Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 4229 No. 1: 8 Feb. 2017
Type: Monograph
Published: 2017-02-07
DOI: 10.11646/zootaxa.4229.1.1
Page range: 1–183
Abstract views: 326
PDF downloaded: 3

Revision of Family Megacalanidae (Copepoda: Calanoida)

National Institute of Water and Atmospheric Research, Private Bag 14901, Wellington 6241, New Zealand.
Department of Marine Sciences, University of Connecticut, 1080 Shennecossett Road, Groton, 06340 CT, USA. Bermuda Institute of Ocean Sciences, St George’s GE 01, Bermuda.
Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, U.K.
Crustacea Megacalanus Bradycalanus Bathycalanus Elenacalanus nom. nov. key morphology genes phylogeny Atlantic Indian Pacific bathypelagic abyssopelagic Megacalanus frosti n. sp. M. ericae n.sp. M. ohmani n. sp. Bradycalanus abyssicolus n. sp. Bathycalanus dentatus n. sp. Ba. milleri n. sp. Ba. tumidus n. sp. Ba. adornatus n. sp. Ba. pustulosus n. sp. Ba. bucklinae n. sp. Elenacalanus tageae n. sp.

Abstract

The Megacalanidae were revised based on new and archived material. Taxonomic confusion that has existed in the family is discussed and a method is suggested for stabilising names. A detailed examination of the morphology of this family, using the light microscope, has added further useful characters that distinguish genera and species. The added, hitherto undescribed species include character states incompatible with aspects of previous generic definitions (e.g. presence or absence of setae on the maxillule coxal endite). Nevertheless, the cladistic and molecular analyses confirmed that there are at least four monophyletic clades mostly with high bootstrap support. These clades represent already defined genera, one of which [Elenacalanus nom. nov. (nomen novum)] replaces the preoccupied name Heterocalanus Wolfenden, 1906. Four previously described species have been re-assigned to Elenacalanus in new combinations: E. princeps (Brady, 1883), E. eltaninae (Björnberg, 1968), E. sverdrupi (Johnson, 1958) and E. inflatus (Björnberg, 1968). Eleven new species are described: three Megacalanus, one Bradycalanus, six Bathycalanus, and one Elenacalanus nom. nov. Bradycalanus pseudotypicus enormis Björnberg, 1968 has been raised to species status based on genetic data although it can be only be distinguished morphologically from Br. typicus by its large size. All four genera are differentially diagnosed and keys are provided to the genera and species. We confirm that all male right antennules are geniculate in the Megacalanidae. Thirteen males are known. Of these males, eight are newly described (M. frosti n. sp., M. ericae n. sp., M. ohmani n. sp., Bathycalanus bradyi (Wolfenden, 1905a), Ba. dentatus n. sp., Ba. milleri n. sp., Ba. unicornis Björnberg, 1968, and Elenacalanus tageae n. sp.). We cannot be absolutely certain that the correct males have been assigned to the appropriate female so our decisions await testing with new data. The cladistic analysis provides the first morphology-based phylogeny. This scheme served as a working hypothesis which was tested and corroborated using the newly gathered molecular data. Vertical and horizontal distributions are summarised.

 

References

  1. Altekar, G., Dwarkadas, S., Huelsenbeck, J.P. & Ronquist, F. (2004) Parallel Metropolis coupled Markov chain Monte Carlo for Bayesian phylogenetic inference. Bioinformatics, 20, 407–415.
    https://doi.org/10.1093/bioinformatics/btg427

    Anderson, F.E. & Swofford, D.L. (2004) Should we be worried about long-branch attraction in real data sets? Investigations using metazoan 18S rDNA. Molecular Phylogenetics and Evolution, 33, 440–451.
    https://doi.org/10.1016/j.ympev.2004.06.015

    Anderson, M.J., Gorley, R.N. & Clarke, K.R. (2008) PERMANOVA+ for PRIMER: guide to software and statistical methods. PRIMER-E, Plymouth.

    Arashkevich, Y.G. (1969) The food and feeding of copepods in the northwestern pacific. Okeanologiya, 9, 857–873. [English translation in Oceanology, 9, 695–709. 1970]

    Barthélémy, R.-M. (1999) Biologie de la reproduction des copépods calanoïdes: Biodiversité morphofonctionnelle et intérêt phylétique des structure génitales femelles données structurales, ultrastructurales et biochimiques sur les grandes associées. These pour obtenir le grade de Docteur de L’Université de Provence. 95 pp., 41 figs.

