Pseudorhabdosynochus
Pseudorhabdosynochus morrhua
Scientific classification e
Kingdom: Animalia
Phylum: Platyhelminthes
Class: Monogenea
Order: Dactylogyridea
Family: Diplectanidae
Genus: Pseudorhabdosynochus
Yamaguti, 1958
Species

See text

Pseudorhabdosynochus is a genus of monopisthocotylean monogeneans, included in the family Diplectanidae. The type-species of the genus is Pseudorhabdosynochus epinepheli (Yamaguti, 1938).

The genus includes more than 80 species, which are all parasitic on the gills of marine fish, especially groupers. Since groupers are mostly abundant in warm waters, species of Pseudorhabdosynochus are found in warm parts of the oceans, especially in coral reefs, but species have also been found in deep-sea groupers. Several species of Pseudorhabdosynochus are pathogens of maricultured groupers.

Morphology

Female (left) and male (right) sclerotized organs of a species of Pseudorhabdosynochus, P. caledonicus
Female (left) and male (right) sclerotized organs of a species of Pseudorhabdosynochus, P. caledonicus
A species of Pseudorhabdosynochus with captions for main parts and organs
A species of Pseudorhabdosynochus with captions for main parts and organs


All species of Pseudorhabdosynochus are small animals, ranging 0.3–1 mm in length.

As most monogeneans, they are flat, with an anterior head bearing four oculi and head glands, a main elongate body and a posterior haptor. The digestive system includes an anterior muscular pharynx, and two lateral intestinal branches (or caeca); as in all Platyhelminthes, there is no anus. The haptor, in the posterior part of the body, is a specialized organ used to attach to the host. The haptor includes sclerotized elements, namely a ventral bar, two lateral (dorsal) bars, two ventral hooks and two dorsal hooks, and fourteen hooklets. As in most diplectanids, the haptor bears special, characteristic, structures called squamodiscs. The squamodiscs (one ventral and one dorsal) of species of Pseudorhabdosynochus are made up of numerous rodlets aligned as concentric rows. All species of the genus have two squamodiscs except Pseudorhabdosynochus sinediscus Neifar & Euzet, 2007 in which these organs are completely lacking.

Adults are hermaphroditic. The reproductive organ include a single ovary and a single testis. As in all diplectanids, the ovary (or germarium) is anterior to the testis and loops around the right intestinal caecum. Species of Pseudorhabdosynochus are characterized by a sclerotized male copulatory organ, or "quadriloculate organ", which has the shape of a bean with four internal chambers. The vagina includes a sclerotized part, which is a complex structure with several chambers and canals, generally used for the diagnosis and differentiation of species.

Etymology and nomenclatural case

A drawing of the type-species, Pseudorhabdosynochus epinepheli
A drawing of the type-species, Pseudorhabdosynochus epinepheli

The origin of the name Pseudorhabdosynochus merits an explanation. ‘Pseudo-’ (from Greek ψευδής, pseudes, "lying, false") is a prefix commonly used in creating a new scientific name for a taxon that superficially appears to be a taxon, but actually is another. The famous Japanese parasitologist Satyu Yamaguti described a ‘membranous plaque’ on the posterior region of Pseudorhabdosynochus epinepheli and created[1] the generic name Pseudorhabdosynochus in reference to the diplectanid genus Rhabdosynochus which, in his interpretation, also had lateral plaques. It has been shown later that ‘plaques’ in P. epinepheli was an erroneous interpretation of damaged specimens.[2][3] Moreover, the diagnosis of Rhabdosynochus was later modified and it is now considered that species of this genus do not have plaques.[4] Therefore, the genus name Pseudorhabdosynochus is the result of both an erroneous observation and an erroneous comparison by Satyu Yamaguti. However, it is valid according to the ICZN.[2]

Pseudorhabdosynochus is also an interesting nomenclatural case. Yamaguti described the same species twice, the first time in 1938 [5] (as Diplectanum epinepheli Yamaguti, 1938) and the second time in 1958 [1] (as Pseudorhabdosynochus epinepheli Yamaguti, 1958). In both papers, the species was described from parasites collected from the same fish species, the grouper Epinephelus akaara off Japan. Kritsky & Beverley-Burton (1986) [2] solved this case with reference to the International Code of Zoological Nomenclature. The type-species of the genus is the name Pseudorhabdosynochus epinepheli Yamaguti, 1958. Pseudorhabdosynochus epinepheli Yamaguti, 1958 is a junior synonym of Pseudorhabdosynochus epinepheli (Yamaguti, 1938) Kritsky & Beverley-Burton, 1986 and Pseudorhabdosynochus epinepheli (Yamaguti, 1938) Kritsky & Beverley-Burton, 1986 is the modern valid combination for Diplectanum epinepheli Yamaguti, 1938. The latter species (a taxon) and the type-species of the genus (a name) are considered to correspond to the same biological taxon but have different, although valid, names. Recent re-examination of type-specimens have confirmed this interpretation. [3]

