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This paper aims to expand understanding of a poorly known group of cestodes that parasitize an intriguingly diverse suite of elasmobranchs. The group’s three currently described members (i.e., Pentaloculum macrocephalum, Pentaloculum hoi, and Zyxibothrium kamienae) parasitize an electric ray, a carpet shark, and a skate, respectively. Pentaloculum grahami n. sp. is described from a second genus of carpet shark, specifically Parascyllium collare, in Australia. Zyxibothrium duffyi n. sp. and Zyxibothrium healyae n. sp. are described from the deep-sea skates Brochiraja asperula and Brochiraja spinifera, respectively off New Zealand. The three new species share distinctive bothridia that bear a small number of large, circular, facial loculi and lateral bands of vitelline follicles that converge posterior to the ovary—features which are found in all other members of these genera. Zyxibothrium healyae n. sp. is unique in possessing three, rather than four or five, facial loculi. Zyxibothrium duffyi n. sp. possesses a combination of five facial loculi and vitelline follicles that stop short of the anterior margin of the proglottid. Pentaloculum grahami n. sp. is the largest member of the group with the greatest number of proglottids. Based on striking similarities in scolex morphology, Pentaloculum and Zyxibothrium have been hypothesized to belong to a distinct subgroup of “tetraphyllideans” provisionally designated as Clade 1. Based on sequence data for the D1–D3 region of the 28S rDNA gene generated for species of Zyxibothrium for the first time, we confirm the reciprocal monophyly of both genera as well as the monophyly of Clade 1 and its status as a distinct lineage among the “Tetraphyllidea”. This work also suggests that the presence of five facial loculi is homoplasious given this character state is found in members of both genera. The new species expand the host associations of Clade 1 to include additional skate and carpet shark genera. Moving forward we would expect to find additional members of this group parasitizing other species of parascyliid carpet sharks as well as other species of the rajid genus Malacoraja and the arhynchobatid genus Brochiraja. Here we have doubled the number of described species in the taxon referred to as Clade 1 while simultaneously expanding our understanding of the morphology and anatomy of its members. This additional information will help inform the ultimate revision of the ordinal classification of the cestodes to address the highly polyphyletic nature of the order “Tetraphyllidea” as it is currently configured. 

Abstract Three new species of ‘tetraphyllideans’ in the family Serendipeidae are described from cownose rays off Senegal. Duplicibothrium jeannettae n. sp. and Duplicibothrium collosum n. sp. parasitize both Rhinoptera marginata and Rhinoptera peli . Duplicibothrium jillae n. sp. parasitizes R. marginata . Duplicibothrium jeannettae n. sp. and D. jillae n. sp. differ from one another and their described congeners in size, number of proglottids, and locular configuration. The generic assignment of D. collosum n. sp. was initially unclear because its scolex more closely resembles that of Serendip than Duplicibothrium . To help inform generic placement, sequence data were generated for the 28S rDNA gene (D1–D3 region) for the three new species as well as for Duplicibothrium minutum and Duplicibothrium n. sp. 2 from Rhinoptera bonasus off North Carolina, Duplicibothrium n. sp. 3 from Rhinoptera brasiliensis off Belize, Duplicibothrium n. sp. 4 and Duplicibothrium n. sp. 5 from Rhinoptera jayakari off Mozambique, and Duplicibothrium n. sp. 6 from Rhinoptera neglecta off Australia. Comparable data were obtained from GenBank for adults and larvae of Duplicibothrium collected from the Gulf of Mexico. The tree resulting from a maximum likelihood analysis (MLA) placed D. collosum n. sp. robustly within Duplicibothrium ; the generic diagnosis is emended accordingly. This raises a question regarding the independence, and thus also validity, of the three genera of the Serendipeidae – a question that must await a molecular analysis that includes Serendip and Glyphobothrium . These results extend the hosts of Duplicibothrium to include R . brasiliensis , R . jayakari , R . marginata , R . neglecta and R . peli , and the distribution to include the western Pacific Ocean, eastern Atlantic Ocean, and western Indian Ocean (Zoobank Registration: 97BB5020-BFFF-4FEA-AE07-B4711D1110FC). 

The Rhoptrobothriidae are one of the more enigmatic families of cestodes of elasmobranchs. Opinions on the taxonomic status of the family’s three original genera (i.e., Myzophyllobothrium, Rhoptrobothrium, and Myzocephalus) have varied over the 115 years since they were erected. Some authors have considered all three valid, others have considered Rhoptrobothrium to be a synonym of Myzopyllobothrium or a genus inquirendum, yet others have considered Myzocephalus to be a synonym of the phyllobothriid genus Thysanocephalum. All three genera were established for specimens collected from eagle rays off Sri Lanka. The erection of Mixophyllobothrium for two specimens from a cowtail stingray off India three decades ago added additional confusion to the situation, with some authors considering it valid and others a synonym of Myzocephalus. These disagreements stem largely from differences in interpretation of the complex morphology of the scolex of members of these genera. Furthermore, with the exception of Rhoptrobothrium comprising four species, each genus is monotypic. All but Rhoptrobothrium has not been considered in detail for nearly a century, largely because of a lack of available material. The taxonomic status of these genera is assessed here based on light and scanning electron microscopy, and molecular data generated from new material collected from eagle rays off Indonesian and Malaysian Borneo, Japan, Sri Lanka, and Viet Nam. Morphological work indicates that the genera differ largely only in the degree of folding of the four remi that extend from the cephalic peduncle. A molecular phylogeny based on sequence data for the D1–D3 region of the 28S rRNA gene, which include new data for eight specimens of four species, indicates that Myzophyllobothrium, Myzocephalus, and Rhoptrobothrium are not mutually monophyletic. The latter two genera and Mixophyllobothrium are considered synonyms of Myzophyllobothrium and five species are transferred to that genus. Myzophyllobothrium okamuri n. comb. is considered a species inquirendum. Myzophyllobothrium nagasawai n. sp. is described from Aetobatus narutobiei off Japan. Myzophyllobothrium narinari n. comb. is re-described based on newly collected cestodes from the type host and locality (i.e., Aetobatus ocellatus off Sri Lanka). Despite consisting of only a single genus, the family status of the group is retained in recognition of the unusual configuration of the scolex, which bears four biloculate bothridia and four remi extending from the cephalic peduncle. The ordinal placement of the family remains uncertain, but affinities with the Phyllobothriidea, rather than “Tetraphyllidea” are considered. 

