Abstract
Early studies during the 1990’s supported a close relationship between the genera Paralaophonte and Loureirophonte. Subsequent studies carried out during the first decade of the 2000’s suggested the monophyly of the latter and that it could be nested within Paralaophonte, rendering the latter paraphyletic. However, efforts to synonymize these two genera were hampered by the lack of phylogenetic analyses at the species level. Here we describe a new species of Paralaophonte from north-western Mexico, P. ullama sp. nov., and we formally synonymize Loureirophonte with Paralaophonte for which we propose two monophyletic subgenera, Pa. (Paralaophonte) and Pa. (Loureirophonte).
The detection of the species most closely related to P. ullama sp. nov. is controversial. Our Bayesian analysis indicates that the new species might be related to Pa. (Pa.) gurneyi, and to Pa. (Pa.) ormieresi and Pa. (Pa.) septemarticulata, with which they form a larger clade supported by a very low Bayesian Posterior Probability value. However, the precise nature of the relationship between the new species and Pa. (Pa.) gurneyi is hampered by the gap in our knowledge of the armature formulae of the swimming legs of the latter, and by the lack of the male of the new species. On the other hand, the new species share several characters with Pa. (Pa.) problematica, Pa. (Pa.) pacificavicinum, Pa. (Pa.) pacificaemulator, and Pa. (Pa.) pacifica, with which it might be related. Our Bayesian analysis of forty-two morphological characters at the species level confirmed earlier suspicions about the relationships between Paralaophonte and Loureirophonte, the latter being nested within the former, and showed the monophyly of the Paralaophonte-Loureirophonte clade with high Bayesian Posterior Probability value, thus confirming the synonymy of these genera. The Paralaophonte-Loureirophonte clade is defined by the synapomorphic dimorphism in the second endopodal segment of the second swimming leg of the male. We did not detect any (aut)apomorphy for Pa. (Paralaophonte). Paralaophonte (Loureirophonte) is defined by the presence of one seta only on the second endopodal segment of the second swimming leg, and the one-segmented endopod of the fourth swimming leg.
References
Apostolov, A. (2004) Paralaophonte livingstoni n. sp. (Crustacea: Copepoda: Harpacticoida)—un nouveau représentant de l’île de Livingstone, Antarctique. Historia Naturalis Bulgarica, 16, 59–67.
Apostolov, A. (2008) Harpacticoïdes (Crustacea, Copepoda) de la mer égée (plages de Kavala, Grèce du nord) [Harpacticoids (Crustacea, Copepoda) from the Aegean Sea (Kavala beaches, northern Greece)]. Historia Naturalis Bulgarica, 19, 5–33.
Brady, G.S. (1899) On the marine Copepoda of New Zealand. Transactions of The Zoological Society of London, 15, 31–54. https://doi.org/10.1111/j.1096-3642.1899.tb00018.x
Brady, G.S. (1910) Die marinen Copepoden der Deutschen Südpolar Expedition 1901-1903. I. Über die Copepoden der Stämme Harpacticoida, Cyclopoida, Notodelphyoida und Caligoida. Deutsche Südpolar-Expedition 1901–1903, 3, 497–594.
Brian, A. (1917) Descrizione di una nuove specie di Laophonte (L. quaterspinata n. sp. mihi) proveniente dai materiali del laboratorio marino di Quarto. Monitore Zoologico Italiano, 28, 174–178.
Burgess, R. (2001) An improved protocol for separating meiofauna from sediments using colloidal silica sols. Marine Ecology Progress Series, 214, 161–165. https://doi.org/10.3354/meps214161
Chislenko, L.L. (1977) Garpaktitsidy (Copepoda, Harpacticoida) s gubok zemli Frantsa-Iosifa. Harpacticids (Copepoda Harpacticoidea) from sponges of Franz Josef Land. Biotsenosy shel’fa zemli Frantsa-Iosifa i fauna sopredel’nykh akuatorii. Issledovaniya Fauny Morei, 14, 237–276.
