The cestode fauna of the darkspotted numbfish, Narcine maculata (Shaw) (Torpediniformes: Narcinidae), from Malaysian Borneo 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.
Recognised for their diversity in apical structure morphology, members of the cestode order Lecanicephalidea Wardle et McLeod, 1952 known to date exhibit relatively mundane and uniform acetabular morphology. A new lecanicephalidean genus, Zanobatocestus gen. n., is proposed for two new species found parasitising the spiral intestine of the striped panray, Zanobatus schoenleinii (Müller et Henle), off Senegal that are highly unusual in acetabular morphology. Unlike the members of the 21 recognised lecanicephalidean genera, which possess simple, uniloculate suckers or bothridia, Zanobatocestus minor sp. n. and Z. major sp. n. possess biloculate bothridia. The form of their apical structures and cocoons readily distinguish the two new species from one another. Zanobatocestus minor sp. n. exhibits an apical modification of the scolex proper that is narrow and elongated, an apical organ that is small and internal, and eggs in cocoons forming linear strands, whereas Z. major sp. n. exhibits an apical modification of the scolex proper that is wide and short, an apical organ that is extensive and primarily external, and eggs in cocoons primarily as doublets with bipolar filaments. Given the typically high host specificity of lecanicephalidean cestodes, as parasites of the only genus and species currently considered valid in the family Zanobatidae, Zanobatocestus gen. n. is likely to remain one of the less specious lecanicephalidean genera.
A new species of tetraphyllidean cestode in the genus Trilocularia is described from an undescribed shark species, Squalus cf. mitsukurii, off the coast of South Africa. Trilocularia eberti sp. n. is the second known member of its genus, and like its congener, T. gracilis (Olsson, 1866-1867) Olsson, 1869, is extremely hyperapolytic, dropping proglottids from its strobila while they are still very immature. Characteristic of the genus, it possesses a distinctive scolex with triloculated bothridia, but differs conspicuously from its congener in its possession of an anterior loculus that is much larger in width relative to the paired posterior loculi, and also in its possession of an anterior, enlarged region of its free proglottids that is triangular with a slit-like ventral aperture, rather than rounded and cup-like. This anterior region of the free proglottid is used in attachment, and its development is described. For assessment of fecundity, an attempt was made to record all free proglottids of all ages found in both host individuals, and yielded an average estimate of 362 free proglottids being produced per individual worm of T. eberti sp. n. Both Trilocularia species parasitize sharks of the genus Squalus, and given the host specificity typically exhibited by tetraphyllideans and preliminary examinations of other members of this shark genus, it is likely that other Squalus species will be found to host additional new Trilocularia species.
Five new species of Acanthobothrium van Beneden, 1850 from the spiral intestine of a specimen of an unusual species of Himantura from the Arafura Sea off northern Australia are described. Acanthobothrium oceanharvestae sp. n. is one of 26 category 1 species (sensu Ghoshroy and Caira 2001) lacking post-ovarian testes; it differs from these in total length, number of proglottids, number of testes, cirrus sac size and details of the terminal genitalia. Acanthobothrium popi sp. n. is unique among category 2 species in its possession of post-ovarian testes. Acanthobothrium rodmani sp. n. is a category 6 species distinct from all congeners in the dense blade-like spinitriches on the distal surfaces of its anterior-most bothridial loculi and conspicuously tapered posterior bothridial margins, which are reflexed anteriorly. Acanthobothrium romanowi sp. n. differs from most other category 1 species in that its genital pore is distinctly posterior. It differs from the remaining category 1 species in size, testis number, cephalic peduncle microthrix form, proglottid shape, and bothridial loculus dimensions. Acanthobothrium zimmeri sp. n. is among the six category 1 species with post-ovarian testes. It differs from these species in total length, ovary shape, number of proglottids and testes and vas deferens extent. This brings the number of Acanthobothrium species with post-ovarian testes to 10, all of which are Indo-Pacific in distribution, and 7 of which parasitize Himantura species. A key to the five new species parasitizing Himantura sp. is provided. Sequence data for the D1-D3 region of 28S rDNA for the five new species and two congeners parasitizing other Himantura species shows no intraspecific variation. Analysis of these and comparable data for two species available in GenBank (Acanthobothrium parviuncinatum and Acanthobothrium sp. 1) showed an interspecific variation of 0.7-11.3% among species pairs. Bayesian, Likelihood and Parsimony phylogenetic analyses of these data for these nine species indicate that the five new species parasitizing Himantura sp. are generally not each others' closest relatives.
