Gardabani, Marneuli, Tetritskaro. WG: Samtredia reported by Burjanadze (1943), Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967) and Kurashvili et al. (1976).

Superfamily Gorgoderoidea Looss, 1901

Family Allocreadiidae Looss, 1902

Dicrocoelium dendriticum (Rudolphi, 1819) Looss, 1899

Nomenclature:

Dicrocoelium lanceatum Stiles & Hassall, 1898

Parasite of:

mammals - Bovidae: Bos taurus, Capra hircus, Ovis aries; Cervidae: Cervus, Capreolus capreolus; Leporidae: Lepus; Suidae: Sus scrofa; Ursidae: Ursus arctos; Muridae: Chionomys.

molluscs (intramolluscan stage) - Enidae: Georginapaeus hohenackeri; Geomitridae: Xeropicta derbentina; Hygromiidae: Harmozica ravergiensis.

Site of infection: gallbladder, hepatic ducts, liver.

Distribution:

Cosmopolitan distribution; in Georgia: EG: Akhmeta, Borjomi, Dedoflistskaro, Dusheti, Kazbegi, Lagodekhi, Tianeti, Highland regions of southern and northern Georgia; WG: Akhalqalaqi, Akhaltsikhe, Batumi, Khobi, Lentekhi, Samtredia, Tsalenjikha, Zugdidi reported by Burdjanadze (1937b), Gamtsemlidze (1941), Kurashvili and Rodonaia (1954), Qoiava (1956a), Qoiava (1961), Kurashvili (1961a), Rodonaia (1962), Rodonaia (1966a), Rodonaia (1966b), Natsvlishvili (1968), Rodonaia (1971) and Matsabaridze (1976).

Dicrocoelium macrostomum Odhner, 1910

Parasite of:

birds - Phasianidae: Coturnix coturnix; Numididae: Numida meleagris.

Site of infection: hepatic ducts, liver.

Distribution:

Occurring in Africa; in Georgia: WG: Samtredia reported by Kurashvili (1957), Kurashvili (1961a) and Kurashvili (1984b).

Family Haplosplanchnidae Poche, 1926

Genus Haplosplanchnus Looss, 1902

Haplosplanchnus pachysoma (Eysenhardt, 1829) Looss, 1902

Parasite of:

fishes - Mugilidae: Chelon auratus, Mugil cephalus.

Site of infection: intestine.

Distribution:

Occurring in Europe, Africa, Oceanian Regions; in Georgia: WG: Paliastomi Lake reported by Chernova (1973) and Murvanidze et al. (2018).

Genus Schikhobalotrema Skrjabin & Guschanskaya, 1955

Schikhobalotrema sparisomae (Manter, 1938) Skrjabin & Guschanskaya, 1955

Parasite of:

fishes - Mugilidae: Chelon auratus.

Site of infection: digestive tract.

Distribution:

Holarctic distribution; in Georgia: WG: Paliastomi Lake reported by Chernova (1973) and Murvanidze et al. (2018).

Superfamily Hemiuroidea Looss, 1899

Family Hemiuridae Looss, 1899

Genus Aphanurus Looss, 1907

Aphanurus stossichii (Monticelli, 1891) Looss, 1907

Parasite of:

Genus Aponurus Looss, 1907

Aponurus tschugunovi Issaitschikov, 1927

Parasite of:

fishes - Mugilidae: Mugil cephalus.

Site of infection: no data

Distribution:

Occurring in the Black sea, European part of Russia; in Georgia: WG: Paliastomi Lake reported by Chernova (1973).

Genus Lecithaster Lühe, 1901

Lecithaster confusus Odhner, 1905

Nomenclature:

Lecithaster musteli Srivastava, 1966

Parasite of:

fishes - Cyprinidae: Alburnus derjugini; Clupeidae: Alosa immaculata.

Site of infection: intestine.

Distribution:

Occurring in the Holarctic Region, Africa; in Georgia: WG: Paliastomi Lake reported by Chernova (1977) and Murvanidze et al. (2018).

Lecithaster tauricus Pigulewsky, 1938

Parasite of:

fishes - Esocidae: Esox lucius; Percidae: Perca fluviatilis.