Hosts

Grouper, such as the blacktip grouper here, harbour Pseudorhabdosynochus species on their gills
Grouper, such as the blacktip grouper here, harbour Pseudorhabdosynochus species on their gills

Species of Pseudorhabdosynochus are parasitic exclusively on the gill filaments of marine fish. Most of these fish belong to the family Serranidae and are commonly known as groupers, such as many species of the genus Epinephelus, and also species of Cephalopholis, Mycteroperca and Variola. A few species have also been described from fish of the genera Alphestes,[6] Paranthias[7] and Paralabrax.[8]

As most diplectanids,[9] species of Pseudorhabdosynochus are generally strictly species-specific, i.e. a species is found on a single species of host and no others. However, several exceptions have been reported.[10] Some species of groupers harbour a wide biodiversity of Pseudorhabdosynochus species (and sometimes other monogeneans as well). Examples are Epinephelus maculatus which has 8 distinct Pseudorhabdosynochus species,[11] and Epinephelus malabaricus which has 7 species.[12][13]

Life cycle

Adult Pseudorhabdosynochus lantauensis (arrows) on the gill filament of a grouper. Scanning electron microscopy.
Adult Pseudorhabdosynochus lantauensis (arrows) on the gill filament of a grouper. Scanning electron microscopy.

The life cycle of Pseudorhabdosynochus species is similar to that of other diplectanid monogeneans.[9] It is monoxenous, i.e. there is a single host, the fish. The life-cycle includes a parasitic phase, the adult worm, and two stages which are free in the environment, the egg and the larva. The adult hermaphroditic worm, which lives on the gills of fish, lays eggs in the water. The egg has an operculum. A larva develops in the egg and hatches through the operculum; the swimming larva is ciliated and called an oncomiracidium. The larva finds its host by swimming to it, first attaches to the skin and migrates to the gills, and eventually transforms itself into a young monogenean which is attached to the gill filament by its haptor. During transformation from larva to adult, the monogenean loses its cilia and produces a new tegument, a phenomenon which is considered a general characteristic of the parasitic Platyhelminthes or Neodermata.[14]

Information is available for the life-cycle of Pseudorhabdosynochus lantauensis, a parasite of the orange-spotted grouper Epinephelus coioides, at a temperature of 30 °C.[15] The adult lays 10-22 eggs/day. The eggs hatch within 2–4 days. The oncomiracidium is viable for 4-8 h and should find a host within this delay. After attachment, the oncomiracidium transforms into an adult in 4–7 days. The parasite produces eggs after 7 days. The life-cycle is thus complete in 13–20 days (at 30 °C).[15]

Certain species of Pseudorhabdosynochus parasitize groupers which show spawning aggregations, i.e. the adult fish gather annually at a precise location and meet other fish of their own species only once per year. It has been suggested that transmission of Pseudorhabdosynochus species occurs preferentially during these spawning aggregations, from older infected fish to younger, uninfected, fish.[16]

Pathology and control in mariculture

Species of Pseudorhabdosynochus are attached to the gills of groupers. No information is available on the pathogen effect of these parasites in wild groupers. Since the life-cycle is monoxenous and thus can be completed with a high rate of success in mariculture condition, species of Pseudorhabdosynochus can proliferate on gills of cultured groupers and become dangerous for the fish, inducing mortality.

