Checklist of The Monogenea (Platyhelminthes) Parasitic in Tunisian Aquatic Vertebrates.

153 species of monogeneans have been recorded in Tunisian aquatic vertebrates (89 hosts). A list of these species with hosts is presented. A comparison of the Monogenea diversity off the coast of Tunisia with other regions of the Mediterranean Sea and the world is provided. The number of parasites depends on the number of hosts examined and their diversity in the region. This list shows that Monopisthocotylea is the richest group. In addition, new records have been reported during a survey of the diversity of monogeneans fish in the southern coast of Tunisia (Gulf of Gabes) including: Benedenia monticellii (Parona and Perugia, 1895) Johnston, 1929, Lamellodiscus bidens Euzet, 1984, Lamellodiscus confusus Amine, Euzet & Kechemir-Issad, 2007, Lamellodiscus ergensi Euzet & Oliver, 1966, Lamellodiscus hilii Euzet, 1984, Lamellodiscus impervius Euzet, 1984, Lamellodiscus knoepffleri Oliver, 1969, Lamellodiscus theroni Amine, Euzet & Kechemir-Issad, 2007, Ligophorus acuminatus Euzet & Suriano, 1977, Ligophorus angustus Euzet & Suriano, 1977, Ligophorus heteronchus Euzet & Suriano, 1977, Ligophorus macrocolpos Euzet & Suriano, 1977, Ligophorus minimus Euzet & Suriano, 1977, Capsala maccallumi Price, 1939 and Pseudanthocotyloides heterocotyle (Van Beneden, 1871) Euzet & Prost, 1969.

Introduction

Tunisian coastal waters are located in the south-central Mediterranean Sea. Tunisia has a 1670- km coastline including 7 coastal lagoons, covering a total surface of 105200 ha and an exceptional continental shelf of 88 000 km2 (Cherif ). It also has a large hydrographic network in the north. The surface water accommodates a relatively high faunal diversity, especially in terms of aquatic vertebrates that can host a high species richness of parasites. Among them, monogeneans (Plathyhelmintes) which are commonly found on fishes can also parasitise freshwater turtles and amphibians. These parasites are ideal for diversity investiga

tions because they are diverse both in terms of morphology and numbers. Their members are generally host specific, and their phylogeny is well resolved (Poulin, 2002). In addition, the study of monogeneans is also fundamental to aquaculture because they are known as agents responsible for many epidemics (Lia ). In Tunisia, these flatworms were first studied by Ktari (1969) who described the species of Atrispinum salpae (Parona & Perugia, 1890) Maillard & Noisy, 1979 from Sarpa salpa (Linnaeus, 1758). After that, more than 50 studies have been published about monogenenans including description of new species and new hosts as well as locality records. Despite these parasitological surveys, there were no attempts to quantify the species richness of Monogenea of Tunisian aquatic vertebrates.

In this paper we present all the available species records of monogeneans in Tunisian waters and we incorporate new data derived from our own work on species recorded from marine fish of the Gulf of Gabes (southern coast of Tunisia) in order to make the first checklist of monogeneans as parasites of Tunisian aquatic vertebrates.

Materials and Methods

The list of Monogenea from Tunisian aquatic vertebrates was prepared based on published data and doctoral and masters theses. It is organized in both a parasite–host table (Table 1 and a host– parasite one (Table 2).The families and species of monogeneans are presented in alphabetical order followed by microhabitat, localities and references (Table 1). The nomenclatures of all taxa are presented according to WoRMS (2021). However, authors of new combinations for some very old taxa of Monogenea are lacking.

Table 1

List of Monogenea parasites of Tunisian aquatic organisms. Records are listed by monogenoidean families and ordered per: parasite species (alphabetically), host species (alphabetically), reference (chronologically). NR, New record. Areas: 1, North; 2, East ; 3, South; 4. hydrographic network in the north.

