The freshwaters of Iraq harbour a high diversity of endemic and phylogenetically unique species. One of the most diversified fish groups in this region is cyprinoids, and although their distribution is relatively well known, their monogenean parasites have only rarely been investigated. Herein, we applied an integrative approach, combining morphology with molecular data, to assess the diversity and phylogeny of cyprinoid-associated monogenean parasites. A total of 33 monogenean species were collected and identified from 13 endemic cyprinoid species. The highest species diversity was recorded for Dactylogyrus (Dactylogyridae, 16 species) and Gyrodactylus (Gyrodactylidae, 12 species). Four species of Dactylogyrus and 12 species of Gyrodactylus were identified as new to science and described. Two other genera, Dogielius (Dactylogyridae) and Paradiplozoon (Diplozoidae), were represented only by 4 and 1 species, respectively. Phylogenetic analyses of the Dactylogyrus and Gyrodactylus species revealed that the local congeners do not form a monophyletic group and are phylogenetically closely related to species from other regions (i.e. Europe, North Africa and Eastern Asia). These findings support the assumption that the Middle East served as an important historical crossroads for the interchange of fauna between these 3 geographic regions.

Department of Botany and Zoology Faculty of Science Masaryk University Brno Czech Republic

Department of Zoology Faculty of Sciences Comenius University in Bratislava Bratislava Slovakia

Zobrazit více v PubMed

Abdullah YS and Abdullah SMA (2013) Monogenean infections on fishes from Darbandikhan Lake in Kurdistan region, Iraq. Basrah Journal of Agricultural Sciences 26, 117–131.

Abdullah SMA and Mhaisen FT (2005) The first record of three species of Dogielius (Monogenea) from three cyprinid fishes from the Greater Zab River, north of Iraq. Ibn Al-Haitham Journal for Pure and Applied Sciences 18, 7–12.

Abdullah SA, Abdulah AHJ and Onda YW (2022) Fish diversity of the Euphrates River between Medina and Qurna cities, southern Iraq. International Journal of Aquatic Biology 10, 429–437.

Acosta AA, Truter M, Malherbe W and Smit NJ (2022) Morphological description and molecular characterisation of Dactylogyrus matlopong sp. n. (Monogenea: Dactylogyridae) from the South African endemic Labeobarbus aeneus (Cyprinidae: Torinae). Folia Parasitologica 69, 021. PubMed

Al-Awadi HMH (2003) Parasitic faunae (Protozoa and Monogenea) of six species of fish from Euphrates River near Kufa district (Najaf Al-Ashraf province), Iraq. Babylon University Journal for Pure and Applied Sciences 8, 529–532.

Al-Ayash YY, Gustinelli A, Al-Nasiri FS and Caffara M (2021) Three new species of Neoechinorhynchus (Acanthocephala: Neoechinorhynchidae) from fresh-water fish collected from Tigris River, Iraq. Systematic Parasitology 98, 667–677. PubMed

Al-Faisal AJ (2020) Updating checklist of freshwater fishes of Iraq. Mesopotamia Environmental Journal 5, 1–7.

Al-Hassan LAJ, Hussain NA and Saoud K (1989) A preliminary, annotated check-list of the fishes of Shatt Al-Arab River, Basrah, Iraq. Polskie Archiwum Hydrobiologii 36, 283–288.

Al-Jawda JM and Ali MH (2020) Monogeneans and Trematodes of some Fishes from Lake Hamrin in Diyala Province, Iraq. Basrah Journal of Agricultural Sciences 33, 218–228.

Al-Rubaie AL, Hussain HT and Abdul-Ameer KN (2007) The external parasites of the common carp (Cyprinus carpio) in Technical Institute of Al-Mussayab fish farm. Journal of University of Babylon, Science 14, 46–50.

Al-Sa'adi BAHE (2007) The parasitic fauna of fishes of Euphrates River (MSc. M. Tech. thesis). Al-Musaib Technical College, Al-Musaib, Iraq. (In Arabic).

Al-Samman A, Molnár K and Székely C (2006) Infection of cultured and freshwater fishes with monogeneans in Syria. Bulletin of European Association of Fish Pathologists 26, 170–173.

Al-Shaikh SM, Mhaisen FT, Al-Khateeb GH, Balasem AN and Mansoor NT (1995) Collection of some fish parasites from the lower reaches of Diyala river, mid Iraq. Journal of Environmental Science and Health. Part A: Environmental Science and Engineering and Toxicology 30, 1707–1715.

Ann D and Zhang J (1988) Monogenea of south China freshwater fishes. V. Four new species of monogenetic trematodes from fishes of Hanjian River. Annual Bulletin of the Society of Parasitology, Guangdong Province 10, 121–123, (In Chinese).

Asmar KR, Balasem AN, Mhaisen FT, Al-Khateeb GH and Al-Jawda JM (1999) Survey of the parasites of some fish species from Al-Qadisiya Dam Lake, Iraq. Ibn AL-Haitham Journal for Pure and Applied Sciences 12, 52–61.

Aydoğdu A, Selver M and Çirak VY (2008) Comparison of helminth species and their prevalence in rudd (Scardinius erythrophthalmus L. 1758) in Gölbaşı Dam Lake and Kocadere stream in Bursa province of Turkey. Turkish Journal of Veterinary & Animal Sciences 32, 10.