Parasite infections are often measured by two variables, prevalence (the proportion of infected hosts) and intensity (the number of parasites per host). Prevalences as high as 100% (i.e. all fish are infected) have been reported for Pseudorhabdosynochus sp. in cage-cultured Epinephelus malabaricus in Thailand,[17] for Pseudorhabdosynochus sp. on Epinephelus fuscoguttatus[18] and Epinephelus coioides[19] maricultured in Indonesia, and for P. beverleyburtonae on Epinephelus marginatus, wild or maricultured, in Brazil.[20] Intensities are as high as up to 1,006 individual Pseudorhabdosynochus sp. per fish (Epinephelus fuscoguttatus) in Indonesia,[18] up to 294 P. coioidesis per fish (Epinephelus coioides) in South China,[21] 327 P. yucatanensis per fish on Epinephelus morio in Mexico,[22] and 500 P. epinepheli per 100 grams of gill filaments of Epinephelus akaara in Japan.[23]

Eradication of gill parasitic monogeneans is a problem because various chemical treatments can be harmful for the fish. Treatments used against Pseudorhabdosynochus species include freshwater, diluted formalin, and diluted hydrogen peroxide (H2O2).[23] Diluted hydrogen peroxide (H2O2) seems the preferred treatment for P. epinepheli on Epinephelus akaara[23] and P. lantauensis on Epinephelus coioides.[citation needed]

List of species

Lists of species have been provided by Kritsky & Beverley-Burton (1986)[2] and Justine (2007). [11] The list below (about 80 species) is based on the World Register of Marine Organisms[24] with additions.