Chislenko, L.L. (1978) New species of harpacticoid copepods (Copepoda, Harpacticoida) from Poss’yet Bay, Sea of Japan. Trudy Zoologicheskogo Instituta, Akademii Nauk SSSR, Leningrad, 61, 161–192.
Claus, C. (1863) Die frei lebenden Copepoden mit besonderer Berucksichtigung der Fauna Deutschlands, der Nordsee und des Mittelmeeres. Verlag von Wilhelm Engelmann, Leipzig, 230 pp. https://doi.org/10.5962/bhl.title.58676
Douwe, C. van (1929) Marine Litoral-Copepoden: zur Verbreitung des Genus Laophonte Philippi im Mittelmeer. Zoologischer Anzeiger, 83, 283–294.
Edwards, C.L. (1891) Beschreibung einiger neuen Copepoden und eines neuen copepodenähnlichen Krebses, Leuckartella paradoxa. Archiv fur Naturgeschichte, Inaugural-Dissertation einer Hohen Philosophischen Faultat der Universität Leipzig, Berlin, 57, 75–104.
Fiers, F. (1986) Harpacticoid copepods from the west Indian Islands: Laophontidae (Copepoda, Harpacticoida). Bijdragen tot de Dierkunde, 56, 132–164. https://doi.org/10.1163/26660644-05601009
Fiers, F. (1993) The laophontid genus Loureirophonte Jakobi, 1953 (Copepoda, Harpacticoida). Zoologische Mededelingen, 67, 207–238.
Gheerardyn, H., Fiers, F., Vincx, M. & De Troch, M. (2006) Paralaophonte harpagone sp. n. (Copepoda: Harpacticoida), a laophontid with an extremely specialised maxilliped. Organisms Diversity and Evolution, 6, 1–9. https://doi.org/10.1016/j.ode.2005.12.004
Gómez, S. & Boyko, C.B. (2006) On a small collection of harpacticoids from Easter Island: the family Laophontidae T. Scott (Crustacea: Copepoda: Harpacticoida). Zootaxa, 1352 (1), 1–70. https://doi.org/10.11646/zootaxa.1352.1.1
Gómez, S. & Morales-Serna, F.N. (2013) On a small collection of Laophontidae T. Scott (Copepoda: Harpacticoida) from Mexico. I. New species and new records of Laophonte Philippi and Paralaophonte Lang. Journal of Natural History, 47, 329–380. https://doi.org/10.1080/00222933.2012.757657
Hamond, R. (1973) Some Laophontidae (Crustacea: Harpacticoida) from off North Carolina. Transactions of the American Microscopical Society, 92, 44–59. https://doi.org/10.2307/3225170
Huys, R. (2009) Unresolved cases of type fixation, synonymy and homonymy in harpacticoid copepod nomenclature (Crustacea: Copepoda). Zootaxa, 2183 (1), 1–99. https://doi.org/10.11646/zootaxa.2183.1.1
Huys, R. & Boxshall, G.A. (1991) Copepod evolution. The Ray Society, London, 468 pp.
Huys, R. & Lee, W. (2000) Basal resolution of laophontid phylogeny and the paraphyly of Esola Edwards. Bulletin of the Natural History Museum, London, Zoology, 66, 49–107.
Huys, R. & Lee, W. (2009) Proposal of Marbefia, gen. n. and Inermiphonte, gen. n., including updated keys to the species of Pseudonychocamptus Lang, 1944 and Paralaophonte Lang, 1948 (Copepoda, Harpacticoida, Laophontidae). ZooKeys, 23, 1–38. https://doi.org/10.3897/zookeys.23.168
Jakobi, H. (1953) Novos Laophontidae (Copepoda-Crustacea) da costa Brasileira (Neue Laophontiden von der Brasilianischen Kueste). Dusenia, 4, 47–60.
Jakubisiak, S. (1932) Sur les harpacticoïdes hébergés par Maja squinado. Bulletin de la Société Zoologique de France, 57, 506–513.
