Untangling the Derogenes varicus species complex in Scandinavian waters and the Arctic: description of Derogenes abba n. sp. (Trematoda, Derogenidae) from Hippoglossoides platessoides and new host records for D. varicus (Müller, 1784) sensu stricto.

Several studies have shown that the euryxenic trematode Derogenes varicus (Müller, 1784) represents a species complex. Four lineages have been designated (DV1-4) with the DV1 clade corresponding to D. varicus sensu stricto. Herein, we investigate newly collected specimens of D. varicus sensu lato from Scandinavian and Arctic waters using integrative taxonomy. The trematodes were collected from Melanogrammus aeglefinus, Eutrigla gurnardus, Trachinus draco, and Merluccius merluccius off the Atlantic coast of Sweden and from Hippoglossoides platessoides from Arctic Svalbard. 28S sequences of derogenids from Sweden were identical to D. varicus sensu stricto, confirming its euryxeny. The 28S sequences of Derogenes sp. from H. platessoides were identical to Derogenes DV2 and differed from D. varicus sensu stricto by 3% and from Derogenes DV3 by 2%. The 28S sequence divergences of Derogenes sp. from H. platessoides with D. ruber and D. lacustris were 3 and 10%, respectively. ITS2 and cox1 divergences between Derogenes sp. from H. platessoides and other Derogenes species/lineages were at levels of interspecific differences. The species from H. platessoides is described here as D. abba n. sp. We also examined the type material of Progonus muelleri (Levinsen, 1881), the type and only species of the genus Progonus, with redescription and designations of paralectotypes. Based on specimens from Theodor Odhner's collections at the Swedish Museum of Natural History, SMNH, Stockholm, we provide novel morphological and anatomical data for D. varicus sensu lato species complex. Lastly, we investigated Arthur Looss's "lost collection" of Trematodes at the SMNH and characterised a putative species Derogenes sp. "limula".

Title: Démêler le complexe d'espèces Derogenes varicus dans les eaux scandinaves et arctiques : description de Derogenes abba n. sp. (Trematoda, Derogenidae) parasite d'Hippoglossoides platessoides et nouveaux signalements d'hôtes pour D. varicus (Müller, 1784) sensu stricto. Abstract: Plusieurs études ont montré que le trématode euryxene Derogenes varicus (Müller, 1784) représente un complexe d'espèces. Quatre lignées ont été désignées (DV1­4), le clade DV1 correspondant à D. varicus sensu stricto. Ici, nous étudions des spécimens nouvellement collectés de D. varicus sensu lato dans les eaux scandinaves et arctiques en utilisant la taxonomie intégrative. Les trématodes ont été collectés de Melanogrammus aeglefinus, Eutrigla gurnardus, Trachinus draco et Merluccius merluccius au large de la côte atlantique de la Suède et d'Hippoglossoides platessoides du Svalbard arctique. Les séquences 28S des Derogenidae de Suède étaient identiques à D. varicus sensu stricto, confirmant son euryxénie. Les séquences 28S de Derogenes sp. de H. platessoides étaient identiques à Derogenes DV2 et différaient de D. varicus sensu stricto par 3% et de Derogenes DV3 par 2%. Les divergences des séquence 28S de Derogenes sp. de H. platessoides avec D. ruber et D. lacustris étaient respectivement de 3 et 10%. Les divergences ITS2 et cox1 entre Derogenes sp. de H. platessoides et d'autres espèces/lignées de Derogenes se situaient à des niveaux de différences interspécifiques. L'espèce de H. platessoides est décrite ici comme Derogenes abba n. sp. Nous avons également examiné le matériel type de Progonus muelleri (Levinsen, 1881), type et seule espèce du genre Progonus, avec une redescription et des désignations de paralectotypes. Sur la base de spécimens des collections de Theodor Odhner au Musée suédois d'histoire naturelle (SMNH), Stockholm, nous fournissons de nouvelles données morphologiques et anatomiques sur le complexe d'espèces de D. varicus sensu lato. Enfin, nous avons étudié la « collection perdue ¼ de Trématodes d'Arthur Looss au SMNH et caractérisé une espèce putative, Derogenes sp. « limula ¼.

"Something old, something new, something borrowed, and the oioxeny is true": description of Plectanocotyle jeanloujustinei n. sp. (Polyopisthocotylea, Plectanocotylidae) from the MNHN Helminthology collection with novel molecular and morphological data for P. gurnardi (Van Beneden & Hesse, 1863) (sensu stricto) from Sweden.

