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.     

Paraorygmatobothrium?taylori n. sp. (Tetraphyllidea: Phyllobothriidae) from the Australian weasel shark Hemigaleus?australiensis White, Last & Compagno (Carcharhiniformes: Hemigaleidae)

Paraorygmatobothrium taylori n. sp. (Tetraphyllidea: Phyllobothriidae) is described from the Australian weasel shark Hemigaleus australiensis White, Last & Compagno in Moreton Bay, off Queensland, Australia. The new species differs from 10 of the 11 described species of Paraorygmatobothrium Ruhnke, 1994 by the possession of prominent, semicircular bothridial muscle bands. From P. barberi Ruhnke, 1994, with which it shares the bothridial muscle bands, it differs in the possession of a cephalic peduncle and vitelline follicles that extend almost to the mid-line of the proglottis and are reduced, rather than completely interrupted, at the level of the ovary. P. janineae Ruhnke, Healy & Shapero, 2006 is recorded from its type–host but in a new locality, Moreton Bay, off Queensland, Australia. P. taylori is the third species of the genus recorded from the Hemigaleidae in Australian waters. Three of the eight known hemigaleid species are now recorded to harbour this genus, and three different species are now known from the two hemigaleids found in Australian waters.     

New collections of blood flukes (Aporocotylidae) from fishes of the tropical Indo-west Pacific,including a new genus,two new species and molecular evidence that Elaphrobates chaetodontis (Yamaguti, 1970) is widespread in the region

We report new collections of the Aporocotylidae from Australia, French Polynesia, and Japan. A new species of Cardicola Short, 1953 is described from Scomberomorus commerson (Lacépède) (Scombridae), off Lizard Island. Cardicola nolani n. sp. can be distinguished from its congeners based on the position of the oötype, the position of the male genital pore, and the absence of an oral sucker. A new species is described from Abalistes stellatus (Anonymous) (Balistidae), also from off Lizard Island. Phylogenetically the new species forms a strongly-supported clade with Cardicola yuelao Yong, Cutmore & Cribb, 2018, which also infects balistids. These two species are distinct from all other aporocotylids in the combination of exceptionally short anterior and long posterior caeca, a lanceolate body, a single testis, an entirely post-ovarian uterus and the position of the oötype; a new genus, Balistidicola, is proposed for them. Balistidicola corneri n. sp. and B. yuelao (Yong, Cutmore & Cribb, 2018) n. comb. are essentially morphologically cryptic, only distinguishable by the form of the spination (B. corneri has five spines per row and B. yuelao has six). Elaphrobates chaetodontis (Yamaguti, 1970) is reported from 21 species of butterflyfishes (Chaetodontidae) from nine locations in tropical Indo-west Pacific; cox1 sequence data demonstrate extensive geographical structuring in this species. Braya jexi Nolan & Cribb, 2006, Elaphrobates milleri (Nolan & Cribb, 2006), and P. corventum Overstreet & Køie, 1989 are each re-reported from their type-hosts, and Pearsonellum pygmaeus Nolan & Cribb, 2004 and Balistidicola yuelao are each reported from a new host.     

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

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

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.     

Comparison of microthrix ultrastructure and morphology on the plerocercoid and adult scolex of Calliobothrium cf. verticillatum (Tetraphyllidea: Onchobothriidae)

