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Bithynids snails are a widespread group of molluscs in European freshwater systems. However, not much information is available on trematode communities from molluscs of this family. Here, we investigate the trematode diversity of Bithynia tentaculata, based on molecular and morphological data. A total of 682 snails from the rivers Lippe and Rhine in North Rhine-Westphalia, Germany, and 121 B. tentaculata from Curonian Lagoon, Lithuania were screened for infections with digeneans. In total, B. tentaculata showed a trematode prevalence of 12.9% and 14%, respectively. The phylogenetic analyses based on 55 novel sequences for 36 isolates demonstrated a high diversity of digeneans. Analyses of the molecular and morphological data revealed a species-rich trematode fauna, comprising 20 species, belonging to ten families. Interestingly, the larval trematode community of B. tentaculata shows little overlap with the well-studied trematode fauna of lymnaeids and planorbids, and some of the detected species (Echinochasmus beleocephalus and E. coaxatus) constitute first records for B. tentaculata in Central Europe. Our study revealed an abundant, diverse and distinct trematode fauna in B. tentaculata, which highlights the need for further research on this so far understudied host–parasite system. Therefore, we might currently be underestimating the ecological roles of several parasite communities of non-pulmonate snail host families in European fresh waters. [ABSTRACT FROM AUTHOR]

Freshwater ecosystems in Lithuania are threatened by the introduction of invasive fish species including Neogobius fluviatilis , N. melanostomus , Perccottus glenii , and Pseudorasbora parva . Data on helminth parasites of these fishes have not been comprehensively studied, with only two reports on parasites of N. melanostomus from the Curonian Lagoon and Baltic Sea, Lithuania. We examined 278 fish individuals representing 4 invasive species from 13 waterbodies. Using morphological and molecular analyses, we identified 29 helminth taxa representing 15 digenean trematodes, 6 nematodes, 4 cestodes, 2 acanthocephalans, and 2 monogeneans. With 18 species, N. fluviatilis had the highest helminth diversity, followed by N. melanostomus (11 species) and Ps. parva (8 species). Perccottus glenii was found to be free from helminth infection. The availability of historical information on the native fish parasites in Lithuania allowed us to conclude that out of the 29 recorded species, invasive fish serve as hosts for 22 local fish helminth species, while 7 helminth species have been reported exclusively in invasive fish. Based on newly obtained and previously published data, a total of 34 helminth species parasitise invasive fish in Lithuania, of which 30 use these fish as intermediate or paratenic hosts.

Trematodes and their snail hosts have developed intimate parasite-host associations, with snails supporting a diverse and often species-specific trematode fauna. In the faucet snail, Bithynia tentaculata (Caenogastropoda, Littorinimorpha), a unique trematode fauna has been recorded recently. However, knowledge of the exact species identity, phylogenetic relationships, and geographical distribution remains limited as many of the species belong to groups with unclear or controversial taxonomical assignment. To contribute to our knowledge of the trematodes, we investigated the trematode fauna of B. tentaculata by examining a total of 556 snails from lakes in County Galway, Ireland. Using an integrative taxonomic approach including DNA sequence data analyses (28S rRNA gene, ITS1-5.8S-ITS2, ITS2, cox 1, nad 1) and morphological tools (taxonomical drawings and measurements), we identified nine trematode species of seven families, with seven species occurring as cercariae ( Cyathocotyle prussica , Lecithodendrium linstowi , Lecithodendrium sp., Asymphylodora progenetica , Sphaerostoma bramae , Metorchis xanthosomus , and Notocotylus sp.) and three species occurring as metacercariae ( A. progenetica , Parasymphylodora parasquamosa , and Sphaeridiotrema sp.). Except for S. bramae , all are new species records for Ireland and provide the most western distribution of these trematodes in Europe. The trematode species recorded are known to use a wide range of definitive hosts and have a wide geographical distribution; among them are species members of genera that are zoonotic ( Metorchis ) and pathogenic to wildlife ( Cyathocotyle , Sphaeridiotrema , and Notocotylus ). There remains an ongoing need for precise identification of the trematode species to ensure that wider ecological contexts are correctly understood and biodiversity and disease threats can be accurately evaluated.

Larval stages of the widely distributed digenean species Proctoeces maculatus (Looss, 1901) were reported 40 years ago from South Africa in the common octopus, Octopus vulgaris Cuvier (Octopodidae). However, the absence of adult specimens and molecular data from this region has hindered a comprehensive understanding of its distribution. In this study, we collected three species of intertidal and near-shore marine fishes [ Clinus superciliosus (L.) (Clinidae), Diplodus capensis (Smith) (Sparidae) and Sparodon durbanensis (Castelnau) (Sparidae)] along the South African coast and discovered adult specimens of P . maculatus at five localities. By employing a combination of morphological and molecular techniques, including 28S rDNA, 18S rDNA and COI mtDNA analyses, the first report of adult P . maculatus from South Africa is presented. The findings encompass a comprehensive morphological description and molecular data, illuminating the true distribution of this species in the region.

Tapeworms of the genus Ophiotaenia La Rue, 1911 parasitize herptiles (= amphibians and 'reptiles') throughout the world, with about 100 species recognised as valid. In the present work, the North American species found in watersnakes (Colubridae) are reviewed. An examination of the holotype of Ophiotaenia perspicua La Rue, 1911, the type species of the genus, and other specimens from Nerodia rhombifer (Hallowell) revealed that two species were used for the species description. The 'true' O. perspicua has a small scolex and small, round suckers. This species is redescribed based on new material from Oklahoma, USA. The other species from N. rhombifer , Ophiotaenia laruei n. sp., has a larger scolex and larger, almost triangular suckers. Examination of the types of O. variabilis (Brooks, 1978) from N. rhombifer and N. cyclopion (Duméril, Bibron et Duméril) from Louisiana, USA has revealed that it is a mixture of two or more species. Because of poor quality of these specimens, it is not possible to adequately characterise O. variabilis , which is considered a species inquirenda . In addition, two new species are described from Nerodia fasciata confluens (Blanchard). Ophiotaenia currani n. sp. from Mississippi, USA is characterised by elongate, narrow proglottids, few testes, and a relatively long cirrus sac. Ophiotaenia tkachi n. sp. from Louisiana, USA is characterised by relatively short and wide proglottids, more testes, and an unusual terminal part of the vagina with folds. Morphologically similar tapeworms of N. fasciata confluens , N. erythrogaster (Förster), N. sipedon (Linnaeus), and Agkistrodon piscivorus (Lacépède) (Viperidae) from Arkansas and Oklahoma, USA, which are genetically nearly identical, are considered to be conspecific with O. tkachi n. sp. The present data suggest a high, previously undescribed species diversity of proteocephalid tapeworms in watersnakes in North America, and generally strict host specificity of these tapeworms., Competing Interests: There is no conflict of interest in the submitted manuscript entitled “Discovering high species diversity of Ophiotaenia tapeworms (Cestoda: Proteocephalidae) of watersnakes (Colubridae) in North America”., (© 2023 The Authors.)

