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The parasites of hosts of conservation concern are often poorly known. This is the case with the iconic group of elasmobranchs known as the sawfish of the genus Pristis, all four species of which are considered as Endangered or Critically Endangered by the International Union for Conservation of Nature (IUCN, Switzerland). Examination of cestodes from three species of sawfish (Pristis pristis, Pristis clavata, and Pristis zijsron) in Australia and one of their close relatives, the also critically endangered widenose guitarfish, Glaucostegus obtusus, in India, collected over the past 25 years, yielded four new species of tapeworms which are described herein. All four belong to the previously monotypic Mixobothrium; the diagnosis of the genus is revised to accommodate the new species. Among the new taxa is a species that had been included in previous molecular phylogenies but whose identity and affinities within the order Rhinebothriidea, and thus also its familial placement, were unclear. This species exhibits the morphological features of Mixobothrium and thus its identity is, at long last, revealed. Sequence data generated for the 28S rDNA gene for three of the new species, as well as an additional new but yet undescribed species from Pristis pectinata from Florida (USA), confirms the uniqueness of this group among the rhinebothriideans. The new family Mixobothriidae is established to house these taxa. The members of this family differ from all but one of the five other families of rhinebothriideans in lacking apical suckers on their bothridia. They are also distinctive in that their bothridia are divided into three regions. The anterior and posterior regions have similar locular configurations to one another and differ from the locular configuration of the middle region. As a consequence, the bothridia are symmetrical along both their vertical and horizontal axes. We predict that a focus on species of guitarfish in the genus Glaucostegus will be the most productive approach for discovering additional diversity in this family of cestodes., (Copyright © 2023 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

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., (© American Society of Parasitologists 2021.)

Here we describe a new species of RhinebothriumLinton, 1890, from Hypanus guttatus (Bloch and Schneider). Rhinebothrium ramosi n. sp. can be differentiated from all 51 valid species of the genus by having 4-5 testes and uterus that extends throughout the entire length of the proglottid. Only 8 of the above species closely resemble R. ramosi in total length (Rhinebothrium bunburyense, Rhinebothrium chollaense, Rhinebothrium corbatai, Rhinebothrium dasyatidis, Rhinebothrium kruppi, Rhinebothrium lintoni, Rhinebothrium margaritense, and Rhinebothrium reydai). Despite the resemblance, R. bunburyense, R. corbatai, R. dasyatidis, R. lintoni, and R. margaritense can be distinguished from the new species by possessing a larger number of proglottids. The remaining 3 species (R. chollaense, R. kruppi, and R. reydai) overlap in total length and number of proglottids with R. ramosi. However, they can be distinguished from the new species by possessing a single posterior-most bothridial loculus instead of arranged as a pair, as found in the new species. This is the first report of the genus from the coastal waters of Brazil and brings to 52 the number of valid species for this genus. Additionally, we use the patterns of infection and distribution for species of Rhinebothrium to make predictions of expected diversity within the genus, especially for unsurveyed hosts in endemic marine ecoregions of the world., (© American Society of Parasitologists 2020.)

Examination of 4 specimens of the leopard whipray Himantura leoparda, a dasyatid stingray from northern Australia, led to the discovery of 3 new species of Rhinebothrium. Rhinebothrium leopardensis n. sp., Rhinebothrium nandoi n. sp., and Rhinebothrium ruhnkei n. sp. are described, increasing the diversity of the genus to 51 species globally. All 3 new species differ from their congeners in terms of testis number, proglottid number, loculus number, and size. With respect to one another, R. leopardensis n. sp. has bothridia that are weakly constricted at their centers and has a greater number of proglottids than the other 2 species (93-108 vs. 11-15, and 48-78, respectively). Rhinebothrium nandoi n. sp. is the smallest of the 3 species found in H. leoparda (3.6-5 vs. 10-15 mm and 10.1-15.8 mm in total length [TL], respectively) and bears bothridia that are constricted at their centers. Rhinebothrium ruhnkei n. sp. bears bothridia that are conspicuously constricted at their centers and has more testes than R. leopardensis and fewer than R. nandoi (7-10 vs. fewer than 7 and 21-33, respectively). Before this study, 56% (27 of 48) of Rhinebothrium species had been described from the freshwater river systems of South America and the marine waters surrounding South and North America. In contrast, despite the remarkably diverse nature of its batoid fauna, only 19 species were known from the Indo-Pacific region. Our work increases this number to 22, emphasizing the highly underestimated nature of Rhinebothrium diversity in this region of the globe. The discovery of these 3 new species was not unexpected, given the relatively poor status of our current knowledge of the cestode faunas of dasyatid stingrays in the Indo-Pacific region, and given the fact that it is common for a single batoid species to host 2 or more species of Rhinebothrium. Our results suggest that additional work on the cestode faunas of the batoids, especially dasyatids, from the Indo-Pacific region is likely to be highly productive in terms of contributing to the knowledge of Rhinebothrium diversity., (© American Society of Parasitologists 2020.)

An understanding of the mechanisms that determine host and parasite relationships is a central aim in parasitology. Association of a parasite species with a host species may be influenced primarily by phylogenetic constraints that cause parasite species to co-speciate with their host species, or predominantly by ecological parameters that influence all other co-evolutionary scenarios. This study aimed to investigate the role of co-speciation as well as other co-evolutionary scenarios in influencing the assemblages of tapeworm parasites (marine cestodes) in skate hosts (Rajiformes) using a modification of the PACo (Procrustean Approach to Cophylogeny) method. The study found that phylogeny and host ecology are both significant predictors of skate-tapeworm relationships, implying that co-speciation as well as other co-evolutionary scenarios are shaping these associations. The study also investigated the key ecological parameters influencing host-switching and found that host diet, distribution depth, average body size and geographical location have a combined effect. Given the importance of parasites in ensuring healthy and stable marine ecosystems, the findings of this study have implications for conservation management worldwide.

Examination of rajid skates off the Algerian coast in the Mediterranean Sea revealed that three of the 33 Dipturus batis Linnaeus, 1758 examined harboured a new tapeworm species: Echeneibothrium algeriensis n. sp. This new species, collected from the anterior half of the spiral valves, is described on the basis of morphological data from light and scanning electron microscopy. The new species differs from previously described Echeneibothrium species by details of the scolex and loculi, total length, the length of the myzorhynchus, the number of proglottides, and the number of testes. Comparison of the diets of the ten skate species common in the Mediterranean basin indicates some varying degree of overlap, suggesting that host specificity in this host-parasite system is determined by other host and/or ecological variables such as adaptations of the parasites to their respective hosts, either on the morpho-anatomical level, in physiological characteristics of the parasite's habitat, in the trophic requirements for the successful transmission of the parasite, or in adaptations to the behavioural characteristics of the host. Furthermore, restricted overlap of E. algeriensis n. sp. with congeners in parasite assemblages of D. batis indicates some structuring according to attachment-site preferences. However, attachment-site preferences are not explained solely by morphological compatibility between bothridia and villi. This study reiterates the need to examine multiple factors synergistically in studies on host specificity of parasites, and the need to examine the parasite fauna of hosts across their entire geographical range in order to truly appreciate the biodiversity they harbour.

