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

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

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Ponovni opis nametnika Axine belones Abildgaard, 1794 (Monogenea, Axinidae), pronađenog na škrgama iglice Belone belone (Linnaeus, 1760) (Teleostei, Belonidae) uz alžirsku obalu u zapadnom Sredozemnom moru

Axine belones Abildgaard, 1794 is a polyopisthocotylean monogenean, parasite of belonid fishes, and there are few accounts with morphological and morphometrical data. Here, we redescribe A. belones based on newly collected specimens from its type-host, the garfish Belone belone (Linnaeus, 1760) collected off Algeria, Western Mediterranean, a new locality for this species. Several anatomical and morphological features (genital apertures and clamps sclerites) are described and illustrated. We provide morpho-anatomical and morphometrical data, along with detailed illustrations of A. belones, and we discuss the host specificity of A. belones. Notes on hosts and localities of this species and other congeners are presented, and host specificity patterns of A. belones revealed stenoxenic specificity to Belonidae Bonaparte, 1835, whilst the genus Axine Abildgaard, 1794 seems to be restricted to fishes of the order Beloniformes. Our attempt to investigate morphometric variations between Mediterranean and oceanic specimens revealed that the two populations differed by the number of testes, body length, and clamp dimensions. However, the limited number of measured organs in the various accounts precluded any distinction between the populations. Molecular data for both Mediterranean and oceanic specimens are needed to determine the existence of cryptic species.Axine belones Abildgaard, 1794 je jednorodni metilj, nametnik na škrgama iglice, o kojem postoji malo radova s morfološkim i morfometrijskim podacima. U ovom radu dajemo ponovni opis vrste A. belones na temelju primjeraka sakupljenih na iglici Belone belone (Linnaeus, 1760), njegovog tipskog domaćina. Primjerci iglica su ulovljeni uz obale Alžira, u zapadnom Sredozemnom moru, što je ujedno i novi lokalitet za ovu vrstu nametnika. Opisano je i ilustrirano nekoliko anatomskih i morfoloških značajki (genitalni otvori i skleriti prijanjaljki). U radu se iznose morfo-anatomski i morfometrijski podatci, zajedno s detaljnim ilustracijama A. belones i raspravlja o specifičnosti domaćina A. belones. Prikazani su i zabilježeni domaćini i lokaliteti nalaza ove vrste i srodnika, a obrasci specifičnosti domaćina kod A. belones ukazali su na stenokseničnu specifičnost prema porodici Belonidae Bonaparte, 1835, dok je rod Axine Abildgaard, 1794 vjerojatno ograničen na ribe reda Beloniformes. Naš pokušaj da istražimo morfometrijske varijacije između sredozemnih i oceanskih jedinki otkrio je da se dvije populacije razlikuju po broju testisa, duljini tijela i dimenzijama prijanjaljki. Međutim, ograničeni broj izmjerenih organa u drugim literaturnim izvorima onemogućio je precizno utvrđivanje razlike između populacija. Postojanje kriptičnih vrsta bi se moglo dokazati korištenjem molekularnih metoda za sredozemne i oceanske uzorke

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

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

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”

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

“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 Plectanocotyle gurnardi 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. 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 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 Plectanocotyle 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

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 hyperparasiteCyclocotyla 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

No vagina, one vagina, or multiple vaginae? An integrative study of

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