Site of infection: intestine.

Distribution:

Occurring in the Black Sea; in Georgia: WG: Paliastomi Lake reported by Chernova (1977) and Murvanidze et al. (2018).

Superfamily Microphalloidea Ward, 1901

Distribution:

Occurring in the Palaearctic; in Georgia: WG: Paliastomi Lake reported by Chernova (1973).

Family Lecithodendriidae Lühe, 1901

Occurring in Europe; in Georgia: WG: Tkibuli reported by Matsaberidze (1963), Matsaberidze and Khotenovskii (1967) and Matsabaridze (1976).

Genus Ophiosacculus Macy, 1935

Ophiosacculus mehelyi (Mödlinger, 1930) Macy, 1935

Nomenclature:

Ophiosacculus eptesicus Matsaberidze & Chotenowsky, 1965

Parasite of:

Prosthodendrium ascidia (van Beneden, 1873) Bhalerao, 1936

Parasite of:

mammals - Vespertilionidae: Eptesicus serotinus, Pipistrellus kuhlii, P. nathusii, Rhinolophidae: Rhinolophus ferrumequinum.

Site of infection: small intestine.

Distribution:

Occurring in Europe; in Georgia: EG: Dedoflistskaro – Shiraqi, Gareji, Sagarejo-Gombori; WG: Tkibuli reported by Matsaberidze and Khotenovskii (1967) and Matsabaridze (1976).

Prosthodendrium chilostomum (Mehlis, 1831) Travassos, 1921

Parasite of:

mammals - Vespertilionidae: Eptesicus serotinus, Pipistrellus nathusii; Rhinolophidae: Rhinolophus ferrumequinum.

Site of infection: small intestine.

Distribution:

Occurring in Europe; in Georgia: EG: Mtskheta, Natakhtari, Sagarejo-Gombori; WG: Tkibuli reported by Matsaberidze and Khotenovskii (1967) and Matsabaridze (1976).

Prosthodendrium parvouterus (Bhalerao, 1926) Skrjabilovich, 1948

Parasite of:

mammals - Vespertilionidae: Eptesicus serotinus, Nyctalus noctula.

Site of infection: small intestine.

Distribution:

Occurring in the Holarctic Region, Africa; in Georgia: EG: Borjomi, Dedoflistskaro – Shiraqi, Sagarejo-Gombori; WG: Tkibuli reported by Matsaberidze and Khotenovskii (1967) and Matsabaridze (1976).

Prosthodendrium skrjabini (Shaldybin in Skarbilovich, 1948)

Nomenclature:

Genus Pleurogenes Looss, 1896

Pleurogenes claviger (Rudolphi, 1819) Looss, 1896

Parasite of:

amphibians - Ranidae: Pelophylax ridibundus, Rana macrocnemis.

Site of infection: intestine.

Distribution:

Occurring in Europe; in Georgia: WG: Bebesiri Lake, Gali, Gudauta reported by Chiaberashvili and Mchedlidze (1961), Kurashvili (1984b), Petriashvili et al. (1985) and Murvanidze et al. (2008a).

Pleurogenes intermedius Issaitschikov, 1926

Skrjabinopsolus semiarmatus (Molin, 1858) Ivanov in Ivanov & Murygin, 1937

Nomenclature:

Skrjabinopsolus acipenseris Ivanov & Murygin, 1937

Skrjabinopsolus minor Bychowskaya-Pavlovskaya & Mikailov, 1969

Parasite of:

fishes - Acipenseridae: Acipenser persicus, A. stellatus, Huso huso.

Site of infection: oesophagus, intestine, stomach.

Distribution:

Occurring in the Holarctic; in Georgia: River Mtkvari; WG: River Rioni (Orpiri Village), surroundings of Batumi reported by Chulkova (1939), Kurashvili et al. (1980) and Murvanidze et al. (2018).

Family Lissorchiidae Magath, 1917

Occurring in Europe; in Georgia: River Mtkvari; WG: Lakes: Japana, Paliastomi reported by Chernova (1973), Kurashvili et al. (1980) and Murvanidze et al. (2018).

