RAJONCHOCOTYLE CERFONTAINE, 1899 (MONOGENEA: HEXABOTHRIIDAE) SPECIES FROM SOUTH AFRICA, WITH DISCUSSION OF THE LITERARY ACCOUNTS OF R. EMARGINATA (OLSSON, 1876).

Rajonchocotyle Cerfontaine, 1899, species are blood-feeding parasites on the gills of rajiforms. Eight species are considered valid, the most recent of which was described just after World War II. Many of the original descriptions of Rajonchocotyle species are limited in diagnostic value, and comparative museum material is scanty. The genus requires revision, and in support thereof, we provide detailed redescriptions for Rajonchocotyle albaCerfontaine, 1899, from the type host Rostroraja alba (Lacepède, 1803) and Rajonchocotyle emarginata (Olsson, 1876), Sproston, 1946, from 2 new host records, Raja straeleni Poll, 1951, and Leucoraja wallacei (Hulley, 1970), from South Africa, a new locality record. The generic diagnosis for Rajonchocotyle is amended to include greater details of the male reproductive system and confirms Paul Cerfontaine's and Nora Sproston's historic observations of the morphology of the male copulatory organ consisting of a separate proximal seminal vesicle and a distal cirrus. The lectotype of Rajonchocotyle kenojeiYamaguti, 1938, is assigned, and a comprehensive Rajonchocotyle-host species list is provided, highlighting species records that require further verification, and the purported global host range of R. emarginata is discussed.

Species of Hexabothriidae (Monogenea) may have extensive distribution ranges reflecting multiple host species: evidence from three new South African records.

Many hexabothriid genera require renewed taxonomic attention to provide additional detailed comparative data from new material to resolve existing species quandaries. Our current study describes and provides additional museum material for three hexabothriid species from three genera from South Africa: Erpocotyle catenulata (Guberlet), which is confirmed as a valid species, Heteronchocotyle gymnurae Neifar, Euzet and Ben Hassine, and Hypanocotyle bullardi Chero, Cruces, Sáez, Carolina, Camargo, Portes Santos and Luque. Evidence of these three hexabothriid species from South Africa suggests that hexabothriids have more globally widespread distributions, and that they are less host-specific than previously considered. A detailed summary of hexabothriids reported from the triakid hosts, Galeorhinus galeus and Mustelus mustelus is provided, and the host-specificity of hexabothriids is discussed.

How do fishes manage disease?

Disease drives the evolution of proactive and reactive mitigation behaviours in fishes as for terrestrial animals. Understanding fish self-remedy behaviours could discover algal bioactives, reveal novel strategies for disease management, identify new habitats or ecosystems critical to population health and conservation, and enhance knowledge of interspecific evolutionary relationships and communication.

Cleaner fish are potential super-spreaders.

Cleaning symbiosis is critical for maintaining healthy biological communities in tropical marine ecosystems. However, potential negative impacts of mutualism, such as the transmission of pathogens and parasites during cleaning interactions, have rarely been evaluated. Here, we investigated whether the dedicated bluestreak cleaner wrasse, Labroides dimidiatus, is susceptible to and can transmit generalist ectoparasites between client fish. In laboratory experiments, L. dimidiatus were exposed to infective stages of three generalist ectoparasite species with contrasting life histories. Labroides dimidiatus were susceptible to infection by the gnathiid isopod Gnathia aureamaculosa, but were significantly less susceptible to the ciliate protozoan Cryptocaryon irritans and the monogenean flatworm Neobenedenia girellae, compared with control host species (Coris batuensis or Lates calcarifer). The potential for parasite transmission from a client fish to the cleaner fish was simulated using experimentally transplanted mobile adult (i.e. egg-producing) monogenean flatworms on L. dimidiatus. Parasites remained attached to cleaners for an average of 2 days, during which parasite egg production continued, but was reduced compared with that on control fish. Over this timespan, a wild cleaner may engage in several thousand cleaning interactions, providing numerous opportunities for mobile parasites to exploit cleaners as vectors. Our study provides the first experimental evidence that L. dimidiatus exhibits resistance to infective stages of some parasites yet has the potential to temporarily transport adult parasites. We propose that some parasites that evade being eaten by cleaner fish could exploit cleaning interactions as a mechanism for transmission and spread.

Sea lice (Copepoda: Caligidae) from South Africa, with descriptions of two new species of Caligus.

