Ectoparasites of the Critically Endangered green sawfish Pristis zijsron and sympatric elasmobranchs in Western Australia.

This study reports the metazoan ectoparasite fauna of juvenile Critically Endangered green sawfish, Pristis zijsron, and sympatric elasmobranchs in Western Australia. Five parasite taxa were found on 76 screened P. zijsron: Caligus furcisetifer (Copepoda: Caligidae), Dermopristis pterophila (Monogenea: Microbothriidae), Branchellion plicobranchus and Stibarobdella macrothela (Hirudinea: Piscicolidae), and praniza larvae of an unidentified gnathiid isopod. Only C. furcisetifer and D. pterophila were common, exhibiting discrepant site-specificity, with C. furcisetifer occurring mostly on the head and rostrum, and D. pterophila around the pectoral and pelvic fins. Intensity of infection for C. furcisetifer and D. pterophila increased with host total length and was influenced by host sex, but in opposite directions; intensity of C. furcisetifer was greater on female P. zijsron, whereas intensity of D. pterophila was greater on males. In the Ashburton River, likelihood of infection for C. furcisetifer and D. pterophila on P. zijsron increased with time since substantial freshwater discharge events, suggesting decreased salinity impacts both taxa. In addition to P. zijsron, five other sympatric elasmobranch species were opportunistically screened for ectoparasites in the study area: the giant shovelnose ray, Glaucostegus typus, the eyebrow wedgefish, Rhynchobatus palpebratus, the nervous shark, Carcharhinus cautus, the lemon shark, Negaprion acutidens, and the graceful shark, Carcharhinus amblyrhynchoides. Caligus furcisetifer was found on R. palpebratus; no other parasites of P. zijsron were found on other sympatric elasmobranch species. Conversely, Perissopus dentatus (Copepoda: Pandaridae) was found on all three carcharhinids but not on batoid rays (P. zijsron, G. typus or R. palpebratus).

A new species of Siphoderina Manter, 1934 (Digenea: Cryptogonimidae) infecting the Dory Snapper Lutjanus fulviflamma (Teleostei: Lutjanidae) from the east coast of South Africa.

Parasitological assessment of marine fishes at Sodwana Bay in the iSimangaliso Marine Protected Area on the east coast of South Africa revealed a new species of cryptogonimid trematode infecting the pyloric caeca of the Dory Snapper, Lutjanus fulviflamma (Forsskål) (Lutjanidae). The new species is morphologically consistent with the concept of the large genus Siphoderina Manter, 1934; its phylogenetic position within this genus was validated through molecular sequencing of the ITS2 and partial 28S ribosomal DNA sub-regions. We name this species Siphoderina nana n. sp. and comment on the current state of understanding for this genus of cryptogonimids.

KANNAPHALLUS RAPHIDIUM N. SP. (MONOGENOIDEA: MAZOCRAEIDEA: HETERAXINIDAE) PARASITIC ON THE GILL LAMELLAE OF THE GOLDEN TREVALLY GNATHANODON SPECIOSUS (CARANGIFORMES: CARANGIDAE) OCCURRING IN THE COASTAL WATERS OF QUEENSLAND AND WESTERN AUSTRALIA.

An undescribed species of KannaphallusUnnithan, 1957 (Monogenoidea: Heteraxinidae) was collected from the gills of the golden trevally Gnathanodon speciosus (Forsskål) (Carangidae) from Moreton Bay, Queensland, during January 2016 and from Ningaloo Reef, Western Australia, during December 2021 and June 2022. The diagnosis for Kannaphallus was emended and the new species, Kannaphallus raphidium, was described. Kannaphallus virilis of Young, nec Unnithan was placed in synonymy with K. raphidium. The distal components of the male reproductive system and the arrangement of the clamp rows of the haptor occurred as mirror images among specimens of K. raphidium, suggesting that the respective antipodes of K. raphidium may have reproductive implications and function in the site selection of the parasite on the host's gills. A specimen of K. raphidium from Western Australia was sequenced for the cytochrome c oxidase subunit 1 (COI) mtDNA and ITS2 rDNA barcoding markers, and the phylogenetically informative 28S rDNA marker. Maximum likelihood and Bayesian inference analyses based on a partial 28S rDNA alignment, including all comparable heteraxinid sequence data available, resolved the Heteraxininae and Cemocotylinae as reciprocatively paraphyletic and provided evidence that Kannaphallus may be paraphyletic. No taxonomic changes concerning the subfamilies and genera of the Heteraxinidae were proposed. Finally, Kannaphallus univaginalisRamalingam, 1960 and Cemocotylelloides univaginalis (Ramalingam, 1960) Nitta, Kondo, Ohtsuka, Kamarudin, and Ismail, 2022 are considered nomen nuda sensu the International Code of Zoological Nomenclature.

