Three new species of <Emphasis Type="Italic">Anthocephalum</Emphasis> Linton, 1890 (Cestoda: Tetraphyllidea) from dasyatid stingrays of the Gulf of California

Three new species of Anthocephalum Linton, 1890 are described from dasyatid stingrays collected in the Gulf of California. Anthocephalum michaeli n. sp. is described from Dasyatis longus (Garman). This species most closely resembles A. alicae Ruhnke, 1994, but differs from this species in proglottid number. A. lukei n. sp. is also described from D. longus. This new species is most similar to A. cairae Ruhnke, 1994, but differs from that species in marginal loculi number and number of proglottids. The third new species, A. currani n. sp., is described from D. brevis (Garman). This species is most similar to A. centrurum (Southwell, 1925) Ruhnke, 1994, but differs from that species in marginal loculi number, number of testes and ovarian length. Phyllobothrium kingae Schmidt, 1978 is also consistent in morphology with species of Anthocephalum and is transferred to this genus, forming the new combination Anthocephalum kingae n. comb. This species most closely resembles A. michaeli n. sp., but differs in testicular shape. This brings the total number of species of Anthocephalum to nine. The transfer of the species Phyllobothrium arctowskii Wojciechowska, 1991, P. georgiense Wojciechowska, 1991, P. rakusai Wojciechowska, 1991 and P. siedleckii Wojciechowska, 1991 to Anthocephalum is not warranted, as these four species lack a posteriorly recurved cirrus-sac and a sinuous vagina, and have vitelline follicles uninterrupted by the ovary. Of the nine known species, all are parasitic in batoid fishes, and six are found in species of Dasyatis Garman. The phylogenetic status of Anthocephalum species in relationship to Rhinebothroides Mayes, Brooks & Thorson, 1981, Pararhineothroides Zamparo, Brooks & Barriga, 1999 and other rhinebothriin taxa is discussed.     

A new species of Calliobothrium (Tetraphyllidea: Onchobothriidae) from the whiskery shark, Furgaleus macki, in Australia

Calliobothrium pritchardae n. sp. is described from the whiskery shark, Furgaleus macki (Whitley, 1943), collected from Young Rocks in South Australia. This species differs from the 7 other known species of Calliobothrium in its possession of 2 rather than 3 posthook loculi. It is a further distinguished from all other Calliobothrium species except Calliobothrium evani in its possession of asymmetrical hooks. The lack of an accessory piece between the bases of the axial hooks and medial axial hook bases that are longer than the lateral axial hook bases further distinguish the new species from C. evani in which an accessory piece is present and the relative lengths of the medial and lateral axial hook bases are reversed. Scanning electron microscopy reveals spiniform microtriches on the neck and proximal bothridial surfaces, filiform microtriches on the apical bothridial surfaces, and a combination of the 2 microthrix types on the distal bothridial surfaces. The extended bases of the axial hooks are densely covered with spiniform microtriches. In addition, the number of testes within worms decreases substantially in progressively posterior (older) segments. The generic diagnosis of Calliobothrium is emended to include this species with 2 rather than 3 posthook loculi.     

Orygmatobothrium spp. (Cestoda: Tetraphyllidea) from triakid sharks in Argentina: redescription of Orygmatobothrium schmitti and description of a new species

Two species of Orygmatobothrium were found inhabiting triakid sharks collected from the coast of Buenos Aires Province, Argentina. Orygmatobothrium schmittii from Mustelus schmitti is redescribed, including new information on the microtrich pattern. Orygmatobothrium juani n. sp. from Mustelus fasciatus can be distinguished from all other species in the genus using the following combination of characters: worm length, number of proglottids, number of testes, testes distribution, size of eggs, ornamented egg shell, shape of bothridial cleft at level of the marginal accessory sucker, and the extension of vitelline follicles. Species in Orygmatobothrium share a common microtrich pattern with the distal bothridial surface covered with maisiform microtriches interspersed with filiform microtriches, a proximal bothridial surface covered with trifid microtriches, with a medial projection conspicuously larger than the lateral basal projections interspersed with filiform microtriches, an inner and outer surface of the accessory sucker and glandulomuscular organ covered with short filiform microtriches, the scolex proper and cephalic peduncle surface covered with bladelike microtriches, and the germinative zone and entire strobila covered with scutes formed by densely packed filiform microtriches. This general configuration is basically similar to the microtrich pattern described in species of Orectolobicestus and Paraorygmatobothrium.     