Monogenoides species	Host species	Microhabitat	Reference
Family Amphibdellatidae Carus, 1885
Amphibdella paronaperugiae Llewellyn, 1960	Torpedo torpedo (Linnaeus, 1758) (1, 3)	gills	Neifar, 1995, 2001
Amphibdelloides benhassinae Tazerouti, Euzet and Neifar, 2006	Torpedo torpedo (Linnaeus, 1758) (1, 3)	gills	Neifar, 1995, 2001 ; Tazerouti et al., 2006
Amphibdelloides kechemirae Tazerouti, Euzet and Neifar, 2006	Torpedo marmorata Risso, 1810 (1, 3)	gills	Neifar, 1995, 2001 ; Tazerouti et al., 2006
Amphibdelloides vallei Llewellyn, 1960	Torpedo marmorata Risso, 1810 (1, 3)	gills	Neifar, 2001
Family Ancyrocephalidae Bychowsky, 1937
Glyphidohaptor plectocirra (Paperna, 1972) Kritsky, Galli and Yang, 2007	Siganus luridus (Rüppell, 1829), S. rivulatus Forsskål and Niebuhr, 1775(3)	gills	Boussellaa, 2020
Haliotrema balisticus (Hargis, 1955) Yamaguti, 1963	Balistes capriscus Gmelin, 1789 (3)	gills	Kacem and Neifar, 2015; present work
Ligophorus acuminatus Euzet and Suriano, 1977	Chelon saliens (Risso, 1810) (3)	gills	present work (NR)
Ligophorus angustus Euzet and Suriano, 1977	Chelon labrosus (Risso, 1827) (3)	gills	present work (NR)
Ligophorus chabaudi Euzet and Suriano, 1977	Mugil cephalus Linnaeus, 1758 (1)	gills	Neifar, 1995
Ligophorus heteronchus Euzet and Suriano, 1977	Chelon saliens (Risso, 1810) (3)	gills	present work (NR)
Ligophorus macrocolpos Euzet and Suriano, 1977	Chelon saliens (Risso, 1810) (3)	gills	present work (NR)
Ligophorus minimus Euzet and Suriano, 1977	Chelon saliens (Risso, 1810) (3)	gills	present work (NR)
Ligophorus szidati Euzet and Suriano, 1977	Chelon auratus (Risso, 1810) (1) (3)	gills	Neifar, 1995 present work
Ligophorus vanbenedenii (Parana and Perugia, 1890) Euzet and Suriano, 1977	Chelon auratus (Risso, 1810) (1) (3)	gills	Neifar, 1995 present work
Pseudempleurosoma sp.	Sphyraena chrysotaenia Klunzinger, 1884 (2, 3)	pharynx and oesophagus	Boussellaa, 2018, 2020
Family Axinidae Monticelli, 1903
Axine belones Abildgaard, 1794	Belone belone (Linnaeus, 1760) (1) (1,2)	gills	Ktari, 1971; Neifar, 1995 Châari, 2013; Châari et al., 2016
Axine sp.	Belone svetovidovi Collette and Parin, 1970 (1, 2)	gills	Châari, 2013; Châari et al., 2016
Axinoides sp.	Tylosurus acus imperialis (Rafinesque, 1810) (1, 2)	gills	Châari, 2013; Châari et al., 2016
Nudaciraxine imperium Châari, Derbel and Neifar, 2010	Tylosurus acus imperialis (Rafinesque, 1810) (1, 2)	gills	Châari, 2013; Châari et al., 2010, 2016
Family Calceostomatidae Parana and Perugia, 1890
Dicrumenia bychowskyi Mamaev, 1969	Pomadasys incisus (Bowdich, 1825) (1)	gills	Euzet and Ktari, 1973
Family Capsalidae Baird, 1853
Benedenia monticellii (Parana and Perugia, 1895) Johnston, 1929	Mugil cephalus Linnaeus, 1758 (3)	skin	present work (NR)
Capsala maccallumi Price, 1939	Euthynnus alletteratus (Rafinesque, 1810) (3)	gills	present work (NR)
Neoentobdella apiocolpos (Euzet and Maillard, 1967) Kearn and Whittington, 2005	Taeniura grabata Geoffroy Saint-Hilaire, 1817 (1, 3)	skin (ventral surface)	Neifar, 2001
Neoentobdella diadema (Monticelli, 1902) Kearn and Whittington, 2005	Dasyatis marmorata (Steindachner, 1892), D. pastinaca (Linnaeus, 1758), D. tortonesei Capapé, 1975 (1,3)	skin (ventral surface)	Neifar, 2001
Trimusculotrema micracantha (Euzet and Maillard, 1967) Whittington and Barton, 1990	Dasyatis marmorata (Steindachner, 1892), D. pastinaca (Linnaeus, 1758), D. tortonesei Capapé, 1975 (1,3)	skin (ventral surface)	Neifar, 2001
Trochopus heteracanthus Massa, 1903	Chelidonichthys lucerna (Linnaeus, 1758) (1)	gills	Neifar, 1995
Trochopus pini (Van Beneden and Hesse, 1863) Massa, 1903	Chelidonichthys lucerna (Linnaeus, 1758) (1)	gills	Ktari, 1971; Neifar, 1995
Family Chauhaneidae Euzet and Trilles, 1960
Cotyloatlantica mediterranea (Euzet and Trilles, 1960) Bravo-Hollis, 1984	Sphyraena sphyraena (Linnaeus, 1758) (1) (2,3)	gills	Ktari, 1971; Neifar, 1995 Boussellaa, 2020; Boussellaa et al. 2018
Family Chimaericolidae Brinkmann, 1942
Chimaericola leptogaster (Leuckart, 1830) Brinkmann, 1942	host unknown (not cited) (1)	gills	Ktari, 1971
Family Dactylogyridae Bychowsky, 1933
Dactylogyrus heteromorphus El Garbi, Birgi and Lambert, 1994	Luciobarbus callensis (Valenciennes, 1842) (4)	gills	El Garbi et al., 1994; Khalil and Polling, 1997; Řehulková et al., 2018
Dactylogyrus tunisiensis El Garbi, Birgi and Lambert, 1994	Luciobarbus callensis (Valenciennes, 1842) (4)	gills	El Garbi et al., 1994; Khalil and Polling, 1997; Řehulková et al., 2018
Family Diclidophoridae Cerfontaine, 1895
Choricotyle chrysophryi Van Beneden and Hesse, 1863	Boops boops (Linnaeus, 1758) (1)	gills	Ktari, 1977; Neifar, 1995
Diclidophora sp.	Trisopterus capelanus (Lacepède, 1800) (1)	gills	Ktari, 1971
Family Dionchidae Bychowsky, 1959
Dionchus agassizi Goto, 1899	Echeneis naucrates Linnaeus, 1758 (1)	gills	Ktari, 1971, 1975, 1977
Dionchus remorae (MacCallum, 1916) Price, 1938	Echeneis naucrates Linnaeus, 1758 (1)	gills	Ktari, 1975, 1977
Family Diplectanidae Monticelli, 1903
Diplectanum aequans (Wagener, 1857)	Dicentrarchus labrax (Linnaeus, 1758) (3) (1)	gills	Euzet, 1984, present work Neifar, 1995
Diplectanum grassei Oliver, 1974	Umbrina cirrosa Linnaeus, 1758 (1)	gills	Neifar, 1995
Diplectanum laubieri Lambert and Maillard, 1974	Dicentrarchus puctatus (Bloch, 1792) (1)	gills	Neifar, 1995
Diplectanum melvillei Oliver and Paperna, 1984	Umbrina cirrosa Linnaeus, 1758 (1)	gills	Neifar, 1995
Diplectanum simile Bychowsky, 1957	Sciaena umbra Linnaeus, 1758 (1)	gills	Neifar, 1995
Echinoplectanum echinophallus (Euzet and Oliver, 1965) Justine and Euzet, 2006	Mycteroperca marginata (Lowe, 1834) (3)	gills	Chaabane et al., 2017
Lamellodiscus abbreviatus Sanfilippo, 1978	Diplodus sargus (Linnaeus, 1758) (1)	gills	Neifar, 1995
Lamellodiscus bidens Euzet, 1984	Diplodus puntazzo (walbaum, 1792) (3)	gills	present work (NR)
Lamellodiscus contusus Amine, Euzet and Kechemir-lssad, 2007	Sarpa salpa (Linnaeus, 1758) (3)	gills	Present work (NR)
Lamellodiscus crampus Neifar, 2008	Dentex maroccanus Valenciennes, 1830 (3)	gills	Neifar, 2008
Lamellodiscus echeneis (Wagener, 1857) Diesing, 1858	Sparus aurata Linnaeus, 1758 (3) (1)	gills	Euzet, 1984, present work Neifar, 1995
Lamellodiscus elegans Bychowsky, 1957	Diplodus annularis (Linnaeus, 1758), D. sargus (Linnaeus, 1758), Oblada melanura (Linnaeus, 1758) (1)	gills	Neifar, 1995
Lamellodiscus ergensi Euzet and Oliver, 1966	Diplodus vulgaris (Geoffroy Saint-Hilaire) (3)	gills	present work (NR)
Lamellodiscus erythrini Euzet and Oliver, 1966	Pagellus erythrinus (Linnaeus, 1758) (3)	gills	Euzet, 1984; present work
Lamellodiscus euzeti Diamanka, Boudaya, Toguebaye and Pariselle,	Dentex gibbosus (Rafinesque, 1810) (3)	gills	Diamanka et al., 2011
2011
Lamellodiscus flagellatus Boudaya, Neifar and Euzet, 2009	Lithognathus mormyrus (Linnaeus, 1758) (3)	gills	Boudaya et al., 2009
Lamellodiscus fraternus Bychowsky, 1957	Diplodus annularis (Linnaeus, 1758) (3)	gills	Euzet, 1984; present work
	(1)		Neifar, 1995
Lamellodiscus furcosus Euzet and Oliver, 1966	Diplodus sargus (Linnaeus, 1758), D. vulgaris (Geoffroy	gills	Neifar, 1995
	Saint-Hilaire, 1817) (1)
Lamellodiscus gracilis Euzet and Oliver, 1966	Oblada melanura (Linnaeus, 1758) (1)	gills	Neifar, 1995