Siphoderina hustoni n. sp. (Platyhelminthes: Trematoda: Cryptogonimidae) from the Maori snapper Lutjanus rivulatus (Cuvier) on the Great Barrier Reef.

A new cryptogonimid trematode, Siphoderina hustoni n. sp., is reported, collected off Lizard Island, Queensland, Australia, from the Maori snapper Lutjanus rivulatus (Cuvier). The new species is moderately distinctive within the genus. It is larger and more elongate than most other species of Siphoderina Manter, 1934, has the shortest forebody of any, a relatively large ventral sucker, a long post-testicular zone, and is perhaps most recognisable for the substantial space in the midbody between the ventral sucker and ovary devoid of uterine coils and vitelline follicles, the former being restricted to largely posterior to the ovary and the latter distributed from the level of the anterior testis to the level of the ovary. In phylogenetic analyses of 28S ribosomal DNA, the new species resolved with the other nine species of Siphoderina for which sequence data are available, all of which are from Queensland waters and from lutjanid and haemulid fishes. Molecular barcode data were also generated, for the ITS2 ribosomal DNA and cox1 mitochondrial DNA markers. The new species is the first cryptogonimid known from L. rivulatus and the first metazoan parasite reported from that fish in Australian waters.

A new microbothriid monogenean Dermopristis pterophilus n. sp. from the skin of the Critically Endangered green sawfish Pristis zijsron Bleeker, 1851 (Batoidea: Pristidae) in Western Australia.

A new microbothriid monogenean Dermopristis pterophilus n. sp. is described from the skin of the Critically Endangered green sawfish Pristis zijsron Bleeker, 1851 in the Ashburton River delta, northern Western Australia. Analyses of the 28S ribosomal DNA marker and the molecular barcoding markers Histone 3 and Elongation Factor 1 α confirmed position among the Microbothriidae, with close affinity to the only other sequenced representative of Dermopristis Kearn, Whittington and Evans-Groing, 2010. The new species is morphologically consistent with the concept of Dermopristis; it has two testes, lacks a male copulatory organ and has a simple haptor. It is smaller than its two congeners D. paradoxus Kearn, Whittington and Evans-Gowing, 2010 and D. cairae Whittington and Kearn, 2011 and is most similar to the former, distinguished only in that it lacks the strong, transverse, parallel ridges on the ventral body surface that characterise that species. It is more easily distinguished from D. cairae, differing in body shape, possession of a seminal receptacle, and relative position and size of the haptor. It may further differ from both species by fine details of the gut diverticula, although these details are difficult to ascertain. Spermatophores were observed in the new species, similar to those previously reported for D. cairae. The new species exhibits site attachment preference: infections were greatest on and immediately adjacent to the host pelvic fins (including male reproductive organs, i.e. claspers), moderate in proximity to the dorsal and pectoral fins, few on the caudal fin and peduncle, and infrequently, isolated worms occurred elsewhere on the dorsal and ventral surfaces of the body. There was no incidence of infection on the head (including rostrum). We presume D. pterophilus is restricted to P. zijsron and thus likely faces the same threat of extinction.

Eight species of Lintonium Stunkard & Nigrelli, 1930 (Digenea: Fellodistomidae) in Australian tetraodontiform fishes.

