Kirsten Jensen

The genus Flapocephalus Deshmukh, 1979, is a little-known group of lecanicephalidean cestodes parasitizing cowtail rays (genus Pastinachus Rüppell) mainly in the Indo-Pacific region. Since the erection of the genus, with Flapocephalus trygonis Deshmukh, 1979, as the type species, and the description of a second species, Flapocephalus saurashtri Shinde and Deshmukh, 1979, both from Pastinachus sephen (Fabricius) from India, reports of this genus have been restricted mainly to brief mentions or discussion of its validity and taxonomic placement. More recently, phylogenetic analyses based on molecular sequence data that included specimens of Flapocephalus have supported Flapocephalus as a distinct genus allied with the Polypocephalidae Meggitt, 1924. Limited by the amount of detail provided in the original descriptions, the unavailability of type material, and the lack of a detailed generic diagnosis, we aimed to fully characterize the genus based on newly collected specimens. A total of 38 specimens of Pastinachus, representing 3 of the 5 described species and 1 undescribed species, from throughout the Indo-Pacific region were examined for specimens of Flapocephalus. Worms were prepared for examination with light and scanning electron microscopy and as histological sections. A subset of specimens was included in a phylogenetic analysis based on data for a portion of the 28S rDNA gene. Overall, these specimens were found to include the type species, 4 new species that are described herein (Flapocephalus monostylorchis Hanselman and Jensen n. sp., Flapocephalus capitonis Hanselman and Jensen n. sp., Flapocephalus rudkinorum Hanselman and Jensen n. sp., and Flapocephalus infundibulapex Hanselman and Jensen n. sp.), and 4 putative new species. Flapocephalus trygonis is redescribed from specimens from Pastinachus ater (Macleay) from Sri Lanka, and a neotype is designated. The newly described species are distinct genetically and can be distinguished morphologically from the existing species and one another based on a combination of total length, the shape of the apical organ, and the arrangement of testes and vitelline follicles. A revised diagnosis of the genus and a key to the described species are presented, as is a hypothesis of the interrelationships resulting from a maximum likelihood analysis including 4 of the 5 described species and the 4 putative new species. The presence of 4 glandular, sac-like structures surrounded by scolex proper, with channels that open to the outside at the base of the apical organ, is reported in lecanicephalidean tapeworms for the first time.

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

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.

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.

The lecanicephalidean cestodes parasitizing the spiral intestine of the endangered giant freshwater whipray, Urogymnus polylepis (Bleeker), are investigated for the first time. Eight host specimens were collected between 2002 and 2008 at 2 collecting sites off the eastern coast of Borneo: 6 from the Kinabatangan River (Malaysia) and 2 from a fish market in Tarakan (Indonesia). Two of these individuals were found to be infected with a total of 3 new species of TetragonocephalumShipley and Hornell, 1905. Tetragonocephalum georgei n. sp. and Tetragonocephalum opimum n. sp. were recovered from a host specimen from the Kinabatangan River, and Tetragonocephalum levicorpum n. sp. was found parasitizing a host specimen purchased at a fish market in Tarakan. Specimens of each of the new species were prepared for light microscopy; specimens of 2 of the new species were prepared for scanning electron microscopy, and histological sections were prepared for 1 of the new species. The 3 new species are distinct from the 9 valid species of Tetragonocephalum and the 1 species inquirendum based on, for example, total length, number of proglottids and testes, and size of the scolex and acetabula. Tetragonocephalum georgei n. sp. and T. levicorpum n. sp. are unusual among their congeners in that they are euapolytic (i.e., gravid proglottids were not observed) rather than apolytic. They differ from one another in scolex and acetabula size. Tetragonocephalum opimum n. sp. is unusual among its congeners in its possession of vitelline follicles arranged in 2, rather than 3, regions in the proglottid. These new species increase the total number of valid species of Tetragonocephalum to 12 and the total number of known cestodes from U. polylepis to 13 species across 6 genera in 4 orders. This is the first account of lecanicephalideans reported from freshwater. The taxonomic status of each of the 32 nominal taxa historically associated with Tetragonocephalum is re-assessed. Type host identities of all valid species are revised and discussed in light of recent taxonomic efforts in the Dasyatidae Jordan and Gilbert.

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.