    Bergsten, J. (2005) A review of long-branch attraction. Cladistics, 21, 163–193.
    https://doi.org/10.1111/j.1096-0031.2005.00059.x

    Björnberg, T.K.S. (1968) Four new species of Megacalanidae (Crustacea: Copepoda). Antarctic Research Series, 11, 73–90.

    Blanco-Bercial, L., Bradford-Grieve, J.M. & Bucklin, A. (2011) Molecular phylogeny of the Calanoida (Crustacea: Copepoda). Molecular Phylogenetics and Evolution, 59, 103–113.
    https://doi.org/10.1016/j.ympev.2011.01.008

    Blanco-Bercial, L., Cornils, A., Copley, N. & Bucklin, A. (2014) DNA Barcoding of marine copepods: assessment of analytical approaches to species identification. PLOS Currents Tree of Life, Edition 1.
    https://doi.org/10.1371/currents.tol.cdf8b74881f87e3b01d56b43791626d2

    Bochdansky, A.B., van Aken, H.M. & Herndl, G.J. (2010) Role of macroscopic particles in deep-sea oxygen consumption. Proceeding of the National Academy of Sciences, 107 (18), 8287–8291.
    https://doi.org/10.1073/pnas.0913744107

    Böttger-Schnack, R. (2009) Taxonomy of Oncaeidae (Copepoda, Cyclopoida s.l.) from the Red Sea. IX. Epicalymma bulbosa sp. nov., first record of the genus in the Red Sea. Journal of Plankton Research, 31, 1027–1043.
    https://doi.org/10.1093/plankt/fbp051

    Böttger-Schnack, R. & Machida, R. (2011) Comparison of morphological and molecular traits for species identification and taxonomic grouping of oncaeid copepods. Hydrobiologia, 666, 111–125.
    https://doi.org/10.1007/s10750-010-0094-1

    Boxshall, G.A. (1985) The comparative anatomy of two copepods, a predatory calanoid and a particle feeding mormonilloid. Philosophical Transactions of the Royal Society, London, Series B, 311, 303–377.
    https://doi.org/10.1098/rstb.1985.0155

    Boxshall, G.A. & Huys, R. (1998) The ontogeny and phylogeny of copepod antennules. Philosophical Transactions of the Royal Society of London Series B, 353, 765–786.
    https://doi.org/10.1098/rstb.1998.0242

    Bradford, J.M., Ohman, M.D. & Jillett, J.B. (1988) Larval morphology and development of Neocalanus tonsus, Calanoides macrocarinatus, and Calanus australis (Copepoda: Calanoida) in the laboratory. New Zealand Journal of Marine and Freshwater Research, 22, 301–320.
    https://doi.org/10.1080/00288330.1988.9516303

    Bradford-Grieve, J.M. (1994) The marine fauna of New Zealand: Pelagic calanoid Copepoda: Megacalanidae, Calanidae, Paracalanidae, Mecynoceridae, Eucalanidae, Spinocalanidae, Clausocalanidae. New Zealand Oceanographic Memoir, 102, 1–160.

    Bradford-Grieve, J.M., Blanco-Bercial, L. & Boxshall, G.A. (2016a) TreeBASE web site hosted by National Evolutionary Synthesis Center. Avaliable from: http://purl.org/phylo/treebase/phylows/study/TB2:S19003 (Accessed 7 Feb. 2017)

    Bradford-Grieve, J.M., Blanco-Bercial, L. & Boxshall, G.A. (2016b) Zenodo web site, European Organization for Nuclear Research Ch-1211 Cern, Genève 23, Switzerland.
    http://dx.doi.org/10.5281/zenodo.46925

    Bradford-Grieve, J.M., Blanco-Bercial, L. & Boxshall, G.A. (2016c) NIWA web site, NIWA, Wellington, available from http://www.niwa.co.nz/static/web/megacalanidae.zip [6MB ZIP]. (accessed 21 September 2016)

    Bradford-Grieve, J.M., Boxshall, G.A. & Blanco-Bercial, L. (2014) Revision of basal calanoid copepod families, with a description of a new species and genus of Pseudocyclopidae. Zoological Journal of the Linnean Society, 171, 507–533.
    https://doi.org/10.1111/zoj.12141

    Brady, G.S. (1883) Report on the Copepoda collected by the H.M.S. “Challenger” during the years 1873–76. Report on the Scientific Results of the Voyage of the H.M.S. Challenger 1873–76, Zoology, 8, 1–142, 55 pls.