We report eight species of Lintonium from tetraodontiform fishes from Australian waters and describe six of them as new. Two species are described from tetraodontids from the Great Barrier Reef (GBR): Lintonium kostadinovae n. sp. from Arothron nigropunctatus (Bloch & Schneider) and Arothron hispidus (Linnaeus); and Lintonium droneni n. sp. from A. nigropunctatus. Two species are described from temperate monacanthids: Lintonium crowcrofti n. sp. from Meuschenia hippocrepis (Quoy & Gaimard) and Meuschenia freycineti (Quoy & Gaimard) off Tasmania and from M. hippocrepis off Glenelg, South Australia and off Fremantle, Western Australia; and Lintonium blendi n. sp. from M. hippocrepis off Stanley, Tasmania. The final two new species are described from tropical monacanthids: Lintonium currani n. sp. from Cantherhines pardalis (Rüppell) from Ningaloo Reef, Western Australia; and Lintonium madhaviae n. sp. from Amanses scopas (Cuvier) from the southern GBR. Two previously described species are reported from tetraodontids: Lintonium pulchrum (Johnston, 1913) Yamaguti, 1954 from Arothron stellatus (Anonymous), A. hispidus, A. manilensis (Marion de Procé) and Lagocephalus lunaris (Bloch & Schneider) from the GBR and southern Queensland; and Lintonium consors (Lühe, 1906) Crowcroft, 1950 from A. nigropunctatus from the southern GBR. Sequence data for three markers (ITS2 and 28S rDNA and cox1 mtDNA) for six of the eight species (L. crowcrofti n. sp., L. currani n. sp., L. droneni n. sp., L. kostadinovae n. sp., L. madhaviae n. sp. and L. pulchrum) are the first for the genus and distinguish each species unambiguously. Many records of species of Lintonium, especially widespread records of the type species, L. vibex (Linton, 1900) Stunkard & Nigrelli, 1930, remain to be clarified. A key finding of the present study is that three fish species (A. hispidus, A. nigropunctatus and M. hippocrepis) are identified as harbouring either two or three species of Lintonium at individual localities.

Retroporomonorchis pansho n. g., n. sp., an unusual monorchiid trematode exploiting an atypical host.

The Monorchiidae Odhner, 1911 are well represented in tropical and subtropical marine fishes worldwide but rarely reported from the Lutjanidae, an important family of tropical fishes that prey mainly on demersal fishes, decapods and cephalopods. Here, we report the first monorchiid from a lutjanid in Australian waters, Retroporomonorchis pansho n. g., n. sp. in Lutjanus fulvus (Forster), off Lizard Island on the northern Great Barrier Reef. It is morphologically and phylogenetically distinct among known monorchiids, resolves within the Monorchiinae Odhner, 1911, and is characterised by a relatively large ventral sucker, a sinistral genital pore immediately posterior to the ventral sucker, unfilamented eggs, a single testis, and vitelline follicles restricted to the level of the gonads. We assessed all previous records of monorchiids from lutjanid fishes and found only one to be convincing, that of Infundiburictus overstreeti (Gupta & Gupta, 1990) Wee, Cutmore, Pérez-del-Olmo & Cribb, 2020 from Lutjanus monostigma (Cuvier) in the Bay of Bengal. Another, Monorcheides xishaensis Shen, 1985, from Lutjanus argentimaculatus (Forsskål) in the Paracel Islands, South China Sea, is here recognised as a cryptogonimid, and is synonymised with Siphoderina asiatica Gu & Shen, 1979. In the remaining reports, the identity of the host is either ambiguous or doubtful, or the identity of the worm is not supported with sufficient evidence.

Opistholobetines (Digenea: Opecoelidae) in Australian tetraodontiform fishes.

