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.

• Keywords
• DNA sequencing, Distribution, elasmobranchs, host associations, pathogens, phylogenetic relationships, ray-finned fish, species diversity, taxonomy,
• MeSH
• Cestoda * genetics   MeSH
• Cestode Infections * epidemiology   veterinary   MeSH
• Diphyllobothrium * MeSH
• Elasmobranchii * MeSH
• Phylogeny MeSH
• Fish Diseases * epidemiology   MeSH
• Fishes MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Review MeSH

The tapeworm order Bothriocephalidea Kuchta, Scholz, Brabec & Bray, 2008 (Platyhelminthes: Eucestoda), which has until recently formed part of the suppressed order Pseudophyllidea, is revised. Four new genera, namely Andycestus n. g. [type- and only species A. abyssmus (Thomas, 1953) n. comb.], Plicocestus n. g. [type- and only species P. janickii (Markowski, 1971) n. comb.] (both Bothriocephalidae), Mesoechinophallus n. g. [type-species M. hyperogliphe (Tkachev, 1979) n. comb.; other species M. major (Takao, 1986) n. comb. (Echinophallidae)] and Kimocestus n. g. [type- and only species K. ceratias (Tkachev, 1979) n. comb.] (Triaenophoridae) are proposed. Parabothriocephaloides Yamaguti, 1934, Penetrocephalus Rao, 1960 and Tetracampos Wedl, 1861 are resurrected as valid genera, whereas Alloptychobothrium Yamaguti, 1968 (newly synonymised with Plicatobothrium Cable & Michaelis, 1967), Capooria Malhotra, 1985 and Coelobothrium Dollfus, 1970 (syns of Bothriocephalus Rudolphi 1808), Fissurobothrium Roitman, 1965 (syn. of Bathybothrium Lühe, 1902), Paratelemerus Gulyaev, Korotaeva & Kurochkin, 1989 (syn. of Parabothriocephaloides Yamaguti, 1934) and Tetrapapillocephalus Protasova & Mordvinova, 1986 (syn. of Oncodiscus Yamaguti, 1934) are considered to be invalid. A recently erected genus, Dactylobothrium Srivastav, Khare & Jadhav, 2006, and its type- and only species, D. choprai Srivastav, Khare & Jadhav, 2006, are considered to be unrecognisable, because their descriptions contain a number of obvious errors and also indicate that a mixture of several taxa, probably of at least two cestode orders, were studied. Parabothriocephaloides wangi nom. nov. is proposed for Parabothriocephalus psenopsis Wang, Liu & Yang, 2004 from Psenopsis anomala in China in order to avoid a secondary homonymy. All 46 genera considered to be valid are revised, with their generic diagnoses amended on the basis of a critical examination of extensive museum and newly collected specimens. Despite apparent paraphyly or polyphyly of some bothriocephalidean families, especially the Triaenophoridae, the current classification restricted to four families, proposed by Bray et al. (1994), namely the Bothriocephalidae, Echinophallidae, Philobythiidae and Triaenophoridae, is provisionally retained with slight modifications until more molecular data on most genera are available. Keys to families and genera are provided.

• MeSH
• Cestoda classification   ultrastructure   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Research Support, U.S. Gov't, Non-P.H.S. MeSH

This paper provides the first ultrastructural data on the spermatozoon of a pseudophyllidean cestode belonging to the family Echinophallidae, Paraechinophallus japonicus, parasitizing deep-sea fish Psenopsis anomala from Japan. The mature spermatozoon contains two axonemes of the 9+"1" trepaxonematan pattern, nucleus, cortical microtubules, and electron-dense granules. Its anterior extremity exhibits a single electron-dense crested body. One of the most interesting features of the mature spermatozoon is the presence of a ring of tubular structures encircling the axoneme. This character, previously reported in species of the families Bothriocephalidae and Triaenophoridae, may be unique to pseudophyllideans of the group "Bothriocephalidea", thus supporting the assumption that the order Pseudophyllidea is formed in fact by two unrelated clades; "Bothriocephalidea" and "Diphyllobothriidea". The posterior extremity contains the posterior part of the nucleus and the disorganized axoneme--a character previously not described in the Eucestoda.

• MeSH
• Cestoda classification   ultrastructure   MeSH
• Echinococcus ultrastructure   MeSH
• Phylogeny MeSH
• Seawater MeSH
• Perciformes parasitology   MeSH
• Spermatozoa ultrastructure   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Spermiogenesis and ultrastructure of the spermatozoon of the pseudophyllidean cestode Triaenophorus nodulosus (Pallas, 1781), a parasite of pike Esox lucius, has been studied by transmission electron microscopy. Spermiogenesis involves firstly the formation of a zone of differentiation with two centrioles associated with striated roots, and an intercentriolar body between them, subsequent growth of the two flagella of unequal length, and a formation of a median cytoplasmic process exhibiting patches of dense material. The nucleus penetrates into spermatid body after flagellar rotation and proximo-distal fusion has started. The mature spermatozoon of T. nodulosus is filiform and contains two axonemes of 9+"1" pattern of the Trepaxonemata, nucleus, cortical microtubules parallel to the spermatozoon axis, and electron-dense granules. The anterior extremity of the gamete contains a single centriole surrounded by numerous electron-dense tubular structures exhibiting spiral arrangement and giving rise to lateral projections, which correspond to the crested body. When the crested body disappears, the spiral pattern of electron-dense tubular structures is changed into a ring, persisting until the centriole of the second axonemes appears. This structure of the crested body of T. nodulosus is unique among the Eucestoda.