Bachellerie J-P and Qu L-H (1993) Ribosomal RNA probes for detection and identification of species. Methods in Molecular Biology 21, 249–263. PubMed

Bahanak D-N-D, Mbondo JA, Bassock Bayiha ED, Pariselle A, Nack J, Bilong Bilong CF and Agnèse J-F (2022) Description of a new species from Clarias maclareni and phylogenetical analysis of Quadriacanthus (Monogenea, Dactylogyridae) species transfers between clariid and non-clariid fish hosts in Cameroon. Parasite 29, 37. PubMed PMC

Bakke TA, Harris PD, Jansen PA and Hansen LP (1992) Host specificity and dispersal strategy in gyrodactylid monogeneans, with particular reference to Gyrodactylus salaris Malmberg (Platyhelminthes, Monogenea). Diseases of Aquatic Organisms 13, 63–74.

Bakke TA, Harris PD and Cable J (2002) Host specificity dynamics: observations on gyrodactylid monogeneans. International Journal for Parasitology 32, 281–308. PubMed

Bakke TA, Cable J and Harris PD (2007) The biology of gyrodactylid monogeneans: the ‘Russian-doll killers’. Advances in Parasitology 64, 161–376. PubMed

Balasem AN, Mhaisen FT, Adday TK, Al-Jawda JM and Asmar KR (2003) A second survey of parasitic infections in freshwater fishes from Al-Qadisiya Dam Lake, Euphrates River, Iraq. Marina Mesopotamica 18, 123–140, (In Arabic).

Barzegar M, Ebrahimzadeh Mousavi H, Rahmati-Holasoo H, Taheri Mirghaed A and Bozorgnia A (2018) Identification of Gyrodactylus (Monogenea, Gyrodactylidae) species in some of Southwest of Caspian Sea Basin fishes. Iranian Journal of Veterinary Research 12, 35–44.

Bashȇ SKR and Abdullah SMA (2010) Parasitic fauna of spiny eel Mastacembelus mastacembelus from Greater Zab River in Iraq. Iranian Journal of Veterinary Research, Shiraz University 11, 18–27.

Benovics M, Desdevises Y, Vukić J, Šanda R and Šimková A (2018) The phylogenetic relationships and species richness of host-specific Dactylogyrus parasites shaped by the biogeography of Balkan cyprinids. Scientific Reports 8, 13006. PubMed PMC

Benovics M, Desdevises Y, Šanda R, Vukić J and Šimková A (2020a) Cophylogenetic relationships between Dactylogyrus (Monogenea) ectoparasites and endemic cyprinoids of the north-eastern European peri-Mediterranean region. Journal of Zoological Systematics and Evolutionary Research 58, 1–21.

Benovics M, Desdevises Y, Šanda R, Vukić J, Scheifler M, Doadrio I, Sousa- Santos C and Šimková A (2020b) High diversity of fish ectoparasitic monogeneans (Dactylogyrus) in the Iberian Peninsula: a case of adaptive radiation? Parasitology 147, 418–430. PubMed PMC

Benovics M, Nejat F, Abdoli A and Šimková A (2021a) Molecular and morphological phylogeny of host-specific Dactylogyrus parasites (Monogenea) sheds new light on the puzzling Middle Eastern origin of European and African lineages. Parasites & Vectors 14, 372. PubMed PMC

Benovics M, Francová K, Volta P, Dlapka V and Šimková A (2021b) Helminth communities of endemic cyprinoids of the Apennine Peninsula, with remarks on ectoparasitic monogeneans, and a description of four new Dactylogyrus Diesing, 1850 species. Parasitology 148, 1003–1018. PubMed PMC

Benovics M, Vukić J, Šanda R, Nejat F, Charmpila EA, Buj I, Shumka S, Porcelloti S, Tarkan SA, Aksu S, Emiroğlu O and Šimková A (2023) Monogeneans and chubs: ancient host-parasite system under the looking glass. Molecular Phylogenetics and Evolution 179, 107667. PubMed

Bi̇reci̇kli̇gi̇l S, Yücel ŞY and Çiçek E (2016) A taxonomic evaluation of Alburnus sellal Heckel, 1843 and Alburnus adanensis Battalgazi, 1944 based on morphological characters and mitochondrial DNA sequences. Pakistan Journal of Zoology 48, 465–473.

Blasco-Costa I, Míguez-Lozano R, Sarabeev V and Balbuena JA (2012) Molecular phylogeny of species of Ligophorus (Monogenea: Dactylogyridae) and their affinities within the Dactylogyridae. Parasitology International 61, 619–627. PubMed

Boeger WA and Kritsky DC (1993) Phylogeny and a revised classification of the Monogenoidea Bychowsky, 1973 (Platyhelminthes). Systematic Parasitology 26, 1–32.

Bogutskaya NG (1997) Contribution to the knowledge of leuciscine fishes of Asia minor. Part 2. An annotated check-list of leuciscine fishes (Leuciscinae, Cyprinidae) of Turkey with descriptions of a new species and two new subspecies. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 94, 161–186.