The genus Coitocaecum Nicoll, 1915 is part of the most speciose digenean family, the Opecoelidae Ozaki, 1925, which is found globally in both freshwater and marine fishes. Fifteen opecoelid species have been reported from marine fishes in South Africa, yet only one species of Coitocaecum has been described from this region: Coitocaecum capense Bray, 1987. During an explorative study of the digeneans of the endemic, intertidal fish Clinus superciliosus (Linnaeus) from the Saldanha Bay area, Cape Town harbour, Hermanus, the Tsitsikamma section of the Garden Route National Park and Chintsa East in South Africa, a total of three distinct species of Coitocaecum were identified based on morphological and molecular (28S rDNA, ITS1-5.8S-ITS2 rDNA and COI mtDNA) data: the previously mentioned C. capense, Coitocaecum brayi sp. n. and a third, unnamed species. We provide the first molecular characterisation of species of Coitocaecum from South Africa, accompanied by detailed morphological descriptions. This study illustrates the importance of an integrated taxonomic approach, especially when studying species with similar morphology. These findings further emphasise the lack of information on the true diversity and molecular data for trematodes of marine fishes in South Africa, creating a great capacity for future explorative taxonomic studies and highlighting the use of intertidal areas for conducting such research.

Proteocephalid tapeworms of frogs of the family Ranidae ('true' frogs) are reviewed with emphasis on their species diversity, host specificity and geographical distribution. New molecular data (nuclear lsrDNA and mitochondrial COI sequences) are presented for tapeworms of four species of ranid frogs in North America, including the poorly known Ophiotaenia saphena Osler, 1931 of Rana clamitans Latreille and R. catesbeiana (Shaw), which is redescribed using new material from Arkansas, USA. Tapeworms of R. sphenocephala (Cope) and R. pipiens Schreber, the latter previously identified as O. saphena, represent another, putative new species, but are not formally described due to insufficient available material. Proteocephalus papuensis Bursey, Goldberg et Kraus, 2008 from Sylvirana supragrisea (Menzies) is transferred to Ophiotaenia La Rue, 1911 as a new combination. After a critical review of the literature, only nine nominal species of Ophiotaenia are recognised as valid, which is in contrast to the large number of ranid frogs (> 440 spp.). The reasons for this striking disparity are briefly discussed, and a key based on morphology is presented for the identification of all species of Ophiotaenia from the Ranidae. Molecular data are available for only two taxa from North America that form a monophyletic group. The relationships among tapeworms of ranid frogs occurring in other zoogeographical regions are not yet known. The taxonomic status of Batrachotaenia Rudin, 1917, which was erected to accommodate proteocephalids from amphibians, is also discussed. To facilitate future studies, a tabulated summary of all 32 species of proteocephalids belonging to three genera reported from amphibians (frogs and salamanders) is presented, with information on their hosts, distribution, and taxonomically important characters, including key measurements.

Cercariae, the free-living larval stages of trematodes, have adopted an amazing variety of transmission strategies. One of them is prey-mimetism, i.e. cercariae mimicking prey to attract motile hosts to be eaten. In a period between 2002 and 2019, we examined small planorbid snails, Bathyomphalus contortus, Gyraulus parvus and Planorbis planorbis from lakes in Finland and Iceland and from the Curonian Lagoon in Lithuania. Cercariae with conspicuously enlarged tails and unusual swimming behaviour, likely mimicking invertebrate prey, were detected and studied by the use of morphological and molecular (cox1, ITS1-5.8S-ITS2 and 28S rDNA) methods. Cercariae of two species belonging to the genus Apatemon (Strigeidae) were recognised. We consider Apatemon sp. 5 ex P. planorbis from the Curonian Lagoon identical to Cercaria globocaudata U. Szidat, 1940. Cercariae ex G. parvus from Iceland and ex B. contortus from Finland were conspecific, and we named them Apatemon sp. 6; these cercariae could not be associated with any known species. For the first time, we verified that cercariae of the Bulbocauda group belong to the genus Apatemon. We provide a mini-review on records of furcocercariae of the family Strigeidae with enlarged tails reported in freshwaters of the northern hemisphere and reveal that it is not only Apatemon but also Australapatemon and most likely Strigea which belong to the Bulbocauda group, rendering it a purely ecological assemblage. Understanding which invertebrate swimming behaviour these cercariae are mimicking will enhance our knowledge of the processes behind trematode transmission and will help to assess evolutionary pathways of host-finding strategies in trematodes., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Brazil is a tropical country with remarkably diverse marine habitats that harbour a rich diversity of fish. Only a small portion of this fish diversity has been investigated for parasites, and thus the diversity of their trematode parasites remains unexplored. Moreover, only 5 out of 184 known digenean trematode species of marine fish in Brazil have been genetically characterised. The Hemiuridae Looss, 1899 is the second most speciose trematode family in marine fishes from Brazil but, in many ways, it remains a neglected group. Forty-three trematode specimens from nine fish species were collected from the coastal zone off Rio de Janeiro, Brazil. Trematodes were found in the stomach of 14 specimens of 9 fish species belonging to 8 families (Carangidae, Clupeidae, Haemulidae, Muraenidae, Percophidae, Pinguipedidae, Trichiuridae, and Triglidae). Trematode specimens were studied using morphological and molecular genetic analyses. A total of eight hemiurid species from four genera, Ectenurus , Lecithochirium , Myosaccium , and Parahemiurus were identified. This paper reports on new host records for four species of hemiurids, adds a new record on the geographical distribution for one species, and provides the first DNA sequence data supplemented with the detailed description of morphology for five species. Phylogenetic analyses supported that the subfamily classifications of the Hemiuridae-based entirely on morphological characters-needs to be reconsidered, taking into account a wider range of information sources.