References

  1. ^ a b Yamaguti, S. 1958: Studies on the helminth fauna of Japan. Part 53. Trematodes of fishes, XII. Publications of the Seto Marine Biological Laboratory, 7, 53-88. Article PDF open access
  2. ^ a b c d e f g h i j k l m n o p q Kritsky, D. C.; Beverley-Burton, M. (1986). "The status of Pseudorhabdosynochus Yamaguti, 1958, and Cycloplectanum Oliver, 1968 (Monogenea: Diplectanidae)". Proceedings of the Biological Society of Washington. 99: 17–20. open access
  3. ^ a b c Justine, Jean-Lou (2009). "A redescription of Pseudorhabdosynochus epinepheli (Yamaguti, 1938), the type-species of Pseudorhabdosynochus Yamaguti, 1958 (Monogenea: Diplectanidae), and the description of P. satyui n. sp. from Epinephelus akaara off Japan". Systematic Parasitology. 72 (1): 27–55. doi:10.1007/s11230-008-9171-5. PMID 19048406. S2CID 9467410.
  4. ^ Kritsky, D. C., Boeger, W. A. & Robaldo, R. B. 2001: Neotropical Monogenoidea. 38. Revision of Rhabdosynochus Mizelle and Blatz, 1941 (Polyonchoinea: Dactylogyridea: Diplectanidae), with descriptions of two new species from Brazil. Comparative Parasitology, 68, 66-75.
  5. ^ a b Yamaguti, S (1938). "Studies on the helminth fauna of Japan. Part 24. Trematodes of fishes, V". Japanese Journal of Zoology. 8: 15–74.
  6. ^ a b c d e f g Mendoza-Franco, Edgar F.; Violante-González, Juan; Rojas Herrera, Agustín A. (2011). "Six new and one previously described species of Pseudorhabdosynochus (Monogenoidea, Diplectanidae) infecting the gills of groupers (Perciformes, Serranidae) from the Pacific coasts of Mexico and Panama". Journal of Parasitology. 97 (1): 20–35. doi:10.1645/GE-2716.1. ISSN 0022-3395. PMID 21348602. S2CID 207251069.
  7. ^ a b Knoff, Marcelo; Cohen, Simone Chinicz; Cárdenas, Melissa Querido; Cárdenas-Callirgos, Jorge M.; Gomes, Delir Corrêa (2015). "A new species of diplectanid (Monogenoidea) from Paranthias colonus (Perciformes, Serranidae) off Peru". Parasite. 22: 11. doi:10.1051/parasite/2015011. PMC 4353888. PMID 25754099.open access
  8. ^ a b Bravo Hollis, Margarita (1954). "Diplectanum amplidiscatum n. sp. Trématode monogénétique des branchies d'un poisson marin". Annales de Parasitologie Humaine et Comparée. 29 (1–2): 37–41. doi:10.1051/parasite/1954291037. open access
  9. ^ a b Oliver, Guy (2015). Les Diplectanidae Bychowsky, 1957 (Monogenea, Monopisthocotylea, Dactylogyridea). Systématique. Biologie. Ontogénie. Écologie. Essai de phylogenèse (thesis). figshare. doi:10.6084/M9.FIGSHARE.1295274.
  10. ^ Schoelinck, Charlotte; Cruaud, Corinne; Justine, Jean-Lou (2012). "Are all species of Pseudorhabdosynochus strictly host specific? — A molecular study" (PDF). Parasitology International. 61 (2): 356–359. doi:10.1016/j.parint.2012.01.009. ISSN 1383-5769. PMID 22326703.
  11. ^ a b c d e f g h i j Justine, Jean-Lou (2007). "Parasite biodiversity in a coral reef fish: twelve species of monogeneans on the gills of the grouper Epinephelus maculatus (Perciformes: Serranidae) off New Caledonia, with a description of eight new species of Pseudorhabdosynochus (Monogenea: Diplectanidae)". Systematic Parasitology. 66 (2): 81–129. doi:10.1007/s11230-006-9057-3. PMID 16972153. S2CID 7737882.
  12. ^ a b c d e f g Justine, J.-L.; Sigura, A. (2007). "Monogeneans of the malabar grouper Epinephelus malabaricus (Perciformes, Serranidae) off New Caledonia, with a description of six new species of Pseudorhabdosynochus (Monogenea: Diplectanidae)" (PDF). Zootaxa. 1543: 1–44. doi:10.11646/zootaxa.1543.1.1. Article (Open Archive)
  13. ^ Justine, Jean-Lou; Beveridge, Ian; Boxshall, Geoff A.; Bray, Rod A.; Moravec, Frantisek; Trilles, Jean-Paul; Whittington, Ian D. (Nov 2010). "An annotated list of parasites (Isopoda, Copepoda, Monogenea, Digenea, Cestoda and Nematoda) collected in groupers (Serranidae, Epinephelinae) in New Caledonia emphasizes parasite biodiversity in coral reef fish" (PDF). Folia Parasitologica. 57 (4): 237–62. doi:10.14411/fp.2010.032. PMID 21344838. open access