Natural history museums worldwide house billions of apposite specimens, offering the potential for cost-free parasitological datasets. Herein, we provide novel morphological and molecular data (28S and cox1) for the polyopisthocotylean Plectanocotyle gurnardi sensu stricto from the type-host Eutrigla gurnardus from Sweden based on newly collected specimens from the Northeast Atlantic, and specimen from T. Odhner's collections at the Swedish Museum of Natural History (Stockholm, Sweden). The newly generated 28S sequences of P. gurnardi from E. gurnardus from the Northeast Atlantic were identical to those from the Western Mediterranean, and nested in a single clade, suggesting the presence of a single species. A 28S sequences of P. gurnardi sensu stricto from Sweden and those from the U.K. (type locality for P. caudata) were identical; we confirm that P. caudata and P. gurnardi are conspecific and formally synonymize them. A single 28S sequence of Plectanocotyle sp. from Chelidonichthys lastoviza off France differed from P. gurnardi from the Northeast Atlantic by 3-4 % and from P. gurnardi from France by 3%. Plectanocotyle sp. ex C. lastoviza off France is clearly not P. gurnardi, suggesting an oioxenic specificity of P. gurnardi to E gurnardus. Careful re-examination of Plectanocotyle cf. gurnardi from C. lastoviza from the Western Mediterranean from the Helminthology collection of Muséum national d'Histoire naturelle (Paris, France) revealed that it differs from all congeners by morphometry (size of clamps, of terminal lappet and its hamuli and uncinuli, and size of atrial spines). The cox1 divergences between P. cf. gurnardi and P. major, P. lastovizae, and P. gurnardi sensu stricto were 10-11 %, 10-11 % and 8 % respectively, falling within the interspecific variations range. Plectanocotyle from the Mediterranean is described as a new species, P. jeanloujustinei n. sp. We apprise nomenclature problems in Plectanocotyle and consider P. elliptica a species inquirenda.

Redescription and Molecular Characterisation of Derogenes ruber Lühe, 1900 (Hemiuroidea: Derogenidae) from Chelidonichthys lastoviza (Scorpaeniformes: Triglidae) in the Western Mediterranean.

PURPOSE: Derogenes ruber Lühe, 1900, the type-species of the genus Derogenes Lühe, 1900, is a poorly known derogenid digenean. The original description of this species was not illustrated and aspects of the morphology of the parasite from the type-host remain scarce. Available records of this species were brief and/or lacked illustrations and were based on morphology alone. Additionally, molecular data for Derogenes spp. are warranted to untangle species complexes as they provide a better assessment of interspecific genetic divergence. METHODS: Derogenes ruber is redescribed based on newly collected specimens from the gall bladder of its type-host Chelidonichthys lastoviza (Bonnaterre, 1788) collected in the Western Mediterranean off the Algerian coast during 2017-2019 and molecular data are provided using a partial fragment of the nuclear 28S ribosomal RNA gene (28S rRNA), the internal transcribed spacer 2 (ITS2) and a fragment of the mitochondrial cytochrome c oxidase subunit 1 (cox1) gene. RESULTS: We herein provide a detailed illustrated redescription and morphometric data of D. ruber from its type-host C. lastoviza. We report a new geographical record (off Algeria) for it. Derogenes ruber is also genetically characterised for the first time. Species/lineages of Derogenes were recovered in five strongly supported reciprocally monophyletic clades: (i) D. ruber from C. lastoviza off Algeria; (ii) D. lacustris from Galaxias maculatus (Jenyns) off Argentina; (iii) Lineage "D. varicus DV1" (D. varicus sensu stricto) from fish hosts in the White and Barents seas and the North Sea; (iv) Lineage "D. varicus DV2" from mollusc hosts in the White Sea; and (v) Lineage "D. varicus DV3" from Eumicrotremus fedorovi Mandrytsa. in the Pacific Ocean. Hence, comparison of the newly generated sequences with other available data for Derogenes species supports the distinction of D. ruber confirming its taxonomic status and helping assess interspecific variation. Comparison of D. ruber with the closely related species Derogenes latus revealed overlaps in morphometric data and the validity of the latter species is questioned. CONCLUSION: The combination of morphological and molecular data provided for D. ruber provides a firm foundation for further investigations of Derogenes spp. Although we do describe herein material of D. ruber from the type-host, given that the occurrence of a single Derogenes species in various hosts has been challenged by molecular data, and both D. lacustris and D. varicus sensu stricto had been genetically proven to occur in various hosts, D. ruber and D. latus may be indeed synonymous. Additional sequencing effort on Derogenes spp. will strengthen systematic comparative studies and evolutionary relationships within the Derogenidae in general.

Morphological and molecular characterization of Encotyllabe vallei Monticelli, 1907 (Monopisthocotylea, Monogenea) from the gilthead seabream Sparus aurata Linnaeus (Teleostei, Sparidae) from the southwestern Mediterranean and notes on host specificity of the genus Encotyllabe Diesing, 1850.