Lamellodiscus gussevi Sanfilippo, 1978	Diplodus sargus (Linnaeus, 1758) (1)	gills	Neifar, 1995
Lamellodiscus hilii Euzet, 1984	Diplodus puntazzo (walbaum) (3)	gills	present work (NR)
Lamellodiscus ignoratus Palombi, 1943	Diplodus sargus (Linnaeus), Sarpa salpa (Linnaeus) (1)	gills	Neifar, 1995
	Diplodus puntazzo (walbaum, 1792), D. vulgaris (Geoffroy
	Saint-Hilaire, 1817), (3)		present work
Lamellodiscus impervius Euzet, 1984	Diplodus puntazzo (walbaum, 1792) (3)	gills	present work (NR)
Lamellodiscus knoepffleri Oliver, 1969	Spicara smaris (Linnaeus, 1758) (3)	gills	present work (NR)
Lamellodiscus mirandus Euzet and Oliver, 1966	Diplodus sargus (Linnaeus, 1758) (1)	gills	Neifar, 1995
Lamellodiscus mormyri Euzet and Oliver, 1967	Lithognathus mormyrus (Linnaeus, 1758) (3)	gills	Euzet, 1984; Boudaya et al., 2009
	(1)		Neifar, 1995
Lamellodiscus parisi Oliver, 1969	Sarpa salpa (Linnaeus, 1758) (3)	gills	Euzet, 1984; present work
	(1)		Neifar, 1995
Lamellodiscus rastellus Neifar, Euzet and Oliver, 2004	Pagrus auriga Valenciennes, 1843 (3)	gills	Neifar et al., 2004
Lamellodiscus sarculus Neifar, Euzet and Oliver, 2004	Pagrus coeruleostictus (Valenciennes, 1830) (3)	gills	Neifar et al., 2004
Lamellodiscus sigilatus Neifar, Euzet and Oliver, 2004	Pagrus coeruleostictus (Valenciennes, 1830) (3)	gills	Neifar et al., 2004
Lamellodiscus theroni Amine, Euzet and Kechemir-lssad, 2007	Diplodus puntazzo (walbaum, 1792) (3)	gills	present work (NR)
Lamellodiscus verberis Euzet and Oliver, 1967	Lithognathus mormyrus (Linnaeus, 1758) (3) (1)	gills	Euzet, 1984; present work Neifar, 1995
Protolamellodiscus serranelli (Euzet and Oliver, 1965) Oliver, 1969	Serranus cabrilla (Linnaeus, 1758), S. scriba (Linnaeus,	gills	Euzet, 1984
	1758) (3)		Neifar, 1995
	(1)
Pseudodiplectanum syrticum Derbel, Boudaya and Neifar, 2007	Synapturichthys kleinii (Risso, 1827) (3)	gills	Derbel et al., 2007; present work
Pseudorhabdosynochus beverleyburtonae (Oliver, 1984) Kritsky and	Mycteroperca marginata (Lowe, 1834) (3)	gills	Chaabane et al., 2016 b
Beverley-Burton, 1986
Pseudorhabdosynochus bouaini Neifar and Euzet, 2007	Mycteroperca costae (Steindachner, 1878) (3)	gills	Neifar and Euzet, 2007; Chaabane et al.,
			2017
Pseudorhabdosynochus dolicocolpos Neifar and Euzet, 2007	Mycteroperca costae (Steindachner, 1878) (3)	gills	Neifar and Euzet, 2007; Chaabane et al.,
			2017
Pseudorhabdosynochus enitsuji Neifar and Euzet, 2007	Mycteroperca costae (Steindachner, 1878) (3)	gills	Neifar and Euzet, 2007; Chaabane et al.,
			2017
Pseudorhabdosynochus hayet Chaabane, Neifar, Gey and Justine,	Mycteroperca rubra (Bloch, 1793) (3)	gills	Chaabane et al., 2016 b
2016
Pseudorhabdosynochus regius Chaabane, Neifar and Justine, 2015	Mycteroperca rubra (Bloch, 1793) (3)	gills	Chaabane et al., 2015
Pseudorhabdosynochus riouxi (Oliver, 1986) Santos, Buchmann and Gibson, 2000	Mycteroperca marginata (Lowe, 1834) (3)	gills	Chaabane et al., 2017
Pseudorhabdosynochus sinediscus Neifar and Euzet, 2007	Mycteroperca costae (Steindachner, 1878) (3)	gills	Neifar and Euzet, 2007; Chaabane et al., 2017
Pseudorhabdosynochus sosia Neifar and Euzet, 2007	Mycteroperca costae (Steindachner, 1878) (3)	gills	Neifar and Euzet, 2007; Chaabane et al., 2017
Pseudorhabdosynochus sulamericanus Santos, Buchmann and Gibson, 2000	Hyporthodus haitensis (Ben-Tuvia, 1953) (3)	gills	Chaabane et al., 2016 a
Family Discocotylidae Price, 1936
Anthocotyle merlucii Van Beneden and Hesse, 1863	Merluccius merluccius (Linnaeus, 1758) (1)	gills	Neifar, 1995
Family Fridericianellidae Gupta and Sachdeva, 1990
Calceostomella inermis (Parana and Perugia, 1889) Palombi, 1943	Sciaena umbra Linnaeus, 1758 (1)	gills	Neifar, 1995
	Umbrina cirrosa Linnaeus, 1758 (1)		Euzet and Ktari, 1970 a, 1973; Neifar, 1995
Family Gastrocotylidae Price, 1943
Gastrocotyle trachuri Van Beneden and Hesse, 1863	Trachurus mediterraneus (Steindachner, 1868), T. trachurus (Linnaeus, 1758), (1, 2, 3)	gills	Feki, 2017
Pseudaxine trachuri Parana and Perugia, 1890	Trachurus mediterraneus (Steindachner, 1868) (1, 2, 3) Trachurus trachurus (Linnaeus, 1758) (1) (1,2,3)	gills	Feki, 2017 Ktari, 1971; Neifar, 1995 Feki, 2017
Pseudaxine sp.	Euthynnus alletteratus (Rafinesque, 1810) (1)	gills	Ktari, 1971
Family Gotocotylidae Yamaguti, 1963
Gotocotyla acanthura (Parana and Perugia, 1896) Meserve, 1938	Brama brama (Bonnaterre, 1788), Dentex dentex (Linnaeus, 1758), Oblada melanura (Linnaeus, 1758), Pomatomus saltatrix (Linnaeus, 1758), Trachinotus ovatus (Linnaeus, 1758) (1)	gills	Ktari, 1971
Gotocotyla sp1. of Neifar, 1995	Trachinotus ovatus (Linnaeus, 1758) (1)	gills	Neifar, 1995
Gotocotyla sp 2. of Neifar, 1995	Pomatomus saltatrix (Linnaeus, 1758) (1)	gills	Neifar, 1995
Family Heteraxinidae Unnithan, 1957
Cemocotyle trachuri Dillon and Hargis, 1965	Trachurus mediterraneus (Steindachner, 1868), T. trachurus (Linnaeus, 1758) (1, 2, 3)	gills	Feki, 2017
Heteraxine seriola (Ishii, 1936) Sproston, 1946	Seriola dumerili (Risso, 1810) (1)	gills	Ktari, 1971
Intracotyle hannibali (Euzet and Ktari, 1970) Mamaev, 1977	Pomadasys incisus (Bowdich, 1825) (1)	gills	Euzet and Ktari, 1970 b
Family Hexabothriidae Price, 1942
Dasyonchocotyle spiniphallus Hargis, 1955	Bathytoshia centroura (Mitchill, 1815) (1, 3)	gills	Neifar, 2001
Epicotyle torpedinis (Price, 1942) Euzet and Maillard, 1974	Torpedo marmorata Risso, 1810, T. torpedo (Linnaeus, 1758) (1,3)	skin	Neifar, 2001
Erpocotyle catenulata (Guberlet, 1933) Yamaguti, 1963	Mustelus mustelus (Linnaeus, 1758) (1, 3)	gills	Neifar, 2001
Erpocotyle sp.	Mustelus punctulatus Risso, 1827 (1)	gills	Neifar, 1995
Heteronchocotyle gymnurae Neifar, Euzet and Ben Hassine, 2001	Gymnura altavela (Linnaeus, 1758) (1, 3)	gills	Neifar, 2001; Neifar et al., 2001 b
Hexabothrium appendiculatum (Kuhn, 1829) von Nordmann, 1840	Scyliorhinus stellaris (Linnaeus, 1758) (1)	gills	Neifar, 2001
Hexabothrium canicula (Cerfontaine, 1899) Price, 1942	host unknown (not cited) (1)	gills	Ktari, 1971
Neonchocotyle pastinacae Ktari and Maillard, 1972	Dasyatis pastinaca(Linnaeus, 1758) (1)	gills	Ktari and Maillard, 1972
Rajonchocotyle sp.	Raja clavata Linnaeus, 1758, R. miraletus Linnaeus, 1758, R. radula Delaroche, 1809, Rostroraja alba (Lacepède, 1803) (1,2,3)	gills	Neifar, 2001
Family Hexostomatidae Hexostoma thynni (Delaroche, 1811) Rafinesque, 1815	Thunnus thynnus (Linnaeus, 1758) (1)	gills	Ktari, 1971
Neohexostoma eufhynni(Meserve, 1938) Price, 1961	Euthynnus alletteratus (Rafinesque, 1810) (1) (3)	gills	Ktari, 1971 present work
Family Mazocraeidae
Grubea cochlear Diesing, 1858	Scomber japonicus Houttuyn, 1782 (3)	gills	present work
	Scomber scombrus Linnaeus, 1758 (1) (1,2,3)		Ktari, 1971 Feki, 2017
Kuhnia scombri (Kuhn, 1829) Sproston, 1945	Scomber scombrus Linnaeus, 1758 (1) (1, 2, 3)	gills	Ktari, 1971 Feki, 2017
	Scombercolias Gmelin, 1789 (1, 2, 3)		Feki, 2017
	Scomber japonicus Houttuyn, 1782 (3)		Present work
Mazocraes pilchardi (Van Beneden and Hesse, 1863) Sproston, 1946	Sardinapilchardus (Walbaum, 1792) (1) (3)	gills	Feki, 2017 present work
Mazocraes sp.	Sardinella aurita Valenciennes, 1847 (3) (2)	gills	Derbel, 2004; present work Feki, 2017
Mazocraeoides sardinellae Ktari, 1982	Sardinella maderensis (Lowe, 1838) (1)	gills	Ktari, 1982; Neifar, 1995