Buchmann K and Lindenstrøm T (2002) Interactions between monogenean parasites and their fish hosts. International Journal for Parasitology 32, 309–319. PubMed

Bueno-Silva M, Boeger WA and Pie MR (2011) Choice matters: incipient speciation in Gyrodactylus corydori (Monogenoidea: Gyrodactylidae). International Journal for Parasitology 41, 657–667. PubMed

Bush AO, Lafferty KD, Lotzs JM and Shostakll AW (1997) Parasitology meets ecology on its own terms: Margolis, et al. revised. Journal of Parasitology 83, 575–583. PubMed

Casal-Lopéz M and Doadrio I (2018) The Messinian imprint on the evolution of freshwater fishes of the genus Luciobarbus Heckel, 1843 (Teleostei: Cyprinidae) in the western Mediterranean. Journal of Biogeography 45, 1593–1603.

Coad BW (2010) Freshwater Fishes of Iraq. Sofia-Moscow, RU: Pensoft Publishers, 294 pp.

Cribb TH, Chisholm LA and Bray RA (2002) Diversity in the Monogenea and Digenea: does lifestyle matter? International Journal for Parasitology 32, 321–328. PubMed

Dağli M and Erdemli AÜ (2009) An investigation on the fish fauna of Balıksuyu stream (Kilis, Turkey). International Journal of Engineering Science 3, 1–2.

Dedić N, Vetešník L and Šimková A (2023) Monogeneans in intergeneric hybrids of leuciscid fish: is parasite infection driven by hybrid heterosis, genetic incompatibilities, or host-parasite coevolutionary interactions? Frontiers in Zoology 20, 5. PubMed PMC

Doadrio I (1990) Phylogenetic relationships and classification of western Palearctic species of the genus Barbus (Osteichthyes, Cyprinidae). Aquatic Living Resources 3, 265–282.

Dos Santos QM and Avenant-Oldewage A (2020) Review on the molecular study of the Diplozoidae: analyses of currently available genetic data, what it tells us, and where to go from here. Parasites and Vectors 13, 539. PubMed PMC

Dos Santos QM, Maina JN and Avenant-Oldewage A (2019) Gyrodactylus magadiensis n. sp. (Monogenea, Gyrodactylidae) parasitizing the gills of Alcolapia grahami (Perciformes, Cichlidae), a fish inhabiting the extreme environment of Lake Magadi, Kenya. Parasite 26, 76. PubMed PMC

Dupont F and Lambert A (1986) Study of the parasitic communities of Monogenea Dactylogyridae from Cyprinidae in Lake Mikri Prespa (Northern Greece). Description of three new species from endemic Barbus: Barbus cyclolepis prespensis Karaman. 1924. Annales de Parasitologie Humaine Comparée 6, 597–616. PubMed

Durand JD, Tsigenopoulos CS, Ünlü E and Berrebi P (2002) Phylogeny and biogeography of the family Cyprinidae in the Middle East inferred from cytochrome b DNA – evolutionary significance of this region. Molecular Phylogenetics and Evolution 22, 91–100. PubMed

Dzika E (2002) The parasites of bream Abramis brama (L.) from lake Kortowskie. Archives of Polish Fisheries 10, 85–96.

El Gharbi S, Renaud F and Lambert A (1992) Dactylogyrids (Platyhelminthes: Monogenea) of Barbus spp. (Teleostei: Cyprinidae) from Iberian Peninsula. Research and Reviews in Parasitology 52, 103–116.

El Gharbi S, Birgi E and Lambert A (1994) Monogenean Dactylogyridae parasites of Cyprinidae of the genus Barbus in North Africa. Systematic Parasitology 27, 45–70.

Ergens R (1970) The parasite fauna of fishes from Montenegro. I. Polyonchoinea (Monogenoidea) of some fishes of the Lakes Skadar and Veliko Crno. Pol'Oprivreda i Shumarstvo 16, 1–44.

Ergens R (1991) Variability of the hard parts of opisthaptor of Gyrodactylus leucisci Žitňan, 1964 (Monogenea, Gyrodactylidae). Folia Parasitologica 38, 23–28. PubMed

Ergens R (1992) Redescription of Gyrodactylus truttae Glaser, 1974 (Monogenea, Gyrodactylidae) from Salmo trutta m. fario L. (Clupeiformes, Salmonidae). Folia Parasitologica 39, 201–206. PubMed

Erk'akan F and Özdemir F (2011) Revision of the fish fauna of the Seyhan and Ceyhan River basins in Turkey. Research Journal of Biological Sciences 6, 1–8.

Froese R and Pauly D (2023) FishBase. World Wide Web electronic publication. Available at http://www.fishbase.org (accessed 12 February 2023).

Gibson DI, Timofeeva TA and Gerasev PI (1996) A catalogue of the nominal species of the monogenean genus Dactylogyrus Diesing, 1850 and their host genera. Systematic Parasitology 35, 3–48.

Gussev AV, Jalali B and Molnár K (1993) New and known species of Dactylogyrus Diesing, 1850 (Monogenea, Dactylogyridae) from Iranian freshwater cyprinid fishes. Systematic Parasitology 25, 221–228.