Opistholebetine opecoelids are reported following examination of 1,041 individual tetraodontiform fishes, comprising 60 species and seven families, collected in Australian waters between 1986 and 2018. Nine species consistent with Opistholebes Nicoll, 1915, Heterolebes Ozaki, 1935 or Maculifer Nicoll, 1915 were recovered. However, phylogenetic analysis of sequence data, generated for some of these species, suggested that five genera, not three, are required to adequately accommodate these taxa. Thus, the concept and composition for each is revised, Pseudoheterolebes Yamaguti, 1959 nec Gupta, 1968 is resurrected and Parallelolebes n. g. is proposed. Of the nine species examined, five are new. Four new species are from fishes endemic to subtropical and temperate Australian waters for which no trematodes have previously been reported: Pa. australis n. sp. and Ps. corazonae n. sp. from the slender-spined porcupinefish Diodon nicthemerus Cuvier (Diodontidae) off Stanley, Tasmania; Pa. virilis n. sp. from the horse-shoe leatherjacket Meuschenia hippocrepis (Quoy & Gaimard) (Monacanthidae) off Stanley; and Ps. stellaglobulus n. sp. from the threebar pocupinefish Dicotylichthys punctulatus Kaup (Diodontidae) in Moreton Bay, south-east Queensland. The fifth new species is M. diodontis n. sp., collected from the spotted porcupinefish Diodon hystrix Linnaeus and the blackblotched porcupinefish D. liturosus Shaw, in tropical waters on the Great Barrier Reef. Species reported previously include the type-species of Opistholebes, O. amplicoelus Nicoll, 1915 from the rough golden puffer Lagocephalus lunaris (Bloch & Schneider) (Tetraodontidae) and the common toadfish Tetractenos hamiltoni (Richardson) (Tetraodontidae) in Moreton Bay, and three species reported for the first time from fishes in Australian waters: H. maculosus Ozaki, 1935, Pa. elongatus Ozaki, 1937 n. comb. and Ps. diodontis (Cable, 1956) n. comb., each from both D. hystrix and D. liturosus on the Great Barrier Reef. Following the revisions, Opistholebes is recognised for two species, Heterolebes for five, Maculifer for eight, Pseudoheterolebes for five and Parallelolebes for three.

Intermediate host switches drive diversification among the largest trematode family: evidence from the Polypipapiliotrematinae n. subf. (Opecoelidae), parasites transmitted to butterflyfishes via predation of coral polyps.

Podocotyloides stenometra Pritchard, 1966 (Digenea: Opecoelidae) is the only trematode known to infect anthozoan corals. It causes disease in coral polyps of the genus Porites Link (Scleractinia: Poritidae) and its life-cycle depends on ingestion of these polyps by butterflyfishes (Perciformes: Chaetodontidae). This species has been reported throughout the Indo-Pacific, from the Seychelles to the Galápagos, but no study has investigated whether multiple species are involved. Here, we recollect P. stenometra from its type-host and type-locality, in Hawaiian waters, and describe four new species from examination of 768 butterflyfishes from French Polynesia. On the basis of morphology, phylogeny and life-history, we propose Polypipapiliotrema Martin, Cutmore & Cribb n. gen. and the Polypipapiliotrematinae Martin, Cutmore & Cribb n. subf., for P. stenometra (Pritchard) n. comb., P. citerovarium Martin, Cutmore & Cribb n. sp., P. hadrometra Martin, Cutmore & Cribb n. sp., P. heniochi Martin, Cutmore & Cribb n. sp., and P. ovatheculum Martin, Cutmore & Cribb n. sp. Given the diversity uncovered here and the ubiquity, abundance and diversity of butterflyfishes on coral reefs, we predict that Polypipapiliotrema will prove to comprise a rich complex of species causing disease in corals across the Indo-Pacific. The unique life-cycle of these taxa is consistent with phylogenetic distinction of the group and provides evidence for a broader basis of diversification among the family. We argue that life-cycle specialisation, in terms of adoption of disparate second intermediate host groups, has been a key driver of the diversification and richness of the Opecoelidae, the largest of all trematode families and the group most frequently encountered in coral reef fishes.

The phylogenetic position of Choerodonicola Cribb, 2005 (Digenea: Opecoelidae) with a partial life-cycle for a new species from the blue-barred parrotfish Scarus ghobban Forsskål (Scaridae) in Moreton Bay, Australia.

