Three new genera of rhinebothriidean cestodes from stingrays in Southeast Asia.

Three genera of rhinebothriideans, previously referred to as New genus 1, New genus 2 and New genus 4, are erected in the the Anthocephaliidae. New genus 1 is established as Divaricobothrium gen. n., with Divaricobothrium tribelum sp. n. as its type species; Echeneibothrium trifidum Shipley et Hornell, 1906 is transferred to the genus as Divaricobothrium trifidum (Shipley et Hornell, 1906) comb. n. This genus is unique among rhinebothriidean genera in bearing bothridia that are posteriorly deeply divided into two lobes with facial loculi but no apical sucker, and a vagina that extends to near the anterior margin of the proglottid. Its species parasitise Indo-Pacific members of the genera Brevitrygon Last, Naylor et Manjaji-Matsumoto, Maculabatis Last, Naylor et Manjaji-Matsumoto and Pateobatis Last, Naylor et Manjaji-Matsumoto. New genus 2 is established as Barbeaucestus gen. n., with Barbeaucestus jockuschae sp. n. as its type species; Barbeaucestus ralickiae sp. n. is also described. Anthobothrium sexorchidum Williams, 1964 and Rhinebothrium shipleyi Southwell, 1912 are transferred to the genus as Barbeaucestus sexorchidus (Williams, 1964) comb. n. and Barbeaucestus shipleyi (Southwell, 1912) comb. n., respectively. This genus is unique among rhinebothriidean genera in that its bothridia are substantially wider than long, bear an apical sucker and at least one row of two or more facial loculi in their anterior half. Its species parasitise the genera Neotrygon Castelnau and Taeniura Müller et Henle. New genus 4 is established as Sungaicestus gen. n. with transfer of Rhinebothrium kinabatanganensis Healy, 2006, as Sungaicestus kinabatanganensis (Healy, 2006) comb. n., as its type species. Among the genera of its order, this genus most closely resembles Rhinebothrium Linton, 1890, however, despite the original description, the bothridia were found to bear, rather than lack, apical suckers. This monotypic genus is known only from the freshwater stingray Urogymnus polylepis (Müller et Henle). The familial diagnosis of the Anthocephaliidae Ruhnke, Caira et Cox, 2015 is emended. The family now houses five genera.

A new genus of rhinebothriidean cestodes from batoid elasmobranchs, with the description of five new species and two new combinations.

Survey work of batoid elasmobranchs in the eastern Atlantic and Indo-Pacific revealed multiple species of a new genus of cestode. Stillabothrium Healy et Reyda gen. n. (Rhinebothriidea: Escherbothriidae) is unique in its possession of an even number of non-medial longitudinal septa in the posterior portion of the bothridia, resulting in a series of loculi that are longer than wide (i.e. vertically oriented) and are arranged in columns. Five new species of Stillabothrium are described, S. ashleyae Willsey et Reyda sp. n., S. davidcynthiaorum Daigler et Reyda sp. n., S. campbelli Delgado, Dedrick et Reyda sp. n., S. hyphantoseptum Herzog, Bergman et Reyda sp. n., S. jeanfortiae Forti, Aprill et Reyda sp. n., and two species are formally transferred to the genus, S. amuletum (Butler, 1987) comb. n., and S. cadenati (Euzet, 1954) comb. n., the latter of which is redescribed. The species differ in the configuration of the other bothridial septa and in proglottid anatomy. Species of Stillabothrium were found parasitising a total of 17 species of batoid elasmobranchs of the genera Dasyatis Rafinesque, Glaucostegus Bonaparte, Himantura Müller et Henle, Pastinachus Rüppell, Rhinobatos Linck and Zanobatus Garman, including several host species that are likely new to science. A phylogenetic hypothesis based on Bayesian analysis of 1 084 aligned positions of the D1-D3 region of 28S rDNA for 27 specimens representing 10 species of Stillabothrium and two outgroup species supported the monophyly of Stillabothrium. These results also supported morphologically determined species boundaries in all cases in which more than one specimen of a putative species was included in the analysis. Host specificity appears to vary across species of Stillabothrium, with the number of host species parasitised by each species of Stillabothrium ranging from one to four. The geographic distribution of species of Stillabothrium spans the eastern Hemisphere, including the eastern Atlantic (coastal Senegal) and several locations in the Indo-Pacific (coastal Vietnam, Borneo and Australia). In addition, Phyllobothrium biacetabulatum Yamaguti, 1960 is formally transferred into family Escherbothriidae, although its generic placement remains uncertain (species incertae sedis).