Pseudanthocotyloides heterocotyle (Van Beneden, 1871) Euzet and Prost, 1969	Engraulis encrasicolus (Linnaeus, 1758) (3)	gills	present work (NR)
Pseudokuhnia minor (Goto, 1984) Rohde and Watson, 1985	Scomber colias Gmelin, 1789 (1, 2, 3)	gills	Feki, 2017
	Scomber japonicus Houttuyn, 1782 (3)		present work
Family Microcotylidae Taschenberg, 1879
Atrispinum salpae (Parana and Perugia, 1890) Maillard and Noisy, 1979	Sarpa salpa (Linnaeus, 1758) (1)	gills	Ktari, 1969, 1971; Neifar, 1995
Bychowskicotyla mormyri (Lorenz, 1878) Unnithan, 1971	Lithognathus mormyrus (Linnaeus, 1758) (1)	gills	Ktari, 1971; Neifar, 1995
Bivagina alcedinis (Parana and Perugia, 1889) Yamaguti, 1963	Spicara maena (Linnaeus, 1758) (1)	gills	Neifar, 1995
Bradyhaptorus trachini (Parana and Perugia, 1890) Unnithan, 1971	Trachinus araneus Cuvier, 1829 T. draco Linnaeus, 1758, T radiatus Cuvier, 1829 (1)	gills	Azizi, 2020; Azizi et al., 2017
Metamicrocotyla cephalus (Azim, 1939) Hargis, 1954	Mugil cephalus Linnaeus, 1758 (1) (3)	gills	Ktari, 1971 present work
Microcotyle erythrini Van Beneden and Hesse, 1863	Pagellus erythrinus (Linnaeus, 1758) (1) (3)	gills	Ktari, 1971; Neifar, 1995 present work
Microcotyle pomatomi Goto, 1899	Pomatomus saltatrix (Linnaeus, 1758) (1)	gills	Neifar, 1995
Pauciconfibula trachini (Parana and Perugia, 1889) Dilllon and	Trachinus araneus Cuvier, 1829 (1)	gills	Ktari, 1971; Neifar, 1995
Hargis, 1965	Trachinus radiatus Cuvier, 1829 (1)		Azizi, 2020; Azizi et al.,2021
Polylabris tubicirrus (Paperna and Kohn, 1964) Mamaev and Parukhin, 1976	Diplodus annularis (Linnaeus, 1758) (1) (3)	gills	Ktari, 1971 present work
Polylabris mamaevi Ogawa and Egusa, 1980	Siganus luridus Rtippell, 1829, S. rivulatus Forsskål and Niebuhr, 1775(3)	gills	Boussellaa, 2020
Pseudoaspinatrium euzeti (Ktari, 1971) Mamaev, 1986	Dentex gibbosus (Rafinesque, 1810) (1)	gills	Ktari, 1971
Pseudoaspinatrium gallieni (Euzet and Ktari, 1971) Mamaev, 1986	Tylosurus acus imperialis (Rafinesque, 1810) (1) (2,3)	Inner gill cover,	Ktari, 1971; Euzet and Ktari, 1971 Châari, 2013; Châari et al., 2016
		gills
Sparicotyle chrysophrii (Van Beneden and Hesse, 1863) Mamaev, 1984	host unknown (not cited) (1)	gills	Ktari, 1971
Sciaenacotyle pancerii (Sonsino, 1891) Mamaev, 1989	Umbrina cirrosa Linnaeus, 1785 (1)	gills	Ktari, 1970, 1971
Serranicotyle labracis (Van Beneden and Hesse, 1863) Maillard, Euzet and Silan, 1988	host unknown (not cited) (1)	gills	Ktari, 1971
Solostamenides mugilis (Vogt, 1879) Unnithan, 1971	Chelon auratus (Risso, 1810), C. labrosus (Risso, 1827), C. ramada (Risso, 1827), Ch saliens (Risso, 1810), Mugil cephalus Linnaeus, 1758 (1)	gills	Ktari, 1971; Neifar, 1995
Family Monocotylidae Taschenberg, 1879
Calicotyle kroyeri Diesing, 1850	Raja miraletus Linnaeus, 1758, R. radula Delaroche, 1809, Rostroraja alba (Lacepède, 1803) (2, 3)	cloaca	Neifar, 2001
Calicotyle palombi Euzet and Williams, 1960	Mustelus punctulatus Risso, 1827 (1, 2, 3)	cloaca	Neifar, 2001
Calicotyle stossichi Braun, 1899	Mustelus mustelus (Linnaeus, 1758), M. punctulatus Risso, 1827 (1,2,3)	rectal gland	Neifar, 2001
Calicotyle vicina Neifar, Euzet and Ben Hassine, 2001	Rhinobatos rhinobatos (Linnaeus, 1758) (3)	cloaca	Neifar, 2001 ; Neifar ef al., 2001a
Cathariotrema sp.	Galeorhinus galeus (Linnaeus, 1758) (1)	nasal capsule	Neifar, 2001
Empruthotrema raiae (MacCallum, 1916) Johnston and Tiegs, 1922	Raja clavata Linnaeus, 1758, R. miraletus Linnaeus, 1758, R. radula Delaroche, 1809, Rostroraja alba (Lacepède, 1803) (1,2,3)	nasal tissues	Neifar, 2001
Heliocotyle kartasi Neifar, Euzet and Ben Hassine, 1999	Aetomylaeus bovinus (Geoffroy Saint-Hilaire, 1817) (1, 3)	gills	Neifar, 2001, Neifar et al., 1999a
Heterocotyle capapei Neifar, Euzet and Ben Hassine, 2000	Dasyatis tortonesei Capapé, 1975 (1,3)	gills	Neifar, 2001; Neifar et al., 2000
Heterocotyle forcifera Neifar, Euzet and Ben Hassine, 1999	Taeniura grabata Geoffroy Saint-Hilaire, 1817 (1, 3)	gills	Neifar, 2001; Neifar et al., 1999 b
Heterocotyle minima (MacCallum, 1916) Price, 1938	Bathytoshia centroura (Mitchill, 1815) (1, 3)	gills	Neifar, 2001
Heterocotyle mokhtarae Neifar, Euzet and Ben Hassine, 1999	Taeniura grabata Geoffroy Saint-Hilaire, 1817 (1, 3)	gills	Neifar, 2001; Neifar et al., 1999 b
Heterocotyle pastinacae Scott, 1904	Dasyatis pastinaca (Linnaeus, 1758) (1, 3) Dasyatis marmorata (Steindachner, 1892) (1, 3)	gills	Neifar, 1995, 2001; Neifar et al., 1998 Neifar, 2001; Neifar et al., 1998
Heterocotyle scotti Neifar, Euzet and Ben Hassine, 1998	Dasyatis marmorata (Steindachne, 1892) (3)	gills	Neifar, 2001; Neifar et al., 1998
	Dasyatis pastinaca (Linnaeus, 1758) (1) (1,3)		Neifar, 1995
			Neifar, 2001; Neifar et al., 1998
Heterocotyle similis Neifar, Euzet and Ben Hassine, 1998	Dasyatis pastinaca (Linnaeus, 1758) (1, 3)	gills	Neifar et al., 1998
Heterocotyle striata Neifar, Euzet and Ben Hassine, 1999	Taeniura grabata Geoffroy Saint-Hilaire, 1817 (1, 3)	skin ventral surface	Neifar, 2001; Neifar et al., 1999 b
Mehracotyle insolita Neifar, Euzet and Ben Hassine, 2002	Rhinobatos cemiculus Geoffroy Saint-Hilaire, 1817 (1, 3)	interbranchial septa	Neifar, 2001; Neifar et al., 2002
Merizocotyle concinna (Scott, 1904) Chisholm, Wheeler and	Dasyatis marmorata (Steindachner, 1892) (1)	nasal tissues	Neifar, 2001
Beverley-Burton, 1995	Dasyatis pastinaca (Linnaeus, 1758) (1, 3)
Monocotyle sp.	Aetomylaeus bovinus (Geoffroy Saint-Hilaire, 1817) (1, 3)	gills	Neifar, 2001
Myliocotyle pteromylaei Neifer, Euzet and Ben Hassine, 1999	Aetomylaeus bovinus (Geoffroy Saint-Hilaire, 1817) (1, 3)	gills	Neifar, 2001 ; Neifar et al., 1999 c
Neoheterocotyle ktarii Neifar Euzet and Ben Hassine, 2001	Rhinobatos rhinobatos (Linnaeus, 1758) (3)	gills	Neifar, 2001 ; Neifar et al., 2001 a
Thaumatocotyle tunisiensis Neifar, Euzet and Ben Hassine, 2000	Dasyatis tortonesei Capapé, 1975 (1, 3)	nasal tissues	Neifar, 2001; Neifar et al., 2000
Triloculotrema euzeti Boudaya and Neifar, 2016	Mustelus punctulatus Risso, 1827 (1, 2, 3)	nasal tissues	Boudaya and Neifar, 2016
Family Plectanocotylidae Monticelli, 1903
Plectanocotyle major Boudaya, Neifar and Euzet, 2006	Chelidonichthys obscurus (Walbaum, 1792) (3)	gills	Boudaya et al., 2006
Plectanocotyle gumardi (Van Beneden and Hesse, 1863) Llewellyn,	Chelidonichthys cuculus (Linnaeus, 1758), C. lastoviza	gills	Ktari, 1971
1941	(Bonnaterre, 1788), C. obscurus (Walbaum, 1792) (1)
Family Polystomatidae Gamble, 1896
Neopolystoma euzeti Combes and Ktari, 1976	Mauremys caspica var. leprosa (Gmelin, 1774) (4)	urinary bladder and rectum	Combes and Ktari, 1976
Polystomoides ocellatus (Rudolphi, 1819) Ozaki, 1935	Mauremys caspica var. leprosa (Gmelin, 1774) (4)	esophagus	Lambert et al., 1978
Family Pseudodactylogyridae Gusev, 1965
Pseudodactylogyrus anguillae (Yin and Sproston, 1948) Gusev, 1965	Anguilla anguilla (Linnaeus, 1758) (1)	gills	Neifar, 1995
Family Pyragraphoridae Yamaguti, 1963
Pyragraphorus hollisae Euzet and Ktari, 1970	Trachinotus ovatus (Linnaeus, 1758) (1)	gills	Euzet and Ktari, 1970 c; Neifar, 1995
Family Tetraonchoididae Bychowsky, 1951
Tetraonchoides paradoxus Bychowsky, 1951	Uranoscopus scaber Linnaeus, 1758 (1)	gills	Neifar, 1995

