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Rhopalocercous Cercaria duplicata von Baer, 1827 develops in an intermediate host, the unionid bivalve Anodonta anatina (L.), but its adult form has been unknown. We examined eight fish species occurring in the presence of a highly infested population of A. anatina in the Zesławice reservoir (S Poland). Gravid Phyllodistomum specimens were obtained from the ureters of ide, Leuciscus idus (L.) and common rudd, Scardinius erythrophthalmus (L.). One of the rudd specimens was doubly infected, a trematode was also found in the urinary bladder. In addition, a gravid Phyllodistomum specimen was found in the ureter of a tench Tinca tinca (L.), caught in Lake Ilmėdas (Lithuania). In order to clarify the phylogenetic position of larval and adult gorgoderids and to establish their life cycle, ITS2 and 28S rDNA sequences were analysed. The analysis showed that adult Phyllodistomum specimens located in the ureters are conspecific with C. duplicata. The trematode found in the urinary bladder of S. erythrophthalmus was P. folium (Olfers, 1816). It is suggested that adult stages of C. duplicata should be referred to as Phyllodistomum duplicatum n. comb. The intercaecal position of the uterus and the deeply-lobed ovary are the main features distinguishing it from other Phyllodistomum species. Host specificity and ecology are discussed., (© 2024. The Author(s).)

Members of the genus Scaphanocephalus mature in accipitrids, particularly osprey, Pandion haliaetus , with metacercaria causing Black Spot Syndrome in reef fishes. In most of the world, only the type species, Scaphanocephalus expansus (Creplin, 1842) has been reported. Recent molecular studies in the Western Atlantic, Mediterranean and Persian Gulf reveal multiple species of Scaphanocephalus , but have relied on 28S rDNA, mainly from metacercariae, which limits both morphological identification and resolution of closely related species. Here we combine nuclear rDNA with mitochondrial sequences from adult worms collected in osprey across North America and the Caribbean to describe species and elucidate life cycles in Scaphanocephalus . A new species described herein can be distinguished from S. expansus based on overall body shape and size. Phylogenetic analysis of the whole mitochondrial genome of Scaphanocephalus indicates a close relationship with Cryptocotyle . We conclude that at least 3 species of Scaphanocephalus are present in the Americas and 2 others are in the Old World. Specimens in the Americas have similar or identical 28S to those in the Mediterranean and Persian Gulf, but amphi-Atlantic species are unlikely in light of divergence in cytochrome c oxidase I and the lack of amphi-Atlantic avian and fish hosts. Our results provide insight into the geographic distribution and taxonomy of a little-studied trematode recently linked to an emerging pathology in ecologically important reef fishes.

Lymnaeid snails are some of the most widespread snails and are the first intermediate host of trematode parasites that affect human and livestock health. A full understanding of the genetic relationship of hosts and parasites is of paramount importance for effective parasite management. The present study assessed the prevalence of trematode larvae in lymnaeid snails and examined the genetic diversity of these snails collected across Thailand. We collected 672 lymnaeid snails from 39 locations in 22 provinces of six regions in Thailand. Subsequently, cercarial infection in the snails was observed by using the shedding method. Lymnaeid snails released 5 types of trematode cercariae, namely, xiphidiocercariae, echinostome cercariae I, echinostome cercariae II, furcocercous cercariae, and strigea cercariae. The phylogenetic analysis based on ITS2 and 28S rDNA sequences revealed 5 cercaria types assigned to four trematode families, of which two belong to the group of human intestinal flukes. Combination of shell morphology and sequence analysis of the mitochondrial COI and 16S rDNA genes, the lymnaeid snails were classified into two species, Radix rubiginosa and Orientogalba viridis. Moreover, the combined dataset of mtDNA genes (COI + 16S rDNA) from R. rubiginosa and O. viridis revealed 32 and 15 different haplotypes, respectively, of which only a few haplotypes were infected with cercariae. The genetic diversity and genetic structure revealed that R. rubiginosa and O. viridis experienced a bottleneck phenomenon, and showed limited gene flow between populations. Population demographic history analyses revealed that R. rubiginosa and O. viridis experienced population reductions followed by recent population expansion. These findings may improve our understanding of parasite-lymnaeid evolutionary relationships, as well as the underlying molecular genetic basis, which is information that can be used for further effective control of the spread of trematode disease., Competing Interests: Declaration of competing interest The authors declare there are no conflicts of interest., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Exploring the phylogenetic signal of morphological traits using geometric morphometry represents a powerful approach to assess the relative weights of convergence and shared evolutionary history in shaping species' forms. We evaluated the phylogenetic signal in shape and size of ventral and dorsal haptoral anchors of 10 species of monogenoids ( Hamatopeduncularia , Chauhanellus and Susanlimocotyle ) occurring in marine catfish (Siluriformes: Ariidae) from the Atlantic coast of South America. The phylogenetic relationships among these species were mapped onto the morphospaces of shape and size of dorsal and ventral anchors. Two different tests (squared change-parsimony and K mult ) were applied to establish whether the spatial positions in the phylomorphospace were influenced by phylogenetic relationships. A significant phylogenetic signal was found between anchor form and parasite phylogeny. Allometric effects on anchor shape were non-significant. Phylogenetically distant species on the same host differed markedly in anchor morphology, suggesting little influence of host species on anchor form. A significantly higher level of shape variation among ventral anchors was also found, suggesting that the evolutionary forces shaping ventral anchor morphology may operate with differing intensities or exhibit distinct mechanisms compared to their dorsal counterparts. Our results suggest that phylogenetic relationships were a key driver of changes in shape (but not size) of anchors of monogenoids of South American ariids. However, it seems that the emergence of the digitiform haptor in Hamatopenducularia and in some species of Chauhanellus played an important role in the reduction in anchor size and may cause secondary losses of anchors in other groups of monogenoids.

Species of Alestidae are known to be parasitised by dactylogyrid monogeneans representing three genera, Afrocleidodiscus Paperna, 1969, Annulotrema Paperna & Thurston, 1969, and Characidotrema Paperna & Thurston, 1968. The objective of the present study was to investigate the species diversity of Monopisthocotylea of African tetras from the Lower Phongolo River and floodplain in South Africa. Four new and two previously described species of Annulotrema were identified from the gills of three species of African tetras, Brycinus imberi, Hydrocynus vittatus, and Micralestes acutidens. The collected parasites were studied using two complementary approaches: morphology of hard sclerotised structures, and molecular markers using rDNA sequence data (28S rDNA, 18S rDNA, and ITS1). Three new species, Annulotrema arcum n. sp., Annulotrema caputfemoris n. sp., and Annulotrema strepsiceros n. sp., were described from B. imberi and one species, Annulotrema retortum n. sp., from M. acuditens. Two previously described species, Annulotrema pikoides Guégan, Lambert and Birgi, 1988 and A. pseudonili Kičinjaová and Řehulková, 2017, were newly recorded from H. vittatus in South Africa. Annulotrema arcum n. sp. and A. caputfemoris n. sp. share similar male copulatory organ morphologies, suggesting a close phylogenetic relationship as sister taxa. Despite weak nodal support, A. strepsiceros n. sp. shows morphological congruence with the former two species, reinforcing their molecular linkage. The present study shows a critical need for the exploration of monopisthocotylean diversity and the paucity of available molecular data of representatives from this group., (© M.L. Kičinjaová et al., published by EDP Sciences, 2024.)

