Small intestinal flukes of the genus Metagonimus (Digenea: Heterophyidae) in Europe and the Middle East: A review of parasites with zoonotic potential.

The heterophyid trematode Metagonimus romanicus (Ciurea, 1915) (Digenea) is redescribed on the basis of type material from domestic dogs (Canis familiaris) in Romania, vouchers from experimentally infected cats (Felis catus) and adults recovered from golden hamsters (Mesocricetus auratus) infected with metacercariae from scales of chub (Squalius cephalus) and common nase (Chondrostoma nasus) (Cypriniformes: Leuciscidae) in Hungary. This trematode, endemic to Europe and neighbouring regions (northwestern Türkiye), was previously misidentified as M. yokogawai (Katsurada, 1912), a zoonotic parasite of humans in East Asia. However, the two species differ considerably both genetically and morphologically, e.g., in the position of the ventral sucker, the presence of the prepharynx, the anterior extent of the vitelline follicles and the posterior extent of the uterus. Metagonimus ciureanus (Witenberg, 1929) (syn. Dexiogonimus ciureanus Witenberg, 1929), described from domestic cats and dogs in Israel, is a valid species distributed in the Middle East and Transcaucasia, which is also confirmed by molecular data. It differs from all Metagonimus species, including M. romanicus, in having symmetrical testes instead of the oblique testes of the other congeners. The zoonotic significance of M. romanicus and M. ciureanus is unclear, but appears to be low in Europe, mainly because raw or undercooked, whole fish with scales are generally not consumed. Accidental infection of fishermen by metacercariae in the scales when cleaning fish is more likely, but has never been reported. Remains of cyprinoids with scales infected with metacercariae of Metagonimus spp. can be an important natural source of infection for dogs, cats, and other carnivores, which can serve as a reservoir for these parasites.

Title: Petits trématodes intestinaux du genre Metagonimus (Digenea : Heterophyidae) en Europe et au Moyen-Orient : revue de parasites à potentiel zoonotique. Abstract: Le trématode Heterophyidae Metagonimus romanicus (Ciurea, 1915) (Digenea) est redécrit sur la base de matériel type provenant de chiens domestiques (Canis familiaris) en Roumanie, de vouchers issus de chats (Felis catus) infectés expérimentalement et d'adultes collectés chez des hamsters dorés (Mesocricetus auratus) infectés par des métacercaires provenant d'écailles de chevesne commun (Squalius cephalus) et de nase commun (Chondrostoma nasus) (Cypriniformes : Leuciscidae) de Hongrie. Ce trématode, endémique d'Europe et des régions voisines (nord-ouest de la Turquie), avait été précédemment identifié à tort comme étant M. yokogawai (Katsurada, 1912), un parasite zoonotique des humains en Asie de l'Est. Cependant, les deux espèces diffèrent considérablement sur le plan génétique et morphologique, par exemple par la position de la ventouse ventrale, la présence du prépharynx, l'étendue antérieure des follicules vitellins et l'étendue postérieure de l'utérus. Metagonimus ciureanus (Witenberg, 1929) (syn. Dexiogonimus ciureanus Witenberg, 1929), décrite chez des chats et des chiens domestiques en Israël, est une espèce valide répartie au Moyen-Orient et en Transcaucasie, ce qui est également confirmé par des données moléculaires. Cette espèce diffère de toutes les espèces de Metagonimus, y compris M. romanicus, par ses testicules symétriques au lieu des testicules obliques des autres congénères. L'importance zoonotique de M. romanicus et M. ciureanus n'est pas claire, mais semble faible en Europe, principalement parce que les poissons n'y sont généralement pas consommés crus ou insuffisamment cuits et entiers avec les écailles. L'infection accidentelle des pêcheurs par des métacercaires présents dans les écailles lors du nettoyage du poisson est plus probable mais n'a jamais été signalée. Les restes de poissons cyprinoïdes avec les écailles, infectés par des métacercaires de Metagonimus spp. peuvent être une source naturelle importante d'infection pour les chiens, les chats et autres carnivores, qui peuvent servir de réservoir à ces parasites.