Three new species of Rhinebothrium (Cestoda: Tetraphyllidea) from the freshwater whipray, Himantura chaophraya, in Malaysian Borneo.

Three new Rhinebothrium Linton, 1890 species, R. kinabatanganensis n. sp., R. megacanthophallus n. sp., and R. abaiensis n. sp., collected from the spiral intestines of 6 freshwater whiprays (Himantura chaophraya, Dasyatidae) in the Kinabatangan River, Malaysian Borneo, are described. This is the first record of cestodes from the Kinabatangan River and from H. chaophraya. The new Rhinebothrium species are differentiated from the 34 valid Rhinebothrium species in addition to 2 species, Echeneibothrium hui Tseng, 1933 and E. oligotesticularis Subramaniam, 1940 that are herein transferred to Rhinebothrium. Each of the new species was examined with light and scanning electron microscopy. Two of the new species possess cilia on their bothridia, which have not previously been reported for members of this genus.

Two new species of Paraorygmatobothrium (Cestoda: Tetraphyllidea) from weasel sharks (Carcharhiniformes: Hemigaleidae) of Australia and Borneo.

Two new species of Paraorygmatobothrium Ruhnke, 1994, P. janineae n. sp. and P. kirstenae n. sp., are described from the spiral intestine of 2 shark species of the Family Hemigaleidae: Hemigaleus microstoma and Hemipristis elongata. The 2 new cestode species differ from other members of Paraorygmatobothrium in vitelline follicle distribution and possession of a cephalic peduncle. The 2 new species differ from 1 another in total length, maximum width, scolex size, number of proglottids per strobila, and number of testes per proglottid. The generic diagnosis of Paraorygmatobothrium is emended to include the new species. The results of this study extend the distribution of Paraorygmatobothrium to include the carcharhinid shark family Hemigaleidae.

A new genus and species of cestode (Rhinebothriidea) from Mobula kuhlii (Rajiformes: Mobulidae) from Malaysian Borneo.

The cestode fauna of the shortfin devilray, Mobula kuhlii (Müller & Henle, 1841) was examined for the first time. The work resulted in the discovery of a new genus and species of rhinebothriidean tapeworm. Crassuseptum pietrafacei, n. gen. n. sp. is erected here on the basis of its unique scolex and proglottid morphology. Histological sections and examination by light and scanning electron microscopy demonstrated that this new genus differs from all other rhinebothriidean genera in its possession of bothridia in which the proximal and distal sides are confluent, i.e., not separated by a rim of tissue, and in its possession of testes that extend to the posterior margin of the ovary. This new species is characterized in part by its possession of stalked, elongate bothridia lacking lateral constrictions, with 13-15 prominent transverse bothridial septa and 4 reduced transverse septa, craspedote proglottids, each with 2-3 layers of testes in cross section, and a vas deferens that joins the cirrus sac at its anterior margin. Histological and optical sections through bothridial septa revealed that the transverse septa are formed by septal muscles separate from bothridial radial musculature, extending from the anterior side to the posterior side of each septum. This is only the second species rhinebothriidean cestode reported from mobulids. This study adds to the number of new species and genera of elasmobranch cestodes discovered off of the island of Borneo.

Proposal for a new tapeworm order, Rhinebothriidea.