We propose and describe Alobophora sandrae Cajiao-Mora & Bullard n. gen., n. sp. (Digenea: Caballerotrematidae) for specimens we collected from arapaima, Arapaima gigas sensu lato (Osteoglossiformes: Arapaimidae) in the Amazon River near Leticia, Colombia. Alobophora differs from Caballerotrema Prudhoe, 1960 by lacking head collar projections and by having clustered corner spines and a narrow head collar (4-5× wider than pharynx), whereas Caballerotrema has head collar projections, lacks clustered corner spines, and has a broad head collar (7-8× wider than pharynx). We reassign Caballerotrema annulatum (Diesing, 1850) Ostrowski de Núñez & Sattmann, 2002 to the new genus, as Alobophora annulata (Diesing, 1850) Cajiao-Mora and Bullard n. comb., and provide a supplemental description of Caballerotrema brasiliense Prudhoe, 1960 based on specimens we collected from arapaima. We also examined the holotype and a paratype of Caballerotrema piscicola (Stunkard, 1960) Kostadinova & Gibson, 2001 and concluded that C. piscicola is a junior subjective synonym of C. brasiliense. Our 28S phylogeny recovered A. sandrae sister to A. annulata, with that clade sister to a clade comprising C. brasiliense and an innominate species of Caballerotrema. Caballerotrematidae was recovered sister to Echinostomatidae. We also provide a dichotomous key to caballerotrematids based on head collar projections, corner spine arrangement, proportional pharynx and head collar breadth, testes shape and arrangement, body surface spine shape and distribution, vitellarium distribution, and abundance of prostatic cells., (© K. Cajiao-Mora et al., published by EDP Sciences, 2024.)

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"., (© C. Bouguerche et al., published by EDP Sciences, 2024.)

A parasitological investigation of Cyprinella venusta and Notropis cf. stramineus sampled in Texas, USA, in the Guadalupe River, revealed the presence of Gyrodactylus crysoleucas Mizelle and Kritsky, 1967 on C. venusta, and Gyrodactylus mediotorus King, Marcogliese, Forest, McLaughlin & Bentzen, 2013 on both fish species. This represents new leuscicid fish hosts and locality records for these two gyrodactylids. Gyrodactylus crysoleucas previously identified from both non-native Californian Notemigonus crysoleucas and from farmed stocks in Minnesota demonstrated intraspecific variability in terms of morphology and genetics as a local adaptation associated with isolation by distance. Results further confirmed G. crysoleucas as alien in the western USA and suggested host-switching involving C. venusta and N. crysoleucas. Conservative morphology and genetics on the part of G. mediotorus from C. venusta and N. cf. stramineus (Guadalupe River) was observed, while higher genetic divergence in the ITS sequences associated with morphological discrepancy was found between the studied G. mediotorus specimens and those of Notropis hudsonius than when considering the parasites of Notropis texanus. The separation of G. mediotorus into geographical subgroups may indicate ongoing speciation linked to the Pleistocene glaciations in North America, and to hydrographic barriers that facilitated separate evolutionary paths leading to speciation. We suggest that deep investigations of Gyrodactylus populations will help to understand the speciation of these parasites and their adaptation to Nearctic fish hosts., (© C. Rahmouni et al., published by EDP Sciences, 2024.)

Parasites have evolved a variety of astonishing strategies to survive within their hosts, yet the most challenging event in their personal chronicles is the passage from one host to another. It becomes even more complex when a parasite needs to pass through the external environment. Therefore, the free-living stages of parasites present a wide range of adaptations for transmission. Parasitic flatworms from the group Digenea (flukes) have free-living larvae, cercariae, which are remarkably diverse in structure and behavior. 1 , 2 One of the cercariae transmission strategies is to attain a prey-like appearance for the host. 3 This can be done through the formation of a swimming aggregate of several cercariae adjoined together by their tails. 4 Through the use of live observations and light, electron, and confocal microscopy, we described such a supposedly prey-mimetic colony comprising cercariae of two distinct morphotypes. They are functionally specialized: larger morphotype (sailors) enable motility, and smaller morphotype (passengers) presumably facilitate infection. The analysis of local read alignments between the two samples reveals that both cercaria types have identical 18S, 28S, and 5.8S rRNA genes. Further phylogenetic analysis of these ribosomal sequences indicates that our specimen belongs to the digenean family Acanthocolpidae, likely genus Pleorchis. This discovery provides a unique example and a novel insight into how morphologically and functionally heterogeneous individuals of the same species cooperate to build colonial organisms for the purpose of infection. This strategy bears resemblance to the cooperating castes of the same species found among insects. 5 ., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2023 The Authors. Published by Elsevier Inc. All rights reserved.)

In recent field studies, suspected gymnophallid metacercariae were histologically located in the mantle of mussels from the Norwegian Sea. Mussels from the sites in which that infection was detected also presented abnormally high pearl numbers. It has been previously described that gymnophallid metacercariae could cause pearl formation processes in mussels, as a host reaction to encapsulate these metacercariae. Given the pathological host reaction these parasites elicit, a study was performed to identify gymnophallid metacercariae found in mussels collected from Tromsø at morphological and molecular level and to assess, by the use of molecular tools, the relationship between the parasite and the biological material inside the pearls. As a result, Gymnophallus bursicola metacercariae infecting Norwegian Mytilus edulis were identified according to morphological characters, along with the first 18S rDNA and COI sequences for this trematode species. In addition, parasite DNA from the core of the pearls was extracted and amplified for the first time, confirming the parasitological origin of these pearls. This procedure could allow identifying different parasitic organisms responsible for the generation of pearls in bivalves.

Five new species of Cosmetocleithrum were described parasitizing the gill filaments of neotropical doradid fishes. Cosmetocleithrum undulatum n. sp., Cosmetocleithrum brachylecis n. sp. and Cosmetocleithrum ludovicense n. sp. are described from Platydoras brachylecis from a market-place of São Luís, State of Maranhão, Brazil. Cosmetocleithrum sacciforme n. sp. and Cosmetocleithrum basicomplexum n. sp. are described from Oxydoras niger from Juruá River, State of Acre, Brazil. Cosmetocleithrum undulatum and Cosmetocleithrum brachylecis resemble Cosmetocleithrum falsunilatum Feronato, Razzolini, Morey & Boeger, 2022 mainly by the unique male copulatory organ (MCO) morphology but differ from these and all congeneric species mainly by the morphology of the MCO, accessory piece and hooks pairs. Cosmetocleithrum ludovicense is closer to Cosmetocleithrum confusus Kritsky, Thatcher & Boeger, 1986 and to Cosmetocleithrum akuanduba Soares, Santos Neto & Domingues, 2018 but differs from those mainly by the morphology of the accessory piece. Cosmetocleithrum sacciforme differs from all congeneric species mainly by the morphology of the accessory piece formed by a single plate of saccular appearance. Cosmetocleithrum basicomplexum also shares morphological characters with Cosmetocleithrum gigas Morey, Cachique & Babilonia, 2019 considering the size of the body and shape of the anchors, but differs mainly in the morphology of the bars and hooks. Besides the new species, new data are presented for Cosmetocleithrum leandroi Soares, Neto & Domingues, 2018, C. akuanduba and C. confusus regarding morphological characteristics and biogeography., (© A.L. Silva et al., published by EDP Sciences, 2023.)

Austrodiplostomum spp. (Platyhelminthes: Digenea) are endoparasites with a broad geographic distribution in South America. During the larval stage, they parasitize the eyes, brains, muscles, gill, kidneys and swim bladder of a wide variety of fishes. The metacercariae of Austrodiplostomum spp. have several morphological characteristics during development, but are very similar among species, which makes it necessary to use molecular tools to contribute to the elucidation during the larval stage. The objective of this study was to perform morphological and molecular analyses of Austrodiplostomum sp. found in specimens of Hypostomus sourced from the Ivaí River in the state of Paraná, Brazil. Of the 93 analyzed specimens (H. hermanni [n = 50], H. albopunctatus [n = 9], Hypostomus sp. 1 [n = 24], and Hypostomus sp. 2 [n = 10]), 60 were parasitized. A total of 577 Austrodiplostomum sp. metacercariae was collected from the infected hosts; DNA from seven of these samples was extracted, amplified, and sequenced. The morphological data associated with the genetic distance values and the relationships observed in the COI gene tree, indicate that all metacercariae were A. compactum. This is the first record of A. compactum parasitizing H. hermanni, H. albopunctatus, Hypostomus sp. 1, and Hypostomus sp. 2 in the Ivaí River.