Table 2

List of Tunisian aquatic host species and their monogenean parasites.

Host species	Monogenean species
Aetomylaeus bovinus (Geoffroy Saint-Hilaire, 1817)	Heliocotyle kartasi Neifar, Euzet and Ben Hassine, 1999; Monocotyle sp.; Myliocotyle pteromylaei Neifer, Euzet and Ben Hassine, 1999
Anguilla anguilla (Linnaeus, 1758)	Pseudodactylogyrus anguillae (Yin and Sproston, 1948) Gusev, 1965
Balistes capriscus Gmelin, 1789	Haliotrema balisticus (Hargis, 1955) Yamaguti, 1963
Bathytoshia centroura (Mitchill, 1815)	Dasyonchocotyle spiniphallus Hargis, 1955; Heterocotyle minima (MacCallum, 1916) Price, 1938
Belone belone (Linnaeus, 1758)	Axine belones Abildgaard, 1794
Belone svetovidovi Collette and Parin, 1970	Axine sp.
Boops boops (Linnaeus, 1758)	Choricotyle chrysophryi Van Beneden and Hesse, 1863
Brama brama (Bonnaterre, 1788)	Gotocotyla acanthura (Parona and Perugia, 1896) Meserve, 1938
Chelidonichthys cuculus (Linnaeus, 1758)	Plectanocotyle gurnardi (Van Beneden and Hesse, 1863) Llewellyn, 1941
Chelidonichthys lastoviza (Bonnaterre, 1788)	Plectanocotyle gurnardi (Van Beneden and Hesse, 1863) Llewellyn, 1941
Chelidonichthys lucerna (Linnaeus, 1758)	Trochopus heteracanthus Massa, 1903; Trochopus pini (Van Beneden and Hesse, 1863) Massa, 1903
Chelidonichthys obscurus (Walbaum, 1792)	Plectanocotyle gurnardi (Van Beneden and Hesse, 1863) Llewellyn, 1941; Plectanocotyle major Boudaya, Neifar and Euzet, 2006
Chelon auratus (Risso, 1810)	Ligophorus szidati Euzet and Suriano, 1977; Ligophorus vanbenedenii (Parona and Perugia, 1890) Euzet and Suriano, 1977; Solostamenides mugilis (Vogt, 1879) Unnithan, 1971
Chelon labrosus (Risso, 1827)	Ligophorus angustus Euzet and Suriano, 1977; Solostamenides mugilis (Vogt, 1879) Unnithan, 1971
Chelon ramada (Risso, 1827)	Solostamenides mugilis (Vogt, 1879) Unnithan, 1971
Chelon saliens (Risso, 1810)	Ligophorus acuminatus Euzet and Suriano, 1977; Ligophorus heteronchus Euzet and Suriano, 1977; Ligophorus macrocolpos Euzet and Suriano, 1977; Ligophorus minimus Euzet and Suriano, 1977; Solostamenides mugilis (Vogt, 1879) Unnithan, 1971
Dasyatis marmorata (Steindachner, 1892)	Heterocotyle pastinacae Scott, 1904; Heterocotyle scotti Neifar, Euzet and Ben Hassine, 1998; Merizocotyle concinna (Scott, 1904) Chisholm, Wheeler and Beverley-Burton, 1995; Neoentobdella diadema (Monticelli, 1902) Kearn and Whittington, 2005; Trimusculotrema micracantha (Euzet and Maillard, 1967) Whittington and Barton, 1990
Dasyatis pastinaca (Linnaeus, 1758)	Heterocotyle pastinacae Scott, 1904; Heterocotyle scotti Neifar, Euzet and Ben Hassine, 1998; Heterocotyle similis Neifar, Euzet and Ben Hassine, 1998; Merizocotyle concinna (Scott, 1904) Chisholm, Wheeler and Beverley-Burton, 1995; Neonchocotyle pastinacae Ktari and Maillard, 1972; Neoentobdella diadema (Monticelli, 1902) Kearn and Whittington, 2005; Trimusculotrema micracantha (Euzet and Maillard, 1967) Whittington and Barton, 1990
Dasyatis tortonesei Capapé, 1975	Heterocotyle capapei Neifar, Euzet and Ben Hassine, 2000; Neoentobdella diadema (Monticelli, 1902) Kearn and Whittington, 2005; Thaumatocotyle tunisiensis Neifar, Euzet and Ben Hassine, 2000; Trimusculotrema micracantha (Euzet and Maillard, 1967) Whittington and Barton, 1990
Dentex dentex (Linnaeus, 1758)	Gotocotyla acanthura (Parona and Perugia, 1896) Meserve, 1938
Dentex gibbosus (Rafinesque, 1810)	Pseudoaspinatrium euzeti (Ktari, 1971) Mamaev, 1986; Lamellodiscus euzeti Diamanka, Boudaya, Toguebaye and Pariselle, 2011
Dentex maroccanus Valenciennes, 1830	Lamellodiscus crampus Neifar, 2008
Dicentrarchus labrax (Linnaeus, 1758)	Diplectanum aequans (Wagener, 1857)
Dicentrarchus puctatus (Bloch, 1792)	Diplectanum laubieri Lambert and Maillard, 1974
Diplodus annularis (Linnaeus, 1758)	Lamellodiscus elegans Bychowsky, 1957; Lamellodiscus fraternus Bychowsky, 1957; Polylabris tubicirrus (Paperna and Kohn, 1964) Mamaev and Parukhin, 1976
Diplodus puntazzo (walbaum, 1792)	Lamellodiscus bidens Euzet, 1984; Lamellodiscus hilii Euzet, 1984; Lamellodiscus ignoratus Palombi, 1943; Lamellodiscus impervius Euzet, 1984; Lamellodiscus theroni Amine, Euzet and Kechemir-Issad, 2007
Diplodus sargus (Linnaeus, 1758)	Lamellodiscus abbreviatus Sanfilippo, 1978; Lamellodiscus elegans Bychowsky, 1957; Lamellodiscus furcosus Euzet and Oliver, 1966; Lamellodiscus gussevi Sanfilippo, 1978; Lamellodiscus ignoratus Palombi, 1943; Lamellodiscus mirandus Euzet and Oliver, 1966
Diplodus vulgaris (Geoffroy Saint-Hilaire, 1817)	Lamellodiscus ergensi Euzet and Oliver, 1966; Lamellodiscus furcosus Euzet and Oliver, 1966; Lamellodiscus ignoratus Palombi, 1943
Echeneis naucrates Linnaeus, 1758	Dionchus agassizi Goto, 1899; Dionchus remorae (MacCallum, 1916) Price, 1938
Engraulis encrasicolus (Linnaeus, 1758)	Pseudanthocotyloides heterocotyle (Van Beneden, 1871) Euzet and Prost, 1969
Euthynnus alletteratus (Rafinesque, 1810)	Capsala maccallumi Price, 1939; Neohexostoma euthynni (Meserve, 1938) Price, 1961
Galeorhinus galeus (Linnaeus, 1758)	Cathariotrema sp.
Gymnura altavela (Linnaeus, 1758)	Heteronchocotyle gymnurae Neifar, Euzet and Ben Hassine, 2001
Hyporthodus haifensis (Ben-Tuvia, 1953)	Pseudorhabdosynochus sulamericanus Santos, Buchmann and Gibson, 2000
Lithognathus mormyrus (Linnaeus, 1758)	Bychowskicotyla mormyri (Lorenz, 1878) Unnithan, 1971; Lamellodiscus flagellatus Boudaya, Neifar and Euzet, 2009; Lamellodiscus verberis Euzet and Oliver, 1967
Luciobarbus callensis (Valenciennes, 1842)	Dactylogyrus heteromorphus El Garbi, Birgi and Lambert, 1994; Dactylogyrus tunisiensis El Garbi, Birgi and Lambert, 1994
Mauremys caspica var. leprosa (Gmelin, 1774)	Neopolystoma euzeti Combes and Ktari, 1976
Merluccius merluccius (Linnaeus, 1758)	Anthocotyle merlucii Van Beneden and Hesse, 1863
Mugil cephalus Linnaeus, 1758	Benedenia monticellii (Parona and Perugia, 1895) Johnston, 1929; Ligophorus chabaudi Euzet and Suriano, 1977; Metamicrocotyla cephalus (Azim, 1939) Hargis, 1954; Solostamenides mugilis (Vogt, 1879) Unnithan, 1971
Mustelus mustelus (Linnaeus, 1758)	Calicotyle stossichi Braun, 1899; Erpocotyle catenulata (Guberlet, 1933) Yamaguti, 1963
Mustelus punctulatus Risso, 1827	Calicotyle palombi Euzet and Williams, 1960; Calicotyle stossichi Braun, 1899; Erpocotyle sp.; Triloculotrema euzeti Boudaya and Neifar, 2016
Mycteroperca costae (Steindachner, 1878)	Pseudorhabdosynochus bouaini Neifar and Euzet, 2007; Pseudorhabdosynochus dolicocolpos Neifar and Euzet, 2007; Pseudorhabdosynochus enitsuji Neifar and Euzet, 2007; Pseudorhabdosynochus sinediscus Neifar and Euzet, 2007; Pseudorhabdosynochus sosia Neifar and Euzet, 2007