  14. ^ Ehlers, U. (1985). Das Phylogenetische System der Plathelminthes. Stuttgart: G. Fischer.
  15. ^ a b Erazo-Pagador, G.; Cruz-Lacierda, E. R. (2010). "The morphology and life cycle of the gill monogenean (Pseudorhabdosynochus lantauensis) on orange-spotted grouper (Epinephelus coioides) cultured in the Philippines". Bulletin of the European Association of Fish Pathologists. 30: 55–64.
  16. ^ a b c d e Sigura, A.; Justine, J.-L. (2008). "Monogeneans of the speckled blue grouper, Epinephelus cyanopodus (Perciformes, Serranidae), from off New Caledonia, with a description of four new species of Pseudorhabdosynochus and one new species of Laticola (Monogenea: Diplectanidae), and evidence of monogenean faunal changes according to the size of fish". Zootaxa. 1695: 1–44. doi:10.11646/zootaxa.1695.1.1.
  17. ^ Ruangpan, L. & Tubkaew, R. (1993). Parasites of the cage cultured grouper Epinephelus malabaricus in Thailand. Paper presented at the Proceedings of Grouper Culture, Kaoseng, Songkhla, Thailand.
  18. ^ a b Palm, H. W.; Kleinertz, S.; Rückert, S. (2011). "Parasite diversity as an indicator of environmental change? An example from tropical grouper (Epinephelus fuscoguttatus) mariculture in Indonesia". Parasitology. 138 (13): 1793–1803. doi:10.1017/S0031182011000011. ISSN 0031-1820. PMID 21320385. S2CID 2413697.
  19. ^ Rückert, Sonja; Klimpel, Sven; Al-Quraishy, Saleh; Mehlhorn, Heinz; Palm, Harry W. (2008). "Transmission of fish parasites into grouper mariculture (Serranidae: Epinephelus coioides (Hamilton, 1822)) in Lampung Bay, Indonesia". Parasitology Research. 104 (3): 523–532. doi:10.1007/s00436-008-1226-7. ISSN 0932-0113. PMID 18855012. S2CID 15956362.
  20. ^ Roumbedakis, K; Marchiori, NC; Paseto, Á; Gonçalves, ELT; Luque, JL; Cepeda, PB; Sanches, EG; Martins, ML (2013). "Parasite fauna of wild and cultured dusky-grouper Epinephelus marginatus (Lowe, 1834) from Ubatuba, Southeastern Brazil". Brazilian Journal of Biology. 73 (4): 871–878. doi:10.1590/S1519-69842013000400025. ISSN 1519-6984. PMID 24789405. open access
  21. ^ Luo, Y.; Yang, T. (2010). "Seasonal population dynamics of the monogeneans Pseudorhabdosynochus coioidesis and P. serrani on wild versus cultured groupers in Daya Bay, South China Sea". Helminthologia. 47 (4): 241–250. doi:10.2478/s11687-010-0038-3. ISSN 1336-9083.
  22. ^ a b Vidal-Martínez, V. M., Aguirre-Macedo, L. & Mendoza-Franco, E. F. 1997: Pseudorhabdosynochus yucatanensis sp. n. (Monogenea: Diplectanidae) from the gills of the red grouper Epinephelus morio (Pisces: Serranidae) of the Yucatan Peninsula, Mexico" Folia Parasitologica 44, 274-278. PDF open access
  23. ^ a b c Isshiki, Tadashi; Nagano, Taizou; Miki, Katsuhiro (2007). "Occurrence of a monogenean gill parasite Pseudorhabdosynochus epinepheli on Red Spotted Grouper Epinephelus akaara and its experimental treatment by Hydrogen Peroxide bathing". Fish Pathology. 42 (1): 71–74. doi:10.3147/jsfp.42.71. ISSN 0388-788X. open access
  24. ^ World Register of Marine Organisms
  25. ^ Price, E. W. (1937). "North American Monogenetic Trematodes. I. The superfamily Gyrodactyloidea". Journal of the Washington Academy of Sciences. 27: 146–164. open access
  26. ^ a b Justine, Jean-Lou (2007). "Pseudorhabdosynochus argus n. sp. (Monogenea: Diplectanidae) from Cephalopholis argus, P. minutus n. sp. and Diplectanum nanus n. sp. from C. sonnerati and other monogeneans from Cephalopholis spp. (Perciformes: Serranidae) off Australia and New Caledonia". Systematic Parasitology. 68 (3): 195–215. doi:10.1007/s11230-007-9096-4. ISSN 0165-5752. PMID 17896188. S2CID 23506802.
  27. ^ Sigura, Aude; Chauvet, Claude; Justine, Jean-Lou (2007). "Pseudorhabdosynochus bacchus sp. nov. (Monogenea, Diplectanidae) from Epinephelus coeruleopunctatus (Perciformes, Serranidae) off New Caledonia". Acta Parasitologica. 52 (3). doi:10.2478/s11686-007-0028-x. ISSN 1896-1851. S2CID 21628540.