Thirteen species of sea lice (family Caligidae) are reported from a range of elasmobranch and actinopterygian fishes caught off South Africa or obtained from public aquaria in South Africa. Two new species of Caligus Müller, 1785 are described: C. linearis n. sp. from Pomatomus saltatrix (Linnaeus) and C. tumulus n. sp. from Chrysoblephus cristiceps (Valenciennes). A supplementary description is provided for both sexes of Caligus tetrodontis Barnard, 1948 taken from Amblyrhynchotes honckenii (Bloch) and previous records of this parasite from South African fishes are critically reviewed. It is concluded that Caligus material from Arothron hispidus Linnaeus was previously misidentified as C. tetrodontis and is in urgent need of re-examination. Morphological and molecular observations on Caligus furcisetifer Redkar, Rangnekar & Murti, 1949 indicate that this copepod is phenotypically and genetically identical to Lepeophtheirus natalensis Kensley & Grindley, 1973, and the latter becomes a junior subjective synonym of C. furcisetifer. We include new geographical distribution records for Caligus longipedis Bassett-Smith, 1898, C. rufimaculatus Wilson, 1905 and Lepeophtheirus spinifer Kirtisinghe, 1937, extending into South African waters, as well as both new distribution and host records for Alebion gracilis Wilson, 1905, Caligus dakari van Beneden, 1892 and Lepeophtheirus acutus Heegaard, 1943. The molecular analysis confirmed the monophyly of the genus Caligus. The South African species of Caligus did not cluster together, but the two included South African species of Lepeophtheirus were recovered as sister taxa.

New monogeneans from the bathydemersal southern African endemic catshark, Holohalaelurus regani (Gilchrist, 1922).

The catshark genus Holohalaelurus Fowler is currently represented by five species distributed off the southern and eastern parts of the African coast. Very few parasitological records exist for any of these five species, representing a significant knowledge gap. We report the first monogenean species from the bathydemersal species Holohalaelurus regani (Gilchrist): a new species of Microbothriidae Price, 1936, Leptomicrobothrium holohalaelure n. sp. from the dorsal skin surface, and a new species of Hexabothriidae Price, 1942 representing a new genus, Scyliorhinocotyle narvaezae n. gen., n. sp., from the gill lamellae. Both monogenean records represent the first for any member of the catshark genus Holohalaelurus. Previously we focused on the identification of monogenean taxa of emerging veterinary importance for public aquaria. We now begin a focused effort to document a generally unexplored monogenean biodiversity from diverse marine habitats off South Africa.

Gyrodactylus molweni sp. n. (Monogenea: Gyrodactylidae) from Chelon richardsonii (Smith, 1846) (Mugilidae) from Table Bay, South Africa.

Gyrodactylus molweni sp. n. is described from the body surface and fins of the South African mullet, Chelon richardsonii (Smith, 1846) collected from Table Bay Harbour, Cape Town and is compared to five other Gyrodactylus species described from grey mullets globally namely G. zhukovi Ling, 1963 and G. mugili Zhukov, 1970 from Planiliza haematocheila (Temminck and Schlegel, 1845); G. mugelus Rawson, 1973 from Mugil cephalus L.; G. curemae Conroy and Conroy, 1985 from Mugil curema Valenciennes, 1836 and G. xiamenensis Zang,Yang and Liu, 2001 from Planiliza macrolepis (Smith, 1846). Morphologically, G. molweni sp. n. has prominent ventral bar processes that near cover the hamulus roots, marginal sickles with large rhomboid heels, slender shafts and fine points that extend beyond the sickle toes. Gyrodactylus molweni sp. n. can, however, be readily differentiated: G. mugili and G. xiamenensis have ventral bars with small ventral processes; G. zhukovi has marginal hooks sickles with slender shafts and proportionately short points and open-faced blades; G. mugelus possesses marginal hook sickles with deep, rounded heels, forward slanting shafts and an angular, square line to the inner face of the blades. Although the length of the marginal hooks of G. curemae are similar to G. molweni sp. n., their hamuli are double the size. A GenBank BlastN search with the 931 bp sequence covering ITS1, 5.8S and ITS2 gave no close hits; the nearest species for which sequences are available is G. nipponensis Ogawa and Egusa, 1978 (identity 96.56%, 899/931 bp). The proposal of G. molweni sp. n. as a new species, therefore, is well supported by both the molecular and morphological analyses presented herein. This Gyrodactylus species is the first to be described from C. richardsonii and only the second Gyrodactylus species to be described from the marine environment off the African continent.

Cleaner shrimp are a sustainable option to treat parasitic disease in farmed fish.