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.)

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., (© C. Bouguerche et al., published by EDP Sciences, 2022.)

Estimates of trematode diversity are inaccurate due to unrecognized cryptic species and phenotypic plasticity within species. Integrative taxonomy (genetics, morphology and host use) increases the clarity of species delineation and improves knowledge of parasite biology. In this study, we used this approach to resolve taxonomic issues and test hypotheses of cryptic species in a genus of trematode, Quinqueserialis. Specimens from throughout North America were field collected from hosts and obtained from museums. We found three morphologically distinct groups and successfully sequenced specimens from two of these groups. DNA sequencing at the 28S and CO1 gene regions revealed that two of the three groups were genetically distinct. One genetic group included two morphological clusters demonstrating host-induced phenotypic plasticity within Quinqueserialis quinqueserialis. The other unique genetic group is a novel species, Quinqueserialis kinsellai n. sp., which is described herein. Our study illustrates the importance of integrating multiple sources of evidence when investigating trematode diversity to account for the influence of cryptic species or phenotypic plasticity. However, further sampling is needed to understand Quinqueserialis spp. diversity as some species have no genetic information associated with them.

Translocation of fishes for aquaculture has resulted in the co-introduction of some of their parasites. African cichlid fishes, generically called "tilapias" have been introduced worldwide, along with their monogenean parasites. In a nation-wide survey, we characterised monogeneans of the genus Gyrodactylus infecting farmed "tilapia" throughout Mexico. We also collected native fishes around farms, to look for potential parasite spillover from cultured fishes. Monogeneans were identified taxonomically using morphological and molecular characters. Originally African, pathogenic Gyrodactylus cichlidarum was recorded in every farm surveyed, infecting different "tilapia" varieties, as well as three native cichlid fish species. Previously, we had shown that G. cichlidarum also infects native, non-cichlid fishes in Mexico. We also recorded that Gyrodactylus yacatli is widely distributed in Mexico, infecting cultured "tilapia" and native fishes; and present data indicating that this is a further translocated African parasite. A third, unidentified gyrodactylid infected farmed and native fishes in Chiapas, southern Mexico; we describe the new species as Gyrodactylus shinni n. sp., and provide evidence that this is a third monogenean translocated with African fish. The wide distribution of exotic parasites co-introduced with "tilapia" and their spillover to native fishes may have an important impact on the ichthyofauna in Mexico, one the world's megadiverse countries.

Amphistome species belonging to the genus Catadiscus are poorly studied intestinal trematodes found primarily in Neotropical anurans. Herein, developmental stages of an amphistome species found during herpetological and malacological surveys in a temporary marsh pond from Brazil were subjected to morphological (light and scanning electron microscopy) and molecular analyses. Adult parasites recovered from anurans were identified as Catadiscus marinholutzi. Amphistome cercariae found in the planorbid snails Drepanotrema depressissimum and Drepanotrema lucidum from the same waterbody were used for experimental and molecular studies. Immature parasites, morphologically compatible with members of Catadiscus, were experimentally obtained in laboratory-reared tadpoles. Sequencing of a partial region of 28S rDNA gene of both adult and cercariae revealed 100% similarity between these developmental stages, confirming their conspecificity. Phylogenetic analyses were attempted for the first time to reveal the position of a species of Catadiscus in the superfamily Paramphistomoidea. Catadiscus marinholutzi falls in a virtual polytomy together with other paramphistomoids, which leaves its phylogenetic relationships within the group unclear. Moreover, the high genetic divergence to Diplodiscus spp. (10.06–10.84%) cast doubts on the placement of Catadiscus within Diplodiscidae. Hence the species composition of the Diplodiscidae should be re-evaluated in further studies using a broader spectrum of related taxa.

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.

Monogeneans rely on firm attachment to often flexible and uneven surfaces and are renowned for their effective posterior attachment structures in the form of adhesives, clamps, hamuli and suckers. Polystomatids do not secrete adhesives and do not have clamps. While only some have hamuli, all have suckers in the adult form. Three different types of haptoral suckers have been described based on basic morphology but have never been studied in depth. Using enzyme digestion and light (differential interference contrast), confocal and scanning electron microscopy, we examined representatives and propose four sucker types. Haptoral sucker Type I are symmetrical soft, flexible, cup- to disk-shaped suckers and are found in all polystomes infecting frogs and salamanders. Type II suckers are symmetrical soft, flexible, cup-shaped suckers with a hollow continuous skeletal ring and no other skeletal elements. They are found in species of Nanopolystoma Du Preez, Wilkinson et Huyse, 2008 infecting caecilians. Type III suckers are symmetrical firm, cup-shaped suckers with elaborate skeletal elements that contribute to a secure grip on the host tissue. This type of sucker is found in all polystomes infecting freshwater turtles and the common hippopotamus. Type IV suckers are asymmetrical with an elaborate series of long, thin sclerites with terminal spines or hooks. This type of sucker is only known from Concinnocotyla australensis (Reichenbach-Klinke, 1966) infecting the Australian lungfish. These different sucker types are crucial for the survival of polystomatid flatworms within their respective microhabitats.

We investigated the prevalence, morphological characters and molecular classifications of trematode cercariae in freshwater snails randomly collected from 59 sampling localities in Bangkok from May 2018 to March 2019. We used a crushing technique to observe the cercarial stage inside each snail body and amplified the internal transcribed spacer 2 regions of cercarial DNA using polymerase chain reaction methodology. The associated phylogenetic tree was reconstructed using Bayesian inference analyses. A total of 517 of 15 621 examined snails were infected with trematode cercariae, and the infected snails were classified into 11 species of seven families with a 3.31% overall prevalence of the infection. The Bithynia siamensis siamensis snail displayed the highest prevalence of infection (16.16%), whereas the Physella acuta snail exhibited the lowest prevalence (0.08%) of infection. Eight morphological types of cercariae were observed. The highest prevalence of infection was observed in mutabile cercaria (1.86%). Based on molecular investigations, the phylogram revealed eight cercaria types assigned to at least nine digenean trematode families, of which five belong to groups of human intestinal flukes. Although, with the exception of schistosome cercaria, trematode cercariae are not known to directly damage humans, understanding the general biology of trematode cercariae (including diversity, distribution, infection rates and host range) is important and necessary for the prevention and control of parasitic transmission that impacts aquatic cultivations, livestock farming and human health.