Superfamily Opecoeloidea Ozaki, 1925

Family Opecoelidae Ozaki, 1925

Site of infection: intestine.

Distribution:

Occurring in Europe; in Georgia: River Mtkvari reported by Kurashvili et al. (1980).

Superfamily Opisthorchioidea Looss, 1899

Family Cryptogonimidae Ward, 1917

Genus Timoniella Rebecq, 1960

Timoniella imbutiformis (Molin, 1859) Brooks, 1980

Parasite of:

fishes - Clupeidae: Alosa tanaica.

Site of infection: no data.

Distribution:

Occurring in the Mediterranean Region, Caspian Sea, Black Sea, northeast Atlantic; in Georgia: WG: Paliastomi Lake reported by Burjanadze (1943) and Murvanidze et al. (2018).

Family Heterophyidae Leiper, 1909

Genus Ascocotyle Looss, 1899

Ascocotyle italica Alessandrini, 1906

Nomenclature:

Parascocotyle italica (Alessandrini, 1906) Price, 1932

Parasite of:

mammals - Canidae: Canis lupus familiaris.

Site of infection: intestine.

Distribution:

Occurring in Europe, Africa; in Georgia: WG: Kolkheti Lowland reported by Burjanadze (1943) and Kurashvili (1984b).

Ascocotyle longa Ransom, 1920

Nomenclature:

Parascocotyle longa (Ransom, 1920) Stunkard & Haviland, 1924

Metascocotyle witenbergi Ciurea, 1933

Parasite of:

Family Diplodiscidae Cohn, 1904

Genus Diplodiscus Diesing, 1836

Diplodiscus mehrai Pande, 1937

Parasite of:

amphibians - Ranidae: Pelophylax ridibundus.

Site of infection: large intestine, rectum.

Distribution:

Occurring in Europe and India; in Georgia: EG: Aragvi River Basin, Bazaleti Lake, Tbilisi – Samgori reported by Chiaberashvili and Mchedlidze (1961), Petriashvili (1964), Burtikashvili and Getzadze (1981), Petriashvili et al. (1985), Sey (2001) and Murvanidze et al. (2008a).

Diplodiscus subclavatus (Pallas, 1760) Diesing, 1836

Parasite of:

reptiles - Colubridae: Natrix natrix.

amphibians - Bufonidae: Bufotes viridis; Ranidae: Pelophylax ridibundus, Rana macrocnemis.

mollusc (intramolluscan stage) - Planorbidae: Planorbis planorbis.

Site of infection: large and small intestine, rectum.

Distribution:

Occurring in Europe, Asia, Africa; in Georgia: EG: Aragvi River Basin, Bazaleti Lake, Borjomi, Jinvali, Kazbegi, Kodjori, Samgori, Sartichala – River Iori, surroundings of Tbilisi; WG: Gali, Khobi, Ozurgeti, Senaki, surroundings of Batumi, Tkibuli Reservoir, Zugdidi reported by Chiaberashvili and Mchedlidze (1961), Petriashvili (1964), Petriashvili (1966), Chiaberashvili (1971a), Burtikashvili and Getzadze (1981), Giorgadze (1985), Jankarashvili (1985), Petriashvili et al. (1985), Sey (2001), Murvanidze et al. (2008a) and Murvanidze et al. (2008b).

Family Paramphistomidae Fischoeder, 1901

Paramphistomum skrjabini Popova, 1927

Parasite of:

mammals - Bovidae: Bos taurus, Bubalus bubalis.

molluscs (intramolluscan stage) - Planorbidae: Planorbis planorbis.

Site of infection: rumen.

Distribution:

Reported from Georgia only; in Georgia: EG: Eastern Georgia reported by Burjanadze (1943), Rodonaia (1960) and Rodonaia (1971).

"fam. Paramphistomidae" gen. sp.

Parasite of:

molluscs (intramolluscan stage) - Planorbidae: Planorbis planorbis.

Site of infection: no data.

Distribution:

Cosmopolitan distribution; in Georgia: EG: Gori, Khashuri, Lagodekhi, Manglisi, Qareli, Tetritskaro, Tsalka reported by Matsaberidze et al. (1989).