Rhinebothrium xiamenensis n. sp. (Eucestoda: Tetraphyllidea) in Dasyatis zugei from the coast of Xiamen, China

A new species of Rhinebothrium inhabiting Dasyatis zugei is reported from the southeast coast of China. This is the first report of Rhinebothrium from D. zugei. It represents a new host and a new location. The new species resembles Rhinebothrium corymbum by having a V-shaped ovary. It differs in the fewer loculi and testes numbers, longer bothrium pedicel and cephalic peduncle, larger cirrus pouch and vitelline follicles, and an aspinose peduncle. Rhinebothrium xiamenensis differs from Rhinebothrium ezuti, Rhinebothrium walga, and Rhinebothrium hawaiiensis by having discrete vitelline follicles, fewer loculi, and ovarian shape.     

Orectolobicestus n. g. (Cestoda: Tetraphyllidea), with the description of five new species and the transfer of Phyllobothrium chiloscyllii to the new genus

ORECTOLOBICESTUS N. G. (CESTODA: Tetraphyllidea) is erected for six cestode species parasitising bamboo sharks (Orectolobiformes: Chiloscyllium). Members of this genus differ from all other phyllobothriid genera in possessing modified maisiform spinitriches on their distal bothridial surfaces. In addition, they are easily recognised in that they share the following unique combination of characters: their vitelline fields are interrupted by the ovary, their necks are scutellate, and their bothridia bear an apical sucker and marginal loculi. Five new species of Orectolobicestus are described, including O. tyleri n. sp. from Chiloscyllium punctatum off Borneo, O. lorettae n. sp. from C. cf. punctatum off Australia, O. mukahensis n. sp. and O. kelleyae n. sp. from C. indicum off Borneo, and O. randyi n.sp. from C. hasselti also from off Borneo. In addition, Phyllobothrium chiloscyllii Subhapradha, 1955 is transferred to the new genus. O. chiloscyllii (Subhapradha, 1955) n. comb. is readily distinguished from all five new species in its greater total length. In addition to a number of proglottid features, O. kelleyae n. sp. and O. randyi n. sp. clearly differ from the other three new species in their possession of trifid, rather than fully serrate, spinitriches on their proximal bothridial surfaces. The latter two species conspicuously differ from one another in total number of proglottids (11-21 vs 27-38). O. tyleri n. sp. generally has fewer proglottids than O. lorrettae n. sp. (7-17 vs 13-23) and, like O. mukahensis n. sp., possesses scutes that are spathate rather than elongate. O. tyleri n. sp. is readily distinguished from O. mukahensis n. sp. in its possession of fewer proglottids (7-17 vs. 19-29). The five new species of Orectolobicestus share derived bothridial microthrix features with Phyllobothrium squali Yamaguti, 1952, Thysanocephalum sp., Orygmatobothrium sp., Ruhnkecestus Caira & Durkin, 2006 and species of Paraorygmatobothrium Ruhnke, 1994. Among these taxa, Orectolobicestus most closely resembles Paraorygmatobothrium and Ruhnkecestus in its vitelline fields being interrupted by the ovary and the possession of a scutellate neck.     

Two new species of Duplicibothrium Williams & Campbell, 1978 (Tetraphyllidea: Serendipidae) from the Pacific cownose ray Rhinoptera steindachneri

Two new tetraphyllidean species, Duplicibothrium cairae 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 Duplicibothriumappear 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.     

Pararhinebothroides hobergi n. gen. n. sp. (Eucestoda: Tetraphyllidea) in Urobatis tumbesensis (Chondrichthyes: Myliobatiformes) from coastal Ecuador.