    Braga, E., Zardoya, R., Meyer, A. & Yen, J. (1999) Mitochondrial and nuclear rRNA based copepod phylogeny with emphasis on the Euchaetidae (Calanoida). Marine Biology, 133, 79–90.
    https://doi.org/10.1007/s002270050445

    Brodsky, K.A., Vyshkvatzeva, N.V., Koc, M.C. & Markhaseva, E.L. 1983. [Copepod crustaceans (Copepoda: Calanoida) of the seas of SSSR and neighbouring waters.] Opredeliteli po Faune SSSR, Isdavayemyye Zoologicheskij Institutom Akademii Nauk, SSSR, 1, 1–356. [In Russian]

    Bucklin, A. & Frost, B.W. (2009) Morphological and molecular phylogenetic analysis of evolutionary lineages within Clausocalanus (Copepoda: Calanoida). Journal of Crustacean Biology, 29, 111–120.
    https://doi.org/10.1651/07-2879.1

    Bucklin, A., Frost, B.W., Bradford-Grieve, J., Allen, L.D. & Copley, N.J. (2003) Molecular systematic and phylogenetic assessment of 34 calanoid copepod species of the Calanidae and Clausocalanidae. Marine Biology, 142, 333–343.
    https://doi.org/10.1007/s00227-002-0943-1

    Bucklin, A., Frost, W.B. & Kocher, T.D. (1992) DNA sequence variation of the mitochondrial 16S rRNA in Calanus (Copepoda: Calanoida): intraspecific and interspecific patterns. Molecular Marine Biology and Biotechnology, 1, 397–407.

    Claus, C. (1863) Die freilebenden Copepoden mit besonderer Berücksichtigung der Fauna Deutschlands, der Nordsee und des Mittelmeeres. Leipzig, 1–230, pls 1–37.

    Coleman, A.W. (2007) Pan-eukaryote ITS2 homologies revealed by RNA secondary structure. Nucleic Acids Research, 35, 3322–3329.
    https://doi.org/10.1093/nar/gkm233

    Coleman, A.W. (2009) Is there a molecular key to the level of “biological species” in eukaryotes? A DNA guide. Molecular Phylogenetics and Evolution, 50, 197–203.
    https://doi.org/10.1016/j.ympev.2008.10.008

    Coleman, C.O. (2003) “Digital inking”: how to make perfect line drawings on computers. Organisms, Diversity and Evolution, 14, 1–14.
    https://doi.org/10.1078/1439-6092-00081

    Coleman, C.O. Lowry J.K. & Macfarlane, T. (2010) DELTA for beginners: An introduction into the taxonomy software package DELTA. Zookeys, 45, 1–75.
    https://doi.org/10.3897/zookeys.45.263

    Colgan, D.J., McLauchlan, A., Wilson, G.D.F., Livingston, S.P., Edgecombe, G.D., Macaranas, J., Cassis, G. & Gray, M.R. (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology, 46, 419–437.
    https://doi.org/10.1071/ZO98048

    Cornils, A. & Blanco-Bercial, L. (2013) Phylogeny of the Paracalanidae Giesbrecht, 1888 (Crustacea: Copepoda: Calanoida). Molecular Phylogenetics and Evolution, 69, 861–872.
    https://doi.org/10.1016/j.ympev.2013.06.018

    Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1993) Users’ guide to the DELTA System: a general system for processing taxonomic descriptions. Edn. 4 136 pp. CSIRO Division of Entomology, Canberra.

    Damkaer, D.M. (2000) Determination and enthusiasm: Richard Norris Wolfenden (1854–1926), his Plankton studies and other things oceanographical. Archives of Natural History, 27 (2), 209–229.
    https://doi.org/10.3366/anh.2000.27.2.209

    Davis, R.E. (2005) Intermediate-depth circulation of the Indian and South Pacific Oceans measured by autonomous floats. Journal of Physical Oceanography, 35, 683–707.
    https://doi.org/10.1175/JPO2702.1

    Farran, G.P. (1908) Second Report on the Copepoda of the Irish Atlantic Slope. Scientific Investigations. Fisheries Branch, Department of Agriculture for Ireland, Appendix II, pp. 19–120, pls 1–11.