Superfamily Plagiorchioidea Lühe, 1901

Family Brachycoeliidae Looss, 1899

Genus Brachycoelium Dujardin, 1845

Brachycoelium salamandrae (Frölich, 1789) Lühe, 1909

Parasite of:

reptiles - Salamandridae: Mertensiella caucasica.

Site of infection: small intestine.

Distribution:

Occurring in Europe, North America, North Africa; in Georgia: EG: Bakuriani reported by Petriashvili et al. (1985) and Murvanidze et al. (2008b).

Family Cephalogonimidae Looss, 1899

reptiles - Colubridae: Natrix natrix, N. tesselata.

Site of infection: intestine.

Distribution:

Occurring in the Palaearctic, Africa; in Georgia: EG: Borjomi, Jinvali, Kazbegi, surroundings of Tbilisi; WG: Khobi, Kulevi, surroundings of Batumi, Zugdidi reported by Sharpilo (1962), Kurashvili (1984b), Jankarashvili (1985) and Kurashvili et al. (1991).

Family Orientocreadiidae Yamaguti, 1958

Genus Orientocreadium Tubangui, 1931

Orientocreadium pseudobagri Yamaguti, 1934

Nomenclature:

Paratormopsolus siluri Dubinina & Bychowsky, 1954

Orientocreadium siluri (Dubinina & Bychowsky in Skrjabin, 1954) Yamaguti, 1958

Parasite of:

fishes - Siluridae: Silurus glanis.

Site of infection: intestine.

Distribution:

Known from Asia (Iraq); in Georgia: WG: Bebesiri Lake, River Tekhura reported by Chiaberashvili (1962b) and Murvanidze et al. (2018).

Family Plagiorchiidae Lühe, 1901

Genus Haplometra Looss, 1899

Haplometra brevicaeca Timon-David, 1962

Parasite of:

Skrjabinoeces similis Looss, 1899

Parasite of:

molluscs (intramolluscan stage) - Planorbidae: Planorbis planorbis.

Site of infection: no data.

Distribution:

Palaearctic distribution; in Georgia: EG: Rustavi reported by Tkachenko (1990).

Family Telorchiidae Looss, 1899

Genus Opisthioglyphe Looss, 1899

Opisthioglyphe ranae (Frölich, 1791) Looss, 1907

Parasite of:

reptiles - Colubridae: Natrix natrix.

amphibians - Bufonidae: Bufo bufo, Bufotes viridis; Hylidae: Hyla arborea; Ranidae: Pelophylax ridibundus, Rana macrocnemis.

molluscs (intramolluscan stage) - Lymnaeidae: Galba palustris, Peregriana peregra.

Site of infection: gall bladder, large intestine, small intestine.

Distribution:

Occurring in Europe, Asia (Iraq), North Africa; in Georgia: EG: Aragvi River Basin, Borjomi, Jinvali, Kazbegi, Kodjori, Samgori, Surroundings of Tbilisi, Tbilisi – Botanic Garden; WG: Batumi, Bebesiri Lake, Gali, Khobi, Lanchkhuti, Ozurgeti, Samtredia, Senaki, Tkibuli Reservoir, Zugdidi reported by Chiaberashvili and Mchedlidze (1961), Javelidze (1964), Burtikashvili and Getzadze (1981), Kurashvili (1984a), Kurashvili (1984b), Giorgadze (1985), Jankarashvili (1985), Petriashvili et al. (1985), Kurashvili et al. (1991) and Murvanidze et al. (2008a).

Opisthioglyphe rastellus (Olsson, 1876) Looss, 1907

Nomenclature:

Dolichosaccus rastellus (Olsson, 1876) Travassos, 1930

Parasite of:

amphibians - Ranidae: Pelophylax ridibundus, Rana macrocnemis.

Parasite of:

reptiles - Lacertidae: Lacerta strigata.

molluscs (intramolluscan stage) - Lymnaeidae: Radix euphratica.

Site of infection: intestine, liver.

