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A new genus and three new species of blood flukes (Aporocotylidae) are described from squirrelfishes (Holocentridae) from the Great Barrier Reef. Holocentricola rufus n. gen., n. sp. is described from Sargocentron rubrum (Forsskål), from off Heron Island, southern Great Barrier Reef, and Lizard Island, northern Great Barrier Reef, Australia. Holocentricola exilis n. sp. and Holocentricola coronatus n. sp. are described from off Lizard Island, H. exilis from Neoniphon sammara (Forsskål) and H. coronatus from Sargocentron diadema (Lacepède). Species of the new genus are distinct from those of all other aporocotylid genera in having a retort-shaped cirrus-sac with a distinct thickening at a marginal male genital pore. The new genus is further distinct in the combination of a lanceolate body, X-shaped caeca, posterior caeca that are longer than anterior caeca, a single, post-caecal testis that is not deeply lobed, a post-caecal, post-testis ovary that is not distinctly bi-lobed, and a post-ovarian uterus. The three new species can be morphologically delineated based on the size and row structure of the marginal spines, as well by total length, oesophagus and caecal lengths, and the position of the male genital pore, testes and ovary relative to the posterior extremity. The three species of Holocentricola are genetically distinct from each other based on cox1 mtDNA and ITS2 rDNA data, and in phylogenetic analyses of 28S rDNA form a well-supported clade sister to species of Neoparacardicola Yamaguti, 1970. This is the first report of aporocotylids from fishes of the family Holocentridae and the order Holocentriformes.

New, well-known and predicted life cycles for trematodes of the Haploporoidea (Haploporidae and Emprostiotrematidae) and three families of the Lepocreadioidea (Enenteridae, Gorgocephalidae, Gyliauchenidae) involve encystment of the metacercaria in the open (usually on vegetation) followed by ingestion by a range of herbivorous or detritivorous fishes. These life cycles appear among relatively highly derived plagiorchiidan trematodes in which three-host life cycles incorporating an animal second intermediate host are dominant. We hypothesise that the two-host life cycles in the Haploporoidea and Lepocreadioidea arose by secondary truncation of a three-host cycle; the second intermediate host was lost in favour of encystment in the open. Modification of a three-host life cycle effective for the infection of carnivores is consistent with the understanding that fishes arose as carnivores and that multiple lineages have secondarily become detritivores and herbivores. Four of the five trematode families involved infect fishes relating to multiple orders, suggesting a complex history of host-switching. In contrast, the Gorgocephalidae, the smallest of the families, has been found only in a single family, Kyphosidae. The timing of the evolutionary events leading to this putative life cycle truncation is yet to be deduced, but the rich developing understanding of the history of the fishes creates a strong template for future analysis.

We describe a new species of Bivesiculidae, Bivesiculoides maiae n. sp., from Hypoatherina tropicalis (Whitley) (Atherinidae) collected from off Heron Island (southern Great Barrier Reef, Queensland, Australia). Bivesiculoides maiae n. sp. is morphologically consistent with Bivesiculoides Yamaguti, 1938 in the entirely pre-testicular position of its uterus, and the possession of caeca and vitelline fields that extend posteriorly to level with the anterior extremity of the testis. The new species is morphologically distinct from the six known Bivesiculoides species in body size and shape, and shape of the pharynx and testis. Bivesiculoides maiae n. sp. is genetically distinct from the only other sequenced Bivesiculoides species, Bivesiculoides fusiformis Cribb, Bray & Barker, 1994, with which it occurs sympatrically at Heron Island. A review of related species allows two systematic recombinations. In view of the pre-testicular position of its uterus, we recombine Bivesicula hepsetiae Manter, 1947 as Bivesiculoides hepsetiae (Manter, 1947) n. comb. In view of its obtriangular body shape, round pharynx, strongly elongated testis, and the position of its ovary opposite the testis, we recombine Bivesiculoides triangularis Machida & Kuramochi, 2000 as Treptodemoides triangularis (Machida & Kuramochi, 2000) n. comb. Host-specificity of species of Bivesiculoides and their geographic distributions are discussed., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Emprostiotrema contains just 3 species: E. fusum , E. kuntzi and E. sigani . As adults, all 3 species infect rabbitfishes (Siganidae: Siganus ). New collections from 11 species of Siganus from northern Australia, Indonesia, New Caledonia, French Polynesia, Palau and Japan enabled an exploration of species composition within this genus. Phylogenetic analyses demonstrate a deep distinction between 2 major clades; clade 1 comprises most of the sequences of specimens from Australia as well as all of those from Japan, Palau and New Caledonia and clade 2 comprises all sequences of specimens from French Polynesia, 2 sequences from Australia and the single sequence from Bali. In all analyses, both major clades have genetic structuring leading to distinct geographic lineages. Morphologically, specimens relating to clades 1 and 2 differ but overlap in body shape, oral sucker and egg size. Principle component analysis shows a general (but not complete) separation between specimens relating to the 2 clades. We interpret the 2 clades as representing 2 species: clade 1 is identified as E. fusum and is reported in this study from 10 species of siganids from Australia, Japan, Palau and New Caledonia; clade 2 is described as E. gotozakiorum n. sp., for all specimens from French Polynesia and rare specimens from Australia and Indonesia. We recognize E. sigani as a junior synonym of E. fusum . Although species of Emprostiotrema occur widely in the tropical Indo-Pacific, they have not been detected from Ningaloo Reef (Western Australia), the southern Great Barrier Reef or Moreton Bay (southern Queensland).

In a parasitological survey of fishes from Moreton Bay (southeastern Queensland, Australia), 169 teleost fishes, representing 54 species from 28 families, were examined for larval cestodes. Of these 54 species, 36 were found to be infected by metacestodes. Metacestodes were characterised by morphological and molecular data (the D1-D3 region of the 28S rDNA gene); these data were analysed in parallel to inform larval type allocation. Metacestodes collected represented eight morphological types, seven previously reported (Types I, II, IV, V, VI, VII, and X) and one novel type (Type XVI). Phylogenetic analyses were conducted to genetically match larval types to adult cestodes. Six of the eight larval types found were matched to adult forms: Type I metacestodes matched species of Phoreiobothrium Linton, 1889 (Onchobothriidae); Type II metacestodes matched species of Acanthobothrium van Beneden, 1849 (Onchobothriidae); Type IV metacestodes matched species of Scyphophyllidium Woodland, 1927 and Alexandercestus Ruhnke & Workman, 2013 (Phyllobothriidae); Type VI metacestodes matched species of Anthobothrium van Beneden, 1850 (Tetraphyllidea incertae sedis); Type X metacestodes matched species of Ambitalveolus Caira & Jensen, 2022 (Tetraphyllidea incertae sedis); and Type XVI metacestodes matched species of Platybothrium Linton, 1890 (Onchobothriidae). Based on phylogenetic topology, Type V metacestodes are inferred to match Pedibothrium Linton, 1909 (Balanobothriidae) and Type VII metacestodes are inferred to match Spongiobothrium Linton, 1889 (Rhinebothriidae). These findings support and extend the unified morphological type system proposed previously, but suggest that morphological types will ultimately be informative to identify metacestodes to a group of related genera rather than any distinct genus., (© 2024. The Author(s).)

