A New Integrated Morpho- and Molecular Systematic Classification of Cenozoic Radiolarians (Class Polycystinea) – Suprageneric Taxonomy and Logical Nomenclatorial Acts

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A New Integrated Morpho- and Molecular Systematic Classification of Cenozoic Radiolarians (Class Polycystinea) – Suprageneric Taxonomy and Logical Nomenclatorial Acts geodiversitas 2021 43 15 Families DIRECTEUR DE LA PUBLICATION / PUBLICATION DIRECTOR : Bruno David, Président du Muséum national d’Histoire naturelle RÉDACTEUR EN CHEF / EDITOR-IN-CHIEF : Didier Merle ASSISTANT DE RÉDACTION / ASSISTANT EDITOR : Emmanuel Côtez ([email protected]) MISE EN PAGE / PAGE LAYOUT : Emmanuel Côtez COMITÉ SCIENTIFIQUE / SCIENTIFIC BOARD : Christine Argot (Muséum national d’Histoire naturelle, Paris) Beatrix Azanza (Museo Nacional de Ciencias Naturales, Madrid) Raymond L. Bernor (Howard University, Washington DC) Alain Blieck (chercheur CNRS retraité, Haubourdin) Henning Blom (Uppsala University) Jean Broutin (Sorbonne Université, Paris, retraité) Gaël Clément (Muséum national d’Histoire naturelle, Paris) Ted Daeschler (Academy of Natural Sciences, Philadelphie) Bruno David (Muséum national d’Histoire naturelle, Paris) Gregory D. Edgecombe (The Natural History Museum, Londres) Ursula Göhlich (Natural History Museum Vienna) Jin Meng (American Museum of Natural History, New York) Brigitte Meyer-Berthaud (CIRAD, Montpellier) Zhu Min (Chinese Academy of Sciences, Pékin) Isabelle Rouget (Muséum national d’Histoire naturelle, Paris) Sevket Sen (Muséum national d’Histoire naturelle, Paris, retraité) Stanislav Štamberg (Museum of Eastern Bohemia, Hradec Králové) Paul Taylor (The Natural History Museum, Londres, retraité) COUVERTURE / COVER : Réalisée à partir des Figures de l’article/Made from the Figures of the article. Geodiversitas est indexé dans / Geodiversitas is indexed in: – Science Citation Index Expanded (SciSearch®) – ISI Alerting Services® – Current Contents® / Physical, Chemical, and Earth Sciences® – Scopus® Geodiversitas est distribué en version électronique par / Geodiversitas is distributed electronically by: – BioOne® (http://www.bioone.org) Les articles ainsi que les nouveautés nomenclaturales publiés dans Geodiversitas sont référencés par / Articles and nomenclatural novelties published in Geodiversitas are referenced by: – ZooBank® (http://zoobank.org) Geodiversitas est une revue en flux continu publiée par les Publications scientifiquesdu Muséum, Paris Geodiversitas is a fast track journal published by the Museum Science Press, Paris Les Publications scientifiques du Muséum publient aussi / The Museum Science Press also publish: Adansonia, Zoosystema, Anthropozoologica, European Journal of Taxonomy, Naturae, Cryptogamie sous-sections Algologie, Bryologie, Mycologie, Comptes Rendus Palevol Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2021 ISSN (imprimé / print) : 1280-9659/ ISSN (électronique / electronic) : 1638-9395 A new integrated morpho- and molecular systematic classification of Cenozoic radiolarians (Class Polycystinea) – suprageneric taxonomy and logical nomenclatorial acts Noritoshi SUZUKI Department of Earth Science Graduate School of Science, Tohoku University 6-3 Aoba Aramaki, Aoba-ku Sendai City, Miyagi 980-8578 (Japan) [email protected] Luis O’DOGHERTY Instituto Universitario de Investigación Marina (INMAR) Facultad de Ciencias del Mar, Universidad de Cádiz 11510 Puerto Real (Spain) [email protected] Jean-Pierre CAULET 242 rue de la Fure, F-38850 Charavines (France) [email protected] Paulian DUMITRICA Institute of Earth Sciences, Université de Lausanne Géopolis, 1015 Lausanne (Switzerland) [email protected] Submitted on 30 April 2020 | accepted on 11 March 2021 | published on 8 July 2021 urn:lsid:zoobank.org:pub:DC259A19-9B35-4B33-AD9F-44F4E1DA9983 Suzuki N., O’Dogherty L., Caulet J.-P. & Dumitrica P. 2021. — A new integrated morpho- and molecular systematic classi- fication of Cenozoic radiolarians (Class Polycystinea) – suprageneric taxonomy,in O’Dogherty L. (ed.), Catalog of Cenozoic radiolarians. Geodiversitas 43 (15): 405-573. https://doi.org/10.5252/geodiversitas2021v43a15. http://geodiversitas.com/43/15 ABSTRACT A revised taxonomy of Cenozoic radiolarian families is of particular importance because exhaustive molecu- lar phylogenetic analyses for Collodaria, Entactinaria, Nassellaria and Spumellaria have shown high level of confidence at family or higher taxonomic ranks. In this sense, this study presents a new comprehensive taxonomy at the family level that integrated a classification based on ribosomal taxonomic marker genes (rDNA) and classical morphological taxonomy. However, many family names commonly used in Cenozoic radiolarians (Polycystinea) are derived from genera whose type species were never illustrated at the time