Distribution:

In North and South America, Europe, Asia; in Georgia: EG: Lakes: Jandari, Turtle reported by Kurashvili et al. (1975) and Arabuli, unpublished data 2021.

Superfamily Pronocephaloidea Looss, 1899

Family Notocotylidae Lühe, 1909

Genus Notocotylus Diesing, 1839

Notocotylus attenuatus (Rudolphi, 1809) Kossack, 1911

Parasite of:

birds - Anatidae: Anas clypeata, A. plathyrynchos, A. plathyrynchos f. domestica, Anser anser, Aythya fuligula, A. ferina, Cygnus cygnus.

molluscs (intramolluscan stage) - Lymnaeidae: Lymnaea stagnalis, Peregriana peregra, Radix auricularia.

Site of infection: caecum, large intestine, rectum (birds), hepatopancreas (molluscs).

Distribution:

Cosmopolitan distribution; in Georgia: EG: Bolnisi, Dedoplistskaro – Qvemo qedi, Dmanisi, Gardabani, Marneuli, Tetritskaro, Tsalka – Khrami Reservoir, Lakes: Bazaleti, Paravani; WG: Lakes: Bebesiri, Madatafa; Samtredia, surroundings of Poti – Paliastomi Lake reported by Burjanadze (1943), Kurashvili (1957), Kurashvili (1961a), Japaridze and Savateeva (1967), Kurashvili et al. (1976), Kurashvili (1984a), Kurashvili (1984b), Tkachenko (1988) and Arabuli et al. (2015).

Notocotylus ephemera (Nitzsch, 1817) Harwood, 1939

Parasite of:

molluscs (intramolluscan stage) - Planorbidae: Planorbis planorbis.

Site of infection: no data.

Distribution:

Occurring in Europe; in Georgia: EG: Gori – Khidistavi reported by Chiaberashvili (1971a).

Notocotylus noyeri Joyeux, 1922

Parasite of:

mammals - Cricetidae: Arvicola terrestris.

Site of infection: small intestine.

Distribution:

Occurrence in Europe; in Georgia: EG: Gori – Khidistavi reported by Matsabaridze (1976).

Notocotylus ponticus Tshiaberashwili, 1964

Parasite of:

molluscs (intramolluscan stage) - Bithyniidae: Bithynia tentaculata.

Site of infection: no data.

Distribution:

Recorded only in Georgia; in Georgia: WG: Bebesiri Lake reported by Chiaberashvili (1971b), Kurashvili (1984a) and Cribb (1991).

Notocotylus zduni Chiaberashvili & Dzavelidze, 1968

Parasite of:

molluscs (intramolluscan stage) - Neritidae: Theodoxus fluviatilis.

Site of infection: no data.

Distribution:

Occurrence in Europe; in Georgia: WG: West Georgia reported by Chiaberashvili and Javelidze (1968).

Analysis

Results

Based on data from literature and our new records, digeneans in Georgia are currently represented by 186 taxa, of which 173 are identified to species level, belonging to 108 genera, 47 families and 17 superfamilies. The majority of trematode species comprised adult stages (160 species), a small proportion being made up by cercariae (33) or metacercariae (24) in their first and second intermediate hosts, respectively. For 28 species (15%), at least two life-cycle stages were recorded (adult in combination with cercariae/metacercariae). Birds and mammals were recorded as definitive hosts only, while the other groups were recorded as definitive and also as second intermediate hosts - fish hosted 32 species as adults and 18 as metacercariae, amphibians hosted 23 species as adults and two as metacercariae and reptiles hosted nine species as adults and three as metacercariae. A total of 35 trematode species were recorded in molluscs, which were used as first or second intermediate hosts. Predominantly, freshwater digeneans were recorded (154 species), while a much lower number of marine (12) and terrestrial (21 species) digeneans was found. A total of 202 free-living vertebrate and mollusc species were recorded as hosts for digeneans – fish of 57 species, amphibians of six species, reptiles of four species, birds of 74 species, mammals of 40 species and molluscs of 21 species.