Life cycle stages, including daughter sporocysts, cercariae, and metacercariae, of Parvatrema duboisi (Dollfus, 1923) Bartoli, 1974 (Digenea: Gymnophallidae) have been found in the Manila clam Ruditapes philippinarum from Aphaedo (Island), Shinan-gun, Jeollanam-do, Korea. The daughter sporocysts were elongated sac-like and 307-570 (av. 395) μm long and 101-213 (av. 157) μm wide. Most of the daughter sporocysts contained 15-20 furcocercous cercariae each. The cercariae measured 112-146 (av. 134) μm in total length and 35-46 (av. 40) μm in width, with 69-92 (av. 85) μm long body and 39-54 (av. 49) μm long tail. The metacercariae were 210-250 (av. 231) μm in length and 170-195 (av. 185) μm in width, and characterized by having a large oral sucker, genital pore some distance anterior to the ventral sucker, no ventral pit, and 1 compact or slightly lobed vitellarium, strongly suggesting P. duboisi. The metacercariae were experimentally infected to ICR mice, and adults were recovered at day 7 post-infection. The adult flukes were morphologically similar to the metacercariae except in the presence of up to 20 eggs in the uterus. The daughter sporocysts and metacercariae were molecularly (ITS1-5.8S rDNA-ITS2) analyzed to confirm the species, and the results showed 99.8-99.9% identity with P. duboisi reported from Kyushu, Japan and Gochang, Korea. These results confirmed the presence of various life cycle stages of P. duboisi in the Manila clam, R. philippinarum, playing the role of the first as well as the second intermediate host, on Aphae-do (Island), Shinan-gun, Korea.

Background: Gyrodactylus salaris Malmberg, 1957 is an OIE (Office International des Epizooties)-listed parasitic pathogen and had until the current study been reported from 19 countries across Europe, although many of these records require confirmation. The last comprehensive evaluation regarding the distribution of G. salaris, however, was made in 2007, although some of the states identified as being G. salaris-positive were ascribed this status based on misidentifications, on partial data resulting from either morphological or molecular tests, or from records that have not been revisited since their early reporting. It is thus important to go through the reports on G. salaris to obtain a status for each country., Methods: To provide a revised update of the G. salaris distribution, a literature review was necessary. This literature, however, was not always readily accessible and, in certain cases, the article only made superficial reference to the parasite without providing details or data to support the identification. In most cases, the original specimens were not deposited in a national collection. Additional Gyrodactylus material for the current study was obtained from selected salmonid populations with the aim to contribute to current understanding regarding the distribution of G. salaris. Additional parasite material collected for this study was processed following standard procedures for species identification in Gyrodactylus [1]., Results: From the work conducted in the current study, G. salaris is reported from a further three regions in Italy, alongside three other species, and appears to occur extensively throughout central Italy without causing significant mortalities to its rainbow trout, Oncorhynchus mykiss (Walbaum), host. The analysis of archive material from G. salaris-positive farms would suggest that G. salaris has been in this country since at least 2000. Material obtained from rainbow trout from Finland and Germany are confirmed as G. salaris, supporting existing data for these countries. No specimens of G. salaris, however, were found in the additional Gyrodactylus material obtained from rainbow trout reared in Portugal and Spain. A morphologically similar species, Gyrodactylus teuchis Lautraite, Blanc, Thiery, Daniel et Vigneulle, 1999, however, was found., Conclusions: Following the present review, Gyrodactylus salaris is reported from 23 out of 50 recognised states throughout Europe; only records from 14 of these states have been confirmed by either morphology and/or by an appropriate molecular test and are considered valid, while only nine of these records have been confirmed by a combination of both methods.

The morphology of sexual adults is the cornerstone of digenean systematics. In addition, life cycle data have always been significant. The integration of these approaches, supplemented with molecular data, has allowed us to detect a new species that many researchers may have previously seen, but not recognized. Sexual adults from common eiders that we found in northern European seas were extremely similar to other notocotylids, but the discovery of their intermediate host, a marine snail, revealed the true nature of this material. Here we describe sexual adults, rediae and cercariae of Catatropis onobae sp. nov. We discuss how 'Catatropis verrucosa' should be regarded, justify designation of the new species C. onobae for our material and explain why it can be considered a cryptic species. The phylogenetic position of C. onobae within Notocotylidae, along with other evidence, highlights the challenges for the taxonomy of the family, for which two major genera appear to be polyphyletic and life cycle data likely undervalued.

Gyrodactylid parasites were observed on non-native populations of North-American freshwater catfishes, Ameiurus nebulosus and Ameiurus melas (Siluriformes: Ictaluridae), at several sites in the Elbe River basin, Czech Republic, Europe. Using a combination of morphological and genetic analyses, the parasites infecting A. nebulosus were determined to be Gyrodactylus nebulosus, a North American parasite co-introduced to Europe along with its Ameiurus fish hosts. Subtle morphometrical differences, as well as seasonal variations, were observed among parasites collected from A. nebulosus and A. melas. The host-related variation was further supported through genetic analysis of the partial 18S rDNA, ITS1-5.8S-ITS2 and COI, showing 0.2, 3.0 and 4.8% divergence, respectively. Consistent genetic differences indicated there were two distinct genotypes. Subtle morphological differences associated with the shape of sickle toe, anchor root and ventral bar membrane, according to host species, also supported the description of a new cryptic species, Gyrodactylus melas n. sp., infecting A. melas. Multivariate morphometrical analysis of haptoral hard parts showed significant differences between the anchor lengths of G. nebulosus and G. melas n. sp. However, the measurements of the haptoral hard structures partially overlapped between species, limiting the usage of these parameters for species delineation.

Background: The flatworms (Lophotrochozoa: Platyhelminthes) are one of the major phyla of invertebrates but their interrelationships are still not well understood including unravelling the most closely related taxon of the Neodermata, which includes exclusively obligate parasites of all main groups of vertebrates with some 60,000 estimated species. Recent phylogenomic studies indicate that the freshwater 'microturbellarian' Bothrioplana semperi may be the closest ancestor to the Neodermata, but this hypothesis receives little morphological support. Therefore, additional morphological and ultrastructural characters that might help understand interrelations within the Neodermata are needed., Methods: Ultrastructure of the excretory ducts of representatives of the most basal parasitic flatworms (Neodermata), namely monocotylid (Monopisthocotylea) and chimaericolid (Polyopisthocotylea) monogeneans, aspidogastreans (Trematoda), as well as gyrocotylidean and amphilinidean tapeworms (Cestoda), were studied using transmission electron microscopy (TEM)., Results: The present study revealed the same pattern of the cytoarchitecture of excretory ducts in all studied species of the basal neodermatans. This pattern is characterised by the presence of septate junctions between the adjacent epithelial cells and lateral ciliary flames along different levels of the excretory ducts. Additionally, a new character was observed in the protonephridial terminal cell of Gyrocotyle urna, namely a septate junction between terminal and adjacent duct cells at the level of the distal extremity of the flame tuft. In Amphilina foliacea, a new type of protonephridial cell with multiple flame bulbs and unique character of its weir, which consists of a single row of the ribs, is described. A remarkable difference has been observed between the structure of the luminal surface of the excretory ducts of the studied basal neodermatan groups and B. semperi., Conclusions: The present study does not provide ultrastructural support for a close relationship between the Neodermata and B. semperi.

Eudiplozoon nipponicum (Goto, 1891) Khotenovsky, 1985 (Monogenea: Diplozoidae), is known to parasitise Cyprinus carpio Linnaeus and species of Carassius. In this study, we conducted a taxonomic re-examination of E. nipponicum using genetic analysis and morphological comparisons from different host species from a single water system. rDNA nucleotide sequences of the internal transcription spacer 2 (ITS-2) region (645 bp) showed interspecific-level genetic differences among diplozoids from species of Carassius and C. carpio (p-distance: 3.1-4.0%) but no difference among those from different species of Carassius (0-0.4%) or between those from C. carpio collected in Asia and Europe (0-1.1%). Large variation was observed among 346 bp cytochrome c oxidase subunit I (COI) sequences (0.3-16.0 %); the topology of the phylogenetic tree showed no relationship to host genera or geographical regions of origin. Morphological observation showed that average clamp size of diplozoids from C. carpio was larger than those from Carassius spp. The number of folds on the hindbody was 10-25 for diplozoids from C. carpio and 12-19 for those from Carassius spp. Thus, our ITS-2 sequence and morphological comparison results indicate that diplozoids from C. carpio and species of Carassius belong to different species. The scientific name E. nipponicum should be applied to the species infected to the type host, Carassius sp. of Nakabo (2013) (Japanese name ginbuna). The diplozoid infecting C. carpio (Eurasian type) should be established as a new species: Eudiplozoon kamegaii sp. n. A neotype of E. nipponicum is designated in this report because the original E. nipponicum specimens are thought to have been lost.

