A checklist of helminths (Acanthocephala, Nematoda and Platyhelminthes) found in rodents (Mammalia, Rodentia) from Slovakia

Yaroslav Syrota; Zuzana Hurníková; Vitaliy Kharchenko; Martina Miterpáková

doi:10.3897/zookeys.1251.154911

Checklist

A checklist of helminths (Acanthocephala, Nematoda and Platyhelminthes) found in rodents (Mammalia, Rodentia) from Slovakia

Yaroslav Syrota^‡§, Zuzana Hurníková^‡, Vitaliy Kharchenko^§, Martina Miterpáková^‡

‡ Institute of Parasitology, Slovak Academy of Sciences, Košice, Slovakia

§ I. I. Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kyiv, Ukraine

Corresponding author: Yaroslav Syrota ( syrota@saske.sk )

Academic editor: Boyko Georgiev

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: Syrota Y, Hurníková Z, Kharchenko V, Miterpáková M (2025) A checklist of helminths (Acanthocephala, Nematoda and Platyhelminthes) found in rodents (Mammalia, Rodentia) from Slovakia. ZooKeys 1251: 61-96. https://doi.org/10.3897/zookeys.1251.154911

ZooBank: urn:lsid:zoobank.org:pub:6643206C-5F65-48B6-83B9-0192BF648B5E

Abstract

Nearly seven decades have passed since the first records of helminths in rodents in Slovakia were published. Thereafter, the number of documented helminth species within the territory of the country has steadily increased. This checklist compiles data from 34 sources, including 27 academic articles, five unpublished reports, and two conference proceedings produced between 1955 and early 2025. Following a critical review of each source, 73 nominal helminth species from 27 species of rodents, comprising 30 nematodes, 29 cestodes, 13 trematodes, and one acanthocephalan, were included in this checklist. Information on helminth diversity is absent for only two of the 29 rodent species registered in the Slovak Republic: Myocastor coypus and Arvicola scherman. Furthermore, our findings suggest that certain rodent species remain under-researched despite the presence of some information on their helminth fauna. This checklist lays the basis for future helminthological research on rodents in Slovakia. The raw dataset generated for this article is available in the Suppl. material 1.

Key words:

Biodiversity, Europe, inventory, micromammal, parasites, review, Slovak Republic

Introduction

Rodents are the most diverse order of Mammalia, with over 2000 species inhabiting terrestrial, subterranean, arboreal, and aquatic environments on all continents except Antarctica (Wolff and Sherman 2007). Recent research estimates that the biomass of all land mammals is 22 million tonnes (wet weight), with rodents comprising 16% of this total (Greenspoon et al. 2023). Due to their wide distribution and significant biomass, rodents play a vital role in the trophic networks in many terrestrial ecosystems. In addition to being prey for various predators, they also host many species of helminths, a subject of considerable research interest, as evidenced by publications summarising numerous studies on helminths of rodents (Smales 1997; Preisser and Falcón-Ordaz 2019; Preisser 2019; Islam et al. 2020; Panti-May et al. 2021; Dursahinhan et al. 2023; Hamzavi et al. 2024).

In Slovakia, as in other countries, studies have also been conducted on the diversity of rodent helminths. To our knowledge, the first summarising work on this topic is a publication indexing a museum collection of helminths from hosts of different taxa, including data on rodent helminths’ findings from the territory of the former Czechoslovakia (Hovorka 1968). Another valuable resource is several papers within the Synopsis of Cestodes in Slovakia series (Macko et al. 1994; Hanzelová et al. 1995; Hanzelová and Ryšavý 1996). These articles, which cover all groups of vertebrate hosts, also include records of rodent cestodes and their hosts. The most recent work providing data on parasites is a comprehensive review of mammals in Slovakia (Krištofík and Danko 2012). Although the monograph offers a list of endo- and ectoparasites found in each mammal species recorded in Slovakia, its primary focus is on the mammals rather than their parasites.

Consequently, the available literature includes five sources with limited inventories on rodent helminth diversity in the country, and none are exclusively dedicated to the helminths of these hosts. Our study aims to fill this gap by compiling up-to-date information on helminths of wild and synanthropic rodents inhabiting Slovakia.

Material and methods

The checklist was compiled based on information obtained from two sources: (1) an online search of academic research databases such as Scopus and Web of Science, and (2) a bibliographical search of literature in the library of the Institute of Parasitology of the Slovak Academy of Sciences. All found sources were critically reviewed before their data were included in this checklist. One subject of this review was eliminating geographical ambiguity, as in several instances, it was unclear whether the data originated from Czechia or Slovakia. This uncertainty arose because part of the research had been conducted in former Czechoslovakia without indicating a precise location. Such sources were excluded. Another criterion for selecting a source was the originality of the data. In some cases, an author had published nearly identical content in two sources: an article and a conference abstract. Only sources with a more comprehensive dataset were selected for such cases. As a result of the selection, 34 sources produced between 1955 and the beginning of 2025 were used to compile the checklist; 27 of these sources are published academic works, five are unpublished reports from the Institute of Parasitology, and two are abstracts from conference proceedings. The raw dataset used to create the checklist is available as the Suppl. material 1.

The studies used for this checklist varied considerably in the precision with which localities were described, from specific sites to large regions; however, most sources provided no exact geographic coordinates. To standardise the presentation of geographical information, we used the Google Maps web application to determine approximate coordinates for each locality. We used approximate centres for larger areas, such as administrative regions or conservation zones. Localities were categorised into three groups based on precision: zone – precise locations; vicinity – areas near landmarks; and area – broad or vaguely defined locations. Detailed information for each locality, including name, coordinates, and precision category, is available in the Suppl. material 1. Coordinates provided by us should be regarded as the best available cartographic proxies for the named sites rather than instrument-logged GPS points. We used QGIS® software to visualise the geographical coverage of helminth research across Slovakia (Fig. 1).

Figure 1.

Approximate geographic locations in Slovakia where rodents were sampled. Note: Sampling localities are categorised based on the precision of their geographic coordinates: zone – precise locations, vicinity – areas near landmarks, area – broad or vaguely defined locations.

The parasite-host list is presented alphabetically, starting with the acanthocephalans, followed by the nematodes and the platyhelminths (cestodes and trematodes). The list includes information on synonyms of helminth scientific names, hosts (listed in alphabetical order), localities (listed in alphabetical order), infection sites (listed in alphabetical order), references, and remarks (if necessary). Scientific names of helminths follow those provided in the Global Biodiversity Information Facility (GBIF.org 2024), Global Cestode Database (Caira et al. 2022), World Register of Marine Species (WoRMS Editorial Board 2024), Nemys: World Database of Nematodes (Nemys 2024), and selected publications (Biserkov 1998; Mahesh Kumar et al. 2006; Jackson and Friberg 2022; Makarikov and Binkienė 2022; Behnke and Jackson 2024). A ‘Synonyms’ entry is provided only when the original binomen in a source differs from the combination accepted in the databases and literature used for name verification.

In addition, a host-parasite list was compiled based on the parasite-host list. This list details each host, including its scientific name and the scientific names of all associated helminths, organised by taxonomic group: Acanthocephala, Nematoda, Cestoda, and Trematoda. Within each taxonomic group, the helminth names are listed in alphabetical order.

Results

Based on the literature analysis, we documented 73 helminth species from 27 rodent host species in Slovakia. Additionally, we considered each of the genera Echinostoma, Mesocestoides, Citellina, and Toxocara to be represented by at least one species, as no species-level identifications were available for these taxa in the checklist. As a result, at least four additional species can be inferred, bringing the total number of detected species to a minimum of 77. These include 32 species of nematodes, 30 species of cestodes, 14 species of trematodes, and one species of acanthocephalans.

Annotated parasite-host list

Phylum Acanthocephala Rudolphi, 1802

Class Archiacanthocephala Meyer, 1931

Order Moniliformida Schmidt, 1972

Family Moniliformidae Van Cleave, 1924

Genus Moniliformis Travassos, 1915

Moniliformis moniliformis (Bremser, 1811)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Glis glis, Microtus arvalis, Mus musculus, Myodes glareolus, Rattus norvegicus, Spermophilus citellus.

Localities. Domica, Jablonov, Košice, Protected Landscape Area (PLA) Poľana, Rožňava, Slovenský kras, Vihorlat, Vysoké Tatry, Žiar nad Hronom.

Infection site. Small intestine.

Reported by. Erhardová (1956, 1958), Baruš and Tenora (1957), Tenora (1963), Mituch (1970, 1975), Mituch et al. (1987).

Phylum Nematoda Cobb, 1932

Class Chromadorea Inglis, 1983

Order Rhabditida Chitwood, 1933

Family Ascarididae Baird, 1853

Genus Porrocaecum Railliet & Henry, 1912

Porrocaecum depressum (Zeder, 1800)

Hosts. Apodemus sylvaticus, Mus musculus, Sciurus vulgaris.

Localities. PLA Poľana, Roháčska dolina.

Infection site. Body cavity, caecum, large intestine.

Reported by. Tenora (1967), Mituch (1975).

Remark. P. depressum has a cosmopolitan distribution and has been reported from approximately 47 species of birds of prey (Atkinson et al. 2008). Rodents serve as paratenic hosts.

Porrocaecum spp.

Hosts. Microtus agrestis, Microtus subterraneus.

Localities. Roháčska dolina, Vysoké Tatry.

Infection site. Caecum, small intestine wall – nodules.

Reported by. Erhardová (1956), Tenora (1967).

Remark. Members of the genus Porrocaecum are widely distributed intestinal parasites of birds, with shrews – and perhaps other small mammals that consume intermediate hosts (e.g., earthworms) – serving as paratenic hosts to transfer the parasite to carnivorous definitive hosts (Anderson 2000).

Genus Toxocara Stiles & Hassall, 1905

Toxocara spp.

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Micromys minutus, Microtus arvalis, Mus musculus, Myodes glareolus.

Localities. Košice, Tatra National Park (TANAP).

Infection site. Antibodies in sera.

Reported by. Dubinský et al. (1995).

Family Heligmosomidae Cram, 1927

Genus Carolinensis Travassos, 1937

Carolinensis minutus (Dujardin, 1845)

Hosts. Microtus arvalis, Microtus subterraneus, Myodes glareolus.

Locality. Čergov Mts.

Infection site. Not specified.

Reported by. Mészáros and Štolmann (1984).

Genus Glistrongylus Durette Desset, Digiani, Kilani & Geffard Kuriyama, 2017

Glistrongylus gracilis (Leuckart, 1842)

Synonym. Longistriata schulzi Schachnasarova, 1949.

Hosts. Apodemus sylvaticus, Dryomys nitedula, Glis glis, Microtus arvalis.

Localities. PLA Poľana, Roháčska dolina, Slovenský kras, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Baruš and Tenora (1957), Tenora (1967), Mituch (1970, 1975).

Genus Heligmosomoides Hall, 1916

Heligmosomoides glareoli Baylis, 1928

Synonym. Heligmosomum tatricum Erchardova, 1955.

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Microtus agrestis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Čergov Mts., PLA Poľana, Roháčska dolina, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Erhardová (1955a), Tenora (1955, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987).

Heligmosomoides polygyrus (Dujardin, 1845)

Synonym. Heligmosomum skrjabini Skrjabin & Schikhobalova, 1952.

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus, Mus musculus, Mus spicilegus, Muscardinus avellanarius, Myodes glareolus, Sicista betulina.

