ПАРАЗИТОЛОГИЯ, 50, 1, 2016

УДК 576.895.121

A CHECKLIST OF PROTOZOAN AND METAZOAN PARASITES OF THE BARBATULA (, BALITORIDAE) © A. E. Zhokhov,1 A. V. Shershneva,2 M. N. Pugacheva1

11. D. Papanin Institute for Biology of Inland Waters, Russian Academy of Sciences, Borok, Yaroslavl, 152742, Russia E-mail: [email protected] 2 P. G. Demidov Yaroslavl State University Sovetskaya, 14, Yaroslavl, 150000, Russia E-mail: [email protected] Submitted 02.01.2015

The present paper summarizes data on parasites of the stoan loach from all its area, ba- sed on published data. The checklist includes all protozoan and metazoan parasites of the stoan loach from different freshwater habitats (rivers, lakes, ponds, reservoirs). A total of 109 parasite / taxa were recorded on the stoan loach (Kinetoplastidea — 4, Diplo- monadea — 1, Coccidea — 2, Microsporea — 1, Myxosporea — 13, Litostomatea — 1, Phyllopharyngea — 2, Oligohymenophorea — 22, Monogenea — 10, Cestoda — 4, Tre- matoda — 31, Nematoda— 11, Acanthocephala — 5, Crustacea — 1, Bivalvia — 1). Most are digenean trematodes, 18 of them being metacercariae. Key words: checklist, parasites, freshwater , Balitoridae, Barbatula barbatula, area.

СПИСОК ПАРАЗИТОВ (PROTOZOA И METAZOA) УСАТОГО ГОЛЬЦА BARBATULA BARBATULA (CYPRINIFORMES, BALITORIDAE) © A. E. Жохов,1 А. В. Шершнева,2 M. H. Пугачева1

1 Институт биологии внутренних вод им. И. Д. Папанина РАН пос. Борок, Ярославская обл., 152742 E-mail: [email protected] 2 Ярославский государственный университет им. П. Г. Демидова ул. Советская, Ярославль, 150000 E-mail: [email protected] Поступила 02.01.2015

Статья суммирует данные по паразитам усатого гольца по всему ареалу его оби- тания, основанные на опубликованных данных. Список включает всех одноклеточ- ных и многоклеточных паразитов усатого гольца из различных пресных водоемов

40 (рек, озер, прудов, водохранилищ). В общей сложности у усатого гольца зарегистри- ровано 109 видов/таксонов паразитов (Kinetoplastidea — 4, Diplomonadea — 1, Cocci- dea — 2, Microsporea — 1, Myxosporea — 13, Litostomatea — 1, Phyllopharyngea — 2, Oligohymenophorea — 22, Monogenea — 10, Cestoda — 4, Trematoda — 31, Nemato- da — 11, Acanthocephala — 5, Crustacea — 1, Bivalvia — 1). Самой многочисленной группой являются трематоды, 18 видов которых представлены метацеркариями. Ключевые слова: список видов, паразиты, пресноводные рыбы, Balitoridae, Barba- tula barbatula, ареал.

Members of the family Balitoridae are small freshwater fishes found in fast streams and rivers of Eurasia. Over 450 species have been described, with the highest diversity in Southeast Asia, but only 5 occured in Europe (Bogutskaya, Naseka, 2004; Nelson, 2006). The stone loach occurs in both lotic and lentic freshwater. Although it prefers well-oxygenated rivers and streams, it can also be found in lakes and smaller ponds. The stone loach possesses wide natural dis- tribution in Europe, from in the west to the Ural and Emba basins in the east, but it does not occur on the Pyrenean, Apennine, and Balkan peninsulas (Vasil'eva, 1998; Freyhof, 2011). In spite of the common presence of the stone loach in many European count- ries, this species is often listed as threatened, vulnerable or endangered. Infor- mation on parasites of species of no economic value is still scanty. To date, a checklist of the protozoan and metazoan parasites of the stone loach has not yet been published. The aim of the present literature review was to summarize all published data on the parasite fauna of the Barbatula barbatula.

