Abstract
The Monniot collection of copepods associated with ascidian hosts was built up over several decades of field collecting and taxonomic research on ascidians by Drs Claude & Françoise Monniot (MNHN, Paris). This paper describes a total of 84 new species of copepods collected from ascidian hosts and five new genera are established. Prior to this study the family Ascidicolidae comprised two genera accommodating five valid species; here we add two new genera, Hamistyelicola gen. nov. and Bathycopola gen. nov., and eight new species in total. The family Buproridae comprised a single genus consisting of three species; here we add a new monotypic genus, Buprorides gen. nov. The family Botryllophilidae comprised 68 valid species in seven genera and here we add 45 new species; 13 of Botryllophilus Hesse, 1864, nine of Schizoproctus Aurivillius, 1885, three of Haplostomides Chatton & Harant, 1924, 12 of Haplostoma Chatton & Harant, 1924, seven of Haplostomella Chatton & Harant, 1924 and a single new species of Haplosaccus Chatton & Harant, 1924. The Enteropsidae comprised 42 species in five genera and here we add two new genera, Monnioticopa gen. nov. and Periboia gen. nov., plus a total of 30 new species; 15 of Enterocola van Beneden, 1860, two of Enterocolides Chatton & Harant, 1922, five of Enteropsis C.W.S. Aurivillius, 1885, five of Monnioticopa gen. nov., two of Mychophilus Hesse, 1865, plus the type species of Periboia gen. nov. Generic diagnoses are provided for all genera represented in the collection. A further 13 known species are also reported and brief supplementary descriptive notes or full redescriptions are provided, as appropriate.
References
Aurivillius, C.W.S. (1885) Krustaceer hos arktiska tunikater. Vega-Expeditionens Vetenskakpliga Iakattagelser, Stockholm, 4, 223–254, pls. 7–9.
Boxshall, G.A. & Halsey, S.H. (2004) An Introduction to Copepod Diversity. The Ray Society, London, 966 pp.
Canu, E. (1892) Le copépodes du Boulonnais. Morphologie, embryologie, taxonomie. Travaux du Laboratoire de Zoologie Maritime de Wimereux-Ambleteuse, 6, 1–354, pls. 1–30.
Chatton, É. & Brément, E. (1910) Sur trois ascidicoles du genre Aplostoma Canu: Aplostoma magellanica sp. nov., A. hibernica (T. et A. Scott), A. sacculus sp. nov. Note préliminaire. Bulletin de la Société Zoologique de France, 35, 80–92.
Chatton, É. & Harant, H. (1924a) Notes sur les copépods ascidicoles XVI. Le nouveau genre Haplostomella. Deux espèces nouvelles de ce genre. Remarques sur les oostegites et la 5” paire de péréipodes. Bulletin de la Société Zoologique de France, 49, 398–406.
Chatton, É. & Harant, H. (1924b) Notes sur les copépods ascidicoles XVII. Le nouveau genre Haplostomides et deux espèces nouvelles de ce genre: Haplostomides scotti et Haplostomides brementi. Bulletin de la Société Zoologique de France, 49, 406–412.
Chatton, É. & Harant, H. (1924c) Notes sur les copépods ascidicoles XVIII. Haplostoma canui sp. nov. Etat actuel de la systématique des Haplosominae n. subf. Le nouveau genre Haplosaccus. Bulletin de la Société Zoologique de France, 49, 413–422.
Conradi, M., López-González, P.J. & García-Gómez, J.C. (1992) A new species of Enterocola van Beneden, 1860 (Copepoda, Ascidicolidae) associated with Sidnyum elegans (Giard, 1872) from the Iberian Peninsula. Crustaceana, 63 (3), 247–256. https://doi.org/10.1163/156854092X00398
Conradi, M., López-González, P.J. & García-Gómez, J.C. (1994) Botryllophilus conicus sp. nov. (Copepoda: Cyclopoida: Ascidicolidae) associated with a compound ascidian from the Strait of Gibraltar. Systematic Parasitology, 29, 97–104. https://doi.org/10.1007/BF00009805
Dudley, P.L. & Illg, P.L. (1991) Marine flora and fauna of the eastern United States. Copepod, Cyclopoida: Archinotodelphyidae, Notodelphyidae, and Ascidicolidae. NOAA Technical Report, 96, 1–39.