Incomplete original descriptions, the unavailability or poor conditions of specimens and the lack of detailed redescriptions have caused the validity of several species of the genus Encotyllabe Diesing, 1850 to be questioned. To date, seven of the recognized species were described upon one or two specimens, hindering study of intraspecific variations. This was made worse by considering few morphoanatomical differences sufficient to erect new species. Among Encotyllabe spp. occurring in Mediterranean waters, E. vallei was first described from the gilt-head bream Sparus aurata (Sparidae) off Italy. Although beautifully illustrated for a paper from that century, morphometric data for E. vallei from the type-host S. aurata remain unavailable. Previous records of E. vallei provided either morphometrical or molecular data, and its validity was questioned. We provide a redescription of E. vallei based on newly collected specimens from the S. aurata from the southwestern Mediterranean (off Algeria) using integrative taxonomy. Analysis of cox1 sequences of E. vallei from S. aurata, compared to sequences from other sparid hosts, mainly Pagellus bogaraveo, revealed a divergence not exceeding 2%, suggesting a stenoxenic specificity for this monogenean. Given that P. bogaraveo is the type-host for Encotyllabe pagelli, we were tempted to suggest a synonymy between E. vallei and E. pagelli. We refrained from doing so because E. pagelli was first described from the Atlantic coast off Brest, France. Morphological data for Encotyllabe from P. bogaraveo are warranted assessing the host specificity of E. vallei and whether there might be a species complex within individual sparid fish species.

Hidden in the fog: morphological and molecular characterisation of Derogenes varicus sensu stricto (Trematoda, Derogenidae) from Sweden and Norway, and redescription of two poorly known Derogenes species.

Derogenes varicus (Müller, 1784) is widely reported as a trematode with exceptionally low host specificity and a wide, bipolar distribution. However, several recent studies have suggested that D. varicus represents a species complex and based on molecular evidence, four genetic lineages (labeled as "DV1-4") have been designated within the D. varicus species complex. This possibility requires improved (ideally molecular) characterisation of specimens from the type-host (Salmo salar) and type-locality (off Denmark). During examination of trematode parasites of fish from Scandinavian and Arctic waters (Sweden and Norway), we found specimens of D. varicus in the stomach of Merlangius merlangus off the coast of Sweden, and in Gadus morhua off the coast of Sweden and Norway; we compared them to D. varicus from the type-host, the Atlantic salmon Salmo salar from Norway, to verify their conspecificity. Newly generated sequences (28S rDNA, ITS2 and cox1) of Scandinavian and Arctic specimens consistent with D. varicus all formed a single clade, DV1. 28S sequences of D. varicus from S. salar from Norway, i.e., close to the Danish type locality, clustered within the DV1 clade along with sequences of D. varicus from various hosts including Limanda limanda, G. morhua and Myoxocephalus scorpius from the White Sea and the Barents Sea (Russia), without any host-related structuring. We thus consider that the lineage DV1 represents D. varicus sensu stricto. Additionally, specimens from M. merlangus had a similar morphology and anatomy to those of D. varicus from L. limanda, G. morhua and M. scorpius from T. Odhner's collection, supporting the presence of a single species in the DV1 lineage designated herein as D. varicus sensu stricto. We redescribe D. varicus sensu stricto, add new morphological characters and provide morphometric data. We infer that D. varicus types DV2-4 all relate to separate species. We also revise type-specimens of Derogenes minor Looss, 1901 from the A. Looss collection in the Swedish Museum of Natural History and provide redescriptions of it and of the type-species of the genus, Derogenes ruber Lühe, 1900. In light of their morphological distinctiveness relative to D. varicus sensu stricto, we reinstate D. parvus Szidat, 1950 and D. fuhrmanni Mola, 1912.