Mycteroperca marginata (Lowe, 1834)	Echinoplectanum echinophallus (Euzet and Oliver, 1965) Justine and Euzet, 2006; Pseudorhabdosynochus beverleyburtonae (Oliver, 1984) Kritsky and Beverley-Burton, 1986; Pseudorhabdosynochus riouxi (Oliver, 1986) Santos, Buchmann and Gibson, 2000
Mycteroperca rubra (Bloch, 1793)	Pseudorhabdosynochus hayet Chaabane, Neifar, Gey and Justine, 2016; Pseudorhabdosynochus regius Chaabane, Neifar and Justine, 2015
Oblada melanura (Linnaeus, 1758)	Gotocotyla acanthura (Parona and Perugia, 1896) Meserve, 1938; Lamellodiscus elegans Bychowsky, 1957; Lamellodiscus gracilis Euzet and Oliver, 1966
Pagellus erythrinus (Linnaeus, 1758)	Lamellodiscus erythrini Euzet and Oliver, 1966; Microcotyle erythrini Van Beneden and Hesse, 1863
Pagrus auriga Valenciennes, 1843	Lamellodiscus rastellus Neifar, Euzet and Oliver, 2004
Pagrus coeruleostictus (Valenciennes, 1830)	Lamellodiscus sarculus Neifar, Euzet and Oliver, 2004; Lamellodiscus sigilatus Neifar, Euzet and Oliver, 2004
Pomadasys incisus (Bowdich, 1825)	Dicrumenia bychowskyi Mamaev, 1969; Intracotyle hannibali (Euzet and Ktari, 1970) Mamaev, 1977
Pomatomus saltatrix (Linnaeus, 1758)	Microcotyle pomatomi Goto, 1899; Gotocotyla acanthura (Parona and Perugia, 1896) Meserve, 1938; Gotocotyla sp 2. of Neifar, 1995
Raja clavata Linnaeus, 1758 Raja miraletus Linnaeus, 1758	Empruthotrema raiae (MacCallum, 1916) Johnston and Tiegs, 1922; Rajonchocotyle sp. Calicotyle kroyeri Diesing, 1850;
Raja radula Delaroche, 1809	Empruthotrema raiae (MacCallum, 1916) Johnston and Tiegs, 1922; Rajonchocotyle sp.
Rhinobatos cemiculus Geoffroy Saint-Hilaire, 1817	Mehracotyle insolita Neifar, Euzet and Ben Hassine, 2002
Rhinobatos rhinobatos (Linnaeus, 1758)	Calicotyle vicina Neifar, Euzet and Ben Hassine, 2001; Neoheterocotyle ktarii Neifar, Euzet and Ben Hassine, 2001
Rostroraja alba (Lacepède, 1803)	Calicotyle kroyeri Diesing, 1850; Empruthotrema raiae (MacCallum, 1916) Johnston and Tiegs, 1922; Rajonchocotyle sp.
Sardina pilchardus (Walbaum, 1792)	Mazocraes pilchardi (Van Beneden and Hesse, 1863) Sproston, 1946
Sardinella aurita Valenciennes, 1847	Mazocraes sp.
Sardinella maderensis (Lowe, 1838)	Mazocraeoides sardinellae Ktari, 1982
Sarpa salpa (Linnaeus, 1758)	Atrispinum salpae (Parona and Perugia, 1890) Maillard and Noisy, 1979; Lamellodiscus confusus Amine, Euzet and Kechemir-Issad, 2007; Lamellodiscus ignoratus Palombi, 1943; Lamellodiscus parisi Oliver, 1969
Sciaena umbra Linnaeus, 1857	Calceostomella inermis (Parona and Perugia, 1889) Palombi, 1943; Diplectanum simile Bychowsky, 1957
Scomber colias Gmelin, 1789	Kuhnia scombri (Kuhn, 1829) Sproston, 1945; Pseudokuhnia minor (Goto, 1984) Rohde and Watson, 1985
Scomber japonicus Houttuyn, 1782	Grubea cochlear Diesing, 1858; Kuhnia scombri (Kuhn, 1829) Sproston, 1945; Pseudokuhnia minor (Goto, 1984) Rohde and Watson, 1985
Scomber scombrus Linnaeus, 1758	Grubea cochlear Diesing, 1858; Kuhnia scombri (Kuhn, 1829) Sproston, 1945
Scyliorhinus stellaris (Linnaeus, 1758)	Hexabothrium appendiculatum (Kuhn, 1829) Von Nordmann, 1840
Seriola dumerili (Risso, 1810)	Heteraxine seriola (Ishii, 1936) Sproston, 1946
Serranus cabrilla (Linnaeus, 1758)	Protolamellodiscus serranelli (Euzet and Oliver, 1965) Oliver, 1969
Serranus scriba (Linnaeus, 1758)	Protolamellodiscus serranelli (Euzet and Oliver, 1965) Oliver, 1969
Siganus luridus (Rüppell, 1829)	Glyphidohaptor plectocirra (Paperna, 1972) Kritsky, Galli and Yang, 2007; Polylabris mamaevi Ogawa and Egusa, 1980
Siganus rivulatus Forsskål and Niebuhr, 1775	Glyphidohaptor plectocirra (Paperna, 1972) Kritsky, Galli and Yang, 2007; Polylabris mamaevi Ogawa and Egusa, 1980
Sparus aurata Linnaeus, 1758	Lamellodiscus echeneis (Wagener, 1857) Diesing, 1858;
Sphyraena sphyraena (Linnaeus, 1758)	Cotyloatlantica mediterranea (Euzet and Trilles, 1960) Bravo-Hollis, 1984
Spicara maena (Linnaeus, 1758)	Bivagina alcedinis (Parona and Perugia, 1889) Yamaguti, 1963
Spicara smaris (Linnaeus, 1758)	Lamellodiscus knoepffleri Oliver, 1969
Synapturichthys kleinii (Risso, 1827)	Pseudodiplectanum syrticum Derbel, Boudaya and Neifar, 2007
Taeniura grabata Geoffroy Saint-Hilaire, 1817	Heterocotyle forcifera Neifar, Euzet and Ben Hassine, 1999; Heterocotyle mokhtarae Neifar, Euzet and Ben Hassine, 1999; Heterocotyle striata Neifar, Euzet and Ben Hassine, 1999; Neoentobdella apiocolpos (Euzet and Maillard, 1967) Kearn and Whittington, 2005
Thunnus thynnus (Linnaeus, 1758)	Hexostoma thynni (Delaroche, 1811) Rafinesque, 1815
Torpedo marmorata Risso, 1810	Amphibdelloides kechemirae Tazerouti, Euzet and Neifar, 2006; Amphibdelloides vallei Llewellyn, 1960; Epicotyle torpedinis (Price, 1942) Euzet and Maillard, 1974
Torpedo torpedo (Linnaeus, 1758)	Amphibdella paronaperugiae Llewellyn, 1960; Amphibdelloides benhassinae Tazerouti, Euzet and Neifar, 2006
Trachinotus ovatus (Linnaeus, 1758)	Gotocotyla acanthura (Parona and Perugia, 1896) Meserve, 1938; Gotocotyla sp1. of Neifar, 1995; Pyragraphorus hollisae Euzet and Ktari, 1970
Trachinus araneus Cuvier, 1829	Pauciconfibula trachini (Parona and Perugia, 1889) Dilllon and Hargis, 1965; Bradyhaptorus trachini (Parona and Perugia, 1890) Unnithan, 1971
Trachinus draco Linnaeus, 1758 Trachinus radiatus Cuvier, 1829	Bradyhaptorus trachini (Parona and Perugia, 1890) Unnithan, 1971 Bradyhaptorus trachini (Parona and Perugia, 1890) Unnithan, 1971; Pauciconfibula trachini (Parona and Perugia, 1889) Dilllon and Hargis, 1965
Trachurus mediterraneus (Steindachner, 1868)	Cemocotyle trachuri Dillon and Hargis, 1965; Gastrocotyle trachuri Van Beneden and Hesse, 1863; Pseudaxine trachuri Parona and Perugia, 1890
Trachurus trachurus (Linnaeus, 1758)	Cemocotyle trachuri Dillon and Hargis, 1965; Gastrocotyle trachuri Van Beneden and Hesse, 1863; Pseudaxine trachuri Parona and Perugia, 1890
Trisopterus capelanus (Lacepède, 1800) Tylosurus acus imperialis (Rafinesque, 1810)	Diclidophora sp. Nudaciraxine imperium Châari, Derbel and Neifar, 2010; Pseudoaspinatrium gallieni (Euzet and Ktari, 1971) Mamaev, 1986
Umbrina cirrosa Linnaeus, 1758	Calceostomella inermis (Parona and Perugia, 1889) Palombi, 1943; Diplectanum grassei Oliver, 1974; Diplectanum melvillei Oliver and Paperna, 1984; Sciaenacotyle pancerii (Sonsino, 1891) Mamaev, 1989
Uranoscopus scaber Linnaeus, 1758	Tetraonchoides paradoxus Bychowsky, 1951