MATERIAL AND METHODS

The checklist (see table) was compiled from the available literature. Parasite species are arranged according to the phylum, class and family, within which the species are presented in the alphabetical order, followed by information on the infection site, the locations where the specimens were collected, and refe- rences. In addition, the checklist includes some parasite species recorded only to the level (undetermined species). Protozoan follows Bauer (1984) and Lom, Dykova (2006); for Trematoda, Gibson et al. (2002) and Jones et al. (2005); Bauer (1985, 1987) for Monogenea, Cestoda, Nematoda, Acantho- cephala, Arthropoda.

RESULTS AND DISCUSSION

A total of 109 parasitic species / taxa were recorded for the stone loach in the area (see table). The protozoan parasite species (46 species / taxa) included re- presentatives of the classes Kinetoplastidea (4), Diplomonadea (1), Coccidea (2), Microsporea (1), Myxosporea (13), Litostomatea (1), Phyllopharyngea (2), and Oligohymenophorea (22). The metazoan parasite species (63 species / taxa) consisted of the platyhelminth classes Monogenea (10), Cestoda (4), Tremato- da (31), and the phyla Nemathelminthes (11), Acanthocephales (5), Ar- thropoda (1), and Mollusca (1). Oligohymenophoreans and trematodes were the most abundant taxonomic groups. Larval helminthes were present in 26 speci-

41 4 2 Parasites of Barbatula barbatula in area including site of infection, drainage (DRA), country and references. § — introduced species

Parasite taxa Site Locality References

Phylum KINETOPLASTIDA Class KINETOPLASTIDEA Family TRYPANOSOMATIDAE Trypanosoma carassii (Mitrophanov, 1883) Bloodstream Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 River Lee (England) Gibson et al., 2005 Family BODONIDAE Ichthyobodo necator (Henneguy, 1884) Gills Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Family CRYPTOBIIDAE Cryptobia branchialis Nie (in: Chen, 1956) Gills River Lososinka (Russia) Yevseeva, 2001 C. varia Leger, 1904 Bloodstream Lake Ladoga (Russia) Rumyantsev et al., 2001 Phylum POLYMASTIGOTA Class DIPLOMONADEA Family HEXAMITIDAE Hexamita sp. Site not given River Lososinka (Russia) Yevseeva, 2001 Phylum SPOROZOA Class COCCIDEA Family EIMERIDAE Eimeria pastuszkoi Jastzebski, 1982 Intestine Locality not given Belova, Krylov, 2006; Duszynski et al., 2000 Eimeria sp. Intestine Lake Ladoga (Russia) Rumyantsev et al., 2001 Phylum MICROSPORA Class MICROSPOREA Family GLUGEIDAE Pleistophora macrospora Cepede, 1906 Abdominal trunk Lake Ladoga (Russia) Rumyantsev et al., 2001 muscles Near Grenoble () Lom, 2002 Rivers flowing into the Black Sea Issi, Voronin, 1984 (Russia) Phylum MYXOZOA Class MYXOSPOREA Family MYXIDIIDAE Myxosporidia gen. sp. Gull bladder Lake Syamozero (Russia) Shul'man, 1962 Mixidium barbatulae Cepede, 1906 Kidney Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Lake Onega (Russia) Rumyantsev etal., 1984 M. pfeifferi Auerbach, 1908 » River Lososinka (Russia) Yevseeva, 2001 Mixidium sp. Site not given Ponds of fish farm (Russia) Kudentsova, 1977 Zchokkella nova Klokacheva, 1914 Gull bladder River Lososinka (Russia) Yevseeva, 2001 Family CHLOROMYXIDAE Chloromyxum truttae Leger, 1906 » » Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Family SPHAEROSPORIDAE Sphaerospora sp. Site not given Lake Ladoga (Russia) Rumyantsev et al., 2001 Mixobilatus fragilicandatus R. Schulman, Site not given River Lososinka (Russia) Yevseeva, 2001 1969 M. legeri (Cepede, 1905) Urinary bladder, Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 renal tubule Lakes Syamozero, Pertozero (Russia) Shul'man et al., 1974; Shul'man, Family MYXOBOLIDAE 1962; Shul'man, Rybak, 1961 Myxobolus nemachili Weisler, 1949 Head connective tis- Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 sue, heart, gills, Ponds of fish farm ( Russia) Kudentsova, 1977 kidney, spleen, vit- River Danube (Ukraine, Slovak Repub- Kulakovskaya, Koval, 1973 reous body lic) Locality not given () Eiras et al., 2005 Thelochanellus furmanni (Auerbach, 1909) Gills, urinary blad- River Lososinka (Russia) Yevseeva, 2001 der Lake Pertozero (Russia) Shul'man et al., 1974 River Danube (Slovak Republic) Kulakovskaya, Koval, 1973 Lake Syamozero (Russia) Shul'man, 1962 Lakes Pertozero (Russia) Shul'man, Rybak, 1961 T. nemachili Diarova, 1967 Vitreous body, mus- Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 culature, gills, kid- Lake Ladoga (Russia) Rumyantsev et al., 2001 ney, heart, cereb- River Lososinka (Russia) Yevseeva, 2001 rum T. pyriformis (Thelochan, 1892) Site not given River Mezen' (Russia) Dorovskich, 1997a Phylum CILIOPHORA Class LITOSTOMATEA Family AMPHILEPTIDAE Hemiophrys branchiarum (Wenrich, 1924) Gills River Lososinka (Russia) Yevseeva, 2001 4 3 Kahl, 1931 4 Continuation of table