    Farran, G.P. (1939) Note on the nomenclature of the copepod genus Megacalanus Wolfenden and allied genera. Annals and Magazine of Natural History, (11) 4 (21), 355–361.
    https://doi.org/10.1080/00222933908527000

    Farris, J.S. (1969) A successive approximation approach to character weighting. Systematic Zoology, 18, 374–385.
    https://doi.org/10.2307/2412182

    Ferrari, F.D. (1995) Six copepodid stages of Ridgewayia klausruetzleri, a new species of copepod crustacean (Ridgewayiidae: Calanoida) from the barrier reef in Belize, with comments on appendage development. Proceedings of the Biological Society of Washington, 108 (2), 180–200.

    Ferrari, F.D. & Dahms, H.-U. 2007. Post-embryonic development of the Copepoda. Crustacean Monographs, 8, 1–229.
    https://doi.org/10.1163/ej.9789004157132.i-230

    Fleminger, A. (1973) Pattern, number, variability, and taxonomic significance of integumental organs (sensilla and glandular pores) in the genus Eucalanus (Copepoda, Calanoida). Fishery Bulletin, 71 (4), 965–1010.

    Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.

    Gelman, A. & Rubin, D.B. (1992) Inference from iterative simulation using multiple sequences. Statistical Science, 457–472.
    https://doi.org/10.1214/ss/1177011136

    Giesbrecht, W. (1888) Mittheilungen über Copepoden 12–14. Mittheilungen aus der Zoologischen Station zu Neapel, 14, 39–42, pls 2–3.

    Goetze, E. (2010) Species discovery in marine planktonic invertebrates through global molecular screening. Molecular Ecology, 19, 952–967.
    https://doi.org/10.1111/j.1365-294X.2009.04520.x

    Goetze, E. & Bradford-Grieve, J. (2005) Genetic and morphological description of Eucalanus spinifer T. Scott, 1894 (Calanoida: Eucalanidae), a circumglobal sister species of the copepod E. hyalinus s.s. (Claus, 1866). Progress in Oceanography, 65, 55–87.
    https://doi.org/10.1016/j.pocean.2005.02.015

    Goetze, E. & Ohman, M.D. (2010) Integrated molecular and morphological biogeography of the calanoid copepod family Eucalanidae. Deep Sea Research Part II: Topical Studies in Oceanography, 57, 2110–2129.
    https://doi.org/10.1016/j.dsr2.2010.09.014

    Gollner, S., Fontaneto, D. & Martínez Arbizu, P. (2011) Molecular taxonomy confirms morphological classification of deep-sea hydrothermal vent copepods (Dirivultidae) and suggests broad physiological tolerance of species and frequent dispersal along ridges. Marine Biology, 158, 221–231.
    https://doi.org/10.1007/s00227-010-1553-y

    Grice, G.D. & Hulsemann, K. (1967) Bathypelagic calanoid copepods of the eastern Indian Ocean. Proceedings of the United States National Museum, 122 (3583), 1–67.
    https://doi.org/10.5479/si.00963801.122-3583.1

    Grice, G.D. & Hulsemann, K. (1968) Calanoid copepods from midwater trawl collections made in the southeastern Pacific Ocean. Pacific Science, 22 (3), 322–335, figs, 1–74.

    Gueredrat, J.-A. (1969) Variations morphologiques de Megacalanus princeps Wolfenden, 1904 (Copepoda, Calanoida). Crustaceana, 17, 64–69, 1 pl.

    Gueredrat, J.-A. & Friess, R. (1971) Importance des migrations nycthémérales de copépodes bathypélagiques. Cahier O.R.S.T.O.M., Série Océanographie, 9 (2), 187–96.

    Hamby, R.K. & Zimmer, E.A. (1988) Ribosomal RNA sequences for inferring phylogeny within the grass family (Poaceae). Plant Systematics and Evolution, 160, 29–37.
    https://doi.org/10.1007/BF00936707

    Hebert, P.D., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences, 270, 313–321.
    https://doi.org/10.1098/rspb.2002.2218

    Hebert, P.D., Stoeckle, M.Y., Zemlak, T.S. & Francis, C.M. (2004) Identification of birds through DNA barcodes. PLoS Biology, 2, E312.
    https://doi.org/10.1371/journal.pbio.0020312

    Heron, G.A. & Bowman, T.E. (1971) Postnaupliar developmental stages of the copepod crustaceas Clausocalanus laticeps, C. brevipes, and Ctenocalanus citer (Calanoida: Pseudocalanidae). Antarctic Research Series, Washington, 17, 141–165.
    https://doi.org/10.1029/AR017p0141