The Diplostomidae Poirier, 1886 is a large family of digeneans within the superfamily Diplostomoidea Poirier, 1886. Members of the family are distributed worldwide and parasitize a diversity of tetrapod definitive hosts. Notably, only 2 mature diplostomids are known from crocodilians and both are suggested to be accidental infections. In this study, we use morphological and molecular data to describe Neofibricola n. gen. from a Nile crocodile Crocodylus niloticus collected in South Africa. We provide a description of adults and metacercariae of the type species, Neofibricola smiti n. sp., and metacercariae of a likely congeneric species. We generated partial 28S and internal transcribed spacer region ribosomal deoxyribonucleic acid (DNA) and cytochrome c oxidase 1 subunit mitochondrial DNA for both species and utilized the newly generated 28S sequences to examine phylogenetic affinities of these new taxa. In addition, we provide a new key to diplostomid genera, considering the substantial systematic changes and newly erected genera since the previously published key to diplostomid genera., (© American Society of Parasitologists 2022.)

In 2014-2015 and 2019-2021, teleost fishes off Galindez Island (Antarctic Peninsula) were examined for trematodes. Combined morphological and molecular analyses revealed the presence of eight trematode species of four families (Hemiuridae, Lecithasteridae, Opecoelidae, Lepidapedidae) from five fish species. Only adult trematodes were found and all of them are Antarctic endemics with their congeners occurring on other continents. The hemiuroids, Elytrophalloides oatesi (Leiper & Atkinson, 1914), Genolinea bowersi (Leiper & Atkinson, 1914), and Lecithaster macrocotyle Szidat & Graefe, 1967 belong to the most common Antarctic species and together with Lepidapedon garrardi (Leiper & Atkinson, 1914) and Neolebouria georgiensis Gibson, 1976 they were recorded as the least host-specific parasites. The originally sub-Antarctic Neolepidapedon macquariensis Zdzitowiecki, 1993 is a new record for the Antarctic Peninsula and Parachaenichthys charcoti (Vaillant), is a new host record. Neolebouria terranovaensis Zdzitowiecki, Pisano & Vacchi, 1993 is considered a synonym of N. georgiensis because of identical morphology and dimensions. The currently known phylogenetic relationships within the studied families are supported, including the polyphyly of Macvicaria Gibson & Bray, 1982 with the future need to accommodate its Antarctic species in a new genus. The validity of M. georgiana (Kovaleva & Gaevskaja, 1974) and M. magellanica Laskowski, Jezewski & Zdzitowiecki, 2013 needs to be confirmed by further analyses. Genetic sequence data are still scarce from Antarctica, and more studies applying integrative taxonomic approaches and large-scale parasitological examinations of benthic invertebrates are needed to match sequences of larval stages to those of well-characterised adults and to elucidate trematode life-cycles., (© 2022. The Author(s), under exclusive licence to Springer Nature B.V.)

DNA sequence data became an integral part of species characterization and identification. Still, specimens associated with a particular DNA sequence must be identified by the use of traditional morphology-based analysis and correct linking of sequence and identification must be ensured. Only a small part of DNA sequences of the genus Diplostomum (Diplostomidae) is based on adult isolates which are essential for accurate identification. In this study, we provide species identification with an aid of morphological and molecular (cox1, ITS-5.8S-ITS2 and 28S) characterization of adults of Diplostomum baeri Dubois, 1937 from naturally infected Larus canus Linnaeus in Karelia, Russia. Furthermore, we reveal that the DNA sequences of our isolates of D. baeri are identical with those of the lineage Diplostomum sp. clade Q , while other sequences labelled as the ‘D. baeri’ complex do not represent lineages of D. baeri. Our new material of cercariae from Radix balthica (Linnaeus) in Ireland is also linked to Diplostomum sp. clade Q. We reveal that D. baeri is widely distributed in Europe; as first intermediate hosts lymnaeid snails (Radix auricularia (Linnaeus), R. balthica) are used; metacercariae occur in eye lens of cyprinid fishes. In light of the convoluted taxonomy of D. baeri and other Diplostomum spp., we extend the recommendations of Blasco-Costa et al. (2016, Systematic Parasitology 93, 295–306) for the ‘best practice’ in molecular approaches to trematode systematics. The current study is another step in elucidating the species spectrum of Diplostomum based on integrative taxonomy with well-described morphology of adults linked to sequences.

The caryophyllidean genus Archigetes Leuckart, 1878 is unique among all tapeworms in that its species can mature in invertebrate hosts (Oligochaeta), i.e., have a monoxenic (direct) life cycle. All five species were described as progenetic plerocercoids in oligochaetes and two of them also as adults from cypriniform fishes. Two species, A. sieboldi Leuckart, 1878 and A. iowensis Calentine, 1962, were found in North America in non-native common carp (Cyprinus carpio). A molecular study of caryophyllideans from the southern United States has revealed the occurrence of three new species in native freshwater fishes (Catostomidae, Ictiobinae): Archigetes loculotruncatus n. sp. from Ictiobus bubalus, I. niger and Carpiodes cyprinus is the largest representative of the genus and differs by a loculotruncate scolex. Archigetes megacephalus n. sp. from Ictiobus niger, I. bubalus and I. cyprinellus is characterised by a prominent, bothrioloculodiscate scolex. Archigetes vadosus n. sp. from I. bubalus is typified by a globular scolex with very shallow loculi; it differs from the closely related A. sieboldi in the shape of the body, with a distinct neck region and a scolex wider than the remaining body. Archigetes iowensis Calentine, 1962 becomes a junior synonym of Paraglaridacris limnodrili (Yamaguti, 1934). The generic diagnosis of Archigetes is amended and a key to identification of North American taxa is provided. Species of Archigetes and Paraglaridacris differ from each other most conspicuously in the structure of the ovary, which is follicular in Archigetes versus compact in Paraglaridacris., (© D. Uhrovič et al., published by EDP Sciences, 2022.)