  28. ^ a b Oliver, G. (1984). "Description de deux nouvelles espèces du genre Cycloplectanum Oliver, 1968". Annales de Parasitologie Humaine et Comparée. 59 (1): 31–39. doi:10.1051/parasite/1984591031. ISSN 0003-4150. open access
  29. ^ a b c d Chaabane, Amira; Neifar, Lassad; Gey, Delphine; Justine, Jean-Lou (2016). "Species of Pseudorhabdosynochus (Monogenea, Diplectanidae) from groupers (Mycteroperca spp., Epinephelidae) in the Mediterranean and Eastern Atlantic Ocean, with special reference to the 'beverleyburtonae group' and description of two new species". PLOS ONE. 11 (8): e0159886. Bibcode:2016PLoSO..1159886C. doi:10.1371/journal.pone.0159886. ISSN 1932-6203. PMC 4988817. PMID 27532108. open access
  30. ^ a b Oliver, G. & Paperna, I. 1984: Diplectanidae Bychowsky, 1957 (Monogenea, Monopisthocotylea), parasites de Perciformes de Méditerranée orientale, de la mer Rouge et de l'océan Indien. Bulletin du Muséum National d'Histoire Naturelle, Paris, 4° série, 6, section A, 49-65.
  31. ^ a b c d e Neifar, Lassad; Euzet, Louis (2007). "Five new species of Pseudorhabdosynochus (Monogenea: Diplectanidae) from the gills of Epinephelus costae (Teleostei: Serranidae)" (PDF). Folia Parasitologica. 54 (2): 117–128. doi:10.14411/fp.2007.017. ISSN 0015-5683. PMID 17886741. open access
  32. ^ a b c d e Chaabane, Amira; Neifar, Lassad; Justine, Jean-Lou (2017). "Diplectanids from Mycteroperca spp. (Epinephelidae) in the Mediterranean Sea: Redescriptions of six species from material collected off Tunisia and Libya, proposal for the "Pseudorhabdosynochus riouxi group", and a taxonomic key". PLOS ONE. 12 (2): e0171392. Bibcode:2017PLoSO..1271392C. doi:10.1371/journal.pone.0171392. ISSN 1932-6203. PMC 5289622. PMID 28152034. open access
  33. ^ a b c d e f g h i j k l m Kritsky, Delane C.; Bakenhaster, Micah D.; Adams, Douglas H. (2015). "Pseudorhabdosynochus species (Monogenoidea, Diplectanidae) parasitizing groupers (Serranidae, Epinephelinae, Epinephelini) in the western Atlantic Ocean and adjacent waters, with descriptions of 13 new species". Parasite. 22: 24. doi:10.1051/parasite/2015024. ISSN 1776-1042. PMC 4536336. PMID 26272242. open access
  34. ^ a b Hinsinger, Damien D.; Justine, Jean-Lou (2006). "The 'Pseudorhabdosynochus cupatus group' (Monogenea: Diplectanidae) on Epinephelus fasciatus, E. howlandi, E. rivulatus and E. merra (Perciformes: Serranidae) off New Caledonia, with descriptions of Pseudorhabdosynochus cyathus n. sp. and P. calathus n. sp". Systematic Parasitology. 64 (2): 69–90. doi:10.1007/s11230-005-9018-2. ISSN 0165-5752. PMID 16773473. S2CID 26881495.
  35. ^ Justine, Jean-Lou (2005). "Species of Pseudorhabdosynochus Yamaguti, 1958 (Monogenea: Diplectanidae) from Epinephelus fasciatus and E. merra (Perciformes: Serranidae) off New Caledonia and other parts of the Indo-Pacific Ocean, with a comparison of measurements of specimens prepared using different methods, and a description of P. caledonicus n. sp". Systematic Parasitology. 62 (1): 1–37. doi:10.1007/s11230-005-5480-0. ISSN 0165-5752. PMID 16132868. S2CID 35119181.
  36. ^ Vidal-Martínez, V. M. & Mendoza-Franco, E. F. 1998: Pseudorhabdosynochus capurroi sp. n. (Monogenea: Diplectanidae) from the gills of Mycteroperca bonaci (Pisces: Serranidae) of the Yucatan Peninsula, Mexico" Folia Parasitologica 45, 221-224. Article PDF open access
  37. ^ Zhang, J. Y., Yang, T. B. & Liu, L. (2001). [Monogeneans of Chinese marine fishes] (in Chinese). Beijing: Agriculture Press.
  38. ^ Bu, S. S. H.; Leong, T. S.; Wong, S. Y.; Woo, Y. S. N.; Foo, R. W. T. (1999). "Three diplectanid monogeneans from marine finfish (Epinephelus spp.) in the Far East". Journal of Helminthology. 73 (4): 301–312. doi:10.1017/S0022149X99000505.