The distal bothridial surfaces of adult triloculate onchobothriids are covered with short structures that have been tentatively classified as very short filitriches, but this hypothesis has never been tested. Scanning electron microscopy (SEM) and transmission electron microscopy (TEM) were used to investigate microthrix morphology in the plerocercoid and adult forms of Calliobothrium cf. verticillatum, a triloculate onchobothriid tapeworm from Long Island Sound (Connecticut). Plerocercoids of C. cf. verticillatum were collected from the anterior midgut ceaca of Pagurus pollicaris Say, 1817 (flat-clawed hermit crab), and adults were collected from the spiral intestine of the dusky smooth hound Mustelis canis (Mitchell, 1815). Two plerocercoids and 2 adults were examined using SEM; 2 plerocercoids and 2 adults were examined using TEM. Microthrix distribution and morphology (including measurements of total length, base length, shaft length, and base width) were investigated on all surfaces of the plerocercoid and adult scolex. Slender filitriches and large bladelike spinitriches were observed extending from the tegument of plerocercoid and adult forms. The filitriches were found to have significantly narrower bases than the spinitriches (65-167 nm vs. 466-1,936 nm, respectively). The scolex proper of the plerocercoid and adult forms were found to have filitriches of medium-length and bladelike spinitriches. The distal bothridial surfaces differed dramatically in microthrix morphology between plerocercoid and adult forms; on the distal surfaces of the plerocercoids were long filitriches and bladelike spinitriches. However, the distal surfaces of the adults had short structures (previously hypothesized to be short filitriches) and a few bladelike spinitriches. Serial transverse sections revealed that the short structures on the distal bothridial surfaces of the adults were homologous with filitriches. They included all of the structural components of a filithrix as well as a base width that conformed to the filitriches found on other surfaces. The bothridial margins of the plerocercoid and adult forms had a microthrix pattern similar to that seen on the proximal bothridial surfaces except that the filitriches on the margins were significantly longer than those found anywhere else on the bothridia. The most dramatic difference between the plerocercoid and adult forms occurred on the distal bothridial surfaces, where the filitriches of the adult cestodes were significantly shorter and narrower, and the spinitriches were almost entirely lacking.     

A new species,new host records and life cycle data for lepocreadiids (Digenea) of pomacentrid fishes from the Great Barrier Reef,Australia

A new species of lepocreadiid, Opechonoides opisthoporus n. sp., is described infecting 12 pomacentrid fish species from the Great Barrier Reef, Australia, with Abudefduf whitleyi Allen & Robertson as the type-host. This taxon differs from the only other known member of the genus, Opechonoides gure Yamaguti, 1940, in the sucker width ratio, cirrus-sac length, position of the testes, position of the pore of Laurer’s canal, and relative post-testicular distance. The new species exhibits stenoxenic host-specificity, infecting pomacentrids from seven genera: Abudefduf Forsskål, Amphiprion Bloch & Schneider, Neoglyphidodon Allen, Neopomacentrus Allen, Plectroglyphidodon Fowler & Ball, Pomacentrus Lacépède and Stegastes Jenyns. Phylogenetic analyses of 28S rDNA sequence data demonstrate that O. opisthoporus n. sp. forms a strongly supported clade with Prodistomum orientale (Layman, 1930) Bray & Gibson, 1990. The life cycle of this new species is partly elucidated on the basis of ITS2 rDNA sequence data; intermediate hosts are shown to be three species of Ctenophora. New host records and molecular data are reported for Lepocreadium oyabitcha Machida, 1984 and Lepotrema amblyglyphidodonis Bray, Cutmore & Cribb, 2018, and new molecular data are provided for Lepotrema acanthochromidis Bray, Cutmore & Cribb, 2018 and Lepotrema adlardi (Bray, Cribb & Barker, 1993) Bray & Cribb, 1996. Novel cox1 mtDNA sequence data showed intraspecific geographical structuring between Heron Island and Lizard Island for L. acanthochromidis but not for L. adlardi or O. opisthoporus n. sp.

     

Two new tapeworms from the goblin shark Mitsukurina owstoni off Australia

Internal parasites are reported from the goblin shark Mitsukurina owstoni for the first time. The shark examined was a 190 kg male collected off of Ulladulla, New South Wales, Australia and deposited at the Australian Museum in Sydney. What appear to be 4 different species of tapeworms were recovered from the spiral intestine. These included two new species, Litobothrium amsichensis n. sp. (order Litobothridea) and Marsupiobothrium gobelinus n. sp. (order Tetraphyllidea), as well as one specimen of a trypanorhynch and one scolex of a second tetraphyllidean. The latter two species are unidentifiable without further material. The generic placement of the new tetraphyllidean is somewhat problematical; this genus was thought to be the most appropriate placement at present. The most conspicuous difference between the new litobothridean and the five other species in the order is that it possesses dorso-ventral projections on five rather than four or three anterior segments. The discovery of a litobothridean parasitising the goblin shark is consistent with the systematic placement of the Mitsukurinidae within the Lamniformes, and may preliminarily indicate close affinities between the Alopidae, Odontaspidae and Mitsukurinidae. The phyllobothriid differs from the five other species of Marsupiobothrium in its possession of extremely long bothridial peduncles, its lack of an arcuate cylindrical pad on the posterior bothridial margins and its possession of a marginal, distinct apical sucker rather than a submarginal, diffuse apical sucker on each bothridium. Four detached specimens of the new phyllobothriid retained tips of mucosal villi within their bothridia. From these specimens it appears that the peduncles allow this species to stretch its scolex for attachment to up to four villi simultaneously.     