    Herrick, C.L. (1884) Final report on the Crustacea of Minnesota, included in the orders Cladocera and Copepoda, together with a synopsis of the described species in North America, and keys to the known species of the more important genera. Reports of the Geological and Natural History Survey of Minnesota, 12 (5), 1–192, pls. 1–29.
    https://doi.org/10.5962/bhl.title.3981

    Herring, P.J. (1988) Copepod luminescence. Hydrobiologia, 167/168, 183–195.
    https://doi.org/10.1007/BF00026304

    Hillis, D.M. & Dixon, M.T. (1991) Ribosomal DNA: molecular evolution and phylogenetic inference. Quarterly Review of Biology, 66, 411–453.
    https://doi.org/10.1086/417338

    Hirai, J., Shimode, S. & Tsuda, A. (2013) Evaluation of ITS2-28S as a molecular marker for identification of calanoid copepods in the subtropical western North Pacific. Journal of Plankton Research, 35, 644–656.
    https://doi.org/10.1093/plankt/fbt016

    Huntley M.E. & Lopez M.D.G. (1992) Temperature-dependent production of marine copepods: a global synthesis. The American Naturalist 140, 201–242.
    https://doi.org/10.1086/285410

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

    Huys, R., Llewellyn-Hughes, J., Conroy-Dalton, S., Olson, P.D., Spinks, J.N. & Johnston, D.A. (2007) Extraordinary host switching in siphonostomatoid copepods and the demise of the Monstrilloida: Integrating molecular data, ontogeny and antennulary morphology. Molecular Phylogenetics and Evolution, 43, 368–378.
    https://doi.org/10.1016/j.ympev.2007.02.004

    Huys, R., Llewellyn-Hughes, J., Olson, P.D. & Nagasawa, K. (2006) Small subunit rDNA and Bayesian inference reveal Pectenophilus ornatus (Copepoda incertae sedis) as highly transformed Mytilicolidae, and support assignment of Chondracanthidae and Xarifiidae to Lichomolgoidea (Cyclopoida). Biological Journal of the Linnean Society, 87, 403–425.
    https://doi.org/10.1111/j.1095-8312.2005.00579.x

    ICZN, (1999) International Code of Zoological Nomenclature, 4th Edition, 306 pp. London, The International Trust for Zoological Nomenclature. Avaliable from; http://iczn.org/iczn/index.jsp (Accessed 7 Feb. 2017)

    Johnson, M.W. (1958) Bathycalanus sverdrupi, n. sp., a copepod crustacean from great depths in the Pacific Ocean. Proceedings of the California Academy of Sciences, 29 (6), 257–265.

    Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance and Usability. Molecular Biology and Evolution, 30, 772–780.
    https://doi.org/10.1093/molbev/mst010

    Kekkonen, M. Standley, P.D.N. (2014) DNA barcode-based delineation of putative species: efficient start for taxonomic workflows. Molecular Ecology Resources, 14, 706–715.
    https://doi.org/10.1111/1755-0998.12233

    Lawson, T.J. & Grice, G.D. (1973) The developmental stages of Paracalanus crassirostris Dahl, 1894 (Copepoda, Calanoida). Crustaceana, 24 (1), 43–56.
    https://doi.org/10.1163/156854073X00056

    Lenz, P.H., Weatherby, T.M., Weber, W. & Wong, K.K. (1996) Sensory specialization along the first antenna of a calanoid copepod, Pleuromamma xiphias (Crustacea). Marine and Freshwater Behaviour and Physiology, 27 (2–3), 213–221.
    https://doi.org/10.1080/10236249609378966

    Leray, M., Yang, J.Y., Meyer, C.P., Mills, S.C., Agudelo, N., Ranwez, V., Boehm, J.T. & Machida, R.J. (2013) A new versatile primer set targeting a short fragment of the mitochondrial COI region for metabarcoding metazoan diversity: application for characterizing coral reef fish gut contents. Frontiers in Zoology, 10, 43.
    https://doi.org/10.1186/1742-9994-10-34

    Lindeque, P.K., Parry, H.E., Harmer, R.A., Somerfield, P.J. & Atkinson, A. (2013) Next Generation Sequencing reveals the hidden diversity of zooplankton assemblages. PLoS ONE, 8, e81327.
    https://doi.org/10.1371/journal.pone.0081327

    Lindsay, D.J., Grossmann, M.M. & Nishikawa, J.U.N. (2015) DNA barcoding of pelagic cnidarians: current status and future prospects. Bulletin of the Plankton Society of Japan, 62, 39–43.