Gotto, R.V. (1970) Haplostomella australiensis n. sp., an ascidicolous copepod from New South Wales. Crustaceana, 19, 267–272.
Gotto, R.V., Holmes, J.M.C., & Lowther, R.P. (1984) Description of the adult male Mychophilus roseus Hesse (Copepoda: Cyclopoida): a copepod with remarkable sensory equipment. Irish Naturalists’ Journal, 21 (7), 305–313.
Humes, A.G. & Gooding, R.U. (1964) A method for studying the external anatomy of copepods. Crustaceana, 6, 238–240.
http://dx.doi.org/10.1163/156854064x00650
Huys, R. & Boxshall, G.A. (1991) Copepod Evolution. The Ray Society, London, 468 pp.
Illg, P.L. & Dudley, P.L. (1980) The family Ascidicolidae and its subfamilies (Copepoda, Cyclopoida), with descriptions of new species. Mémoires du Muséum National d’Histoire Naturelle, Paris, Série A, Zoologie, 117, 1–192.
Kim, I.-H. & Boxshall, G.A. (2020a) A revision of the family Archinotodelphyidae Lang, 1949 (Copepoda: Cyclopoida: Oithonida), with the recognition of 15 new species. Zootaxa, 4801 (1), 1–56.
https://doi.org/10.11646/zootaxa.4801.1.1
Kim, I.-H. & Boxshall, G.A. (2020b) Untold diversity: the astonishing species richness of the Notodelphyidae (Copepoda: Cyclopoida), a family of symbiotic copepods associated with ascidians (Tunicata). Megataxa, 4, 1-660.
https://doi.org/10.11646/megataxa.4.1.1
Kim, I.-H. & Moon, S.Y. (2011) Eight new species of ascidicolous copepods from the eastern coast of Korea (Crustacea, Copepoda, Cyclopoida). Ocean Science Journal, 46 (1), 23–46.
https://doi.org/10.1007/s12601-011-0003-1
Kim, I.-H., Cruz-Rivera, E., Sherif, M.-E.-D. & El-Sahhar, S. (2016) Cyclopoid copepods (Ascidicolidae, Notodelphyidae) associated with Phallusia nigra Savigny, 1816 (Ascidiacea) in the Red Sea: A new ascidicolid and first descriptions of the males from two notodelphyids. Journal of Crustacean Biology, 36 (4), 553–566.
https://doi.org/10.1163/1937240X-00002439
Lang, K. (1948) Copepoda Notodelphyoida from the Swedish west-coast with an outline on the systematics of the Copepoda. Arkiv for Zoologi, 40A (14), 1–36.
López-González, P.J. & Conradi, M. 1996. Mychophilus palmatus, a new species (Copepoda: Cyclopoida: Ascidicolidae) associated with the compound ascidian Botryllus leachi Savigny, with remarks on congeneric species and related genera. Hydrobiologia, 330, 67–72.
López-González, P.J., Conradi, M. & García-Gómez, J.C. (1993) Enterocola africanus, a new species (Copepoda: Ascidicolidae) associated with a compound ascidian Synoicum species from North Africa (Strait of Gibraltar). Proceedings of the Biological Society of Washington, 106 (1), 131–136.
Lützen, J. (1968) Styelicola bahusia n. g., sp. nov. (family Ascidicolidae), a commensal copepod from Styela atlantica and S. gelainosa from the Skagerrak. Crustaceana, Supplement 1 (Studies on Copepoda), pp. 96–102.