As part of a metazoan parasite survey of elasmobranchs from Malaysian Borneo, specimens of Rhoptrobothrium Shipley et Hornell, 1906 were collected from the eagle rays Aetomylaeus maculatus (Gray) and Aetomylaeus niehofii (Bloch et Schneider). The type species is redescribed from its type host, and a neotype specimen is designated. In addition, three new species of Rhoptrobothrium are described: R. chongi sp. n., R. gambangi sp. n. and R. limae sp. n. Rhoptrobothrium myliobatidis conspicuously differs from the three new species in its lack of a secondary areola; R. limae is distinguished from R. chongi and R. gambangi based on its greater total length; R. chongi possesses conspicuously stalked remi, while R. gambangi possesses short remi, often folded anteriorly. Rhoptrobothrium is somewhat unusual among tetraphyllideans in its possession of a ''metascolex,'' a character it shares with other taxa in the Thysanocephalinae (i.e., Myzocephalus Shipley et Hornell, 1906, Myzophyllobothrium Shipley et Hornell, 1906 and Thysanocephalum Linton, 1889). The morphology of the ''metascolex'' of Rhoptrobothrium is investigated and new terminology is suggested to standardise the names given to structures constituting a metascolex. As a result, Rhoptrobothrium is considered to possess cephalic peduncle extensions, termed remi. In Rhoptrobothrium, each remus bears, at its distal end, a primary areola, and, in the case of the three new species, also a secondary areola proximal to the primary areola. Myzocephalus and Myzophyllobothrium are tentatively considered to possess remi; the configuration of the ''metascolex'' of Thysanocephalum, however, is not considered homologous to the condition in the other three genera currently placed in the Thysanocephalinae.
Examination of seven specimens of an undescribed species of bamboo shark (Orectolobiformes: Hemiscylliidae), currently referred to as Chiloscyllium sp. 1, from Sri Lanka yielded three new species of 'tetraphyllidean' cestodes, one each in the genera Carpobothrium Shipley et Hornell, 1906, Spiniloculus Southwell, 1925, and Yorkeria Southwell, 1927. Carpobothrium marjorieae Caira, Otto et Jensen sp. n. differs from its three valid congeners in total length, number of testes, and size of the apical sucker of its anterior bothridial flap. Like Carpobothrium eleanorae Koontz et Caira, 2016 it possesses spherical eggs with highly elongate bipolar filaments, but its eggs are conspicuously larger than those of the latter species. Spiniloculus akshayi Caira, Otto et Jensen sp. n. can be distinguished from its five congeners in total length and number of proglottids. Yorkeria sachiniae Caira, Otto et Jensen sp. n. differs from its 16 congeners in number of proglottids and testes, total length, terminal proglottid, cirrus sac, and cephalic peduncle length, hook size, genital pore position, and bothridial shape. The eggs of Y. sachiniae Caira, Otto et Jensen sp. n. are spindle-shaped and bear a single long, polar filament. All three species were found to exhibit microthrix patterns on their scolices like those of the majority of their congeners that have been examined to date with scanning electron microscopy. Microtriches on the distal surfaces of the bothridial pouches of a species of Carpobothrium were characterised for the first time; C. marjorieae Caira, Otto et Jensen sp. n. was found to possess a modified form of gladiate spinithrix on this surface. Among the seven members of the genus, this is the first species of Chiloscyllium Müller et Henle that has been found to host representatives of all three of these 'tetraphyllidean' genera. However, unlike most of its congeners, a representative of the non-hooked genus Scyphophyllidium Woodland, 1927 was not found infecting Chiloscyllium sp. 1.