  39. ^ a b c Schoelinck, Charlotte; Justine, Jean-Lou (2011). "Four species of Pseudorhabdosynochus (Monogenea: Diplectanidae) from the camouflage grouper Epinephelus polyphekadion (Perciformes: Serranidae) off New Caledonia". Systematic Parasitology. 79 (1): 41–61. doi:10.1007/s11230-010-9289-0. ISSN 0165-5752. PMID 21487947. S2CID 23523617.
  40. ^ a b c Young, P. C. (1969). "Some Monogenoideans of the Family Diplectanidae Bychowsky, 1957 from Australian Teleost Fishes". Journal of Helminthology. 43 (1–2): 223–254. doi:10.1017/S0022149X00004053. ISSN 0022-149X. PMID 5381127.
  41. ^ a b Justine, Jean-Lou; Henry, Émilie (2010). "Monogeneans from Epinephelus chlorostigma (Val.) (Perciformes: Serranidae) off New Caledonia, with the description of three new species of diplectanids". Systematic Parasitology. 77 (2): 81–105. doi:10.1007/s11230-010-9263-x. ISSN 0165-5752. PMID 20852982. S2CID 36351344.
  42. ^ a b c d Santos, Cláudia Portes; Buchmann, Kurt; Gibson, David I. (2000). "Pseudorhabdosynochus spp. (Monogenea: Diplectanidae) from the gills of Epinephelus spp. in Brazilian waters". Systematic Parasitology. 45 (2): 145–153. doi:10.1023/A:1006232029426. ISSN 0165-5752. PMID 10743859. S2CID 26949491.
  43. ^ Justine, Jean-Lou (2005). "Pseudorhabdosynochus hirundineus n. sp. (Monogenea: Diplectanidae) from Variola louti (Perciformes: Serranidae) off New Caledonia". Systematic Parasitology. 62 (1): 39–45. doi:10.1007/s11230-005-5481-z. ISSN 0165-5752. PMID 16132869. S2CID 2967245.
  44. ^ Justine, Jean-Lou (2008). "Pseudorhabdosynochus inversus sp. nov. (Monogenea, Diplectanidae) from the halfmoon grouper Epinephelus rivulatus (Perciformes, Serranidae) off New Caledonia". Acta Parasitologica. 53 (4). doi:10.2478/s11686-008-0057-0. ISSN 1896-1851.
  45. ^ Zeng, Bijian; Yang, Tingbao (2006). "Description of Pseudorhabdosynochus justinei n. sp. (Monogenea: Diplectanidae) and redescription of P. vagampullum (Young, 1969) Kritsky & Beverley-Burton, 1986 from the gills of the longfin grouper Epinephelus quoyanus (Valenciennes) (Perciformes: Serranidae) in Dapeng Bay, South China Sea". Systematic Parasitology. 66 (3): 223–235. doi:10.1007/s11230-006-9067-1. ISSN 0165-5752. PMID 17143574. S2CID 29694108.
  46. ^ a b Justine, Jean-Lou; Dupoux, Cyndie; Cribb, Thomas (2009). "Resolution of the discrepant host-specificity of Pseudorhabdosynochus species (Monogenea, Diplectanidae) from serranid fishes in the tropical Indo-Pacific". Acta Parasitologica. 54 (2). doi:10.2478/s11686-009-0027-1. ISSN 1896-1851.
  47. ^ Saengpheng, Chompunooch; Purivirojkul, Watchariya (2020-01-07). "Pseudorhabdosynochus kasetsartensis n. sp. (Monogenea: Diplectanidae) from the cloudy grouper Epinephelus erythrurus (Valenciennes) (Perciformes: Serranidae) in the lower Gulf of Thailand". Systematic Parasitology. 97 (1): 99–106. doi:10.1007/s11230-019-09899-z. ISSN 0165-5752.
  48. ^ Dyer, William G.; Williams, Ernest H.; Bunkley-Williams, Lucy (1995). "Pseudorhabdosynochus kritskyi n. sp. (Monogenea: Diplectanidae) on Gag from the Gulf of Mexico". Journal of Aquatic Animal Health. 7 (4): 337–340. doi:10.1577/1548-8667(1995)007<0337:KNSDOG>2.3.CO;2. ISSN 0899-7659.
  49. ^ Beverley-Burton, Mary; Suriano, Delia Mabel (1981). "A revision of Cycloplectanum Oliver, 1968 (Monogenea: Diplectanidae) and descriptions of C.hongkongensis n.sp. and C.lantauensis n.sp. from Epinephelus spp. (Serranidae) in the South China Sea". Canadian Journal of Zoology. 59 (7): 1276–1285. doi:10.1139/z81-180. ISSN 0008-4301.
  50. ^ Aljoshkina, L. D. (1984). "[New species of the family Diplectanidae (Monogenea) in the South-West Atlantic]". Zoologicheskii Zhurnal. 63: 1253–1256.
  51. ^ Laird, Marshall (1958). "Parasites of South Pacific fishes II. Diplectanum melanesiensis n. sp., a monogenetic trematode from Fiji and the New Hebrides". Canadian Journal of Zoology. 36 (2): 167–173. doi:10.1139/z58-017. ISSN 0008-4301.