Title: Caché dans le brouillard : caractérisation morphologique et moléculaire de Derogenes varicus sensu stricto (Trematoda, Derogenidae) de Suède et de Norvège, et redescription de deux espèces de Derogenes peu connues. Abstract: Derogenes varicus (Müller, 1784) est largement signalé comme un trématode avec une spécificité d'hôte exceptionnellement faible et une large distribution bipolaire. Cependant, plusieurs études récentes ont suggéré que D. varicus représente un complexe d'espèces. Cette possibilité nécessite une meilleure caractérisation (idéalement moléculaire) des spécimens de l'hôte-type (Salmo salar) et de la localité-type (au large du Danemark). Lors de l'examen des trématodes parasites des poissons des eaux scandinaves et arctiques (Suède et Norvège), nous avons trouvé des spécimens de D. varicus dans l'estomac de Merlangius merlangus au large de la Suède, et dans Gadus morhua au large de la Suède et de la Norvège ; nous les avons comparés à D. varicus de l'hôte-type, le saumon atlantique Salmo salar de Norvège, pour vérifier leur conspécificité. Les séquences nouvellement générées (ADNr 28S, ITS2 et cox1) de spécimens scandinaves compatibles avec D. varicus ont toutes formé un seul clade, DV1. Les séquences 28S de D. varicus de S. salar de Norvège, c'est-à-dire proches de la localité-type au Danemark, se regroupent dans le clade DV1 avec des séquences de D. varicus de divers hôtes dont Limanda limanda, G. morhua et Myoxocephalus scorpius de la mer Blanche et la mer de Barents (Russie), sans aucune structuration liée à l'hôte. Nous considérons donc que la lignée DV1 représente D. varicus sensu stricto. De plus, les spécimens de M. merlangus avaient une morphologie et une anatomie similaires à celles de D. varicus de L. limanda, G. morhua et M. scorpius de la collection de T. Odhner, confirmant la présence d'une seule espèce dans la lignée DV1 désignée ici comme D. varicus sensu stricto. Nous redécrivons D. varicus sensu stricto, ajoutons de nouveaux caractères morphologiques et fournissons des données morphométriques. Nous en déduisons que les types DV2-4 de D. varicus se rapportent tous à des espèces distinctes. Nous révisons également les spécimens-types de Derogenes minor Looss, 1901 de la collection d'A. Looss du Muséum Suédois d'Histoire Naturelle, et fournissons des redescriptions de cette espèce et de l'espèce-type du genre, Derogenes ruber Lühe, 1900. À la lumière de leur distinction morphologique par rapport à D. varicus sensu stricto, nous rétablissons D. parvus Szidat, 1950 et Derogenes fuhrmanni Mola, 1912.

Morphological and molecular characterisation of Tristoma integrum Diesing, 1850 (Monogenea, Capsalidae), including its complete mitogenome.

Capsalids are monopisthocotylean monogenean parasites found on the skin and gills of fish. Capsalines (subfamily Capsalinae) are large-sized capsalids, parasitic on highly prized gamefish, and species of Tristoma parasitise only the gills of swordfish (Xiphias gladius). We obtained specimens of Tristoma integrum Diesing, 1850 from swordfish collected off Algeria in the Mediterranean Sea. Here, we describe the specimens, including the key systematics characters of dorsolateral body sclerites. One specimen was used for a next generation sequencing analysis but a part of it, including the sclerites, was mounted on a permanent slide, drawn, and deposited in a curated collection. We characterised the complete mitogenome, the ribosomal cluster (including 18S and 28S) and additional genes such as Elongation factor 1 alpha (EF1α) and Histone 3. We also retrieved molecular information from the host tissue present in the gut of the monogenean and provide the sequence of the complete rRNA cluster of the host, X. gladius. The mitogenome of T. integrum is 13 968 bp in length and codes for 12 protein, 2 rRNA and 22 tRNA. Phylogenies of capsalids were generated from 28S sequences and concatenated mitochondrial protein-coding genes, respectively. In the 28S phylogeny, most subfamilies based on morphology were not found to be monophyletic, but the Capsalinae were monophyletic. In both phylogenies, the closest member to Tristoma spp. was a member of the Capsaloides. In an Appendix, we report the complex nomenclatural history of Tristoma Cuvier, 1817 and its species.

Title: Caractérisation morphologique et moléculaire de Tristoma integrum Diesing, 1850 (Monogenea, Capsalidae) y compris son mitogénome complet. Abstract: Les Capsalidae sont des monogènes Monopisthocotylea parasites de la peau et des branchies des poissons. Les Capsalinae sont des Capsalidae de grande taille qui parasitent les poissons de pêche sportive, très prisés, et les espèces de Tristoma ne parasitent que les branchies de l'espadon (Xiphias gladius). Nous avons obtenu des spécimens de Tristoma integrum Diesing, 1850 à partir d'espadons collectés au large de l'Algérie en Méditerranée. Nous décrivons les spécimens, y compris les caractères systématiques clés des sclérites dorsolatéraux du corps. Un spécimen a été utilisé pour une analyse de séquençage de nouvelle génération, mais une partie de celui-ci, qui comprend des sclérites, a été montée sur une lame permanente, dessinée et déposée dans une collection. Nous avons caractérisé le mitogénome complet, le cluster ribosomal (y compris 18S et 28S) et des gènes supplémentaires tels que le facteur d'élongation 1 alpha (EF1α) et histone 3. Nous avons également récupéré des informations moléculaires à partir du tissu de l'hôte présent dans l'intestin du monogène et fournissons la séquence du cluster d'ARNr complet de l'hôte, X. gladius. Le mitogénome de T. integrum a une longueur de 13 968 pb et code pour 12 gènes de protéines, 2 ARNr et 22 ARNt. Des phylogénies des Capsalidae ont été générées, respectivement à partir des séquences 28S et des gènes codant pour les protéines mitochondriales concaténés. Dans la phylogénie 28S, la plupart des sous-familles basées sur la morphologie n'ont pas été trouvées monophylétiques, à l'exception notable des Capsalinae. Dans les deux phylogénies, les membres les plus proches de Tristoma spp. étaient des espèces de Capsaloides. Dans une annexe, nous rapportons l'histoire nomenclaturale complexe de Tristoma Cuvier, 1817 et de ses espèces.