Parasites are important components of natural systems, and among their various roles, parasites strongly influence the flow of energy between and within food webs. Over 1000 tapeworm species are known to parasitise elasmobranchs, although full life cycles are resolved for fewer than 10 of them. The lack in resolution stems from the inability to distinguish larval from adult stages using morphology alone. Molecular elucidation of trophic transmission pathways is the next step in understanding the role of hosts and parasites within food webs. We investigated the parasite assemblage of New Zealand's rough skate, Zearaja nasuta. Skates and their prey items (obtained from the skates' stomachs) were dissected for the recovery of adult and larval tapeworms, respectively. A fragment of the 28S rDNA region was amplified for worm specimens with the aim to confirm species identity of parasites within rough skates and to uncover trophic transmission pathways that exploit the predation links between rough skates and their prey. We identified seven species of tapeworms from four tapeworm orders. Four trophic transmission pathways were resolved between three prey items from skates stomachs and skates, and one pathway between larval tapeworm sequence from a New Zealand sole and skate, i.e. a genetic match was found between larval tapeworms in prey and adult worms in skates. We report the first case of an adult trypanorhynch parasitising rough skate. These findings contribute to our limited understanding of cestode life cycles as well as providing insights into the importance of predator-prey relationships for parasite transmission.

Species of the eutetrarhynchid genus Hispidorhynchus Schaeffner and Beveridge, 2012 possess an uncinate macrohook on the bothrial surface of the basal swelling of each tentacle. This unique feature of the oncotaxy is only shared with the closely related genus Oncomegas Dollfus, 1929 . A new species of Hispidorhynchus is described from specimens infecting Styracura schmardae (Werner, 1904) (Potamotrygonidae) from the western Caribbean Sea off the coast of Belize and Panama. Hispidorhynchus styracurae n. sp. differs from its 3 congeners in the possession of smaller and narrower bulbs, fewer principle hooks in the metabasal armature, size of the macrohook, and different scolex proportions. Scanning electron microscopy reveals new information on the microthrix morphology of the genus. A diagnostic key for the differentiation of species of Hispidorhynchus is provided. In addition, new host and locality records are reported for Oncomegas wageneri ( Linton, 1890 ) Dollfus, 1929 , collected from Hypanus guttatus (Bloch and Schneider, 1801) (Dasyatidae) off the coast of Maceió, Alagoas (Brazil). This considerably expands the geographical distribution of this species in the western Atlantic Ocean.

The spiral intestines of a total of 16 round fantail stingrays Taeniurops grabata from the Mediterranean Sea off Tunisia were examined for cestodes. A new genus is erected in the Anthocephaliidae (Rhinebothridea) as Alveobothrium gen. n., with Alveobothrium grabatum sp. n. as its type species; the new genus differs from the other genera in the order in that its members possess bothridia with an apical sucker, marginal loculi and multiple staggered rows of facial loculi. Alveobothrium zarzisense sp. n. is also described. The species differ in the number of marginal loculi and in proglotid anatomy. Another anthocephaliid belonging to the genus Anthocephalum is also described from T. grabata. Anthocephalum jeancadenati sp. n. is most similar to A. alicae and A. michaeli, but differs in size of terminal proglottid and number of proglottids. All these new species are also found in formalin-preserved cestodes from T. grabata collected at Gorée Island (Senegal) between 1946 and 1954 by the French ichthyologist J. Cadenat and conserved in the personal collection of the late L. Euzet. The presence of the same rhinebothriideans species parasitizing T. grabata in both the Mediterranean (Tunisia) and the eastern Atlantic (Senegal) is discussed.

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 helminthological examination of nine individuals of Aetobatus cf. narinari (spotted eagle ray; raya pinta; arraia pintada) revealed the presence of an undescribed species of cestode of the genus Acanthobothrium. The stingrays were collected from four locations in México: Laguna Términos, south of Isla del Carmen and the marine waters north of Isla del Carmen and Champotón, in the State of Campeche, and Isla Holbox, State of Quintana Roo. The new species, nominated Acanthobothrium marquesi, is a category 3 species (i.e, the strobila is long, has more than 50 proglottids, the numerous testicles greater than 80, and has asymmetrically-lobed ovaries); at the present, the only category 3 species that has been reported in the Western Atlantic Ocean is Acanthobothrium tortum. Acanthobothrium marquesi n. sp. can be distinguished from A. tortum by length (26.1 cm vs. 10.6 cm), greater number of proglottids (1,549 vs. 656), a larger scolex (707 µm long by 872 µm wide vs. 699 µm long by 665 µm wide), larger bothridia (626 µm long by 274 µm wide vs. 563 µm long by 238 µm wide). This is the first report of a species of Acanthobothrium from the Mexican coast of the Gulf México.

Much progress has recently been made in revising the taxonomic assignments of genera originally classified in the polyphyletic "Tetraphyllidea." Many of these genera, including Echeneibothrium, were accommodated in the order Rhinebothriidea. However, beyond this larger taxonomic action, little work has been conducted on this genus over the past 50 yr. Consequently, the criteria used for characterizing species of Echeneibothrium have lagged behind those typically used in more modern descriptions of elasmobranch-hosted cestode taxa. A series of collecting trips to Chile to obtain cestodes from the yellownose skate, Dipturus chilensis , provided a unique opportunity to apply modern morphological and molecular methods to investigate the 3 species of Echeneibothrium reported parasitizing this skate, specifically Echeneibothrium megalosoma, Echeneibothrium multiloculatum, and Echeneibothrium williamsi. In addition to redescribing all 3 species, using morphological data from light and scanning electron microscopy, maximum likelihood and bayesian inference phylogenetic analyses of the D1-D3 regions of the 28S rDNA gene were conducted to assess their relationships among other echeneibothriids for which comparable data are available. Sequencing of 59 specimens representing these 3 species of Echeneibothrium allowed us to assess the intra- and interspecific variation in the 28S rDNA gene. The redescriptions use standardized terminology for scolex morphology, proglottid anatomy, and microthrix forms and pattern; they also expand on the original descriptions to include data on scolex size, ovary size, vas deferens and vaginal configurations, testes arrangement, and genital pore position. Our morphological work led to a major reinterpretation of the scolex morphology with the recognition that all 3 species bear an apical bothridial sucker, rather than an apical loculus, prompting emendation of the diagnosis for the family Echeneibothriidae. The presence of a band of spinitriches at the apex of the apical modification of the scolex proper seems to represent an important feature for distinguishing the 2 portions of the myzorhynchus across species. Intraspecific variation ranged from 0 to 7 bp across species and interspecific variation ranged from a low of 39-46 bp between E. williamsi and E. multiloculatum to a high of 61-66 bp between E. multiloculatum and E. megalosoma. Phylogenetic analyses indicate that the 3 species of Echeneibothrium hosted by the yellownose skate are not each other's closest relatives, suggesting multiple colonization events of D. chilensis have occurred. Further phylogenetic investigation is also likely to confirm the status of the genus Pseudanthobothrium as a synonym of Echeneibothrium because its species generally group among members of Echeneibothrium.

Three new species of Aberrapex Jensen, 2001 (Cestoda: Lecanicephalidea) have been collected from species of Myliobatis Cuvier along the coast of Argentina. Aberrapex ludmilae sp. n. parasitises M. goodei Garman in the San Matías Gulf. This species is unique in a combination of features including the shape and extent of the uterus, scolex size, testis distribution, and by lacking an external seminal vesicle and postovarian vitelline follicles. Aberrapex sanmartini sp. n. from M. goodei in San Blas Bay and A. vitalemuttiorum sp. n. from M. ridens Ruocco, Lucifora, Díaz de Astarloa, Mabragaña et Delpiani in coastal waters off Buenos Aires Province, are distinguished from all other congeners by the microthrix pattern on the scolex surface, uterus shape and extension, position of the connection of the uterine duct to the uterus, presence and extension of an external seminal vesicle, and its overall size and number of proglottids. The specimens of M. goodei and M. ridens show locally distinct cestode faunas, which are correlated with well-defined biogeographic regions. Some of these areas correspond with mating and nursery zones for species of Myliobatis.