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.     

First steps to restructuring the problematic genus Lasiotocus Looss, 1907 (Digenea: Monorchiidae) with the proposal of four new genera

Lasiotocus Looss, 1907 is the largest genus within the Monorchiidae Odhner, 1911, with 52 species currently considered valid. Species belonging to this genus exhibit wide morphological variation and it is likely that many of them belong in other genera; however, testing the validity of the group has been hampered by a lack of molecular sequence data, especially for the type-species. Here, we report the first DNA sequences for Lasiotocus mulli (Stossich, 1883) Odhner, 1911, the type-species of the genus, and only the sixth Lasiotocus species to be sequenced. Sequences were generated for three ribosomal DNA markers, the ITS2, 18S and 28S regions, and for one mitochondrial DNA marker, the cox1 region. Phylogenetic analyses show that the six sequenced species form four clearly distinct clades, each of which we argue require separate genera. On the basis of these relationships and review of the morphology of all species in the genus, we propose a system of six genera. Four genera, Lasiotocus, the resurrected concept of Ancylocoelium Nicoll, 1912, Infundiburictus n. gen., and Sinistroporomonorchis n. gen., represent the four molecular clades. Two genera, Paralasiotocus n. gen. and Alloinfundiburictus n. gen., are proposed on the basis of morphology only. Ancylocoelium is resurrected for Ancylocoelium typicum Nicoll, 1912, Infundiburictus is erected for Lasiotocus arrhichostoma Searle, Cutmore & Cribb, 2014, Sinistroporomonorchis is erected for Lasiotocus glebulentus Overstreet, 1971and Lasiotocus lizae Liu, 2002, Paralasiotocus is erected for Lasiotocus okinawaensis Machida, 2011, and Alloinfundiburictus is erected for Lasiotocus cacuminatus Nicoll, 1915. Of the 52 species presently recognised in Lasiotocus, 43 are distributed among these genera; nine are considered species inquirendae or are transferred to other genera. We think it likely that more genera will be required for the species reviewed here but more sequence data are essential to further refine the classification.     

Heterobucephalopsine and prosorhynchine trematodes (Digenea: Bucephalidae) from teleost fishes of Moreton Bay,Queensland, Australia,with the description of two new species

Eight species of the trematode family Bucephalidae Poche, 1907 are reported from teleost fishes in Moreton Bay, Queensland, Australia. Heterobucephalopsis yongi n. sp. is described from Gymnothorax eurostus (Muraenidae); the new form is distinguished from its congeners in the possession of a tiny cirrus-sac relative to body length, the length of the caecum, the position of the mouth and pharynx, and the position of the testes and ovary. Two known species of Dollfustrema Eckmann, 1934, D. durum Nolan, Curran, Miller, Cutmore, Cantacessi & Cribb, 2015 and D. gibsoni Nolan & Cribb, 2010, are reported from Gymnothorax pseudothyrsoideus (Bleeker) (Muraenidae); although both species were described from Australian waters, this represents the first reports from Moreton Bay and G. pseudothyrsoideus. Four species of Prosorhynchus Odhner, 1905 are reported, including one new, P. brayi n. sp., which is described from Epinephelus coioides (Hamilton) (Serranidae); P. brayi n. sp. is distinguished from its congeners in the possession of vitelline follicles in a confluent arc distinctly posterior to a conical rhynchus, uterine coils that do not extend anterior to the vitelline arc, contiguous testes, a cirrus-sac that reaches anteriorly to at least the level of the posterior testis and a short excretory vesicle. Three known species of Prosorhynchus are reported from Australia, for the first time: P. luzonicus Velasquez, 1959 and P. maternus Bray & Justine, 2006 from E. coioides and Prosorhynchus platycephali (Yamaguti, 1934) Srivastava, 1938 from Ambiserrula jugosa (McCulloch) and Inegocia japonica (Cuvier) (Platycephalidae). Skrjabiniella Issaitschikow, 1928 is re-recognised for new specimens of Skrjabiniella uniporus (Ozaki, 1924) n. comb. collected from Conger cinereus Rüppell (Congridae); three additional species of Prosorhynchus are considered members of this genus, two of which are synonymised with S. uniporus.     