Parasite taxa Site Locality References

Class PHYLLOPHARYNGEA Family CHILODONELLIDAE Chilodonella piscicola (Zacharias, 1894) Jankowski, Body surface River Lososinka (Russia) Yevseeva, 2001 1980 Ponds of fish farm (Russia) Kudentsova, 1977 Family TRICHOPHRYIDAE Capriniana piscium (Bütschli, 1889) Jankowski, Gills Ponds of fish farm (Russia) Kudentsova, 1977 1973 Class OLIGOHYMENOPHOREA Family EPISTYLIDIDAE Apiosoma amoeba (Grenfell, 1887) Body surface Ponds of fish farm (Russia) Kudentsova, 1977 A. baueri (Kaschkowski, 1965) Body surface, fins River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 Lake Onega (Russia) Rumyantsev et al., 1984 A. campanulatum (Timofeev, 1962) Gills, skin Rivers of Upper Volga DRA Shershneva, Zhokhov, 2013 (Russia) A. phoxini Lom, 1966 Body surface River Lososinka (Russia) Yevseeva, 2001 Lake Onega (Russia) Rumyantsev et al., 1984 A. piscicolum Blanchard, 1885 Gills Lake Ladoga (Russia) Rumyantsev et al., 2001 A. poteriformis (Timofeev, 1962) Body surface Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 Apiosoma sp. Gills Lake Ladoga (Russia) Rumyantsev et al., 2001 Epistylis Iwoffi Faure-Fremiet, 1943 » Rivers of Upper Volga DRA Shershneva, Zhokhov, 2013 (Russia) River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 Family TRICHODINIDAE Trichodina cobitis Lom, 1960 Nasal fosse, skin Rivers of Upper Volga DRA Shershneva, Zhokhov, 2013 (Russia) T. domerguei domerguei (Wallendrenn, 1897) Gills, fins Lake Pestovo (Russia) Pozdnyakova, 1957 River Mezen' (Russia) Dorovskich, 1997a Rivers of Danube DRA (Slovak Re- Kulakovskaya, Koval, 1973 public) Т. fultoni Davis, 1947 Skin Danube DRA (Czech and Slovak Re- Kulakovskaya, Koval, 1973; Lom, public) 1961 Т. nemachili Lom, 1960 Gills, body surface Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 Danube DRA (Czech, Slovak Repub- Kulakovskaya, Koval, 1973 lic) Т. nigra Lom, 1960 Site not given River Osoblahy (Czech Republic) Ergens, 1965 Т. schulmani Chan, 1961 Gills Lakes of Karelia (Russia) Chan, 1961; Shul'man et al., 1974 Trichodina sp. Lake Ladoga (Russia) Rumyantsev et al., 2001 Skin, gills Lake Pertozero ( Russia) Shul'man, Rybak, 1961 Paratrichodina incisa (Lom, 1959) Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Gills Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 P. phoxini Lom, 1963 Site not given River Lososinka (Russia) Yevseeva, 2001 Lake Onega (Russia) Rumyantsev et al., 1984 Tripatriella copiosa (Lom, 1959) Gills Ponds of fish farm (Russia) Kudentsova, 1977 T. lata Lom, 1963 Body surface Ponds of fish farm (Russia) Kudentsova, 1977 Trichodinella epizootica (Raabe, 1950) » » River Tisa, Danube DRA (Ukraine, Ergens et al., 1975 Slovak Republic) Family TETRAHYMENIDAE Tetrachimena pyriformis (Ehrenberg, 1830) Gills Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Family ICHTHYOPHTHIRIIDAE Ichthyophthirius miltifillis Fouquet, 1876 Gills, fins Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Ponds of fish farm (Russia) Kudentsova, 1977 River Osoblaha (Czech Republic) Ergens, 1965 Phylum PLATHELMINTHES Class MONOGENOIDEA Family ANCYROCEPHALIDAE Ancyrocephalus cruciatus (Wedl, 1857) Gills River Severskiy Donets, Don DRA Shevchenko, 1956 4 5 (Russia) 4 6 Continuation of table