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 lecanicephalidean genus is erected for cestodes previously recognised as "New Genus 12" (Polypocephalidae) in a phylogenetic analysis of the interrelationship of members of this order. Examination of the cestode fauna of the mangrove whipray, Urogymnus granulatus (Macleay) (Myliobatiformes: Dasyatidae) from the Solomon Islands and northern Australia revealed the existence of specimens representing two new species, consistent in morphology with "New Genus 12." Corollapex gen. n. is unique among the 24 valid lecanicephalidean genera in its possession of an apical organ in the form of an external retractable central disk surrounded by eight concave muscular, membrane-bound pads and an internal heterogeneous glandular component. The two new species described herein, Corollapex cairae sp. n. (type species) and Corollapex tingoi sp. n., differ from one another in overall size and number of mature and immature proglottids, and are noted to demonstrate a differential distribution between mature and juvenile host individuals. Additional species diversity in the new genus, beyond C. cairae sp. n., C. tingoi sp. n., and "New Genus 12 n. sp. 1" of Jensen et al. (2016) is suggested. Corollapex gen. n. appears to be restricted to dasyatid hosts in the Indo-West Pacific region.

Acanthobothrium ningdense n. sp. and Acanthobothrium guanghaiense n. sp. are described from the spiral intestine of the whip stingray, Dasyatis akajei (Müller & Henle). Acanthobothrium ningdense n. sp. is reported based on 38 cestode specimens collected at five locations along the Chinese coast, i.e. Taizhou, Zhejiang Province, Ningde and Xiamen, Fujian Province, Taishan, Guangdong Province and Sanya, Hainan Province between 2012 and 2015. Acanthobothrium ningdense n. sp. belongs to the "species category 4" characterised by Ghoshroy & Caira. Among category 4 Acanthobothrium species, A. ningdense n. sp. most closely resembles A. micracantha Yamaguti, 1952 and A. latum Yamaguti, 1952, both from the same host species. Acanthobothrium ningdense n. sp. differs from A. micracantha by lacking a long robust hook handle that connects the bases of medial and lateral hooks, and from A. latum by having much longer axial prongs than abaxial prongs. Acanthobothrium guanghaiense n. sp., only found in Taishan, Guangdong Province in 2014, is a category 2 Acanthobothrium species. Among category 2, A. guanghaiense n. sp. most closely resembles A. semnovesiculum Verma, 1928, but can be differentiated from it by the arrangement of testes (in two staggered columns rather than in two tandem columns), and the shape of abaxial prong (nearly straight instead of with a conspicuous curve in the proximal portion). Acanthobothrium macrocephalum Wang & Yang, 2001 is redescribed based on new specimens collected from D. akajei in Guanghai and Sanya. With this study, the total number of Acanthobothrium species reported from D. akajei is brought to nine. It is possible that some of the species of Acanthobothrium previously reported from D. akajei not encountered in this study may have geographically restricted distributions, as was observed here for A. guanghaiense n. sp..

Acanthocotyle urolophi sp. nov. is described from the skin of the banded stingaree, Urolophus cruciatus (Lacépède, 1804). This is the first acanthocotylid to be described from Australian waters. Acanthocotyle urolophi sp. nov. is distinguished from other species of Acanthocotyle by a combination of the number of vitelline follicles 38 (33-46) and the number of rows 35 (32-37) of sclerites on the pseudohaptor. In addition, Acanthocotyle urolophi sp. nov. has no germarial appendix and no uterine "arm". A uterine receptaculum seminis was not identified in whole mounts. There is no penis papilla and no penis sclerite associated with the male reproductive opening. A brief description of the larva is provided. The diagnosis of the Acanthocotylidae Price, 1936 is amended and we review the Allacanthocotylinae Yamaguti, 1963, Lophocotylinae Yamaguti, 1963 and Pseudacanthocotylinae Yamaguti, 1963. We deem that these subfamilies are invalid and that the family now comprises only the subfamily Acanthocotylinae and the genus Acanthocotyle. The validity of species previously assigned to the Acanthocotylidae (sensu Yamaguti, 1963) is discussed and a key to what we consider to be the valid species in the family is also provided.

This study presents the first comprehensive phylogenetic analysis of the interrelationships of the morphologically diverse elasmobranch-hosted tapeworm order Lecanicephalidea, based on molecular sequence data. With almost half of current generic diversity having been erected or resurrected within the last decade, an apparent conflict between scolex morphology and proglottid anatomy has hampered the assignment of many of these genera to families. Maximum likelihood and Bayesian analyses of two nuclear markers (D1-D3 of lsrDNA and complete ssrDNA) and two mitochondrial markers (partial rrnL and partial cox1) for 61 lecanicephalidean species representing 22 of the 25 valid genera were conducted; new sequence data were generated for 43 species and 11 genera, including three undescribed genera. The monophyly of the order was confirmed in all but the analyses based on cox1 data alone. Sesquipedalapex placed among species of Anteropora and was thus synonymized with the latter genus. Based on analyses of the concatenated dataset, eight major groups emerged which are herein formally recognised at the familial level. Existing family names (i.e., Lecanicephalidae, Polypocephalidae, Tetragonocephalidae, and Cephalobothriidae) are maintained for four of the eight clades, and new families are proposed for the remaining four groups (Aberrapecidae n. fam., Eniochobothriidae n. fam., Paraberrapecidae n. fam., and Zanobatocestidae n. fam.). The four new families and the Tetragonocephalidae are monogeneric, while the Cephalobothriidae, Lecanicephalidae and Polypocephalidae comprise seven, eight and four genera, respectively. As a result of their unusual morphologies, the three genera not included here (i.e., Corrugatocephalum, Healyum and Quadcuspibothrium) are considered incertae sedis within the order until their familial affinities can be examined in more detail. All eight families are newly circumscribed based on morphological features and a key to the families is provided. Aspects of morphological evolution and host associations are discussed in a phylogenetic context for each family and for the order as a whole. Lecanicephalidean genera lacking apical structures were confirmed as the earliest diverging lineages. Proglottid anatomy was determined to be much more conserved and indicative of phylogenetic affinities than scolex morphology. Collectively, the Lecanicephalidea parasitize three of the four orders of Batoidea-their almost exclusive absence from skates (Order Rajiformes) appears to be real; only a few records from sharks exist. At the family level, the breadth of host associations is correlated with taxonomic diversity of the family. The degree to which factors such as intermediate host use or host specificity at any stage in the life-cycle shape these patterns is currently unknown., (Copyright © 2016 Australian Society for Parasitology Inc. Published by Elsevier Ltd. All rights reserved.)