We intended to describe 2 digenean trematodes found from a Chinese sea snake, Laticauda semifasciata, as the new fauna in the Republic of Korea. The snake was caught offshore of Aewol-eup, Jeju-do, in August 2017. Two species of fluke were found in the lung and intestinal tract of the snake in the process of necropsy. They were identified as Pulmovermis cyanovitellosus Coil and Kuntz, 1960 and Harmotrema laticaudae Yamaguti, 1933, respectively based on the morphological characters. Pulmovermis cyanovitellosus showed elongated body with well-developed and elongated male genital system and compact vitelline. And H. laticaudae was characterized by linguiform body with heavily armed cirrus with excretory system. This is the first time both species have been reported and described off the Korean coast. We provide morphological descriptions with some comments on their biology and geographical distribution. In addition, the taxonomic validity of the genus Hydrophitrema Sandars, 1960 was discussed, in terms of morphological descriptions and host ranges. This study provides novel insight into digenean fluke species existing off the coast of Korea.

Ruavermis mikebargeri gen. et sp. n. infects the yellow-headed temple turtles Heosemys annandalii (Boulenger et Robinson) in the Mekong River Basin. It resembles Platt Roberts et Bullard, 2018 and Coeuritrema Mehra, 1933 by having the anterior to posterior anatomical sequence of a ventral sucker, external seminal vesicle, cirrus sac, anterior testis, ovary, transverse vitelline duct, and posterior testis. These genera are further similar by having the combination of an elongate/ovoid aspinous body, a ventral sucker at the level of the body constriction, an oesophagus that terminates in the anterior 1/5 of the body and that is ventral to the anterior nerve commissure, intestinal caeca that bifurcate in the anterior 1/3 of the body (not immediately anterior to the ventral sucker), a sinistral caecum that bends toward the midline at level of the cirrus and common genital pore, an external seminal vesicle that abuts the anterodextral margin of the cirrus sac, an oviduct that emerges from the dextral margin of the ovary, and an oviducal seminal receptacle that comprises the middle portion of the oviduct. These genera lack lateral oesophageal diverticulae and a median oesophageal diverticulum. The new genus is unique by having a papillate ventral body surface, an external seminal vesicle lateral to the cirrus sac, vasa efferentia that are ventral to the gonads, an oviduct that is convoluted, a Laurer's canal pore that is preovarian, a Laurer's canal that extends anterolaterad, and an excretory vesicle that is Y-shaped. The 28S rDNA phylogenetic analysis recovered the new species sister to Coeuritrema platti Roberts et Bullard, 2016, with that clade sister to Hapalorhynchus spp. and Platt spp. The new turtle blood fluke is the fourth from Vietnam, second from a Vietnam geomydid, and first from Heosemys Stejneger as well as the first endohelminth from the yellow-headed temple turtle.

Background: Lake Tanganyika is considered a biodiversity hotspot with exceptional species richness and level of endemism. Given the global importance of the lake in the field of evolutionary biology, the understudied status of its parasite fauna is surprising with a single digenean species reported to date. Although the most famous group within the lake's fish fauna are cichlids, the pelagic zone is occupied mainly by endemic species of clupeids (Actinopterygii: Clupeidae) and lates perches (Actinopterygii: Latidae, Lates Cuvier), which are an important commercial source for local fisheries. In this study, we focused on the lake's four lates perches and targeted their thus far unexplored endoparasitic digenean fauna., Methods: A total of 85 lates perches from four localities in Lake Tanganyika were examined. Cryptogonimid digeneans were studied by means of morphological and molecular characterisation. Partial sequences of the nuclear 28S rRNA gene and the mitochondrial cytochrome c oxidase subunit 1 (cox1) gene were sequenced for a representative subset of the specimens recovered. Phylogenetic analyses were conducted at the family level under Bayesian inference., Results: Our integrative approach revealed the presence of six species within the family Cryptogonimidae Ward, 1917. Three out of the four species of Lates were found to be infected with at least one cryptogonimid species. Two out of the three reported genera are new to science. Low interspecific but high intraspecific phenotypic and genetic diversity was found among Neocladocystis spp. Phylogenetic inference based on partial 28S rDNA sequences revealed a sister group relationship for two of the newly erected genera and their close relatedness to the widely distributed genus Acanthostomum Looss, 1899., Conclusions: The present study provides the first comprehensive characterisation of the digenean diversity in a fish family from Lake Tanganyika which will serve as a baseline for future explorations of the lake's digenean fauna. Our study highlights the importance of employing an integrative approach for revealing the diversity in this unique host-parasite system.

A revisionary study revealed two species of monogeneans, Dactylogyrus crucis Rogers, 1967 and Dactylogyrus lythruri sp. n., parasitising Lythrurus Jordan (formerly a subgenus of Notropis Rafinesque, 1818). New records and updated taxonomy of seven of 12 known minnows of the genus of Lythrurus are provided for D. crucis. A record of Dactylogyrus attenuatus Mizelle et Klucka, 1953 (syn. Dactylogyrus umbratilus [Kimpel, 1939], nomen nudum) on Lythrurus umbratilis (Girard) is referrable to D. crucis. Dactylogyrus lythruri is described from eight species of Lythrurus. It most closely resembles Dactylogyrus beckeri Cloutman, 1987, but is distinguished by having a smaller base of the male copulatory organ (MCO) and lacking a ventral enlargement of the distal end of the basal process. Previous reports of Dactylogyrus banghami Mizelle et Donahue, 1944 and Dactylogyrus cf. beckeri Cloutman, 1987 from Lythrurus atrapiculus (Snelson) and Lythrurus bellus (Hay) from Alabama, and Dactylogyrus confusus Mueller, 1938 from L. umbratilis in Illinois are herein relegated to D. lythruri. Four species of Lythrurus appeared not to be infected with Dactylogyrus.

This is the first record of the trematode Choledocystus incurvatum and Choledocystus elegans in Argentinean amphibians collected in two localities of the Chacoan region. The first trematode species was found infecting the small intestine of Pseudis paradoxa Linnaeus, captured near the city of Corrientes, Corrientes Province; the second species was found infecting the gall-bladder of Lepidobatrachus laevis Budgget, captured in Ingeniero Juárez, Formosa Province. Morphology of these species was studied in detail using light microscopy and scanning electron microscopy (SEM), and compared with previous studies of Brazilian and Venezuelan specimens. Morphological descriptions of these parasites are provided to supplement existing data. Choledocystus incurvatum was characterized using molecular methods by sequencing and analyzing rDNA. Regarding metric characters of C. incurvatum, the Argentinean specimens show some differences in body and egg size (some larger than Venezuelan specimens). For C. elegans the specimen shows smaller body size than Brazilian specimens. SEM observations of C. incurvatum showed tegument covered with triangular spines and confirmed presence of a sucker-like structure around the genital atrium. The morphological and genetic data on C. incurvatum contribute to a more thorough diagnosis of the genus. Moreover, this report represents the first occurrence of this genus in the Argentine Chacoan region.

