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Published: 2023-12-29
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The taxonominal status of the nomina Lesueuria Milne Edwards, 1841 and Lesueuriidae Chun, 1880, and introduction of a new genus and a new family for Lesueuria pinnata Ralph & Kaberry, 1950, as well as an additional new species of the new genus (Ctenophora, Lobata)

Friday Harbor Laboratories and Department of Biology; University of Washington; 620 University Road; Friday Harbor; Washington 98250; U.S.A.
Institut de Systématique; Évolution; Biodiversité; ISYEB - UMR 7205 - CNRS; MNHN; UPMC; EPHE; Muséum national d’Histoire naturelle; Sorbonne Université; 57 rue Cuvier; CP 30; 75005; Paris; France
Lesueuria Lesueuriidae Bolina Bolinopsis Bolinopsidae onomatophore type specimen type species nomen dubium nomenclatural stability new species new genus new family


The nomenclatural problems posed by the nomen of the ctenophore species Lesueuria vitrea Milne Edwards, 1841, type species of the genus Lesueuria Milne Edwards, 1841 which is type genus of the family Lesueuriidae Chun, 1880, and by a few other species long referred to this genus and family, are addressed, and new solutions are proposed for some of them. For the species Lesueuria pinnata Ralph & Kaberry, 1950, an unusual ‘finned’ lobate ctenophore from the southern Pacific Ocean, a new genus and a new family are introduced. It is shown that the nomenclatural problems discussed in this paper raise difficulties for four distinct reasons, related to four different basic concepts of zoological nomenclature: those of ‘name-bearing type’ or onomatophores, of ‘type species’ of genera or nucleospecies, of availability, validity and nomen dubium, and of ‘nomenclatural stability’. The fact that specimens of this group are ‘fragile’ and difficult or impossible to fix and keep in collections requires recourse to indirect methods (detailed description, good iconography and molecular sequencing) applied to ‘ephemeral’ type specimens (holotypes or neotypes) to objectify and stabilize the nomenclature in this group.


  1. Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological Nomenclature. ‘Fourth edition’. London (International Trust for zoological Nomenclature): i–xxix + 1–306.
  2. Anonymous [International Commission on Zoological Nomenclature] (2012) Amendment of Articles 8, 9, 10, 21 and 78 of the International Code of Zoological nomenclature to expand and refine methods of publication. Bulletin of zoological Nomenclature, 69 (3): 161‒169. <>.
  3. Anonymous [International Commission on Zoological Nomenclature] (2017) Declaration 45. Addition of Recommendations to Article 73 and of the term ‘specimen, preserved’ to the Glossary. Bulletin of zoological Nomenclature, 73 (2‒4): 96‒97. <>.
  4. Anonymous (2021) Lesueuria pinnata. <>; <>.
  5. Anonymous [WoRMS Editorial Board] (2022) World Register of Marine Species. <> at VLIZ.
  6. Agassiz, A. (1865) Illustrated catalogue of the Museum of comparative Zoology, at Harvard College. No. 2. North American Acalephae. Cambridge (Sever & Francis): i‒xiv + 1–234. <>.
  7. Agassiz, L. (1850) Contributions to the natural history of the Acalephae of North America. Part II. On the beroid medusae of the shores of Massachusetts, in their perfect state of development. Memoirs of the American Academy of Arts and Sciences, 4: 313‒374, pl. 6‒8. <>.
  8. Agassiz, L. (1860) Contributions to the natural history of the United States of America. Second monograph, Vol. 3. Boston (Little, Brown & Co.) & London (Trübner & Co.): i‒xi + 1‒301 + 1‒26, pl. 1‒19.
  9. Bigelow, H. B. (1904) Medusae from the Maldive Islands. Bulletin of the Museum of comparative Zoology at Harvard College, 39 (9): 245–269.
  10. Bigelow, H. B. (1912) Reports on the scientific results of the expedition to the eastern tropical Pacific, in charge of Alexander Agassiz, by the U. S. Fish Commission Steamer ‘Albatross’, from October, 1904, to March, 1905, Lieut. Commander L. M. Garrett, U. S. N., commanding. XXVI. The ctenophores. Bulletin of the Museum of comparative Zoology at Harvard College, 54 (12): 369–404, pl. 1–2.
  11. Brünnichius, M. T. (1772) Zoologiae fundamenta, praelectionibus academicis accommodata. Hafniae et Lipsiae (Frider, Christ, Pelt): [i‒iii] + 1‒254. <>.