Localities. Bratislava, Čergov Mts., Domica, Gabčíkovo, Košice, Liptovský Mikuláš, Orava, PLA Poľana, Roháčska dolina, Rozhanovce, South Eastern Slovakia, South Slovakia, Veľké Kapušany, Vysoké Tatry, Zádiel, Žiar nad Hronom.

Infection site. Intestines, small intestine, small intestine – muscularis propria.

Reported by. Erhardová (1955a, 1955b, 1956), Tenora (1963, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987), Ribas et al. (2009), Ondríková et al. (2010).

Genus Heligmosomum Railliet & Henry, 1909

Heligmosomum costellatum (Dujardin, 1845)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus.

Localities. Bratislava, Čergov Mts., Domica, Kráľovský Chlmec, PLA Poľana, Roháčska dolina, Vysoké Tatry.

Infection site. Small intestine, stomach.

Reported by. Erhardová (1955a, 1956), Tenora (1955, 1963, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987).

Heligmosomum halli (Schulz, 1926)

Hosts. Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Intestines, small intestine.

Reported by. Mituch (1970, 1975).

Heligmosomum mixtum Schulz, 1929

Host. Myodes glareolus.

Locality. Čergov Mts.

Infection site. Small intestine.

Reported by. Mészáros and Štolmann (1984).

Family Heterakidae Railliet & Henry, 1912

Genus Heterakis Schrank, 1790

Heterakis spumosa Schneider, 1866

Synonym. Ganguloterakis spumosa (Schneider, 1866).

Hosts. Apodemus agrarius, Rattus norvegicus.

Localities. Banská Bystrica, Bratislava, Bytča, Galanta, Levice, Púchov, Rajec, Rozhanovce, Sečovce, Senec, Trenčín, Trnava, Turčianske Teplice.

Infection site. Caecum, large intestine, small intestine.

Reported by. Mituch (1957), Erhardová (1958), Ondríková et al. (2010).

Family Heteroxynematidae Skrjabin & Shikhobalova, 1948

Genus Aspiculuris Schulz, 1927

Aspiculuris dinniki Schulz, 1927

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Chionomys nivalis, Microtus agrestis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. PLA Poľana, Roháčska dolina, Vysoké Tatry.

Infection site. Caecum, large intestine, small intestine.

Reported by. Erhardová (1955a), Tenora (1955, 1967), Mituch (1970, 1975), Mituch et al. (1987).

Aspiculuris tetraptera Nitzsch, 1821

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Mus musculus.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Caecum, large intestine.

Reported by. Erhardová (1955a), Mituch (1970, 1975), Mituch et al. (1987).

Family Oxyuridae Weinland, 1858

Genus Citellina Prendel, 1928

Citellina spp.

Host. Spermophilus citellus.

Localities. Boleráz – airport, Bratislava – Zelená voda, Spišský hrad – castle.

Infection site. Intestines.

Reported by. Štrkolcová et al. (2024).

Genus Enterobius Leach, 1853

Enterobius (Enterobius) sciuri (Cameron, 1932)

Host. Sciurus vulgaris.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Caecum, large intestine.

Reported by. Mituch (1970, 1975), Mituch et al. (1987).

Genus Syphacia Seurat, 1916

Syphacia (Seuratoxyuris) frederici (Roman, 1945)

Host. Apodemus flavicollis.

Locality. Čergov Mts.

Infection site. Not specified.

Reported by. Mészáros and Štolmann (1984).

Syphacia (Syphacia) montana (Yamaguti, 1943)

Hosts. Marmota marmota, Microtus agrestis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Čergov Mts., Roháčska dolina, Vysoké Tatry.

Infection site. Caecum, small intestine.

Reported by. Tenora (1967), Mituch (1970), Mészáros and Štolmann (1984), Mituch et al. (1987).

Syphacia (Syphacia) nigeriana (Baylis, 1928)

Host. Microtus arvalis.

Locality. Čergov Mts.

Infection site. Not specified.

Reported by. Mészáros and Štolmann (1984).

Syphacia (Syphacia) obvelata (Rudolphi, 1802)

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus, Mus musculus, Mus spicilegus, Muscardinus avellanarius, Myodes glareolus.

Localities. Domica, Gabčíkovo, Košice, Kráľovský Chlmec, Liptovský Mikuláš, Medzilaborce, Michalovce, Nízke Tatry, Orava, PLA Poľana, Prešov, Rožňava, Slovenské Nové Mesto, South Eastern Slovakia, Vysoké Tatry, Žiar nad Hronom.

Infection site. Caecum, intestines, large intestine, small intestine.

Reported by. Erhardová (1955a, 1956, 1958), Tenora (1955, 1963), Mituch (1970, 1975), Mituch et al. (1987), Ribas et al. (2009).

Remark. Genetic evidence indicates that S. obvelata is associated with murine rodents (Okamoto et al. 2007; Stewart et al. 2018); therefore, reports of its occurrence in other rodent groups are doubtful and should likely be considered misidentifications or incidental infections.

Syphacia (Syphacia) stroma (Linstow, 1884)

Hosts. Apodemus agrarius, Apodemus flavicollis.

Locality. Rozhanovce.

Infection site. Not specified.

Reported by. Ondríková et al. (2010).

Family Rhabditidae

Genus Pelodera Schneider, 1866

Pelodera orbitalis (Sudhaus & Schulte, 1986)

Synonym. Rhabditis orbitalis Sudhaus & Schulte, 1986.

Hosts. Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Belianske Tatry, Nízke Tatry, Roháče, Slovenský kras, Vysoké Tatry.

Infection site. Conjunctival sacs, orbit.

Reported by. Baruš and Hrabě (1991), Kocianová-Adamcová et al. (2006).

Family Rictulariidae Railliet, 1916

Genus Rictularia Froelich, 1802

Rictularia proni Seurat, 1915

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Dryomys nitedula, Eliomys quercinus, Microtus arvalis, Mus musculus, Myodes glareolus.

Localities. Bratislava, Jablonov, Roháčska dolina, Slovenský kras, Vysoké Tatry.

Infection site. Small intestine, stomach.

Reported by. Baruš and Tenora (1957), Erhardová (1958), Tenora (1967), Mituch (1970).

Rictularia spp.

Host. Myodes glareolus.

Locality. Roháčska dolina.

Infection site. Small intestine.

Reported by. Tenora (1967).

Family Spirocercidae Chitwood & Wehr, 1932

Genus Mastophorus Diesing, 1853

Mastophorus muris (Gmelin, 1790)

Hosts. Apodemus agrarius, Chionomys nivalis, Microtus subterraneus, Mus musculus, Myodes glareolus, Rattus norvegicus.

Localities. PLA Poľana, Roháčska dolina, Rozhanovce, Vysoké Tatry.

Infection site. Stomach.

Reported by. Tenora (1967), Mituch (1970, 1975), Mituch et al. (1987), Ondríková et al. (2010).

Family Strongyloididae Chitwood & McIntosh, 1934

Genus Strongyloides Grassi, 1879

Strongyloides ratti Sandground, 1925

Host. Rattus norvegicus.

Localities. Dunajská Streda, Galanta, Košice, Kráľovský Chlmec, Levice, Nové Zámky, Šahy, Šurany.

Infection site. Small intestine.

Reported by. Mituch (1957).

Family Trichostrongylidae Leiper, 1912

Genus Travassosius Khalil, 1922

Travassosius rufus Khalil, 1922

Host. Castor fiber.

Localities. Danube River, Laborec River, Topľa River.

Infection site. Intestines.

Reported by. Bystrianska et al. (2021).

Genus Trichostrongylus Looss, 1905

Trichostrongylus retortaeformis (Zeder, 1800)

Hosts. Microtus arvalis, Sciurus vulgaris.

Locality. Vysoké Tatry.

Infection site. Intestines.

Reported by. Mituch (1970).

Remark. Lagomorphs are the primary hosts of T. retortaeformis, as shown by Audebert et al. (2002); rodent records likely represent accidental spill-over infections rather than stable host–parasite associations.

Trichostrongylidae gen. sp.

Host. Marmota marmota, Spermophilus citellus.

Localities. Bratislava – Štefánikova mohyla, Bratislava – Zelená voda, Chtelnica, Detvianska Huta, Gemerské Dechtáre, Jesenské, Košice – airport, Kružná, Moldava – hill, Moldava nad Bodvou, Petrovce – airport, Pieskovec – east, Pieskovec zásoby, Poráč, Silica pod Búčnikom, Spišská Nová Ves – Hrádok, Velická dolina.

Infection site. Intestines.

Reported by. Jászayová et al. (2023), Štrkolcová et al. (2024).

Class Enoplea Inglis, 1983

Order Trichinellida Hall, 1916

Family Capillariidae Railliet, 1915

Genus Aonchotheca López-Neyra, 1947

Aonchotheca annulosa (Dujardin, 1845)

Synonyms. Capillaria annulosa (Dujardin, 1845), Capillaria murisylvatici (Diesing, 1851).

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Microtus arvalis, Mus musculus, Myodes glareolus, Rattus norvegicus.

Localities. Čergov Mts., Gabčíkovo, Kráľovský Chlmec, Nové Mesto nad Váhom, PLA Poľana, Rajec, Rozhanovce, Sečovce, Vihorlat, Vysoké Tatry.

Infection site. Intestines, small intestine, stomach.

Reported by. Erhardová (1956), Mituch (1957, 1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987), Ondríková et al. (2010).

Aonchotheca myoxinitelae (Diesing, 1851)

Synonym. Skrjabinocapillaria myoxi-nitelae (Diesing, 1851).

Host. Eliomys quercinus.

Locality. Slovenský kras.

Infection site. Not specified.

Reported by. Baruš and Tenora (1957).

Genus Calodium Dujardin, 1845

Calodium hepaticum (Bancroft, 1893)

Synonyms. Capillaria hepatica Bancroft, 1893, Hepaticola hepatica (Bancroft, 1893).

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Arvicola amphibius, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Myodes glareolus, Rattus norvegicus.

Localities. Košice – zoo, Lučenec, PLA Poľana, Sečovce, TANAP, Trenčín, Vihorlat, Vysoké Tatry.

Infection site. Liver.

Reported by. Erhardová (1956), Mituch (1957, 1970, 1975), Mituch et al. (1987), Miterpáková et al. (2024).

Genus Capillaria

Capillaria spp.

Host. Spermophilus citellus.

Localities. Bratislava – airport, Chtelnica, Ďurkovec – camp, Ďurkovec – skilift, Jesenské, Košice – airport, Kružná, Kuchyňa vývrat, Moldava – hill, Moldava nad Bodvou habitation, Muráň Biele Vody, Petrovce – airport, Pieskovec – east, Pieskovec zásoby, Poráč, Silica pod Búčnikom, Spišská Kapitula, Spišská Nová Ves – airport, Spišská Nová Ves – Hrádok, Spišské Podhradie, Spišský hrad – castle, Teplička, Vyšný Slavkov.

Infection site. Intestines.

Reported by. Štrkolcová et al. (2024).

Genus Liniscus Dujardin, 1845

Liniscus papillosus (Polonio, 1860)

Synonym. Capillaria papillosa (Polonio, 1860).

Host. Rattus norvegicus.

Localities. Bytča, Kráľovský Chlmec, Ružomberok, Trebišov.

Infection site. Urinary bladder.

Reported by. Mituch (1957).

Family Trichinellidae Ward, 1907

Genus Trichinella Railliet, 1895

Trichinella britovi Pozio, La-Rosa, Murrell & Lichtenfels, 1992

Host. Myodes glareolus.

Locality. Vysoké Tatry.

Infection site. Muscles.