Hahn C, Weiss SJ, Stojanovski S and Bachmann L (2015) Co-speciation of the ectoparasite Gyrodactylus teuchis (Monogenea, Platyhelminthes) and its salmonid hosts. PLoS ONE 10, e0127340. PubMed PMC

Hansen H, Alvestad AH, MacKenzie K, Darrud M, Earlsbakk E, Hemmingsen W and Arneberg P (2020) Gyrodactylus triglopsi n. sp. (Monogenea: Gyrodactylidae) from the gills of Triglops nybelini Jensen, 1944 (Teleostei: Cottidae) in the Barents Sea. Acta Parasitologica 65, 796–803. PubMed PMC

Harris PD, Shinn AP, Cable J and Bakke TA (2004) Nominal species of the genus Gyrodactylus von Nordmann 1832 (Monogenea: Gyrodactylidae), with a list of principal host species. Systematic Parasitology 59, 1–27. PubMed

Hashim DS, Abdullah SMA and Hassan HF (2015) Investigation of parasitic helminthes in freshwater fishes in higher Zab River in Aski kalak, Erbil, Iraq. Kirkuk University Journal-Scientific Studies 10, 309–329.

Hassouna N, Michot B and Bachellerie JP (1984) The complete nucleotide sequence of mouse 28S rRNA gene. Implications for the process of size increase of the large subunit rRNA in higher eukaryotes. Nucleic Acids Research 12, 3563–3583. PubMed PMC

Herzog P (1969) Untersuchungen über die Parasiten der Süßwasserfische des Irak. Archiv für Fischereiwissenschaft 20, 132–147.

Hoffman GL (1999) Parasites of North American Freshwater Fishes, 2nd Edn. Ithaca, NY: Cornell University Press, 539 pp.

Hussain HT (2008) Study on the external ectoparasites of mosquito fish (Gambusia affinis) in Al-Hilla River, Babylon province. Journal of University of Babylon, Pure and Applied Science 15, 245–248.

Huyse T and Volckaert FAM (2002) Identification of a host-associated species complex using molecular and morphometric analyses, with the description of Gyrodactylus rugiensoides n. sp. (Gyrodactylidae, Monogenea). International Journal for Parasitology 32, 907–919. PubMed

Huyse T and Volckaert FAM (2005) Comparing host and parasite phylogenies: Gyrodactylus flatworms jumping from Goby to Goby. Systematic Biology 54, 710–718. PubMed

Huyse T, Audenaert V and Volckaert FAM (2003) Speciation and host-parasite relationships in the parasite genus Gyrodactylus (Monogenea, Platyhelminthes) infecting gobies of the genus Pomatoschistus (Gobiidae, Teleostei). International Journal for Parasitology 33, 1679–1689. PubMed

International Commission on Zoological Nomenclature (2012) Amendment of articles 8, 9, 10, 21 and 78 of the international code of zoological nomenclature to expand and refine methods of publication. Bulletin of Zoological Nomenclature 69, 161–169. PubMed PMC

Jalali B and Molnár K (1990) Occurrence of monogeneans on freshwater fishes of Iran: Dactylogyridae from fish of natural waters and description of Dogielius mokhayeri sp.n. Acta Veterinaria Hungarica 38, 239–242. PubMed

Jawad LA (2006) Fishing gear and methods of the Lower Mesopotamian Plain with reference to fisheries management. Marina Mesopotamica 1, 1–39.

Jin X, Li W, Cheng Y, Li M, Wu S, Zou H and Wang G (2022) Description of Gyrodactylus banmae n. sp. (Monogenea, Gyrodactylidae) parasitic on zebrafish, Danio rerio. Parasitology International 87, 102531. PubMed

Katoh K, Misawa K, Kuma K and Miyata T (2002) MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30, 3059–3066. PubMed PMC

Khalaf KT (1961) The Marine and Freshwater Fishes of Iraq. Baghdad, IQ: Al-Rabitta Press, 164 pp.

Kmentová N, Gelnar M, Mendlová M, Van Steenberge M, Koblmüller S and Vanhove MPM (2016) Reduced host-specificity in a parasite infecting non-littoral Lake Tanganyika cihclids evidenced by intraspecific morphological and genetic diversity. Scientific Reports 6, 39605. PubMed PMC

Kottelat M and Freyhof J (2007) Handbook of European Freshwater Fishes. Berlin, Germany: Kottelat, Cornol and Freyhof, 646 pp.

Koyee QMK and Abdullah SMA (2019) Phylogenetic and secondary RNA structure analysis of monogenean gill ectoparasites (Dactylogyrus spp.) parasitizing certain freshwater fishes. Polish Journal of Veterinary Sciences 22, 667–675. PubMed

Koyun M (2011) Occurrence of monogeneans on some cyprinid fishes from Murat River in Turkey. African Journal of Biotechnology 10, 18285–18293.

Koyun M, Ulupinar M and Gül A (2015) Seasonal distribution of metazoan parasites of Kura barbell (Barbus lacerta) in Eastern Anatolia, Turkey. Pakistan Journal of Zoology 47, 1253–1261.

Koyun M, Korkut N and Gül A (2019) Occurrence of endo and ectoparasites on Capoeta trutta (Heckel, 1843) and Acanthobrama marmid Heckel, 1843 (Cypriniformes: Cyprininae) inhabiting in Göynük Stream Eastern Anatolia. Biharean Biologist 13, 94–100.

Koyuncu CE, Yilmaz H, Secer FS, Kaya D and Genc E (2021) Fish parasites of Tigris and Euphrates River systems. In Jawad LA (ed), Tigris and Euphrates Rivers: Their Environment From Headwaters to Mouth. Cham, CH: Springer, pp. 1389–1418.