  12. Carus, J. V. (1885) 2. Class. Ctenophorae. In: Prodromus Faunae Mediterraneae, Volume 1, Stuttgart (E. Schweizerbart’sche Verhandlung, E. Koch): 54–57.
  13. Carus, J. V. & Gerstaecker, C. E. A. (1863) II. Classe. Ctenophorae Eschsch., Rippenquallen. In: Handbuch der Zoologie, Volume 2, Leipzig (Wilhelm Engelmann): 537–542.
  14. Ceríaco, L. M. P., Gutiérrez, E. E., Dubois, A. et al. [490 signatories] (2016) Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences. Zootaxa, 4196 (3): 435–445. <>.
  15. Christianson, L. M., Johnson, S. B., Schultz, D. T., & Haddock, S. H. D. (2022) Hidden diversity of Ctenophora revealed by new mitochondrial COI primers and sequences. Molecular Ecology Resources, 22: 283–294. .
  16. Chun, C. (1879) Die im Golf von Neapel erscheinenden Rippenquallen. Mittheilungen aus der zoologischen Station zu Neapal, 1: 180–217, pl. 6.
  17. Chun, C. (1880) Die Ctenophoren des Golfes von Neapel und der angrenzenden Meeres-abschnitte. In: Fauna und Flora des Golfes von Neapel, und der angrenzenden Meeres-abschnitte herausgegeben von der zoologischen Station in Neapel, Leipzig (Wilhelm Engelmann): i‒xviii + 1‒314, pl. 1‒18. <>.
  18. Chun, C. (1898) Die Ctenophoren der Plankton-Expedition. Ergebnisse der Plankton-Expedition der Humboldt-Stiftung, 2 K.a.: 1– 31, pl. 1–3.
  19. Dawydoff, C. (1946) Contribution a la connaissance des cténophores pélagiques des eaux de l’Indochine. Bulletin biologique de la France et de la Belgique, 80 (2): 116–170.
  20. Dayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85: 407–415. <>.
  21. Dickinson, E. C., Overstreet, L. K., Dowsett, R. J. & Bruce, M. D. (2011) Priority! The dating of scientific names in ornithology: a directory to the literature and its reviewers. Northampton, U. K. (Aves Press): 1–320.
  22. Dubois, A. (2000) Synonymies and related lists in zoology: general proposals, with examples in herpetology. Dumerilia, 4 (2): 33‒98.
  23. Dubois, A. (2005) Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code of Zoological Nomenclature. 1. Some general questions, concepts and terms of biological nomenclature. Zoosystema, 27 (2): 365–426. <>.
  24. Dubois, A. (2009) Endangered species and endangered knowledge. Zootaxa, 2201: 26‒29. <>.
  25. Dubois, A. (2011) The International Code of Zoological Nomenclature must be drastically improved before it is too late. Bionomina, 2: 1–104. <>.
  26. Dubois, A. (2017a) The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related nomina (Amphibia, Anura), with general comments on zoological nomenclature and its governance, as well as on taxonomic databases and websites. Bionomina, 11: 1–48. <>.
  27. Dubois, A. (2017b) The need for reference specimens in zoological taxonomy and nomenclature. Bionomina, 12: 4–38. <>.
  28. Dubois, A. (2020) Allocation of nomina to taxa in zoological nomenclature. Bionomina, 18: 1‒43. <>.
  29. Dubois, A. (2021) Recommendations concerning the presentation of synonymic and related lists in zootaxonomy: the example of the family Cryptobranchidae Fitzinger, 1826 (Amphibia, Urodela). Bionomina, 25: 52‒69. <>.
  30. Dubois, A. (2022) Type species of genera in zoological nomenclature. Bionomina, 26: 1‒33. <>.
  31. Dubois, A., Bauer, A. M., Ceríaco, L. M. P., Dusoulier, F., Frétey, T., Löbl, I., Lorvelec, O., Ohler, A., Stopiglia, R. & Aescht, E. (2019) The Linz Zoocode project: a set of new proposals regarding the terminology, the Principles and Rules of zoological nomenclature. First report of activities (2014‒2019). Bionomina, 17: 1‒111. <>.
  32. Dubois, A., Frétey, T. & Ohler, A. (2022) The nomenclature of Eurasian giant salamanders (Amphibia, Urodela, Andrias). Bionomina, 31: 70‒87. <>.
  33. Dubois, A. & Nemésio, A. (2007) Does nomenclatural availability of nomina of new species or subspecies require the deposition of vouchers in collections? Zootaxa, 1409: 1–22. <>.
  34. Dubois, A. & Ohler, A. (1997) Early scientific names of Amphibia Anura. I. Introduction. Bulletin du Muséum national d’Histoire naturelle, (4), 18 (3–4): 297–320. <>.
