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Enigmas from the past: M’Intosh’s (1885) annelidicolous copepods from the voyage of H.M.S. Challenger



The type material of four monotypic genera, Leaniricola, Oestrella, Praxillinicola and Trophoniphila (Copepoda, Cyclopoida), described by M’Intosh (1885) from deep water polychaete hosts collected during the H.M.S. Challenger expedition, is re-examined. Leaniricola rotundata M’Intosh (1885) is removed from its floating status as species inquirenda in the Nereicolidae and fixed as the type of a new family, Leaniricolidae fam. nov., based on the presence of an oral cone and massive, three-dimensionally expanded, mandibular gnathobases which are used to anchor the parasite in the parapodial integument of its host. The ectoparasitic Praxillinicola kroyeri (M’Intosh, 1885), previously treated as a species inquirenda in the Clausiidae, cannot be placed in any of the currently recognized poecilostome families and is here fixed as the type of a new family, Praxillinicolidae fam. nov. Females are characterized by unique bilobate antennules, contained within anterior sockets shared with the reduced antennae, and by paired labral hooks, both of which probably serve as auxiliary attachment organs. Despite its highly transformed body P. kroyeri has retained the plesiomorphic condition of the female genital system, with the gonopores positioned dorsolaterally and the paired copulatory pores lying close together on the midventral surface. The position of the mesoparasitic Trophoniphila bradyi M’Intosh, 1885 in the Bradophilidae is confirmed based on the presence of a median copulatory pore in the female, the disproportionately large egg sacs, the shape of the ectosoma and host utilization. Both sexes of Bradophila pygmaea Levinsen, 1878 are re-examined and the male is redescribed in detail. Flabellicola Gravier, 1918a is treated as a genus incertae sedis in the Bradophilidae. The anal prominence, previously reported in the female of Oestrella (= Melinnacheres) levinseni M’Intosh, 1885 (Saccopsidae), proved upon re-examination to be one of the paired genital apertures. The third and fourth pairs of appendages reported in male saccopsids are here interpreted as the maxillae and maxillipeds, respectively.



Crustacea, Bradophila, Flabellicola, Leaniricola, Oestrella, Praxillinicola, Trophoniphila, Leaniricolidae fam. nov., Praxillinicolidae fam. nov., Cyclopoida

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Bocquet, C. & Stock, J.H. (1956) Travaux de la Station biologique de Roscoff, XXXV. Copépodes parasites d’invertébrés des côtes de la Manche. I. Endocheres obscurus, nov. gen., n. sp., parasite de Calliostoma zizyphinum. Archives de Zoologie expérimentale et générale, 93 (notes et revue 3), 113–122.

Bocquet, C., Stock, J.H. & Kleeton, G. (1963) Copépodes parasites d’invertébrés des côtes de la Manche. X. Cyclopoïdes Poecilostomes associés aux Annélides Polychètes, dans la région de Roscoff. Archives de Zoologie expérimentale et générale, 102 (notes et revue 1), 20–40.

Boxshall, G.A. (1977) On the synonymy of the genera Oestrella McIntosh, 1885, and Melinnacheres M. Sars, 1870. Crustaceana, 32, 214–215.

Boxshall, G.A. & Halsey, H.S. (2004) An Introduction to Copepod Diversity. The Ray Society, London, xv + 966 pp.

Boxshall, G.A. & Huys, R. (1998) The ontogeny and phylogeny of copepod antennules. Proceedings of the Royal Society of London, Biological Sciences, 353, 765–786.

Boxshall, G.A., O’Reilly, M., Sikorski, A. & Summerfield, R. (2015) A new genus and family of copepods (Crustacea: Copepoda) parasitic on polychaetes of the genus Jasmineira Langerhans, 1880 (family Sabellidae) in the northeastern Atlantic. Zootaxa, 4018 (3), 426–436.

Bresciani, J. (1964) Sigecheres brittae gen. et sp. nov., a parasitic copepod from the polychaete Sige fusigera Malmgren. Ophelia, 1, 295–301.