We examined gelatinous zooplankton from off eastern Australia for lepocreadiid trematode metacercariae. From 221 specimens of 17 species of cnidarian medusae and 218 specimens of four species of ctenophores, infections were found in seven cnidarian and two ctenophore species. Metacercariae were distinguished using cox1 mtDNA, ITS2 rDNA and morphology. We identified three species of Prodistomum Linton, 1910 [P. keyam Bray & Cribb, 1996, P. orientale (Layman, 1930), and Prodistomum Type 3], two species of Opechona Looss, 1907 [O. kahawai Bray & Cribb, 2003 and O. cf. olssoni], and Cephalolepidapedon saba Yamaguti, 1970. Two species were found in cnidarians and ctenophores, three only in cnidarians, and one only in a ctenophore. Three Australian fishes were identified as definitive hosts; four species were collected from Scomber australasicus and one each from Arripis trutta and Monodactylus argenteus. Transmission of trematodes to these fishes by ingestion of gelatinous zooplankton is plausible given their mid-water feeding habits, although such predation is rarely reported. Combined morphological and molecular analyses of adult trematodes identified two cox1 types for C. saba, three cox1 types and species of Opechona, and six cox1 types and five species of Prodistomum of which only two are identified to species. All three genera are widely distributed geographically and have unresolved taxonomic issues. Levels of distinction between the recognised species varied dramatically for morphology, the three molecular markers, and host distribution. Phylogenetic analysis of 28S rDNA data extends previous findings that species of Opechona and Prodistomum do not form monophyletic clades., Competing Interests: Declaration of competing interest The authors declare that they have no conflict of interest., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

Pleustonic organisms form an important part of pelagic ecosystems by contributing to pelagic trophic chains and supporting connectivity between oceanic habitats. This study systematically analysed the trematode community harboured by pleustonic molluscs and cnidarians from offshore Queensland, Australia. Four mollusc and three cnidarian species were collected from beaches of North Stradbroke Island, Queensland. Two mollusc species and all three cnidarians harboured large numbers of hemiuroid metacercariae (Trematoda: Hemiuroidea). Eight taxa from four hemiuroid families (Accacoeliidae, Didymozoidae, Hemiuridae and Sclerodistomidae) were distinguished via molecular sequencing. Four of those taxa were identified to species. All trematode taxa except one didymozoid were shared by two or more host species; five species occurred in both gastropods and cnidarians. It is hypothesised that the life-cycles of these hemiuroids are highly plastic, involving multiple opportunistic pathways of metacercarial transmission to the definitive hosts. Transmission and the use of pleuston by hemiuroids likely varies with sea surface use and ontogenetic trophic shifts of apex predators. The small number of trematode species found in pleuston is consistent with significant ecological specificity, and the inference that other pelagic trematodes use alternative pathways of transmission that do not involve pleustonic organisms. Such pathways may involve i) pelagic hosts exclusively; ii) benthic or demersal hosts exclusively, consumed by apex predators during their dives; or iii) both benthic and pelagic hosts in transmission chains dependent on vertical migrations of prey. The influence of the connectivity of open-ocean ecosystems on parasite transmission is identified as an area in critical need of research., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Geographical distribution plays a major role in our understanding of marine biodiversity. Some marine fish trematodes have been shown to have highly restricted geographical distributions, while some are known to occur over very wide ranges; however, very few of these wide distributions have been demonstrated genetically. Here, we analyse species of the genus Schikhobalotrema (Haplosplanchnidae) parasitizing beloniforms from the tropical west Pacific, the eastern Pacific and the Gulf of Mexico (GoM). We test the boundaries of these trematodes by integrating molecular and morphological data, host association, habitat of the hosts and geographical distribution, following a recently proposed and standardized delineation method for the recognition of marine trematode species. Based on the new collections, Schikhobalotrema huffmani is here synonymized with the type-species of the genus, Schikhobalotrema acutum ; Sch. acutum is now considered to be widely distributed, from the GoM to the western Pacific. Additionally, we describe a new species, Schikhobalotrema minutum n. sp., from Strongylura notata and Strongylura marina (Belonidae) from La Carbonera coastal lagoon, northern Yucatán, GoM. We briefly discuss the role of host association and historical biogeography of the hosts as drivers of species diversification of Schikhobalotrema infecting beloniforms.

Species of Transversotrema Witenberg, 1944 (Transversotrematidae) occupy a unique ecological niche for the Trematoda, living externally under the scales of their teleost hosts. Previous studies of the genus have been impeded partly by limited variation in ribosomal DNA sequence data between closely related species and partly by a lack of morphometrically informative characters. Here, we assess richness of the tropical Indo-west Pacific species through parallel phylogenetic and morphometric analyses, generating cytochrome c oxidase subunit 1 mitochondrial sequence data and morphometric data for hologenophore specimens from Australia, French Polynesia, Japan and Palau. These analyses demonstrate that molecular data provide the only reliable basis for species identification; host distribution, and to a lesser extent morphology, are useful for identifying just a few species of Transversotrema. We infer that a combination of morphological simplicity and infection site constraint has led to the group displaying exceptionally low morphological diversification. Phylogenetic analyses of the mitochondrial data broadly support previous systematic interpretations based on ribosomal data, but also demonstrate the presence of several morphologically and ecologically cryptic species. Ten new species are described, eight from the Great Barrier Reef, Australia (Transversotrema chrysallis n. sp., Transversotrema daphnidis n. sp., Transversotrema enceladi n. sp., Transversotrema hyperionis n. sp., Transversotrema iapeti n. sp., Transversotrema rheae n. sp., Transversotrema tethyos n. sp., and Transversotrema titanis n. sp.) and two from off Japan (Transversotrema methones n. sp. and Transversotrema panos n. sp.). There are now 26 Transversotrema species known from Australian marine fishes, making it the richest trematode genus for the fauna., (Crown Copyright © 2023. Published by Elsevier Ltd. All rights reserved.)

We report nine species of the Schistorchiinae Yamaguti, 1942 (Apocreadiidae Skrjabin, 1942) from Indo-Pacific marine fishes. Molecular data (ITS2 and 28S rDNA and cox1 mtDNA) are provided for all species and the genus-level classification of the subfamily is revised. For Schistorchis Lühe, 1906, we report the type-species Sch. carneus Lühe, 1906 and Sch. skrjabini Parukhin, 1963. For Sphinteristomum Oshmarin, Mamaev & Parukhin, 1961 we report the type-species, Sph. acollum Oshmarin, Mamaev & Parukhin, 1961. We report and re-recognise Lobatotrema Manter, 1963, for the type and only species, L. aniferum Manter, 1963, previously a synonym of Sph. acollum. Lobatotrema aniferum is phylogenetically distant from, but morphologically similar to, Sph. acollum and Lobatotrema is recognised as a 'cryptic genus'. We propose Blendiella n. gen. for B. trigintatestis n. sp. and B. tridecimtestis n. sp. These species are broadly consistent with the present morphological concept of Schistorchis but are phylogenetically distant from the type-species; a larger number of testes and some other subtle morphological characters in species of Blendiella serve to distinguish the two genera. We report three species of Paraschistorchis Blend, Karar & Dronen, 2017: P. stenosoma (Hanson, 1953) Blend, Karar & Dronen, 2017 (type-species), P. seychellesiensis (Toman, 1989) Blend, Karar & Dronen, 2017, and P. zancli (Hanson, 1953) Blend, Karar & Dronen, 2017. Lobatotrema aniferum, P. stenosoma, and Sch. carneus each have two distinct cox1 populations either over geographical range or in sympatry. Available evidence suggests that most of these species, but not all, are widespread in the tropical Indo-Pacific., (© 2023. The Author(s).)