  35. Dubois, A. & Ohler, A. (2018) The Hyla quoyi-Hyla prasina case (Amphibia, Anura), with comments on bibliographic and taxonomic databases and on Article 23.9 of the Code. Zoosystema, 40 (23): 501‒506. <>.
  36. Dubois, A., Ohler, A. & Pyron, R. A. (2021) New concepts and methods for phylogenetic taxonomy and nomenclature in zoology, exemplified by a new ranked cladonomy of recent amphibians (Lissamphibia). Megataxa, 5 (1): 1‒738. <>.
  37. Dufay, P. (2009) Le roman du Jardin du Roy. Monaco (Editions du Rocher): 1‒218.
  38. Eschscholtz, F. (1825) Bericht über die zoologische Ausbeute während der Reise von Kronstadt bis St. Peter und Paul. Tafel V. Isis von Oken, 16 (6): 733–747, pl. 5.
  39. Eschscholtz, F. (1829) System der Acalephen. Eine ausführliche Beschreibung aller Medusenartigen Strahlthiere. Berlin (Ferdinand Dümmler): i‒iv + 1‒190, pl. 1‒16. <>.
  40. Frétey, T. (2022) The authorship and date of the familial nomen Dermochelyidae (Chelonii). Bionomina, 31: 46‒69. <>.
  41. Frétey, T., Dewynter, M. & Ohler, A. (2018) Onymotopes in zoological nomenclature: some additional terms, with fixation of a lectonymotope for Xenopus petersii Bocage, 1895 (Amphibia, Anura). Bionomina, 13: 37–50. <>.
  42. Garraffoni, A. R. S. & Freitas, A. V. L. (2017) Photos belong in the taxonomic Code. Science, 355 (6327): 805. <>.
  43. Gegenbaur, C. (1856) Studien über Organisation un Systematik der Ctenophoren. Archiv für Naturgeschichte, 22 (1): 163–205, pl. 7‒8.
  44. Gershwin, L., Zeidler, W. & Davie, P. J. F. (2010) Ctenophora of Australia. Memoirs of the Queensland Museum–Nature, 54 (3): 1–45.
  45. Johnson, S. B., Winnikoff , J. R., Schultz, D. T., Christianson, L. M., Patry, W. L., Mills, C. E. & Haddock, S. H. D. (2022) Speciation of pelagic zooplankton: invisible boundaries can drive isolation of oceanic ctenophores. Frontiers in Genetics, 13 [970314]: 1–17. <>.
  46. Komai, T. (1918) On ctenophores of the neighborhood of Misaki. Annotationes zoologicae japonensis, 9: 451–474, pl. 7.
  47. Krell, F.-T., Klimeš, P., Rocha, L. A., Fikaček, M. & Miller, S. E. (2016) Preserve specimens for reproducibility. Nature, 539: 168. <>.
  48. Laval, P., Braconnot, J.-C., Carré, C., Goy, J., Morand, P. & Mills, C. E. (1989) Small-scale distribution of macroplankton and micronekton in the Ligurian Sea (Mediterranean Sea) as observed from the manned submersible Cyana. Journal of Plankton Research, 11: 665–685. <>.
  49. Lendenfeld, R. von (1885) The metamorphosis of Bolina chuni Nov. Spec. Proceedings of the Linnean Society of New South Wales, 9 (4): 929–931, pl. 44‒45. <>.
  50. Lesson, R. P. (1843) Histoire naturelle des Zoophytes. Acalèphes. Paris (Roret): i‒viii + 1‒596 + [i‒ii], pl. 1‒12. <>.
  51. Lučić, D., Pestorić, B., Malej, A., Lopez-Lopez, L., Drakulović, D., Onofri, V., Miloslavić, M., Gangai, B., Onofri, I., & Benović, A. (2012) Mass occurrence of the ctenophore Bolinopsis vitrea (L. Agassiz, 1860) in the nearshore southern Adriatic Sea (Kotor Bay, Montenegro). Environmental Monitoring and Assessment, 184: 4777–4785. <>.
  52. Madin, L. P & Harbison, G. R. (1978) Bathocyroe fosteri gen nov., sp. nov.: a mesopelagic ctenophore observed and collected from a submersible. Journal of the biological Association of the United Kingdom, 58: 559–564. <>.
  53. Matsumoto, G. I. (1988) A new species of lobate ctenophore, Leucothea pulchra sp. nov., from the California Bight. Journal of Plankton Research, 10: 301–311. <>.