Krasnovyd V, Vetešník L, Gettová L, Civáňová K and Šimková A (2017) Patterns of parasite distribution in the hybrids of non-congeneric cyprinid fish species: is asymmetry in parasite infection the result of limited coadaptation? International Journal for Parasitology 47, 471–483. PubMed

Krasnovyd V, Vetešník L and Šimková A (2020) Distribution of host-specific parasites in hybrids of phylogenetically related fish: the effects of genotype frequency and maternal ancestry? Parasites & Vectors 13, 402. PubMed PMC

Kritsky DC, Ali AH and Khamees NR (2013) Gyrodactylus aff. mugili Zhukov, 1970 (Monogenoidea: Gyrodactylidae) from the gills of mullets (Mugiliformes: Mugilidae) collected from the inland waters of southern Iraq, with an evaluation of previous records of Gyrodactylus spp. on mullets in Iraq. Folia Parasitologica 60, 441–447. PubMed

Kuchta R, Řehulková E, Francová K, Scholz T, Morand S and Šimková A (2020) Diversity of monogeneans and tapeworms in cypriniform fishes across two continents. International Journal for Parasitology 50, 771–786. PubMed

Kuru M (2004) Türkiye İçsu Balıklarının Son Sistematik Durumu. GÜ, Gazi Eğitim Fakültesi Dergisi 24, 1–21.

Kvach Y, Ondračková M, Seifertová M and Hulak B (2019) Gyrodactylus ginestrae n. sp. (Monogenea: Gyrodactylidae), a parasite of the big-scale sand smelt, Atherina boyeri Risso, 1810 (Actinopterygii: Atherinidae) from the Black Sea. Parasitology Research 118, 3315–3325. PubMed

Lang S (1981) Monogenetic fauna of freshwater fishes of Hainan Island. Acta Zoologica Sinica 27, 75–86, (In Chinese).

Loot G, Reyjol Y, Poulet N, Šimková A, Blanchet S and Lek S (2007) Effects of small weirs on fish parasite communities. Parasitology Research 101, 1265–1276. PubMed

Lymbery AJ, Morine M, Kanani HG, Beatty SJ and Morgan DL (2014) Co-invaders: the effects of alien parasites on native hosts. International Journal for Parasitology: Parasites and Wildlife 3, 171–177. PubMed PMC

Malmberg G (1957) Om forekomsten av Gyrodactylus pa svenska fiskar. Skrifter Utgivna av Sodra Sveriges Fiskeriforening. Arsskift 1956, 19–76.

Malmberg G (1970) The excretory systems and the marginal hooks as a basis for the systematics of Gyrodactylus (Trematoda, Monogenea). Arkiv för Zoology 23, 1–235.

Matějusová I, Gelnar M, McBeath AJA, Collins CM and Cunningham CO (2001a) Molecular markers for gyrodactylids (Gyrodactylidae: Monogenea) from five fish families (Teleostei). International Journal for Parasitology 31, 738–745. PubMed

Matějusová I, Koubková B, D'Amelio S and Cunningham CO (2001b) Genetic characterization of six species of diplozoids (Monogenea; Diplozoidae). Parasitology 123, 465–474. PubMed

Mendlová M, Desdevides Y, Civáňová K, Pariselle A and Šimková A (2012) Monogeneans of West African cichlid fish: evolution and cophylogenetic interactions. PLoS ONE 7, e37268. PubMed PMC

Mendoza-Palmero CA, Blasco-Costa I and Scholz T (2015) Molecular phylogeny of Neotropical monogeneans (Platyhelminthes: Monogenea) from catfishes (Siluriformes). Parasites & Vectors 8, 164. PubMed PMC

Mhaisen FT (1995) Parasites of fishes of Basrah province marshy area, Iraq. Journal of Environmental Science Health A 30, 41–49.

Mhaisen FT and Abdul-Ameer KN (2013) Checklists of Gyrodactylus species (Monogenea) from fishes of Iraq. Basrah Journal of Agricultural Sciences 26, 8–25.

Mhaisen FT and Abdullah SMA (2017) Parasites of fishes of Kurdistan region, Iraq: checklists. Biological and Applied Environmental Research 1, 131–218.

Mhaisen FT and Al-Nasiri FS (2012) Literature review on the parasites of fishes of Salah Al-Deen Province, Iraq. Tikrit Journal for Agricultural Sciences 12, 209–219.

Mhaisen FT and Al-Rubaie AL (2016) Checklists of parasites of farm fishes of Babylon province, Iraq. Journal of Parasitology Research 2016, 7170534. PubMed PMC

Mhaisen FT, Balasem AN, Al-Khateeb GH and Asmar KR (2003) Recording of five monogenetic trematodes for the first time from fishes of Iraq. Bulletin of the Iraq Natural History Museum 10, 31–38.

Mhaisen FT, Al-Rubae AL and Al-Sa'adi BA (2015) Monogenean parasites of fishes from the Euphrates River at Al-Musaib City, Mid Iraq. American Journal of Biomedical and Life Sciences 3, 50–57.

Mhaisen FT, Abdul-Ameer KN and Hamdan ZK (2018) Checklists of parasites of fishes of Salah Al-Din Province, Iraq. Biological and Applied Environmental Research 2, 180–218.