Monogenea were reported from hosts of different localities in the coast of Tunisia, at the northern area (Gulf of Tunis 37°N, 10°30’E), at the eastern area (Gulf of Hammamet 36°5’N, 10°45’E) and at the southern area (Gulf of Gabes 34°05’N, 10°26’E) (Fig. 1). In addition, fish were sampled from the Gulf of Gabes at Skhira (34°05’N; 10°01’E), Kerkennah (34°45’N; 11°17’E) and Sidi Mansour (34°46’N; 10°48’E). This fish has been caught by locals fishermen. The specimens, coming from the coastal fishing, were identified using Fisher and Whitehead . These fish were dissected as soon as they died and examined for Monogenea. Living parasites were detached then partially compressed between slide and coverslip. They were examined using an optical microscope. Some parasites were fixed with 70 % alcohol and stained with Semichon’s acetic carmine. After dehydration using a graded ethanol series, the parasites were cleared in clove oil and mounted in Canada balsam for identification. Several specimens were fixed, stained and mounted directly in ammonium picrate glycerine after Malmberg (1957). The slides were sealed with Canada balsam. The host-parasite list of our work is arranged in alphabetical order of Monogenea-family (Table 3). Prevalence (P), mean abundance (MA) and mean intensity (MI) are determined according to Bush .