Parasite taxa Site Locality References

Family GYRODACTYLIDAE Gyrodactylus barbatuli Achmerov, 1952 Body surface Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 River Mezen' (Russia) Dorovskich, 1997b River Tisa, Danube DRA (Ukraine, Ergens et al., 1975; Kulakovskaya, Slovak Republic) Koval, 1973 Rivers of Danube DRA (Czech Repub- Moravec, 2001 lic) Rivers of Visla DRA () Prost, 1981 G. cobitis Bychowsky, 1933 Fins Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 G. jiroveci Ergens et Bychowsky, 1967 Fins, skin Rivers of Danube DRA (Czech Repub- Prikrylovä et al., 2008 lic) Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Rivers Northern Dvina, Mezen', Dorovskich, 1997b Pechora (Russia) River Tisa, Danube DRA (Ukraine, Ergens et al., 1975; Kulakovskaya, Slovak Republic) Koval, 1973 Rivers of Danube DRA (Czech, Slovak Moravec, 2001 Republic) River Skawa, Visla DRA (Poland) Prost, 1981 Rivers of Danube DRA (Czech Repub- Prikrylovä et al., 2005 lic) G. laevis Malmberg, 1957 Gills Ponds of fish farm (Russia) Kudentsova, 1977 G. menshikowi Gvosdev, 1950 Fins, skin, gills Ponds of fish farm (Russia) Kudentsova, 1977 River of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Rivers of Danube DRA, Elbe DRA Moravec, 2001 (Czech, Slovak Republic) Rivers of Visla DRA (Poland) Prost, 1981 Rivers of Danube DRA (Czech Repub- Prikrylovä et al., 2005 lic) G. nemachili Bychowsky, 1936 Gills, fins Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 River Osoblahy (Czech Republic) Ergens, 1965 Rivers of Danube DRA (Czech and Moravec, 2001; Prikrylovä et al., Slovak Republic) 2005 G. pavlovskyi Ergens et Bychowsky, 1967 Fins Rivers of Visla DRA (Poland) Prost, 1981 Rivers of Danube DRA (Czech and Moravec, 2001; Prikrylovä et al., Slovak Republic) 2005 G. pseudonemachili Ergens et Bychowsky, Fins, skin Rivers of Danube DRA (Czech Repub- Prikrylovä et al., 2005, 2008 1967* lic) G. sedelniekovi Gvosdev, 1950 Gills, fins, skin Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Ponds of fish farm (Russia) Kudentsova, 1977 Rivers Mezen', Pechora (Russia) Dorovskich, 1997b Rivers of Danube DRA (Czech, Slovak Moravec, 2001; Prikrylovä et al., Republic) 2005 Danube DRA () Kulakovskaya, Koval, 1973 Rivers of Visla DRA (Poland) Prost, 1981 Danube DRA (Bulgaria) Kakacheva-Avramova, 1970 Class CESTODA Family TRIAENOPHORIDAE Triaenophorus nodulosus (Pallas, 1781) plero- Liver River Northern Dvina (Russia) Dorovskich, 2000 cercoid River Avon (England) Rumpus, 1975 Family LIGULIDAE Schistocephcdus nemachili Dubinina, 1959 Abdominal cavity River Mezen' (Russia) Dorovskich, 2000 plerocercoid Ligiila colymbi Zeder, 1803 plerocercoid » » Tayganak reservoir (Crimea) Reshetnikova, 1957 Family PROTEOCEPHALIDAE Intestine Proteocephalus scigittus (Grimm, 1872) Lake Pertozero (Russia) Anikieva, Rumyantsev, 2008 Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Kudentsova, 1977 4 7 Ponds of fish farm (Russia) Lake Pestovo (Russia) Pozdnyakova, 1957 4 8 Continuation of table