Truly a hyperparasite, or simply an epibiont on a parasite? The case of Cyclocotyla bellones (Monogenea, Diclidophoridae).

Cyclocotyla bellones Otto, 1823 (Monogenea, Diclidophoridae) is one of the few monogenean species reported as hyperparasitic: the worms dwell on cymothoid isopods, themselves parasites of the buccal cavity of fishes. We present here observations based on newly collected monogenean specimens from Ceratothoa parallela (Otto, 1828), an isopod parasite of Boops boops off Algeria and also investigated its diet to address whether Cy. bellones is indeed a hyperparasite, i.e., whether it feeds on the isopod. We also compared the body shape of various monogeneans belonging to the same family as Cy. bellones, the Diclidophoridae, including Choricotyle cf. chrysophryi Van Beneden & Hesse, 1863, collected from Pagellus acarne off Algeria. No morphological character of the anterior organs suggested any special adaptation in Cy. bellones to the perforation of the crustacean cuticle. The wall of the oesophagus and of the intestine of Cy. bellones was lined with a dark pigment similar to what is usually observed in haematophagous polyopisthocotyleans, and which is derived from ingested fish blood. We noticed that an anterior elongate stem exists only in diclidophorids dwelling on parasitic isopods and never in those attached to the gills. We hypothesize that the anterior stem of the body of Cy. bellones is an anatomical adaptation for the monogenean to feed on the fish while dwelling on the isopod. We thus consider that Cy. bellones is an epibiont of the parasitic crustacean, as it uses it merely as an attachment substrate, and is not a true hyperparasite.

Title: Vraiment un hyperparasite, ou simplement un épibionte sur un parasite ? Le cas de Cyclocotyla bellones (Monogenea, Diclidophoridae). Abstract: Cyclocotyla bellones Otto, 1823 (Monogenea, Diclidophoridae) est l'une des rares espèces de monogènes signalées comme hyperparasites : les vers vivent sur des isopodes cymothoïdes, eux-mêmes parasites de la cavité buccale des poissons. Nous présentons ici des observations basées sur des spécimens de monogènes nouvellement collectés de Ceratothoa parallela (Otto, 1828), un isopode parasite de Boops boops au large de l'Algérie et avons également étudié son régime alimentaire pour déterminer si Cy. bellones est bien un hyperparasite (c'est-à-dire, se nourrit-il de l'isopode ?). Nous avons également comparé la morphologie de divers monogènes appartenant à la même famille que Cy. bellones, les Diclidophoridae, dont Choricotyle cf. chrysophryi Van Beneden & Hesse, 1863, collecté sur Pagellus acarne au large de l'Algérie. Aucun caractère morphologique des organes antérieurs ne suggérait d'adaptation particulière à la perforation de la cuticule des crustacés chez Cy. bellones. La paroi de l'œsophage et de l'intestin de Cy. bellones était tapissée d'un pigment foncé semblable à ce que l'on observe habituellement chez les Polyopisthocotylea hématophages, et qui est issu du sang de poisson ingéré. Nous avons remarqué qu'une partie allongée antérieure n'existe que chez les Diclidophoridae vivant sur des isopodes parasites et jamais chez ceux attachés aux branchies. Nous émettons l'hypothèse que la partie antérieure du corps de Cy. bellones est une adaptation anatomique permettant au monogène de se nourrir du poisson tout en vivant sur l'isopode. Nous considérons donc que Cy. bellones est un épibionte du crustacé parasite, puisqu'il ne l'utilise que comme substrat pour son attachement, et n'est pas un véritable hyperparasite.

Triple barcoding for a hyperparasite, its parasitic host, and the host itself: a study of Cyclocotyla bellones (Monogenea) on Ceratothoa parallela (Isopoda) on Boops boops (Teleostei).