    Machida, R.J., Kweskin, M. & Knowlton, N. (2012) PCR Primers for Metazoan Mitochondrial 12S Ribosomal DNA Sequences. PLoS ONE, 7, e35887.
    https://doi.org/10.1371/journal.pone.0035887

    Maddison, D.R. & Maddison, W.P. (2000) MacClade 4: Analysis of phylogeny and character Evolution. CD-ROM. Sinauer Associates, Sunderland, Massachusetts.

    Markhaseva, E.L. (1983) In: Brodsky, K.A., Vyshkvatzeva, N.V., Koc, M.C., Markhaseva, E.L. 1983. [Copepod crustaceans (Copepoda: Calanoida) of the seas of SSSR and neighbouring waters.] Opredeliteli po Faune SSSR, Isdavayemyye Zoologicheskij Institutom Akademii Nauk, SSSR, 1, 1–356. [In Russian]

    Marukawa, H. (1921) Plankton List and some new species of copepods from the northern waters of Japan. Bulletin de l’Institut Océanographique, 384, 1–13, 4 pls.

    Mauchline, J. & Gordon, J.D.M. (1991) Oceanic pelagic prey of benthopelagic fish in the benthic boundary layer of a marginal oceanic region. Marine Ecology Progress Series, 74, 109–115.
    https://doi.org/10.3354/meps074109

    McKinnon, A.D. & Arnott, G.H. (1985) The developmental stages of Gladioferens pectinatus (Brady, 1899) (Copepoda: Calanoida). New Zealand Journal of Marine and Freshwater Research, 19, 21–42.
    https://doi.org/10.1080/00288330.1985.9516072

    Michel, H. (1994) Antarctic Megacalanidae (Copepoda: Calanoida) and the distribution of the family. Journal of the Marine Biological Association of the United Kingdom, 74, 175–192.
    https://doi.org/10.1017/S0025315400035748

    Miller, C.B. (2002) A variant form of Megacalanus longicornis (Copepoda: Megacalanidae) from deep waters off Southern California. Hydrobiologia, 480, 129–143.
    https://doi.org/10.1023/A:1021245320351

    Ortman, B.D. (2008) DNA Barcoding the Medusozoa and Ctenophora. Doctoral Dissertations AAI3345202. Avaliable from: http://digitalcommons.uconn.edu/dissertations/AAI3345202 (Accessed 7 Feb. 2017)

    Owre, H. B. & Foyo, M. (1967) Copepods of the Florida Current with illustrated keys to genera and species. Fauna Caribaea 1. Crustacea, Part 1. Copepoda, 1, 1–137.

    Pantin C.F.A. (1964) Notes on microscopical technique for zoologists. Cambridge University Press, 76 pp.

    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

    Rose, M. (1929) Copépodes pélagiques particulièrement de surface provenant des campagnes scientifique du Prince Albert Ier de Monaco. Résultats des Campagne Scientifique accomplies sur son yacht par Prince Albert Ier, 78, 1–123, figs 1–6.

    Sars, G.O. (1900) Crustacea. Scientific Results. Norwegian North Polar Expedition, 1893–1896, 1 (50), 1–141, pls 1–36.

    Sars, G.O. (1905) List préliminaire des Calanoïdés recueillis pendant les campagnes de S.A.S. le Prince Albert de Monaco, avec diagnoses des genres et des espèces nouvelles. Bulletin du Musée Océanographique de Monaco, 26, 1–22.

    Sars, G.O. (1920) Calanoïdés recueillis pendant les campagnes de S.A.S. le Prince Albert de Monaco (Nouveau supplëment). Bulletin de l’Institut Océanographique, 377, 1–20.

    Sars, G.O. (1924, 1925) Copépodes particulièrement bathypélagique provenant des campagnes scientifique du Prince Albert 1er de Monaco. Résultats des Campagnes Scientifiques accompliés par le Prince Albert 1, Monaco, 69, Atlas, 1924, 127 pls; text, 1925, 1–408.

    Schlick-Steiner, B.C., Arthofer, W. & Steiner, F.M. (2014) Take up the challenge! Opportunities for evolution research from resolving conflict in integrative taxonomy. Molecular Ecology, 23, 4192–4194.
    https://doi.org/10.1111/mec.12868

    Schultz, J., Maisel, S., Gerlach, D., Müller, T. & Wolf, M. (2005) A common core of secondary structure of the internal transcribed spacer 2 (ITS2) throughout the Eukaryota. RNA, 11(4), 361–364.
    https://doi.org/10.1261/rna.7204505

    Scott, A. (1909) The copepods of the Siboga Expedition. Part 1. Free-swimming, littoral and semi-parasitic Copepoda. Siboga-Expeditie, 27 (29a), 1–324, 69 pls.