Chemical use is widespread in aquaculture to treat parasitic diseases in farmed fish. Cleaner fish biocontrols are increasingly used in fish farming as an alternative to medicines. However, cleaner fish are susceptible to some of their clients' parasites and their supply is largely dependent on wild harvest. In comparison, cleaner shrimp are not susceptible to fish ectoparasites and they can be reliably bred in captivity. The effectiveness of shrimp in reducing parasites on farmed fish remained unexplored until now. We tested four cleaner shrimp species for their ability to reduce three harmful parasites (a monogenean fluke, a ciliate protozoan, and a leech) on a farmed grouper. All shrimp reduced parasites on fish and most reduced the free-living early-life environmental stages - a function not provided by cleaner fish. Cleaner shrimp are sustainable biocontrol candidates against parasites of farmed fish, with the peppermint cleaner shrimp reducing parasites by up to 98%.

Anoplodiscus Sonsino, 1890 (Monogenea: Anoplodiscidae): a new Australian species, and the first African record from South African hosts.

Species of Anoplodiscus Sonsino, 1890 were previously only known from host members of Sparidae. A new species, Anoplodiscus hutsonae n. sp. is proposed for museum specimens originally collected from species of Scolopsis Cuvier (Nemipteridae) off Heron Island and Lizard Island, Australia. Additionally, Anoplodiscus tai Ogawa, 1994 is synonymised with Anoplodiscus cirrusspiralis Roubal, Armitage & Rohde, 1983 due to a lack of support for differential characters, and Anoplodiscus richiardii is considered a species inquirenda. Anoplodiscus cirrusspiralis causes visible lesions on the skin and fins of its host, and may also contribute to poor food conversion rates in sparid aquaculture. Anoplodiscus cirrusspiralis has been recorded from cultured sparids in Australia, Japan, South Africa, and South Korea, and was implicated as a disease agent in fish from the former two countries. However, the discovery of A. cirrusspiralis on Chrysoblephus gibbiceps (Valenciennes), Ch. laticeps (Valenciennes) and Cymatoceps nasutus (Castelnau) in South Africa, ?Pagrus major (Temminck & Schlegel) in South Korea, and P. auratus (Forster) in Australia, New Zealand and Japan suggests that this species may have a wide distribution and low host-specificity within the Sparidae. In South Africa, A. cirrusspiralis was first encountered on a morbid C. nasutus and Ch. gibbiceps from two public aquaria in 2009 (Two Oceans Aquarium, Cape Town and uShaka Sea World, Durban, respectively). Additional material was collected from C. laticeps kept at an abalone farm in Hermanus that originated from Struisbaai on the South African south coast. Anoplodiscus cirrusspiralis is redescribed from the South African specimens. This is the first record of a member of Anoplodiscidae Tagliani, 1912 from Africa.

Evidence of multiple species of Chilodonella (Protozoa, Ciliophora) infecting Australian farmed freshwater fishes.

Parasitic Chilodonella species, Chilodonella piscicola and Chilodonella hexasticha, cause considerable economic losses globally to freshwater farmed fish production. Some genetic studies of Chilodonella spp. have indicated that many species within the genus may form cryptic species complexes. To understand the diversity of Chilodonella spp. infecting Australian freshwater farmed fish, specimens were isolated from infected barramundi (Lates calcarifer) and Murray cod (Maccullochella peelii) from fish farms in tropical north Queensland (QLD), temperate Victoria (Vic) and New South Wales (NSW) for genetic and morphological analysis. Parasites were stained and measured for morphological description and comparative phylogenetic analyses were performed using the mitochondrial small subunit (mtSSU) rDNA marker. Morphological analyses revealed four distinct morphotypes of Chilodonella infecting farmed barramundi and Murray Cod. Three putative species were isolated from barramundi (Chilodonella hexasticha, C. acuta and C. uncinata) and one from Murray cod (C. piscicola). However, phylogenetic analyses detected only three distinct genotypes, with the putative C. hexasticha and C. piscicola sharing 100% sequence identity. This suggests that Australian isolates of C. hexasticha and C. piscicola could represent the same species and may exhibit phenotypic plasticity. Further molecular analysis, including isolates from the type localities, should be performed to support or refute the synonymy of these species.

Electrocotyle whittingtoni n. gen., n. sp. (Monogenea: Monocotylidae: Heterocotylinae) from the gills of a captive onefin electric ray, Narke capensis (Narkidae) at Two Oceans Aquarium, Cape Town, South Africa.