Reported by. Hurníková et al. (2025)

Trichinella pseudospiralis Garkavi, 1972

Hosts. Apodemus flavicollis, Rattus norvegicus.

Localities. Michalovce, Vysoké Tatry.

Infection site. Muscles.

Reported by. Hurníková et al. (2005, 2025).

Trichinella spp.

Hosts. Apodemus flavicollis, Rattus norvegicus.

Localities. Humenné, Košice, Prešov, Stará Ľubovňa.

Infection site. Muscles.

Reported by. Mituch (1956, 1957).

Family Trichosomoididae Yorke & Maplestone, 1926

Genus Trichosomoides Railliet, 1895

Trichosomoides crassicauda (Bellingham, 1840)

Host. Rattus norvegicus.

Localities. Bratislava, Galanta, Hlohovec.

Infection site. Urinary bladder.

Reported by. Mituch (1957), Erhardová (1958).

Family Trichuridae Railliet, 1915

Genus Trichuris Roederer, 1761

Trichuris muris (Schrank, 1788)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Mus musculus, Rattus norvegicus.

Localities. Bratislava, Domica, Gabčíkovo, Košice, Kráľovský Chlmec, Levice, PLA Poľana, Rožňava, Ružomberok, Sečovce, Trebišov, Vysoké Tatry.

Infection sites. Caecum, intestines, large intestine.

Reported by. Erhardová (1955a, 1956, 1958), Mituch (1957, 1970, 1975), Tenora (1963), Mituch et al. (1987).

Remark. Trichuris arvicolae Feliu et al., 2000 was described from arvicoline rodents, whereas T. muris is suggested to be specialised to murine rodents (Feliu et al. 2000). Accordingly, the status of the specimens from arvicoline rodents should be re-evaluated since they probably belong to T. arvicolae.

Phylum Platyhelminthes Minot, 1876

Class Cestoda van Beneden, 1849

Order Cyclophyllidea van Beneden in Braun, 1900

Family Anoplocephalidae Cholodkovsky, 1902

Genus Anoplocephaloides Baer, 1923

Anoplocephaloides dentata (Galli-Valerio, 1905)

Synonyms. Paranoplocephala dentata Galli-Valerio, 1905, Paranoplocephala brevis Kirschenblatt, 1938.

Hosts. Apodemus flavicollis, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Čergov Mts., Domica, Nízke Tatry, PLA Poľana, Roháče, Roháčska dolina, Vysoké Tatry.

Infection sites. Caecum, intestines, small intestine.

Reported by. Tenora (1955, 1967), Erhardová (1955a, 1956, 1958), Mituch (1970, 1975), Tenora and Murai (1980), Mészáros and Štolmann (1984), Mituch et al. (1987).

Genus Ctenotaenia Railliet, 1893

Ctenotaenia marmotae (Frölich, 1802)

Host. Marmota marmota.

Locality. Velická dolina, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Tenora (1961), Jászayová et al. (2023).

Genus Eurotaenia Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Eurotaenia gracilis (Tenora & Murai, 1980)

Synonym. Paranoplocephala gracilis Tenora & Murai, 1980.

Hosts. Chionomys nivalis, Microtus agrestis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Mutné, Roháče.

Infection site. Small intestine.

Reported by. Tenora and Murai (1980).

Genus Microticola Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Microticola blanchardi (Moniez, 1891)

Synonym. Paranoplocephala blanchardi (Moniez, 1891).

Hosts. Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Locality. Roháče.

Infection site. Small intestine.

Reported by. Tenora and Murai (1980).

Genus Paranoplocephala Lühe, 1910

Paranoplocephala macrocephala (Douthitt, 1915)

Synonym. Andrya macrocephala Douthitt, 1915.

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Arvicola amphibius, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus.

Localities. Gabčíkovo, PLA Poľana, Roháčska dolina, Veľký Meder, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Erhardová (1956, 1958), Tenora (1963, 1967), Mituch (1970, 1975).

Remark. According to Haukisalmi and Henttonen (2003), P. macrocephala is a parasite of microtine and geomyid rodents in North America. The taxonomic position of cestodes from Europe identified as P. macrocephala requires further studies.

Paranoplocephala microti (Hansen, 1947)

Synonym. Andrya microti Hansen, 1947.

Hosts. Microtus arvalis, Myodes glareolus.

Localities. Domica, Košice.

Infection site. Small intestine.

Reported by. Erhardová (1956).

Remark. According to Haukisalmi et al. (2014), P. microti is a parasite of voles in North America only. The taxonomic position of cestodes from Slovakia identified as P. microti needs to be confirmed by further examinations.

Paranoplocephala omphalodes (Hermann, 1783)

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Myodes glareolus, Rattus norvegicus.

Localities. Bratislava, Domica, Eastern Slovakia, Orava, PLA Poľana, Roháče, Vysoké Tatry.

Infection site. Intestines, large intestine, small intestine.

Reported by. Erhardová (1955a, 1956, 1958), Tenora (1955), Mituch (1970, 1975), Tenora and Murai (1980), Mituch et al. (1987).

Paranoplocephala spp.

Hosts. Marmota marmota, Microtus subterraneus, Myodes glareolus.

Localities. Čergov Mts., Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mészáros and Štolmann (1984), Mituch et al. (1987).

Family Catenotaeniidae Spasskii, 1950

Genus Catenotaenia Janicki, 1904

Catenotaenia dendritica (Goeze, 1782)

Host. Sciurus vulgaris.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mituch (1970, 1975), Mituch et al. (1987).

Catenotaenia henttoneni Haukisalmi & Tenora, 1993

Synonym. Catenotaenia cricetorum f. glareolica Tenora, 1959.

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus.

Localities. Čergov Mts., Domica, PLA Poľana, Roháčska dolina, Vysoké Tatry.

Infection site. Intestines, small intestine.

Reported by. Tenora (1963, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987).

Catenotaenia pusilla (Goeze, 1782)

Localities. Bratislava, Eastern Slovakia, Gabčíkovo, Košice, Kráľovský Chlmec, Nízke Tatry, PLA Poľana, Rožňava, South Slovakia, Vysoké Tatry.

Infection site. Intestines, small intestine.

Reported by. Erhardová (1955a, 1956, 1958), Mituch (1970, 1975).

Genus Spasskijela Tenora, 1959

Spasskijela lobata (Baer, 1925)

Synonym. Skrjabinotaenia lobata (Baer, 1925).

Hosts. Apodemus agrarius, Apodemus flavicollis, Microtus arvalis, Mus musculus, Myodes glareolus.

Localities. Bratislava, Domica, Gabčíkovo, Košice, PLA Poľana, Rozhanovce, Zádiel, Žiar nad Hronom.

Infection site. Intestines, small intestine.

Reported by. Erhardová (1956, 1958), Tenora (1963), Mituch (1975), Ondríková et al. (2010).

Family Dilepididae Railliet & Henry, 1909

Genus Sobolevitaenia Spasskaya & Makarenko, 1965

Sobolevitaenia unicoronata (Fuhrmann, 1908)

Synonym. Choanotaenia unicoronata (Fuhrmann, 1908).

Hosts. Apodemus flavicollis, Microtus arvalis.

Localities. Roháčska dolina, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Tenora (1967), Mituch (1970).

Remark. The definitive hosts of S. unicoronata are passeriform birds (Spasskaya and Spasskii 1977). Nonetheless, Tenora (1967) and Mitúch (1970) each reported larvae of the species recovered from rodents. These occurrences likely represent paratenic parasitism.

Family Hymenolepididae Ariola, 1899

Genus Armadolepis Spasskii, 1954

Armadolepis myoxi (Rudolphi, 1819)

Hosts. Dryomys nitedula, Glis glis.

Localities. PLA Poľana, Slovenský kras, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Tenora (1965), Mituch (1970, 1975).

Remark. Tenora (1965) reported H. sulcata from G. glis. Makarikov and Georgiev (2020) re-examine the archived material of Tenora (1965) and re-identified it as A. myoxi (sensu stricto).

Armadolepis spasskyi Tenora & Baruš, 1958

Hosts. Dryomys nitedula, Eliomys quercinus, Glis glis.

Locality. Slovenský kras.

Infection site. Small intestine.

Reported by. Tenora and Baruš (1958), Tenora (1965).

Remark. The species was initially described by Tenora and Baruš (1958) from D. nitedula, E. quercinus, and G. glis. Makarikov and Georgiev (2020) redescribed the species based on the type series, adding corrections to the diagnostic characters of the species. They also re‐examined archived cestode specimens recovered from G. glis in Slovakia and found no confirmed occurrences of A. spasskyi. Therefore, the authors concluded that the records of A. spasskyi from G. glis are doubtful.

Genus Arostrilepis Mas-Coma & Tenora, 1997

Arostrilepis horrida (Linstow, 1901)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Arvicola amphibius, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus, Rattus norvegicus, Sicista betulina.

Localities. Čergov Mts., Nové Zámky, PLA Poľana, Roháčska dolina, TANAP, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mituch (1957, 1970, 1975), Tenora (1967), Mészáros and Štolmann (1984), Mituch et al. (1987).

Genus Hymenolepis Weinland, 1858

Hymenolepis diminuta (Rudolphi, 1819)

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Eliomys quercinus, Glis glis, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus, Microtus tatricus, Mus musculus, Mus spicilegus, Myodes glareolus, Rattus norvegicus, Spermophilus citellus.

Localities. Bratislava, Čergov Mts., Domica, Gabčíkovo, Galanta, Košice, Kráľovský Chlmec, Levice, Medzilaborce, Michalovce, Nové Zámky, PLA Poľana, Prešov, Rozhanovce, Sečovce, Senec, Slovakia, Slovenský kras, Snina, South Slovakia, Šurany, Svit, Turčianske Teplice, Veľké Kapušany, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Erhardová (1955a, 1955b, 1956, 1958), Mituch (1957, 1970, 1975), Tenora (1963, 1965), Mészáros and Štolmann (1984), Mituch et al. (1987), Ribas et al. (2009), Ondríková et al. (2010).

Hymenolepis sulcata (von Linstow, 1879)

Host. Glis glis.

Locality. Rozhanovce.

Infection site. Small intestine.

Reported by. Sałamatin et al. (2005).

Hymenolepis spp.

Hosts. Dryomys nitedula, Eliomys quercinus, Glis glis, Spermophilus citellus.

Locality. Bratislava – airport, Ďurkovec – camp, Chtelnica, Jánovce, Roháčska dolina, Slovenský kras, Teplička.

Infection site. Small intestine.

Reported by. Baruš and Tenora (1957), Tenora (1967), Štrkolcová et al. (2024).

Remark. Tenora (1967) reported a single specimen of Hymenolepis sulcata from D. nitedula. However, Makarikov and Georgiev (2020) re-examined the original archived material. They concluded that the specimen cannot be identified to the species level due to the absence of its scolex and the poor condition of the strobila. Therefore, we include this record as Hymenolepis sp. in the checklist.

Genus Kontrimavichusia Makarikov & Binkienė, 2022

Kontrimavichusia asymmetrica (Janicki, 1904)

Synonyms. Rodentolepis asymmetrica (Janicki, 1904), Hymenolepis ampla Erhardová, 1955, Rodentolepis ampla (Erhardová, 1955).

Localities. Bratislava, Čergov Mts., Eastern Slovakia, Gabčíkovo, Nízke Tatry, PLA Poľana, Roháčska dolina, TANAP, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Erhardová (1955a, 1956, 1958), Tenora (1955, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987).