• MeSH
• Models, Biological MeSH
• Cell Nucleus ultrastructure   MeSH
• Centrioles ultrastructure   MeSH
• Cestoda physiology   MeSH
• Cytoplasm ultrastructure   MeSH
• Flagella ultrastructure   MeSH
• Microtubules ultrastructure   MeSH
• Morphogenesis physiology   MeSH
• Organelles ultrastructure   MeSH
• Spermatogenesis physiology   MeSH
• Spermatozoa ultrastructure   MeSH
• Microscopy, Electron, Transmission MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Vitellogenesis in two spathebothriidean tapeworms, dixenous adult Cyathocephalus truncatus and monoxenous progenetic Diplocotyle olrikii, has been examined using transmission electron microscopy and cytochemical staining with periodic acid-thiosemicarbazide-silver proteinate for glycogen. Each vitelline follicle consists of vitellocytes at various stages of development and one irregularly shaped interstitial cell. Projections of the interstitial cell enclose the vitellocytes and extend as a cytoplasmic sheath on the follicular periphery. An outer thin fibrous layer (= extracellular lamina) covers the cytoplasmic sheath in C. truncatus, but lacks in D. olrikii. Maturing and mature vitellocytes contain vitelline material in the form of single small shell globules that gradually fuse and give rise to the large shell globule clusters. Morphology of shell globule clusters differs slightly in both species. In addition, single "lamellar" granules are present in the cytoplasm of vitellocytes of C. truncatus, but not in D. olrikii. Both electron lucent and electron dense lipid droplets are present in the vitellocytes of C. truncatus, whereas only electron dense lipids occur in D. olrikii. A single lipid droplet turns up occasionally in the nuclei of some of the vitellocytes of C. truncatus. The ultrastructural features of vitellogenesis in spathebothriideans resemble those reported previously in "lower" cestodes, especially in pseudophyllideans.

• MeSH
• Cestoda chemistry   physiology   ultrastructure   MeSH
• Glycogen analysis   MeSH
• Oocytes chemistry   physiology   ultrastructure   MeSH
• Periodic Acid-Schiff Reaction MeSH
• Microscopy, Electron, Transmission methods   MeSH
• Vitellogenesis physiology   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Names of Substances
• Glycogen MeSH

The fine structure of the mature spermatozoon of the corallobothriine tapeworm Corallobothrium solidum Fritsch, 1886 (Cestoda: Proteocephalidea) from the electric catfish Malapterurus electricus from the Nile River in Egypt was studied by transmission electron microscopy for the first time. The filiform spermatozoon of C. solidum contains two axonemes of unequal length and a typical 9 + "1" trepaxonematan pattern. A single helicoidal crested body (30-200 nm thick) is localized at the anterior extremity of the gamete. The cortical microtubules line the periphery of the cell, largely parallel to the long axis of the spermatozoon and exhibiting signs of twisting at the beginning of region II. The nucleus, in the form of an electron-dense (largely in gametes of testes) and/or fibrous cord (largely in gametes from male reproductive ducts and seminal vesicle), coils in a spiral through the middle part (region III) of the spermatozoon. The cytoplasm contains electron-dense granules in regions II, III and partly in region IV. The cytoplasm of some spermatozoa exhibits an apparently higher electron-density at the end of the nucleated region (III), and continuously toward the middle part of region IV. The anterior and posterior extremities of the spermatozoa have a single axoneme. The ultrastructural features of the mature spermatozoon of C. solidum mostly coincide with those of the spermatozoon of other proteocephalideans, especially the gangesiine Electrotaenia malopteruri parasitizing the same host.

• MeSH
• Cestoda ultrastructure   MeSH
• Cestode Infections parasitology   veterinary   MeSH
• Fish Diseases parasitology   MeSH
• Spermatozoa ultrastructure   MeSH
• Catfishes parasitology   MeSH
• Microscopy, Electron, Transmission MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Phylogenetic analysis of sequences of the ITS-2 rRNA genes of 20 samples of pseudophyllidean cestodes of the family Diphyllobothriidae (Ligula, Digramma, Diphyllobothrium, and Schistocephalus) from different hosts and geographical regions revealed that: (1) the inclusion of ligulids (previously family Ligulidae) to the Diphyllobothriidae is correct; (2) Schistocephalus appears as the most basal taxon of the Diphyllobothriidae, well separated from Ligula and Digramma, thus supporting the validity of Schistocephalinae Dubinina, 1962; (3) Digramma belonged with samples of Ligula, thus suggesting its invalidity as a genus; and (4) isolates of Ligula, presumably belonging to Ligula intestinalis, are paraphyletic, indicating that this species may represent a complex of separate taxa. Our results indicate the necessity for a taxonomic revision of the family Diphyllobothriidae.