Although some parasitological efforts have focused on the frigate tuna Auxis thazard (Lacepède) (Scombriformes, Scombridae) in Brazil, its digenean fauna remains poorly known. Combining morphological and molecular methods, we investigated the diversity of digenean trematodes of A. thazard collected from the coastal waters off the state of Rio de Janeiro, Brazil in 2021. Six species belonging to four families were recorded: the bucephalid Rhipidocotyle cf. angusticolle Chandler, 1941, the didymozoid Didymocystis sp. 6 sensu Louvard et al. (2022), the fellodistomid Tergestia sp., and three hemiurids, Dinurus euthynni Yamaguti, 1934, Lecithochirium floridense (Manter, 1934), and L. synodi Manter, 1931. The current study brings the total number of digenean trematode species parasitising A. thazard in Brazil up to eight, with hemiuroid trematodes being most diverse. Auxis thazard is a new host record for L. floridense, L. synodi and potentially for R. angusticolle. The geographic distribution of several species found in our study appeared to be wider than previously known. Our study is the first to apply a DNA-based approach to digenean diversity in marine fishes in Brazil and we believe that both morphological descriptions and molecular sequence data provided in our study will aid future research assessing the diversity of digenean trematodes of A. thazard and other marine fishes in Brazil., (© C. Pantoja et al., published by EDP Sciences, 2022.)

Monozoic tapeworms (Caryophyllidea) are dominant components of parasite communities of suckers (Catostomidae) in North America, with Biacetabulum Hunter, 1927 representing one of the more species-rich genera. Molecular (28S rDNA) and morphological (including scanning electron microscopy and histology) evaluation of newly collected tapeworms from different fish hosts revealed the existence of four similar (and three closely related) species of Biacetabulum. These four species differ from their congeners by having a long body (up to 48 mm long) with a very long, slender neck (its length represents ≥30% of total body length), a large, globular scolex with a prominent central acetabulum-like loculus on the dorsal and ventral sides, two pairs of shallow lateral loculi and a distinct, slightly convex apical disc, and a cirrus-sac that is situated between the anterior arms of the ovarian wings. Taken together, the morphological and molecular data and the host associations of these species provide evidence of their host specificity. Biacetabulum isaureae n. sp. occurs in notch clip redhorse, Moxostoma collapsum, in South Carolina (USA), B. longicollum n. sp. in silver redhorse, Moxostoma anisurum (type host), and golden redhorse, M. erythrurum, in Manitoba (Canada) and West Virginia (USA), B. overstreeti n. sp. in a spotted sucker, Minytrema melanops, in Mississippi, and B. hypentelii n. sp. in northern hogsucker, Hypentelium nigricans, in Tennessee (USA). The new species differ from each other in the number of postovarian vitelline follicles, the posterior extent of preovarian vitelline follicles and relative size of the cirrus sac.

A new genus, Megancestus n. gen., is proposed to accommodate the caryophyllidean tapeworm Biacetabulum carpiodi Mackiewicz, 1969 from carpsuckers and quillback (Carpiodes spp.) in North America. This species is not closely related to other species of Biacetabulum Hunter, 1927 and is transferred to a newly erected genus. This new genus is typified by the possession of a small body (total length of 3.1-7.5 mm) with a scolex that bears a pair of large acetabulum-like loculi, two pairs of shallow lateral loculi, and a slightly convex apical disc, testes arranged in one or two layers, oval, thick-walled cirrus-sac, well-developed external seminal vesicle, separate gonopores, H-shaped ovary, few median vitelline follicles, and the uterus extending by a single loop anterior to the cirrus-sac. Megancestus differs from all Nearctic caryophyllidean genera (family Capingentidae), including Biacetabulum, by vitelline follicles dorsal to the ovary that connect the preovarian and postovarian vitelline fields. The most closely related Hunterella Mackiewicz et McCrae, 1962 differs by shape of the scolex (tholate, i.e., devoid of any loculi), dumbbell-shaped ovary and the uterus not extending anterior to the cirrus-sac. Megancestus carpiodi (Mackiewicz, 1969) n. comb. is the only species of the genus and it is a stenoxenous parasite, which has been found only in the river carpsucker (Carpiodes carpio-type host), quillback (Carpiodes cyprinus) and highfin carpsucker (Carpiodes velifer) (Catostomidae: Ictiobinae) in the lower and middle Mississippi basin.

South African clinids are a major component of the temperate intertidal regions that are also known to participate in life cycles and transmission of several groups of parasites. However, the knowledge of trematode diversity of these fishes is incomplete. In this study, two species of Clinus Cuvier, the super klipfish Clinus superciliosus (Linnaeus) and the bluntnose klipfish Clinus cottoides Valenciennes, were collected from six localities along the South African coast and examined for the presence of trematodes. Metacercariae of Cardiocephaloides Sudarikov, 1959 were found in the eye vitreous humour and brain of C. superciliosus and in the eye vitreous humour of C. cottoides. Detailed analyses integrating morphological and molecular sequence data (28S rDNA, ITS2 rDNA-region, and COI mtDNA) revealed that these belong to two species, Cardiocephaloides physalis (Lutz, 1926) and an unknown species of Cardiocephaloides. This study provides the first report of clinid fishes serving as intermediate hosts for trematodes, reveals that the diversity of Cardiocephaloides in South Africa is higher than previously recorded, and highlights the need for further research to elucidate the life cycles of these trematode species. The broad geographical distribution of Cardiocephaloides spp. was confirmed in the present study based on molecular sequence data. The host-parasite interactions between clinid fishes and metacercariae of Cardiocephaloides are yet to be explored.