Mhaisen FT, Al-Mayali HMH and Al-Abodi HRJ (2019) Checklist of parasites of fishes of Al-Diwaniyah province, Iraq. Bulletin of the Iraq Natural History Museum 15, 293–318.

Mizelle JD (1936) New species of trematodes from the gills of Illinois fishes. American Midland Naturalist 17, 785–806.

Mohammad MK (2016) The parasitic fauna of the exotic fish Tilapia zillii in the Middle and South of Iraq. International Journal of Current Microbiology and Applied Science 5, 93–96.

Molnár K (2012) Fifty years of observations about changes of Dactylogyrus infection of European common carp (Cyprinus carpio carpio L.) in Hungary. Magyar Allatorvosok Lapja 134, 111–118, (In Hungarian).

Molnár K and Jalali B (1992) Further monogeneans from Iranian freshwater fishes. Acta Veterinaria Hungarica 40, 55–61. PubMed

Muhammad IM, Dhahir SF, Bilal SJ and Abdullah SMA (2013) Parasitic fauna of some freshwater fishes from Greater Zab River, Kurdistan region, Iraq. Science Journal of University of Zakho 1, 620–627.

Musilová N, Řehulková E and Gelnar M (2009) Dactylogyrids (Platyhelminthes: Monogenea) from the gills of the African carp, Labeo coubie Rüppell (Cyprinidae), from Senegal, with descriptions of three new species of Dactylogyrus and the redescription of Dactylogyrus cyclocirrus Paperna, 1973. Zootaxa 2241, 47–68.

Myers N, Mittermeier RA, Mittermeier CG, de Fonseca GAB and Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403, 853–858. PubMed

Nasraddin MO (2013) Some ecological aspects of monogenean infections on some fishes from Lesser Zab River near Koysinjaq city, Kurdistan region, Iraq (MSc thesis). University of Salahaddin, Erbil, Iraq.

Nitta M (2023) Capsalid monogeneans of fishes from the Seto Inland Sea, Japan: Description of Benedenia kobudai n. sp. parasitic on Semicossyphus reticulatus (Perciformes: Labridae). Parasitology International 92, 102677. PubMed

Öktener A (2021) The parasites of fishes of the Euphrates and Tigris Rivers: Iraq and Turkey. In Jawad LA (ed), Tigris and Euphrates Rivers: Their Environment From Headwaters to Mouth. Cham, CH: Springer, pp. 1419–1444.

Pakosta T, Vetešník L and Šimková A (2018) A long temporal study of parasitism in asexual-sexual populations of Carassius gibelio: does the parasite infection support coevolutionary Red Queen dynamics? BioMed Research International 2018, 6983740. PubMed PMC

Paperna I (1961) Monogenetic trematodes of the Cyprinidae and Claridae of the Lake of Galilee. Bamidgeh, Bulletin of Fish Culture in Israel 13, 14–29.

Paperna I (1964) Parasitic helminths of inland-water fishes in Israel. Israel Journal of Zoology 13, 1–20.

Pazooki J, Jalali Jafari B and Ghobadian M (2006) Monogenean species from freshwater fishes of Zanjan province, Iran. Iranian Journal of Fisheries Sciences 6, 103–112.

Perea S, Böhme M, Zupančič P, Freyhof J, Šanda R, Özuluğ M, Abdoli A and Doadrio I (2010) Phylogenetic relationships and biogeographical patterns in CircumMediterranean subfamily Leuciscinae (Teleostei, Cyprinidae) inferred from both mitochondrial and nuclear data. BMC Evolutionary Biology 10, 265. PubMed PMC

Pleijel F, Jondelius U, Norlinder E, Nygren A, Oxelman B, Schander C, Sunberg P and Thollesson M (2008) Phylogenies without roots? A plea for the use of vouchers in molecular phylogenetic studies. Molecular Phylogenetics and Evolution 48, 369–371. PubMed

Přikrylová I, Vanhove MPM, Janssens SB, Billeter PA and Huyse T (2013) Tiny worms from a mighty continent: high diversity and new phylogenetic lineages of African monogeneans. Molecular Phylogenetics and Evolution 67, 43–52. PubMed

Pugachev ON, Gerasev PI, Gussev AV, Ergens R and Khotenowsky I (2009) Guide to Monogenoidea of Freshwater Fish of Palearctic and Amur Regions. Milan, Italy, Ledizione-Ledi Publishing, 564 pp.

Rahemo ZIF (1980) Diplozoon kasimii new species from a freshwater teleost fish, Cyprinion macrostomum Heckel. Bulletin of the Biological Research Center 12, 109–114.

Rahemo ZIF (1982) Two new species of Ergasilus (Copepoda: Cyclopoida) from the gills of two Iraqi freshwater fishes. Bulletin of Basrah Natural History Museum 5, 39–59.