[https://www.jstor.org/stable/25027368]
Marchenkov, A.V. (1994) A description of new species of the parasitic copepod Enteropsis arcticus sp. n. from ascidians of White Sea. Parazitologiya, 28 (3), 222–229. [in Russian, with English summary]
Marchenkov, A. & Boxshall, G.A. (2002) The Buproridae Thorell, 1859, a family of ascidicolous copepods (Copepoda: Cyclopoida). Systematic Parasitology, 53, 191–198.
https://doi.org/10.1023/A:1021147124817
Marchenkov, A. & Boxshall, G.A. (2003) Copepods of the genera Haplostomella and Haplostomides (Cyclopoida: Ascidicolidae) associated with ascidians from the White Sea and Russian Far East coastal waters. Hydrobiologia, 510, 1–15.
https://doi.org/10.1023/B:HYDR.0000008471.26364.e9
Marchenkov, A. & Boxshall, G. A. (2005) Three new species of Enterocola van Beneden, 1860 (Crustacea: Copepoda: Cyclopoida) from ascidian hosts. Systematic Parasitology, 61, 223–234.
http://doi.org/10.1007/s11230-005-3171-5
Monniot, C. (1982). Paulillgia polycarpae n. g., n. sp., Copépode parasite d’un Polycarpa (Ascidiacea) de la pente du plateau continental du Golfe de Gascogne. Crustaceana, 43, 21–24.
Ohtsuka, S., Kitazawa, K. & Boxshall, G.A. (2010) A new genus of the endoparasitic copepods (Cyclopoida: Enterognathidae) forming a gall in the calyx of deep-sea crinoids. Zoological Science, 27, 689–696.
Ooishi, S. (1994) Haplostomides hawaiiensis, new species (Copepoda: Cyclopoida: Ascidicolidae), associated with the ascidian Polyclinum constellatum at Honolulu, Hawaii. Hydrobiologia, 292/294, 89–96.
https://doi.org/10.1007/BF00229927
Ooishi, S. (1996) Two ascidicolid copepods, Botryllophilus macropus Canu and B. norvegicus Schellenberg, from British waters. Journal of Crustacean Biology, 16, 169–191.
https://doi.org/10.2307/1548939
Ooishi, S. (1998) Haplostoma dudleyae sp. nov. (Cyclopoida: Ascidicolidae), parasitic in Eudistoma olivaceum from the Indian River in southern Florida. Journal of Marine Systems, 15, 273–279.
https://doi.org/10.1016/S0924-7963(97)00068-7
Ooishi, S. (1999) Female and male Botryllophilus ruber (Copepoda: Cyclopoida) associated with the compound ascidian Botryllus schlosseri. Journal of Crustacean Biology, 19, 556–577.
https://doi.org/10.2307/1549262
Ooishi, S. (2000) Botryllophilus bamfieldensis, new species (Copepod: Cyclopoida: Ascidicolidae), living in a compound ascidian from the west coast of Vancouver Island, Canada. Journal of Crustacean Biology, 20 (3), 571–588.
https://doi.org/10.1651/0278-0372(2000)020[0571:BBNSCC]2.0.CO;2
Ooishi, S. (2002a) Female and male Haplostomides scotti (Copepoda: Cyclopoida) living in the compound ascidian Polyclinum aurantium. Journal of Crustacean Biology, 22 (2), 249–267.
https://doi.org/10.1163/20021975-99990232
Ooishi, S. (2002b) Redescription of Botryllophilus inaequipes Hansen (Crustacea: Copepoda: Cyclopoida: Ascidicolidae). Proceedings of the Biological Society of Washington, 115 (3), 636–649.
Ooishi, S. (2002c) Description of Botryllophilus sarsi, new species (Copepoda: Cyclopoida: Ascidicolidae), replacing B. brevipes Sars, 1921. Journal of Crustacean Biology, 22 (4), 819–833.
https://doi.org/10.1163/20021975-99990295
Ooishi, S. (2004a) Redescription of Haplostoma banyulensis (Copepoda: Cyclopoida: Ascidicolidae) living in the compound ascidian Trididemnum tenerum from Strangford Lough, Northern Island. Journal of Crustacean Biology, 24 (1), 1–8.
https://doi.org/10.1651/C-2414
Ooishi, S. (2004b) Female and male Haplostoma brevicauda (Copepoda: Cyclopoida: Ascidicolidae), living in compound ascidians. Journal Crustacean Biology, 24 (3), 422–439.
https://doi.org/10.1651/C-2465
Ooishi, S. (2006) Two species of Botryllophilus (Copepoda: Cyclopoida) living in compound ascidians, and a revision of female morphotype A of the genus. Journal of Crustacean Biology, 26, 23–47.