of KEY WORDS the generic definition. Obviously, in the vast majority of those cases, the “Principle of Typification” regulated Cenozoic, in the International Code of Zoological Nomenclature (ICZN 1999: Art. 61) cannot be logically applied. Radiolaria, This has contributed to a century-long misunderstanding about the validity of Cenozoic taxa (species, genera Molecular taxonomy, Morphological and/or family-group names) erected without any illustration or drawing of their types, in particular the huge taxonomy, contribution of Ernst Haeckel from samples of the Challenger expedition (1872-1876). Reexamination of Taxonomic revision, Haeckel’s collection definitively confirmed that all the original types series (the specimens on which Haeckel new synonyms, new status, established the nominal species-group taxon) being nonextant; in other words, all name-bearing specimens new families. (the types) are restricted to the illustrations given in Haeckel’s drawings. Because “types” in taxonomy are GEODIVERSITAS • 2021 • 43 (15) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 405 Suzuki N. et al. precious things, a nominal species-group taxon lacking at all of illustration (or indication to a repository) do not ensure the recognition of the species. Following the rules and recommendations of the ICZN, these names should be excluded from all nomenclatorial and taxonomical acts. This revision presents the state of the art of all proposed family-group names (with full synonymy lists) for Cenozoic Polycystinea. The list of family-group nominal taxa and their names was inventoried from 6694 publications (89% of the whole known references on radiolarians). The references were examined in order to clarify and fix the status of family names; hence these family-group names were rigidly classified as: valid, junior synonym, nomen dubium, nomen nudum, homonym, and invalid names. A total of 372 family-group names were proposed for the Cenozoic. These consist of 94 valid family-groups, 118 junior synonym family-groups, 111 nomen dubium family-groups (mainly artificially created in a hypothetical conceptual framework), 6 junior homonym family-groups, 19 nomen nudum family-groups, as well as 24 invalid names. In addition, one nomen novum et four new families are presented. The description of 25 families have been also emended. This study also outlines the advantages of an integrated approach to taxonomy of Polycystinea by the combination of both morphological and molecular systematics. Based on molecular phylogenetic studies, the systematic classification proposed at suprageneric level is arranged as follows: a) Order Spumellaria: three Phylogenetic Molecular Lineages (PM Lineages = suborders), 13 su- perfamilies and 42 families; b) Order Entactinaria: one PM Lineage, five superfamilies and nine families; c) Order Nassellaria: four PM Lineages, 16 superfamilies and 37 families; d) Order Collodaria: three superfamilies and six families. RÉSUMÉ Une nouvelle classification systématique intégrée basée sur la phylogénèse moléculaire et le classement morphologique des radiolaires du Cénozoïque (Classe des Polycystinea) – taxonomie supragénérique et actes logiques de nomenclature. Une révision de la taxonomie des familles de radiolaires du Cénozoïque est particulièrement importante, car de nouvelles analyses phylogénétiques moléculaires pour Collodaria, Entactinaria, Nassellaria et Spu- mellaria ont montré d’excellents résultats pour les rangs taxonomiques familiaux ou supérieurs. En ce sens, cette étude présente une nouvelle taxonomie complète au niveau familial, qui intègre une classification fondée sur les gènes marqueurs taxonomiques ribosomiques (ADNr) et la taxonomie classique fondée sur des caractéristiques morphologiques. De plus, de nombreux noms de familles communément utilisés pour les radiolaires polycystines du Cénozoïque dérivent de genres dont les espèces types n’ont jamais été illustrées au moment de la définition du genre. Apparemment, dans la plupart de ces cas, le « Principe de Typification » défini dans le Code international de Nomenclature zoologique (ICZN 1999: Art. 61) ne peut être logiquement utilisé. Cela a généré une longue incompréhension (un siècle) quant à la validité des taxa cénozoïques (noms d’espèces, genres, et/ou familles) érigés à partir de types non illustrés ou dessinés, en particulier dans l’immense travail de Ernst Haeckel sur les échantillons récoltés par l’Expédition du « Challenger » (1872-1876). Le réexamen de la collection d’Haeckel a définitivement confirmé que tous les originaux des espèces types d’Haeckel (les spécimens à partir desquels Haeckel a établi les taxa des groupes d’espèces) n’existent pas, en d’autres termes que les spécimen-types sont réduits aux illustrations des planches dessinées
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