Background: Monogenea van Beneden, 1858 is a group of parasitic flatworms, commonly found infecting bony fish. Several genera, such as Cichlidogyrus Paperna, 1960, are reported to include potential pathogenic species that can negatively impact aquaculture fish stocks. They can switch from introduced to native fish and vice versa. In Africa (and all over the world), fish species belonging to Cichlidae are often kept in aquaculture and represent a major source of food. Thus, research on the biodiversity and occurrence of monogenean species on these fish is of importance for aquaculture and conservation. The present study is a survey of the diversity of species of Cichlidogyrus in the south of the Democratic Republic of the Congo (DRC) on three cichlid species: Orthochromis sp. 'Lomami', Serranochromis cf. macrocephalus, and Tilapia sparrmanii Smith, 1840., Methods: Specimens of Cichlidogyrus were isolated from the gills and mounted on glass slides with Hoyer's medium. The genital and haptoral hard parts were measured and drawn using interference contrast., Results: In total, six species of Cichlidogyrus were found, all new to science: C. bulbophallus n. sp. and C. pseudozambezensis n. sp. on S. cf. macrocephalus, C. flagellum n. sp. and C. lobus n. sp. on T. sparrmanii, C. ranula n. sp. on S. cf. macrocephalus and Orthochromis sp. 'Lomami', and C. maeander n. sp. found on Orthochromis sp. 'Lomami' and T. sparrmanii. The first four species are considered to be strict specialists, C. ranula n. sp. an intermediate generalist and C. maeander n. sp. a generalist. These parasite species show morphological similarities to species found in the Lower Guinea and Zambezi ichthyofaunal provinces, which might be explained by past river capture events between river systems of the Congo Province and both these regions., Conclusions: Serranochromis cf. macrocephalus and Orthochromis sp. 'Lomami' can harbour respectively three and two species of Cichlidogyrus, all described in this study. Tilapia sparrmanii can harbour seven species, of which three are described in the present study. These results highlight the species diversity of this parasite genus in the Congo Basin.

Gymnophallid metacercariae found in the Manila clam Ruditapes philippinarum ('Banjirak' in Korean) from Gochang-gun, Jeollabuk-do, Korea were morphologically and molecularly confirmed to be Parvatrema duboisi (Dollfus, 1923) Bartoli, 1974. The metacercariae were morphologically characterized by having a large oral sucker, small ventral sucker, genital pore some distance anterior to the ventral sucker, no ventral pit, and 1 compact or slightly lobed vitellarium, which were all compatible with P. duboisi. Some of the metacercariae were experimentally fed to mice, and adult flukes were recovered at day 7 post-infection. The morphology of the adult flukes was basically the same as that of the metacercariae except for the presence of uterine eggs; the uterus was filled with up to 40 eggs. The nucleotide sequences (1,193 bp) from ITS regions (ITS1, 5.8S rDNA, and ITS2) of the metacercariae showed 99.7% identity with P. duboisi and 75.7% identity with Gymnophalloides seoi deposited in GenBank. These results confirmed the presence of P. duboisi metacercariae in the Manila clam R. philippinarum in an estuary region of Gochang-gun, Korea.

Background: Microcotyle erythrini van Beneden & Hesse, 1863 (Platyhelminthes: Monogenea) and other closely related species of the genus are often considered as cryptic. Records in hosts other than the type-host with no species confirmation by molecular analyses have contributed to this situation., Methods: Gill parasites of five sparid fishes, Boops boops (L.), Pagellus erythrinus (L.), P. acarne (Risso), Dentex dentex (L.) and Pagrus pagrus (L.), from the Western Mediterranean off Spain were collected. Specimens of Microcotyle spp. were characterised both molecularly and morphologically. Partial fragments (domains D1-D3) of the 28S rRNA gene and the cytochrome c oxidase subunit 1 (cox1) gene were amplified and used for molecular identification and phylogenetic reconstruction. Principal components analysis was used to look for patterns of morphological separation., Results: Molecular analyses confirmed the identity of three species: M. erythrini ex P. erythrinus and Pa. pagrus; M. isyebi Bouguerche, Gey, Justine & Tazerouti, 2019 ex B. boops; and a species new to science described herein, M. whittingtoni n. sp. ex D. dentex. The specific morphological traits and confirmed hosts (P. erythrinus and Pa. pagrus) are delimited here in order to avoid misidentifications of M. erythrini (sensu stricto). Microcotyle erythrini (s.s.) is mostly differentiated by the shape of its haptor, which is also longer than in the other congeners. New morphological and molecular data are provided for M. isyebi from the Spanish Mediterranean enlarging the data on its geographical range. Microcotyle whittingtoni n. sp. is described from D. dentex and distinguished from the remaining currently recognised species of the genus by the number and robustness of the clamps., Conclusions: New diagnostic morphological traits useful to differentiate Microcotyle spp. are suggested: (i) haptor dimensions including lobes; (ii) the thickness of the clamps; (iii) the size and shape of spines of the genital atrium; (iv) the extension of the posterior extremities of vitelline fields; and (v) the shape of egg filaments. The use of new morphological approaches may allow considering these species of Microcotyle as being pseudocryptic. The use of representative undamaged specimens that have been genetically confirmed as conspecific is considered crucial to avoid abnormally wide morphological ranges that prevent species differentiation.

More than 1.450 marine fish species are commercialized as ornamental fish and approximately 90% of these are wild-caught. Rainford's butterflyfish, Chaetodon rainfordi (McCulloch, 1923), is endemic to the western Pacific. This fish species belongs to one of the main ornamental fish families imported into Brazil. Haliotrema is a genus of the family Dactylogyridae. Species of this genus have very interesting host associations and geographic distributions. The aim of this study was to report on occurrences of exotic Haliotrema aurigae in diseased Rainford butterflyfish that were imported to Brazil. One Rainford's butterflyfish with suspected parasitic disease was euthanized and subjected to skin and gill scraping. The parasite specimens were mounted on semi-permanent slides with Hoyer's solution for morphological analysis of sclerotized structures. The hard-sclerotized structures of these specimens were compared and were identified as those of Haliotrema aurigae (Yamaguti, 1968). These results emphasize the importance of implementation of correct control measures at the country's borders, to avoid the introduction of exotic parasites. In addition, it can be emphasized that the easy adaptation of this parasite to new environments is a concern for ornamental fish farmers.

A new species of Tereancistrum Kritsky, Thatcher & Kayton, 1980 collected of Prochilodus lineatus gills from the Batalha River, Tietê-Batalha basin, São Paulo State, Brazil is described. The new species can be distinguished from its congeners mainly by the configuration of the ventral bar, which has an anvil-shaped characteristic with corrugated anterior projection. Tereancistrum takemotoi n. sp. is morphologically similar to T. toksonum Lizama, Takemoto & Pavanelli, 2004 in terms of their dorsal bars (Y-shaped), their dorsal anchors with divergent roots (superficial and deep) wherein their deep root rather elongated, and by the fact that they both have the male copulatory organ counterclockwise. However, only T. takemotoi n. sp. presents the male copulatory organ with 2¼ rings and shows undulations in the anterior margin of the dorsal bar. These undulations are absent in T. toksonum (which only has 1¼ rings). This is the fourth Tereancistrum species described for P. lineatus and the first described for the region from the Tietê-Batalha basin.

Parasite biodiversity of fish in coral reefs of the South China Sea is still incompletely explored. We describe here a new species of Neohexostoma (Monogenea: Hexostomatidae) from the gill filaments of the dogtooth tuna Gymnosarda unicolor (Scombridae), collected off Yongshu Reef, South China Sea. Neohexostoma gymnosardae n. sp. is distinguished from its congeners by the following features: (i) haptor clearly marked from body proper by a strongly constricted peduncle, divided in its posterior margin into two symmetrical lobes, (ii) vagina armed with scattered small blunt spines, (iii) eggs tied by their long polar filaments, (vi) esophagus with several lateral diverticula, (v) intestinal ceca unfused and extending into the haptor. We present an analysis of the relationships of this monogenean based on partial 28S rDNA sequences. An identification key for species of Neohexostoma is provided. This is the first member of the genus Neohexostoma known to parasitize a species of Gymnosarda., (© P.-W. Zhu et al., published by EDP Sciences, 2020.)