A new species of tetraphyllidean eucestode inhabiting Urobatis tumbesensis from inshore waters of southeastern Ecuador shares 3 synapomorphies with Rhinebothroides spp.: apical bothridial suckers poorly differentiated from the marginal loculi, internal seminal vesicles, and insertion of the vas deferens dorsally closer to the poral than the aporal end of the cirrus sac. The new species differs from Rhinebothroides spp. by lacking medial bothridial septa and loculi and having symmetrical ovarian arms, and possesses an apparent autapomorphic trait by having the vas deferens tapering to a narrow tube before entering the cirrus sac, extending posteriorly to the posterior end of the cirrus sac where it expands into an external seminal vesicle running ventral to the cirrus sac anteriorly to anterior to the vagina. In Rhinebothroides spp., the vas deferens is expanded into an external seminal vesicle near the insertion into the cirrus sac As the sister group of Rhinebothroides, we propose a new genus to accommodate the new species. Phylogenetic evaluation of phyllobothriids recently assigned to Anthocephalum shows that they represent a paraphyletic assemblage of species of varying degrees of relatedness to Rhinebothroides spp. and the new species. Uncovering the relationships of the new species and the various species assigned to Anthocephalum permitted reevaluation of character transformations used in previous phylogenetic analysis of Rhinebothroides. Transformation series for 3 characters, previously based on functional outgroup comparisons, changed and a new character, length of cirrus sac, was added. The new phylogenetic analysis differs from the previous hypothesis only in placing R. scorzai as the sister species of R. circularisi + R. venezuelae + R. moralarai rather than of R. freitasi + R. glandularis + R. mclennanae. The occurrence of the sister species of Rhinebothroides in a Pacific Ocean stingray adds additional support to the hypothesis of Pacific origins of South American freshwater stingrays.     

Microtriches of tetraphyllidean metacestodes from Western Mediterranean striped dolphins (Stenella coeruleoalba)

The tegumental structures of two types of tetraphyllidean plerocercoids and two types of merocercoids (Phyllobothrium delphini and Monorygma grimaldii) from Mediterranean striped dolphins, Stenella coeruleoalba, are described for the first time using scanning electron microscopy. The tegument of all of the specimens was fully covered with microtriches. Four basic types were found: filitriches, blade-like spinitriches of different sizes and shapes, cone-shaped spinitriches (with two parallel small projections of equal length at the apex), and crowned cylindrical spinitriches (with 6-7 small papillae forming a crown at the apex); the two latter types are newly described. The two types of plerocercoids had a similar morphology and distribution of microtriches; in addition, cilium-like projections appeared interspersed among the microtriches on the apical sucker, accessory suckers, and distal bothridial loculus. Merocercoids exhibited a greater variety of tegumental structures, especially on the distal bothridial loculus. Both merocercoid types had regularly spaced papillae or "buttons" on the accessory suckers and the distal bothridial loculus that were composed of a central cilium-like projection surrounded by numerous filitriches. However, crowned cylindrical spinitriches were specific to P. delphini and cone-shaped spinitriches were specific to M. grimaldii. Differences in the morphology and distribution of scolex microtriches of adult cestodes have been considered useful for species identification. A previous molecular study has suggested that P. delphini and M. grimaldii are actually different congeneric species. Our study has shown that significant differences in the morphology and distribution of microtriches occur between these species at the merocercoid stage.     

A new tetraphyllidean genus and species, Caulopaterapagei n. g., n. sp. (Tetraphyllidea: Phyllobothriidae), from the grey carpetshark Chiloscyllium punctatum Müller & Henle (Orectolobiformes: Hemiscylliidae)

A new genus and species of tetraphyllidean cestode, Caulopatera pagei n. g., n. sp., is described from the grey carpetshark Chiloscyllium punctatum Müller & Henle in Moreton Bay, Australia. The new genus is placed in the Phyllobothriidae, subfamily Phyllobothriinae. Caulopatera n. g. is distinct from all other phyllobothriine genera in having stalked, circular, non-loculate bothridia that lack an apical sucker, testes that are restricted to the region anterior to the cirrus-sac and circum-medullary vitelline follicles. The new genus most closely resembles Carpobothrium Shipley & Hornell, 1906, with which it shares non-loculate, stalked, unhooked bothridia without an accessory sucker and testes that are entirely anterior to the cirrus-sac, but differs from it in that it lacks a slit-like opening in each bothridium and flaps surrounding the opening. The possession of bothridial stalks is consistent with two cestode orders, the Tetraphyllidea and the Rhinebothriidea. The morphology of the bothridial stalks is consistent with other tetraphyllidean genera, in that Caulopatera possesses triangular bothridial stalks surrounding the back of the bothridia, indicating that it belongs in the Tetraphyllidea senso stricto, rather than in the recently recognised Rhinebothriidea.     