The biodiversity of freshwater ecosystems globally still leaves much to be discovered, not least in the trematode parasite fauna they support. Echinostome trematode parasites have complex, multiple-host life-cycles, often involving migratory bird definitive hosts, thus leading to widespread distributions. Here, we examined the echinostome diversity in freshwater ecosystems at high latitude locations in Iceland, Finland, Ireland and Alaska (USA). We report 14 echinostome species identified morphologically and molecularly from analyses of nad1 and 28S rDNA sequence data. We found echinostomes parasitising snails of 11 species from the families Lymnaeidae, Planorbidae, Physidae and Valvatidae. The number of echinostome species in different hosts did not vary greatly and ranged from one to three species. Of these 14 trematode species, we discovered four species (Echinoparyphium sp. 1, Echinoparyphium sp. 2, Neopetasiger sp. 5, and Echinostomatidae gen. sp.) as novel in Europe; we provide descriptions for the newly recorded species and those not previously associated with DNA sequences. Two species from Iceland (Neopetasiger islandicus and Echinoparyphium sp. 2) were recorded in both Iceland and North America. All species found in Ireland are new records for this country. Via an integrative taxonomic approach taken, both morphological and molecular data are provided for comparison with future studies to elucidate many of the unknown parasite life cycles and transmission routes. Our reports of species distributions spanning Europe and North America highlight the need for parasite biodiversity assessments across large geographical areas., (© C. Pantoja et al., published by EDP Sciences, 2021.)

Reliable data on the diversity of the genus Diplostomum (Digenea: Diplostomidae) parasitising freshwater fishes in South Africa, as well as in Africa, is almost non-existent. Most of the morphology-based identifications of species within this genus reported from Africa require critical revision. The aim of the present study was to determine the diversity of Diplostomum metacercariae in South African fishes applying molecular and traditional morphological techniques. To achieve this aim, a total of 216 fishes belonging to 21 species collected in the Rivers Phongolo, Riet, Usuthu and Mooi in three provinces of South Africa were examined. Metacercariae of Diplostomum were recovered from the eye lenses of 38 fishes belonging to five species of the families Anguillidae, Cichilidae and Mochokidae, with an overall low prevalence of infection (18%). Metacercariae were subjected to morphological study and molecular sequencing of the partial mithochondrial cox 1 and ribosomal 28S rDNA genes as well as of ribosomal ITS1-5.8S-ITS2 region. Morphological and phylogenetic analyses revealed the presence of three species which matched those previously reported from Nigeria, Iraq and China, therefore those from Tilapia sparrmanii and Synodontis zambezensis were named Diplostomum sp.; those from Anguilla labiata , Oreochromis mossambicus and S . zambezensis were named Diplostomum sp. 14; and those from Pseudocrenilabrus philander were named Diplostomum sp. 16. Geographic distribution of several species of Diplostomum appeared to be wider than expected. Morphological description and novel sequence data generated during this study will contribute to the elucidation of the life cycles of Diplostomum sp., Diplostomum sp. 14 and Diplostomum sp. 16 and advance further research of diplostomids in Africa., Competing Interests: The authors declare no conflicts of interest., (© 2019 The Authors.)

Four species of previously known nematodes from the family Camallanidae were found from different hosts in South Africa: Batrachocamallanus xenopodis from the frog Xenopus muelleri , Paracamallanus cyathopharynx and Procamallanus pseudolaeviconchus from the catfish Clarias gariepinus and Spirocamallanus daleneae from the catfish Synodontis zambezensis . In the material collected from various marine fishes, several specimens of nematodes from the genus Camallanus clearly differed from all previously known species. Based on morphological differences these specimens are assigned to a new species, C. sodwanaensis . Molecular data of 18S and 28S rDNA and COI sequences are provided for the collected species and a phylogenetic analyses based on 28S gene fragmets are presented., Competing Interests: No conflict of interest., (© 2019 The Authors.)

Three new species of acanthocephalans are described from marine fishes collected in Sodwana Bay, South Africa: Rhadinorhynchus gerberi n. sp. from Trachinotus botla (Shaw), Pararhadinorhynchus sodwanensis n. sp. from Pomadasys furcatus (Bloch et Schneider) and Transvena pichelinae n. sp. from Thalassoma purpureum (Forsskål). Transvena pichelinae n. sp. differs from the single existing species of the genus Transvena annulospinosa Pichelin et Cribb, 2001, by the lower number of longitudinal rows of hooks (10-12 vs 12-14, respectively) and fewer hooks in a row (5 vs 6-8), shorter blades of anterior hooks (55-63 vs 98), more posterior location of the ganglion (close to the posterior margin of the proboscis receptacle vs mid-level of the proboscis receptacle) and smaller eggs (50-58 × 13 µm vs 62-66 × 13-19 µm). Pararhadinorhynchus sodwanensis n. sp. differs from all known species of the genus by a combination of characters. It closely resembles unidentified species Pararhadinorhynchus sp. sensu Weaver and Smales (2014) in the presence of a similar number of longitudinal rows of hooks on the proboscis (16-18 vs 18) and hooks in a row (11-13 vs 13-14), but differs in the position of the lemnisci (extend to the level of the posterior end of the proboscis receptacle or slightly posterior vs extend to the mid-level of the receptacle), length of the proboscis receptacle (910-1180 µm vs 1,460 µm) and cement glands (870-880 µm vs 335-350 µm). Rhadinorhynchus gerberi n. sp. is distinguishable from all its congeners by a single field of 19-26 irregular circular rows of the tegumental spines on the anterior part of the trunk, 10 longitudinal rows of hooks on the proboscis with 29-32 hooks in each row, subterminal genital pore in both sexes, and distinct separation of the opening of the genital pore from the posterior edge of the trunk (240-480 μm) in females. Sequences for the 18S rDNA, 28S rDNA and cox1 genes were generated to molecularly characterise the species and assess their phylogenetic position. This study provides the first report based on molecular evidence for the presence of species of Transvena Pichelin et Cribb, 2001 and Pararhadinorhynchus Johnston et Edmonds, 1947 in African coastal fishes.

Corynosoma obtuscens Lincicome, 1943 (Acanthocephala: Polymorphidae) is synonymised with Corynosoma australe Johnston, 1937 based on combined morphological and molecular evidence. Morphological comparison of C. obtuscens (24 males and 27 females) collected from a California sea lion Zalophus californianus (Lesson) in California, USA, with the type-specimens of C. obtuscens and C. australe, and with published data on C. australe collected from different hosts and regions showed no significant differences. The levels of genetic divergence in the cox1 sequences obtained from C. obtuscens from a California sea lion in the present study and C. australe from otariid seals from Argentina and penguins from Brazil ranged between 1.4-1.6% and was considered to represent intraspecific variability. Additionally, cox1 sequences were generated for Andracantha phalacrocoracis (Yamaguti, 1939), Corynosoma semerme (Forssell, 1904), C. strumosum (Rudolphi, 1802), C. validum Van Cleave, 1953 and C. villosum Van Cleave, 1953. Our results revealed inconsistency in the identification of material used as a source of the previously published sequence data for C. obtuscens and C. magdaleni Montreuil, 1958.