Bresciani, J. (1967) Redescription du mâle de Selioides bolbroei Levinsen avec une note sur la répartition géographique du genre Selioides (Copepoda Cyclopoida). Crustaceana, 13, 220–226.

Bresciani, J. & Lützen, J. (1961) The anatomy of a parasitic copepod, Saccopsis steenstrupi n. sp. Crustaceana, 3, 9–23.

Bresciani, J. & Lützen, J. (1966) The anatomy of Aphanodomus terebellae (Levinsen) with remarks on the sexuality of the family Xenocoelomidae nov. fam. (Parasitic Copepoda). Bulletin de la Muséum national d’Histoire naturelle, Paris, Série 2, 37, 787–806.

Bresciani, J. & Lützen, J. (1974) On the biology and development of Aphanodomus Wilson (Xenocoelomidae), a parasitic copepod of the polychaete Thelepus cincinnatus. Videnskabelige Meddelelser fra dansk Naturhistorisk Forening i Kjøbenhavn, 137, 25–63.

Bresciani, J. & Lützen, J. (1975) Melinnacheres ergasiloides M. Sars, a parasitic copepod of the polychaete Melinna cristata, with notes on multiple infections caused by annelidicolous copepods. Ophelia, 13, 31–42.

Brumpt, E. (1897) Sur un Copépode nouveau (Saccopsis Alleni, nova species, parasite de Polycirrus aurantiacus Grube). Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, Paris, 124, 1464–1467.

Caullery, M. & Mesnil, F. (1919) Xenocoeloma brumpti C. & M., Copépode parasite de Polycirrus arenivorum C. Bulletin biologique de la France et de la Belgique, 53, 161–233.

Carton, Y. (1965) Description de Selioides bocqueti n. sp., Copépode Cyclopoïde parasite de Scalisetosus assimilis MacIntosh, Aphroditidae commensal d’Echinus esculentus L. Archives de Zoologie expérimentale et générale, 104, notes et revue, 83–103.

Carton, Y. & Laubier, L. (1974) Description de Selioides guineensis sp. n., Copépode Cyclopoïde parasite d’Aphroditidae. Archives de Zoologie expérimentale et générale, 115, 129–139.

Cheng, Y.-r., Ho, J.-s. & Dai, C.-F. (2014) Description of Indoclausia helioporae sp. nov. (Copepoda, Clausiidae) from blue coral, associated with a number of polychaetes. Crustaceana, 87, 627–633.

Conradi, M., Bandera, M.E., Marin, I. & Martin, D. (2015) Polychaete-parasitizing copepods from the deep-sea Kuril–Kamchatka Trench (Pacific Ocean), with the description of a new Ophelicola species and comments on the currently known annelidicolous copepods. Deep-Sea Research Part II: Topical Studies in Oceanography, 111, 147–165.

Delamare-Deboutteville, C. & Laubier, L. (1960a) Les Phyllocolidae, une famille nouvelle de Copépodes parasites d’Annélides Polychètes. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, Paris, 251, 2083–2085.

Delamare-Deboutteville, C. & Laubier, L. (1960b) Les Phyllocolidae Delamare et Laubier, Copépodes parasites d’Annélides Polychètes Phyllodocides, et leur rapports avec les Copépodes annélidicoles. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, Paris, 251, 2231–2233.

Ehlers, E. (1874) Annulata nova vel minus cognita in Expeditione ‘Porcupine’ capta. Annals and Magazine of natural History, Series 4, 13, 292–298.

Filippova, A.V., Tzetlin, A.B. & Purschke, G. (2003) Morphology and ultrastructure of the anterior end of Diplocirrus longisetosus Marenzeller, 1890 (Flabelligeridae, Polychaeta, Annelida). In: Sigvaldadóttir, E., Mackie, A.S.Y., Helgason, G.V., Reish, D.J., Svavarsson, J., Steingrímsson, S.A. & Guðmundsson, G. (Eds.), Advances in Polychaete Research. Hydrobiologia, 496, 215–223.