Electrocotyle whittingtoni n. gen., n. sp. (Monogenea: Monocotylidae) is described from the gills of a captive female onefin electric ray, Narke capensis, collected for exhibition at Two Oceans Aquarium in Cape Town, South Africa. Electrocotyle n. gen. is most similar to the heterocotyline genera Heterocotyle and Potamotrygonocotyle but could not be accommodated easily in either of these groups. The new genus is characterised by a haptor with one central and eight peripheral loculi, four unsclerotised structures on the dorsal surface of the haptor, a single unsclerotised non-sinous ridge on the ventral surface of the haptoral septa, large hamuli with a long handle and reduced guard, a vagina with sclerotised walls, and tetrahedral eggs. Molecular phylogenetic analyses based on 28S sequences strongly support the separate genus status of Electrocotyle n. gen and thus support our morphological conclusion. The Heterocotylinae is amended to accommodate the new genus, and the new species is fully described and illustrated herein. This is the first record of a monocotylid from the Narkidae. Electrocotyle whittingtoni n. gen. n. sp. is considered potentially pathogenic given its negative impact on the health of its captive host kept in the quarantine facility at Two Oceans Aquarium.

A new species of Neoheterocotyle Hargis, 1955 (Monogenea: Monocotylidae) from the gills of Rhinobatos annulatus Müller & Henle (Rhinobatidae) off the southern tip of Africa.

Neoheterocotyle robii n. sp. is described from the gills of four female Rhinobatos annulatus Müller & Henle caught as bycatch during routine demersal research trawls off the Southern Cape coast of South Africa. The new species can be distinguished from all other members of Neoheterocotyle Hargis, 1955 by the morphology of the accessory piece associated with the male copulatory organ, the presence of a distinct horseshoe-shaped muscle associated with the male copulatory organ and the presence of a sclerotised structure in the proximal portion of the vagina. This is the first Neoheterocotyle species described from South African waters.

Gyrodactylus eyipayipi sp. n. (Monogenea: Gyrodactylidae) from Syngnathus acus (Syngnathidae) from South Africa.

Gyrodactylus eyipayipi sp. n. is described from the skin, gills, flute and male brood pouch of captive specimens of the greater pipefish Syngnathus acus L., collected for and maintained at the Two Oceans Aquarium in Cape Town, South Africa. It is the first marine Gyrodactylus species reported from the African continent. The new species is compared to the three known Gyrodactylus species affecting syngnathiform hosts (G. pisculentus Williams, Kritsky, Dunnigan, Lash et Klein, 2008, G. shorti Holliman, 1963, and G. syngnathi Appleby, 1996). Although all four species have similar-sized and shaped attachment hooks with some overlap, separation of the species is possible using marginal hook morphology. The marginal hooks of G. eyipayipi measure (mean) 30 pm in total length and are larger than those of the three other species (mean, 24-28 tm). Gyrodactylus eyipayipi can also be discriminated based on differences in the shape of the marginal hook sickle notably by its long sickle point which extends far beyond the toe, its blunt rounded toe and, by the approximate rectangular shape to the base of the sickle. By comparison, the sloping toe regions of G. pisculentus and G. syngnathi give the sickle bases an approximately triangular shape, whilst the short sickle point and open aperture to the sickles of G. shorti allow for their discrimination from G. eyipayipi.

Three Dendromonocotyle species (Monogenea: Monocotylidae) reported from captive rays, including D. lotteri sp. n. from Himantura gerrardi (Elasmobranchii: Dasyatidae) in the public aquarium at the Atlantis resort, Dubai.

Dendromonocotyle lotteri sp. n. is described from the dorsal skin surface of the stingray Himantura gerrardi (Gray) on exhibit in the public aquarium at the Atlantis resort in Dubai. It is differentiated from all other Dendromonocotyle species by the unique morphology of the distal portion of the sclerotised male copulatory organ. Dendromonocotyle lotteri is the second representative in the genus with 56 marginal haptoral papillae having a papillae to loculus association represented numerically as 6-6-8-8. We found Dendromonocotyle colorni Chisholm, Whittington et Kearn, 2001 on the same host specimens at the Atlantis resort public aquarium and Dendromonocotyle kuhlii Young, 1967 on Neotrygon kuhlii (Müller et Henle) kept at Burgers' Zoo Aquarium in Arnhem, The Netherlands. Supplemental information is provided for both D. colorni and D. kuhlii. The presence of Dendromonocotyle infections in public aquaria and host specificity are discussed. A key to the 17 species of Dendromonocotyle is also provided.