  54. Mayer, A. G. (1912) Ctenophores of the Atlantic coast of North America. Washington D.C. (Carnegie Institution), Publication No. 162: [i‒ii] + 1‒58, pl. 1‒17. <>.
  55. Mertens, H. (1833) Beobachtungen und Untersuchungen über die Beroartigen Akalephen. Mémoires de l’Académie impériale des Sciences de St. Pétersbourg, (6), 2 (5): 479‒543, pl. 1‒13.
  56. Mianzan, H., Dawson, E. W. & Mills, C. E. (2009) Phylum Ctenophora: comb jellies. In: D. P. Gordon (ed.), New Zealand inventory of biodiversity, Volume 1, Kingdom Animalia: Radiata, Lophotrochozoa, and Deuterostomia, Christchurch (Canterbury University Press): 49–58.
  57. Mills, C. E. (1995) Ctenophora. In: A. Minelli, S. Ruffo & La Posta. S. (ed.), Checklist delle specie della fauna italiana, Vol. 3, Bologna (Edizioni Calderini): 32–38.
  58. Mills, C. E. (2022) Phylum Ctenophora: list of all valid species names. <>. [Web page established in March 1998, last updated on 12 June 2017; accessed 3 May 2022].
  59. Mills, C. E., Haddock, S. H. D. & Bezio, N. (2023) Chapter 5: Kingdom Animalia, phylum Ctenophora (comb jellies). In: M. Kelly, S. Mills, M. Terezow, C. Sim-Smith & W. Nelson (ed.), The marine biodiversity of Aotearoa New Zealand, Updating our marine biodiversity, NIWA Biodiversity Memoir 136: 111–129, 3 pl.
  60. Mills, C. E., Pugh, P. R., Harbison, G. R. & Haddock, S. H. D. (1996) Medusae, siphonophores and ctenophores of the Alborán Sea, south western Mediterranean. Scientia marina, 60: 145–163.
  61. Milne Edwards, M. H. (1841) Observations sur la structure et les fonctions de quelques Zoophytes, Mollusques et Crustacés des côtes de la France. Annales des Sciences naturelles, (2), 16 (Zoologie): 193–232, pl. 1‒10.
  62. M’Intosh, W. (1888) LXI. Notes from the St. Andrews Marine Laboratory (under the Fishery Board for Scotland). No. IX. 1. On Lesueuria vitrea, M.-Edwards. Annals and Magazine of natural History, (6), 2: 464–466. <>.
  63. M’Intosh, W. (1890) VI. Notes from the St. Andrews Marine Laboratory (under the Fishery Board for Scotland). No. X. 2. On the occurrence of the ctenophores throughout the year. Annals and Magazine of natural History, (6), 5: 43–47. <>.
  64. Mortensen, T. (1912) Ctenophora. The Danish Ingolf-Expedition. Volume 5. Copenhagen (Bianco Luno): 1–97, pl. 1–3.
  65. Moser, F. (1907) Neues über Ctenophoren. Mitteilung II. Zoologischer Anzeiger, 32: 449–454.
  66. Moser, F. (1908) Beiträge zur Naturgeschichte Ostasiens: Japanische Ctenophoren. Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie der Wissenschaften, Supplement-Band 1, 4: 1–78, pl. 1‒2.
  67. Moser, F. (1909) Die Ctenophoren der Deutschen Südpolar-Expedition 1901–1903. Deutschen Südpolar-Expedition, 2 (Zoologie 3): 116–192, pl. 20–22.
  68. Müller, O. F. (1776) Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Havniae (Typis Hallageriis): i‒xxxii + 1‒282. <>.
  69. Pape, T., Allison, A., Bickel, D. J., Carlton, J. T., Dikow, T., Donegan, T., Duszynski, D. W., El-Hawagry, M. S., Evenhuis, N. L., Fautin, D. G., Gaimari, S. D., Gharali, B., Greenwalt, D. E., Kaiser, H., Kirk-Spriggs, A. H., Lamas, G., Lonsdale, O., Mah, C., Marshall, S. A., Meier, R., Ohl, M., Patterson, D. J., Penev, L., Pentcheff, N. D., Pyle, R. L., Rubinoff, D. J., Runyon, J., Tallowin, O., Thorpe, S., Wang, B., Welter-Schultes, F., Yanega, D., Yang, D., Yao, G. & Yeung, N. W. (2016) Species can be named from photos. Nature, 537: 307. <>.
  70. Poplin, F. (1981) Sur deux autographes présumés de Buffon et de Daubenton dans la grande grotte d’Arcy-sur-Cure et sur l’os de girafe du cabinet de Gaston d’Orléans. In: Association bouguignonne des sociétés savantes, Cinquante et unième congrès, Montbard, 6, 7, 8 juin 1980, Actes du congrès de Montbard, Montbard (Association des Amis de la Cité de Montbard): 99‒105.