The polyphyletic nature of the tapeworm order Tetraphyllidea Carus, 1863 is addressed in part with the establishment of the new order Rhinebothriidea for a subset of the taxa formerly comprising the phyllobothriid subfamily Rhinebothriinae (Platyhelminthes: Eucestoda). Support for the order comes from Bayesian, maximum likelihood, and parsimony analyses of complete ssrDNA and partial (D1-D3) lsrDNA sequence data for 58 cestode species. These data consisted of novel data generated for 40 species in 15 genera of candidate rhinebothriines and the cathetocephalidean species Sanguilevator yearsleyi as well as comparable data taken from GenBank for an additional 18 cestode species in 17 genera. In total, the species analyzed consisted of two Cathetocephalidea, two Litobothriidea, two Lecanicephalidea, three Proteocephalidea, and 49 Tetraphyllidea. The tetraphyllideans consisted of three Onchobothriidae, three Serendipidae, and 43 Phyllobothriidae (one Thysanocephalinae, one Echeneibothriinae, five Phyllobothriinae, 35 candidate Rhinebothriinae and the poorly known Spongiobothrium). This work suggests that some elements of current membership in the group are in need of revision. For example, while inclusion of the echeneibothriine genus Echeneibothrium and the phyllobothriine genera Rhodobothrium and Anthocephalum, and also Spongiobothrium, in the Rhinebothriidea is supported, inclusion of Duplicibothrium and Caulobothrium in the new order is not. Histological sections and scanning electron microscopy of selected members of the study group suggest that the presence of bothridial stalks may serve as an effective morphological feature to characterise the order. The group is restricted to elasmobranchs, and appears to have a particular affinity for Myliobatiformes. The new order includes at least 13 genera. Intraordinal relationships were determined to be insufficiently stable to justify the formal reorganization of rhinebothriidean families at this time.

A revision of Platybothrium Linton, 1890 (Tetraphyllidea: Onchobothriidae), with a phylogenetic analysis and comments on host-parasite associations.

A revision of Platybothrium Linton, 1890 is presented, based on available type and voucher material, as well as extensive new collections from elasmobranchs belonging to the Carcharhinidae (requiem sharks) and Sphyrnidae (hammerhead sharks). All 10 nominal Platybothrium species are treated or redescribed herein. Four of these 10 nominal species, in addition to one former member of Dicranobothrium Euzet, 1953, are considered valid members of Platybothrium. Five new Platybothrium species are described: P. angelbahiense n. sp. ex Carcharhinus leucas, P. coshtaprum n. sp. ex C. plumbeus, P. jondoeorum n. sp. ex Negaprion acutidens and C. melanopterus, P. kirstenae n. sp. ex C. obscurus, and P. tantulum n. sp. ex Sphyrna lewini and S. zygaena. Sixty-three morphological characters were employed in cladistic analyses of nine Platybothrium species and five outgroup species. Coding and analysis strategies were varied to assess the effects of coding inapplicable characters as missing or as a separate character state, and of excluding characters for which data are missing in more than 10% of the taxa. The analysis in which inapplicable characters were coded as a separate character state and no characters were excluded produced the best-supported and most conservative estimate of the interrelationships of Platybothrium spp. Platybothrium appears to be a monophyletic assemblage, with the most basal species being P. spinulifera Southwell, 1912. The group of species possessing an accessory piece between the hooks forms a clade within the genus. Species lacking an accessory piece, which had previously been placed in Dicranobothrium Euzet, 1953, do not appear to be each other's closest relatives; thus, Dicranobothrium is considered a synonym of Platybothrium. An examination of host associations indicated that Platybothrium species are broadly distributed among, and entirely restricted to, carcharhinid and sphyrnid shark species. Most Platybothrium species exhibit oioxenous host-specificity, with all but two species each parasitising only a single host species. In several host species, multiple Platybothrium congeners parasitise the same host individual, a phenomenon not previously reported for Platybothrium.