Giesbrecht, W. (1895) Mitteilungen über Copepoden. 10 und 11. 10. Über Miracia minor Th. Scott. 11. Über Seridium rugosum, einen neuen Anneliden-Parasiten. Mittheilungen aus der Zoologischen Station zu Neapel, 12, 217–226, plate 9.

Gooding, R.U. (1963) External morphology and classification of marine poecilostome copepods belonging to the families Clausidiidae, Clausiidae, Nereicolidae, Eunicicolidae, Synaptiphilidae, Catiniidae, Anomopsyllidae, and Echiurophilidae. Ph.D. dissertation, University of Washington, Seattle, iv + 247 pp, 26 plates, 2 tables. [plates and tables unpaginated]

Gotto, R.V. (1979) The association of Copepoda with marine invertebrates. Advances in marine Biology, 16, 1–109.

Gotto, R.V. & Leahy, Y. (1988) A new annelidicolous copepod, Cyclorhiza megalova n. sp., with comments on its functional biology and possible phylogenetic relationships. In: Boxshall, G.A. & Schminke, H.K. (Eds.), Biology of Copepods. Proceedings of the Third International Conference on Copepoda. Hydrobiologia, 167/168, 533–538.

Gravier, C.J. (1918a) Sur un nouveau Copépode (Flabellicola n.g. neapolitana n. sp.) parasite d’un Annélide polychète [Flabelligera diplochaitos (Otto)]. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences, 166, 502–505. [published 25 March]

Gravier, C.J. (1918b) Contribution à l’étude d’un Copépode (Flabellicola neapolitana Gravier) parasite d’un Annélide Polychète [Flabelligera (Siphonostoma) diplochaitos (Otto)]. Pubblicazioni della Stazione zoologica di Napoli, 2, 209–222. [published 15 October]

Gunther, A.E. (1977) The life of William Carmichael M’Intosh, M.D., F.R.S. of St. Andrews, 1938–1931. A pioneer of marine biology. Scottish Academic Press for University of St. Andrews, Edinburgh, 214 pp.

Haddon, K. (1912) Herpyllobius arcticus. Quarterly Journal of microscopical Science, new series, 58, 385–410.

Hansen, H.J. (1892) Rhizorhina Ampeliscæ n. gen., n. sp. En ny til Herpyllobiidæ, n. fam., hørende Copepod, snyltende paa Amp. lævigata Lilljb. Entomologiske Meddelelser, 3, 207–234, plate III.

Hansen, H.J. (1897) The Choniostomatidæ, a Family of Copepoda, Parasites on Crustacea Malacostraca. A.F. Host & Son, Copenhagen, 205 pp., 13 plates.

Hansen, H.J. (1923) Crustacea Copepoda. II. Copepoda parasita and hemiparasita. Danish Ingolf Expedition, 3 (7), 1–92, plates 1–5.

Heegaard, P.E. (1942) Cyclorhiza eteonicola n. gen., n. sp., a new parasitic copepod. Det Kongelige norske Videnskabers Selskab Forhandlinger, 15 (14), 53–54.

Hesse, E. (1869) Observations sur les Crustacés rares ou nouveaux des côtes de France (dix-septième article). Annales des Sciences naturelles, Série Zoologie et Paléontologie ,Série 5, 11, 275–308, plate I.

Ho, J.-S. (1984) New family of poecilostomatoid copepods (Spiophanicolidae) parasitic on polychaetes from Southern California, with a phylogenetic analysis of nereicoliform families. Journal of crustacean Biology, 4, 134–146.

Humes, A.G. (1973) Tychidion guyanense n. gen., n. sp. (Copepoda, Cyclopoida) associated with an annelid off Guyana. Zoologische Mededelingen, Leiden, 46, 189–196.

Humes, A.G. & Gooding, R.U. (1964) A method for studying the external anatomy of copepods. Crustaceana, 6, 238–240.