Parasite taxa Site Locality References

Lake Pertozero (Russia) Shul'man et al., 1974 Rivers of Danube DRA, Oder DRA, El- Moravec, 2001; Scholz et al., 2003 be DRA (Czech, Slovak Republic) Lake Konchozero (Russia) Shul'man, Rybak, 1961 River Danube (Bulgaria) Kakacheva-Avramova, 1969 Class TREMATODA Family DIPLOSTOMIDAE Diplostomum chromatophorum Brown, 1931 Lens Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 metacercariae Lake Ladoga (Russia) Lebedeva, 2005 D. rutili Razmashkin, 1969 metacercariae Lens Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 River Mezen' (Russia) Dorovskich, 1997c D. helveticum (Dubois, 1929) metacercariae Lens Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 River Mezen' (Russia) Dorovskich, 1997c D. mergi Dubois, 1932 metacercariae Lens Ponds of fish farm (Russia) Kudentsova, 1977 D. phoxini (Faust, 1918) Arvy et Buttner, 1954 River Pechora (Russia) Dorovskich, 1997c metacercariae Cerebrum D. paraspathaceum Shigin, 1965 metacercariae Ponds of fish farm (Russia) Kudentsova, 1977 D. petromyzifluviatilis Diesing, 1860 metacer- Lens River Mezen' (Russia) Dorovskich, 1997c cariae D. pussilum (Dubois, 1928) metacercariae Cerebrum Lake Ladoga (Russia) Rumyantsev et al., 2001 D. spathaceum (Rudolphi, 1819) metacercariae Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lens River Mezen' (Russia) Dorovskich, 1997c Lens, vitreous Rivers, reservoirs of England Wootten, 1974 body Rivers of Danube DRA; river Oder Moravec, 2001 (Czech, Slovak Republic) Lakes of Karelia (Russia) Shul'man, 1962; Shul'man, Rybak, 1961 Ponds of fish farm (Russia) Kudentsova, 1977; Shigin, 1980 D. volvens Nordmann, 1832 metacercariae Lens River Mezen' (Russia) Dorovskich, 1997c Lake Ladoga (Russia) Lebedeva, 2005 Tylodelphys clavata Nordmann, 1832 Vitreous body River Lososinka (Russia) Yevseeva, 2001 metacercariae Lakes Pertozero, Syamozero (Russia) Shul'man et al., 1974 Rivers of Danube DRA (Czech, Slovak Moravec, 2001 Republic) Lake Syamozero (Russia) Shul'man, 1962 Posthodiplostomum cuticola (Nordmann, Skin, musculature Rivers of Danube DRA (Czech Repub- Moravec, 2001 1832) metacercariae lic) Family CYATHOCOTYLIDAE Holostephanus cobitidis Opravilova, 1968 Abdominal cavity Rivers of Upper Volga DRA (Russia) Zhokhov, 2000 metacercariae Paracoenogonimus ovatus Katsurada, 1914 Fins, musculature, Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 metacercariae gills, eyes, go- nads Family STRIGEIDAE Ichthyocotylurus pileatus (Rudolphi, 1802) Swim bladder Lake Ladoga (Russia) Rumyantsev et al., 2001; Lebedeva, metacercariae wall, muscula- 2005 ture, hardroe, Lake Pertozero (Russia) Shul'man et al., 1974 spleen Rivers of Danube DRA (Czech, Slovak Moravec, 2001 Republic) River Wieprz (Poland) Adamczyk, 1980 Lakes Pertozero, Konchozero (Russia) Shul'man, Rybak, 1961 I. variegatus (Creplin, 1825) metacercariae Abdominal cavity River Roding (England) Kennedy, 1974 Apatemon gracilis (Rudolphi, 1819) Szidat, » » Rivers Elbe DRA, Danube DRA, Oder Moravec, 2001, 2003; Ergens et al., 1928 metacercariae DRA (Ukraine, Slovak, Czech Re- 1975 public) Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 River Lososinka (Russia) Yevseeva, 2001 Rivers Mezen', Pechora (Russia) Dorovskich, 1997c Rivers of (Czech, Slovak Republic) Moravec, 2001 Family SANGUINICOLIDAE Sangunicola sp. Bloodstream River Severskiy Donets, Don DRA Shevchenko, 1956 (Russia) Family ALLOCREADIIDAE Crepidostomum metoecus (Braun, 1900) Intestine River Kamenice, Elbe DRA (Czech Re- Moravec, 2003 4 9 public) 5 0 Continuation of table