Gaps in parasitological research in the molecular era.

We live in the age of molecular biology and '-omics', and molecular methods have opened up unimagined possibilities for biological research, including parasitology. However, too one-sided a focus on new approaches can lead to major gaps as less 'cool' topics are neglected. Selected areas of research are briefly discussed to highlight the gaps caused by the current excessive focus on molecular and '-omics' methods. It is crucial to combine both 'classical' and modern methods without neglecting the complexity of the interactions of parasites with their hosts and the environment (One Health concept), which is even more urgent in today's rapidly changing world. Parasitologists should be more involved in field studies and multidisciplinary assessment of parasites.

Discovering high species diversity of Ophiotaenia tapeworms (Cestoda: Proteocephalidae) of watersnakes (Colubridae) in North America.

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.

Human diphyllobothriosis in Taiwan: A review of cases and molecular evidence of Dibothriocephalus nihonkaiensis.

Diphyllobothriosis is an infectious disease caused by the consumption of raw freshwater or marine fish containing larvae of broad tapeworms (Diphyllobothriidae). In the present study, we critically reviewed all cases of human diphyllobothriosis reported from Taiwan, including unpublished reports from hospitals in Taipei. Genotyping based on mitochondrial DNA marker (cox1) confirmed that two of the recent cases were caused by Dibothriocephalus nihonkaiensis, which is not native to Taiwan and was probably imported with Pacific salmon infected with larvae of D. nihonkaiensis. The causative species previously reported in Taiwan could not be definitively confirmed. However, considering the distribution of Dibothriocephalus latus, which is not endemic in Taiwan, past cases diagnosed as D. latus are questionable.

Zoonotic Marine Nematode Infection of Fish Products in Landlocked Country, Slovakia.

Fish products in Slovakia have been heavily infected with Anisakis spp. larvae, which causes human anisakiasis. We found larvae in all tested samples of frozen Atlantic herring. Anisakid allergen t-Ani s7 testing revealed 2 positive cases in humans, signaling need for health authorities to closely monitor zoonotic marine parasites, even in inland areas.

TAPEWORMS (CESTODA: OPHIOTAENIA) FROM THE NORTHERN COTTONMOUTH (AGKISTRODON PISCIVORUS).

The northern cottonmouth, Agkistrodon piscivorus (Lacépède, 1789) (Viperidae: Crotalinae), occurs in the southeastern United States and is one of the few semiaquatic vipers in the world. Three proteocephalid tapeworms of the genus OphiotaeniaLa Rue, 1911 (Cestoda), have been described from this venomous snake. A critical evaluation of type specimens and tapeworms recently collected from A. piscivorus has revealed that only 2 species, Ophiotaenia marenzelleri (Barrois, 1898) and Ophiotaenia grandisLa Rue, 1911 (new synonym Ophiotaenia agkistrodontis [Harwood, 1933]), are specific parasites of this crotaline viper; both species are redescribed here. Ophiotaenia grandis was originally described from a mixture of 2 species: 'true' O. grandis, which is indistinguishable from O. agkistrodontis described 22 yr later and thus considered to be a junior synonym of O. grandis, and O. marenzelleri. Ophiotaenia marenzelleri, also reported from the pygmy rattlesnake, Sistrurus miliarius (Linnaeus, 1766), is a larger tapeworm with a massive scolex, a large cirrus sac, a very large, annular vaginal sphincter near the equatorial gonopore, and an oval, 3-layered embryophore surrounded by a nearly square hyaline outer membrane. Ophiotaenia grandis is much smaller and more slender and has a narrower scolex, a smaller cirrus sac and vaginal sphincter, a 2-layered embryophore, and a distinctly pre-equatorial gonopore. In addition to O. marenzelleri and O. grandis, other Ophiotaenia species typical of semiaquatic snakes (Colubridae: Natricinae) may be present in the northern cottonmouth, which serves only as a postcyclic or occasional host. There is also a tabular summary of 18 species of Ophiotaenia from semiaquatic snakes worldwide, with information on their hosts, distribution, and taxonomically important characters, including key measurements.