Choerodonicola Cribb, 2005 is a minor genus of opecoelid trematodes defined for species with exceptionally small eggs but otherwise generalised morphology. Four species are currently recognised, all from fishes collected in Japanese waters but each from different perciform families: a labrid, a scarid, a sparid and pinguipeds. We report on a new species, Choerodonicola arothokoros n. sp., from the blue-barred parrotfish Scarus ghobban Forsskål (Scaridae) collected in subtropical waters of Moreton Bay, south-east Queensland, Australia. Using genetic sequence data for the ITS2 rDNA marker, we matched adult C. arothokoros to intramollsucan stages discovered in an intertidal gastropod Herpetopoma atratum (Gmelin) (Vetigastropoda: Chilodontidae) collected in close proximity to the fish hosts. Notably, the cercariae lack a penetration stylet and are among the smallest known in the Opecoelidae. We provide the first assessment of the phylogenetic position of Choerodonicola based on sequence data generated for the phylogenetically informative 18S and 28S rRNA coding regions, for C. arothokoros and also C. renko Machida, 2014, which we recollected from the yellowback seabream Dentex hypselosomus Bleeker from the fish market in Minabe, Wakayama Prefecture, Japan. In our analyses, species of Choerodonicola resolved to neither of the major marine Plagioporinae (sensu lato) clades, clustering instead with Trilobovarium parvvatis Martin, Cutmore & Cribb, 2017, Podocotyloides parupenei (Manter, 1963) Pritchard, 1966 and Macvicaria magellanica Laskowski, Jezewski & Zdzitowiecki, 2013. This clade is phylogenetically distinctive such that it has the potential to be recognised as a new opecoelid subfamily, but further investigation is required to establish the bounds for such a grouping and to determine the morphological and/or life-history patterns reflected by the phylogeny. Finally, we propose C. interruptus (Manter 1954) n. comb. for a species previously recognised in Plagioporus Stafford, 1904 and known only from Pseudolabrus miles (Schneider & Forster), a labrid endemic to New Zealand.

Revision of Podocotyloides Yamaguti, 1934 (Digenea: Opecoelidae), resurrection of Pedunculacetabulum Yamaguti, 1934 and the naming of a cryptic opecoelid species.

Despite morphological and ecological inconsistencies among species, all plagioporine opecoelids with a pedunculate ventral sucker are currently considered to belong in the genus Podocotyloides Yamaguti, 1934. We revise the genus based on combined morphological and phylogenetic analyses of novel material collected from haemulid fishes in Queensland waters that we interpret to represent species congeneric with the type-species, Pod. petalophallus Yamaguti, 1934, also known from a haemulid, off Japan. Our phylogenetic analysis demonstrates polyphyly of Podocotyloides; prompts us to resurrect Pedunculacetabulum Yamaguti, 1934; and suggests that Pod. brevis Andres & Overstreet, 2013, from a deep-sea congrid in the Caribbean, and Pod. parupenei (Manter, 1963) Pritchard, 1966 and Pod. stenometra Pritchard, 1966, from mullids and chaetodontids, respectively, on the Great Barrier Reef, may each represent a distinct genus awaiting recognition. Our revised concept of Podocotyloides requires a pedunculate ventral sucker, but also a uterine sphincter prior to the genital atrium, a petalloid cirrus appendage, restriction of the vitelline follicles to the hindbody, and for the excretory vesicle to reach to the level of the ventral sucker. Of about 20 nominal species, we recognise just three in Podocotyloides (sensu stricto): Pod. petalophallus, Pod. gracilis (Yamaguti, 1952) Pritchard, 1966 and Pod. magnatestes Aleshkina & Gaevskaya, 1985. We provide new records for Pod. gracilis, and propose two new species of Podocotyloides, Pod. australis n. sp. and Pod. brevivesiculatus n. sp., and one new Pedunculacetabulum species, Ped. inopinipugnus n. sp., all from haemulids. Podocotyloides australis is morphologically indistinguishable from Pod. gracilis, and exploits the same definitive host, but is genetically and biogeographically distinct. It is thus a cryptic species, the first such opecoelid to be formally named.

An updated concept and revised composition for Hamacreadium Linton, 1910 (Opecoelidae: Plagioporinae) clarifies a previously obscured pattern of host-specificity among species.

The present concept of the trematode genus Hamacreadium Linton, 1910 encompasses considerable morphological variability and includes species reported from a broad range of fishes. These include herbivores and planktivores, despite the life-cycle of the type-species, Hamacreadium mutabile Linton, 1910, being known to use fishes as intermediate hosts. Reports of H. mutabile are numerous, spanning the west Atlantic, east Pacific and Indo-west Pacific, whereas other nominal species are infrequently reported and several inadequately described. Following a comprehensive review, a strict revised morphological definition is proposed for the genus. Several nominal species are excluded, but, conversely, finer distinctions are recognised among the species concluded to genuinely belong in the genus. Justified records for species retained in the genus are overwhelmingly from fishes of the families Lutjanidae Gill (snappers) and Lethrinidae Bonaparte (emperors), revealing a previously concealed pattern of host-specificity. For H. mutabile, it is argued that only records from western Atlantic lutjanid fishes should be considered genuine; those from plausible Indo-Pacific fishes most likely represent different species. In addition to H. mutabile, eight species are recognised: Hamacreadium cribbi Bray & Justine, 2016, Hamacreadium hainanense Shen, 1990, Hamacreadium interruptum Nagaty, 1941, Hamacreadium lethrini Yamaguti, 1934, Hamacreadium longivesiculum (Yamaguti, 1952) n. comb., Hamacreadium lutiani (Shen, 1990) n. comb., Hamacreadium morgani Baz,1946 and Hamacreadium phyllorchis (Bilqees, 1976) Cribb, 2005. A key to species of Hamacreadium and comprehensive lists of all host-locality records are included.