Parasite taxa Site Locality References

С. farionis (Müller, 1784) Intestine Lake Onega (Russia) Rumyantsev et al., 1984 Rivers of Danube DRA (Czech Repub- Moravec, 2001 lic) С. oschmarini Zhokhov et Pugacheva, 1998 Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013; Zhokhov, Pugacheva, 1998 A. isoporum (Looss, 1894) Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 River Pechora (Russia) Dorovskich, 1997c Rivers of Danube DRA, Elbe DRA Moravec, 2001 (Czech, Slovak Republic) Lake Konchozero (Russia) Shul'man, Rybak, 1961 A. transversale (Rudolphi, 1802) Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Konchozero (Russia) Shul'man et al., 1974 Allocreadiiim sp. Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Family OPECOELIDAE Nicolla gallica (Dollfus, 1941) Intestine River Avon (England) Rumpus, 1975 N. skrjabini (Iwanitzky, 1928) Intestine River Severskiy Donets, Don DRA Shevchenko, 1956 (Russia) Sphaerostomum bramae (Müller, 1776) Intestine Lake Konchozero (Russia) Shul'man, Rybak, 1961 Sphaerostomum globiporum (Rudolphi, 1802) Intestine Lake Pertozero (Russia) Shul'man et al., 1974 Family HETEROPHYIDAE Metagonimus yokogawai Katsurada, 1912 meta- Fins River Danube (Slovak Republic) Moravec, 2001 cercariae Family GORGODERIDAE Phyllodistomum folium (Olfer, 1926) Ureters, urinary Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 bladder River Northern Dvina (Russia) Dorovskich, 1997c Lake Ladoga (Russia) Rumyantsev et al., 2001; Lebedeva, 2005 River Lososinka (Russia) Yevseeva, 2001 River Morava (Czech Republic) Moravec, 2001; Petkeviciüte et al., 2015 Family ZOOGONIDAE Pseudochetosoma salmonicola Dollfus, 1951 Gull bladder River Nishava, Danube DRA (Bulga- Kakacheva-Avramova, 1969 ria) Phylum NEMATHELMINTHES Class NEMATODA Family CAPILLARIIDAE Pseudocapillaria tomentosa Dujardin, 1843 Intestine Rivers of Danube DRA (Czech Repub- Moravec, 2001 lic) Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 Schulmanella petruschewskii (Schulman, 1948) Liver River Tisa, Danube DRA (Ukraine, Ergens et al., 1975 Slovak Republic) Family ANISAKIDAE Raphidascaris acus (Bloch, 1799) larvae Liver, abdominal Rivers of Elbe DRA (Czech Republic) Moravec, 2003, 2004 cavity, intestine, Rivers of Danube DRA, Oder DRA, Koubkovä et al., 2004; Moravec, 2001 stomach Lake Mächa (Czech Republic) Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Lake Pestovo (Russia) Pozdnyakova, 1957 Lake Pertozero (Russia) Shul'man et al., 1974 Rivers Mezen', Northern Dvina Dorovskich, 1999 (Russia) River Wieprz (Poland) Adamczyk, 1980 Lakes of Karelia (Russia) Shul'man, Rybak, 1961 Family CAMALLANIDAE Camallanus truncatus (Rudolphi, 1814) Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Family PHILOMETRIDAE Philometroides cyprini (Ishii, 1931) § Body cavity Ponds of fish farm (Russia) Kudentsova, 1977 Family RHABDOCHONIDAE Rhabdochona denudata (Dujardin, 1845) Intestine Rivers of Oder DRA (Czech Republic) Moravec, 2001 Lake Pestovo (Russia) Pozdnyakova, 1957 Rivers Mezen', Pechora (Russia) Dorovskich, 1999 R. ergensi Moravec, 1968 Intestine Rivers of Elbe DRA, Danube DRA, Moravec, 2001, 2003 5 1 Oder DRA (Czech, Slovak Republic) 5 2 Continuation of table