Humes, A.G. & Huys, R. (1992) Copepoda (Poecilostomatoida and Siphonostomatoida) from deep-sea hydrothermal vent areas off British Columbia, including Amphicrossus altalis, a new species of Erebonasteridae, with notes on the taxonomic position of the genus Tychidion Humes. Canadian Journal of Zoology, 70, 1369–1380.

Huys, R. (1996) Superornatiremidae fam. nov. (Copepoda: Harpacticoida): An enigmatic family from North Atlantic anchihaline caves. Scientia marina, 60, 497–542.

Huys, R. (2001) Splanchnotrophid systematics: a case of polyphyly and taxonomic myopia. Journal of crustacean Biology, 21, 106–156.[0106:SSACOP]2.0.CO;2

Huys, R. & Boxshall, G.A. (1990) Discovery of Centobnaster humesi, new genus, new species (Erebonasteridae), the most primitive poecilostomatoid copepod known, in New Caledonian deep waters. Journal of crustacean Biology, 10, 504–519.

Huys, R. & Boxshall, G.A. (1991) Copepod Evolution. The Ray Society, London, 468 pp.

Huys, R. & Iliffe, T.M. (1998) Novocriniidae, a new family of harpacticoid copepods from anchihaline caves in Belize. Zoologica Scripta, 27, 1–15.

Keferstein, W. (1863) Ueber einen neuen Schmarotzerkrebs (Nereicola ovata Kef.) von einer Annelide. Zeitschrift für wissenschaftliche Zoologie, 12, 461–464, plate XLII.

Krøyer, H. (1837) Om Snyltekrebsene, isaer med hensyn til den Danske fauna. Naturhistorisk Tidsskrift, 1, 476–506, plate V.

Laubier, L. (1961) Phyllodicola petiti (Delamare et Laubier, 1960) et la famille des Phyllodicolidae, Copépodes parasites d’Annélides Polychètes en Méditerranée occidentale. Crustaceana, 2, 228–242.

Laubier, L. (1978) Ophelicola drachi gen. sp. n., un nouveau Copépode Cyclopoïde abyssal ectoparasite d’Annélides Polychètes Opheliidae. Archives de Zoologie expérimentale et générale, 119, 39–50.

Leigh-Sharpe, W.H. (1926) The Herpyllobiidae. A family of Copepoda parasitic on polynoid worms. Parasitology, 18, 269–276.

Levinsen, G.M.R. (1878) Om nogle parasitiske Krebsdyr, der snylte hos Annelider. Videnskabelige Meddelelser fra den naturhistoriske Forening i Kjöbenhavn, 1877–1878, 351–380, plate 6.

López-González, P.J., Bresciani, J. & Conradi, M. (2000) Two new species of Herpyllobius Steenstrup & Lütken, 1861 and a new record of Herpyllobius antarcticus Vanhöffen, 1913 (parasitic Copepoda) from the Weddell Sea, Antarctica. Polar Biology, 23, 265–271.

López-González, P.J., Bresciani, J. & Conradi, M. (2006) New genus, three new species and new records of Herpyllobiidae Hansen, 1892 (Crustacea, Copepoda), parasites of polychaetes from Antarctica. Scientia marina, 70, 243–259.

Lützen, J. (1964) A revision of the family Herpyllobiidae (parasitic copepods) with notes on hosts and distribution. Ophelia, 1, 241–274.

Lützen, J. (1966) The anatomy of the family Herpyllobiidae (parasitic copepods). Ophelia, 3, 45–64, plate 1.

M’Intosh, W.C. (1876) VIII. On the Annelida of the ‘Porcupine’ Expeditions of 1869 and 1870. Part I. Euphrosynidæ, Amphinomidæ, Aphroditidæ, Polynoidæ, Acoëtidæ, and Sigalionidæ. Transactions of the zoological Society of London, 9 (7), 395–416.