The population of California sea lion Zalophus californianus (CSL) has steadily increased during the last several decades. Despite extensive research addressing CSL biology and ecology performed during the last decades, there has been a minimal number of published papers documenting their parasite fauna. Our objective was to analyze the actual list of the metazoan parasites reported from CSLs and add new data on the age-related differences in the prevalence and biodiversity of the parasite community. There have been 33 species recorded but this study considers only 24 of them valid. Among them, 11 species are specific parasites of CSLs and 13 species are not specific. Additional species represent accidental infections or misidentifications. In total, 6653 helminths and 847 mites were collected and identified from 34 CSLs for this study. Six species of nematodes, Anisakis simplex sensu lato s . l . (prevalence 41%; intensity 7.6), Contracaecum ogmorhini s . l . (38%; 269.6), Pseudoterranova decipiens s . l . (29%; 33), P. azarazi (9%; 2.7), Acanthocheilonema odendhali (15%; 3.5) and Parafilaroides decorus were found. Two species of cestodes, Diphyllobothrium sp. (38%; 8.5) and Anophryocephalus sp. (15%; 14.6) represent novel undescribed species. Two species of trematodes, Apophallus zalophi (18%; 19.7) and Zalophotrema hepaticum (12%; 39.2), and five species of acanthocephalans, Corynosoma obtuscens (68%; 100.8), C. strumosum (53%; 4.6), Andracantha phalacrocoracis (3%; 1), Andracantha sp. (9%; 4.3) and Profilicollis altmani (6%; 8.5) were found. Mites Orthohalarchne attenuata (prevalence 85%) were found in the nasal cavity, while O. diminuata (21%) parasitized in the trachea and bronchi. The highest levels of infection with nematodes and trematodes were found in adult CSLs (3-16 years old), whereas the highest level of infection with acanthocephalans was found in young CSLs (pups and yearlings).

Background: Metacercariae of Diplostomum are important fish pathogens, but reliable data on their diversity in natural fish populations are virtually lacking. This study was conducted to explore the species diversity and host-parasite association patterns of Diplostomum spp. in a large riverine system in Europe, using molecular and morphological data., Methods: Twenty-eight species of fish of nine families were sampled in the River Danube at Nyergesújfalu in Hungary in 2012 and Štúrovo in Slovakia in 2015. Isolates of Diplostomum spp. were characterised morphologically and molecularly. Partial sequences of the 'barcode' region of the cytochrome c oxidase subunit 1 (cox1) and complete sequences of the nicotinamide adenine dinucleotide dehydrogenase subunit 3 (nad3) mitochondrial genes were amplified for 76 and 30 isolates, respectively. The partial cox1 sequences were used for molecular identification of the isolates and an assessment of haplotype diversity and possible host-associated structuring of the most prevalent parasite species. New primers were designed for amplification of the mitochondrial nad3 gene., Results: Only lens-infecting Diplostomum spp. were recovered in 16 fish species of five families. Barcoding of representative isolates provided molecular identification for three species/species-level genetic lineages, D. spathaceum, D. pseudospathaceum and 'D. mergi Lineage 2', and three single isolates potentially representing distinct species. Molecular data helped to elucidate partially the life-cycle of 'D. mergi Lineage 2'. Many of the haplotypes of D. spathaceum (16 in total), D. pseudospathaceum (15 in total) and 'D. mergi Lineage 2' (7 in total) were shared by a number of fish hosts and there was no indication of genetic structuring associated with the second intermediate host. The most frequent Diplostomum spp. exhibited a low host-specificity, predominantly infecting a wide range of cyprinid fishes, but also species of distant fish families such as the Acipenseridae, Lotidae, Percidae and Siluridae. The nad3 gene exhibited distinctly higher levels of interspecific divergence in comparison with the cox1 gene., Conclusions: This first exploration of the species diversity and host ranges of Diplostomum spp., in natural fish populations in the River Danube, provided novel molecular, morphological and host-use data which will advance further ecological studies on the distribution and host ranges of these important fish parasites in Europe. Our results also indicate that the nad3 gene is a good candidate marker for multi-gene approaches to systematic estimates within the genus.

Metacercariae of two species of Posthodiplostomum Dubois, 1936 (Digenea: Diplostomidae) were subjected to morphological and molecular studies: P. brevicaudatum (von Nordmann, 1832) from Gasterosteus aculeatus (L.) (Gasterosteiformes: Gasterosteidae), Bulgaria (morphology, cox1 and ITS1-5.8S-ITS2) and Perca fluviatilis L. (Perciformes: Percidae), Czech Republic (morphology, cox1, ITS1-5.8S-ITS2 and 28S); and P. centrarchi Hoffman, 1958 from Lepomis gibbosus (L.) (Perciformes: Centrarchidae), Bulgaria (morphology, cox1 and ITS1-5.8S-ITS2) and Slovakia (cox1 and ITS1-5.8S-ITS2). In addition, cercariae of P. cuticola (von Nordmann, 1832) from Planorbis planorbis (L.) (Mollusca: Planorbidae), Lithuania (morphology and cox1) and metacercariae of Ornithodiplostomum scardinii (Schulman in Dubinin, 1952) from Scardinius erythrophthalmus (L.) (Cypriniformes: Cyprinidae), Czech Republic, were examined (morphology, cox1, ITS1-5.8S-ITS2 and 28S). These represent the first molecular data for species of Posthodiplostomum and Ornithodiplostomum Dubois, 1936 from the Palaearctic. Phylogenetic analyses based on cox1 and ITS1-5.8S-ITS2, using O. scardinii as the outgroup and including the three newly-sequenced Posthodiplostomum spp. from Europe and eight published unidentified (presumably species-level) lineages of Posthodiplostomum from Canada confirmed the distinct status of the three European species (contrary to the generally accepted opinion that only P. brevicaudatum and P. cuticola occur in the Palaearctic). The subspecies Posthodiplostomum minimum centrarchi Hoffmann, 1958, originally described from North America, is elevated to the species level as Posthodiplostomum centrarchi Hoffman, 1958. The undescribed "Posthodiplostomum sp. 3" of Locke et al. (2010) from centrarchid fishes in Canada has identical sequences with the European isolates of P. centrarchi and is recognised as belonging to the same species. The latter parasite, occurring in the alien pumpkinseed sunfish Lepomis gibbosus in Europe, is also supposed to be alien for this continent. It is speculated that it colonised Europe long ago and is currently widespread (recorded in Bulgaria, Slovakia and Spain); based on the cox1 sequence of an adult digenean isolate from the Ebro Delta, Spain, only the grey heron (Ardea cinerea L.) (Ciconiiformes: Ardeidae) is known to be its definitive host in Europe.