Invasive parasites and global change: Evidence for the recent and rapid spillover of a potential pathogen of tilapias with a complex, three-host life cycle.

Biological invasions pose a serious threat to local flora and fauna and have negative impacts on ecosystems. Invasive parasites can also cause severe losses in aquaculture. In this article, we provide evidence of the recent spillover of an African parasite with a complex, three-host life cycle that has rapidly and successfully established itself in the Middle East, most likely due to the recent migration of its final hosts (great cormorant) from Africa. This case of parasite introduction into a country with intensive aquaculture is also important from an economic point of view, since large (up to 2 cm long) larvae of this parasite, the cyclophyllidean tapeworm Amirthalingamia macracantha (Cestoda) localised in the liver, can be pathogenic to their fish hosts, including farmed and wild fish, as shown by our histopathological examination of heavily infected fish. Since its first detection in Israel in November 2020, the parasite has spread rapidly and is currently found in both migratory (great cormorant, Phalacrocorax carbo) and non-migratory birds (pygmy cormorant, Microcarbo pygmaeus), as well as in fish intermediate hosts, including farmed tilapia in several farms in Israel and wild cichlids. There are numerous examples of the spillover of introduced parasites, including those that parasitise fish of commercial importance, but have a direct life cycle or use only a single intermediate host. Tilapines are the second most important group of farmed fish in the world after carps and are produced mainly in Southeast Asia, Central and South America. The global spread of great cormorants and the early evidence that pygmy cormorant may also harbour A. macracantha pose the risk of further spread of this invasive parasite to other countries and areas. In addition, global warming and reductions in foraging and resting areas near these waters may allow the parasite to complete its life cycle in new hosts.

The evolution of endoparasitism and complex life cycles in parasitic platyhelminths.

Within flatworms, the vast majority of parasitism is innate to Neodermata, the most derived and diversified group of the phylum Platyhelminthes.1,2 The four major lineages of Neodermata maintain various combinations of life strategies.3 They include both externally (ecto-) and internally feeding (endo-) parasites. Some lineages complete their life cycles directly by infecting a single host, whereas others succeed only through serial infections of multiple hosts of various vertebrate and invertebrate groups. Food sources and modes of digestion add further combinatorial layers to the often incompletely understood mosaic of neodermatan life histories. Their evolutionary trajectories have remained molecularly unresolved because of conflicting evolutionary inferences and a lack of genomic data.4 Here, we generated transcriptomes for nine early branching neodermatan representatives and performed detailed phylogenomic analyses to address these critical gaps. Polyopisthocotylea, mostly hematophagous ectoparasites, form a group with the mostly hematophagous but endoparasitic trematodes (Trematoda), rather than sharing a common ancestor with Monopisthocotylea, ectoparasitic epithelial feeders. Phylogenetic placement of the highly specialized endoparasitic Cestoda alters depending on the model. Regardless of this uncertainty, this study brings an unconventional perspective on the evolution of platyhelminth parasitism, rejecting a common origin for the endoparasitic lifestyle intrinsic to cestodes and trematodes. Instead, our data indicate that complex life cycles and invasion of vertebrates' gut lumen, the hallmark features of these parasites, evolved independently within Neodermata. We propose the demise of the traditionally recognized class Monogenea and the promotion of its two subclasses to the class level as Monopisthocotyla new class and Polyopisthocotyla new class.

ERECTION OF BOTHRIOCESTUS N. GEN. (CESTODA: BOTHRIOCEPHALIDEA) AND REDESCRIPTION OF BOTHRIOCESTUS CUSPIDATUS (COOPER, 1917) (SYN. BOTHRIOCEPHALUS CUSPIDATUS) FROM WALLEYE, SANDER VITREUS, (PERCIFORMES: PERCIDAE) IN NORTH AMERICA.