Cyclocotyla bellones Otto, 1823 (Diclidophoridae) is a monogenean characterised by an exceptional way of life. It is a hyperparasite that attaches itself to the dorsal face of isopods, themselves parasites in the buccal cavity of fishes. In this study, Cy. bellones was found on Ceratothoa parallela (Otto, 1828), a cymothoid isopod parasite of the sparid fish Boops boops off Algeria in the Mediterranean Sea. We provide, for the first time, molecular barcoding information of a hyperparasitic monogenean, the parasitic crustacean host, and the fish host, with COI sequences.

TITLE: Triple code-barre pour un hyperparasite, son hôte parasite et l'hôte lui-même : une étude de Cyclocotyla bellones (Monogenea) sur Ceratothoa parallela (Isopoda) sur Boops boops (Teleostei). ABSTRACT: Cyclocotyla bellones Otto, 1823 (Diclidophoridae) est un monogène caractérisé par un mode de vie exceptionnel. C'est un hyperparasite qui se fixe sur la face dorsale d'isopodes, eux-mêmes parasites dans la cavité buccale de poissons. Dans cette étude, Cy. bellones a été trouvé sur Ceratothoa parallela (Otto, 1828), un isopode cymothoïdé parasite du poisson sparidé Boops boops au large de l'Algérie, en mer Méditerranée. Nous fournissons, pour la première fois, des informations de codes-barres moléculaires d'un monogène hyperparasite, de l'hôte crustacé parasite et de l'hôte poisson, avec des séquences de COI.

Redescription and molecular characterization of two species of Pauciconfibula (Monogenea, Microcotylidae) from trachinid fishes in the Mediterranean Sea.

Many Pauciconfibula spp. have a long and complicated taxonomic history. The remaining unsolved taxonomic confusion in this genus is impelled by the host range and status of Pauciconfibula spp. from trachinid fishes: Pauciconfibula trachini and Pauciconfibula draconis, from Trachinus radiatus and Trachinus draco (Trachinidae), respectively. Pauciconfibula trachini was reported on Trachinus draco, type host of Pauciconfibula draconis suggesting thus a stenoxenic specificity for the former monogenean and the occurrence of two congeneric polyopisthocotyleans on a single host. Moreover, the validity of Pauciconfibula draconis was repeatedly questioned by several authors, unjustified synonymy between the two species was proposed, and the delimitations between the two species remained unsolved. Original descriptions were also incomplete and poorly illustrated. In this study, we provide a detailed illustrated redescription of both species based on newly collected specimens of Pauciconfibula trachini and Pauciconfibula draconis collected from their type hosts from off three Mediterranean localities: Algeria, Tunisia, and Italy. Integrative taxonomy using COI sequences was applied to resolve the delimitation between Pauciconfibula trachini and P. draconis. This study provides the first DNA barcoding for members of this genus.

Four polyopisthocotyleans (Platyhelminthes: Monogenea) from carangid fishes in the Mediterranean, off the Algerian coasts.

Four polyopisthocotyleans were collected from the gill filaments of carangids from off the Algerian coast, southern Mediterranean. Specimens of Gastrocotyle trachuri van Beneden & Hesse, 1863 (Gastrocotylidae) and Cemocotyle cf. trachuri Dillon & Hargis, 1965 (Heteraxinidae) from the Mediterranean horse mackerel Trachurus mediterraneus (Steindachner), Zeuxapta seriolae (Meserve, 1938) (Heteraxinidae) from the greater amberjack Seriola dumerili (Risso) and Pyragraphorus hollisae Euzet & Ktari, 1970 (Pyragraphoridae) from the pompano Trachinotus ovatus (Linnaeus) are redescribed based on newly collected specimens. Their taxonomically important morphological features (male copulatory organ and clamp sclerites) are described and illustrated, and the morphometric variation between Mediterranean and oceanic specimens is highlighted. Careful examination of the specimens of Cemocotyle Sproston, 1946 from the Mediterranean revealed that they exhibited unusual features compared with Cemocotyle trachuri Dillon & Hargis, 1965 from the Pacific, mainly the absence of the terminal lappet, thus questioning previous records of this species in the Mediterranean. New geographical locality records are provided for Z. seriolae and P. hollisae. The presence of C. cf. trachuri and Z. seriolae in the Mediterranean is noteworthy as these monogeneans were initially described in the Pacific Ocean. This study extends the geographical range of Z. seriolae to the southern Mediterranean.

Redescription of Flexophora ophidii Prost & Euzet, 1962 (Monogenea: Diclidophoridae) from Ophidion barbatum (Ophidiidae) off the Algerian coast, Mediterranean Sea.

The diclidophorid Flexophora ophidii Prost & Euzet, 1962, the type- and only species of the genus Flexophora Prost & Euzet, 1962, has never been redescribed neither reported since its original description. The latest lacked a diagnosis of the genus that remains unavailable to date. Flexophora ophidii is redescribed and illustrated based on examination of new material from the type-host Ophidion barbatum L. (Ophidiiformes: Ophidiidae) collected off the Algerian coast (new geographical record), southern Mediterranean. A diagnosis of the genus is provided.