Revision of Neolebouria Gibson, 1976 (Digenea: Opecoelidae), with Trilobovarium n. g., for species infecting tropical and subtropical shallow-water fishes.

A new opecoelid trematode is reported from fishes of the Lethrinidae, Lutjanidae and Nemipteridae off Lizard Island on the northern Great Barrier Reef, Australia. The new species keys to Neolebouria Gibson, 1976 and shows strong similarity to several species of that genus, but is not consistent with the type-species, N. georgiensis Gibson, 1976, or others known from temperate/polar and/or deep-sea fishes. The new species is also phylogenetically distant from N. lanceolata (Price, 1934) Reimer, 1987, the only representative of the genus for which molecular data are available. A new genus, Trilobovarium n. g., is proposed for the new species, T. parvvatis n. sp. Eight morphologically similar species, previously recognised as belonging to Neolebouria, from shallow-water, mostly tropical/subtropical fishes, are transferred to Trilobovarium: T. diacopae (Nagaty & Abdel Aal, 1962) n. comb.; T. ira (Yamaguti, 1940) n. comb.; T. khalili (Ramadan, 1983) n. comb.; T. krusadaiense (Gupta, 1956) n. comb.; T. lineatum (Aken'Ova & Cribb, 2001) n. comb.; T. moretonense (Aken'Ova & Cribb, 2001) n. comb.; T. palauense (Machida, 2014) n. comb.; and T. truncatum (Linton, 1940) n. comb. Paramanteriella Li, Qiu & Zhang, 1988 is resurrected for five species of Neolebouria with a post-bifurcal genital pore: P. cantherini Li, Qiu & Zhang, 1988; P. capoori (Jaiswal, Upadhyay, Malhotra, Dronen & Malhotra, 2014) n. comb.; P. confusa (Overstreet, 1969) n. comb.; P. leiperi (Gupta, 1956) n. comb.; and P. pallenisca (Shipley & Hornell, 1905) n. comb. Neolebouria georgenascimentoi Bray, 2002, a species with an exceptionally long cirrus-sac, is transferred to Bentholebouria Andres, Pulis & Overstreet, 2004 as B. georgenascimentoi (Bray, 2002) n. comb., and N. maorum (Allison, 1966) Gibson 1976, an unusual species known from cephalopods, is designated a species incertae sedis. Eleven species are retained in a revised concept of Neolebouria.

Trematodes of fishes of the Indo-west Pacific: told and untold richness.

The Indo-west Pacific is a marine bioregion stretching from the east coast of Africa to Hawaii, French Polynesia and Easter Island. An assessment of the literature from the region found reports of 2,582 trematode species infecting 1,485 fish species. Reports are concentrated in larger fishes, undoubtedly reflecting the tendency for larger hosts to be infected by more species of parasites as well as a collecting bias. Many hundreds of fish species, including many from families known to be rich in trematodes, have yet to be reported as hosts. Despite some areas (the Great Barrier Reef, Hawaii and the waters off China, India and Japan) receiving sustained attention, none can be considered to be comprehensively known. Several regions, most importantly in East Africa, French Polynesia and the Coral Triangle, are especially poorly known. The fauna of the Indo-west Pacific has been reported so unevenly that we consider it impossible to predict the true trematode richness for the region. We conclude that the greatest gap in our understanding is of the geographical distribution of species in the Indo-west Pacific. This is highlighted by the fact that 87% of trematodes in the region have been reported no more than five times. The reliable recognition of species is a major problem in this field; molecular approaches offer prospects for resolution of species identification but have been little adopted to date.