  52. ^ Dyer, William G.; Williams, Ernest H.; Bunkley-Williams, Lucy (1994). "Pseudorhabdosynochus monaensis n. sp. (Monogenea: Diplectanidae) on Rock Hind from Mona Island, Puerto Rico". Journal of Aquatic Animal Health. 6 (1): 59–63. doi:10.1577/1548-8667(1994)006<0059:PMNSMD>2.3.CO;2. ISSN 0899-7659.
  53. ^ a b Justine, Jean-Lou (2008). "Two new species of Pseudorhabdosynochus Yamaguti, 1958 (Monogenea: Diplectanidae) from the deep-sea grouper Epinephelus morrhua (Val.) (Perciformes: Serranidae) off New Caledonia". Systematic Parasitology. 71 (2): 145–158. doi:10.1007/s11230-008-9156-4. ISSN 0165-5752. PMID 18716902. S2CID 23245863.
  54. ^ Justine, Jean-Lou; Vignon, Matthias (2008). "Monogeneans of the grouper Epinephelus tauvina (Perciformes, Serranidae) off Moorea, French Polynesia, with a description of Pseudorhabdosynochus pai n. sp. (Monogenea: Diplectanidae)". Systematic Parasitology. 72 (2): 113–125. doi:10.1007/s11230-008-9159-1. ISSN 0165-5752. PMID 19115085. S2CID 7679605.
  55. ^ Schoelinck, Charlotte; Justine, Jean-Lou (2011). "Pseudorhabdosynochus quadratus n. sp. (Monogenea: Diplectanidae) from the white-streaked grouper Epinephelus ongus (Bloch) (Perciformes: Serranidae) off New Caledonia". Systematic Parasitology. 79 (1): 77–80. doi:10.1007/s11230-011-9295-x. ISSN 0165-5752. PMID 21487950. S2CID 22318648.
  56. ^ Yamaguti, S. (1968). Monogenetic Trematodes of Hawaiian Fishes. Honolulu: University of Hawaii Press.
  57. ^ Chaabane, Amira; Neifar, Lassad; Justine, Jean-Lou (2015). "Pseudorhabdosynochus regius n. sp. (Monogenea, Diplectanidae) from the mottled grouper Mycteroperca rubra (Teleostei) in the Mediterranean Sea and Eastern Atlantic". Parasite. 22: 9. doi:10.1051/parasite/2015005. ISSN 1776-1042. PMC 4325681. PMID 25674913.open access
  58. ^ Oliver, Guy (1986). "Cycloplectanum riouxi n. sp., une nouvelle espèce de Diplectanidae (Monogenea, Monopisthocotylea) parasite d'Epinephelus guaza (Linnaeus, 1758) (Pisces, Serranidae)". Systematic Parasitology. 8 (4): 317–322. doi:10.1007/BF00009739. ISSN 0165-5752. S2CID 39530925.
  59. ^ Yamaguti, S. 1953: Parasitic worms mainly from Celebes. Part 2. Monogenetic trematodes of fishes. Acta Medicinae Okayama, 8, 203-256 + 209 Pl. Article Archived 2016-03-04 at the Wayback Machine PDF Archived 2015-02-09 at the Wayback Machine open access
  60. ^ Yang, Tingbao; Zeng, Bijian; Gibson, David I. (2005). "Description of Pseudorhabdosynochus shenzhenensis n. sp. (Monogenea: Diplectanidae) and redescription of P. serrani Yamaguti, 1953 from Epinephelus coioides off Dapeng Bay, Shenzhen, China". Journal of Parasitology. 91 (4): 808–813. doi:10.1645/GE-518R.1. ISSN 0022-3395. PMID 17089747. S2CID 19512689.
  61. ^ Chaabane, Amira; Justine, Jean-Lou; Gey, Delphine; Bakenhaster, Micah D.; Neifar, Lassad (2016). "Pseudorhabdosynochus sulamericanus (Monogenea, Diplectanidae), a parasite of deep-sea groupers (Serranidae) occurs transatlantically on three congeneric hosts (Hyporthodus spp.), one from the Mediterranean Sea and two from the western Atlantic". PeerJ. 4: e2233. doi:10.7717/peerj.2233. ISSN 2167-8359. PMC 4991870. PMID 27602259. open access
  62. ^ Yang, Tingbao; Gibson, David I.; Zeng, Bijian (2005). "Pseudorhabdosynochus summanoides n. sp. (Monogenea: Diplectanidae) from Epinephelus coioides in Dapeng Bay, South China Sea, with observations on several similar species of Pseudorhabdosynochus Yamaguti, 1958". Systematic Parasitology. 62 (3): 221–239. doi:10.1007/s11230-005-5497-4. ISSN 0165-5752. PMID 16315082. S2CID 7907556.
  63. ^ Hinsinger, Damien D.; Justine, Jean-Lou (2006). "Pseudorhabdosynochus venus n. sp. (Monogenea: Diplectanidae) from Epinephelus howlandi (Perciformes: Serranidae) off New Caledonia". Systematic Parasitology. 63 (2): 153–158. doi:10.1007/s11230-005-9012-8. ISSN 0165-5752. PMID 16718595. S2CID 23278926.