Onchobothrium malakhovi n. sp. (Cestoda: Onchoproteocephalidea) ex Bathyraja (Arctoraja) sexoculata (Rajiformes: Arhynchobatidae) from Kuril Islands (Russia), with comments on the status of the genus Onchobothrium

Onchobothrium malakhovi n. sp. was found in the spiral valve of the softnose skate Bathyraja (Arctoraja) sexoculata off the Simushir Island (Kuril Islands, Russia). The new species has bothridia with three loculi and no additional suckers on bothridia, single-toothed hooks unconnected by their bases, no spines at the bases of the hooks, dense matrix around the hook bases shaped as an unpaired butterfly wing, and a short and wide ovary. Onchobothrium malakhovi n. sp. differs from O. antarcticum and O. magnum in having a smaller total length, cirrus sac and ovary, smaller testes and eggs. Additionally, the new species differs from O. antarcticum by the absence of a vaginal sphincter and shorter bothridia; differs from O. magnum in having fewer proglottids and smaller vitelline follicles. It differs from O. farmeri, O. convolutum, and O. pseudouncinatum, by the absence of a small spine at the base of the hooks and the absence of accessory suckers on bothridia; from O. pseudouncinatum, additionally, by unconnected hooks; from O. schizacanthium, by the number of testes and by the presence of a postvaginal group of testes. Onchobothrium malakhovi n. sp. was placed among other members of the Onchoproteocephalidea with a high support based on the sequence data for the D1-D3 region of the 28S rDNA and cox1 gene. The phylogenetic position of the genus Onchobothrium sensu lato remains ambiguous. We suggest that Onchobothrium sensu lato is a complex genus containing at least two morphologically different groups of species. Onchobothrium farmer, O. convolutum, O. schizacanthium, and O. pseudouncinatum, for which there are no molecular genetic data, are considerably different morphologically from O. malakhovi n. sp., O. antarcticum, and O. magnum. A new genus might have to be established for the latter three species after the accumulation of genetic data.     

Prochristianella spinulifera n. sp. (Cestoda: Trypanorhyncha) from Australian dasyatid and rhinobatid rays

Prochristianella spinulifera n. sp. (Cestoda: Trypanorhyncha: Eutetrarhynchidae) is described from the spiral valves of the rays Rhinobatos typus (Rhinobatidae) and Himantura fai (Dasyatidae) from Heron Island, Queensland, Australia. The new species is distinguished from all congeners by the deltoid microtriches covering the anterior 80% of the scolex and the presence of a dorsoventrally elongate genital atrium. The species occurred in the anteriormost section of the spiral valve of R. typus. The orientation of the armature of this and other congeners is such that principal rows of hooks begin on the bothridial surface of the tentacle and end on the antibothridial surface.     

Surface ultrastructure of the elasmobranchia parasitizing <Emphasis Type="Italic">Grillotiella exilis</Emphasis> and <Emphasis Type="Italic">Pseudonybelinia odontacantha</Emphasis> (Trypanorhyncha,Cestoda)

The surface ultrastructure of two monotypic trypanorhynch genera is described based on new material of Grillotiella exilis (Linton, 1909) and type material of Pseudonybelinia odontacantha Dollfus 1966. In G. exilis, spiniform microtriches cover the bothrial surfaces and the anterior part of the pars vaginalis posterior to the bothria. Bifurcate microtriches adorn the bothrial margins, filiform microtriches the scolex peduncle, and capilliform microtriches the posterior scolex end. This microthrix pattern resembles that found in, e.g., Grillotia erinaceus (van Beneden, 1858), with the difference that the anterior part of the pars vaginalis is covered with a collar of multidigitate palmate microtriches. The position of Grillotiella within the Grillotiinae, Lacistorhynchidae is supported based on these data. The bothria and scolex peduncle of P. odontacantha are covered with acerosate and unciniform microtriches on the distal bothrial surface and capilliform microtriches on the scolex peduncle. Short filiform microtriches cover the appendix. The microthrix pattern resembles that of the Tentaculariidae but with unciniform and acerosate microtriches densely covering the entire distal bothrial surface. Tegumental grooves are present on the posterior bothrial margin. They can be distinguished from bothrial pits in otobothrioid trypanorhynchs in having similar unciniform microtriches compared to the other parts of the bothrial surface and in lacking any spiniform microtriches. With the absence of bothrial pits as characteristic for the otobothrioids and its characteristic microthrix pattern, P. odontacantha together with Paranybelinia otobothrioides Dollfus 1966, both belonging to the Paranybeliniidae change their position in the most recent system from the Otobothrioidea into the Tentacularioidea.     