Diphterostomum brusinae (Zoogonidae) is a digenean species that has been recorded worldwide parasitizing marine fishes. Several species have been synonymized with D. brusinae because they lack conspicuous morphological differences. However, due to the breadth of its geographic distribution and the variety of hosts involved in the life cycles, it is likely to be an assemblage of cryptic species. Diphterostomum flavum n. sp. is described here as a morphologically cryptic relative of D. brusinae, in the fish Pinguipes brasilianus (Pinguipedidae) off the Patagonian coast, Southwestern Atlantic Ocean, and its life cycle is elucidated through morphology and molecular analysis. This species uses the gastropod Buccinanops deformis (Nassariidae) as first and second intermediate host with metacercariae encysting within sporocysts. They also, however, use the polychaete Kinbergonuphis dorsalis (Onuphidae) as second intermediate host. No morphological differences were found between adults of D. flavum n. sp. and D. brusinae; however, the number of penetration glands of the cercariae, a diagnostic feature, differed (9 vs. 3 pairs), as well as the ITS2 sequences for the two species. This work provides morphological and molecular evidence of cryptic diversification among species described as D. brusinae, in which the only clear differences are in larval morphology and host spectrum. The strict specificity to the snail acting as the first intermediate host and the variety of fishes with different feeding habits acting as definitive hosts support the likely existence of multiple cryptic species around the world., (© C. Gilardoni et al., published by EDP Sciences, 2020.)

The edible land snail Cornu aspersum, native to the Mediterranean coastlines of North Africa, is widely distributed on most continents and often invasive in areas where introduction is recent. This species could contribute to the geographic spread of parasites as demonstrated for Brachylaima spp. These cosmopolitan trematodes may represent a threat to human health, like in Australia where Brachylaima cribbi infects humans. In this study, we demonstrate for the first time the occurrence of Brachylaima spp. in two French populations of C. aspersum, Thorigné-Fouillard (Ille-et-Vilaine), and Arçais (Deux-Sèvres), with an overall prevalence of 10.4% (Thorigné-Fouillard) and 73.3% (Arçais), respectively and a metacercarial intensity on average three times higher in Thorigné-Fouillard (37) than in Arçais (11). Cornu aspersum may act as a first and second intermediate host, as demonstrated in Arçais. The morphometrics of metacercariae, particularly the great body length about 2 mm, discriminate our Brachylaima species from those already described in C. aspersum (B. cribbi in Australia, and B. aspersae, B. llobregatensis and B. mascomai in Europe). Molecular analysis, based on 28S and COI, suggests the occurrence of two species in our study, one of which is probably Brachylaima mesostoma, an intestinal parasite of passeriform birds described in Central Europe. We underline the need for further research to identify species of Brachylaima in France and measure the health hazard of consuming field-collected snails., (© C. Gérard et al., published by EDP Sciences, 2020.)

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., (© C. Bouguerche et al., published by EDP Sciences, 2020.)

Paracaesicola n. gen., is erected herein to accommodate a new microcotylid species, Paracaesicola nanshaensis n. sp., collected from the Yongshu Reef, South China Sea. This species is the first monogenean to be recorded from the gills of Paracaesio sordida. The new species is characterized by the following features: (i) haptor short, with clamps arranged in two equal bilateral rows; (ii) testes numerous, arranged in two roughly alternating longitudinal rows, extending into the haptor; (iii) genital atrium armed with 16 robust spines, which are vertically arranged on top of the sausage shaped muscular male copulatory organ; and (iv) single vagina, bottle-shaped, with a distinctly bulbous vaginal atrium. The terminals of the reproductive system discriminate Paracaesicola n. gen. from all other genera in the Microcotylidae. Molecular phylogenetic analyses, based on partial 28S rDNA, places Paracaesicola nanshaensis n. sp. within the microcotylid clade, but its sequence differs from all known available microcotylid sequences., (© Z.-H. Zhou et al., published by EDP Sciences, 2020.)

Two species of Cosmetocleithrum Kritsky, Thatcher & Boeger, 1986 (both new) and two species of Demidospermus Suriano, 1983 (one new) are reported from the gills of the catfish Auchenipterus nuchalis, popularly known as "mapará", from the Tocantins River and tributaries, North Region of Brazil. Cosmetocleithrum berecae n. sp. differs from all other species presently known in the genus by the morphology of the anchors presenting an elongate shaft and short recurved point, a coiled male copulatory organ (MCO) with three rings, and an elongate slender accessory piece with a bifurcated distal end. Cosmetocleithrum nunani n. sp. differs from its congeners by the combination of the following features: (1) Ventral and dorsal anchors with moderately long curved shaft and short point; (2) Hooks with poorly developed thumb; (3) Hook pairs 5 and 6 similar to each other, but morphologically distinct from remaining hook pairs; and (4) MCO coiled, with approximately 1.5 rings. Demidospermus tocantinensis n. sp. is easily distinguished from other species of the genus by presenting an inverted-G-shaped MCO with a median knee-like expansion. Demidospermus osteomystax Tavernari, Takemoto, Lacerda & Pavanelli, 2010 is redescribed based on paratypes and specimens from the gills of A. nuchalis from the Tocantins River, a new host and locality records for this species. The monotypic Paracosmetocleithrum Acosta, Scholz, Blasco-Costa, Alves & Silva, 2017, the only other Neotropical genus reported in siluriforms besides Cosmetocleithrum with species presenting two ribbon-like projections on the posterior margin of the dorsal bar, is considered a junior subjective synonym of Cosmetocleithrum., (© S.C. Cohen et al., published by EDP Sciences, 2020.)

Clarias gariepinus is a fish from North of South Africa and was later introduced in several countries, including Brazil. The present study aimed to describe the first report of Diplostomidae metacercariae in C. gariepinus in Brazil. For this, 30 C. gariepinus were captured in a lake in the city of Campos dos Goytacazes, RJ, Brazil. Fishes were euthanized using freezing and necropsied for collection of parasites. The organs were dissected and analyzed for the presence of parasites that were processed for light and scanning electron microscopy. Trematodes were stained with Semichon's Carmine and Gomori's Trichrome, observed and schematized under a light microscope with image analysis software. A total of 190 trematodes were collected from the gills, suprabranchial organs, heart, stomach, intestinal mesentery, liver and body cavity of the fish. The parasites had a foliaceous body divided by a discrete constriction, without genital primordia, and a holdfast organ present at the posterior region, typical of metacercariae of the family Diplostomidae. It was classified as the 'Diplostomulum' morphotype, based on the morphology of the reserve bladder structure. This is the first report of the metacercariae of Diplostomidae parasitizing C. gariepinus in the Americas. This fish acts as an intermediate or paratenic host of this digenean in Brazil.

Avian trematodes, Urogonimus turdi (Digenea: Leucochloridiidae), were collected from the intestine of wild birds, Zoothera aurea, 2013-2017 in the Daejeon Metropolitan City, Korea. The body was ellipsoidal, attenuated and/or round ends, 1,987-2,120 long and 819-831 µm wide. The oral sucker was subterminal, rounded anteriorly, and 308- 425×351-432 µm in size; the prepharynx and esophagus were almost lacking; pharynx was well-developed, 142- 179×78-170 µm in size; intestine narrow, bifurcating just after pharynx, ascending to the oral sucker before looping posteriorly and terminating near the posterior end; ventral sucker larger, in almost median, 536-673×447-605 µm and approximately 1.5 times larger than oral sucker. A phylogenetic tree constructed with 18S ribosomal RNA showed inter- and intraspecific relationships. Based on these morphological and molecular findings, we report here a U. turdi from White's thrushes in Korea.