Integration of morphological and molecular approaches to species delineation has become an essential part of digenean trematode taxonomy, particularly when delimiting cryptic species. Here, we use an integrated approach to distinguish and describe two morphologically cryptic species of Hysterolecitha Linton, 1910 (Trematoda: Lecithasteridae) from fishes of Moreton Bay, Queensland, Australia. Morphological analyses of Hysterolecitha specimens from six fish species demonstrated a complete overlap in morphometric data with no reliable differences in their gross morphological characters that suggested the presence of more than one species. Distinctions in ITS2 rDNA and cox1 mtDNA sequence data for corresponding specimens suggested the presence of two forms. A principal component analysis on an imputed dataset showed clear separation between the two forms. These two forms are partially separated on the basis of their host's identity. Therefore, we describe two morphologically cryptic species: Hysterolecitha melae n. sp. from three species of Abudefduf Forsskål and one species of Parma Günther (Pomacentridae), with the Bengal sergeant, Abudefduf bengalensis (Bloch), as the type-host; and Hysterolecitha phisoni n. sp. from species of Pomacentridae (including A. bengalensis), Pomatomidae and Siganidae, with the black rabbitfish, Siganus fuscescens (Houttuyn), as the type-host., (© 2023. The Author(s).)

Neospirorchis Price, 1934 is a genus of blood flukes that infect the cardiovascular system, including vessels surrounding the nervous systems of marine turtles. Although the genus comprises just two named species, the available molecular data suggest substantial richness which has not yet been formally described. The lack of description of species of Neospirorchis is probably explained by their small, slender, elongate bodies, which allow them to infect numerous organs and vessels in their hosts, such as the heart and peripheral vessels of nervous system, endocrine organs, thymus, mesenteric vessels, and gastrointestinal submucosa. This morphology and site of infection means that collecting good quality, intact specimens is generally difficult, ultimately hampering the formal description of species. Here we supplement limited morphological samples with multi-locus genetic data to formally describe four new species of Neospirorchis infecting marine turtles from Queensland, Australia and Florida, USA; Neospirorchis goodmanorum n. sp. and Neospirorchis deburonae n. sp. are described from Chelonia mydas, Neospirorchis stacyi n. sp. is described from Caretta caretta, and Neospirorchis chapmanae n. sp. from Ch. mydas and Ca. caretta. The four new species are delineated from each other and the two known species based on the arrangement of the male and female reproductive organs, on the basis of cytochrome c oxidase subunit 1 (cox1), internal transcribed spacer 2 (ITS2), and 28S ribosomal DNA (rDNA) molecular data, site of infection, and host species. Molecular evidence for three further putative, presently undescribable, species is also reported. We propose that this integrated characterisation of species of Neospirorchis, based on careful consideration of host, molecular and key morphological data, offers a valuable solution to the slow rate of descriptions for this important genus. We provide the first known life cycle data for Neospirorchis in Australian waters, from Moreton Bay, Queensland; consistent with reports from the Atlantic, sporocysts were collected from a terebellid polychaete and genetically matched to an unnamed species of Neospirorchis infecting Ch. mydas from Queensland and Florida., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

A new genus, Doorochen n. gen., is erected for four species of Postlepidapedon Zdzitowiecki, 1993, all of which inhabit members of the labroid genus Choerodon Bleeker, the tuskfishes, and which molecular phylogenies have indicated are not congeneric with the type-species, P. opisthobifurcatum (Zdzitowiecki, 1990) Zdzitowiecki, 1993. Doorochen secundum (Durio & Manter, 1968) n. comb. from Choerodon graphicus (De Vis), the Graphic tuskfish, from the Great Barrier Reef (GBR) and New Caledonia is designated the type-species of the new genus. Other species recognised are Doorochen spissum (Bray, Cribb & Barker, 1997) n. comb. from C. venustus (De Vis), the Venus tuskfish, C. cyanodus (Richardson), the Blue tuskfish, and C. graphicus from the GBR; D. uberis (Bray, Cribb & Barker, 1997) n. comb. from C. schoenleinii (Valenciennes), the Blackspot tuskfish, and C. venustus from the GBR and Moreton Bay; and D. philippinense (Machida, 2004) n. comb. from C. anchorago (Bloch), the Orange-dotted tuskfish, from Philippine waters. In addition to these four species, two new species are described: D. zdzitowieckii n. sp. from C. fasciatus (Günther), the Harlequin tuskfish, and C. graphicus from the GBR; and D. goorchana n. sp. from C. anchorago from the GBR and Palau. The genus Postlepidapedon is now considered to comprise just two species, P. opisthobifurcatum and P. quintum Bray & Cribb, 2001. The relationships of Doorochen, Postlepidapedon, Myzoxenus Manter, 1934 and Intusatrium Durio & Manter, 1968 in the family Lepidapedidae Yamaguti, 1958 are discussed., Competing Interests: Declaration of Competing Interest The authors declare that they have no conflict of interest., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The family Aporocotylidae is recognized as having the widest intermediate host usage in the Digenea. Currently, intermediate host groups are clearly correlated with definitive host groups; all known life cycles of marine teleost-infecting aporocotylids involve polychaetes, those of freshwater teleost-infecting aporocotylids involve gastropods, and those of chondrichthyan-infecting aporocotylids involve bivalves. Here we report the life cycle for a marine elopomorph-infecting species, Elopicola bristowi Orélis-Ribeiro & Bullard in Orélis-Ribeiro, Halanych, Dang, Bakenhaster, Arias & Bullard, 2017, as infecting a bivalve, Anadara trapezia (Deshayes) (Arcidae), as the intermediate host in Moreton Bay, Queensland, Australia. The cercaria of E. bristowi has a prominent finfold, distinct anterior and posterior widenings of the oesophagus, a tail with symmetrical furcae with finfolds, and develops in elongate to oval sporocysts. We also report molecular data for an unmatched aporocotylid cercaria from another bivalve, Megapitaria squalida (G. B. Sowerby I) (Veneridae), from the Gulf of California, Mexico, and six unmatched cercariae from a gastropod, Posticobia brazieri (E. A. Smith) (Tateidae), from freshwater systems of south-east Queensland, Australia. Phylogenetic analyses demonstrate the presence of six strongly-supported lineages within the Aporocotylidae, including one of elopomorph-infecting genera, Elopicola Bullard, 2014 and Paracardicoloides Martin, 1974, now shown to use both gastropods and bivalves as intermediate hosts. Of a likely 14 aporocotylid species reported from bivalves, six are now genetically characterised. The cercarial morphology of these six species demonstrates a clear distinction between those that infect chondrichthyans and those that infect elopomorphs; chondrichthyan-infecting aporocotylids have cercariae with asymmetrical furcae that lack finfolds and develop in spherical sporocysts whereas those of elopomorph-infecting aporocotylids have symmetrical furcae with finfolds and develop in elongate sporocysts. This morphological correlation allows predictions of the host-based lineage to which the unsequenced species belong. The Aporocotylidae is proving exceptional in is propensity for major switches in intermediate host use, with the most parsimonious interpretation of intermediate host distribution implying a minimum of three host switches within the family., (Crown Copyright © 2022. Published by Elsevier Ltd. All rights reserved.)

The Didymozoidae (Trematoda: Hemiuroidea) is among the most speciose trematode families, known from a wide range of marine teleost fishes. Despite their richness, however, didymozoid life cycles are unusually poorly known; only two first intermediate hosts are known, a marine bivalve ( Anadara trapezia ) and a pelagic gastropod ( Firoloida desmarestia ). This study uses multi-locus molecular sequence data to identify a novel first intermediate host for the family, a sessile gastropod of the genus Thylacodes Guettard (Vermetidae). The didymozoid infection is not identified to species but, based on molecular phylogenetic analyses, it is close to Saccularina magnacetabula Louvard et al. , 2022, which uses a bivalve as a first intermediate host. The distribution of known first intermediate hosts of didymozoids (a bivalve, a holoplanktonic gastropod and a sessile gastropod that feeds with the use of mucus nets) suggests that first intermediate host use within the Didymozoidae has been opportunistically driven by the trophic ecology of potential mollusc hosts and has involved significant host-switching events.

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., (Copyright © 2022 Elsevier B.V. All rights reserved.)

Plesiochorus Looss, 1901 is a genus of Gorgoderidae infecting the urinary bladders of marine turtles globally. Currently, just two morphologically similar species are recognised, Plesiochorus cymbiformis (Rudolphi, 1819) Looss, 1901 and Plesiochorus elongatus Pigulevsky, 1953, which have been distinguished by molecular data and subtle morphological differences. Here we describe a new species, Plesiochorus irwinorum n. sp., infecting hawksbill turtles (Eretmochelys imbricata (L.)), which is primarily distinguished from the other two species of Plesiochorus on the basis of ITS2, cox1 and 28S sequence data. Morphometric data for specimens examined during this study overlap between P. cymbiformis and P. irwinorum n. sp. for every measured feature, rendering them functionally cryptic. However, principal components analysis clearly distinguishes the two species. Additionally, we report new specimens of P. cymbiformis, and provide new sequence data for specimens from Australian loggerhead (Caretta caretta (L.)) and hawksbill turtles. There is little understanding of the host-specificity or geographical distribution of the three species of Plesiochorus, and it remains possible that some of the previously reported sequences have been attributed to the wrong species., (© 2022. The Author(s).)

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., (© 2022. The Author(s).)

Species of the digenean genus Enenterum Linton, 1910 (Lepocreadioidea: Enenteridae) are characterised primarily by their elaborate oral suckers, which are divided into varying numbers of anteriorly directed lobes, and their host-restriction to herbivorous marine fishes of the family Kyphosidae. We describe Enenterum petrae n. sp. from the brassy chub Kyphosus vaigiensis (Quoy Gaimard) collected off Lizard Island, Great Barrier Reef, Queensland, Australia. Enenterum petrae n. sp. is readily differentiated from congeners by its unique oral sucker morphology, in having a minute pharynx, and the combination of a genital cap and accessory sucker. We also provide the first record of Enenterum kyphosi Yamaguti, 1970 from Australia based on material obtained from the blue sea chub Kyphosus cinerascens (Forsskl) collected off Lizard Island and North Stradbroke Island, Queensland. Morphologically, our specimens of E. kyphosi agree closely with descriptions of this species from Hawaii and South Africa, and despite lack of molecular data from outside of Australian waters, we consider all three reports to represent a single, widespread species. The first ITS2 and COI mtDNA gene sequences for species of Enenterum are provided and molecular phylogenetic analyses of 28S rDNA gene sequences place these species in a strongly-supported clade with the type-species of the genus, Enenterum aureum Linton, 1910. The oral suckers of both E. kyphosi and E. petrae n. sp. can be interpreted as having varying numbers of lobes depending on the particular specimen and how the division between lobes is defined. Scanning electron microscopical images improves understanding of the morphology of the enenterid oral sucker, and permits speculation regarding the evolutionary history leading to its specialisation in this lineage.

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., (© 2022. The Author(s).)

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., (Copyright © 2022 Elsevier B.V. All rights reserved.)

The first first-intermediate host for a species of Didymozoidae (Trematoda: Hemiuroidea), a bivalve of the family Arcidae, is identified using multi-loci molecular data. First intermediate, (likely) third intermediate, and adult stages of a new didymozoid taxon (Saccularina magnacetabula n. gen. n. sp.) from Moreton Bay, Queensland, Australia were collected from the Sydney cockle Anadara trapezia (Deshayes) (Arcoidea: Arcidae), Sillago sp. (Sillaginidae) and Elops hawaiensis Regan (Elopiformes: Elopidae), respectively, and genetically matched. Infections in A. trapezia were present as sporocysts and cystophorous cercariae, and infected tissue at the base of the gills. Morphologically, S. magnacetabula is distinctive relative to all other didymozoids in the combination of hermaphroditism, mate-pairing, filiform body shape, the presence of a ventral sucker, a single testis, and a saccular excretory vesicle at the posterior extremity. Molecular sequence data were generated for S. magnacetabula and 42 other putative didymozoid species to explore relationships within the Didymozoidae and Hemiuroidea. In molecular phylogenetic analyses of the 28S rDNA region, the new genus forms a clade with an undescribed taxon from the redthroat emperor, Lethrinus miniatus (Bloch & Schneider) (Perciformes: Lethrinidae), from the Great Barrier Reef, and another uncharacterised taxon from E. hawaiensis. This clade is sister to a moderately well-supported clade comprising all other didymozoid species for which sequences are available, including representatives of five of the six presently recognised subfamilies. The infection of a bivalve by a didymozoid is discussed in the context of the overwhelming use of gastropod molluscs as first intermediate hosts by the Hemiuroidea., (Copyright © 2022 Australian Society for Parasitology. All rights reserved.)

We report on Neoechinorhynchus aldrichettae Edmonds, 1971 (Acanthocephala: Neoechinorhynchidae), obtained from yellow-eye mullet Aldrichetta forsteri (Valenciennes) (Mugiliformes: Mugilidae) from the Huon River, Tasmania, Australia. We provide new 18S and 28S rDNA gene sequence data for N. aldrichettae, assess its phylogenetic position relative to other species of Neoechinorhynchus and provide an updated morphological account of this species including detail of features omitted in the type-description, specifically of the apical organ, a collar at the base of the neck and a para-receptacle structure associated with the proboscis receptacle. We determine that eggs in this species are ovoid, without polar prolongations of fertilisation membrane, which permits assignment of N. aldrichettae to the subgenus Neoechinorhynchus. Our phylogenetic analyses place N. (N.) aldrichettae in a clade with other species of Neoechinorhynchus which parasitise mullets in marine and estuarine waters. We find that, in terms of monophyletic clades, the current subgeneric classification system for Neoechinorhynchus is not reflected in our phylogenetic analyses., (© 2022. The Author(s), under exclusive licence to Springer Nature B.V.)

The taxonomy of species of Bivesicula Yamaguti, 1934 is analysed for samples from holocentrid, muraenid and serranid fishes from Japan, Ningaloo Reef (Western Australia), the Great Barrier Reef (Queensland), New Caledonia and French Polynesia. Analysis of three genetic markers (cox1 mtDNA, ITS2 and 28S rDNA) identifies three strongly supported clades of species and suggests that Bivesicula as presently recognized is not monophyletic. On the basis of combined morphological, molecular and biological data, 10 species are distinguished of which five are proposed as new. Bivesicula Clade 1 comprises seven species of which three are effectively morphologically cryptic relative to each other; all seven infect serranids and four also infect holocentrids. Bivesicula Clade 2 comprises three species of which two are effectively morphologically cryptic relative to each other; all three infect serranids and one also infects a muraenid. Bivesicula Clade 3 comprises two known species from apogonids and a pomacentrid, and forms a clade with species of Paucivitellosus Coil, Reid & Kuntz, 1965 to the exclusion of other Bivesicula species. Taxonomy in this genus is made challenging by the combination of low resolving power of ribosomal markers, the existence of regional cox1 mtDNA populations, exceptional and unpredictable host-specificity and geographical distribution, and significant host-induced morphological variation.

Blood flukes of the family Spirorchiidae Stunkard, 1921 are significant pathogens of marine turtles, both in the wild and in captivity. Despite causing considerable disease and mortality, little is known about the life cycles of marine species, with just four reports globally. No complete life cycle has been elucidated for any named species of marine spirorchiid, but the group is reported to use vermetid and fissurellid gastropods, and terebelliform polychaetes as intermediate hosts. Here we report molecular evidence that nine related spirorchiid species infect vermetid gastropods as first intermediate hosts from four localities along the coast of Queensland, Australia. ITS2 rDNA and cox1 mtDNA sequence data generated from vermetid infections provides the first definitive identifications for the intermediate hosts for the four species of Hapalotrema Looss, 1899 and Learedius learedi Price, 1934. Additionally, we provide a new locality report for larval stages of Amphiorchis sp., and evidence of three additional unidentified spirorchiid species in Australian waters. Based on the wealth of infections from vermetids during this study, we conclude that the previous preliminary report of a fissurellid limpet as the intermediate host for L. learedi was likely mistaken. The nine species found infecting vermetids during this study form a strongly supported clade exclusive of species of the other two marine spirorchiid genera for which sequence data are available; Carettacola Manter & Larson, 1950 which falls sister to the vermetid-infecting clade + a small clade of freshwater spirorchiids, and Neospirorchis Price, 1934 which is distantly related to the vermetid-infecting clade. We provide further evidence that spirorchiid transmission can occur in closed system aquaria and show that spirorchiid transmission occurs at both an important turtle rookery (Heron Island, southern Great Barrier Reef, Australia) and foraging ground (Moreton Bay, Australia). We discuss the implications of our findings for the epidemiology of the disease, control in captivity, and the evolution of vermetid exploitation by the Spirorchiidae., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Molecular data have transformed approaches to trematode taxonomy by providing objective evidence for the delineation of species. However, although the data are objective, the interpretation of these data regarding species boundaries is subjective, especially when different markers conflict. Conserved markers can lead to an underestimation of richness and those used for finer species delineation have the capacity to inflate species recognition, perhaps unrealistically. Here we examine molecular and morphological evidence for species recognition in an especially confusing system, the lepocreadiid genus Preptetos Pritchard, 1960 in acanthuriform fishes of the tropical Indo-west Pacific. We consider species boundaries within this genus based on combined data (ITS2 and 28S rDNA; cox1 mtDNA and morphometrics) for substantial new collections. Delineation of species using only morphological data suggest fewer species than analysis of the sequence data; the latter suggests the presence of potential cryptic species and analysis of different markers suggests the presence of differing numbers of species. We conclude that an integrative interpretation creates the most satisfying taxonomic hypothesis. In the light of the new data, we have chosen and propose a model of trematode species recognition that demands reciprocal monophyly in the most discriminating available molecular marker plus distinction in morphology or host distribution. By invoking these criteria, we distinguish eight species in our new tropical Indo-west Pacific collections. Six of these are new (Preptetos allocaballeroi n. sp., Preptetos paracaballeroi n. sp., Preptetos pearsoni n. sp., Preptetos prudhoei n. sp., Preptetos quandamooka n. sp. and Preptetos zebravaranus n. sp.) and we continue to recognise Preptetos cannoni Barker, Bray & Cribb, 1993 and Preptetos laguncula Bray & Cribb, 1996. Notably; two of the new species, P. allocaballeroi n. sp. and P. paracaballeroi n. sp., are morphologically cryptic relative to each other. Our criteria lead us to recognise, as species, populations with unvarying morphology and similar host relationships but which may have a complex population structure over their range. In our view, this paradigm has the capacity to render tractable the interpretation of the species status of the huge trematode fauna of the tropical Indo-west Pacific., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

We report specimens of monorchiids infecting Haemulidae from the waters off Japan and Australia; these specimens represent five species of Helicometroides Yamaguti, 1934, three of which are unambiguously new. Helicometroides murakamii n. sp. infects Diagramma pictum pictum from off Minabe, Japan; Helicometroides gabrieli n. sp. infects Plectorhinchus chrysotaenia from off Lizard Island, Australia; and Helicometroides wardae n. sp. infects Plectorhinchus flavomaculatus and Plectorhinchus multivittatus from off Heron Island, Australia. Helicometroides murakamii n. sp. and H. gabrieli n. sp. conform to the most recent diagnosis of Helicometroides in lacking a terminal organ, but H. wardae n. sp. possesses a terminal organ with distinct, robust spines; despite this morphological distinction, the three form a strongly-supported clade in phylogenetic analyses. We also report specimens morphologically consistent with Helicometroides longicollis Yamaguti, 1934, from D. pictum pictum from off Minabe, Japan, and Diagramma pictum labiosum on the Great Barrier Reef, Australia. Genetic analyses of ITS2 rDNA, 28S rDNA and cox1 mtDNA sequence data for the Japanese specimens reveal the presence of two distinct genotypes. Specimens of the two genotypes were discovered in mixed infections and are morphologically indistinguishable; neither genotype can be associated definitively with H. longicollis as originally described. We thus identify them as H. longicollis lineage 1 and 2, pending study of further fresh material. Genetic analyses of specimens from the Great Barrier Reef are consistent with the presence of only H. longicollis lineage 1. This species thus has a range that incorporates at least Australia and Japan, localities separated by over 7000 km.

We genetically characterised larval and adult specimens of species of Echinocephalu s Molin, 1858 (Gnathostomatidae) collected from various hosts found within Australian waters. Adult specimens of Echinocephalus were collected from a dasyatid stingray [ Pastinachus ater (Macleay); n = 2] from Moreton Bay, Queensland and larvae from a hydrophiine sea snake [ Hydrophis peronii (Duméril); n = 3] from Cape York Peninsula, Queensland, from an octopus ( Octopus djinda Amor & Hart; n = 3) from Fremantle, Western Australia and from a lucinid bivalve [ Codakia paytenorum (Iredale); n = 5] from Heron Island, Queensland Australia. All nematode samples were identified morphologically and genetically characterised using the small subunit nuclear ribosomal DNA (SSU). Some morphological differences were identified between previous studies of Echinocephalus spp. and those observed herein but the significance of these differences remains unresolved. Molecular phylogenetic analyses revealed that larval Echinocephalus sp. from H. peronii and C. paytenorum in Australia were very similar (with strong nodal support) to larval Echinocephalus sp. infecting two fish species from Egypt, Saurida undosquamis (Richardson) (Synodontidae) and Pagrus pagrus (Linnaeus) (Sparidae). The SSU sequences of larval Echinocephalus sp . from O. djinda and adults from P. ater formed a well-supported clade with that of adult E. overstreeti Deardorff and Ko, 1983 from the Port Jackson shark, Heterodontus portusjacksoni (Meyer), as well as that of the larval Echinocephalus sp., from the common carp ( Cyprinus carpio Linnaeus) from Egypt. This study extends the intermediate host range of Echinocephalus larvae by including a sea snake for the first time. Findings of this study highlight the importance of genetic characterisation of larval and adult specimens of Echinocephalus spp. to resolve the current difficulties in the taxonomy of this genus., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2022 The Authors.)

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., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Trematodes are the richest class of platyhelminths in the marine environment, infecting all classes of marine vertebrates as sexual adults and many phyla of marine invertebrates as part of their life cycles. Despite the cryptic nature of their existence (almost all marine trematodes are internal parasites), they have been the focus of study for almost 250 years, with the first species described in 1774. Here we review progress in the study of the "biodiversity" of these parasites, contrasting the progress made in the last 50 years (post-1971) to that in the almost 200 years before it (pre-1972). We consider an understanding of biodiversity to require knowledge of the species present in the system, an understanding of their evolutionary relationships (which informs higher classification), and, specifically for trematodes, an understanding of their complex life cycles. The fauna is now large, comprising well over 5,000 species. Although species description continues, we see evidence of a slow-down in all aspects of discovery. There has been only one completely new family identified since 1984 and the proposal of new genera is in decline as is the description of new species, especially for those of tetrapods. However, the extent to which this slow-down reflects an approach to the richness asymptote is made uncertain by changes in the field; reduced effort and difficulty of study may be important components of the effect. Regardless of how close we are to a complete description of the fauna, we infer that the outline is well-understood although the details are not. Adoption of molecular methodologies over the last 40 years have complemented morphometric analyses to facilitate objective recognition of species; however, despite these objective data, there is still inconsistency between authors on species delimitation. Molecular methodologies have also completely revolutionised inference of relationships at all levels, from within genera to between orders, and underpinned elucidation of novel life cycles. We expect the next 50 years to produce further dividends from technological innovations. The backdrop to the field will be global environmental concerns and the growing problem of funding for basic biodiversity studies., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Ankistromeces Nolan & Cribb, 2004 and Phthinomita Nolan & Cribb, 2006 are sister genera of threadlike blood flukes (Trematoda: Aporocotylidae) infecting teleost fishes of the tropical Indo-west Pacific. Here, we report new collections of these genera from Australia, Indonesia, and Japan. A new species of Ankistromeces, Ankistromeces kawamurai n. sp., is described from Siganus spinus (Linnaeus) off Okinawa, Japan, and a new species of Phthinomita, Phthinomita abdita n. sp., from Choerodon cephalotes (Castelnau), in Moreton Bay, Australia; the new species are morphologically cryptic within their respective genera and are delineated by molecular and ecological data. Ankistromeces olsoni Nolan & Cribb, 2006 is reported from Siganus fuscescens (Houttuyn) off Heron Island (southern Great Barrier Reef), Lizard Island (northern Great Barrier Reef), and Okinawa and Wakayama Prefectures, Japan and from Siganus spinus (Linnaeus) from off Bali, Indonesia. Ankistromeces mariae Nolan & Cribb, 2004 is re-reported from the type-host, Meuschenia freycineti (Quoy & Gaimard), from a new location, Gypsy Bay, Tasmania. Phthinomita poulini Nolan & Cribb, 2006 is re-reported from its type-locality, Lizard Island, from a range of mullids, including five new host species, and its range is extended to include Moreton Bay. Phthinomita symplocos Nolan & Cribb, 2006 is reported from Bali and P. hallae Nolan & Cribb, 2006, P. jonesi Nolan & Cribb, 2006, P. littlewoodi Nolan & Cribb, 2006, and P. munozae Nolan & Cribb, 2006 are each re-reported from their type-host and type-localities. New cox1 mtDNA data were generated for all known species of these two genera from new and archival material. Analyses of these data enabled an evaluation of all known Phthinomita species; P. robertsthomsoni Nolan & Cribb, 2006 is synonymised with P. adlardi Nolan & Cribb, 2006, and P. brooksi Nolan & Cribb, 2006 is synonymised with P. sasali Nolan & Cribb, 2006. We highlight the failure of ITS2 data to delineate closely related aporocotylid species. In contrast, cox1 sequence data are proving reliable and effective in this context and we recommend their incorporation in future studies of blood fluke taxonomy., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Novel multi-locus sequence data were used to assess the molecular phylogenetic relationships of fish blood flukes showing similarity to the genus Cardicola Short, 1953 (Trematoda: Aporocotylidae). Analyses of three ribosomal (ITS2, 28S & 18S) subregions and one mitochondrial (cox1) DNA subregion shows that the hitherto-monophyletic clade formed by species of Cardicola Short, 1953 also includes species of three other genera - Braya Nolan & Cribb, 2006, Elaphrobates Bullard & Overstreet, 2003 and Rhaphidotrema Yong & Cribb, 2011 - as well as a new, morphologically distinct species discovered from the heart of the yellowfin tripodfish, Tripodichthys angustifrons (Tetraodontiformes: Triacanthidae). In the context of conflicting morphological, molecular and ecological data, we argue that the recognition of seven genera produces a more satisfactory taxonomy for these parasites than considering them all as species of Cardicola. We thus recognise Cardicola (as an explicitly polyphyletic taxon) together with Braya, Elaphrobates, Rhaphidotrema and three new genera. We propose Allocardicola n. gen. for A. johnpagei n. sp. from T. angustifrons, Chanicola n. gen. for three species of Cardicola that infect the chanid Chanos chanos, and Spirocaecum n. gen. for six species of Cardicola that infect siganid fishes. We interpret the pattern of diversification seen in the clade of these seven genera as one of multiple host-switching events followed by diversification among closely-related hosts and differing levels of morphological divergence., (Copyright © 2021 Elsevier Inc. All rights reserved.)

The trematode superfamily Monorchioidea comprises three families of teleost parasites: the Monorchiidae Odhner, 1911, Lissorchiidae Magath, 1917, and Deropristidae Cable & Hunninen, 1942. All presently known lissorchiid and deropristid life cycles have gastropods as first intermediate hosts, whereas those of monorchiids involve bivalves. Here, we report an unexpected intermediate host for monorchiids; two species of Hurleytrematoides Yamaguti, 1954 use gastropods as first intermediate hosts. Sporocysts and cercariae were found infecting two species of the family Vermetidae, highly specialised sessile gastropods that form calcareous tubes, from two locations off the coast of Queensland, Australia. These intramolluscan infections broadly corresponded morphologically to those of known monorchiids in that the cercariae have a spinous tegument, oral and ventral suckers, a simple tail and distinct eye-spots. Given the simplified morphology of intramolluscan infections, genetic data provided a definitive identification. ITS2 rDNA and cox1 mtDNA sequence data from the gastropod infections were identical to two species of Hurleytrematoides, parasites of butterflyfishes (Chaetodontidae); Hurleytrematoides loi McNamara & Cribb, 2011 from Moreton Bay (south-eastern Queensland) and Heron Island (southern Great Barrier Reef) and Hurleytrematoides morandi McNamara & Cribb, 2011 from Heron Island. Notably, species of Hurleytrematoides are positioned relatively basal in the phylogeny of the Monorchiidae and are a sister lineage to that of species known to infect bivalves. Thus, the most parsimonious evolutionary hypothesis to explain infection of gastropods by these monorchiids is that basal monorchiids (in our analyses, species of Cableia Sogandares-Bernal, 1959, Helicometroides Yamaguti, 1934 and Hurleytrematoides) will all prove to infect gastropods, suggesting a single host switching event into bivalves for more derived monorchiids (17 other genera in our phylogenetic analyses). A less parsimonious hypothesis is that the infection of vermetids will prove to be restricted to species of Hurleytrematoides, as an isolated secondary recolonisation of gastropods from a bivalve-infecting lineage. Regardless of how their use arose, vermetids represent a dramatic host jump relative to the rest of the Monorchiidae, one potentially enabled by their specialised feeding biology., (Copyright © 2021 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

Of over 250 species of Monorchiidae Odhner, 1911, just four are known from gerreid fishes. Here, we report adult specimens of a new species infecting Gerres oyena (Forsskål) and Gerres subfasciatus Cuvier from off Heron Island and North Stradbroke Island, Queensland, Australia. The species is morphologically most similar to the concept of Lasiotocus Looss, 1907, which currently comprises eight species, in the possession of an unspined genital atrium, bipartite terminal organ, round oral sucker and unlobed ovary. However, phylogenetic analyses of the 28S ribosomal DNA gene region shows the species to be distantly related to the two sequenced species of Lasiotocus - Lasiotocus mulli (Stossich, 1883) Odhner, 1911 and Lasiotocus trachinoti Overstreet & Brown, 1970 - and that it clearly requires a distinct genus; thus, we propose Gerricola queenslandensis n. g., n. sp. Morphologically, G. queenslandensis n. g., n. sp. differs significantly from L. mulli and L. trachinoti only in the possession of distinctly longer caeca, which terminate in the post-testicular region, and in the absence of a distinct gap in the terminal organ spines. The remaining species of Lasiotocus possess caeca that also terminate in the post-testicular region, which might warrant their transfer to Gerricola n. g. However, doubt about their monophyly due to a combination of significant morphological variation, a lack of information on some features and infection of a wide range of hosts, lead us to retain these taxa as species of Lasiotocus until molecular sequence data are available to better inform their phylogenetic and taxonomic positions. Sporocysts and cercariae of G. queenslandensis n. g., n. sp. were found in a lucinid bivalve, Codakia paytenorum (Iredale), from Heron Island. Sexual adult and intramolluscan stages were genetically matched with the ITS2 ribosomal DNA and cox1 mitochondrial DNA regions. This is the second record of the Lucinidae as a first intermediate host for the Monorchiidae. Additionally, we report sporocysts and cercariae of another monorchiid infection in a tellinid bivalve, Jactellina clathrata (Deshayes), from Heron Island. Molecular sequence data for this species do not match any sequenced species and phylogenetic analyses do not suggest any generic position.

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., Competing Interests: Declaration of Competing Interest The authors declare that they have no conflict of interest., (Copyright © 2020 Elsevier B.V. All rights reserved.)

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.

Three new species of the family Bucephalidae Poche, 1907 (Trematoda: Digenea) are described from the yellowtail pike, Sphyraena obtusata Cuvier (Sphyraenidae), from Moreton Bay, Queensland, Australia. The three species are morphologically consistent with the present broad concept of the genus Bucephalus Baer, 1827, but significant phylogenetic and ecological differences relative to the type-species of Bucephalus require the proposal of a new genus. Aenigmatrema n. g. is proposed for A. undecimtentaculatum n. sp. (type-species), A. inopinatum n. sp. and A. grandiovum n. sp. In addition, based on morphological, ecological and biogeographical similarities, we recombine two existing species of Bucephalus as Aenigmatrema kaku (Yamaguti, 1970) n. comb. and Aenigmatrema sphyraenae (Yamaguti, 1952) n. comb. Although the three species described in this study are extremely morphologically similar, they can be differentiated from each other, and from A. kaku and A. sphyraenae, morphometrically on the basis of egg size, tentacle number and a combination of the caecum and vitelline field lengths. Complete ITS2 rDNA, partial 28S rDNA and partial cox1 mtDNA sequence data were generated for the three new species, which formed a well-supported clade in all 28S phylogenetic analyses. An expanded phylogenetic tree for the subfamily Bucephalinae Poche, 1907 is presented, demonstrating unresolved issues with the morphology-based taxonomy of the subfamily. The three largest genera, Bucephalus, Rhipidocotyle Diesing, 1858 and Prosorhynchoides Dollfus, 1929 remain extensively polyphyletic, indicating the need for significant further systematic revision.

Prior to the present study, species of the trematode family Monorchiidae Odhner, 1911 had been reported from four of the ten families of tetraodontiform fishes: the Balistidae, Monacanthidae, Ostraciidae and Tetraodontidae. Here we report the first monorchiid from the family Triacanthidae, Pseudohurleytrema yolandae n. sp. infecting Tripodichthys angustifrons (Hollard), from Moreton Bay, Queensland, Australia. The species conforms well to the morphological concept of the subfamily Hurleytrematinae Yamaguti, 1958, and the genus Pseudohurleytrema Yamaguti, 1954, in the possession of filamented eggs, a bipartite terminal organ, and a unipartite seminal vesicle. Relative to the other three recognised species of Pseudohurleytrema, the present species is distinctive in the size of the testis and eggs, position of the ovary, and the form of the vitellarium and excretory vesicle. We consider Pseudohurleytrema magnum Kaikabad & Bilqees in Bilqees, 1991 as a species inquirenda. Sequence data for the 28S ribosomal RNA gene and cox1 mitochondrial gene were generated for P. yolandae, providing the first molecular data for the genus. Phylogenetic analysis showed that P. yolandae does not form a clade with the other three hurleytrematine genera for which there are molecular data (Helicometroides Yamaguti, 1934, Hurleytrematoides Yamaguti, 1953 and Provitellus Dove & Cribb, 1998), forming a poorly-supported clade with Proctotrema addisoni Searle, Cutmore & Cribb, 2014 within the clade of the subfamily Monorchiinae Odhner, 1911. The four hurleytrematine genera resolved as four distinct clades, indicating that the current subfamilial classification requires comprehensive revision.

The diversity of color vision systems found in extant vertebrates suggests that different evolutionary selection pressures have driven specializations in photoreceptor complement and visual pigment spectral tuning appropriate for an animal's behavior, habitat, and life history. Aquatic vertebrates in particular show high variability in chromatic vision and have become important models for understanding the role of color vision in prey detection, predator avoidance, and social interactions. In this study, we examined the capacity for chromatic vision in elasmobranch fishes, a group that have received relatively little attention to date. We used microspectrophotometry to measure the spectral absorbance of the visual pigments in the outer segments of individual photoreceptors from several ray and shark species, and we sequenced the opsin mRNAs obtained from the retinas of the same species, as well as from additional elasmobranch species. We reveal the phylogenetically widespread occurrence of dichromatic color vision in rays based on two cone opsins, RH2 and LWS. We also confirm that all shark species studied to date appear to be cone monochromats but report that in different species the single cone opsin may be of either the LWS or the RH2 class. From this, we infer that cone monochromacy in sharks has evolved independently on multiple occasions. Together with earlier discoveries in secondarily aquatic marine mammals, this suggests that cone-based color vision may be of little use for large marine predators, such as sharks, pinnipeds, and cetaceans., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The superfamily Pronocephaloidea Looss, 1899 comprises digeneans occurring in the gut and respiratory organs of fishes, turtles, marine iguanas, birds and mammals. Although many life cycles are known for species of the Notocotylidae Lühe, 1909 maturing in birds and mammals, relatively few are known for the remaining pronocephaloid lineages. We report the cercariae of five pronocephaloid species from marine gastropods of the Queensland coast, Australia. From Lizard Island, northern Great Barrier Reef, we report three cercariae, two from Rhinoclavis vertagus (Cerithiidae) and one from Nassarius coronatus (Nassariidae). From Moreton Bay, southern Queensland, an additional two cercariae are reported from two genotypes of the gastropod worm shell Thylacodes sp. (Vermetidae). Phylogenetic analysis using 28S rRNA gene sequences shows all five species are nested within the Pronocephaloidea, but not matching or particularly close to any previously sequenced taxon. In combination, phylogenetic and ecological evidence suggests that most of these species will prove to be pronocephalids parasitic in marine turtles. The Vermetidae is a new host family for the Pronocephaloidea.

We explore the growing issue of cryptic speciation in the Digenea through study of museum material and newly collected specimens consistent with the enenterid genus Koseiria from five species of the Kyphosidae and Chaetodontoplus meredithi Kuiter (Pomacanthidae) collected in the Indo-Pacific. We use an integrated approach, employing traditional morphometrics, principal components analysis (PCA), and molecular data (ITS2 and 28S rDNA). Our results support recombination of Koseiria allanwilliamsi Bray & Cribb, 2002 as Proenenterum allanwilliamsi (Bray & Cribb, 2002) n. comb. and transfer of Koseiria huxleyi Bray & Cribb, 2001 to a new genus as Enenterageitus huxleyi (Bray & Cribb, 2002) n. comb. Molecular data indicate the presence of four further species consistent with Koseiria, one from Western Australia (sequence data only) and three from eastern Australia. All three eastern Australian species are morphologically consistent with Koseiria xishaensis Gu & Shen, 1983, but distinct from all other previously described species. Although K. xishaensis has been reported from Australia, we conclude that the similarity of the present forms to the original description of K. xishaensis means records of this species from Japan, Palau and Australia are unreliable. Because the eastern Australian forms cannot be reliably ascribed to K. xishaensis, we describe Koseiria argalea n. sp., Koseiria laiphopharophora n. sp., and Koseiria pyknophora n. sp., following application of PCAs and iterative refinement of species concepts and type series. These analyses did not allow convincing identification hypotheses for all specimens examined. In this genus, both morphological and molecular data, together with reliable host identifications, are essential for species recognition, and thus we refrain from attempting to name samples lacking molecular data. The issues presented by these taxa encapsulate those of trematodes in the region as a whole. Many records require dramatically improved supporting data, leading to substantial uncertainly in the identification of this fauna., (Copyright © 2019 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

The Enenteridae Yamaguti, 1958 and Gyliauchenidae Fukui, 1929 exhibit an interesting pattern of host partitioning in herbivorous fishes of the Indo-West Pacific. Enenterids are known almost exclusively from fishes of the family Kyphosidae, a group of herbivorous marine fishes common on tropical and temperate reefs. In contrast, gyliauchenids are found in most of the remaining lineages of marine herbivorous fishes, but until the present study, had never been known from kyphosids. Here we report on the first species of gyliauchenid known from a kyphosid. Endochortophagus protoporus gen. nov., sp. nov. was recovered from the Western buffalo bream, Kyphosus cornelii (Whitley, 1944), collected off Western Australia. Kyphosus cornelii also hosts an enenterid, Koseiria allanwilliamsi Bray & Cribb, 2002, and is thus the first fish known in which enenterids and gyliauchenids co-occur. Molecular phylogenetic analyses place the new species close to those of Affecauda Hall & Chambers, 1999 and Flagellotrema Ozaki, 1936, but there is sufficient morphological evidence, combined with the unusual host, to consider it distinct from these genera. We discuss factors which may have contributed to the host partitioning pattern observed between enenterids and gyliauchenids.

Four new monorchiid trematodes are reported from Moreton Bay, Australia; three new species of Provitellus Dove & Cribb, 1998 and one species of Ovipusillus Dove & Cribb, 1998, are described from Gnathanodon speciosus (Forsskål) (Carangidae), the golden trevally. Provitellus chaometra n. sp., Provitellus infrequens n. sp. and Provitellus infibrova n. sp. differ significantly from the only other species of this genus, Provitellus turrum Dove & Cribb, 1998, in the structure of the eggs, vitelline follicles and terminal organ. The four species are united, however, in the possession of short caeca and a long genital atrium, a combination not reported previously. Ovipusillus geminus n. sp. strongly resembles its only congener, Ovipusillus mayu Dove & Cribb, 1998, but differs in the morphology of the diverticulum in the cirrus-sac and the shape of the pharynx. Complete ITS2 and partial 28S rDNA sequence data were generated for all four species, as well as for two known species of Hurleytrematoides Yamaguti, 1953, Hurleytrematoides galzini McNamara & Cribb, 2011 and Hurleytrematoides loi McNamara & Cribb, 2011. These sequences were analysed with those for other monorchiids available on GenBank, and phylogenetic analyses showed that the four species of Provitellus and two species of Ovipusillus each form strongly supported clades. As with previous monorchiid phylogenetic studies, however, the overall resolution of the phylogeny of the Monorchiidae is poor.

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., (Copyright © 2018 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.)

A rich fauna of digenetic trematodes has been documented from the Great Barrier Reef (GBR), yet little is known of the complex life-cycles of these parasites which occur in this diverse marine ecosystem. At Heron Island, a small coral cay at the southern end of the GBR, the intertidal marine gastropod Clypeomorus batillariaeformis Habe & Kusage (Cerithiidae) is especially abundant. This gastropod serves as an intermediate host for 12 trematode species utilising both fish and avian definitive hosts. However, 11 of these species have been characterised solely with morphological data. Between 2015 and 2018 we collected 4870C. batillariaeformis from Heron Island to recollect these species with the goal of using molecular data to resolve their phylogenetic placement. We found eight of the 12 previously known species and two new forms, bringing the total number of digenean species known to parasitise C. batillariaeformis to 14. The families of this trematode community now include the Atractotrematidae Yamaguti, 1939, Bivesiculidae Yamaguti, 1934, Cyathocotylidae Mühling, 1898, Hemiuridae Looss, 1899, Heterophyidae Leiper, 1909, Himasthlidae Odhner, 1910, Microphallidae Ward, 1901, and Renicolidae Dollfus, 1939. Molecular data (ITS and 28S rDNA) were generated for all trematode species, and the phylogenetic position of each species was determined. The digenean community parasitising C. batillariaeformis includes several common species, as well as multiple species which are uncommon to rare. Although most of those trematodes in the community which exploit fishes as definitive hosts have remained common, the composition of those which utilise birds appears to have shifted over time., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Fishes of the elasmobranch superorder Batoidea and the basal teleost superorder Elopomorpha were assessed for blood flukes (Digenea: Aporocotylidae) during a parasitological survey conducted in Moreton Bay, Queensland, Australia. A new blood fluke genus and species, Ogawaia glaucostegi n. gen., n. sp., is described from the Giant shovelnose ray, Glaucostegus typus (Anonymous [Bennett]) (Rhinopristiformes: Glaucostegidae). Ogawaia glaucostegi differs from species of all other aporocotylid genera in the combination of the absence of anterior caeca and oral sucker, having a pronounced distal oesophageal chamber, a strongly coiled testis and a common genital pore. The new species most closely resembles Myliobaticola richardheardi Bullard & Jensen, 2008, from which it differs in lacking an oral sucker and in possessing a straight (rather than coiled) oesophagus, longer caeca in proportion to the oesophageal and total body length, and a much longer testis relative to body length. Ogawaia glaucostegi is just the eighth aporocotylid described from chondrichthyans, of which four belong to monotypic genera. This is the first description of a blood fluke from the order Rhinopristiformes, and the first of a chondrichthyan-infecting aporocotylid from Australian waters. Elopicola bristowi Orélis-Ribeiro & Bullard, 2017 is reported from Australia for the first time, from the type-host, Elops hawaiensis Regan (Elopiformes: Elopidae). This species is identified by morphological and molecular data and distinctions between our specimens and those of the original description are discussed., (Copyright © 2018 Elsevier B.V. All rights reserved.)

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.