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  1. 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. 
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  2. Worsaae, Katrine (Ed.)
    The three members of the lecanicephalidean tapeworm family Eniochobothriidae are unusual among tapeworms in that they lack a vagina and possess a series of expanded proglottids forming a trough at the anterior end of their body. They exclusively parasitise cownose rays of the genus Rhinoptera (Myliobatiformes: Rhinopteridae). New collections from six of the nine known species of cownose rays from the waters off Australia, Mexico, Mozambique, Senegal, Taiwan and the United States (off Mississippi, Louisiana and South Carolina) yielded eight new species and a new genus of eniochobothriids. Here we erect Amiculucestus, gen. nov. and describe six of the eight new species – four in the new genus and two in Eniochobothrium – expanding the number of genera in the family to two and the number of described species in the family to nine. Morphological work was based on light and scanning electron microscopy. The tree resulting from a maximum likelihood analysis of sequence data for the D1–D3 region of the 28S rDNA gene for 11 species of eniochobothriids supports the reciprocal monophyly of both genera. The mode of attachment to the mucosal surface of the spiral intestine of the host was investigated using histological sections of worms in situ. These cestodes appear to use the anterior trough-like portion of their body, which consists of an unusual series of barren proglottids, rather than their scolex, to attach to the mucosal surface. Based on our new collections, we estimate that the total number of eniochobothriids across the globe does not exceed 27 species. ZooBank LSID: urn:lsid:zoobank.org:pub:0740EC72‐AC3F‐43AA‐BD41‐B9820BA9D0CE 
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  3. 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). 
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  4. Since 2010, the trypanorhynch tapeworm family Rhinoptericolidae Carvajal & Campbell, 1975 has housed just two distinctive, monotypic genera ( Rhinoptericola Carvajal & Campbell, 1975 and Nataliella Palm, 2010). However, global collections of tapeworms from sharks and rays over the last more than three decades brought to light the need for major revision of the family by suggesting a much greater species-level diversity for the nominal genus Rhinoptericola . Through synonymy and the description of new species, the number of species in the genus is increased from one to eight. A phylogenetic analysis of the D1–D3 gene region of 28S rRNA (28S), including seven of the now nine species of rhinoptericolids, and a broad sampling of the other Trypanobatoida is the first to recover a monophyletic Rhinoptericolidae. In addition to systematic revision, this study allowed for the first evaluation of the degree of intraspecific vs interspecific variation in 28S for adult trypanorhynchs across the various hosts and geographic localities from which they have been reported, suggesting a relatively consistent boundary for Rhinoptericola . It is further suggested that detailed scanning electron microscopy (SEM) images of both the basal and metabasal armatures greatly aid in the interpretation of hook arrangement and shape. A schematic to streamline determination of the tentacular surface presented in scanning electron micrographs and line drawings of trypanorhynchs is presented for species with both two and four bothria. In combination, these methodological refinements can now be used as a model to resolve issues of classification and non-monophyly within both major lineages of the Trypanorhyncha. As a result of the taxonomic work, Rhinoptericola megacantha Carvajal & Campbell, 1975 (previously only known from the American cownose ray from the Chesapeake Bay and the Ticon cownose ray from the Gulf of Mexico, Venezuela, and Brazil) is now known from an additional species of cownose ray and a species of stingray, and is revealed to have a transatlantic distribution. Data from SEM suggest a simpler interpretation of hook arrangement in the metabasal armature for Rhinoptercola and—in combination with 28S sequence data—support Shirleyrhynchus Beveridge & Campbell, 1988 (a former rhinoptericolid) as its junior synonym. The three species formerly assigned to Shirleyrhynchus are thus transferred to Rhinoptericola . Data from light microscopy on whole-mounted specimens and histological sections, SEM, and 28S showed the eutetrarhynchid Prochristianella jensenae Schaeffner & Beveridge, 2012b to be morphologically consistent with species of Rhinoptericola and it is thus transferred to the genus. The type series of P. jensenae was determined to be mixed, representing two distinct species which are here redescribed and described as new, respectively. Two additional novel species of Rhinoptericola are described from cownose rays from off Mozambique and the Gulf of California. 
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  5. 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. 
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  6. null (Ed.)
    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. 
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  7. Ruiz-Rodriguez, Magdalena (Ed.)
    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. 
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