Based on previous molecular phylogenetic analyses, Bothriocestus n. gen. is erected to accommodate bothriocephalid tapeworms that have an elongate scolex, a well-developed apical disc, and a narrow neck region, parasitize freshwater fishes in the Holarctic, and were previously placed in the polyphyletic genus Bothriocephalus Rudolphi, 1808 (Cestoda: Bothriocephalidea). Bothriocestus claviceps (Goeze, 1782) n. comb., a parasite of eels (Anguilla spp.) in the Holarctic region, is designated as the type species. Another species of the new genus, Bothriocestus cuspidatus (Cooper, 1917) (syn. Bothriocephalus cuspidatusCooper, 1917) is redescribed from type and voucher specimens, and new material from the type host, the walleye, Sander vitreus (Mitchill, 1818) (Perciformes: Percidae), in Manitoba and Ontario (where the type locality is located) (Canada) and in New York state and Wisconsin. Bothriocestus cuspidatus of S. vitreus is characterized primarily by the possession of a narrow, long strobila (total length up to 18 cm) composed of distinctly craspedote, trapezoidal proglottids, with primary, secondary, and tertiary proglottids differing in size, and by an arrow-shaped (=cuspidatus) scolex that is distinctly broader than the first proglottids, widest near the base in lateral view and gradually becoming broader toward the anterior end in dorsoventral view. A "dwarf" form of B. cuspidatus (total length of 9-27 mm) from Johnny darter, Etheostoma nigrum Rafinesque, 1820, and tessellated darter, Etheostoma olmstedi Storer, 1842 (both Percidae: Etheostominae), is also characterized morphologically in the present paper.

First insight in element localisation in different body parts of the acanthocephalan Dentitruncus truttae using TEM and NanoSIMS.

Acanthocephalans, intestinal parasites of vertebrates, are characterised by orders of magnitude higher metal accumulation than free-living organisms, but the mechanism of such effective metal accumulation is still unknown. The aim of our study was to gain new insights into the high-resolution localization of elements in the bodies of acanthocephalans, thus taking an initial step towards elucidating metal uptake and accumulation in organisms under real environmental conditions. For the first time, nanoscale secondary ion mass spectrometry (NanoSIMS) was used for high-resolution mapping of 12 elements (C, Ca, Cu, Fe, N, Na, O, P, Pb, S, Se, and Tl) in three selected body parts (trunk spines, inner part of the proboscis receptacle and inner surface of the tegument) of Dentitruncus truttae, a parasite of brown trout (Salmo trutta) from the Krka River in Croatia. In addition, the same body parts were examined using transmission electron microscopy (TEM) and correlated with NanoSIMS images. Metal concentrations determined using HR ICP-MS confirmed higher accumulation in D. truttae than in the fish intestine. The chemical composition of the acanthocephalan body showed the highest density of C, Ca, N, Na, O, S, as important and constitutive elements in living cells in all studied structures, while Fe was predominant among trace elements. In general, higher element density was found in trunk spines and tegument, as body structures responsible for substance absorption in parasites. The results obtained with NanoSIMS and TEM-NanoSIMS correlative imaging represent pilot data for mapping of elements at nanoscale resolution in the ultrastructure of various body parts of acanthocephalans and generally provide a contribution for further application of this technique in all parasite species.

A global survey of tapeworms (Cestoda: Proteocephalidae) of 'true' frogs (Amphibia: Ranidae), including a tabulated list of all proteocephalids parasitising amphibians.

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.

Mining various genomic resources to resolve old alpha-taxonomy questions: A test of the species hypothesis of the Proteocephalus longicollis species complex (Cestoda: Platyhelminthes) from salmonid fishes.