Rahmouni I, Řehulková E, Pariselle A, Rkhami OB and Šimková A (2017) Four new species of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae) parasitising the gills of northern Moroccan Luciobarbus Heckel (Cyprinidae): morphological and molecular characterisation. Systematic Parasitology 94, 575–591. PubMed

Rahmouni C, Vanhove MPM, Koblmüller S and Šimková A (2022) Molecular phylogeny and speciation patterns in host-specific monogeneans (Cichlidogyrus, Dactylogyridae) parasitizing cichlid fishes (Cichliformes, Cichlidae) in Lake Tanganyika. International Journal for Parasitology 52, 359–375. PubMed

Rahmouni C, Seifertová M and Simková A (2023) Revealing the hidden diversity of Gyrodactylus communities (Monogenea: Gyrodactylidae) from Nearctic Catostomidae and Leuciscidae fish hosts (Teleostei: Cypriniformes), with description of ten new species. Parasite 30, 40. PubMed PMC

Rambaut A, Drummon AJ, Xie D, Baele G and Suchard MA (2018) Posterior summarization in Bayesian phylogenetic using Tracer 1.7. Systematic Biology 67, 901–904. PubMed PMC

Rasheed AR and Al-Saadi AAJJ (2018) The first record of four species of Gyrodactylus (Monogenea) from gills of four freshwater fish species in Iraq. Biological and Applied Environmental Research 2, 1–11.

Řehulková E, Seifertová M, Přikrylová I and Francová K (2018) Monogenea. In Scholz T, Vanhove MPM, Smit N, Jayasundera Z and Gelnar M (eds). A Guide to the Parasites of African Freshwater Fishes. Brussels, BE: AbcTaxa, pp. 185–243.

Řehulková E, Benovics M and Šimková A (2020) Uncovering the diversity of monogeneans (Platyhelminthes) on endemic cypriniform fishes of the Balkan Peninsula: new species of Dactylogyrus and comments on their phylogeny and host-parasite associations in a biogeographic context. Parasite 27, 66. PubMed PMC

Řehulková E, Rahmouni I, Pariselle A and Šimková A (2021) Integrating morphological and molecular approaches for characterizing four species of Dactylogyrus (Monogenea: Dactylogyridae) from Moroccan cyprinids, with comments on their host specificity and phylogenetic relationships. PeerJ 9, e10867. PubMed PMC

Rohde K (1979) A critical evaluation of intrinsic and extrinsic factors responsible for niche restriction in parasites. The American Naturalist 114, 648–671.

Rohlenová K, Morand S, Hyršl P, Tolarová S, Flajšhans M and Šimková A (2011) Are fish immune systems really affected by parasites? An immunoecological study of common carp (Cyprinus carpio). Parasites & Vectors 4, 120. PubMed PMC

Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA and Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across large model space. Systematic Biology 61, 539–542. PubMed PMC

Sala OE, Chapin FS, Armesto JJ, Berlow R, Bloomfield J, Dirzo R, Huber-Sanwald E, Huenneke LF, Jackson RB, Kinzig A, Leemans R, Lodge D, Mooney HA, Oesterheld M, Poff NL, Sykes MT, Walker BH, Walker M and Wall DH (2000) Global biodiversity scenarios for the year 2100. Science 287, 1770–1774. PubMed

Sasal P, Trouvé S, Müller-Graf C and Morand S (1999) Specificity and host predictability: a comparative analysis among monogenean parasites of fish. Journal of Animal Ecology 68, 437–444.

Schönhuth S, Vukić J, Šanda R, Yang L and Mayden RL (2018) Phylogenetic relationships and classification of the Holarctic family Leuciscidae (Cypriniformes: Cyprinoidei). Molecular Phylogenetics and Evolution 127, 781–799. PubMed

Seifertová M, Vyskočilová M, Morand S and Šimková A (2008) Metazoan parasites of freshwater cyprinid fish (Leuciscus cephalus): testing biogeography hypotheses of species diversity. Parasitology 135, 1417–1435. PubMed

Shigoley M, Rahmouni I, Louizi H, Pariselle A and Vanhove MPM (2023) First study on Gyrodactylus (Monogenea: Gyrodactylidae) in Morocco, with description of a new species from Luciobarbus pallaryi and Luciobarbus ksibi (Actinopterygii: Cyprinidae). Animals 13, 1624. PubMed PMC

Šimková A and Morand S (2008) Co-evolutionary patterns in congeneric monogeneans: a review of Dactylogyrus species and their cyprinid hosts. Journal of Fish Biology 73, 2210–2227.

Šimková A, Plaisance L, Matějusová I, Morand S and Verneau O (2003) Phylogenetic relationships of the Dactylogyridae Bychowsky, 1933 (Monogenea: Dactylogyridea): the need for the systematic revision of the Ancyrocephalinae Bychowsky, 1937. Systematic Parasitology 54, 1–11. PubMed

Šimková A, Morand S, Jobet E, Gelnar M and Verneau O (2004) Molecular phylogeny of congeneric monogenean parasites (Dactylogyrus): a case of intrahost speciation. Evolution 58, 1001–1018. PubMed

Šimková A, Verneau O, Gelnar M and Morand S (2006) Specificity and specialization of congeneric monogeneans parasitizing cyprinid fish. Evolution 60, 1023–1037. PubMed

Šimková A, Serbielle C, Pariselle A, Vanhove MPM and Morand S (2013) Speciation in Thaparocleidus (Monogenea: Dactylogyridae) parasitizing Asian pangasiid catfishes. BioMed Research International 2013, 353956.. PubMed PMC

Šimková A, Řehulkova E, Jean Robertin R, Jorissen M, Scholz T, Faltynková A, Mašová S and Vanhove MPM (2018) Transmission of parasites from introduced tilapias: a new threat to endemic Malagasy ichthyofauna. Biological Invasions 21, 803–819.