Lang, K. (1934) Marine harpacticiden von der Campbell-Insel und einigen anderen südlichen Inseln. Acta Universitatis Lundensis, New Series, Avd. 2 (=Kungl. Fysiografiska Sällskapets Handlingar, 45 (14), 1–56.
Lang, K. (1944) Monographie der Harpacticiden (vorläufige Mitteilung). Almqvist & Wiksells Boktryckeri AB, Uppsala, 39 pp.
Lang, K. (1948) Monographie der Harpacticiden. Vols. I & II. Nordiska Bokhandeln, Stockholm, 1682 pp.
Lang, K. (1965) Copepoda Harpacticoidea from the Californian Pacific coast. Kungliga Svenska Vetenskapsakademiens Handlingar, Series 4, 10, 1–560.
Lee, W. & Huys, R. (1999) Bathylaophonte gen. nov. from deep-sea hydrothermal vents and the polyphyly of Paronychocamptus (Copepoda: Harpacticoida). Cahiers de Biologie Marine, 40, 293–328. https://doi.org/10.21411/CBM.A.36920EDF
Mielke, W. (1974) Eulitorale Harpacticoidea (Copepoda) von Spitzsbergen. Mikrofauna des Meeresbodens, 37, 1–52.
Mielke, W. (1981) Interstitielle Fauna von Galapagos. XXVIII. Laophontinae (Laophontidae), Ancorabolidae (Harpacticoida). Mikrofauna des Meeresbodens, 84, 1–106.
Mielke, W. (1982) Einige Laophontidae (Copepoda, Harpacticoida) von Panama. Crustaceana, 42, 1–11. https://doi.org/10.1163/156854082X00641
Mielke, W. (2001) Loureirophonte psammophila, a new species of Laophontidae T. Scott, 1904 (Copepoda: Harpacticoida) from Baja California, Mexico. Hydrobiologia, The Hague, 445, 77–83. https://doi.org/10.1023/A:1017546905481
Milne Edwards, H. (1840) Ordre des copépodes. Histoire naturelle des crustacés, comprenant l’anatomie, la physiologie et la classification de ces animaux, 3, 411–529.
Monard, A. (1928) Les harpacticoïdes marins de Banyuls. Archives de Zoologie Expérimentale et Générale, 67, 259–443.
Monard, A. (1935a) Étude sur la faune des harpacticoïdes marins de Roscoff. Travaux de la Station Biologique de Roscoff, 13, 5–88.
Monard, A. (1935b) Les harpacticoïdes marins de la région de Salammbô. Bulletin de la Station Océanographique de Salammbô, 34, 1–94.
Monard, A. (1936) Note préliminaire sur la fauna des Harpacticoïdes marins d’Alger. Bulletin de la Station d’Aquiculture et de Pêche Castiglione, 1935, 45–85.
Nicholls, A.G. (1944) Littoral Copepoda from the Red Sea. Annals and Magazine of Natural History, Series 11, 11 (80), 487–503. https://doi.org/10.1080/00222934408527451
Petkovski, T.K. (1964) Bemerkenswerte Entomostraken aus Jugoslavien. Acta Musei Macedonici Scientiarum Naturalium, Skopje, 9, 147–181.
Philippi, A. (1840) Zoologische Bemerkungen. Archiv für Naturgeschichte, 6, 181–195.
Por, F.D. (1964) A study of the Levantine and Pontic Harpacticoida (Crustacea, Copepoda). Zoologische Verhandelingen, 64, 1–128.
Raibaut, A. (1968) Paralaophonte ormieresi n. sp., copépode harpacticoide trouvé sur les branchies de Maja squinado (Herbst) (Crustacea, Decapoda). Bulletin de la Société Zoologique de France, 93, 451–457.