Discussion

In the present study, we compiled data on digenean trematode fauna of Georgia, which is represented by 186 species. Amongst these, 37 species belong to the order Diplostomida and 149 species to Plagiorchiida. Predominantly, trematode species using birds as hosts were recorded (62 species), followed by those parasitising fishes (50), mammals (33) and amphibians (25) and the fewest species were reported from reptiles (12), a trend common in trematodes (see Yamaguti (1971)). Adult digeneans, recorded together with a metacercarial and/or cercarial stage, comprised 28 species (15%), which indicates that the Georgian fauna of trematodes and their life-cycles are understudied and large-scale investigations of molluscs are needed to assess the trematode species spectrum completing their life-cycles in Georgia. The majority of species were recorded as associated with freshwater, most probably because of lack of investigation in the marine realm.

Apart from well-identified species, there are taxa of dubious identity appearing in the Georgian scientific literature, which we decided not to include in the checklist above. One of them is Clinostomum sp. of Kurashvili (Kurashvili 1950, Kurashvili 1953a, Kurashvili 1957, Kurashvili 1961a), which was not identified to species level, but it was claimed to be closest to Clinostomum hornum; however, there is no voucher specimen available for checking, leaving the question of the identity of this record unanswered. The next record is Petasiger jubilarum (Elperina in Skrjabin, Petrov & Baschkirova, 1947) Skrjabin & Baschkirova, 1956 found in Anas plathyrynchos from Paliastomi Lake (Kurashvili 1957, Kurashvili 1961a, Kurashvili 1984b); however, this species is currently considered a species inquirendum (see Faltýnková et al. (2008)). In another study, Javelidze (1976) obtained an adult experimentally from Melanopsis praemorsa and presumed it could be Philophtalmus sp. (nyrocae? Yamaguti, 1934); however, there is no clear species delimitation.

A specific topic in the literature on digeneans are reports of cercariae (the motile larval stages of digenean trematodes emerging from mollusc hosts). Due to the historical development of understanding of trematode life-cycles, the nomenclature and taxonomy of cercariae was originally separated from that of adult trematodes. This resulted in a substantial amount of cercarial names, many of which now have to be regarded as provisional, unless they can be assigned to a valid species described, based on an adult. In Georgia, surveys of larval trematodes from snails were carried out and some of the cercariae found can be identified to genus level or only to family: Cercaria monostomi Linstow, 1884 ex Ampullaceana balthica and A. lagotis from surroundings of Sartichala (Chiaberashvili 1971a) belongs to the Notocotylidae; Cercaria stylosa Linstow, 1884 (Chiaberashvili 1971a) could possibly belong to Ichtyocotylurus sp.; while Cercaria limnaeae ovatae Linstow, 1884 ex A. balthica (Chiaberashvili 1971a) most probably belongs to Plagiorchiidae (for all compare with Cichy et al. (2011). The following cercariae most probably belong to the family Lecithodendriidae: Cercaria ksaniensis ex Melanopsis praemorsa from the River Qsani (Eastern Georgia) (Javelidze 1973), Cercaria thezamiensis from River Thezami (Eastern Georgia) (Javelidze 1973), Cercaria rosetae and Cercaria ginetzinskaiae ex M. praemorsa from the River Qsani (Javelidze and Chiaberashvili 1973) and Cercaria colchica I ex Peregriana peregra from Kolkheti lowland water reservoirs (Kurashvili 1984a). There were more cercariae recorded which could not be assigned to any genus or family and, because of lack of information, we do not make any attempt for grouping into families; they are the following: Cercaria samgoriensis ex A. balthica from surroundings of Samgori (Chiaberashvili 1971a); Cercaria joriensis ex A. balthica and P. planorbis from Sartichala and Gardabani (Chiaberashvili 1971a); Cercaria burti ex P. planorbis from Samgori (Chiaberashvili 1971a); Cercaria gracilis ex A. balthica from surroundings of Sartichala and Gardabani (Chiaberashvili 1971a); Cercaria rhionica I from River Pichori (Kurashvili 1984a); Cercaria rhionica VIII Olenov & Doborovolskiy, 1975 and Cercaria rhionica XII Olenov & Doborovolskiy, 1975 ex M. praemorsa from rivers of Western Georgia (Ataev and Dobrovolskiy 1992); Cercaria ascanica Chiaberashvili & Javelidze 1977 ex M. praemorsa (Chiaberashvili and Javelidze 1977). Many of them were designated by the authors (Chiaberashvili 1971a, Javelidze 1973, Javelidze and Chiaberashvili 1973, Kurashvili 1984a) as new species, however, without providing any adults as vouchers.