A complex of the blood fluke genus Psettarium (Digenea: Aporocotylidae) infecting tetraodontiform fishes of east Queensland waters

Seven species of Psettarium (Digenea: Aporocotylidae), including four new species, are reported from tetraodontiform fishes from off coastal east Queensland. Psettarium pandora n. sp. infects the yellow boxfish, Ostracion cubicus (Ostraciidae), the first known aporocotylid to infect this family of fishes. Three new species are reported from pufferfishes of the genus Arothron (Tetraodontidae): Psettarium yoshidai n. sp. infects the map puffer (Arothron mappa), Psettarium hustoni n. sp. infects the black-spotted puffer (A. nigropunctatus) and Psettarium martini n. sp. infects the starry puffer (A. stellatus). We also report three species of Psettarium from Australian waters for the first time. Paracardicola hawaiensis Martin, 1960, the sole species of Paracardicola, is redescribed based on specimens collected from the type-host, the stars-and-stripes puffer, Arothron hispidus. Paracardicola is synonymised with Psettarium and P. hawaiensis is recombined as Psettarium hawaiiense (Martin, 1960) n. comb. Psettarium pulchellum Yong, Cutmore, Bray, Miller, Semarariana, Palm & Cribb, 2016, described from the narrow-lined puffer (Arothron manilensis) from off Bali, Indonesia, is reported from the same fish species at two locations on the Queensland coast, significantly extending the range of this species. Psettarium nolani (Bray, Cribb & Littlewood, 2013), originally described from French Polynesia, is reported from A. hispidus, A. manilensis and A. stellatus, representing both new host and locality records for this species. Molecular phylogenetic analysis shows these species to all be closely related, such that they cannot be considered to represent separate genera despite their differing morphology. Analysis of 28S sequence data for Psettarium anthicum Bullard & Overstreet, 2006, a non-tetraodontiform-infecting species, shows it to be distantly related to all other species of Psettarium for which sequence data are available. The species is re-assigned to a new genus, Cardallagium n. gen., as Cardallagium anthicum (Bullard & Overstreet, 2006) n. comb. We think it likely that the host range of species of Psettarium is limited to tetraodontiform fishes. We assessed the infection biology of two species, P. nolani and P. hawaiiense n. comb. infecting A. hispidus, using histology to assess the pathways of egg release for these species. Eggs of both species were observed in both circulatory and visceral organs of infected hosts, often in high numbers. Eggs were seen trapped in the mucosal layer of the intestine and, in rare instances, causing lesions in the laminar epithelium, providing the strongest evidence yet that they pass through the gut wall and escape the host via the faeces. Lastly, we discuss the biogeographical implications of our findings, noting that some Psettarium species now show very wide geographical distributions.     

Four new monorchiids from marine teleost fishes of Moreton Bay and the Great Barrier Reef,Australia, including the proposal of a new genus

We report four new species of monorchiids infecting teleost fishes from Australian waters. Two new species of Paralasiotocus Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020, Pa. abstrusus n. sp. and Pa. tectus n. sp., are described from haemulids of the Great Barrier Reef. The two species are morphologically cryptic and occur in sympatry but differ significantly in cox1 mtDNA and ITS2 rDNA sequence data. Paralasiotocus tectus n. sp. is found only in Plectorhinchus albovittatus (Rüppell) whereas Pa. abstrusus n. sp. infects Pl. albovittatus, Plectorhinchus flavomaculatus (Cuvier) and Plectorhinchus lineatus (Linnaeus). The two species differ from all known species of Paralasiotocus in the possession of a clear gap in the spines of the terminal organ. A new species is described from a mullid, Parupeneus spilurus (Bleeker), from off Heron Island and Moreton Bay. The new species is morphologically broadly consistent with the concept of Paralasiotocus in the possession of an unspined genital atrium, bipartite terminal organ, and lobed ovary. However, it possesses a highly lobed cirrus and is phylogenetically widely separated from the two species of Paralasiotocus characterized here, and thus we propose Lobucirruatus infloresco n. g., n. sp. Proctotrema prominens n. sp., is described from Pl. albovittatus. It is differentiated from all other species of Proctotrema in the combination of a prominent metraterm, slightly fusiform body, slightly funnel-shaped oral sucker, elongate cirrus-sac, unlobed ovary, and caeca that terminate in the post-testicular region.