M’Intosh, W.C. (1885) Report on Annelida Polychaeta collected by H.M.S. Challenger during the years 1873–1876. Report of the scientific Results of the Voyage of H.M.S. Challenger 1873–76, Zoology, 12, i–xxxvi, 1–554, plates I–LV, IA–XXXIXA.

Malmgren, A.J. (1866) Nordiska Hafs-Annulater. Öfversigt af Königlich Vetenskaps-Akademiens Förhandlingar, 22 (5), 355–410, plates XVIII–XXIX.

Marchenkov, A.V. (1997) Fauna of symbiotic and parasitic crustaceans (Copepoda, Isopoda, Cirripedia, Amphipoda) from the benthic invertebrates of the White Sea and Arctic Seas. Parazitologija, 31, 24–44. [in Russian with English summary]

Marchenkov, A.V. (1998) Redescription of Bradophyla [sic] pigmaea [sic] Levinsen, 1887 [sic], a copepod parasitic on the Brada villosa (Rathke) from the White Sea: new signs and taxonomic perspectives. In: Abstracts Special International Conference New Methods in Copepod Taxonomy, Saint Petersburg, 04–08 May 1998, p. 15.

Marchenkov, A.V. (1999a) Symbiotic and parasitic Crustacea off benthic marine invertebrates from the Arctic Seas of Russia. Unpublished Ph.D. dissertation, Zoological Institute, Russian Academy of Sciences, Saint Petersburg, 193 pp. [in Russian]

Marchenkov, A.V. (1999b) Symbiotic and parasitic Crustacea of benthic marine invertebrates from the Arctic Seas of Russia. Synopsis of Ph.D. dissertation, Saint Petersburg, 23 pp.

Marchenkov, A.V. (2001) Peculiarities of parasitism in the Copepoda and Rhizocephala. Parazitologiya, 35, 89–97. [in Russian with English summary]

Marchenkov, A.V. (2002) Bradophilidae fam. nov.—the new family of mesoparasitic copepod collected from the polychaete Brada villosa from the White Sea. Parazitologiya, 36, 514–516. [in Russian with English summary]

Martínez Arbizu, P. (1997) The monophyly of Erebonasteridae, with the description of Centobnaster severnicus sp. n. (Copepoda: Poecilostomatoida) from the Laptev Sea (Arctic Ocean). Zoologischer Anzeiger, 235, 263–270.

Monod, T. & Laubier, L. (1996) Les Crustacés dans la Biosphère. Traité de Zoologie. Anatomie, Systématique, Biologie. Tome VII. Crustacés. Fascicule 2. Généralités (suite) et Systématique. Masson, Paris, 76 pp. [pp. 90–166]

Nordmann, A. von. (1864) Neue Beiträge zur Kenntnis parasitischer Copepoden. Bulletin de la Société des Naturalistes de Moscou, 37 (4), 461–520, plates V–VIII.

O’Reilly, M.G. (1995) A new genus of copepod (Copepoda: Poecilostomatoida) commensal with the maldanid polychaete Rhodine gracilior, with a review of the family Clausiidae. Journal of natural History, 29, 47–64.

O’Reilly, M.G. (2000) Notes on copepod parasites of phyllodocid polychaete worms in Scottish waters; including the first UK records of the Mediterranean copepod Phyllodicola petiti (Delamare-Deboutteville & Laubier, 1960). The Glasgow Naturalist, 23 (5), 39–44.

O’Reilly (2016) Parasitic and commensal polychaetes (Fams. Arabellidae and Sphaerodoridae) and copepods (Fam. Saccopsidae) associated with lamella-worms (Terebellides spp.) in Scottish, and nearby, waters. The Glasgow Naturalist, 26, 61–70.

Otto, A.G. (1821) Animalium maritimorum nondum editorum genera duo. Nova Acta physico-medica Academiae caesareae leopoldino-carolinae naturae curiosorum, 10, 617–634, plates L–LI.