Fig. 1

Map of Tunisia showing sites of monogenean records.

Table 3

Monogenean species diversity in the Gulf of Gabes.

Monogenean family	Monogenean species	Fish species	Number of fishes examined	P (%)	MA	Ml
Ancyrocephalidae	Haliotrema balisticus	Batistes capriscus	2	100	6	6
	Ligophorus vanbenedenii	Chelon auratus	21	80	6.4	8
	Ligophorus szidati			80	4.8	6
	Ligophorus angustus	Chelon labrosus	4	100	3	3
	Ligophorus macrocolpos	Chelon saliens	12	50	0.75	1
	Ligophorus minimus			75	2	2
	Ligophorus heteronchus			100	8.33	8
	Ligophorus acuminatus			100	10.3	10
Capsalidae	Benedenia monticellii	Mugil cephalus	22	13.6	0.22	1
	Capsala maccallumi	Euthynnus alletteratus	4	50	3	6
Diplectanidae	Diplectanum aeguans	Dicentrarchus labrax	2	100	10	10
	Pseudodiplectanum syrticum	Synapturichthys kleinii	30	90	15.3	17
	Lamellodiscus knoepffleri	Spicara smaris	9	88	4.4	5
	Lamellodiscus elegans	Diplodus annularis	82	22	0.5	2
	Lamellodiscus fraternus			71.4	4.7	6
	Lamellodiscus ignoratus	Diplodus puntazzo	6	83.3	18	21
		Diplodus vulgaris	5	100	28	28
	Lamellodiscus theroni	Diplodus puntazzo	6	66	13	20.5
	Lamellodiscus hilii			50	1	2
	Lamellodisscus bidens			66	2.2	4
	Lamellodiscus impervius			33	1.6	4
	Lamellodiscus ergensi	Diplodus vulgaris	5	100	3.6	3
	Lamellodiscus verberis	Lithognatus mormyrus	4	100	38	38
	Lamellodiscus erythrini	Pagellus erythrinus	7	85.7	15.28	20
	Lamellodiscus rastellus	Pagrus auriga	2	100	24	24
	Lamellodiscus confusus	Sarpa salpa	14	100	12	12
	Lamellodiscus parisi			28	0.5	2
	Lamellodiscus echeneis	Sparus aurata	4	75	1.5	2
Hexostomatidae	Neohexostoma euthynni	Euthynnus alletteratus	4	75	6	8
Microcotylidae	Metamicrocotyla cephalus	Mugil cephalus	22	13	0.6	7
	Polylabris tubicirrus	Diplodus annularis	82	2.5	1	1
	Microcotyle erythrini	Pagellus erythrinus	7	42	0.85	2
Mazocraeidae	Grubea cochlear	Scomber japonicus	21	19	0.1	1
	Pseudokuhnia minor			55	2.35	4
	Kuhnia scombri			18	0.36	2
	Pseudanthocotyloides heterocotyle	Engraulis encrasicolus	31	54	2.4	4
	Mazocraes sp.	Sardinella aurita	46	8.6	0.08	1
	Mazocraes pilchardi	Sardina pilchardus	63	15	0.19	1

Ethical Approval and/or Informed Consent

All applicable institutional, national and international guidelines for the care and use of animals were followed.

Results and Discussion

The checklist of Monogenea species recorded from Tunisian waters includes 153 species from 89 host species. Among them, 37 species are reported from elasmobranchs and two species, Neopolystoma euzeti Combes & Ktari, 1976 and Polystomoides ocellatus (Rudolphi, 1819) Ozaki, 1935 from the freshwater tortoise Mauremys caspica var. leprosa (Gmelin, 1774).

The diversity of Monogenea in the coast of Tunisia is similar to that of other regions of the Mediterranean. For example, in the coast of Italy, there are 141 species of Monogenea reported from 93 host species (Strona ). However, this diversity is lower than the other regions of the World. In fact, 367 Monogenea are reported from 363 Mexican aquatic vertebrates (Mendoza-Garfias ) and 581 species from 263 inland aquatic vertebrates in China (Xia ). In general, the number of parasites reported must be dependent on the number of host examined as well as the number of host in the regions. Thus, areas with the highest host diversity have the highest diversity of parasites reported if the research efforts into fish parasites discovery are important. Jorge and Poulin (2018) note that there is a poor match between the host species richness and the parasite species discovery. Some areas with hotspots of host diversity, such as the tropical regions, suffer from a research deficit in parasite discovery.

In this list, twenty six families are recorded; among them the Diplectanidae Monticelli, 1903 which represents the major mono-genean family. Most species belong to the genus Lamellodiscus Johnston & Tiegs, 1922. Many phylogenetic analyses suggest that this group seem to be able to speciate rapidly, even in sympatric condition (Desdevises ) leading to a high diversity.

This list shows that there are more species of Monopisthocotylea (93 species) than Polyopisthocotylea (60 species). Several studies of Monogenea diversity of fish from many localities show that the Monopisthocotylea is the richest group (Justine , 2012; Mendoza-Garfias ). This difference is probably due to the differences in speciation rates between the two groups. In fact, sympatric speciation is important factor in parasite diversity, thus multiple congeners species in hosts are the product of speciation and diversification within the host (Poulin, 1999). Therefore, it appears that polyopisthocotyleans exhbit a lower level of speciation than monopisthocotyleans.

This difference can be also explained by the sampling effort. Indeed, examining additional hosts can probably change the number of monogeneans records and may also change the Monopisthocotylea/Polyopisthocotylea balance of recorded species. Moreover, Mendoza-Garfias mentioned that the inventory of the monogeneans fauna in Mexico is not completed because a small number of aquatic vertebrates have been studied for monogeneans in this area.

We noted that the number of Monogenea recorded in the coast of Tunisia has increased in the past 2 decades and led to the description of many new species. The first species were reported by Ktari and Euzet & Ktari in the 1970s. After that, the number has increased with the studies of Neifar (1995, 2001). Since 2001, only 61 species have been reported. This diversity is related to the research effort in terms of the number of studies and the number of fish species examined each year.

The species accumulation curve of this group off the Tunisian waters shows a rising curve that is not reaching the asymptote (Fig. 2). This means that new species (or new records) of monogeneans will continue to be found when other hosts and other areas are examined. This number does not represent the real monogenean fauna of Tunisia but only the number of species reported by the few researchers who have worked in this region. Those described so far represent only a very small proportion of the real richness considering the potential host fauna. It is difficult to have an estimate of relative parasites biodiversity.

Fig. 2

Cumulative curve of monogeneans species recorded from Tunisia.

The discovery of cryptic species affects estimates of parasites diversity. Several cases show that what was once thought to be a single parasite species infecting a few host species turns out to represent on complex of cryptic species, each specific to a single host species (Poulin & Keeney, 2008). Their discovery depends mostly on the application of molecular methods.

Most Monogenea included in the checklist are restricted to fishes with high commercial value. The parasitological study of fishes with minor commercial interest allows us to complete the list of Monogenea from the coast of Tunisia. Moreover, the parasiofauna of some area such as the eastern coast is not well studied.

This list includes results from our study of Monogenea parasite of fish in the Gulf of Gabes. During this study, we have examined 22 teleost fish species belonging to nine families. A total of 37 species of Monogenea belonging to 16 genera and six families were found (Table 3). Among them 15 species are recorded for the first time off the coast of Tunisia (Table 1). The number of helminth species per host species was variable. Diplodus puntazzo (Walbaum) has the richest Monogenea fauna (5 species).