High-throughput sequencing strategies became commonly employed to study non-model parasites, but the corresponding genomes and transcriptomes were seldom mined following the original publication. Similar to the data generated with genome skimming techniques based on shallow-depth shotgun genomes, various genomic and transcriptomic resources can be screened for useful molecular phylogenetic markers traditionally characterised with Sanger sequencing. Here, we provide an example of a strategy using reduced-representation genomic as well as transcriptomic data to obtain broad insights into the molecular diversity of the cestode Proteocephalus longicollis, a common parasite of salmonids distributed throughout the Holarctic region. We extract popular mitochondrial and nuclear ribosomal markers from various genomic resources for hundreds of parasite specimens from multiple European whitefish populations and compare those with Proteocephalus representatives from other species of salmonids and various geographical regions. In contrast with the previous morphology-based assessments, molecular phylogeny reveals a high degree of genetic divergence between Proteocephalus isolates from different salmonids, contrastingly low genetic differentiation within the parasite's populations hosted by the European whitefish (Coregonus lavaretus species complex), and a sister species relationship of Proteocephalus from European whitefish and Proteocephalus percae, a parasite of European perch (Perca fluviatilis). Proteocephalus spp. from North American lake whitefish, brown trout and Arctic charr each formed clearly distinct lineages. These results advance our understanding of the interrelationships of the Proteocephalus-aggregate, a well-recognized clade of Holarctic freshwater fish proteocephalids, and support resurrection of some of the nominal species of Proteocephalus, including Proteocephalus exiguus La Rue, 1911 from North American coregonids and Proteocephalus fallax La Rue, 1911 from European C. lavaretus, reserving Proteocephalus longicollis (Zeder, 1800) exclusively for parasites of Salmo trutta.

Historical dispersal and host-switching formed the evolutionary history of a globally distributed multi-host parasite - The Ligula intestinalis species complex.

Studies on parasite biogeography and host spectrum provide insights into the processes driving parasite diversification. Global geographical distribution and a multi-host spectrum make the tapeworm Ligula intestinalis a promising model for studying both the vicariant and ecological modes of speciation in parasites. To understand the relative importance of host association and biogeography in the evolutionary history of this tapeworm, we analysed mtDNA and reduced-represented genomic SNP data for a total of 139 specimens collected from 18 fish-host genera across a distribution range representing 21 countries. Our results strongly supported the existence of at least 10 evolutionary lineages and estimated the deepest divergence at approximately 4.99-5.05 Mya, which is much younger than the diversification of the fish host genera and orders. Historical biogeography analyses revealed that the ancestor of the parasite diversified following multiple vicariance events and was widespread throughout the Palearctic, Afrotropical, and Nearctic between the late Miocene and early Pliocene. Cyprinoids were inferred as the ancestral hosts for the parasite. Later, from the late Pliocene to Pleistocene, new lineages emerged following a series of biogeographic dispersal and host-switching events. Although only a few of the current Ligula lineages show narrow host-specificity (to a single host genus), almost no host genera, even those that live in sympatry, overlapped between different Ligula lineages. Our analyses uncovered the impact of historical distribution shifts on host switching and the evolution of host specificity without parallel host-parasite co-speciation. Historical biogeography reconstructions also found that the parasite colonized several areas (Afrotropical and Australasian) much earlier than was suggested by only recent faunistic data.

Fish parasites (special issue).

Fish (Elasmobranchia and Actinopterygii) inhabit the majority of aquatic habitats globally. They are crucial for human nutrition but they may be negatively affected by parasitic protists and metazoan parasites. Fish parasites are also an extraordinary group of animals because of their ecological and evolutionary importance and unique adaptations to parasitism. They also play a key role in ecosystem functioning. In the present special issue, 13 review and research articles on major groups of fish parasites are provided to document the current advancement in our understanding of different aspects of their biology, ecology and associations with their fish hosts. The existing gaps in our knowledge of these peculiar animals are mapped and future trends in their research outlined.

The first new species of European Ascocotyle Looss, 1899 (Digenea: Heterophyidae) described in more than half a century.

Ascocotyle (Phagicola) trentinii n. sp. is described based on adults from experimentally infected ducklings (Anas platyrhynchos domesticus) fed with metacercariae from the visceral serosa of the Mediterranean banded killifish, Aphanius fasciatus (Cyprinodontiformes: Aphaniidae), from coastal lagoons in northeastern Italy (Emilia-Romagna Region). The new species is placed into the subgenus Phagicola because of the presence of a single row of circumoral spines, vitelline follicles being confined between the ventral sucker and testes, and uterine loops not reaching anterior to the ventral sucker. Ascocotyle (P.) trentinii n. sp. differs from other members of the subgenus Phagicola, as well as other species of Ascocotyle, by the number (27-33) of circumoral spines which are 13.5-17 µm long and 3.5-5 µm wide, and by the morphology of a gonotyl which is composed of about 8 large refractile pockets. The occurrence of metacercariae in A. fasciatus indicates that the life cycle of the new species is completed in brackish water lagoons. It is the fourth species of Ascocotyle described in Europe and may be endemic to the Mediterranean region because its second (fish) intermediate host is endemic to this region.