Šimková A, Řehulková E, Choudhury A and Seifertová M (2022) Host-specific parasites reveal the history and biogeographical contacts of their hosts: the Monogenea of Nearctic cyprinoid fishes. Biology 11, 229. PubMed PMC

Soylu E (2012) Monogenean parasites of white bream (Blicca bjoerkna Linnaeus, 1758) in Lake Sapanca, Turkey. Kafkas Universitesi Veteriner Fakultesi Dergisi 18, A23–A28.

Stamatakis A (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 2688–2690. PubMed

Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analyses and post-analysis of large phylogenies. Bioinformatics 30, 1312–1313. PubMed PMC

Steininger FF and Rögl F (1984) Paleogeography and palinspastic reconstruction of the Neogene of the Mediterranean and paratethys. In Dixon JE and Robertson AH (eds), The Geological Evolution of the Eastern Mediterranean. Oxford, UK: Blackwell, pp. 659–668.

Stiassny MLJ and Getahun A (2007) An overview of labeonin relationships and the phylogenetic placement of the Afro-Asian genus Garra Hamilton, 1922 (Teleostei: Cyprinidae), with the description of five new species of Garra from Ethiopia, and a key to all African species. Zoological Journal of Linnaean Society 150, 41–83.

Stojanovski S, Kulišić Z, Baker R, Hristovski N, Cakić P and Hristovski M (2004) Fauna of monogenean trematods – parasites of some cyprinid fishes from Lake Prespa (Macedonia). Acta Veterinaria (Beograd) 54, 73–82.

Strayer DL and Dudgeon D (2010) Freshwater biodiversity conservation: recent progress and future challenges. Journal of the North American Benthological Society 29, 344–358.

Tan M and Armbruster JW (2018) Phylogenetic classification of extant genera of fishes of the order Cypriniformes (Teleostei: Ostariophysi). Zootaxa 4476, 6–39. PubMed

Teimori A, Esmaeili HR, Sayyadzadeh G, Zarei N and Gholamhosseini A (2015) Molecular systematics and distribution review of the endemic cyprinid species, Persian chub, Acanthobrama persidis (Coad, 1981) in Southern Iran (Teleostei: Cyprinidae). Molecular Biology Research Communications 4, 189–206. PubMed PMC

Tkach VV, Littlewood DTJ, Olson PD, Kinsella JM and Swiderski ZP (2003) Molecular phylogenetic analysis of the Microphalloidea Ward, 1901 (Trematoda: Digenea). Systematic Parasitology 56, 1–15. PubMed

Truter M, Smit NJ, Malherbe W and Přikrylová I (2022) Description of Gyrodactylus paludinosus sp. nov. (Monogenea: Gyrodactylidae) from the straightfin barb, Enteromius paludinosus (Peters, 1852), in South Africa. Acta Parasitologica 67, 446–453. PubMed

Tunç A and Koyun M (2018) Seasonal infection of metazoan parasites on Mosul Bleak (Alburnus mossulensis) inhabiting Murat River and its tributaries in Eastern Anatolia, Turkey. Turkish Journal of Agricultural and Natural Sciences 5, 153–162.

Turan D, Kaya C, Aksu İ and Bektaş Y (2022) Paracapoeta, a new genus of the Cyprinidae from Mesopotamia, Cilicia and Levant (Teleostei, Cypriniformes). Zoosystematics and Evolution 98, 201–212.

Viñuela-Rodríguez N, Šanda R, Zogaris S and Vukić J (2021) Evolutionary history of the Pelasgus minnows (Teleostei: Leuciscidae), an ancient endemic genus from the Balkan Peninsula. Molecular Phylogenetics and Evolution 164, 107274. PubMed

Whittington ID and Kearn GC (2011) Hatching strategies in monogenean (platyhelminth) parasites that facilitate host infection. Integrative and Comparative Biology 51, 91–99. PubMed

Whittington ID, Chisholm LA and Rohde K (1999) The larvae of Monogenea. Advances in Parasitology 44, 139–232.

Whittington ID, Bronwen WC, Hamwood TE and Halliday JA (2000) Host-specificity of monogenean (platyhelminth) parasites: a role for anterior adhesive areas? International Journal for Parasitology 30, 305–320. PubMed

Wilson JR, Saunders RJ and Hutson KS (2019) Parasites of the invasive tilapia Oreochromis mossambicus: evidence for co-introduction. Aquatic Invasions 14, 332–349.

Yang L, Sado T, Hirt MV, Pasco-Viel E, Arunachalam M, Li J, Wang X, Freyhof J, Saitoh K, Simons AM, Miya M, He S and Mayden RL (2015) Phylogeny and polyploidy: resolving the classification of cyprinine fishes (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution 85, 97–116. PubMed

Zhang J and Ji G (1980) Eleven new species of Dactylogyrus (Trematoda: Monogenea) from China. Acta Zootaxonomica Sinica 5, 341–352, (In Chinese).

Ziȩtara MS and Lumme J (2003) The crossroads of molecular, typological and biological species concepts: two new species of Gyrodactylus Nordmann, 1832 (Monogenea: Gyrodactylidae). Systematic Parasitology 55, 39–52. PubMed