Genus Rodentolepis Spasskii, 1954

Rodentolepis fraterna (Stiles, 1906)

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus uralensis, Microtus arvalis, Microtus tatricus, Mus musculus, Myodes glareolus, Rattus norvegicus.

Localities. Banská Bystrica, Dolný Kubín, Košice, Kráľovský Chlmec, Liptovský Mikuláš, Martin, Nové Mesto nad Váhom, PLA Poľana, Poprad, Púchov, Rajec, Rozhanovce, Rožňava, Ružomberok, Sečovce, Senec, Šurany, Svit, Topoľčany, Trebišov, Turčianske Teplice, Veľké Kapušany, Vysoké Tatry, Žilina, Zvolen.

Infection site. Small intestine.

Reported by. Erhardová (1955a, 1956), Mituch (1957, 1970, 1975), Ondríková et al. (2010).

Rodentolepis microstoma (Dujardin, 1845)

Host. Cricetus cricetus.

Locality. Košice.

Infection site. Not specified.

Reported by. Jarošová et al. (2020).

Rodentolepis straminea (Goeze, 1782)

Hosts. Apodemus agrarius, Apodemus flavicollis, Apodemus sylvaticus, Microtus arvalis, Myodes glareolus, Rattus norvegicus.

Localities. Domica, PLA Poľana, Rozhanovce, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Tenora (1963), Mituch (1970, 1975), Mituch et al. (1987), Ondríková et al. (2010).

Family Mesocestoididae Fuhrman, 1907

Genus Mesocestoides Vaillant, 1863

Mesocestoides spp.

Hosts. Apodemus agrarius, Apodemus flavicollis, Myodes glareolus.

Localities. Čergov Mts., Rozhanovce.

Infection site. Not specified.

Reported by. Mészáros and Štolmann (1984), Ondríková et al. (2010).

Remark. Mesocestoides spp. are zoonotic cestodes found as adults in carnivorous domestic and wild definitive hosts and as metacestodes in several taxa of intermediate hosts, including Rodentia (Jesudoss Chelladurai and Brewer 2021).

Family Paruterinidae Fuhrmann, 1907

Genus Cladotaenia Cohn, 1901

Cladotaenia cylindracea (Bloch, 1782)

Hosts. Apodemus flavicollis, Myodes glareolus.

Localities. PLA Poľana, TANAP, Vysoké Tatry.

Infection site. Liver.

Reported by. Mituch (1970, 1975), Mituch et al. (1987).

Remark. In Slovakia, species from the genus Cladotaenia are recorded in birds of prey (Komorová et al. 2017). Rodents are intermediate hosts for these cestodes.

Family Taeniidae Ludwig, 1886

Genus Echinococcus Rudolphi, 1801

Echinococcus multilocularis Leuckart, 1863

Host. Ondatra zibethicus.

Locality. Prešov.

Infection site. Liver.

Reported by. Miterpáková et al. (2006).

Remark. Definitive hosts of E. multilocularis are canids. Rodents serve as intermediate hosts (Caira et al. 2022).

Genus Hydatigera Lamarck, 1816

Hydatigera taeniaeformis (Batsch, 1786)

Synonym. Strobilocercus fasciolaris.

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Apodemus uralensis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus, Ondatra zibethicus, Rattus norvegicus, Sciurus vulgaris.

Localities. Čergov Mts., Domica, Gabčíkovo, Galanta, Košice, Lúč na Ostrove, Nízke Tatry, PLA Poľana, Roháčska dolina, Rozhanovce, Rožňava, Slovakia, Veľké Kapušany, Vysoké Tatry.

Infection site. Liver, spleen.

Reported by. Erhardová (1955a, 1956, 1958), Tenora (1963, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Rajský (1985), Mituch et al. (1987), Ondríková et al. (2010).

Remark. A recent study demonstrated that H. taeniaeformis exclusively uses murine rodents (rats and mice) as intermediate hosts, with felids as definitive hosts (Lavikainen et al. 2016). Therefore, historical records from non-murine rodents should most likely be attributed to Hydatigera kamiyai Iwaki in Lavikainen et al. 2016.

Genus Taenia Linnaeus, 1758

Taenia crassiceps (Zeder, 1800)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Dryomys nitedula, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus, Microtus tatricus, Mus musculus, Myodes glareolus.

Localities. Domica, Gabčíkovo, PLA Poľana, Roháčska dolina, Vysoké Tatry.

Infection site. Body cavity, subcutaneous tissue.

Reported by. Erhardová (1955a, 1956), Tenora (1955, 1963, 1967), Mituch (1970, 1975), Mituch et al. (1987).

Remark. T. crassiceps is a tapeworm described from Vulpes vulpes L. (Caira et al. 2022). Carnivorous mammals are definitive hosts. While rodents are intermediate hosts, they develop the metacestode (larval) stage of the parasite.

Taenia martis (Zeder, 1803)

Host. Myodes glareolus.

Locality. Čergov Mts.

Infection site. Not specified.

Reported by. Mészáros and Štolmann (1984).

Remark. T. martis is described from Martes martes L. (Caira et al. 2022). Rodents are intermediate hosts.

Taenia pisiformis (Bloch, 1780)

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Mus musculus, Myodes glareolus.

Localities. Košice, PLA Poľana, Roháčska dolina, Sklené Teplice, Vysoké Tatry.

Infection site. Body cavity, liver, mesentery.

Reported by. Erhardová (1956), Tenora (1967), Mituch (1970, 1975).

Remark. Definitive hosts of T. piriformis are canids, intermediate hosts – lagomorphs and rodents (Caira et al. 2022).

Taenia polyacantha Leuckart, 1856

Hosts. Apodemus flavicollis, Apodemus sylvaticus, Chionomys nivalis, Dryomys nitedula, Microtus agrestis, Microtus subterraneus, Microtus tatricus, Myodes glareolus.

Localities. Čergov Mts., Roháčska dolina, South Slovakia, Vysoké Tatry.

Infection site. Abdominal cavity.

Reported by. Erhardová (1955a, 1956), Tenora (1967), Mituch (1970), Mészáros and Štolmann (1984), Mituch et al. (1987).

Remark. T. polyacantha is described from V. vulpes (Caira et al. 2022). Rodents are intermediate hosts.

Genus Versteria Nakao, Lavikainen, Iwaki, Haukisalmi, Konyaev, Oku, Okamoto & Ito, 2013

Versteria mustelae (Gmelin, 1790)

Synonyms. Taenia tenuicollis Rudolphi, 1819, Taenia mustelae Gmelin, 1790.

Hosts. Apodemus agrarius, Apodemus flavicollis, Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus subterraneus, Microtus tatricus, Mus musculus, Muscardinus avellanarius, Myodes glareolus, Rattus norvegicus.

Localities. Čergov Mts., Domica, Michalovce, Nízke Tatry, PLA Poľana, Prešov, Roháčska dolina, Vysoké Tatry.

Infection site. Liver.

Reported by. Erhardová (1955a, 1956, 1958), Tenora (1955, 1963, 1967), Mituch (1970, 1975), Mészáros and Štolmann (1984), Mituch et al. (1987).

Remark. The known definitive hosts are representatives of the genus Martes. Intermediate hosts are primarily rodents (Caira et al. 2022).

Class Trematoda Rudolphi, 1808

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

Family Brachylaimidae Joyeux & Foley, 1930

Genus Brachylaima Dujardin, 1843

Brachylaima recurvum Dujardin, 1845

Hosts. Eliomys quercinus, Glis glis.

Localities. Jablonov, Slovenský kras.

Infection site. Small intestine.

Reported by. Baruš and Tenora (1957), Erhardová (1958).

Brachylaima spp.

Host. Apodemus agrarius.

Locality. Rozhanovce.

Infection site. Small intestine.

Reported by. Ondríková et al. (2010).

Family Diplostomidae Poirier, 1886

Genus Alaria Greville, 1830

Alaria alata (Goeze, 1782)

Host. Apodemus flavicollis.

Locality. Vysoké Tatry.

Infection site. Body cavity.

Reported by. Mituch et al. (1987).

Remark. A. alata requires freshwater snails as its first intermediate host and amphibians as its second intermediate host, uses various vertebrates (e.g., rodents, wild boar) as paratenic hosts, and reaches maturity in carnivorous definitive hosts (Möhl et al. 2009).

Order Plagiorchiida La Rue, 1957

Family Cladorchiidae Fischoeder, 1901

Genus Neostichorchis Jones, 2005

Neostichorchis subtriquetus (Rudolphi, 1814)

Host. Microtus arvalis.

Locality. Vysoké Tatry.

Infection site. Stomach.

Reported by. Mituch et al. (1987).

Genus Stichorchis Fischoeder, 1901

Stichorchis subtriquetrus (Rudolphi, 1814)

Host. Castor fiber.

Localities. Danube River, Hanušovce nad Topľou, Laborec River, Topľa River.

Infection site. Caecum, colon, intestines.

Reported by. Bystrianska et al. (2021), Lazár et al. (2023).

Family Dicrocoeliidae Looss, 1899

Genus Dicrocoelium Dujardin, 1845

Dicrocoelium dendriticum (Rudolphi, 1819)

Synonym. Dicrocoelium lanceatum Stiles & Hassall, 1898.

Host. Rattus norvegicus.

Locality. Košice.

Infection site. Stomach.

Reported by. Mituch (1957).

Genus Lyperosomum Looss, 1899

Lyperosomum armenicum Shcherbakova, 1942

Host. Dryomys nitedula.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Liver.

Reported by. Mituch (1970, 1975).

Family Echinostomatidae Looss, 1899

Genus Echinostoma Rudolphi, 1809

Echinostoma spp.

Hosts. Apodemus flavicollis, Myodes glareolus.

Locality. Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mituch (1970).

Family Notocotylidae Lühe, 1909

Genus Notocotylus Diesing, 1839

Notocotylus noyeri Joyeux, 1922

Hosts. Chionomys nivalis, Microtus agrestis, Microtus arvalis, Microtus oeconomus, Microtus subterraneus.

Localities. Gabčíkovo, PLA Poľana, South Slovakia, Vysoké Tatry.

Infection site. Caecum, small intestine.

Reported by. Erhardová (1955a, 1955b, 1956), Mituch (1970, 1975), Mituch et al. (1987).

Genus Quinqueserialis Skvortsov, 1935

Quinqueserialis quinqueserialis (Barker & Laughlin, 1911)

Host. Microtus arvalis.

Locality. PLA Poľana.

Infection site. Caecum.

Reported by. Mituch (1975).

Family Plagiorchiidae Lühe, 1901

Genus Plagiorchis Lühe, 1899

Plagiorchis elegans (Rudolphi, 1802)

Hosts. Apodemus flavicollis, Myodes glareolus.

Locality. Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mituch et al. (1987).

Plagiorchis muris Tanabe, 1922

Hosts. Microtus oeconomus, Myodes glareolus.

Localities. Gabčíkovo, PLA Poľana, South Slovakia, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Tenora (1955), Erhardová (1956), Mituch (1970, 1975).

Plagiorchis proximus Barker, 1915

Host. Apodemus flavicollis.

Localities. PLA Poľana, Vysoké Tatry.

Infection site. Small intestine.

Reported by. Mituch (1970, 1975).

Family Psilostomidae Looss, 1900

Genus Psilotrema Odhner, 1913

Psilotrema pharyngeatum Grabda, 1954

Host. Microtus oeconomus.

Locality. Gabčíkovo.

Infection site. Small intestine.

Reported by. Erhardová (1958).