Three new psilostomid genera, Byrdtrema n. g., Longisaccus n. g. and Macracetabulum n. g., each with a single species, are described from ducks, Aix sponsa (L.) and Bucephala albeola (L.) in North America. Byrdtrema n. g. and Macracetabulum n. g. possess a bipartite seminal vesicle and share this character with four psilostomid genera, Grysoma Byrd, Bogitsh & Maples, 1961, Neopsilotrema Kudlai, Pulis, Kostadinova & Tkach, 2016, Psilostomum Looss, 1899 and Psilotornus Byrd & Prestwood, 1969. Byrdtrema n. g. differs from Macracetabulum n. g. in the shape of the body (elongate vs elongate-oval); the position of the ventral sucker (in first third of body vs just pre-equatorial); the shorter forebody; as well as in the smaller size of the eggs in relation to body length. Both new genera differ from (i) Grysoma by the nature of the vitellarium (large, compact follicles with small vitelline cells vs weakly defined follicles with large vitelline cells, respectively) and the smaller size of the eggs in relation to body length; (ii) Psilostomum in the posterior extend of the cirrus-sac in relation to ventral sucker (slightly posterior vs more posterior), the location of the genital pore (at the level of oesophagus vs just postbifurcal), the shorter length of uterine and longer post-testicular fields in relation to body length, and the anterior limits of vitellarium (at the level of ventral sucker vs posterior to ventral sucker); (iii) Psilotornus by the presence of a muscular pharynx (vs absent or rudimentary) and the location of the cirrus-sac (antero-dorsal to ventral sucker or more posterior vs entirely anterior to ventral sucker) and ovary (in hindbody vs in forebody). Byrdtrema n. g. differs from Neopsilotrema in the shape of the body (elongate vs subspherical to elongate-oval) and ventral sucker (elongate-oval vs subspherical to transversely oval), the shorter forebody and smaller eggs in relation to body length. Macracetabulum n. g. differs from Neopsilotrema by the shape of the ventral sucker (elongate-oval vs subspherical to transversely oval), the anterior limits of vitellarium (level of middle of ventral sucker vs level of intestinal bifurcation or anterior testis); and the slightly smaller size of eggs in relation to body length. Among the psilostomid genera, Longisaccus n. g. shows close affinities to Psilochasmus Lühe, 1909 in the presence of the long cirrus-sac and tubular internal seminal vesicle but can be clearly distinguished from the latter by the absence of the retractile tail-like process. In combination with molecular data, the above differences justify the recognition of three new genera. A key to the genera of the Psilostomidae is provided.

A new species of microphallid, Longiductotrema tethepae n. sp., is reported from a muraenid eel Gymnothorax pseudothyrsoideus (Bleeker) on the northern Great Barrier Reef. The new species is described based on adults from Gy. pseudothyrsoideus and metacercariae from a grapsid crab, Grapsus albolineatus Latreille in Milbert, collected from off Lizard Island, Queensland, Australia. The new species is assigned to Longiductotrema Deblock & Heard, 1969 based on morphological characters (presence of a cirrus-sac; a long metraterm, intensively ensheathed by gland-cells; an entirely postcaecal uterus; vitellarium composed of two lateral clusters each of about ten follicles, situated in the testicular and post-testicular areas). Longiductotrema tethepae n. sp. is the third species assigned to the genus, differing from its congeners in having a distinctly larger body dimensions, a smaller pharynx in relation to oral sucker, the anterior limits of the vitelline fields at the level of the testes (vs at the level of the ovary) and in its parasitism of a muraenid fish (vs birds). Additionally, the new species differs from L. floridense Deblock & Heard, 1969 in having a shorter metraterm and from L. scandolensis Deblock & Bartoli, 1988 in having a less elongate forebody in relation to body length, shorter caeca and prepharynx, and slightly larger eggs. Phylogenetic analyses, based on partial 28S rRNA gene sequences, showed that the present species is sister to all other microphallids for which sequence data are available. This is the fourth report of a microphallid from a marine eel, the first from the Muraenidae Rafinesque and the first from a marine fish in the Indo-west Pacific. A summary of all species of the Microphallidae parasitising fish is provided.

Neopsilotrema n. g. (Digenea: Psilostomidae) and three new species of psilostomid digeneans are described from birds in North America and Europe: Neopsilotrema lakotae n. sp. from Aythya americana (Eyton) in North Dakota, USA, Neopsilotrema affine n. sp. from Aythya affinis (Eyton) in Minnesota, USA and Neopsilotrema lisitsynae n. sp. from Anas crecca L. in Kherson Region, Ukraine. Neopsilotrema n. g. shares a bipartite seminal vesicle with only three genera within the Psilostomidae, Psilotornus Byrd & Prestwood, 1969, Psilostomum Looss, 1899 and Grysoma Byrd, Bogitsh & Maples, 1961. The new genus differs from Psilotornus in the presence of a muscular pharynx and a massive ventral sucker; the location of the cirrus-sac in relation to the ventral sucker and more posterior location of ovary; the nature of the vitellarium (i.e. comprising large, compact follicles with small vitelline cells vs weakly defined follicles with large vitelline cells); a proportionately shorter forebody; and in parasitisation in anseriform (vs passeriform) birds. Differences between the new genus and Psilostomum comprise the shape of the body, the relative size of the suckers, somewhat longer forebody and a more anterior location of the testes. Neopsilotrema n. g. differs from Grysoma in the relative size of the suckers, the degree of development of prostatic cells, the nature of the vitellarium and the size of the eggs in relation to body length. The European species Neopsilotrema lisitsynae n. sp. is distinguished from its congeners in having a longer, narrower and distinctly more elongate body with a longer post-testicular region and anterior limits of the vitelline fields posterior to ventral sucker. The two North American forms, Neopsilotrema lakotae n. sp. and Neopsilotrema affine n. sp., are cryptic species with largely overlapping metrical data; these are distinguished by comparing genetic data. The phylogenetic hypotheses for the Psilostomidae developed from sequence data analyses based on partial 28S rDNA support the erection of the new genus and the distinction of the three new species. Grysoma marilae (Price, 1942) agrees more closely with the generic diagnosis of Neopsilotrema, especially in relation to the size and shape of the body, the relative length of the forebody and post-testicular field, the structure of the vitellarium, the location of the reproductive organs and the sucker ratio. Consequently, it is here transferred to the new genus as Neopsilotrema marilae (Price, 1942) n. comb.