A total of 17 species of the genus Bifurcohaptor Jain, 1958 have been reported from two fish families namely Bagridae Bleeker, 1858 (Mystus vittatus (Bloch, 1794), M. tengara (Hamilton, 1822), M. keletius (Valenciennes, 1840), Hemibagrus nemurus (Valenciennes, 1840), Rita rita (Hamilton, 1822) and Sperata seenghala (Sykes, 1839)) and Sisoridae Bleeker, 1858 (Bagarius bagarius (Hamilton, 1822)). Out of these, only two species viz. B. indicus and B. giganticus are found valid in India, parasitizing gills of Mystus spp. and Bagarius sp. Taxonomic studies suggest, present specimen of B. indicus and B. giganticus, both are morphologically close to species described by Jain (1958), except morphometric variations and posses 7 pairs of marginal hooks instead of 6 pairs. Present manuscript delves with the characterization of B. indicus and B. giganticus reported from India, using molecular techniques. Partial mt COI nucleotide sequence based insilico protein analysis and partial 28S and ITS-1 rDNA based phylogenetic analysis, estimated by Neighbour-joining (NJ) and Minimum Evolution (ME) methods revealed that the species of the genus Bifurcohaptor are genetically distinct and valid. The grouping of Bifurcohaptor spp. with other representatives of family Dactylogyridae supports morphology based placement into family Dactylogyridae. Present and previous host-parasite information suggests both Bifurcohaptor spp. are species specialist however, the genus Bifurcohaptor is generalist at generic level.

Hippocrepis hippocrepis is a notocotylid that has been widely reported in capybaras; however, the molluscs that act as intermediate hosts of this parasite remain unknown. Furthermore, there are currently no molecular data available for H. hippocrepis regarding its phylogenetic relationship with other members of the family Notocotylidae. In the present study, we collected monostome cercariae and adult parasites from the planorbid Biomphalaria straminea and in the large intestine of capybaras, respectively, from Belo Horizonte, Minas Gerais, Brazil. We subjected them to morphological and molecular (amplification and sequencing of partial regions of 28S and cox-1 genes) studies. Adult parasites collected from the capybaras were identified as H. hippocrepis and the sequences obtained for both molecular markers showed 100% similarity with monostome cercariae found in B. straminea. The sequences obtained for H. hippocrepis were compared with data available in public databases; analysis revealed this species differs from other notocotylids with available sequences (1.5-3.8% with respect to 28S and 11.4%-13.8% with respect to cox-1). On the phylogenetic analyses, H. hippocrepis appeared to be a distinct lineage in relation to other notocotylids. Some ecological aspects related to the infection of capybaras with H. hippocrepis are briefly discussed., Competing Interests: The authors have declared that no competing interests exist.

The first molecular assessment of phylogenetic relationships of cladorchiid digeneans (superfamily Paramphistomoidea Fischoeder, 1901) from freshwater fishes based on 28S rDNA, ITS2 and cox1 sequences reveals the subfamilies Dadayiinae Fukui, 1929 and Kalitrematinae Travassos, 1933 as non-monophyletic, whereas Dadaytrema Travassos, 1931 represented by three species is monophyletic. Fourteen species of cladorchiids were found in characiform, perciform and siluriform fishes in the Neotropical Region (Brazil and Peru), with numerous new host and geographical records. The first scanning electron micrographs of seven species are presented. Two new species of dadayiine and one new species of kalitrematine paramphistomes are described. Microrchis macrovarium sp. n. from Pimelodella cristata (Müller et Troschel) (type host), Tetranematichthys quadrifilis (Kner) and Pterodoras granulosus (Valenciennes) in Brazil and Peru differs from all three congeners in that the testes are directly tandem, not oblique, and that the ovary, between the caecal ends, is widely separated from the testes. Pronamphistoma philippei sp. n. from Heros sp. in Brazil is distinguished from the type and only species, Pronamphistoma cichlasomae Thatcher, 1992, by the absence of the anterior collar-like expansion present in the type species, the presence of extramural rather than intramural pharyngeal sacs, and the unusual development of the dorsal and ventral exterior circular muscle fibre series in the acetabulum. Pseudocladorchis romani sp. n. from P. granulosus (type host), Brachyplatystoma vaillantii (Valenciennes), Calophysus macropterus (Lichtenstein), Megalodoras uranoscopus (Eigenmann et Eigenmann) and Oxydoras niger (Valenciennes) in Brazil and Peru, is most similar to Pseudocladorchis nephrodorchis Daday, 1907 but differs in the shape of the testes (irregular, versus reniform in the latter species) and the size of the ovary (as large as, or larger than, the testes in the new species). The generic diagnosis of Pronamphistoma Thatcher, 1992 is amended. Dadaytremoides parauchenipteri Lunaschi, 1989 is transferred to Doradamphistoma Thatcher, 1979 as D. parauchenipteri (Lunaschi, 1989) comb. n. based on morphological and molecular evidence.

Synthesium elongatum (Brachycladiidae) is an intestinal digenean described from the finless porpoise (Neophocaena asiaeorientalis) in Japan. Few records of this species exist and there is a remarkable morphological similarity between S. elongatum and S. tursionis, such that a synonymy between the species has been suggested previously. However, no morphological and/or molecular analysis has been carried out to clarify the taxonomic status of S. elongatum. In this study, we collected specimens of Synthesium sp. from N. asiaeorientalis in western Japan. The specimens possess lobed testes within the third quarter of the body, a round ovary, and vitellaria extending to level of uterine field, which are diagnostic characters for both S. elongatum and S. tursionis. They were morphologically identified to S. elongatum or S. tursionis due to the fact that the available morphometric data for both species overlap remarkably. A molecular analysis of the mitochondrial ND3 gene showed that the pairwise nucleotide distances between these specimens and S. tursionis were small, and phylogenetic analysis showed that these specimens and S. tursionis are in the same clade. Therefore, it was indicated that S. elongatum and S. tursionis are the same species and, consequently, S. elongatum is a synonym of S. tursionis.

A 46-year old man visited our outpatient clinic with complaint of foreign body sensation in throat after consuming raw freshwater fish 5 days ago. Laryngoscopic examination revealed a motile worm attached on posterior pharyngeal wall. The worm was removed using biopsy forceps under transnasal endoscopy and evidently identified as Clinostomum complanatum after microscopic examination. Patient's subjective foreign body sensation of throat and hyperemia of laryngeal mucosa remained for approximately 2 weeks post-removal, which were eventually resolved after administration of non-steroidal anti-inflammatory drug and anti-refluxant drug for 2 weeks. Treatment was ended at three weeks since the first visit. C. complanatum infections in humans are rare, and only four cases have been reported in Korea. Symptoms resembling pharyngitis or laryngitis occurs by consumption of raw, infected freshwater fish and treatment is done by mechanically removing the parasite.

A stray female cat of unknown age, presenting bright red watery diarrhea, was submitted to the Animal and Plant Quarantine Agency for diagnosis. In the small intestines extracted from the necropsied cat, numerous white oval-shaped organisms were firmly embedded in the mucosa and there was thickening of intestinal wall. Histopathological analysis revealed severe necrotizing enteritis, together with atrophied intestinal villi, exfoliated enterocytes, and parasitic worms. Recovered worms were identified as Pharyngostomum cordatum by morphological observation and genetic analysis. Although P. cordatum is known to occur widely in Korea, this is the first clinical description of an infection by P. cordatum causing severe feline enteritis.