https://doi.org/10.2988/11-34.1
Ooishi, S. (2007a) Female Ascidicola rosea Thorell (Copepoda: Cyclopoida) living in solitary ascidians. Journal of Crustacean Biology, 27 (2), 327–338.
https://doi.org/10.1651/S-2673.1
Ooishi, S. (2007b) Enterocola fulgens van Beneden (Crustacea: Copepoda: Cyclopoida) living in the compound ascidian Polyclinum aurantium Milne Edwards. Proceedings of the Biological Society of Washington, 120 (3), 293–310.
https://doi.org/10.2988/0006-324X(2007)120[293:EFVBCC]2.0.CO;2
Ooishi, S. (2008a) Haplostomides gottoi, new species (Crustacea: Copepoda: Cyclopoida: Ascidicolidae), living in a compound ascidian from Madagascar. Proceedings of the Biological Society of Washington, 121 (1), 85–93.
https://doi.org/10.2988/07-33.1
Ooishi, S. (2008b) Female Enteropsis roscoffensis Chatton & Brément (Crustacea: Copepoda: Cyclopoida: Ascidicolidae) living in the solitary ascidian Dendrodoa grossularia. Proceedings of the Biological Society of Washington, 121 (4), 501–511.
https://doi.org/10.2988/07-45.1
Ooishi, S. (2008c) Female and male Mychophilus roseus Hesse (Crustacea: Copepoda: Cyclopoida: Ascidicolidae) living in the compound ascidian Botryllus schlosseri. Proceedings of the Biological Society of Washington, 121 (4), 512–530.
https://doi.org/10.2988/07-46.1
Ooishi, S. (2009a) Enteropsis fusiformis, new species (Copepoda: Cyclopoida: Ascidicolidae), living in a compound ascidian from Madagascar. Proceedings of the Biological Society of Washington, 122 (3), 333–341.
https://doi.org/10.2988/08-46.1
Ooishi, S. (2009b) Four new species of Haplostoma (Copepoda: Cyclopoida: Ascidicolidae) living in compound ascidians from Madagascar. Proceedings of the Biological Society of Washington, 122 (4), 426–448.
https://doi.org/10.2988/09-06.1
Ooishi, S. (2010a) Enterocola hessei Chatton & Harant (Copepoda: Cyclopoida: Ascidicolidae) living in the compound ascidian Clavelina lepadiformis (Müller). Proceedings of the Biological Society of Washington, 123 (2), 137–148.
https://doi.org/10.2988/09-23.1
Ooishi, S. (2010b) Enterocola brementi Illg & Dudley (Copepoda: Cyclopoida: Ascidicolidae) living in the compound ascidian Aplidium pallidum (Verrill), compared with E. mammiferus Chatton & Harant living in A. sperum Drasche. Proceedings of the Biological Society of Washington, 123 (4), 312–323.
https://doi.org/10.2988/09-27.1
Ooishi, S. (2011) Female and male Enterocola sydnii Chatton & Harant (Copepoda: Cyclopoida: Ascidicolidae) living in compound ascidians. Proceedings of the Biological Society of Washington, 124 (4), 358–377.
https://doi.org/10.2988/11-12.1
Ooishi, S. (2012) Two new species of Botryllophilus (Copepoda: Cyclopoida: Ascidicolidae) living in compound ascidians from Madagascar. Proceedings of the Biological Society of Washington, 125, 122–144.
https://doi.org/10.2988/11-34.1
Ooishi, S. (2014a) Three species of Enterocola van Beneden (Copepoda: Cyclopoida: Ascidicolidae) living in compound ascidians from Madagascar, with a revision of Enterocola setiferus Hansen. Proceedings of the Biological Society of Washington, 126 (4), 343–368.
https://doi.org/10.2988/0006-324X-126.4.343
Ooishi, S. (2014b) Botryllophilus symmetricus, new species (Copepoda: Cyclopoida: Ascidicolidae), living in an undescribed compound ascidian (Synoicum) from Madagascar. Proceedings to the Biological Society of Washington, 127 (2), 340–352.