Parasite taxa Site Locality References

Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 River Lososinka (Russia) Yevseeva, 2001 Rivers Mezen', Northern Dvina Dorovskich, 1999 (Russia) Rhabdochona hellichi (Sramek, 1901) Intestine Lakes of Karelia (Russia) Shul'man, Rybak, 1961; Shul'man et al., 1974; Shul'man, 1962 R. phoxini Moravec, 1968 larvae Intestine Locality not given (Czech Republic) Moravec, 2001 Family CYSTIDICOLIDAE Salmonema ephemeridarum (Linstow, 1872) Stomach Rivers of Elbe DRA, Danube DRA Moravec, 2001, 2003 larvae (Czech Republic) River Lososinka (Russia) Yevseeva, 2001 Family ACUARIIDAE Streptocara crassicaudata (Creplin, 1829) Stomach, intesti- Rivers of Danube DRA (Czech Repub- Moravec, 2001 larvae ne wall, liver lic) Locality not given (Poland) Czaplinski, 1963 Phylum ACANTHOCEPHALES Class ACANTHOCEPHALA Family ARHYTHMACANTHIDAE Acanthocephalus anguillae (Muller, 1780) Intestine Rivers of Oder DRA, Danube DRA Moravec, 2001 (Czech, Slovak Republic) A. lucii (Müller, 1776) Intestine Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Rivers of Danube DRA; river Oder Moravec, 2001 (Czech, Slovak Republic) Lakes of Karelia (Russia) Shul'man, Rybak, 1961 Paracanthocephalus sp. Intestine River Lososinka (Russia) Yevseeva, 2001 Neoechinorhynchus rutili (Müller, 1780) Intestine Lake Pestovo (Russia) Pozdnyakova, 1957 Rivers Mezen', Northern Dvina Dorovskich, 1999 (Russia) Rivers of Danube DRA, Oder DRA Moravec, 2001 (Czech, Slovak Republic) River Chusovaya, Kama DRA (Russia) Kostarev, 2003 Family ILLIOSENTIDAE Pomphorhynchus laevis (Müller, 1776) Intestine Rivers of England Kennedy, 1999; Wanstall et al., 1990; Rumpus, 1975 River Tisa, Danube DRA (Ukraine, Ergens et al., 1975; Moravec, 2001 Czech, Slovak Republic) Phylum ARTHROPODA Class COPEPODA Family ERGASILIDAE Ergasilus sieboldi von Nordmann, 1832 Gills Lake Pertozero (Russia) Shul'man et al., 1974 Rivers of Danube DRA; river Oder Moravec, 2001 (Czech, Slovak Republic) Lake Pertozero (Russia) Shul'man, Rybak, 1961 Phylum MOLLUSCA Class BIVALVIA Family UNIONIDA Glochidium sp. Fins Rivers of Upper Volga DRA (Russia) Shershneva, Zhokhov, 2013 Lake Ladoga (Russia) Rumyantsev et al., 2001 Rivers of Danube DRA (Slovak Repub- Moravec, 2001 lic) River Prut (Ukraine) Kulakovskaya, Koval, 1973 River Dnestr (Ukraine) Kulakovskaya, 1959 5 3 es / taxa; most (18) were metacercariae of trematodes. The introduced nematode Philometroides cyprini (Ishii, 1931) was recorded for the stone loach only in Russia (Kudentsova, 1977). In general, the number of metazoan parasite species was significantly higher than the number of protozoan parasite species. Oligohymenophorea was the most abundant group among protozoan parasites. Myxosporidia were also abun- dant. The other groups of protozoan were represented by a small number of spe- cies. Trematodes had the highest number of species among metazoan parasites of the stone loach. Monogeneans and nematodes were also abundant groups. Relatively few cestode and acanthocephalans species were found. Copepods and molluscs were the least abundant taxonomic groups of parasites in the stone loach. The genus Gyrodactyliis is the most numerously represented monogene- an taxon in the stone loach.

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