    Scott, T. (1894) Additions to the fauna of the Firth of Forth. Part VI. Annual Report of the Fishery Board for Scotland, Edinburgh, 12 (3), 231–271, pls. 5–10.

    Sewell, R.B.S. (1929) The Copepoda of Indian Seas. Calanoida. Memoir of the Indian Museum, 10, 1–221, figs. 1–21.

    Sewell, R.B.S. (1945) Introduction and list of stations. Scientific Reports. The John Murray Expedition 1933–34, 1, 1–14, 1pl., 1 chart.

    Sewell, R.B.S. (1947) The free-swimming planktonic Copepoda. Scientific Reports. The John Murray Expedition 1933–34, 8, 1–303.

    Soltis, P.S. & Soltis, D.E. (2003) Applying the bootstrap in phylogeny reconstruction. Statistical Science, 18 (2), 256–267.
    https://doi.org/10.1214/ss/1063994980

    Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690.
    https://doi.org/10.1093/bioinformatics/btl446

    Stamatakis, A. (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics, 30, 1312–1313.
    https://doi.org/10.1093/bioinformatics/btu033

    Stocsits, R.R., Letsch, H.. Hertel, J., Misof, B. & Stadler, P.F. (2009) Accurate and efficient reconstruction of deep phylogenies from structured RNAs. Nucleic Acids Research, 37 (18), 6184–6193.
    https://doi.org/10.1093/nar/gkp600

    Swofford, D.L. (2002) Phylogenetic Analysis Using Parsimony. (*and other methods) Version 4. Sinauer Associates, Sunderland, Massachusetts. Available from: http://paup.csit.fsu.edu/Cmd_ref_v2.pdf (Accessed 7 Feb. 2017)

    Tamura, K., Nei, M. & Kumar, S. (2004) Prospects for inferring very large phylogenies by using the neighbor-joining method. Proceedings of the National Academy of Sciences of the United States of America, 101, 11030–11035.
    https://doi.org/10.1073/pnas.0404206101

    Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
    https://doi.org/10.1093/molbev/mst197

    Tanaka, O. (1956) The pelagic copepods of the Izu region, middle Japan. Systematic Account I. Families Calanidae and Eucalanidae. Publications of the Seto Marine Biological Laboratory, 5, 251–272.

    Taniguchi, M., Kanehisa, T., Sawabe, T., Christen, R. & Ikeda, T. (2004) Molecular phylogeny of Neocalanus copepods in the subarctic Pacific Ocean, with notes on non-geographical genetic variations for Neocalanus cristatus. Journal of Plankton Research, 26, 1249–1255.
    https://doi.org/10.1093/plankt/fbh115

    Thompson, J.D., Higgins, D.G. & Gibson, T.J. (1994) CLUSTAL W: improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research, 22, 4673–4680.
    https://doi.org/10.1093/nar/22.22.4673

    Thuesen, E.V., Miller, C.B. & Childress, J.J. (1998) Ecophysiological interpretation of oxygen consumption rates and enzymatic activities of deep-sea copepods. Marine Ecology Progress Series, 168, 95–107.
    https://doi.org/10.3354/meps168095

    Thum, R.A. (2004) Using 18S rDNA to resolve diaptomid copepod (Copepoda: Calanoida: Diaptomidae) phylogeny: an example with the North American genera. Hydrobiologia, 519, 135–141.
    https://doi.org/10.1023/B:HYDR.0000026500.27949.e9

    Toggweiler, J.R. & Key R.M. (2001) Thermohaline circulation. In: Encyclopedia of Ocean Sciences, 2941–2947.
    https://doi.org/10.1006/rwos.2001.0111

    Tomczak, M. & Godfrey, J.S. (1994) Regional Oceanography: An Introduction. Oxford, Elsevier, 422 pp.

    Vervoort, W. (1946) The bathypelagic Copepoda Calanoida of the Snellius Expedition I. Families Calanidae, Eucalanidae, Paracalanidae and Pseudocalanidae. In: Biological Results of the Snellius Expedition, XV. Temminckia, 8, 1–181, figs 1–10.