  71. Quoy [J. R. C.] & Gaimard [J. P.] (1824) Zoologie. In: L. de Freycinet (ed.), Voyage autour du monde, entrepris par ordre du Roi, sous le Ministère et conformément aux instructions de S. Exc. le Vicomte du Bouchage, Secrétaire d’État au Département de la Marine, exécuté sur les corvettes de S. M. l’Uranie et la Physicienne, pendant les années 1817, 1818, 1819 et 1820..., Paris (Pillet Aîné): [i‒vii] + 1‒712. <>.
  72. Rafinesque, C. S. (1815) Analyse de la nature ou tableau de l’univers et des corps organisés. Palerme (Jean Barravecchia): 1–124, 1 pl. <>.
  73. Ralph, P. M. & Kaberry, C. (1950) New Zealand Coelenterates: Ctenophores from Cook Strait. Zoology Publications from Victoria University College, 3: 1–11.
  74. Sars, M. (1835) Beskrivelser og iagttagelser over nogle mærkelige eller nye i havet ved den bergenske kyst levende dyr af Polypernes, Acalephernes, Radiaternes, Annelidernes og Molluskernes classer, med en kort oversigt over de hidtil af forfatteren sammesteds fundne arter og deres forekommen. Bergen (Thorstein Hallagers): i–xii + 1–81, pl. 1‒15. <>.
  75. Sars, M. (1857) Bidrag til kündskaben om middelhavets littoral-fauna: reisebemaerknonger fra Italien. Christiania (Johan Dahls Forlag): 57–155, 2 pl.
  76. Schiariti, A., Dutto, M. S., Oliveira, O. M., Siquier, G. F., Puente Tapia, F. A. & Chiaverano, L. (2021) Overview of the comb jellies (Ctenophora) from the south-western Atlantic and sub Antarctic region (32–60°S; 34–70°W). New Zealand Journal of marine and freshwater Research, 55: 286–310. <>.
  77. Shiganova, T. & Malej, A. (2009) Native and non-native ctenophores in the gulf of Trieste, northern Adriatic sea. Journal of Plankton Research, 31: 61–71. <>.
  78. Spagnolini, A. (1870) Di una specie del gen. Bolina Mertens nel Golfo di Napoli. Bullettino dell’ Associazione dei Naturalisti e Medici di Napoli, 1 (4): 61–64, pl. 1.
  79. Tokioka, T. (1964) Bolinopsis rubripunctata n. sp., a new lobatean ctenophore from Seto. Publications of the Seto marine biological Laboratory, 12 (1): 93–99. <>.
  80. Trégouboff, G. (1958) Prospection biologique sous-marine dans la région de Villefranche-sur-Mer au cours de l’année 1957. I. Plongées en bathyscaphe. Bulletin de l’Institut océanographique, 1117: 1–37.
  81. Trégouboff, G. & Rose, M. (1957) Manuel de planctonologie méditerranéenne. Tome 1. Texte. Paris (Centre National de la Recherche Scientifique): 1‒587.
  82. Vanhöffen, E. (1895) Zoologische Ergebnisse der von der Gesellshaft für Erdkunde zu Berlin unter Leitung Dr. von Drygalski’s ausgesandten Grönland-expedition, nach Dr. Vanhöffen’s Sammlungen bearbeitet. II. Die grönländischen Ctenophoren. Bibliotheca zoologica, Stuttgart, 20 [Not seen].
  83. Vanhöffen, E. (1903) Ctenophoren. Nordisches Plankton, 6 (11): 1–7.
  84. Whelan, N. V., Kocot, K. M., Moroz, T. P., Mukherjee, K., Williams, P., Paulay, G., Moroz, L. L. & Halanych, K. M. (2017) Ctenophore relationships and their placement as the sister group to all other animals. Nature Ecology & Evolution, 1: 1737–1746. <>.
  85. Winnikoff, J. R., Francis, W. R., Thuesen, E. V. & Haddock, S. H. D. (2019) Combing transcriptomes for secrets of deep-sea survival: environmental diversity drives patterns of protein evolution. Integrative and comparative Biology, 59 (4): 786–798. <>.
  86. Winnikoff, J. R., Haddock, S. H. D. & Budin, I. (2021) Depth- and temperature-specific fatty acid adaptation in ctenophores from extreme habitats. Journal of experimental Biology, 224 (21) [jeb242800]: 1–10. <>.