Rafinesque, C.S. (1815) Analyse de la Nature ou Tableau de l’Univers et des Corps organisés. Imp. Jean Barravecchia, Palermo, 224 pp.

Rathke, H. (1843) Beiträge zur Fauna Norwegens. Novorum Actorum Academiae caesareae leopoldino-carolinae naturae curiosorum, 20, 1–264c, plates I–XII.

Rouse, G.W. (2001) A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera. Zoological Journal of the Linnean Society, 132, 55–80.

Sars, G.O. (1921) Copepoda Monstrilloida and Notodelphyoida. Parts V & VI. Botryllophilidae, Enterocolidae, Supplement. An Account of the Crustacea of Norway, with short Descriptions and Figures of all the Species, 8, 69–90, index, title, preface, plates XXXIII–XXXVII.

Sars, M. (1829) Bidrag til Söedyrenes Naturhistorie. Vol. 1. Chr. Dahl, Bergen, 60 pp., 6 plates.

Sars, M. (1870) VII. Bidrag til Kundskab om Christianiafjordens Fauna. II. Crustacea. Beskrivelse af nye, paa annelider snyltende Copepodeformer. Nyt Magazin for Naturvidenskaberne, 17 (2), 113–160, plates 8–10.

Southward, E.C. (1964) On three new cyclopoid copepods associated with deep-water polychaetes in the north-east Atlantic. Crustaceana, 6, 207–219.

Steenstrup, J.J.S. & Lütken, C.F. (1861) Bidrag til Kundskab om det aabne Havs Snyltekrebs og Lernæer samt om nogle andre nye eller hidtil kun ufuldstændigt kjendte parasitiske Copepoder. Det Kongelige Danske videnskabernes Selskabs Skrifter. Naturvidenskabelig og mathematisk Afdeling, Series 5, 5, 341–432, plates 1–15.

Stock, J.H. (1968) Vectoriella marinovi, un Copépode nouveau, parasite d’une Annélide Polychète pontique. Crustaceana, supplement 1, 186–192.

Stock, J.H. (1979) Serpulidicolidae, a new family of Copepoda associated with tubicolous polychaetes, with descriptions of a new genus and species from the Gulf of Mexico. Memoirs of the Hourglass Cruises, 5 (2), 1–11

Stock, J.H. (1982) Description de Cotylemyzon vervoorti gen. et sp. nov., un Copépode Cyclopoïde très original, parasite d’un polychète d’Amboine. Netherlands Journal of Zoology, 32, 364–373.

Stock, J.H. (1986) Cases of hyperassociation in the Copepoda (Herpyllobiidae and Nereicolidae). Systematic Parasitology, 8, 71–81.

Stock, J.H. (1988) Lamippidae (Copepoda: Siphonostomatoida) parasitic in Alcyonium. Journal of the marine biological Association of the United Kingdom, 68, 351–359.

Stock, J.H. (1996) Two new species of Copepoda parasitic on polynoid polychaetes. Crustaceana, 69, 438–445.

Tauber, P. (1879) Annulata Danica. En Kritisk Revision af de i Danmark Fundne Annulata Chaetognatha, Gephyrea, Balanoglossi, Discophoreae, Oligochaeta, Gymnocopa og Polychaeta. Reitzel, Kopenhagen, 143 pp.

Willey, A. (1905) Report on the Polychæta collected by Professor Herdman, at Ceylon, in 1902. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar by W.A. Herdman, with supplementary Reports upon the marine Biology of Ceylon, by other Naturalists, 4 (Supplementary Report 30), 243–324, plates I–VIII.

Wilson, C.B. (1923) A new genus and species of parasitic copepod from lower California. American Museum Novitates, 81, 1–4.

Wilson, C.B. (1932) The copepods of the Woods Hole region, Massachusetts. Bulletin of the United States national Museum, 158, 1–635.

Wilson, M.S. & Illg, P.L. (1955) The family Clausiidae (Copepoda, Cyclopoida). Proceedings of the biological Society of Washington, 68, 129–142.



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