The Echinostomatoidea is a large, cosmopolitan group of digeneans currently including nine families and 105 genera, the vast majority parasitic, as adults, in birds with relatively few taxa parasitising mammals, reptiles and, exceptionally, fish. Despite the complex structure, diverse content and substantial species richness of the group, almost no attempt has been made to elucidate its phylogenetic relationships at the suprageneric level based on molecules due to the lack of data. Herein, we evaluate the consistency of the present morphology-based classification system of the Echinostomatoidea with the phylogenetic relationships of its members based on partial sequences of the nuclear lsrRNA gene for a broad diversity of taxa (80 species, representing eight families and 40 genera), including representatives of five subfamilies of the Echinostomatidae, which currently exhibits the most complex taxonomic structure within the superfamily. This first comprehensive phylogeny for the Echinostomatoidea challenged the current systematic framework based on comparative morphology. A morphology-based evaluation of this new molecular framework resulted in a number of systematic and nomenclatural changes consistent with the phylogenetic estimates of the generic and suprageneric boundaries and a new phylogeny-based classification of the Echinostomatoidea. In the current systematic treatment: (i) the rank of two family level lineages, the former Himasthlinae and Echinochasminae, is elevated to full family status; (ii) Caballerotrema is distinguished at the family level; (iii) the content and diagnosis of the Echinostomatidae (sensu stricto) (s. str.) are revised to reflect its phylogeny, resulting in the abolition of the Nephrostominae and Chaunocephalinae as synonyms of the Echinostomatidae (s. str.); (iv) Artyfechinostomum, Cathaemasia, Rhopalias and Ribeiroia are re-allocated within the Echinostomatidae (s. str.), resulting in the abolition of the Cathaemasiidae, Rhopaliidae and Ribeiroiinae, which become synonyms of the Echinostomatidae (s. str.); and (v) refinements of the generic boundaries within the Echinostomatidae (s. str.), Psilostomidae and Fasciolidae are made., (Copyright © 2015 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The Indo-west Pacific is a marine bioregion stretching from the east coast of Africa to Hawaii, French Polynesia and Easter Island. An assessment of the literature from the region found reports of 2,582 trematode species infecting 1,485 fish species. Reports are concentrated in larger fishes, undoubtedly reflecting the tendency for larger hosts to be infected by more species of parasites as well as a collecting bias. Many hundreds of fish species, including many from families known to be rich in trematodes, have yet to be reported as hosts. Despite some areas (the Great Barrier Reef, Hawaii and the waters off China, India and Japan) receiving sustained attention, none can be considered to be comprehensively known. Several regions, most importantly in East Africa, French Polynesia and the Coral Triangle, are especially poorly known. The fauna of the Indo-west Pacific has been reported so unevenly that we consider it impossible to predict the true trematode richness for the region. We conclude that the greatest gap in our understanding is of the geographical distribution of species in the Indo-west Pacific. This is highlighted by the fact that 87% of trematodes in the region have been reported no more than five times. The reliable recognition of species is a major problem in this field; molecular approaches offer prospects for resolution of species identification but have been little adopted to date.

Two cystocercous gorgoderid cercariae of the genus Phyllodistomum Braun, 1899, Phyllodistomum umblae (Fabricius, 1780) and Phyllodistomum folium (Olfers, 1816), developing in sphaeriid bivalves from Norway and Lithuania, were studied and compared. Our previous molecular studies revealed very close phylogenetic relation of these two species and proved that cystocercous cercariae of P. folium develop in sporocysts in the gills of sphaeriid bivalves. In the present study morphological descriptions are given for the two cercariae, together with karyological and molecular characteristics. Comparative karyological and sequence analysis using ITS2 and 28S rDNA revealed the conspecificity of cercariae emitted from Pisidium hibernicum and Sphaerium corneum with adult P. umblae from Thymallus thymallus and Coregonus albula. The cercariae of P. folium and P. umblae are very similar morphologically with main differences in the structure of the tail. The two species clearly differ in karyotype structure. The diploid set of P. folium is composed of 18 chromosomes, but 16 chromosomes with one pair of large metacentrics were found in mitotic cells of P. umblae. Interspecific karyotypic difference presumably arose from Robertsonian fusion of two uni-armed chromosomes., (Copyright © 2015 Elsevier Ireland Ltd. All rights reserved.)

Cercariae and metacercariae of three species of the Microphallidae Travassos, 1920 were found in snails and crustaceans from tributaries of the Brisbane River, Queensland, Australia. Specimens of Maritrema brevisacciferum Shimazu et Pearson, 1991 and Microphallus minutus Johnston, 1948, which have previously been reported in Queensland, were found as cercariae in the tateid gastropod Posticobia brazieri (Smith) and as metacercariae of M. brevisacciferum in the atyid shrimp Caridina indistincta Calman and of M. minutus in the parastacid crayfish Cherax dispar Reik. Combined analysis of morphological and molecular data, based on newly generated ITS2 and partial 28S rDNA data, linked cercariae and metacercariae for both species. This is the first report of the first intermediate hosts of M. brevisacciferum and M. minutus. Infections of another unidentified microphallid metacercariae, Microphallidae gen. sp., were found in P. brazieri and C. indistincta. The sequences of metacercarial isolates from both hosts were identical. The data on the Microphallidae from Australia and species that develop in freshwater invertebrates were examined in detail.