The microthrix pattern of adult Parachristianella indonesiensis Palm, 2004 is described for the first time based on the standardized terminology for microtriches, and the generic diagnosis of Parachristianella Dollfus, 1946, regarding microtriches, is also amended. The surface ultrastructure of P. indonesiensis consists of three kinds of microtriches so that the proximal and distal bothrial surfaces are covered respectively with acicular to short capilliform filitriches and aristate gladiate spinitriches interspersed with capilliform filitriches. The remainder of the scolex bears capilliform filitriches. The surface of the proglottid is adorned with acicular filitriches. There was no significant difference in the length of the capilliform filitriches covering the different parts of the scolex peduncle. For the first time, the uniciliated papillae adorned with filitriches as sensory receptors are reported from this parasitic species. The partial rrnl of P. indonesiensis is also presented as the only known sequence of this gene in the superfamily Eutetrarhynchoidea Dollfus, 1969. The pairwise comparisons of rrnl between P. indonesiensis and the trypanorhynch species for which rrnl sequence is available were in accordance with the last phylogeny presented for the order Trypanorhyncha. The infection parameters (prevalence, mean abundance, and mean intensity) of P. indonesiensis parasitizing Pastinachus sephen (Forsskål) from the eastern Persian Gulf were also compared during approximately 5 years, and no significant differences were observed. This may be interpreted as the stability of the factors playing important roles in the life cycle of P. indonesiensis in the eastern Persian Gulf during this period.

The sensory organs in tegument of two trypanorhynchean species--Nybelinia surmenicola (plerocercoid) and adult Parachristianella sp. (Cestoda, Trypanorhyncha)--were studied with the aim of ultrastructural description and a comparative analysis. The Nybelinia surmenicola plerocercoid lacks papillae with sensory cilia on the bothria adhesive surface. We found an unciliated sensory organ within the median bothria fold. This unciliated free nerve ending contains the central electron-dense disc, three dense supporting rings, and broad root. The nerve ending locates in the basal matrix under the tegument. The tegument of N. surmenicola has a number of ultrastructural features which make it significantly different from other Trypanorhyncha: (i) the tegumental cytoplasm has a plicated constitution in a form of high apical and deep basal folds, (ii) numerous layers of the basal matrix are presented in the subtegument, and (iii) the squamiform and bristlelike microtriches N. surmenicola lack the base and the basal plate. In contrast, numerous ciliated and unciliated receptors were found in Parachristianella sp.: six types on the bothria and one type in the strobila tegument. Ultrastructural constitution of sensory organs in the form of ciliated free nerve endings as well as unciliated basal nerve endings of Parachristianella sp. has many common features inside Eucestoda. In comparison with other Trypanorhyncha, all Nybelinia species studied have less quantity of the bothrial sensory organs. This fact may reflect behavioral patterns of Nybelinia as well as phylogenetic position into Trypanorhyncha. Our observations of living animals conventionally demonstrate the ability of N. surmenicola plerocercoids to locomote in forward direction on the Petri dish surface. The participation of the bothrial microtriches in a parasite movement has been discussed.

Four new species are described in the lecanicephalidean genus Hornellobothrium from the ocellated eagle ray, Aetobatus ocellatus, from Australia and Indonesian Borneo. The species possess flat, laterally expanded immature proglottids and a small internal, glandular apical organ diagnostic of the genus. To date, only 2 valid species are recognized in the genus: Hornellobothrium cobraformis Shipley & Hornell, 1906 , from " Aetobatus narinari " from Sri Lanka, and Hornellobothrium extensivum Jensen, 2005 , from A. ocellatus , also from Australia. The new species differ from H. cobraformis in the shape of the apical organ and mature proglottids, and from H. extensivum in testes number and scolex dimensions. The new species can be distinguished from one another based on the following combination of features: Hornellobothrium gerdaae n. sp. possesses testes arranged in 2 layers and lacks post-ovarian vitelline follicles; Hornellobothrium iotakotta n. sp. has a single column of 4 testes and lacks spiniform microtriches on the distal bothridial surface; both Hornellobothrium kolossakotta n. sp. and Hornellobothrium najaforme n. sp. possess a single column of 6 testes, but H. najaforme n. sp. is a longer worm with a greater number of laterally expanded immature proglottids. Host associations suggest this genus is potentially limited to A. ocellatus . While reports of up to 9 lecanicephalidean congeners in a single host species exist, they are considered dubious and in need of verification, mainly because of host identification issues. This study demonstrates the presence of at least 5 species of Hornellobothrium in A. ocellatus . Insufficient sampling of this host species across the Indo-Pacific and at each locality may account for the apparent restricted distribution of species of Hornellobothrium.

Floriparicapitus n. gen. (Cestoda: Lecanicephalidea), with Floriparicapitus euzeti n. gen., n. sp. as its type, is erected to house 3 new tapeworm species and 2 known species that are transferred to the new genus, all parasitizing sawfishes and guitarfishes (order Rhinopristiformes) in Indo-Pacific waters. The new genus differs from the 21 valid lecanicephalidean genera in its possession of a large scolex bearing a laterally expanded apical organ in the form of a rugose sheet in combination with a cirrus conspicuously armed with spinitriches and 3 pairs of excretory vessels. It most closely resembles Lecanicephalum, but differs conspicuously in its possession of 3, rather than 1, pair of excretory vessels. Two new species are described from sawfishes: Floriparicapitus euzeti n. sp., from Pristis clavata and Floriparicapitus juliani n. sp. from Pristis pristis, both from Australia. Floriparicapitus plicatilis n. sp. is described from the guitarfish Glaucostegus typus in Australia and the guitarfish Glaucostegus thouin in Malaysian Borneo. Two species formerly assigned to Cephalobothrium are transferred to the new genus; Floriparicapitus variabilis ( Southwell, 1911 ) n. comb. from the sawfish Anoxypristis cuspidata in Sri Lanka and Floriparicapitus rhinobatidis ( Subhapradha, 1955 ) n. comb. from the guitarfish Glaucostegus granulatus in India. The species from guitarfish differ conspicuously from those parasitizing sawfish in their possession of only 4 ( F. plicatilis n. sp.) or 5 (F. rhinobatidis n. comb.) testes per proglottid versus 9 or more in the 3 sawfish-parasitizing species. The latter 3 species differ from one another in scolex width, acetabular size, number of proglottids, and cirrus sac size. As it stands, the new genus appears to be restricted to a subclade of the Rhinopristiformes consisting of the sawfishes and species of Glaucostegus.

Two morphologically disparate undescribed species of diphyllidean cestodes from the Yellowspotted skate, Leucoraja wallacei , from South Africa were included in a recent molecular phylogenetic study aimed at revising diphyllidean classification. From a molecular standpoint, these species were determined to be only distantly related to one another. One (originally referred to as Echinobothrium n. sp. 2) showed affinities with species of the batoid-parasitizing Echinobothrium sensu stricto and is described here as Echinobothrium marquesi n. sp. This species most closely resembles Echinobothrium joshuai, an affinity supported by the previous molecular study, but differs in the form of its "B" hooks and degree of overlap between bothria and cephalic peduncle. The other species (originally referred to as New genus n. sp. 1), although exhibiting the full complement of scolex armature, grouped among primarily shark-hosted genera, most of which lack or exhibit reduced scolex armatures. That unexpected result suggested that erection of a novel genus might be warranted, but morphological grounds supporting the action were not apparent at that time. The present study aimed to explore the morphology of this taxon in more detail. Light and scanning electron microscopy revealed this taxon to be exceptional in that its lateral hooklets, which are arranged in 2 clusters like those of Echinobothrium and Coronocestus, are unique in being arranged in anterior and posterior rows, rather than in a single row. Andocadoncum n. gen. is erected, with Andocadoncum meganae n. sp. as its type, to accommodate this taxon. A minor adjustment to the existing hook formula by presenting counts for the anterior hooklets (a) separately from the posterior hooklets (b) readily accommodates this new form. Leucoraja wallacei is distinctive among skates in hosting 2 different genera of diphyllideans. In the context of the previous molecular phylogenetic analyses, it appears that the presence of Andocadoncum n. gen. in this skate species may be the result of a host-switching event involving a weakly or unarmed shark-hosted taxon moving to a batoid-a shift that was likely accompanied by a return to the fully armed condition seen, for example, in other batoid-parasitizing members of the order.

Two new species of diphyllidean cestodes of the genus Echinobothrium, each hosted by a different skate species in the Raja miraletus complex, are described. Echinobothrium mercedesae n. sp. is described from R. cf. miraletus 2 off Senegal. Echinobothrium yiae n. sp. is described from R. cf. miraletus 1 off South Africa. Both species are small worms that differ from their 29 described congeners in the combination of number of cephalic peduncle spines per column, hook formula, number and arrangement of testes, and arrangement of vitelline follicles. They are easily distinguished from one another in that whereas the vitelline follicles of E. yiae n. sp. are circumcortical, they are lateral in E. mercedesae n. sp., and also in number of cephalic peduncle spines per column (14-17 vs. 10-12). Echinobothrium yiae n. sp. is also unusual in that the cephalic peduncle spines stop short of the anterior margin of the peduncle. In addition, although the paucity of available material precluded their formal description, evidence of 2 additional new species parasitizing R. miraletus also from Senegal is presented. In combination these worms provide support for the interpretation that what is currently recognized as Raja miraletus actually consists of a complex of geographically restricted species, rather than a polymorphic species of multiple parapatric or allopatrically distributed populations. This interpretation is not only supported by previously published molecular data, but also by newly collected morphological data involving differences in the color patterns of disc ocelli among host specimens of the 3 forms available as a result of digital efforts to ensure the accuracy of host identifications, which are also presented here.

A wild-caught specimen of the zonetail butterfly ray, Gymnura zonura (Bleeker), harboured numerous specimens of Hexacanalis folifer Cielocha & Jensen, 2011 (Systematic Parasitology, 79, 1-16; Cestoda: Lecanicephalidea) within its spiral intestine. The cestodes were primarily attached in single rows along the base of mucosal folds, each associated with a nodular mucosal thickening. Microscopically, the scolex was embedded within the submucosa and muscularis; the attachment sites were marked by ulceration and necro-proliferative inflammation demarcating the parasite from normal host tissues. Physical attachment of the cestode was restricted to the anterior portion of its scolex where presumed tegumental secretions from the apical organ contributed to a cementing intermediate layer blending with necrotic host cells. The presence of tegumental differentiation between the apical organ and the scolex proper, associated with presumed different roles in attachment, correlated with ultrastructural observations of the surface modifications on the scolex. Despite the locally severe pathological change, insignificant morbidity owing to this particular host-parasite relationship is suggested., (© 2012 Blackwell Publishing Ltd.)

A new species of Grillotia was found from teleosts and elasmobranchs along the Patagonian shelf off Argentina. Grillotia patagonica n. sp. is described from plerocerci recovered from gadiform (Moridae) and perciform fish (Cheilodactylidae, Bovichtidae, and Nototheniidae) and adults from the smallthorn sand skate Psammobatis rudis (Rajiformes, Rajidae). Grillotia patagonica most closely resembles species in the Grillotia erinaceus species complex (viz., Grillotia borealis, Grillotia brayi, Grillotia dollfusi, G. erinaceus, and Grillotia musculara) in having 4 hooks per principal row and 2 or more intercalary rows in the metabasal region, a band of hooks on the external surface of the tentacle, numerous proglottids, a hermaphroditic sac, internal and external seminal vesicles, uterine pore, and attachment of the retractor muscle in the mid region of the tentacular bulb. Grillotia patagonica n. sp. is unique among all valid species in the genus by having the hooks on the first row reduced and of a different shape from the rest of the metabasal rows. Despite the diversity of elasmobranchs that are available as definitive hosts for species in Grillotia, most species (10/14) are oioxenous or mesostenoxenous. The specificity for the last intermediate host is variable among species of Grillotia, with most plerocerci being oioxenous (5/15) or euryxenous (8/15). Host specificity is higher for the adult stage in the definitive host (mean HS(S) = 4.07) than for the plerocerci (mean HS(S) = 7.30). All the species with oioxenous specificity (either larvae or adults) have limited geographic distributions. Some previous records of species of Grillotia from fishes caught off Argentina require reconsideration, i.e., G. erinaceus, Grillotia minuta (reported as Grillotia bothridiopunctata) and G. borealis . An updated host-parasite checklist for the valid species of Grillotia is also presented along with new host records for Grillotia carvajalregorum.

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.

Three new species of diphyllidean cestodes are described from rhinobatid elasmobranchs of the genus Glaucostegus from the Indian and Pacific Oceans. All 3 new species have 11 apical hooks (6 type-A and 5 type-B hooks) and, therefore, differ in this feature from all but 5 of the 39 valid species of Echinobothrium. In addition, Echinobothrium tetabuanense n. sp. from Glaucostegus cf. typus in the Sulu Sea, Borneo differs from all of its congeners in the number and arrangement of hooklets, number of spines on the cephalic peduncle, and the number and distribution of testes. Echinobothrium sematanense n. sp. from Glaucostegus thouin in the South China Sea, Borneo, can be distinguished from its congeners based on the following combination of characters, i.e., small size (worms less than 1 mm long), number of hooklets, spines per column on the cephalic peduncle, and number of testes. Echinobothrium weipaense n. sp. from northern Australia is unique in the position of the cirrus sac and genital pore, both structures being well posterior and not overlapping the ovary. An emended description of the microthrix pattern on the scolex of Echinobothrium chisholmae from G. typus in Australia is also presented.

A new species of Oncomegas Dollfus, 1929 is described from the cowtail stingray, Pastinachus atrus (Macleay), collected in the Makassar Strait, Indonesian Borneo. Oncomegas trimegacanthus n. sp. possesses 2 oval bothria, gland cells within the bulbs, prebulbar organs, a distinctive basal armature with a single macrohook on the bothrial surface of the asymmetrical basal swelling, and a heteroacanthous, heteromorphous metabasal armature with 10 hooks per principle row. It differs from congeners by its possession of additional enlarged hooks at the base of the tentacle. Because of apparent morphological similarities, we suppress the genus Oncomegoides Beveridge & Campbell, 2005 with Oncomegas, and place the type and only species, Oncomegoides celatus Beveridge & Campbell, 2005 , within Oncomegas as Oncomegas celatus n. comb. Three species of Oncomegas , namely, Oncomegas paulinae Toth, Campbell & Schmidt, 1992, Oncomegas australiensis Toth, Campbell & Schmidt, 1992, and Oncomegas aetobatidis Campbell & Beveridge, 2009, differ from other species, possessing testes posterior to the ovary and a metabasal armature consisting of tiny, relatively homeomorphous hooks, with more than 14 hooks per principle row. Based on these morphological differences, a new genus, Hispidorhynchus n. gen., is erected, with Hispidorhynchus australiensis n. comb. as the type species.

New lecanicephalidean cestodes inhabiting the spiral intestine were investigated in 4 of the 6 known species of eagle rays of the genus Aetomylaeus Garman. Hosts examined consisted of 5 specimens of Aetomylaeus vespertilio from northern Australia, 5 of Aetomylaeus maculatus from Borneo, 10 of Aetomylaeus nichofii sensu stricto from Borneo, and 7 of Aetomylaeus cf. nichofii 2 from northern Australia. As a result of these new collections, 3 new genera and 6 new species of lecanicephalideans are formally described. Aetomylaeus vespertilio hosted the new genera and species Collicocephalus baggioi n. gen., n. sp. and Rexapex nanus n. gen., n. sp., as well as Aberrapex weipaensis n. sp. Aetomylaeus maculatus and A. nichofii sensu stricto hosted 3 new species of the novel genus Elicilacunosus , with the former eagle ray hosting Elicilacunosus sarawakensis n. sp. and the latter hosting both Elicilacunosus dharmadii n. sp. and Elicilacunosus fahmii n. sp. No new lecanicephalideans were described from A. cf. nichofii 2. Collicocephalus n. gen. is conspicuously unique among the genera of its order in possessing a large, retractable apical organ that, in cross-section, is transversely oblong, rather than round. Rexapex n. gen. is distinctive in its possession of an apical organ that bears 18 papilliform projections around its perimeter, and Elicilacunosus n. gen. is unlike any other known lecanicephalidean, or eucestode, in its possession of a region of musculo-glandular tissue along the midline of the dorsal and ventral surfaces of its proglottids, manifested externally as a tandem series of depressions. Among other features, A. weipaensis n. sp. differs from its congeners in its lack of post-ovarian vitelline follicles. All 6 new species were each restricted to a single species of Aetomylaeus . These records formally establish species of Aetomylaeus as hosts of lecanicephalideans. A summary of cestodes of myliobatid rays is presented.

Two new species of Rhinebothrium (Cestoda, Rhinebothriidea) from potamotrygonid stingrays in the Neotropical region are herein described, which raises the total number of Rhinebothrium species known from Neotropical freshwaters to five. Rhinebothrium corbatai sp. n., and Rhinebothrium mistyae sp. n. were both collected from Potamotrygon motoro (Müller et Henle), captured in four tributaries of the Paraná River in Santa Fé Province, Argentina. The new species can be distinguished from their congeners, and from each other, by a combination of various features, including worm size (length and number of proglottids), number of loculi per bothridium, microthrix pattern, size of the cirrus sac, and the extent of the vas deferens. The discovery of these new species from P. motoro supports the pattern of high host specificity in this cestode genus, and reinforces the notion that some of the previous records of the enigmatic R. paratrygoni may correspond to new species of Rhinebothrium yet to be described.

The cestode fauna of the darkspotted numbfish, Narcine maculata (Shaw) (Torpediniformes: Narcinidae), from Malaysian Bomrneo was examined for the first time. This work resulted in the discovery of a new genus and two new species of Anteroporidae (Lecanicephalidea). Sesquipedalapex comicus gen. n., sp. n. was erected on the basis of the peculiarities of its scolex, in particular its possession of an extremely long apical modification of the scolex proper, which readily distinguishes it from the other genus in the family. The genus is also distinct in its possession of acetabula that are in the form of suckers, rather than bothridiate in form. This species was found to deeply embed its elongate apical structure for much of its length within the intestinal mucosa, provoking a papilliform expansion of the outer wall of the spiral intestine at the site of attachment. The second new species, Anteropora klosmamorphis sp. n., is readily distinguished from its congeners on the basis of testis number and bothridial shape. Both new species are hyperapolytic. The diagnosis of Anteroporidae is amended to accommodate both new taxa. This increases the total number of genera in the family to two, and the total number of species to five.

We describe lesions associated with a seemingly intense infection of trypanorhynch plerocerci (Platyhelminthes: Cestoda: Trypanorhyncha) in the gastric wall of a female cownose ray, Rhinoptera bonasus (Myliobatiformes: Rhinopteridae) captured in the northern Gulf of Mexico. Grossly, the multitude of encapsulated, encysted plerocerci imparted a bumpy and cobbled appearance to the serosa of the stomach, and none was observed in any other tissue during routine parasitological necropsy. Histologically, the plerocerci were associated with severe intramural granulomatous gastritis, vascular ectasia and mesothelial polyposis with the exclusion of the mucosa. To our knowledge, this is the first published case study documenting platyhelminth-associated histopathological changes in the gastrointestinal tract of R. bonasus as well as that of the efficacy of immunocytochemical markers for smooth muscle actin, Factor VIII, S-100, and proliferating cell nuclear antigen in Myliobatiformes. It also may serve as a potential primer for much needed ecological investigations regarding the potential role of elasmobranchs as intermediate or 'paratenic' hosts in the life cycles of trypanorhynch cestodes., (© 2011 Blackwell Publishing Ltd.)

Rhinebothrium paranaensis n. sp. (Tetraphyllidea: Cestoda) is described from the freshwater stingray Potamotrygon falkneri collected in the Colastiné River, a tributary to the Lower Parana River in Argentina. The specimens were studied in detail using light and scanning electron microscopy and histology. Rhinebothrium paranaensis can be distinguished from all valid species in the genus using the following combination of characters: worm length, number of proglottids, diamond-shaped bothridia lacking constriction, number of loculi on the distal surface of bothridium, and the lack of vaginal sphincter and external seminal vesicle.

In total, 20 fanrays, Platyrhina sinensis (Rajiformes: Rhinobatidae), collected from the Taiwan Strait, were examined. A new phyllobothriid genus and species (Tetraphyllidea: Phyllobothriidae) was found and described. Biotobothrium platyrhina n. gen., n. sp. is assigned to the Rhinebothriinae Euzet, 1953, because it possesses 4 stalked bothridia that are each divided by septa into numerous small facial loculi but that lack apical suckers. The new genus and species is clearly different from any other in Rhinebothriinae in that each bothridium bears 2 sets of 5 small facial loculi, with 1 set located at each of the distal ends of the bothridium, rather than facial loculi throughout the entire distal surface of the bothridium, as is seen in all other genera in the subfamily. This character has not been previously reported for members of Rhinebothriinae. In addition, the testicular arrangement in the anterior quarter of the proglottid is unique. The posterior range of the vas deferens almost reaches the ovarian bridge, and the morphology of the cirrus sac and several other morphological characters are at variance with presently described species in the subfamily.

The phylogeography and host specificity of three monogenean species infecting different sites on the southern fiddler ray, Trygonorrhina fasciata (Rhinobatidae) in South Australia (SA) were studied: Branchotenthes octohamatus (Hexabothriidae: gills), Calicotyle australis (Monocotylidae: cloaca) and Pseudoleptobothrium aptychotremae (Microbothriidae: skin). Five rhinobatid species (Aptychotrema vincentiana, T. fasciata, Trygonorrhina sp. A, Aptychotrema rostrata and Rhinobatos typus) with distributions spanning west, south and east Australian coastal waters, were surveyed for monogeneans resembling the three species documented from T. fasciata in SA. The identities of hosts and parasites collected were investigated using the mitochondrial genes ND4 and Cytochrome b (cytb), respectively, in addition to the nuclear marker, Elongation factor 1-alpha (EF1a) for Pseudoleptobothrium. Genetic analyses confirmed that B. octohamatus is geographically widespread and displays little genetic structure, suggesting high levels of gene flow. It was collected from four rhinobatid species throughout its distribution and is not, therefore, host specific. For C. australis, genetic analyses revealed two discrete populations with a genetic divergence of ∼4%, one population occurring west of Bass Strait on two sympatric host species and the other population on the east coast, also occurring on sympatric host species. Similarly, for Pseudoleptobothrium, specimens collected west of Bass Strait were genetically distinct (∼3.5%) from those collected to the east. However, on the east coast, a third Pseudoleptobothrium population was revealed, separated by a genetic distance of >11%, indicating a morphologically cryptic species. Host preferences were indicated for each Pseudoleptobothrium lineage. These genetic discoveries are discussed in relation to life history characteristics of each monogenean species, highlighting the value of phylogeographic analyses to understand the parasite-host relationship.

Spiral intestines of 12 specimens of the dwarf whipray, Himantura walga, collected from Malaysian Borneo in 2002 and 2003, were examined for cestodes. These yielded a new species of Acanthobothrium (Tetraphyllidea) and a new species of Echinobothrium (Diphyllidea), both of which are described. Acanthobothrium marymichaelorum n. sp. is a category 1 species. It differs from all but 4 of its category 1 congeners in its possession of postovarian testes. It also differs from these 4 species in its possession of fewer testes, shorter length, fewer proglottids, and/or shorter posterior loculus. Echinobothrium minutamicum n. sp. differs from its congeners in its possession of outer hooks in the dorsal and ventral rostellar groups that are trifid; it is also the smallest member of its genus. The spiral intestine of H. walga consisted of 12 mucosal chambers. Most (89%) of the 35 specimens of E. minutamicum n. sp. for which chamber data were generated were found in chambers 2-4. In contrast, the 57 specimens of A. marymichaelorum n. sp. occurred throughout chambers 5-12, with 86% in chambers 6-10. The modes of attachment of both cestode species were similar, i.e., both embedded their scolex within the lumen of a mucosal crypt with the hooks and/or spines penetrating the lamina propria. Both also eroded the epithelial lining of the crypts and caused modest expansion of crypt diameter. Although the configuration of the mucosal surface may explain sites in which both species were able to attach, it does not explain their absence from other regions; histological sections and scanning electron microscopy showed the mucosal surface to be similar in configuration throughout the length of the spiral intestine. The cestode fauna of H. walga also included at least 1 species of rhinebothriine, 2 lecanicephalidean species, a trypanorhynch species, and 1-2 additional new species of Acanthobothrium. However, formal description of these species must await the collection of additional material, mature material, and/or the erection of the new genera. It is of note that the fauna of the dwarf whipray consists of a suite of unusually small taxa. Although the cestode genera reported here are generally consistent with those reported from other Himantura species, they are completely inconsistent with previous records from H. walga (as Trygon walga) in Sri Lanka. This suggests that either the original host identifications are suspect or that differences exist in the faunas of H. walga between these 2 localities.

The objectives of this study were to (1) describe the host range for 6 tetraphyllidean species and quantify their host specificity using 5 specificity indices; (2) determine the role of morphological determinants in the host specificity of tetraphyllideans by comparing villar and bothridial measurements of species examined herein; and (3) determine the role of a physiological component in the host specificity of tetraphyllideans by exposing tetraphyllideans to blood sera from different fish species and other solutions. Our results indicate that Echeneibothrium dubium abyssorum (ex Amblyraja radiata), Echeneibothrium canadensis (ex A. radiata), and Zyxibothrium kamienae (ex Malacoraja senta) exhibit the highest degree of specificity, followed by Echeneibothrium vernetae (ex Leucoraja erinacea and Leucoraja ocellata), Pseudanthobothrium hanseni (ex A. radiata and M. senta), and Pseudanthobothrium purtoni (ex Leucoraja erinacea and L. ocellata). However, these results vary based on the specificity index used. Compatible bothridial and villar measurements indicate that there is no morphological determinant of host specificity but that there is a morphological determinant to attachment site specificity. Our data indicate that attachment site specificity may also be phylogenetically determined. Additionally, the exposure of parasites to blood sera from various hosts confirms that host specificity in this system has a physiological determinant. Therefore, host specificity in this system is determined, at least in part, by physiological factors, whereas attachment site specificity is an extension of host specificity and is phylogenetically determined.

A new cestode, Macrobothridium sinensis n. sp., is described from the spiral intestine of Platyrhina sinensis from coast of Xiamen, China. It is the first record of the order Diphyllidea in China. The new species can be distinguished from the 3 previously described species of Macrobothridium by the testes number (16-24 vs. 29-37, 27-46, and 5-6, in M. rhynchobati, M. euterpes, and M. syrtensis, respectively) and a single-row arrangement of testes; M. sinensis is most similar to M. euterpes in strobila length (1.77-6.23 vs. 2-4.5 mm) and number of segments (6-8 vs. 5-9). In addition, M. sinensis differs from M. rhynchobati and M. syrtensis in the strobila length, ovary shape, and number of apical hooks. The uterine pore is the first described in the Diphyllidea; thus, the present uterine pore can now be used to distinguish Macrobothridium from Echinobothrium and Ditrachybothridium in the Diphyllidae.

Guidus n. gen. (Cestoda, Tetraphyllidea) is proposed for 3 cestode species from skates (Bathyraja spp.: Rajiformes, Rajidae). Members of Guidus differ from those of all other phyllobothriid genera in possessing 4 sessile and conspicuously muscular, saclike bothridia; a bothridial aperture toward anterior end of scolex, and perpendicular to scolex axis; aperture surrounded by a continuous sphincter of circular muscles, and 1 marginal accessory sucker. A new species, Guidus argentinense n. sp., is described, the diagnosis of Marsupiobothrium antarcticum is emended, and it is transferred to the new genus along with Marsupiobothrium awii. These species can be easily distinguished from G. argentinense by the presence of lappets in the bothridial margin. In addition, the diagnosis of Marsupiobothrium is emended based on the redescription of the type species, Marsupiobothrium alopias.

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.

Five new species of Acanthobothrium (Tetraphyllidea: Onchobothriidae) are described from the spiral intestine of the Freshwater whipray, Himantura chaophraya, in the Kinabatangan River in Malaysian Borneo. Based on criteria set forth in a previous categorization scheme for species of Acanthobothrium, these consist of 3 Category 1 species, Acanthobothrium asnihae n. sp., Acanthobothrium saliki n. sp., and Acanthobothrium zainali n. sp.; a Category 8 species, Acanthobothrium etini n. sp.; and a Category 2 species, Acanthobothrium masnihae n. sp.. Acanthobothrium asnihae n. sp. differs from all Category 1 species in its possession of a horizontal band of weak musculature that divides the posterior loculus in half. Among Category 1 species, A. saliki n. sp. differs from all but Acanthobothrium southwelli in its possession of postovarian testes. It differs from A. southwelli in its possession of fewer testes and a greater number of proglottids. Acanthobothrium zainali n. sp. differs from the 25 other Category 1 species in a combination of overall size, muscular pad and hook shape, arrangement and number of testes, ovary configuration in cross section, position of ovarian isthmus, and genital pore position. Acanthobothrium etini n. sp. is distinguished from all 5 other Category 8 species in its lack of testes from the proglottid antiporal and postporal regions and in testis number. Acanthobothrium masnihae n. sp. differs from the 35 other Category 2 species in its possession of fewer testes, postporal testes, or a greater number of proglottids. A key to Acanthobothrium species parasitizing H. chayophraya is presented. This represents the first report of Acanthobothrium from freshwater stingrays belonging to a family other than the Potamotrygonidae.

Examination of the spiral intestines of 44 freshwater stingrays, Potamotrygon motoro, from tributary rivers of the Parana River in Argentina, allowed for the collection of specimens of an undescribed species of Acanthobothrium. Acanthobothrium ramiroi n. sp. can be distinguished from all other congeners by the combination of the following characters: asymmetrical hooks (medial and lateral hooks conspicuously different in size and form, with axial prong of medial hooks stouter than abaxial prong), hook size (total length of medial hooks up to 242 microm, total length of lateral hooks up to 239 microm), bothridia not fused to the scolex proper at posterior ends, worm size (51-84 mm long), and the presence of a conspicuous vaginal sphincter. The new species is different from all other species of Acanthobothrium in freshwater potamotrygonids, except Acanthobothrium terezae, in having conspicuous asymmetrical hooks. The main differences that allow for the distinction between A. ramiroi and A. terezae include hook size, the way the bothridia are attached to the scolex proper, and the shape of the older gravid segments. The discovery of a new species of Acanthobothrium from a potamotrygonid extends our understanding of the diversity of the genus in freshwater stingrays in South America.

Adult tetraphyllidean tapeworms (Platyhelminthes: Eucestoda) from the spiral intestines of 3 species of potamotrygonid stingrays (Paratrygon aiereba, Potamotrygon castexi, and Portamotrygon motoro) in the Madre de Dios river in Peru were found to host numerous cysts embedded in their parenchymal tissues. Histological sections of the cysts revealed the presence of a scolex bearing 4 suckers and an unarmed apical organ consistent with larval stages of both Cyclophyllidea and Proteocephalidea. To further elucidate their identities, partial 28S ribosomal DNA (rDNA) sequences were characterized from 3 cysts and 4 adult Rhinebothrium spp. 'host' worms and screened against all available cestode 28S rDNA data. Initial BLAST screening and subsequent alignment ruled out the possibility that the cysts were cyclophyllidean, and the cyst and adult sequences were thus aligned together with all available lecanicephalidean, litobothriidean, proteocephalidean, and tetraphyllidean sequences. Sequences from all 3 cysts were identical, and phylogenetic analysis clearly placed them among derived members of the Proteocephalidea, although no exact match was found. Sequences from the adult host worms formed 2 identical pairs and grouped together with other tetraphyllidean species from rays. These results are compared with records of hyperparasites of South American catfish cestodes. This is the first confirmed record of a proteocephalidean cestode parasitizing a tetraphyllidean cestode.

Extensive new collections of specimens of Rhinebothroides spp. from a wide range of Neotropical freshwater potamotrygonid stingrays in numerous localities permitted critical evaluation of the validity of the 7 nominal species in the genus. Some qualitative characters previously used in the taxonomy of this group are highly variable within and among populations, and most morphometric and meristic characters thought to have taxonomic value for this group overlapped among several previously defined nominal species. Characters that were consistent throughout populations and that clearly define morphotypes supported recognition of 4 species: R. freitasi (syn. R. circularisi and R. venezuelensis), R. glandularis (syn. R. mclennanae), R. moralaurai, and R. scorzai. We provide a taxonomic review of the genus, 8 additional host records, 7 new localities, and an identification key.

Scalithrium gen. n. (Cestoda, Tetraphyllidea) is proposed with Scalithrium minimum (Van Beneden, 1850) n. comb., parasite of Dasyatis pastinaca (Elasmobranchii, Dasyatidae) as type-species. The new genus Scalithrium (Tetraphyllidea, Phyllobothriidae, Rhinebothriinae) is erected for several species previously included in the genus Rhinebothrium. These species have a scolex with four bothridia, the distal surface of which is divided by transverse septa in a single row of loculi. Scalithrium minimum (Van Beneden, 1850) n. comb. is redescribed from specimens collected from the type-host Dasyatis pastinaca in Tunisia and becomes the type-species of the new genus. After Braun (1900) Echeneibothrium variabile Van Beneden, 1850 is considered as type-species of the genus Echeneibothrium. Species of Rhinebothriinae to be transferred into the genus Scalithrium are discussed and a key is proposed for the eight species.

Echinobothrium chisholmae n. sp. is described from Rhinobatos typus Bennett (Rhinobatidae), collected from Heron Island, Great Barrier Reef, Australia. E. chisholmae differs from all congeners in possessing 11 hooks in each dorsal and ventral group on the rostellum and groups of 3-6 hooklets on either side of the hooks. A single metacestode of E. chisholmae was collected from the decapod crustacean Penaeus longistylus Kubo. Yellow pigmentation of the cephalic peduncle in immature adults is caused by the accumulation of large vesicles in the distal cytoplasm of the tegument. The vesicles probably provide materials for spine formation. Ultrastructural examination of the rostellar musculature revealed that the muscles are stratified (striated-like), consisting of a periodic repetition of sarcomeres separated by perforated Z-like lines that are oblique to the long axes of the myofilaments.

Anindobothrium n. gen. is proposed to accommodate Caulobothrium anacolum inhabiting Himantura schmardae from Colombia, and 2 new species, one inhabiting Potamotrygon orbigny in Brazil and the other inhabiting Paratrygon aereiba in Venezuela. Members of the new genus resemble members of Pararhinebothroides, Rhinebothroides, and Anthocephalum by having bothridia with poorly differentiated apical suckers and vasa deferentia expanded into external seminal vesicles. It further resembles Pararhinebothroides, Rhinebothroides, and Anthocephalum cairae by having vas deferens inserted near the poral rather than aporal end of the cirrus sac. The 3 species assigned to the new genus form an apparent monophyletic group, based on the possession of 3 putative synapomorphies: (1) genital pores in the anterior 1/4 of the proglottid, a trait that is unusual, but not unique, among phyllobothriids; (2) anteroventral ovarian lobes converging to the center of the proglottid, a character not previously reported for phyllobothriids; and (3) ovarian lobes comprising a loose network of digitiform processes.

As a result of an investigation of metazoan parasites of elasmobranch fishes in the Gulf of Gabès, Tunisia, we discovered 2 new species of diphyllidean cestodes. Macrobothridium euterpes n. sp. is described from the spiral intestine of Rhinobatos rhinobatos, and Macrobothridium syrtensis n. sp. from the spiral intestine of Rhinobatos cemiculus. Macrobothridium euterpes is distinguished from the only other species in the genus (Macrobothridium rhynchobati) by the number of rostellar hooks, size, genital pore position, vagina position, and ovary shape. Macrobothridium syrtensis is distinguished from M. rhynchobati by the hook morphology, testis number, and overall size, and from M. euterpes by the number of rostellar hooks, testis number, genital pore position, vagina position, and ovary shape. These are the first 2 species added to the genus since its establishment in 1989. A standardized formula for representing the number and arrangement of rostellar hooks in diphyllidean species is presented.

Two new tetraphyllidean species, Duplicihothrium caircae n. sp. and D. paulum n. sp., are described from Pacific cownose rays Rhinoptera steindachneri collected from the Gulf of California. D. cairae n. sp. differs from the only other known species in this genus, D. minutum, in its possession of two posteriorly bifurcating longitudinal septa on each bothridium and having a greater number of loculi per bothridium, wider bothridia and a greater number of segments per worm. D. paulum n. sp. differs from the above two species in being shorter, having two continuous longitudinal septa on each bothridium, rather than posteriorly bifurcating or absent longitudinal septa, having a greater number of loculi per bothridium and having fewer segments per worm. The generic diagnosis of Duplicibothrium is amended to reflect the inclusion of the two new species. Species of Duplicibothrium appear to be restricted to host species of the genus Rhinoptera. Systematic information gained from the study of these two new species supports the monophyly of the tetraphyllidean family Serendipidae and suggests that Duplicibothrium shares a phylogenetic heritage with species in the genera Glyphobothrium and Serendip, taxa that are also parasitic in cownose rays. Evidence for a phylogenetic relationship between serendipid species and species of Dioecotaenia is discussed, as are potential molluscan hosts for cestode species in Duplicibothrium and Dioecotaenia.