Three new species of diphyllidean cestodes are described from stingrays of the genus Pastinachus from the Indian and Pacific Oceans. All three new species differ from all but 10 of the 36 valid species of Echinobothrium van Beneden, 1849 in that they bear lateral hooklets arranged in a continuous band across each lateral surface of the rostellum joining the groups of dorsal and ventral apical hooks, rather than arranged in distinct dorsal and ventral groups, on each side of the apical hooks. Echinobothrium nataliae sp. n. ex Pastinachus solocirostris off Borneo differs from other relatives in the following combination of characters: number of spines in each column on the cephalic peduncle, number of lateral hooklets and number of apical hooks. Echinobothrium reginae sp. n. ex Pastinachus cf. sephen off Madagascar differs from all congeners in the following combination of characters: number of hooklets and number of spines on the cephalic peduncle. Echinobothrium vojtai sp. n. from an undescribed species of Pastinachus off Borneo differs from its congeners in the following combination of characters: number of hooklets, number of apical hooks and number of spines in each column on the cephalic peduncle. The hook formula was modified to accommodate species with lateral hooklets arranged in a continuous band as follows: {(LH) AH(A)/AH(B)}, where (LH) is the total number of lateral hooklets in a band. A table of all nominal species of Echinobothrium, showing their taxonomic status, type host and locality, number of peduncle spines in a column and hook formula, is presented. Macrobothridium djeddensis and Macrobothridium sinensis are transferred to Echinobothrium. In addition, Echinobothrium deeghai is considered as a species inquirenda.
The type species (Spiniloculus mavensis Southwell, 1925) of the previously monotypic tetraphyllidean genus Spiniloculus Southwell, 1925 is redescribed from the type material from Moreton Bay, Australia. As a consequence the identity of this species is definitively resolved. Three new species in the genus, all collected from Chiloscyllium punctatum Müller et Henle (brownbanded bambooshark), in Borneo, are described. Spiniloculus calhouni sp. n. conspicuously differs from all three of its congeners in its possession of post-poral testes. Spiniloculus fylerae sp. n. and Spiniloculus paigeae sp. n. differ from their two other congeners in that they are relatively small worms (4-6.5 and 2.2-5 mm in total length, respectively) with fewer than 30 proglottids. They can be distinguished from one another in that, while the vitelline follicles are interrupted at the level of its ovary in S. fylerae, this is not the case in S. paigeae. Furthermore, whereas the cirrus sac of the former species is pyriform, it is elongate-oval in the latter species. This brings the total number of species in the genus to four, and lends support to the suggestion that the original identity of the type host of S. mavensis as Mustelus sp. was in error. This work also extends the range of the genus to include the island of Borneo. A key to the species of Spiniloculus is provided. Morphological data generated here, using both light and scanning electron microscopy, support the suggested close affinities between Spiniloculus and Yorkeria Southwell, 1927, both of which parasitize bamboosharks.
Recent collections of cestode parasites from two species of the myliobatid genus Aetomylaeus Garman from several localities in the Pacific Ocean resulted in the discovery of two new species of Halysioncum Caira, Marques, Jensen, Kuchta et Ivanov, 2013. Halysioncum gibsoni sp. n. from Aetomylaeus maculatus (Gray) in the South China Sea off Borneo differs from all of its congeners in having the following combination of characters: 27 apical hooks (14 type A and 13 type B hooks), 11-12 lateral hooklets, 22-28 spines per column on the cephalic peduncle, testes distributed in a single column and an internal seminal vesicle. Halysioncum arafurense sp. n., recovered from Aetomylaeus cf. nichofii 2 (sensu Naylor et al. 2012b) in the Arafura Sea off the Wessel Islands, Northern Territory, Australia, can be distinguished from its congeners based on the following combination of characters: 23 apical hooks (12 type A and 11 type B hooks), the number of lateral hooklets (9-11), the number of spines per column on the cephalic peduncle (20-24), the number and distribution of the testes (13-15 testes in two irregular columns), and the distribution of vitelline follicles (interrupted dorsally at the level of the ovarian lobes). Both species represent the first verified records of diphyllideans from eagle rays of the genus Aetomylaeus and formally extend the host associations of diphyllideans to include a third genus of Myliobatiformes. The myliobatiforms are indeed an understudied group of available hosts for diphyllideans and represent interesting target hosts if the diversity of diphyllidean tapeworms is to be fully estimated and understood.