Rambaut, A. (2018) FigTree. Version 1.4.4. Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 24 August 2021)
Rohal, M., Thistle, D. & Easton, E.E. (2018) Extraction of metazoan meiofauna from muddy deep-sea samples: operator and taxon effects on efficiency. Journal of Experimental Marine Biology and Ecology, 502, 105–110. https://doi.org/10.1016/j.jembe.2017.01.006
Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) Mrbayes 3.2: Efficient bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. https://doi.org/10.1093/sysbio/sys029
Sars, G.O. (1903) Copepoda Harpacticoida. Parts I & II. Misophriidae, Longipediidae, Cerviniidae, Ectinosomidae. An account of the Crustacea of Norway, with short descriptions and figures of all the species, 5, 1–28.
Sars, G.O. (1908) Copepoda Harpacticoida. Parts XXIII & XXIV. Laophontidae (continued). An account of the Crustacea of Norway, with short descriptions and figures of all the species, 5, 257–276.
Sars, G.O. (1909) Crustacea. Report of the Second Norwegian Arctic Expedition in the “FRAM” 1898–1902, 3, 1–47.
Sars, G.O. (1911) Copepoda Harpacticoida. Parts XXXV & XXXVI. Supplement (concluded), index etc. An account of the Crustacea of Norway, with short descriptions and figures of all the species, 5, 421–449.
Sars, G.O. (1920) Copepoda Supplement. Parts V & VI. Harpacticoida (continued). An account of the Crustacea of Norway, with short descriptions and figures of all the species, 7, 53–72.
Scott, A. (1902) On some Red Sea and Indian Ocean Copepoda. Proceedings and Transactions of the Liverpool Biological Society, 16, 397–428.
Scott, T. (1892) III Additions to the fauna of the Firth of Forth. Part IV. Annual Report of the Fishery Board for Scotland, 10, 244–272.
Scott, T. (1894) Report on Entomostraca from the Gulf of Guinea, collected by John Rattray, B. Sc. Transactions of the Linnean Society of London, Series 2, Zoology, 6, 1–161. https://doi.org/10.1111/j.1096-3642.1894.tb00660.x
Scott, T. (1899) Report on the marine and freshwater Crustacea from Franz-Josef Land, collected by Mr. William S. Bruce, of the Jackson-Harmsworth Expedition. Journal of the Linnean Society, Zoology, 27, 60–126. https://doi.org/10.1111/j.1096-3642.1899.tb01421.x
Scott, T. (1904) On some new and rare Crustacea from the Scottish seas. Reports of the Fishery Board for Scotland, 23, 141–153.
Scott, T. & Scott, A. (1893) On some new or rare Scottish Entomostraca. Annals and Magazine of Natural History, 6, 210–215. https://doi.org/10.1080/00222939308677499
Vervoort, W. (1962) Report on some Copepoda collected during the Melanesia Expedition of the Osaka Museum of Natural History. Publications of the Seto Marine Biological Laboratory, 10, 393–470. https://doi.org/10.5134/175305
Vervoort, W. (1964) Free-living Copepoda from Ifaluk Atoll in the Caroline Islands with notes on related species. Bulletin of the United States National Museum, 236, 1–431. https://doi.org/10.5479/si.03629236.236.1
Wells, J.B.J. (2007) An annotated checklist and keys to the species of Copepoda Harpacticoida (Crustacea). Zootaxa, 1568 (1), 1–872. https://doi.org/10.11646/zootaxa.1568.1.1
Wells, J.B.J., Hicks, G.R.F. & Coull, B.C. (1982) Common harpacticoid copepods from New Zealand harbours and estuaries. New Zealand Journal of Zoology, 9, 151–184. https://doi.org/10.1080/03014223.1982.10423847
Willen, E. (1995) Archilaophonte maxima gen. n., spec. n., a new taxon of the Laophontidae (Copepoda, Harpacticoida) from the high Antartic (Weddell Sea). Hydrobiologia, 302, 241–255. https://doi.org/10.1007/BF00032113
Willey, A. (1932) Copepod phenology. Observations based on new material from Canada and Bermuda. Archivio di Zoologia Italiana, Atti dell’ XI Congresso Internazionale Zoologia, 16, 601–617. [1930]