• MeSH
• Cestoda classification   genetics   MeSH
• Cestode Infections parasitology   veterinary   MeSH
• Diphyllobothrium classification   genetics   MeSH
• Phylogeny * MeSH
• DNA, Ribosomal Spacer analysis   genetics   MeSH
• Molecular Sequence Data MeSH
• Fish Diseases parasitology   MeSH
• Fishes parasitology   MeSH
• Sequence Analysis, DNA * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Names of Substances
• DNA, Ribosomal Spacer MeSH

The fine structure of the mature spermatozoon of the tapeworm Electrotaenia malopteruri (Fritsch, 1886), a member of a supposedly primitive group of proteocephalidean tapeworms (Proteocephalidae: Gangesiinae), was studied by transmission electron microscopy for the first time. The mature spermatozoon of E. malopteruri is filiform, tapers at both extremities, and contains two axonemes of 9+"1" trepaxonematan type. A helicoidal crested body (60-150 nm thick) is present at the anterior extremity of the gamete. The twisting cortical microtubules (CM) line the periphery of the spermatozoon continuously. The nucleus, a fine cord of moderately condensed chromatin, occupies the middle part (region III) of the spermatozoon. The slightly electron-dense cytoplasm contains electron-dense granules in regions II and III and becomes more electron-dense at the end of region IV. The anterior and posterior extremities of the spermatozoon contain a single axoneme. Two modes of disorganisation of the axoneme at the posterior end of the mature spermatozoa are described for the first time in cestodes. The present data also indicate that the sperm ultrastructure of E. malapteruri shows some characters typical for onchobothriid tetraphyllideans, but it resembles that of Cyclophyllidea in the arrangement of twisting of the CM.

• MeSH
• Cestoda ultrastructure   MeSH
• Cestode Infections parasitology   veterinary   MeSH
• Microscopy, Electron MeSH
• Fish Diseases parasitology   MeSH
• Fresh Water MeSH
• Spermatozoa ultrastructure   MeSH
• Catfishes parasitology   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Geographicals
• Egypt MeSH

The reinvestigation of the fine structure of the mature spermatozoon of the proteocephalidean tapeworm Proteocephalus longicollis, a parasite of salmonid fish, has shown new characters not described previously for this species. They include a helicoidal crested body, 60-100 nm thick and localised at the anterior extremity of the spermatozoon. Also new is the finding that cortical microtubules in P. longicollis spermatozoon are of two types. The nucleus situated in the middle part of the gamete is rod-shaped anteriorly, but it is wound around one of the axonemes posteriorly. It is newly observed that both the anterior and posterior extremities of the spermatozoon of P. longicollis contain a single axoneme. The ultrastructural characteristics of the mature spermatozoon of P. longicollis coincide in their main features with those of the spermatozoon of P. torulosus, a parasite of cyprinid fish.

• MeSH
• Cestoda ultrastructure   MeSH
• Cestode Infections veterinary   MeSH
• Microscopy, Electron veterinary   MeSH
• Fish Diseases parasitology   MeSH
• Oncorhynchus mykiss parasitology   MeSH
• Intestinal Diseases, Parasitic veterinary   MeSH
• Spermatozoa ultrastructure   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

Spermiogenesis of the proteocephalidean cestode Proteocephalus torulosus (Batsch, 1786) was examined for the first time using transmission electron microscopy. Spermiogenesis begins with the formation of a distal cytoplasmic protrusion, a differentiation zone, at the periphery of the early spermatid. This differentiation zone is lined with cortical microtubules and contains two centrioles aligned along the same axis. Subsequently, each centriole is associated with the striated root and the intercentriolar body appears between them. A flagellar bud arises from each centriole, growing later as a free flagellum. Simultaneously, a median cytoplasmic process (MCP) develops distally to the flagella. The two flagella, which are of unequal length, become longer and rotate towards the MCP. At this stage, two arching membranes appear at the base of the differentiation zone. The nucleus elongates and when both flagella are fused with the MCP, the nucleus subsequently migrates into the MCP. Finally, the advanced spermatids detach from a condensing residual cytoplasm at the level of the arching membranes.

• MeSH
• Cestoda physiology   ultrastructure   MeSH
• Cestode Infections parasitology   veterinary   MeSH
• Cyprinidae parasitology   MeSH
• Microscopy, Electron MeSH
• Fish Diseases parasitology   MeSH
• Intestinal Diseases, Parasitic parasitology   veterinary   MeSH
• Spermatids ultrastructure   MeSH
• Spermatogenesis MeSH
• Spermatozoa ultrastructure   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH