The oceanic pleuston community as a potentially crucial life-cycle pathway for pelagic fish-infecting parasitic worms.

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.

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.

Novel insights into the genetics, morphology, distribution and hosts of the global fish parasitic digenean Proctoeces maculatus (Looss, 1901) (Digenea: Fellodistomidae).

Larval stages of the widely distributed digenean species Proctoeces maculatus (Looss, 1901) were reported 40 years ago from South Africa in the common octopus, Octopus vulgaris Cuvier (Octopodidae). However, the absence of adult specimens and molecular data from this region has hindered a comprehensive understanding of its distribution. In this study, we collected three species of intertidal and near-shore marine fishes [Clinus superciliosus (L.) (Clinidae), Diplodus capensis (Smith) (Sparidae) and Sparodon durbanensis (Castelnau) (Sparidae)] along the South African coast and discovered adult specimens of P. maculatus at five localities. By employing a combination of morphological and molecular techniques, including 28S rDNA, 18S rDNA and COI mtDNA analyses, the first report of adult P. maculatus from South Africa is presented. The findings encompass a comprehensive morphological description and molecular data, illuminating the true distribution of this species in the region.

First elucidation of a didymozoid life cycle: Saccularina magnacetabula n. gen. n. sp. infecting an arcid bivalve.

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.

Molecular phylogenetic analysis of the problematic genus Cardicola (Digenea: Aporocotylidae) indicates massive polyphyly, dramatic morphological radiation and host-switching.

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.

Parasitic copepods of the family Lernanthropidae Kabata, 1979 (Copepoda: Siphonostomatoida) from Australian fishes, with descriptions of seven new species.

The total number of species of Lernanthropidae previously recorded from Australian waters is 15 (i.e., one species each of Aethon Krøyer, 1837, Lernanthropodes Bere, 1936, and Lernanthropsis Do, in Ho Do, 1985; 10 species of Lernanthropus de Blainville, 1822; and two species of Sagum Wilson, 1913), and all of these records are reviewed. We report here the presence of three species of Aethon. One species, A. garricki Hewitt, 1968, is reported from Australian waters for the first time and a new species, A. bicamera sp. nov., is described from the latrid, Latris lineatus (Forster, 1801) caught off South Australia. The genus Lernanthropodes is represented by a single species, L. trachinoti Pillai, 1962. We recognize Chauvanium Kazachenko, Kovaleva, Nguyen Ngo, 2017 as a subjective synonym of Lernanthropodes and transfer its type and only species C. chauvani Kazachenko, Kovaleva, Nguyen Ngo, 2017 which becomes Lernanthropodes chauvani (Kazachenko, Kovaleva, Nguyen Ngo, 2017) n. comb. Lernanthropsis mugilii (Shishido, 1898) is reported here from Mugil cephalus Linnaeus, 1758 sampled in Queensland and in New South Wales. The genus Lernanthropus is the most species rich and we report the presence of 20 nominal species on Australian marine fishes. This total includes six new species: L. alepicolus sp. nov. from Alepes apercna Grant, 1987, L. elegans sp. nov. from Atractoscion aequidens (Cuvier, 1830), L. gnathanodontus sp. nov. from Gnathanodon speciosus (Forsskål, 1775), L. paracruciatus sp. nov. from Protonibea diacanthus (Lacepède, 1802), L. pemphericola sp. nov. from Pempheris compressa (White, 1790), and L. selenotoca sp. nov. from Selenotoca multifasciata (Richardson, 1846). In addition, we report the presence of another four species in Australian waters for the first time: L. abitocephalus Tripathi, 1962, L. cadenati Delamare Deboutteville Nuñes-Ruivo, 1954, L. microlamini Hewitt, 1968, and L. pomadasysis Rangnekar Murti, 1961. After reexamination of the types of L. paenulatus Wilson, 1922 held in the USNM, we relegate this species to subjective synonymy with L. seriolii Shishido, 1898. Previous records of L. paenulatus from Australian Seriola species should be reassigned to L. seriolii. Lernanthropus ecclesi Kensley Grindley, 1973 is recognized as a junior subjective synonym of L. micropterygis Richiardi, 1884, and L. delamarei Marques, 1960, which is based on the male only, is tentatively considered to be a junior subjective synonym of L. micropterygis. Males are described for the first time for three species; L. breviculus Kabata, 1979, L. microlamini and L. mollis Kabata, 1979. A member of the genus Mitrapus Song Chen, 1976, M. oblongus (Pillai, 1964), is reported from Australia for the first time, on Herklotsichthys castelnaui (Ogilby, 1897) caught off Queensland and New South Wales. Finally, two species of Sagum were previously known from Australia and here we add three more. Two of the newly reported species were originally described as species of Lernanthropus but we formally transfer them here to Sagum as S. lativentris (Heller, 1865) n. comb. and S. sanguineus (Song, in Song Chen, 1976) n. comb. The males of S. lativentris and S. vespertilio Kabata, 1979 are described for the first time. A key to the females of the 31 species of lernanthropids found in Australian waters is provided.

Aporocotylids from batoid and elopomorph fishes from Moreton Bay, Queensland, Australia, including a new genus and species of blood fluke infecting the Giant shovelnose ray, Glaucostegus typus (Rhinopristiformes: Glaucostegidae).

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.

Redescription of Philometra pellucida (Jägerskiöld, 1893) (Nematoda: Philometridae) parasitic in the abdominal cavity of the blackspotted puffer Arothron nigropunctatus (Bloch & Schneider) (Teleostei: Tetraodontidae) off Australia and Japan.

The nematode Philometra pellucida (Jägerskiöld, 1893) (Philometridae) is redescribed from female specimens obtained from the abdominal cavity of the marine tetraodontiform fish (blackspotted puffer) Arothron nigropunctatus (Bloch & Schneider) (Tetraodontidae) off Lizard Island, Australia and off Okinawa, Japan. This is just the second record of this parasite since its description 125 years ago. Light and scanning electron microscopical examination of these nematodes revealed some taxonomically important, previously unreported morphological features, such as the number and arrangement of cephalic papillae, presence of the oesophageal cell nucleus and the size of first-stage larvae in the uterus. This new report of P. pellucida infecting A. nigropunctatus off Australia and Japan represents new host and geographical records. A key to the species of Philometra Costa, 1845 parasitising tetraodontiform fishes is provided.

Two new and one known species of Tergestia Stossich, 1899 (Trematoda: Fellodistomidae) with novel molecular characterisation for the genus.

Combined morphological and molecular analyses are employed to characterise three species of Tergestia Stossich, 1899 (Digenea: Fellodistomidae) from fishes of Moreton Bay, Queensland, Australia. Tergestia clonacantha Manter, 1963 is reported here for the first time from the halfbeak (Beloniformes: Hemiramphidae) species Arrhamphus sclerolepis krefftii (Steindachner), Hyporhamphus australis (Steindachner), H. quoyi (Valenciennes) and H. regularis ardelio (Whitley). Two new species, both infecting trevally (Perciformes: Carangidae) species, are described: T. maryae n. sp. from Alepes apercna Grant and T. henryi n. sp. from Pantolabus radiatus (MacLeay). Complete ITS2 and partial 28S ribosomal DNA data were generated for each of the new taxa. The three species differ from each other by 47-58 base pairs (bp) in the ITS2 rDNA region. Phylogenetic analysis of 28S rDNA supports Tergestia as a reliable generic concept, with our analyses showing that some species of the genus form a well-supported clade to the exclusion of all other fellodistomids for which sequence data are available.

A re-evaluation of diversity of the Aporocotylidae Odhner, 1912 in Siganus fuscescens (Houttuyn) (Perciformes: Siganidae) and associated species.

The aporocotylid fauna of the mottled spinefoot, Siganus fuscescens (Houttuyn), from Moreton Bay, Queensland, Australia, was characterised using a combined morphological and molecular approach. Four aporocotylid species were identified, three belonging to the genus Ankistromeces Nolan & Cribb, 2006 and one to Cardicola Short, 1953. Specimens of Cardicola matched an undescribed species from the same host and locality; this species is described as Cardicola mogilae n. sp. Phylogenetic analyses of ITS2 and 28S data showed that C. mogilae n. sp. forms a strongly supported clade with other Cardicola species from siganid fishes. We record Ankistromeces olsoni Nolan & Cribb, 2006 in Moreton Bay for the first time, redescribe A. dunwichensis Nolan & Cribb, 2006 on the basis of new specimens and sequence data and re-report Ankistromeces sp. X from Moreton Bay based on molecular data. We review the status of the ten putative species of aporocotylids reported from siganids. Small variation in ITS2 rDNA sequences, in association with different geographic localities, was previously used to separate Cardicola lafii Nolan & Cribb, 2006 from C. parilus Nolan & Cribb, 2006, C. bartolii Nolan & Cribb, 2006 from C. watsonensis Nolan & Cribb, 2006, C. tantabiddii Nolan & Cribb, 2006 from Cardicola sp. 2, Ankistromeces sp. Y from A. olsoni and Ankistromeces sp. X from Ankistromeces sp. Z. These five combinations are reinterpreted as each representing a single species; Cardicola lafii is recognised as the senior synonym of C. parilus and C. bartolii as the senior synonym of C. watsonensis. This study thus suggests that six, rather than ten, species should be recognised as infecting S. fuscescens. This richness remains greater than is known for any other fish species and siganids are, so far, unique among fishes in harbouring two strongly radiated lineages of aporocotylids.

A complex of Cardicola Short, 1953 (Digenea: Aporocotylidae) species infecting the milkfish Chanos chanos Forsskål (Gonorynchiformes), with descriptions of two new species.

Two new species of Cardicola Short, 1953 are described from the milkfish, Chanos chanos Forsskål (Gonorynchiformes: Chanidae), obtained from off Lizard Island on the northern Great Barrier Reef (GBR) and North Stradbroke Island in southeast Queensland. These are the first known blood flukes from this order of fishes. Cardicola suni n. sp. differs from all other Cardicola spp. by a combination of a large ovoid oral sucker surrounding a subterminal mouth, recurved tegumental spines up to 16 µm long, anterior caeca occupying 25.1-31.3% (28.7%) of total body length and a mostly-intercaecal ovary. Cardicola jiigurru n. sp. differs from C. suni n. sp. and all other species of Cardicola by a combination of a narrowly lanceolate body, weakly-muscularised and poorly-demarcated oral sucker, minute tegumental spines <1 µm in length, and anterior caeca occupying 15.9-22.0% (19.4%) of total body length, an almost entirely post-caecal ovary and the male genital pore terminal on a dorsolateral protuberance. A third species, closely resembling C. suni n. sp., was also discovered off Wangetti Beach, north Queensland, but is not described due to lack of material. Molecular phylogenetic analysis, based on both ITS2 and partial 28S rDNA regions, shows that these three species form a clade nested within that formed by existing species of Cardicola.

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.

Pseudobacciger cheneyae n. sp. (Digenea: Gymnophalloidea) from Weber's chromis (Chromis weberi Fowler & Bean) (Perciformes: Pomacentridae) at Lizard Island, Great Barrier Reef, Australia.

A new species of digenean, Pseudobacciger cheneyae n. sp., is described from the intestines of Weber's chromis (Chromis weberi Fowler & Bean) from off Lizard Island, Great Barrier Reef, Australia. This species differs from the three described species of Pseudobacciger Nahhas & Cable, 1964 [P. cablei Madhavi, 1975, P. harengulae Yamaguti, 1938 and P. manteri Nahhas & Cable, 1964] in combinations of the size of the suckers and the length of the caeca. The host of the present species is a perciform (Family Pomacentridae) which contrasts with previous records of the genus which are almost exclusively from clupeiform fishes. The genus Pseudobacciger is presently recognised within the family Faustulidae (Poche, 1926) but phylogenetic analyses of 28S and ITS2 rDNA show that the new species bears no relationship to species of four other faustulid genera (Antorchis Linton, 1911, Bacciger Nicoll, 1924, Paradiscogaster Yamaguti, 1934 and Trigonocryptus Martin, 1958) but that instead it is nested within the Gymnophalloidea (Odhner, 1905) as sister to the Tandanicolidae (Johnston, 1927). This result suggests that the Faustulidae is polyphyletic.

Rhaphidotrema kiatkiongi, a new genus and species of blood fluke (Digenea: Aporocotylidae) from Arothron hispidus (Osteichthyes: Tetraodontidae) from the Great Barrier Reef, Australia.

A new genus of fish blood flukes (Aporocotylidae Odhner, 1912) is proposed for a species found on reefs surrounding Lizard Island on the northern Great Barrier Reef. Rhaphidotrema kiatkiongi gen. et sp. n. was recovered from the heart of the stars-and-stripes pufferfish, Arothron hispidus (L.) (Tetraodontidae). Rhaphidotrema kiatkiongi is notable as the first digenean reported to possess a penis stylet. It also differs from all other aporocotylid flukes in having a combination of 18-19 testes in a group at the ends of the intestinal caeca, a broad lanceolate body shape with a dextrally-directed posterior bend at the level of the male genital pore, and separate genital pores, with the female genital pore distinctly sinistral and the male genital pore slightly dextral to midline. This is the second species of aporocotylid fluke reported from this pufferfish.