A NEW SPECIES OF BOTHRIOCEPHALUS (CESTODA: BOTHRIOCEPHALIDAE) FROM LEPOMIS SPP. (ACTINOPTERYGII: CENTRARCHIDAE) IN NORTH AMERICA.

Based on morphological and molecular data, a new species of tapeworm, Bothriocephalus kupermani n. sp., is described from pumpkinseed, Lepomis gibbosus (Linnaeus, 1758) (type host), and green sunfish, L. cyanellus Rafinesque, 1819 (Actinopterygii: Centrarchidae) in the United States. The new species differs from its North American congeners mainly in the shape of its scolex, which is almost rectangular in dorsoventral and lateral views (nearly parallel margins in the bothrial region), with shallow, wide bothria, and by an extensive field of vitelline follicles that are also variably present, albeit more sparsely, medially in the cortex of the ovarian and uterine areas. This tapeworm appears to be typically associated with Lepomis sunfishes. Examination of museum specimens of Bothriocephalus species from Lepomis spp. indicates that the tapeworm also parasitizes bluegill, Lepomis macrochirus, longear sunfish, Lepomis megalotis, and that previous records of Bothriocephalus cuspidatus and Bothriocephalus claviceps in these fishes are of the new species, B. kupermani.

Fish tapeworms (Cestoda) in the molecular era: achievements, gaps and prospects.

The tapeworms of fishes (Chondrichthyes and Actinopterygii) account one-third (1670 from around 5000) of the total tapeworm (Platyhelminthes: Cestoda) species diversity. In total 1186 species from 9 orders occur as adults in elasmobranchs (sharks, rays and chimaeras), and 484 species from 8 orders mature in ray-finned fishes (referred to here as teleosts). Teleost tapeworms are dominated by freshwater species (78%), but only 3% of elasmobranch tapeworms are known from freshwater rays of South America and Asia (Borneo). In the last 2 decades, vast progress has been made in understanding species diversity, host associations and interrelationships among fish tapeworms. In total, 172 new species have been described since 2017 (149 from elasmobranchs and 23 from teleosts; invalidly described taxa are not included, especially those from the Oriental region). Molecular data, however, largely limited to a few molecular markers (mainly 28S rDNA, but also 18S and cox1), are available for about 40% of fish tapeworm species. They allowed us to significantly improve our understanding of their interrelationships, including proposals of a new, more natural classification at the higher-taxonomy level (orders and families) as well as at the lower-taxonomy level (genera). In this review, we summarize the main advances and provide perspectives for future research.

Cercariae of a Bird Schistosome Follow a Similar Emergence Pattern under Different Subarctic Conditions: First Experimental Study.

The emergence of cercariae from infected mollusks is considered one of the most important adaptive strategies for maintaining the trematode life cycle. Short transmission opportunities of cercariae are often compensated by periodic daily rhythms in the cercarial release. However, there are virtually no data on the cercarial emergence of bird schistosomes from freshwater ecosystems in northern latitudes. We investigated the daily cercarial emergence rhythms of the bird schistosome Trichobilharzia sp. "peregra" from the snail host Radix balthica in a subarctic lake under both natural and laboratory seasonal conditions. We demonstrated a circadian rhythm with the highest emergence during the morning hours, being seasonally independent of the photo- and thermo-period regimes of subarctic summer and autumn, as well as relatively high production of cercariae at low temperatures typical of northern environments. These patterns were consistent under both field and laboratory conditions. While light intensity triggered and prolonged cercarial emergence, the temperature had little effect on cercarial rhythms but regulated seasonal output rates. This suggests an adaptive strategy of bird schistosomes to compensate for the narrow transmission window. Our results fill a gap in our knowledge of the transmission dynamics and success of bird schistosomes under high latitude conditions that may serve as a basis for elucidating future potential risks and implementing control measures related to the spread of cercarial dermatitis due to global warming.