No vagina, one vagina, or multiple vaginae? An integrative study of Pseudaxine trachuri (Monogenea, Gastrocotylidae) leads to a better understanding of the systematics of Pseudaxine and related genera.

The presence/absence and number of vaginae is a major characteristic for the systematics of the Monogenea. Three gastrocotylid genera share similar morphology and anatomy but are distinguished by this character: Pseudaxine Parona & Perugia, 1890 has no vagina, Allogastrocotyle Nasir & Fuentes Zambrano, 1983 has two vaginae, and Pseudaxinoides Lebedev, 1968 has multiple vaginae. In the course of a study of Pseudaxine trachuri Parona & Perugia 1890, we found specimens with structures resembling "multiple vaginae"; we compared them with specimens without vaginae in terms of both morphology and molecular characterisitics (COI barcode), and found that they belonged to the same species. We also investigated the male copulatory organ (MCO) of this species, the accuracy of the original description of which is known to be a matter of debate. We found that the genital atrium is armed with 12 hooks arranged as a single circle and a central hollow stylet which is probably involved in traumatic insemination. We redescribed Pseudaxine trachuri based on newly collected specimens from off the coast of Algeria and Museum specimens from off France. Specimens from the type-host, Trachurus trachurus, were found to be similar, for both molecular sequences and morphology, to those found on Boops boops. We can therefore confirm, for the first time with molecular evidence, that B. boops is a host of this parasite. We consider that Pseudaxinoides was erected on the basis of an erroneous interpretation of structures which are not vaginae and, consequently, propose the transfer of most of its species to Pseudaxine, as P. australis (Lebedev, 1968) n. comb., P. bychowskyi (Lebedev, 1977) n. comb., P. caballeroi (Lebedev, 1977) n. comb., P. cariacoensis (Nasir & Fuentes-Zambrano, 1983) n. comb., and P. vietnamensis (Lebedev, Parukhin & Roitman, 1970) n. comb. We also propose Allogastrocotyle dillonhargisorum nom. nov. for Pseudaxine bivaginalis Dillon & Hargis, 1965 to avoid a secondary homonymy.

TITLE: Pas de vagin, un vagin ou plusieurs vagins ? Une étude intégrative de Pseudaxine trachuri (Monogenea, Gastrocotylidae) mène à une meilleure compréhension de la systématique de Pseudaxine et des genres apparentés. ABSTRACT: La présence/absence et le nombre de vagins est une caractéristique majeure de la systématique des Monogènes. Trois genres de Gastrocotylidae partagent une morphologie et une anatomie similaires mais se distinguent par ce caractère : Pseudaxine Parona & Perugia, 1890 n'a pas de vagin, Allogastrocotyle Nasir & Fuentes Zambrano, 1983 a deux vagins et Pseudaxinoides Lebedev, 1968 a plusieurs vagins. Au cours d'une étude de Pseudaxine trachuri Parona & Perugia 1890, nous avons trouvé des spécimens avec des structures ressemblant à des « vagins multiples ¼. Nous les avons comparés, pour la morphologie et les molécules (code à barres COI), à des spécimens sans vagin et avons constaté qu'ils appartenaient à la même espèce. Nous avons également examiné l'organe copulateur mâle (OCM) de cette espèce, dont l'exactitude de la description originale est connue pour être un sujet de débat. Nous avons constaté que l'atrium génital est armé de 12 crochets disposés en un seul cercle et d'un stylet central creux qui est probablement impliqué dans l'insémination traumatique. Nous avons redécrit Pseudaxine trachuri à partir de spécimens nouvellement collectés des côtes algériennes et de spécimens de Musée des côtes de France. Les spécimens de l'hôte type, Trachurus trachurus, se sont avérés similaires, pour les molécules et la morphologie, à ceux trouvés sur Boops boops. Nous pouvons donc confirmer, pour la première fois avec des preuves moléculaires, que B. boops est un hôte de ce parasite. Nous considérons que Pseudaxinoides a été érigé sur la base d'une interprétation erronée de structures qui ne sont pas des vagins et, par conséquent, proposons le transfert de la plupart de ses espèces à Pseudaxine, comme P. australis (Lebedev, 1968) n. comb., P. bychowskyi (Lebedev, 1977) n. comb., P. caballeroi (Lebedev, 1977) n. comb., P. cariacoensis (Nasir et Fuentes-Zambrano, 1983) n. comb. et P. vietnamensis (Lebedev, Parukhin & Roitman, 1970) n. comb. Nous proposons également Allogastrocotyle dillonhargisorum nom. nov. pour Pseudaxine bivaginalis Dillon & Hargis, 1965, pour éviter une homonymie secondaire.