Nutritionally-based mutualisms with bacteria are known to occur in a wide array of invertebrate phyla, although less commonly in the Platyhelminthes. Here we report what appears to be a novel example of this type of association in two geographically disparate and phylogenetically distant species of tapeworms of eagle rays—the lecanicephalidean Elicilacunosus dharmadii off the island of Borneo and the tetraphyllidean Caulobothrium multispelaeum off Senegal. Scanning and transmission electron microscopy revealed that the grooves and apertures on the outer surfaces of both tapeworms open into expansive cavities housing concentrations of bacteria. This led us to reject the original hypothesis that these structures, and their associated mucopolysaccharides, aid in attachment to the host mucosa. The cavities were found to be specialized in-foldings of the tapeworm body that were lined with particularly elongate filitriches. Given tapeworms lack a gut and employ filitriches to assist in nutrient absorption, enhanced nutrient uptake likely occurs in the cavities. Each tapeworm species appeared to host different bacterial monocultures; those in E . dharmadii were coccoid-like in form, while those in C . multispelaeum were bacillus-like. The presence of bacteria in a specialized structure of this nature suggests the structure is a symbiotic organ. Tapeworms are fully capable of obtaining their own nutrients, and thus the bacteria likely serve merely to supplement their diet. Given the bacteria were also extracellular, this structure is more consistent with a mycetome than a trophosome. To our knowledge, this is not only the first evidence of an external symbiotic organ of any type in a nutritionally-based mutualism, but also the first description of a mycetome in a group of invertebrates that lacks a digestive system. The factors that might account for the independent evolution of this unique association in these unrelated tapeworms are unclear—especially given that none of their closest relatives exhibit any evidence of the phenomenon. 

Abstract New genera are erected for three clades of tapeworms originally discovered using molecular sequence data. The morphological features of each are characterized after examination of specimens with light and scanning electron microscopy. Rockacestus gen. nov. parasitizes skates. Ruhnkebothrium gen. nov. parasitizes hammerhead sharks. Yamaguticestus gen. nov. parasitizes small squaliform sharks and catsharks. The novelty of these genera is supported by a taxonomically comprehensive molecular phylogenetic analysis of the D1–D3 region of the 28S rDNA gene, which, with the addition of newly generated sequence data, is the first to include representation of 15 of the 18 genera of phyllobothriideans plus the three new genera. Five new species are described from elasmobranchs in the western Atlantic Ocean, the Gulf of California, Chile, the Falkland Islands and South Africa to help circumscribe the new genera. Two of the genera provide appropriate generic homes for ten species of phyllobothriideans from catsharks and skates with uncertain generic affinities and thus resolve longstanding taxonomic issues. Given that these genera parasitize some of the most poorly sampled groups of elasmobranchs (i.e. hammerhead sharks, squaliform sharks, catsharks and skates), based on the strict degree of host specificity observed, we predict that further work on other members of these groups will yield as many as 200 additional species in these three genera of tapeworms globally. This brings the total number of genera in the Phyllobothriidea to 21. 

Two new species of the cestode genus Caulobothrium, collected from the duckbill eagle ray, Aetomylaeus bovinus, off Senegal, are described. Although postulated as sister taxa in an earlier molecular phylogenetic analysis, Caulobothrium multispelaeum n. sp. and Caulobothrium katzi n. sp., respectively, are among the smallest and largest members of the genus. The smaller species is unique among its congeners in possessing unusual medial longitudinal grooves along the dorsal and ventral surfaces of its strobila that develop into a tandem series of elliptical apertures on the posterior proglottids. The inner surfaces of these apertures stained positively with McManus’ periodic acid Schiff in a manner similar to that seen in members of the distantly related lecanicephalidean genus Elicilacunosus. The larger species differs from its congeners in size, number of proglottids, and arrangement of bothridial loculi. Both new species were found to possess a small apical sucker on the anterior margin of each of their bothridia. Scanning electron microscopy (SEM) and frontal sections of a bothridium of Caulobothrium tetrascaphium suggests that this species also bears an apical sucker. Examination of the hologenophore of the species provisionally referred to as Caulobothrium n. sp. 5 in the earlier molecular analysis indicates it is conspecific with the recently described Caulobothrium pedunculatum, which was also determined to possess bothridial apical suckers. This leads us to suspect that this feature may be found to occur in all members of the genus. SEM of specimens of Caulobothrium for the first time indicates their bothridial surfaces are covered with filitriches of various sizes but lack spinitriches; spinitriches were seen only on the cephalic peduncle of C. katzi n. sp. The geographic distribution and host associations of Caulobothrium are expanded to include data now available for all species. The diagnosis of the genus is revised to include all of this information.