Host-parasite list

Apodemus agrarius (Pallas, 1771)

Nematoda – Heligmosomoides polygyrus, Heterakis spumosa, Mastophorus muris, Syphacia (Syphacia) obvelata, Syphacia (Syphacia) stroma, Toxocara spp.; Cestoda – Catenotaenia pusilla, Hymenolepis diminuta, Mesocestoides spp., Paranoplocephala omphalodes, Rodentolepis fraterna, Rodentolepis straminea, Spasskijela lobata, Versteria mustelae; Trematoda – Brachylaima spp.

Apodemus flavicollis (Melchior, 1834)

Acanthocephala – Moniliformis moniliformis; Nematoda – Aonchotheca annulosa, Aspiculuris dinniki, Aspiculuris tetraptera, Calodium hepaticum, Heligmosomoides glareoli, Heligmosomoides polygyrus, Heligmosomum costellatum, Rictularia proni, Syphacia (Seuratoxyuris) frederici, Syphacia (Syphacia) obvelata, Syphacia (Syphacia) stroma, Toxocara spp., Trichinella pseudospiralis, Trichinella spp., Trichuris muris; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Cladotaenia cylindracea, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Mesocestoides spp., Paranoplocephala macrocephala, Paranoplocephala omphalodes, Rodentolepis fraterna, Rodentolepis straminea, Sobolevitaenia unicoronata, Spasskijela lobata, Taenia crassiceps, Taenia pisiformis, Taenia polyacantha, Versteria mustelae; Trematoda – Alaria alata, Echinostoma spp., Plagiorchis elegans, Plagiorchis proximus.

Apodemus sylvaticus (Linnaeus, 1758)

Acanthocephala – Moniliformis moniliformis; Nematoda – Aonchotheca annulosa, Aspiculuris dinniki, Aspiculuris tetraptera, Calodium hepaticum, Glistrongylus gracilis, Heligmosomoides glareoli, Heligmosomoides polygyrus, Heligmosomum costellatum, Porrocaecum depressum, Rictularia proni, Syphacia (Syphacia) obvelata, Toxocara spp., Trichuris muris; Cestoda – Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Paranoplocephala macrocephala, Paranoplocephala omphalodes, Rodentolepis straminea, Taenia crassiceps, Taenia pisiformis, Taenia polyacantha.

Apodemus uralensis (Pallas, 1811)

Nematoda – Aspiculuris dinniki, Heligmosomoides polygyrus, Heligmosomum costellatum, Toxocara spp.; Cestoda – Arostrilepis horrida, Hydatigera taeniaeformis, Hymenolepis diminuta, Rodentolepis fraterna.

Arvicola amphibius (Linnaeus, 1758)

Cestoda – Arostrilepis horrida, Paranoplocephala macrocephala; Nematoda – Calodium hepaticum.

Castor fiber Linnaeus, 1758

Nematoda – Travassosius rufus; Trematoda – Stichorchis subtriquetrus.

Chionomys nivalis (Martins, 1842)

Nematoda – Aspiculuris dinniki, Calodium hepaticum, Heligmosomoides glareoli, Heligmosomum costellatum, Mastophorus muris, Pelodera orbitalis, Syphacia (Syphacia) obvelata; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Eurotaenia gracilis, Kontrimavichusia asymmetrica, Paranoplocephala macrocephala, Paranoplocephala omphalodes, Taenia crassiceps, Taenia polyacantha, Versteria mustelae; Trematoda – Notocotylus noyeri.

Cricetus cricetus (Linnaeus, 1758)

Cestoda – Rodentolepis microstoma.

Dryomys nitedula (Pallas, 1778)

Nematoda – Glistrongylus gracilis, Rictularia proni; Cestoda – Armadolepis myoxi, Armadolepis spasskyi, Hymenolepis spp., Taenia crassiceps, Taenia polyacantha; Trematoda – Lyperosomum armenicum.

Eliomys quercinus (Linnaeus, 1766)

Nematoda – Aonchotheca myoxinitelae, Rictularia proni; Cestoda – Armadolepis spasskyi, Hymenolepis diminuta, Hymenolepis spp.; Trematoda – Brachylaima recurvum.

Glis glis (Linnaeus, 1766)

Nematoda – Glistrongylus gracilis; Acanthocephala – Moniliformis moniliformis; Cestoda – Armadolepis myoxi, Armadolepis spasskyi, Hymenolepis diminuta, Hymenolepis sulcata, Hymenolepis spp.; Trematoda – Brachylaima recurvum.

Marmota marmota (Linnaeus, 1758)

Nematoda – Syphacia (Syphacia) montana, Trichostrongylidae gen. sp.; Cestoda – Ctenotaenia marmotae, Paranoplocephala spp.

Micromys minutus (Pallas, 1771)

Nematoda – Toxocara spp.

Microtus agrestis (Linnaeus, 1761)

Nematoda – Aspiculuris dinniki, Calodium hepaticum, Heligmosomoides glareoli, Heligmosomum costellatum, Heligmosomum halli, Pelodera orbitalis, Porrocaecum spp., Syphacia (Syphacia) montana, Syphacia (Syphacia) obvelata, Trichuris muris; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Eurotaenia gracilis, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Paranoplocephala macrocephala, Paranoplocephala omphalodes, Taenia crassiceps, Taenia polyacantha, Versteria mustelae; Trematoda – Notocotylus noyeri.

Microtus arvalis (Pallas, 1779)

Acanthocephala – Moniliformis moniliformis; Nematoda – Aonchotheca annulosa, Calodium hepaticum, Carolinensis minutus, Glistrongylus gracilis, Heligmosomoides polygyrus, Heligmosomum costellatum, Heligmosomum halli, Pelodera orbitalis, Rictularia proni, Syphacia (Syphacia) nigeriana, Syphacia (Syphacia) obvelata, Toxocara spp., Trichostrongylus retortaeformis, Trichuris muris; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Paranoplocephala macrocephala, Paranoplocephala microti, Paranoplocephala omphalodes, Rodentolepis fraterna, Rodentolepis straminea, Sobolevitaenia unicoronata, Spasskijela lobata, Taenia crassiceps, Versteria mustelae; Trematoda – Neostichorchis subtriquetus, Notocotylus noyeri, Quinqueserialis quinqueserialis.

Microtus oeconomus (Pallas, 1776)

Nematoda – Heligmosomoides polygyrus, Syphacia (Syphacia) obvelata, Trichuris muris; Cestoda – Hymenolepis diminuta, Paranoplocephala macrocephala, Taenia crassiceps; Trematoda – Notocotylus noyeri, Plagiorchis muris, Psilotrema pharyngeatum.

Microtus subterraneus (Selys-Longchamps, 1836)

Nematoda – Aspiculuris dinniki, Carolinensis minutus, Heligmosomoides glareoli, Heligmosomoides polygyrus, Heligmosomum costellatum, Heligmosomum halli, Mastophorus muris, Pelodera orbitalis, Porrocaecum spp., Syphacia (Syphacia) montana, Syphacia (Syphacia) obvelata; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Eurotaenia gracilis, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Microticola blanchardi, Paranoplocephala macrocephala, Paranoplocephala omphalodes, Paranoplocephala spp., Taenia crassiceps, Taenia polyacantha, Versteria mustelae; Trematoda – Notocotylus noyeri.

Microtus tatricus Kratochvíl, 1952

Nematoda – Aspiculuris dinniki, Heligmosomoides glareoli, Heligmosomum costellatum, Heligmosomum halli, Pelodera orbitalis, Syphacia (Syphacia) montana; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Eurotaenia gracilis, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Microticola blanchardi, Paranoplocephala macrocephala, Paranoplocephala omphalodes, Rodentolepis fraterna, Taenia crassiceps, Taenia polyacantha, Versteria mustelae.

Mus musculus Linnaeus, 1758

Acanthocephala – Moniliformis moniliformis; Cestoda – Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Paranoplocephala macrocephala, Rodentolepis fraterna, Spasskijela lobata, Taenia crassiceps, Taenia pisiformis, Versteria mustelae; Nematoda – Aonchotheca annulosa, Aspiculuris tetraptera, Heligmosomoides polygyrus, Heligmosomum costellatum, Mastophorus muris, Porrocaecum depressum, Rictularia proni, Syphacia (Syphacia) obvelata, Toxocara spp., Trichuris muris.

Mus spicilegus Petényi, 1882

Nematoda – Heligmosomoides polygyrus, Syphacia (Syphacia) obvelata; Cestoda – Hymenolepis diminuta.

Muscardinus avellanarius (Linnaeus, 1758)

Nematoda – Heligmosomoides polygyrus, Syphacia (Syphacia) obvelata; Cestoda – Versteria mustelae.

Myodes glareolus (Schreber, 1780)

Acanthocephala – Moniliformis moniliformis; Nematoda – Aonchotheca annulosa, Aspiculuris dinniki, Calodium hepaticum, Carolinensis minutus, Heligmosomoides glareoli, Heligmosomoides polygyrus, Heligmosomum costellatum, Heligmosomum halli, Heligmosomum mixtum, Mastophorus muris, Pelodera orbitalis, Rictularia proni, Rictularia spp., Syphacia (Syphacia) montana, Syphacia (Syphacia) obvelata, Toxocara spp.; Cestoda – Anoplocephaloides dentata, Arostrilepis horrida, Catenotaenia henttoneni, Catenotaenia pusilla, Cladotaenia cylindracea, Eurotaenia gracilis, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Mesocestoides spp., Microticola blanchardi, Paranoplocephala macrocephala, Paranoplocephala microti, Paranoplocephala omphalodes, Paranoplocephala spp., Rodentolepis fraterna, Rodentolepis straminea, Spasskijela lobata, Taenia crassiceps, Taenia martis, Taenia pisiformis, Taenia polyacantha, Versteria mustelae; Trematoda – Echinostoma spp., Plagiorchis elegans, Plagiorchis muris.

Ondatra zibethicus (Linnaeus, 1766)

Cestoda – Echinococcus multilocularis, Hydatigera taeniaeformis.

Rattus norvegicus (Berkenhout, 1769)

Acanthocephala – Moniliformis moniliformis; Nematoda – Aonchotheca annulosa, Calodium hepaticum, Heterakis spumosa, Liniscus papillosus, Mastophorus muris, Strongyloides ratti, Trichinella pseudospiralis, Trichinella spp., Trichosomoides crassicauda, Trichuris muris; Cestoda – Arostrilepis horrida, Catenotaenia pusilla, Hydatigera taeniaeformis, Hymenolepis diminuta, Kontrimavichusia asymmetrica, Paranoplocephala omphalodes, Rodentolepis fraterna, Rodentolepis straminea, Versteria mustelae; Trematoda – Dicrocoelium dendriticum.

Sciurus vulgaris Linnaeus, 1758

Nematoda – Enterobius (Enterobius) sciuri, Porrocaecum depressum, Trichostrongylus retortaeformis; Cestoda – Catenotaenia dendritica, Hydatigera taeniaeformis.

Sicista betulina (Pallas, 1779)

Nematoda – Heligmosomoides polygyrus; Cestoda – Arostrilepis horrida.

Spermophilus citellus (Linnaeus, 1766)

Acanthocephala – Moniliformis moniliformis; Cestoda – Hymenolepis diminuta, Hymenolepis spp.; Nematoda – Capillaria spp., Citellina spp., Trichostrongylidae gen. sp.

Discussion

The presented checklist summarises information on the helminth fauna of rodents inhabiting Slovakia. Although the list includes extensive data on helminth species from 27 rodent species, encompassing the entire available research period and information from across the country, its completeness may be limited for several reasons. These include incomplete taxonomic coverage of hosts, uneven spatial and temporal coverage of research, and methodological changes in studies.

Thirty-one rodent species have been reported in Slovakia, but two of them, Sicista subtilis (Pallas, 1773) and Rattus rattus (Linnaeus, 1758), are currently unconfirmed in the country (Krištofík and Danko 2012). Thus, of the 29 rodent species presently found in Slovakia, data on their helminths are available for 27 of them. Information on the helminths of two rodent species – Myocastor coypus (Molina, 1782) and Arvicola scherman (Shaw, 1801) – is absent, which is related to the limited distribution of these species in the country. In particular, M. coypus is an introduced species from South America with a fragmented distribution in Slovakia, and A. scherman is a rare species with only a single documented occurrence (Krištofík and Danko 2012).

Although data on helminths of rodents were collected across Slovakia, the intensity of research varied significantly between regions. The highest number of studies was conducted in Eastern Slovakia, with the Tatra National Park being the most extensively studied area, particularly in the 1950s and 1980s (see Suppl. material 1). Furthermore, research effort was unevenly distributed across host species, with most rodent species being extensively studied, while the helminth diversity of others remains poorly documented.

Different methods were used for helminth identification in the studies underlying this checklist, including microscopic examination of morphological features, coprological analysis, enzyme-linked immunosorbent assay (ELISA), and polymerase chain reaction (PCR). However, in most of these studies, morphological identification was the only approach used. Although morphology is fundamental to the taxonomy and systematics of helminths, morphological identification has certain limitations (Miller et al. 2024). This limitation is especially acute for cryptic and closely related species, which often remain unrecognised or misidentified when morphological features are the only criteria used. Molecular techniques have greatly expanded diagnostic capabilities; nonetheless, morphological examination often remains the most economical and rapid means of identifying well-characterised helminths.

Our comparison with the most comprehensive Polish helminth catalogue (Pojmańska et al. 2007) revealed 71 helminth species recorded in Poland from 17 rodent hosts: 23 cestodes, 11 trematodes, 36 nematodes, and one acanthocephalan. Although the overall helminth species richness is similar to that of Slovakia, host coverage differs. It is worth noting that Slovakia and Poland share an almost identical rodent fauna (Krištofík and Danko 2012; Tykarski 2022); M. spicilegus occurs only in Slovakia, whereas Spermophilus suslicus (Güldenstaedt, 1770) is found exclusively in Poland. In Slovakia, helminthological data are available for 27 of the 31 rodent species known to occur in the country, whereas in Poland, records exist for only 17 of the 31 species. Subsequent comparison reveals 44 helminth species shared by both countries, 29 unique to Slovakia, and 27 unique to Poland, resulting in a Jaccard similarity of 44%. Although our comparison is crude, likely influenced by biases discussed above, it suggests a high probability of finding additional helminths of rodents in Slovakia, particularly in border areas.

Since Krištofík and Danko (2012), five additional helminths (R. microstoma, S. subtriquetrus, Citellina spp., T. britovi, and T. rufus) have been recorded in Slovakia, highlighting the potential for further discoveries. We expect that helminth species new to Slovakia will most likely be found in their larval stages. Currently, of the 77 helminth taxa documented in Slovak rodents, 13 are documented at larval stages (Porrocaecum spp., Toxocara spp., C. cylindracea, all representatives of the family Taeniidae, Mesocestoides spp., and A. alata), and one appears to be a case of accidental infection (T. retortaeformis).

As previously mentioned, this checklist is partly based on reports produced at the Institute of Parasitology, Slovak Academy of Sciences (Mituch 1956, 1957, 1970, 1975; Mituch et al. 1987). Voucher material generated during those studies was deposited in the Institute’s collection. In 1996, the entire collection, including numerous helminths from hosts other than rodents, was transferred to the East-Slovak Museum (Košice, Slovakia), where it is still curated. The most valuable specimens, notably type material, have since been catalogued by Dudiňák and Špakulová (2005). Although the present review relies solely on literature sources, we mention the collection here for the benefit of future researchers interested in examining the original material.

In conclusion, although the present checklist provides a comprehensive synthesis of available data on rodent helminths in Slovakia, our knowledge of their diversity remains incomplete. Further research involving broader host sampling, geographically balanced study designs, and modern molecular techniques for parasite identification is needed to advance understanding in Slovakia.

Acknowledgements

We gratefully acknowledge Dr Christopher Glasby for providing linguistic editing and valuable improvements to the manuscript. We thank the anonymous reviewers for the time they devoted to evaluating our work. We also thank Dr Boyko Georgiev, the subject editor, for his additional guidance and comments throughout the review process.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

During the preparation of this article, the authors used Grammarly (https://www.grammarly.com/) to correct grammatical errors and enhance readability. After using this tool, the authors reviewed and edited the content as needed. The authors accept full responsibility for the content of the published article.

Funding

The research was financially supported by the Slovak Research and Development Agency under the contract No. APVV-21-0166 and Tubitak-2024-01, and by the NextGenerationEU through the Recovery and Resilience Plan for Slovakia under project No. 09I03-03-V01-00046.

Author contributions

Conceptualization: YS. Data curation: YS. Resources: MM. Validation: MM, YS, VK, ZH. Writing – original draft: YS. Writing – review and editing: MM, VK, ZH.

Author ORCIDs

Yaroslav Syrota https://orcid.org/0000-0002-8070-9823

Zuzana Hurníková https://orcid.org/0000-0002-6816-2992

Vitaliy Kharchenko https://orcid.org/0000-0002-3824-2078

Martina Miterpáková https://orcid.org/0000-0002-3742-948X

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

References

Anderson RC (2000) Nematode parasites of vertebrates: Their development and transmission, 2^nd ed. CABI International, Wallingford, 650 pp. https://doi.org/10.1079/9780851994215.0000

Atkinson CT, Thomas NJ, Hunter DB [Eds] (2008) Parasitic Diseases of Wild Birds. Wiley-Blackwell, Ames, Iowa, 595 pp. https://doi.org/10.1002/9780813804620

Audebert F, Hoste H, Durette-Desset MC (2002) Life cycle of Trichostrongylus retortaeformis in its natural host, the rabbit (Oryctolagus cuniculus). Journal of Helminthology 76(3): 189–192. https://doi.org/10.1079/JOH2002126

Baruš V, Hrabě V (1991) Larvae of Rhabditis orbitalis Sudhaus et Schulte, 1986 in the mountain rodents in Czech and Slovak Federative Republic. Helminthologia 28: 93–97.

Baruš V, Tenora F (1957) Příspěvek k helmintofauně plchovitých (Myoxidae) jižního Slovenska. [Contribution to the helminth fauna of dormice (Myoxidae) in southern Slovakia. ] Československá parasitology 4: 53–56. [In Czech]

Behnke JM, Jackson JA (2024) Aonchotheca annulosa and Aonchotheca murissylvatici, which is which? A reappraisal of the gastrointestinal Aonchotheca (Nematoda: Capillariidae) species common in wood mice and bank voles. Parasitology 151(14): 1–11. https://doi.org/10.1017/S0031182024001471

Biserkov V (1998) Heligmosomoides glareoli Baylis, 1928 (Nematoda: Heligmosomidae): redescription and synonymy. Systematic Parasitology 41(3): 179–186. https://doi.org/10.1023/A:1006095525983

Bystrianska J, Papajová I, Šmiga Ľ, Šoltys J, Majláthová V, Majláth I, Gomboš F, Kleban J (2021) First report on parasites of European beavers in the Slovak Republic. Parasitology Research 120(1): 355–358. https://doi.org/10.1007/s00436-020-06943-6

Caira JN, Jensen K, Barbeau E (2022) Global Cestode Database. https://tapewormdb.uconn.edu/ [Accessed July 2024]

Dubinský P, Havasiová-Reiterová K, Peťko B, Hovorka I, Tomašovičová O (1995) Role of small mammals in the epidemiology of toxocariasis. Parasitology 110(2): 187–193. https://doi.org/10.1017/S0031182000063952

Dudiňák V, Špakulová M (2005) A survey of helminth species originally described from the territory of Slovakia supplemented by a list of the type material deposited in the East-Slovakian Museum (Východoslovenské múzeum) in Košice. Helminthologia 42(4): 233–245.

Dursahinhan A, Kenkel D, Gardner S (2023) Helminth and Protozoan parasites of subterranean rodents (Chordata, Mammalia, Rodentia) of the World. ZooKeys 1151: 159–203. https://doi.org/10.3897/zookeys.1151.97126

Erhardová B (1955a) Helmintofauna hrabošů a myší Tatranského národního parku. Zoologicke a Entomologicke Listy 4(4): 353–366. [Helminth fauna of voles and mice from the Tatra National Park.] [In Czech]

Erhardová B (1955b) Příspěvek k poznání motolic u Microtus oeconomus mehelyi (Rodentia, Muridae). [Contribution to the knowledge of flukes in Microtus oeconomus mehelyi (Rodentia, Muridae). ] Československá parazitologie 2: 38–40. [In Czech]

Erhardová B (1956) Parasitičtí červi našich myšovitých hlodavců II. [Parasitic worms of our murine rodents II. ] Československá parasitologie 3: 49–66. [In Czech]

Erhardová B (1958) Parasitičtí červi hlodavců Československa. [Parasitic worms of the rodents of Czechoslovakia. ] Československá parasitologie 5(1): 27–93. [In Czech]

Feliu C, Špakulová M, Casanova J, Renaud F, Morand S, Hugot J-P, Santalla F, Durand P (2000) Genetic and morphological heterogeneity in small rodent whipworms in southwestern Europe: characterization of Trichuris muris and description of Trichuris arvicolae n. sp. (Nematoda: Trichuridae). The Journal of Parasitology 86(3): 442–449. https://doi.org/10.1645/0022-3395(2000)086[0442:GAMHIS]2.0.CO;2

GBIF.org (2024) GBIF Home Page. https://www.gbif.org [Accessed July 2024]

Greenspoon L, Krieger E, Sender R, Rosenberg Y, Bar-On YM, Moran U, Antman T, Meiri S, Roll U, Noor E, Milo R (2023) The global biomass of wild mammals. Proceedings of the National Academy of Sciences of the United States of America 120(10): e2204892120. https://doi.org/10.1073/pnas.2204892120

Hamzavi Y, Khodayari MT, Davari A, Shiee MR, Karamati SA, Raeghi S, Jabarmanesh H, Bashiri H, Bozorgomid A (2024) A systematic review and meta-analysis on prevalence of gastrointestinal helminthic infections in rodents of Iran: An emphasis on zoonotic aspects. Heliyon 10(11): e31955. https://doi.org/10.1016/j.heliyon.2024.e31955

Hanzelová V, Ryšavý B (1996) Synopsis of cestodes in Slovakia IV. Hymenolepididae (continued). Helminthologia 33: 213–222.

Hanzelová V, Ryšavý B, Šnábel V (1995) Synopsis of cestodes in Slovakia III. Cyclophyllidea: Amabiliidae, Acoleidae, Catenotaeniidae, Davaineidae, Hymenolepididae (in part). Helminthologia 32: 67–73.

Haukisalmi V, Henttonen H (2003) What is Paranoplocephala macrocephala (Douthitt, 1915) (Cestoda: Anoplocephalidae)? Systematic Parasitology 54(1): 53–69. https://doi.org/10.1023/A:1022141809571

Haukisalmi V, Hardman LM, Hoberg EP, Henttonen H (2014) Phylogenetic relationships and taxonomic revision of Paranoplocephala Lühe, 1910 sensu lato (Cestoda, Cyclophyllidea, Anoplocephalidae). Zootaxa 3873(4): 371–415. https://doi.org/10.11646/zootaxa.3873.4.3

Hovorka J (1968) Index helminthum. [Index of helminths.] In: Studia helminthologica II. Institutio Helminthologiae Academiae Scientiarum Slovacae Cassoviae ČSSR. Košice, 221–279. [In Latin]

Hurníková Z, Šnábel V, Pozio E, Reiterová K, Hrčková G, Halásová D, Dubinský P (2005) First record of Trichinella pseudospiralis in the Slovak Republic found in domestic focus. Veterinary Parasitology 128(1): 91–98. https://doi.org/10.1016/j.vetpar.2004.11.013

Hurníková Z, Syrota Y, Komorová P, Chovancová G, Miterpáková M (2025) Contribution to host diversity, genetic diversion, and epidemiology of Trichinella pseudospiralis in Slovakia, central Europe. Veterinary Parasitology 333: 110332. https://doi.org/10.1016/j.vetpar.2024.110332

Islam MM, Farag E, Hassan M, Bansal D, Awaidy SA, Abubakar A, Al-Romaihi H, Mkhize-Kwitshana Z (2020) Helminth parasites among rodents in the Middle East countries: A systematic review and meta-analysis. Animals (Basel) 10(12): 2342. https://doi.org/10.3390/ani10122342

Jackson JA, Friberg IM (2022) Carolinensis minutus (Dujardin, 1845) Travassos 1937 (Nematoda, Heligmonellidae) in Microtus agrestis in the United Kingdom. Check List 18(4): 845–848. https://doi.org/10.15560/18.4.845

Jarošová J, Šnábel V, Cavallero S, Chovancová G, Hurníková Z, Antolová D (2020) The mouse bile duct tapeworm, in free-living small mammals in Slovakia: Occurrence and genetic analysis. Helminthologia 57(2): 120–128. https://doi.org/10.2478/helm-2020-0022

Jászayová A, Režnarová J, Chovancová G, Kostygov AY, Yurchenko V, Antolová D, Zwijacz-Kozica T, Csanády A, Hurníková Z (2023) A study of oribatid mites as potential intermediate hosts of anoplocephalid tapeworms of Tatra chamois and Tatra marmots from the Tatra Mountains, Central Europe, and report of a new intermediate host for Andrya cuniculi, the parasite of Leporidae. Life (Basel, Switzerland) 13(4): 955. https://doi.org/10.3390/life13040955

Jesudoss Chelladurai JRJ, Brewer MT (2021) Global prevalence of Mesocestoides infections in animals – A systematic review and meta-analysis. Veterinary Parasitology 298: 109537. https://doi.org/10.1016/j.vetpar.2021.109537

Kocianová-Adamcová M, Špakulová M, Kocianová E (2006) Long-term variation in an occurrence of Rhabditis orbitalis parasitic larvae (Nematoda, Rhabditidae) in the eyes of montane rodents. Helminthologia 43(4): 232–236. https://doi.org/10.2478/s11687-006-0043-8

Komorová P, Sitko J, Špakulová M, Hurníková Z, Sałamatin R, Chovancová G (2017) New data on helminth fauna of birds of prey (Falconiformes, Accipitriformes, Strigiformes) in the Slovak Republic. Helminthologia 54(4): 314–321. https://doi.org/10.1515/helm-2017-0038

Krištofík J, Danko Š (2012) Cicavce Slovenska: rozšírenie, bionómia a ochrana. [Mammals of Slovakia: distribution, bionomy, and conservation. ] VEDA, Bratislava, 712 pp. [In Slovak]

Lavikainen A, Iwaki T, Haukisalmi V, Konyaev SV, Casiraghi M, Dokuchaev NE, Galimberti A, Halajian A, Henttonen H, Ichikawa-Seki M, Itagaki T, Krivopalov AV, Meri S, Morand S, Näreaho A, Olsson GE, Ribas A, Terefe Y, Nakao M (2016) Reappraisal of Hydatigera taeniaeformis (Batsch, 1786) (Cestoda: Taeniidae) sensu lato with description of Hydatigera kamiyai n. sp. International Journal for Parasitology 46(5): 361–374. https://doi.org/10.1016/j.ijpara.2016.01.009

Lazár J, Šmigová J, Šmiga Ľ, Lazár P, Čurlík J, Papajová I (2023) Morphological and molecular identification of adult Stichorchis subtriquetrus in beaver in Slovakia. Veterinary Research Communications 47(4): 2161–2164. https://doi.org/10.1007/s11259-023-10074-5

Macko J, Ryšavý B, Hanzelová V, Králová I (1994) Synopsis of cestodes in Slovakia II. Cyclophyllidea: Mesocestoididae, Tetrabothriidae, Nematotaeniidae, Taeniidae. Helminthologia 31: 95–103.

Mahesh Kumar J, Reddy PL, Aparna V, Srinivas G, Nagarajan P, Venkatesan R, Sreekumar C, Sesikaran B (2006) Strobilocercus fasciolaris infection with hepatic sarcoma and gastroenteropathy in a Wistar colony. Veterinary Parasitology 141(3–4): 362–367. https://doi.org/10.1016/j.vetpar.2006.05.029

Makarikov AA, Binkienė R (2022) Redescription and taxonomic position of Rodentolepis (sensu lato) asymmetrica (Janicki, 1904), with the erection of Kontrimavichusia n. g. (Eucestoda: Hymenolepididae) from arvicoline rodents (Rodentia: Cricetidae). Journal of Helminthology 96: e63. https://doi.org/10.1017/S0022149X22000505

Makarikov AA, Georgiev BB (2020) Review of records of hymenolepidids (Eucestoda: Hymenolepididae) from dormice (Rodentia: Gliridae) in Europe, with a redescription of Armadolepis spasskyi Tenora & Baruš, 1958 and the description of A. genovi n. sp. Systematic Parasitology 97(1): 83–98. https://doi.org/10.1007/s11230-019-09891-7

Mészáros F, Štolmann A (1984) Results of complex parasitological investigations on small mammals in the Cergov Mountains (Western Carpathians, Czechoslovakia). Miscellanea zoologica hungarica 2: 7–11.

Miller ML, Rota C, Welsh A (2024) Transforming gastrointestinal helminth parasite identification in vertebrate hosts with metabarcoding: A systematic review. Parasites & Vectors 17(1): 311. https://doi.org/10.1186/s13071-024-06388-1

Miterpáková M, Antolová D, Ševčíková Z, Stanko M, Dinkel A, Gašpar V, Dubinský P (2006) Echinococcus multilocularis in musk rat (Ondatra zibethicus): The first finding of the parasite in naturally infected rodent in the Slovak Republic. Helminthologia 43(2): 76–80. https://doi.org/10.2478/s11687-006-0015-z

Miterpáková M, Hurníková Z, Komorová P, Stanko M, Chovancová G, Syrota Y (2024) Micromammals as a reservoir for the zoonotic nematode Calodium hepaticum (syn. Capillaria hepatica) in recreational areas of Slovakia. Current Research in Parasitology & Vector-borne Diseases 6: 100214. https://doi.org/10.1016/j.crpvbd.2024.100214

Mituch J (1956) Výskum ohnísk trichinelózy na Slovensku. Záverečná zpráva. [Research on trichinellosis foci in Slovakia. Final report.] Helmintologický ústav SAV [Institute of Parasitology of the Slovak Academy of Sciences], Košice. [In Slovak]

Mituch J (1957) Helmintofauna potkana severného na Slovensku. Záverečná zpráva. [Helminth fauna of the northern rat in Slovakia. Final report.] Helmintologický ústav SAV [Institute of Parasitology of the Slovak Academy of Sciences], Košice. [In Slovak]

Mituch J (1970) Helmintofauna drobných cicavcov a mäsožravcov TANAP. Záverečná zpráva. [Helminth fauna of small mammals and carnivores in TANAP. Final report.] Helmintologický ústav SAV [Institute of Parasitology of the Slovak Academy of Sciences], Košice. [In Slovak]

Mituch J (1975) Helmintocenózy cicavcov masívu Poľany. Záverečná správa. [Helminth communities of mammals in the Poľana Massif. Final report.] Helmintologický ústav SAV [Institute of Parasitology of the Slovak Academy of Sciences], Košice. [In Slovak]

Mituch J, Hovorka J, Hovorka I, Tenkacova I (1987) Helminty a helmintocenózy cicavcov karpatského oblúka. Záverečná správa. [Helminths and helminth communities of mammals in the Carpathian Arc. Final report.]. Helmintologický ústav SAV [Institute of Parasitology of the Slovak Academy of Sciences], Košice. [In Slovak]

Möhl K, Große K, Hamedy A, Wüste T, Kabelitz P, Lücker E (2009) Biology of Alaria spp. and human exposition risk to Alaria mesocercariae – a review. Parasitology Research 105(1): 1–15. https://doi.org/10.1007/s00436-009-1444-7

Nemys (2024) Nemys: World Database of Nematodes. Available from https://nemys.ugent.be/ Accessed July 2024. https://doi.org/10.14284/366

Okamoto M, Urushima H, Iwasa M, Hasegawa H (2007) Phylogenetic relationships of rodent pinworms (genus Syphacia) in Japan inferred from mitochondrial CO1 gene sequences. The Journal of Veterinary Medical Science 69(5): 545–547. https://doi.org/10.1292/jvms.69.545

Ondríková J, Miklisová D, Ribas A, Stanko M (2010) The helminth parasites of two sympatric species of the genus Apodemus (Rodentia, Muridae) from south-eastern Slovakia. Acta Parasitologica 55(4): 369–378. https://doi.org/10.2478/s11686-010-0043-1

Panti-May JA, Duarte-Jiménez A, Hernández-Batancourt SF, Rodríguez-Vivas RI (2021) A checklist of the helminth parasites of invasive murid rodents in Mexico. Therya 12(1): 85–85. https://doi.org/10.12933/therya-21-1043

Pojmańska T, Niewiadomska K, Okulewicz A (2007) Pasożytnicze helminty Polski: gatunki, żywiciele, białe plamy. [Parasitic helminths of Poland: species, hosts, and knowledge gaps. ] Polskie Towarzystwo Parazytologiczne, Warszawa, 360 pp. [In Polish]

Preisser W (2019) Latitudinal gradients of parasite richness: A review and new insights from helminths of cricetid rodents. Ecography 42(7): 1315–1330. https://doi.org/10.1111/ecog.04254

Preisser WC, Falcón-Ordaz J (2019) A checklist of the parasitic helminths of cricetid and heteromyid rodents in Mexico. Therya 10(3): 329–329. https://doi.org/10.12933/therya-19-787

Rajský D (1985) Ondatra pižmová ako medzihostiteľ vo vývinovom cykle pásomnice mačacej. Veterinarÿstvi 35: 138. [The muskrat as an intermediate host in the life cycle of the cat tapeworm.] [In Slovak]

Ribas A, Ondríková J, Stanko M (2009) Helminths of the mound-building mouse Mus spicilegus (Rodentia) in Slovakia. In: Nevo E, Mares M, Johnson C, Marroig G, Marque P (Eds) 10^th International Mammalogical Congress, Mendoza (Argentina), August 9–14, 182–182.

Sałamatin R, Cielecka D, Karbowiak G, Stanko M (2005) Hymenolepis sulcata (von Linstow, 1879): occurrence in dormice, Glis glis (Rodentia) in Slovak Republic. Helminthologia 42(3): 185–186. [Conference abstract: Proceedings of the Thirteenth Helminthological Days, Jedovnice, Czech Republic, May 9–13, 2005.]

Smales LR (1997) A review of the helminth parasites of Australian rodents. Australian Journal of Zoology 45(5): 505–521. https://doi.org/10.1071/ZO97013

Spasskaya LP, Spasskii AA (1977) Cestody ptic SSSR. Dilepididy sukhoputnykh ptic [Cestodes of birds in the USSR. Dilepididae of terrestrial birds]. Nauka, Moscow, 301 pp. [In Russian]

Stewart A, Lowe A, Smales L, Bajer A, Bradley J, Dwużnik D, Franssen F, Griffith J, Stuart P, Turner C, Zaleśny G, Behnke JM (2018) Parasitic nematodes of the genus Syphacia Seurat, 1916 infecting Muridae in the British Isles, and the peculiar case of Syphacia frederici. Parasitology 145(3): 269–280. https://doi.org/10.1017/S0031182017001470

Štrkolcová G, Ciho A, Kaduková M, Janošková N, Schreiberová A (2024) Intestinal parasites of the endangered European ground squirrel (Spermophilus citellus) populations in Slovakia. Veterinary Parasitology. Regional Studies and Reports 50: 101014.

Tenora F (1955) Predbezná zpráva o cizopasnych cervech hraboše tatranského Microtus (Chionomys) mirhanreini. Zoologicke a Entomologicke Listy 4(2): 194–200. [Preliminary report on parasitic worms of the Tatra vole Microtus (Chionomys) mirhanreini.] [In Slovak]

Tenora F (1961) Nález tasemnice Ctenotaenia marmotae (Fröhlich, 1862) Railliet, 1893 v ČSSR. [Finding of the tapeworm Ctenotaenia marmotae (Fröhlich, 1862) Railliet, 1893 in ČSSR (Czechoslovak Socialist Republic).] Zoologické listy 10: 396. [In Czech]

Tenora F (1963) Přehled cizopasných červů myšic rodu Apodemus v ČSSR. [Overview of parasitic worms of wood mice of the genus Apodemus in the ČSSR (Czechoslovak Socialist Republic). ] Zoologické listy 12: 331–336. [In Czech]

Tenora F (1965) Supplementary notes on hymenolepidis tapeworms parasitizing glirid dormice in south Slovakian limestone area. Československá parazitologie 12: 299–303.

Tenora F (1967) The helminthfauna of small rodents of the Rohacská Dolina Valley (Liptovské Hole Mts., Slovakia). Acta Scientiarum Naturalium Brno 1: 29–68.

Tenora F, Baruš V (1958) Armadolepis spasskyi sp. n., eine neue Bandwurmart aus Nagern der Fam. Myoxidae. Folia Zoologica 4: 339–342. [In German]

Tenora F, Murai E (1980) The genera Anoplocephaloides and Paranoplocephala (Cestoda) parasites of Rodentia in Europe. Acta Zoologica Academiae Scientiarum Hungaricae 26: 263–284.

Tykarski P [Ed.] (2022) Digital Catalogue of Biodiversity of Poland – Animalia: Chordata: Vertebrata: Mammalia. https://ipt.gbif.pl/resource?r=dcbp-animalia-chordata-vertebrata-mammalia [Accessed June 2025]

Wolff JO, Sherman PW (2007) Rodent Societies as Model Systems. In: Wolff JO, Sherman PW (Eds) Rodent Societies: An Ecological and Evolutionary Perspective. University of Chicago Press, Chicago, 3–7. https://doi.org/10.7208/chicago/9780226905389.001.0001

WoRMS Editorial Board (2024) World Register of Marine Species. Available from: https://www.marinespecies.org Accessed July 2024. https://doi.org/10.14284/170

Supplementary material

Supplementary material 1

Detailed information for each locality

Yaroslav Syrota, Zuzana Hurníková, Vitaliy Kharchenko, Martina Miterpáková

Data type: (measurement/occurrence/multimedia/etc.)

Explanation note: This file is an Excel workbook. It consists of three sheets: data, locations, and references. The data sheet systematically compiles the information presented in the main text of the article into a tabular format optimised for computational processing. The locations sheet contains geographic information, including approximate coordinates, for each locality mentioned in the data sheet. The references sheet lists the bibliographic sources used in compiling the dataset.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Download file (70.40 kb)

﻿Abstract

Key words:

﻿Introduction

﻿Material and methods

﻿Results

﻿Annotated parasite-host list

﻿Phylum Acanthocephala Rudolphi, 1802

Moniliformis moniliformis (Bremser, 1811)

﻿Phylum Nematoda Cobb, 1932

Porrocaecum depressum (Zeder, 1800)

Porrocaecum spp.

﻿Genus Toxocara Stiles & Hassall, 1905

Toxocara spp.

﻿Family Heligmosomidae Cram, 1927

Carolinensis minutus (Dujardin, 1845)

﻿Genus Glistrongylus Durette Desset, Digiani, Kilani & Geffard Kuriyama, 2017

Glistrongylus gracilis (Leuckart, 1842)

﻿Genus Heligmosomoides Hall, 1916

Heligmosomoides glareoli Baylis, 1928

Heligmosomoides polygyrus (Dujardin, 1845)

﻿Genus Heligmosomum Railliet & Henry, 1909

Heligmosomum costellatum (Dujardin, 1845)

Heligmosomum halli (Schulz, 1926)

Heligmosomum mixtum Schulz, 1929

﻿Family Heterakidae Railliet & Henry, 1912

Heterakis spumosa Schneider, 1866

﻿Family Heteroxynematidae Skrjabin & Shikhobalova, 1948

Aspiculuris dinniki Schulz, 1927

Aspiculuris tetraptera Nitzsch, 1821

﻿Family Oxyuridae Weinland, 1858

Citellina spp.

﻿Genus Enterobius Leach, 1853

Enterobius (Enterobius) sciuri (Cameron, 1932)

﻿Genus Syphacia Seurat, 1916

Syphacia (Seuratoxyuris) frederici (Roman, 1945)

Syphacia (Syphacia) montana (Yamaguti, 1943)

Syphacia (Syphacia) nigeriana (Baylis, 1928)

Syphacia (Syphacia) obvelata (Rudolphi, 1802)

Syphacia (Syphacia) stroma (Linstow, 1884)

﻿Family Rhabditidae

Pelodera orbitalis (Sudhaus & Schulte, 1986)

﻿Family Rictulariidae Railliet, 1916

Rictularia proni Seurat, 1915

Rictularia spp.

﻿Family Spirocercidae Chitwood & Wehr, 1932

Mastophorus muris (Gmelin, 1790)

﻿Family Strongyloididae Chitwood & McIntosh, 1934

Strongyloides ratti Sandground, 1925

﻿Family Trichostrongylidae Leiper, 1912

Travassosius rufus Khalil, 1922

﻿Genus Trichostrongylus Looss, 1905

Trichostrongylus retortaeformis (Zeder, 1800)

Trichostrongylidae gen. sp.

﻿Class Enoplea Inglis, 1983

Aonchotheca annulosa (Dujardin, 1845)

Aonchotheca myoxinitelae (Diesing, 1851)

﻿Genus Calodium Dujardin, 1845

Calodium hepaticum (Bancroft, 1893)

﻿Genus Capillaria

Capillaria spp.

﻿Genus Liniscus Dujardin, 1845

Liniscus papillosus (Polonio, 1860)

﻿Family Trichinellidae Ward, 1907

Trichinella britovi Pozio, La-Rosa, Murrell & Lichtenfels, 1992

Trichinella pseudospiralis Garkavi, 1972

Trichinella spp.

﻿Family Trichosomoididae Yorke & Maplestone, 1926

Trichosomoides crassicauda (Bellingham, 1840)

﻿Family Trichuridae Railliet, 1915

Trichuris muris (Schrank, 1788)

﻿Phylum Platyhelminthes Minot, 1876

Anoplocephaloides dentata (Galli-Valerio, 1905)

﻿Genus Ctenotaenia Railliet, 1893

Ctenotaenia marmotae (Frölich, 1802)

﻿Genus Eurotaenia Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Eurotaenia gracilis (Tenora & Murai, 1980)

﻿Genus Microticola Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Microticola blanchardi (Moniez, 1891)

﻿Genus Paranoplocephala Lühe, 1910

Paranoplocephala macrocephala (Douthitt, 1915)

Abstract

Introduction

Material and methods

Results

Annotated parasite-host list

Phylum Acanthocephala Rudolphi, 1802

Phylum Nematoda Cobb, 1932

Genus Toxocara Stiles & Hassall, 1905

Family Heligmosomidae Cram, 1927

Genus Glistrongylus Durette Desset, Digiani, Kilani & Geffard Kuriyama, 2017

Genus Heligmosomoides Hall, 1916

Genus Heligmosomum Railliet & Henry, 1909

Family Heterakidae Railliet & Henry, 1912

Family Heteroxynematidae Skrjabin & Shikhobalova, 1948

Family Oxyuridae Weinland, 1858

Genus Enterobius Leach, 1853

Genus Syphacia Seurat, 1916

Family Rhabditidae

Family Rictulariidae Railliet, 1916

Family Spirocercidae Chitwood & Wehr, 1932

Family Strongyloididae Chitwood & McIntosh, 1934

Family Trichostrongylidae Leiper, 1912

Genus Trichostrongylus Looss, 1905

Class Enoplea Inglis, 1983

Genus Calodium Dujardin, 1845

Genus Capillaria

Genus Liniscus Dujardin, 1845

Family Trichinellidae Ward, 1907

Family Trichosomoididae Yorke & Maplestone, 1926

Family Trichuridae Railliet, 1915

Phylum Platyhelminthes Minot, 1876

Genus Ctenotaenia Railliet, 1893

Genus Eurotaenia Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Genus Microticola Haukisalmi, Hardman, Hoberg & Henttonen, 2014

Genus Paranoplocephala Lühe, 1910

Family Catenotaeniidae Spasskii, 1950

Genus Spasskijela Tenora, 1959

Family Dilepididae Railliet & Henry, 1909

Family Hymenolepididae Ariola, 1899

Genus Arostrilepis Mas-Coma & Tenora, 1997

Genus Hymenolepis Weinland, 1858

Genus Kontrimavichusia Makarikov & Binkienė, 2022

Genus Rodentolepis Spasskii, 1954

Family Mesocestoididae Fuhrman, 1907

Family Paruterinidae Fuhrmann, 1907

Family Taeniidae Ludwig, 1886

Genus Hydatigera Lamarck, 1816

Genus Taenia Linnaeus, 1758

Genus Versteria Nakao, Lavikainen, Iwaki, Haukisalmi, Konyaev, Oku, Okamoto & Ito, 2013

Class Trematoda Rudolphi, 1808

Family Diplostomidae Poirier, 1886

Order Plagiorchiida La Rue, 1957

Genus Stichorchis Fischoeder, 1901

Family Dicrocoeliidae Looss, 1899

Genus Lyperosomum Looss, 1899

Family Echinostomatidae Looss, 1899

Family Notocotylidae Lühe, 1909

Genus Quinqueserialis Skvortsov, 1935

Family Plagiorchiidae Lühe, 1901

Family Psilostomidae Looss, 1900

Host-parasite list

Apodemus agrarius (Pallas, 1771)

Apodemus flavicollis (Melchior, 1834)

Apodemus sylvaticus (Linnaeus, 1758)