https://doi.org/10.2988/0006-324X-127.2.340
Ooishi, S. (2014c) Botryllophilus kozloffi, new species (Copepoda: Cyclopoida: Ascidicolidae), living in the compound ascidian Clavelina lepadiformis (Müller) from Roscoff, France. Proceedings of the Biological Society of Washington, 127 (3), 483–495.
https://doi.org/10.2988/0006-324X-127.3.483
Ooishi, S. (2014d) Botryllophilus millari, new species (Copepoda: Cyclopoida: Ascidicolidae), living in the compound ascidian Eudistoma caeruleum (Sluiter) from Madagascar. Proceedings to the Biological Society of Washington, 127 (3), 496–509.
https://doi.org/10.2988/0006-324X-127.3.496
Ooishi, S. & Illg, P.L. (1977) Haplostominae (Copepoda, Cyclopoida) associated with compound ascidians from the San Juan Archipelago and vicinity. Special Publications from the Seto Marine Biological Station, Series 5, 1–154.
Ooishi, S. & Illg, P.L. (1988) Two morpho-types of Botryllophilus (Cyclopoida: Ascidicolidae). Hydrobiologia, 167/168, 561–566.
https://doi.org/10.1007/978-94-009-3103-9_65
Ooishi, S. & Illg, P.L. (1989) Botryllophilus abbotti new species (Copepoda: Cyclopoida: Ascidicolidae) associated with a compound ascidian from the Monterey Peninsula. Bulletin of Marine Science, 45, 447–466.
Ooishi, S. & O’Reilly, M.G. (2004) Redescription of Haplostoma eruca (Copepoda: Cycloppoida: Ascidicolidae) living in the intestine of Ciona intestinalis from the Clyde Estuary, Scotland. Journal of Crustacean Biology, 24 (1), 9–16.
https://doi.org/10.1651/C-2415
Sars, G.O. (1921) An account of the Crustacea of Norway with short descriptions and figures of all the species. Vol. 8. Copepoda. Monstrilloida and Notodelphyoida. Bergen Museum, Bergen, 91 pp., 37 pls.
Schellenberg, A. (1922) Neue Notodelphyiden des Berliner und Hamburger Museums mit einer Übersicht der ascidienbewohnenden Gattungen und Arten. 2. Mitteilungen aus dem Zoologischen Museum in Berlin, 10 (2), 275–298.
Seo, I.S. & Lee, K.S. (1995) Copepod associated with ascidians from Korea. II. A new species of Botryllophilus (Copepoda, Cyclopoida, Ascidicolidae) from simple ascidians at Chindo Island in the South Sea, Korea. Korean Journal of Systematic Zoology, 11 (1), 79–86.
Seo, I.S. & Lee, K.S. (2001) Haplostoma kimi, a new species and a redescription of Haplostomella halocynthiae (Fukui) from Korea (Copepoda: Cyclopoida: Ascidicolidae). Proceedings of the Biological Society of Washington, 114 (1), 229–236.
Stock, J.H. (1967) Mychophilus fallax n. sp., a new vermiform copepod parasite of a Red Sea tunicate. Reports of the Israel South Red Sea Expedition, 24, 9–12.
Stock, J.H. (1970) Notodelphyidae and Botryllophilidae (Copepoda) from the West Indies. Studies on the Fauna of Curaçao and other Caribbean Islands, 34 (123), 1–45.
Thorell, T. (1859) Bidrag til Kännedomen om Krustaceer som lefva i arter af Slägtet Ascidia L. Kongliga Svenska vetenskaps-akademiens handlingar, 3 (8), 1–84.
Tovar, M.A., Suárez-Morales, E. & Yáñez, B. (2010) The parasitic copepod Haplostomides hawaiiensis (Cyclopoida) from the invasive ascidian Polyclinum constellatum in the Southern Gulf of California. Bulletin of Marine Science, 86 (3), 637–648.
Walter, T.C. & Boxshall, G.A. (2020) World of Copepods database. Cyclopoida. Accessed through: World Register of Marine Species. Available from: ahttp://www.marinespecies.org/aphia.php?p=taxdetails&id=1101 (accessed 16 July 2020)
WoRMS Editorial Board (2020) World Register of Marine Species. VLIZ. Available from: http://www.marinespecies.org (accessed 18 February 2020)