<Emphasis Type="Italic">Empruthotrema longipenis</Emphasis> n. sp. (Monogenoidea: Monocotylidae: Merizocotylinae) from the olfactory sacs of the smooth butterfly ray <Emphasis Type="Italic">Gymnura micrura</Emphasis> (Bloch &amp; Schneider) (Myliobatiformes: Gymnuridae) in the Gulf of Mexico

A new species of Empruthotrema Johnston & Tiegs, 1922 is described based on specimens collected from the olfactory sacs of smooth butterfly rays Gymnura micrura (Bloch & Schneider) captured in Mobile Bay (northcentral Gulf of Mexico), Alabama, USA. Empruthotrema longipenis n. sp. is most similar to the type-species Empruthotrema raiae (MacCallum, 1916) Johnston & Tiegs, 1922 by having 12 marginal and two interhamular loculi with members of haptoral hook pair 1 located midway along the periphery of each interhamular loculus and those of hook pair 2 located at the marginal termini of the bilateral septa flanking the interhamular loculi. Empruthotrema longipenis n. sp. differs from E. raiae by having a much longer male copulatory organ and from its remaining congeners by the sinistral and extracecal ejaculatory bulb flanking the pharynx, the number of interhamular and marginal septa, and the distribution of hook pairs 1 and 2 along the haptoral margin. This is the first report of a monocotylid from the smooth butterfly ray and from Mobile Bay. The diversity of haptoral morphotypes among the currently accepted species of Empruthotrema is detailed and discussed in the context of monophyly of the genus.     

Phyllobothrium discopygi n. sp. (Cestoda: Tetraphyllidea) from Chile,with a critical comparison of the affinities of P. auricula van Beneden, 1858 and P. foliatum Linton, 1890

Phyllobothrium discopygi n. sp. is described from the spiral valve of Discopyge tschudi Heckel, 1846 (Torpedinidae) taken in the Pacific Ocean off Coquimbo, San Antonio and Antofagasta, Chile, Specimens are described from whole-mounts, sections and SEM. Phyllobothrium discopygi is distinguished from other tetraphyllidean cestodes of the genus Phyllobothrium van Beneden, 1849 by a bifurcate scolex with bothridia joined in pairs, short neck, testes number and distribution, and morphology of the vitelline follicles and ovary. Phyllobothrium foliatum Linton, 1890 is considered a junior synonym of P. auricula van Beneden, 1858 following comparisons of P. auricula from D. pastinaca, Linton's holotype of P. foliatum from D. centroura and fresh specimens from D. centroura taken in waters near Woods Hole, Massachusetts. A meristic comparison of the adult worms is presented.     

Rhinebothrium biorchidum n. sp., a tetraphyllidean cestode from a yellow-spotted stingray, Urolophus jamaicensis, in Jamaica

Rhinebothrium biorchidum n. sp. (Tetraphyllidea: Phyllobothriidae) is described from the spiral valve of the yellow-spotted stingray, Urolophus jamaicensis, from Jamaica. It was most similar to R. spinicephalum Campbell 1970 but differed by having fewer proglottids (15 to 26 vs. 36 to 49), smaller peduncle (110 to 146 vs. 330 to 470) and pedicels (100 to 180 vs. 170 to 370), fewer transverse septa (6 to 8 vs. 16 to 17), fewer total loculi per bothridium (22 to 30 vs. 32 to 34) and larger ovarian lobes (148 to 310 vs. 88 to 176). A key to bitestate species of Rhinebothrium is included.