Redescription and molecular characterisation of Allogastrocotyle bivaginalis Nasir & Fuentes Zambrano, 1983 (Monogenea: Gastrocotylidae) from Trachurus picturatus (Bowdich) (Perciformes: Carangidae) off the Algerian coast, Mediterranean Sea.

Allogastrocotyle bivaginalis Nasir & Fuentes Zambrano, 1983, the sole species of Allogastrocotyle Nasir & Fuentes Zambrano, 1983, was described from Trachurus lathami Nichols off Venezuela and never recorded since. We found monogeneans on Trachurus picturatus (Bowdich) (Carangidae) off the Algerian coast, Mediterranean Sea, which had the characteristics of the genus, including, especially, paired vaginae. We compared them to the single available specimen of A. bivaginalis, the holotype. Unfortunately, this holotype does not show clamp structure and soft internal anatomy. Our specimens were similar to A. bivaginalis in most characteristics such as body shape, vaginal openings, and number of testes, clamps and hooks in genital atrium. We detected minor differences in clamp structure but could not ascertain if these were the result of incomplete observations in the original description of real morphological differences. The host fish of our specimens were barcoded (cox1) confirming their specific identity. A phylogenetic analysis of cox1 sequences showed that our sequences of A. bivaginalis were distinct from those of Pseudaxine trachuri Parona & Perugia, 1889 (distance > 15%) and of several other gastrocotylids. Finally, we could not distinguish our Mediterranean specimens from A. bivaginalis, neither on the base of morphology (because the original description is incomplete) nor on molecules (because molecular information is lacking on A. bivaginalis from Venezuela) and ascribe them to A. bivaginalis. However, hosts are different, and localities are widely separated, so it is likely that future study will show that the species from the Mediterranean is a distinct, new, species.

Towards the resolution of the Microcotyle erythrini species complex: description of Microcotyle isyebi n. sp. (Monogenea, Microcotylidae) from Boops boops (Teleostei, Sparidae) off the Algerian coast.

The monogenean Microcotyle erythrini is atypical because it has been recorded from several fish host species in the Mediterranean Sea and Atlantic Ocean, in contrast to many species which are considered strictly specific. This could indicate a true lack of specificity or that several cryptic species are involved. This paper is a partial attempt to solve this problem. Specimens of a monogenean resembling M. erythrini were collected from bogues, Boops boops, caught off Algeria. A comparison with published descriptions and with museum specimens of M. erythrini did not yield any clear morphological difference. However, sequences of cytochrome c oxidase subunit I (COI) differed by 16.3% from that of M. erythrini (from GenBank, material collected from the type-host Pagellus erythrinus), indicating that the species was different. The species from B. boops is therefore described here as Microcotyle isyebi n. sp. and differential diagnoses with Microcotyle species from the Mediterranean and from sparids are provided. These results suggest that a molecular re-evaluation of other M. erythrini-like specimens from various fish hosts could reveal the existence of additional parasite biodiversity.

Microcotyle visa n. sp. (Monogenea: Microcotylidae), a gill parasite of Pagrus caeruleostictus (Valenciennes) (Teleostei: Sparidae) off the Algerian coast, Western Mediterranean.

Parasite biodiversity of fish of the southern part of the Mediterranean sea is still incompletely explored. We describe here Microcotyle visa n. sp. from the gill filaments of the bluespotted seabream Pagrus caeruleostictus (Valenciennes) (Sparidae) collected off the Algerian coast. The identity of fish hosts was confirmed by barcoding. Microcotyle visa n. sp. is herein described and illustrated. Analysis of the cox1 gene of the monogeneans revealed minor intraspecific variation (1.4%), an order of magnitude lower than the distance between this species and other Microcotyle species (10-15 %). Microcotyle visa n. sp. is distinguished from Microcotyle erythrini van Beneden & Hesse, 1863, a congener infesting sparids, on the basis of morphological (size of clamps, number of testes) and molecular (cox1) differences. This is the fourth member of the genus known to parasitise a sparid host. A species of Paramicrocotyle sp. included in the molecular analysis was nested within a robust Microcotyle + Paramicrocotyle clade; in the absence of demonstrated molecular and morphological differences, we consider that Paramicrocotyle Caballero & Bravo-Hollis, 1972 is a junior synonym of Microcotyle van Beneden & Hesse, 1863 and transfer two species of Paramicrocotyle as Microcotyle danielcarrioni (Martinez & Barrantes, 1977) n. comb. and Microcotyle moyanoi (Villalba & Fernandes, 1986) n. comb.