Our revision of digenean species listed in the monograph by Kurashvili et al. (1980) revealed that some of the data on species distribution are vague and inaccurate. This is mainly because the transboundary River Mtkvari (Kura) and the mountain ranges (Caucasus Mountains), which stretch over all Caucasian countries, are frequently considered a continuous distributional area for the taxa and their locations were not distinguished by country. As a result, subsequent sources, citing Kurashvili et al. (1980) and listing the Caucasian countries for species distribution, are frequently imprecise. For instance, according to the NHM London database (NHML 2022), all digenean species recorded in the South Caucasus, based on Kurashvili et al. (1980), are also indicated for Georgia. However, after checking the original literary sources (Kurashvili et al. 1951, Chiaberashvili 1955, Chiaberashvili 1957, Chiaberashvili 1959, Qoiava 1966, Chiaberashvili 1968, Petriashvili 1971), we found out that the following digenean species (that were actually recorded in Azerbaijan) were not reported from Georgia: Acanthocreadium araxicus Mikailov, 1969; A. talischiensis Mikailov, 1969; Ascocotyle coleostoma (Looss, 1896) Looss, 1899; Asymphylodora imitans (Muhling, 1898) Looss, 1899; A. kurensis Paschajev, 1970; Bunocotyle cingulata Odhner, 1928; Bychowskycreadium bychowsky Mikailov, 1968; B. schiliani Mikailov, 1967; Hysteromorpha triloba (Rudolphi, 1819) Lutz, 1931; Ichthyocotylurus pileatus (Rudolphi, 1802) Odening, 1969 and Sanguinicola inermis Plehn, 1905. Similar results were found for host species and their distribution. Careful checking of the original literature (Petriashvili 1964, Petriashvili 1966) cited by Sey (2001) revealed that R. esculenta is not a host for D. mehrai in Georgia; while Emys orbicularis and Triturus vulgaris are not hosts of D. subclavatus; however, they are listed as such in the database of NHM London. This highlights the continuing importance of assessing primary sources.

In recent years, we recorded trematode intramolluscan stages (sporocysts, rediae, cercariae and metacercariae) in snails, which were new host records for trematodes in Georgia: Helix lucorum for Brachylaimidae (Murvanidze et al. 2010, Arabuli et al. 2019); Oxychilus mingrelicus for Dicrocoelium sp. (Arabuli 2018); Lymnaea stagnalis for Diplostomum spathaceum, Moliniella anceps and Notocotylus attenuatus; Radix euphratica for Telorchis sp., Tylodelphys sp. and Plagiorchis sp. (Arabuli, unpublished data). Moliniella anceps (Molin, 1859) Hübner, 1939 was recorded for the first time in Georgia by Arabuli et al. (2015).

Conclusion

Data on the digeneans of Georgia are mainly represented by adults in their definitive hosts and they are predominantly from freshwater. Most of the research was conducted many decades ago and was based solely on morphology and was published predominantly in Georgian or Russian language. Since there is a striking shortage of studies on intermediate hosts (i.e. molluscs), large-scale and site-intensive investigations are needed to explore species diversity and distribution and to obtain data on life-cycles and transmission pathways to be further used for ecological and epidemiological studies, as well as for biodiversity conservation in Georgia.

Acknowledgements

We would like to thank our colleague Ketevan Asatiani for helping with fieldwork and laboratory assistance and Ani Bikashvili for freshwater snail species identification. This research has been supported by Shota Rustaveli National Science Foundation of Georgia (SRNSFG) - Ph.D. Grant to L.A. (№PHDF-21-260, “Parasitological research of molluscs of Lisi and Turtle lakes”). We would also like to express our gratitude to anonymous reviewers and the editor, Simona Georgieva, for their valuable comments and suggestions.

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Supplementary material

Endnotes