Three new cestode species are described from the crocodile shark (Pseudocarcharias kamoharai) in Ecuador. All three were examined with light and scanning electron microscopy. The unique combination of morphological features in one of the new species prompted formal investigation of the non-monophyly ofParaorygmatobothriumrelative to the morphologically similar generaDoliobothrium,Guidus,Marsupiobothrium,Nandocestus,Orectolobicestus,RuhnkecestusandScyphophyllidium. Sequence data generated for part of the 28S rDNA gene were subjected to maximum likelihood (ML) analysis. The resulting tree led to the synonymization of six of these seven genera withScyphophyllidium, and transfer of their species to the latter genus. With the new species, the number of described members ofScyphophyllidiumis now 45. The diagnosis ofScyphophyllidiumis revised to accommodate these species. In addition, to expedite future descriptions, eight categories ofScyphophyllidiumspecies are circumscribed, based largely on bothridial features.Scyphophyllidium timvickiorumn. sp. is a category 1 species. Beyond being the smallest category 1 species, it bears, rather than lacks, apical suckers and lacks, rather than bears, strobilar scutes. The two other new species are members ofClistobothrium.Clistobothrium amyaen. sp. differs from its congeners in bothridial shape, elongate cephalic peduncle and tiny size.Clistobothrium gabywalterorumn. sp. differs from the two of its congeners that also possess foliose bothridia in overall size and testis number. Despite their substantial morphological differences, the ML tree indicates they are sister taxa. Both are unique among their congeners in possessing cephalic peduncle spinitriches. The diagnosis ofClistobothriumis revised accordingly.

Five new species of the elasmobranch tapeworm genus Anthocephalum Linton, 1890 (Rhinebothriidea: Anthocephaliidae Ruhnke, Caira and Cox, 2015) are described from the mangrove whipray, Urogymnus granulatus (Macleay) from the Solomon Islands and northern Australia. Anthocephalum blairi n. sp., Anthocephalum gravisi n. sp., Anthocephalum haroldsoni n. sp., Anthocephalum mounseyi n. sp., and Anthocephalum ruhnkei n. sp. differ from one another and their congeners based on quantitative and qualitative features of the scolex and proglottid anatomy. Species boundaries recognized by these distinguishing morphological features are supported by a maximum likelihood phylogenetic analysis based on combined 18S rDNA and 28S rDNA (D1-D3) sequence data showing the new species as independent lineages among 13 of the 18 species of Anthocephalum described to date. Several morphological features (the possession of columns of vitelline follicles that are interrupted by the ovary, a uterus that does not extend to the anterior margin of the proglottid but stops short in the anterior region of the field of testes, proximal surfaces of the marginal loculi covered with acicular filitriches only throughout, or with gladiate spinitriches and acicular filitriches throughout, and proximal bothridial surfaces with gladiate spinitriches and acicular filitriches) are collectively exhibited by the 5 new species and have not been documented previously in other species of Anthocephalum. The genus is herein amended to reflect these features. Additionally, the replacement name Anthocephalum centrurum (Southwell, 1925) Ruhnke, 1994 is reinstated as the valid name for the type species of the genus, Anthocephalum gracile Linton, 1890, with the latter being the junior secondary homonym of A. gracile (Wedl, 1855) Ruhnke, 1994. The description of these new species increases the number of valid species of Anthocephalum from 18 to 23.

The Invertebrate Zoology Collection at the University of Kansas (KU) Biodiversity Institute is one of KU’s smaller collections, with just over 2,000 lots. Its taxonomic strength are hexacorallians (Cnidaria: Anthozoa) from across the globe. Holdings also include earthworms primarily from Southeast Asia and the Caribbean, as well as crayfish and molluscs from the United States, notably from Kansas. The collection has seen little loan activity over the past decade, in part due to the fact that collection records are not digitally available. Moreover, the collection has been virtually untouched for several years as research activities on hexacorallians has ceased following curator retirement. In an initial inventory, physical holdings were checked against original catalog data, while simultaneously re-curating to ensure proper storage containers and maximal levels of either ethanol or formalin. Preliminary comparison of the catalogued data with original and secondary label data housed with the specimens suggest that across these sources, the captured and entered data is somewhat inconsistent and incomplete. In an attempt to remedy such issues, the next phase of the project will involve digitally capturing label data to verify collection information. Once the data has been validated, the working data in spreadsheet format will be imported into Specify, and published to a list of aggregators including Global Biodiversity Information Facility (GBIF), Integrated Digitized Biocollections (iDigBio), Biodiversity Information Serving Our Nation (BISON), and Ocean Biogeographic Information System (OBIS), for visibility and use outside of KU. The hope through such efforts is an accessible and easily searchable collection that is properly preserved for future research.