    Vervoort, W. (1949) Some new and rare Copepoda Calanoida from East Indian Seas. Zoologische Verhandelingen, Leiden, 5, 1–53.

    Vervoort, W. (1957) Copepods from Antarctic and subantarctic plankton samples. Report of the British, Australian and New Zealand Antarctic Research Expedition 1929–31, series B3, 1–160, Figs 1–138.

    Vinogradov, M.E. (1968) Vertical distribution of the oceanic zooplankton. Nauka, Moscow. 320 pp.

    Vinogradov, M.E. & Tseiltin, V.B. (1983) Deep-sea pelagic domain (aspects of bioenergetics). In: Rowe, G.T. (Ed.), The Sea, 8, 123–140.

    Wang, X.-C., Liu, C., Huang, L., Bengtsson-Palme, J., Chen, H., Zhang, J.-H., Cai, D.& Li, J.-Q. (2015) ITS1: a DNA barcode better than ITS2 in eukaryotes? Molecular Ecology Resourses, 15, 573–586.
    https://doi.org/10.1111/1755-0998.12325

    Watling, L., Guinotte, J., Clark, M.R. & Smith, C.R. (2013) A proposed biogeography of the deep ocean floor. Progress in Oceanography, 111, 91–112.
    https://doi.org/10.1016/j.pocean.2012.11.003

    White, T.J., Bruns, T., Lee, S. & Taylor, J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. Chapter 38. In: Innis, M., Gelfand, D., Sninsky J. & White, T. (Eds.), PCR protocols: a guide to methods and applications, 18: 315–322. Academic Press, Orlando, Florida.
    https://doi.org/10.1016/b978-0-12-372180-8.50042-1

    Wiebe, P.H., Burt, K.H., Boyd, S.H. & Morton, A.W. (1976) A multiple opening/closing and environmental sensing system for sampling zooplankton. Journal of Marine Research, 34, 313–326.

    Wiebe, P.H., Bucklin, A., Madin, L., Angel, M.V., Sutton, T., Pagés, F., Hopcroft, R.R. & Lindsay, D. (2010) Deep-Sea sampling on CMarZ curises in the Atlantic Ocean—an introduction. Deep-Sea Research II, 57, 2157–2166.
    https://doi.org/10.1016/j.dsr2.2010.09.018

    With, C. (1915) Copepoda I. The Danish Ingolf-Expedition. Vol III, Part 4, 1–260, pls I–VIII.

    Wolf, M. (2015) ITS so much more. Trends in Genetics, 31 (4), 175–176.
    https://doi.org/10.1016/j.tig.2015.02.005

    Wolfenden, R.N. (1904) Notes on the Copepoda of the North Atlantic Sea and the Faröe Channel. Journal of the Marine Biological Association of the United Kingdom, 7 (1), 110–146, pl. IX.
    https://doi.org/10.1017/S0025315400072805

    Wolfenden, R.N. (1905a) Plankton Studies: preliminary notes upon new or interesting species. Part 1. Copepoda [original version, May 1905] Rebman, London, pp. 1–24, pls I–VII.

    Wolfenden, R.N. (1905b) Plankton Studies: preliminary notes upon new or interesting species. Part 1. Copepoda [amended version, 13 November 1905] Rebman, London, pp 1–24, pls I–VII.

    Wolfenden, R.N. (1906) Plankton Studies: preliminary notes upon new or interesting species. Part 1I. Copepoda. Rebman, London, pp. 25–44, pls VIII–XIV.

    Wolfenden, R.N. (1911) Die marinen Copepoden II. Die pelagischen Copepoden der Westwinddrift und des südlichen Eismeers mit Beschreibung mehrer neuer Arten aus dem Atlantischen Ozean. Deutsche Südpolar-Expedition 1901–1903, 12 (Zoology 4), 181–380, pls XXII–XLI.

    Wyngaard, G.A., Hołyńska, M. & Schulte J.A. (2010) Phylogeny of the freshwater copepod Mesocyclops (Crustacea: Cyclopidae) based on combined molecular and morphological data, with notes on biogeography. Molecular Phylogenetics and Evolution, 55, 753–764.
    https://doi.org/10.1016/j.ympev.2010.02.029

    Zagoskin, M.V., Lazareva, V.I., Grishanin, A.K. & Mukha, D.V. (2014) Phylogenetic Information Content of Copepoda Ribosomal DNA Repeat Units: ITS1 and ITS2 Impact. BioMed Research International, 2014, 926342.
    https://doi.org/10.1155/2014/926342