{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,14]],"date-time":"2026-05-14T20:27:09Z","timestamp":1778790429233,"version":"3.51.4"},"reference-count":122,"publisher":"Geological Society of London","issue":"4","license":[{"start":{"date-parts":[[2022,3,10]],"date-time":"2022-03-10T00:00:00Z","timestamp":1646870400000},"content-version":"unspecified","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.lyellcollection.org"],"crossmark-restriction":true},"short-container-title":["JGS"],"published-print":{"date-parts":[[2022,7]]},"abstract":"<jats:p>\n            Ecdysozoans (Phyla Arthropoda, Kinorhyncha, Loricifera, Nematoda, Nematomorpha, Onychophora, Priapulida, Tardigrada) are invertebrates bearing a tough, periodically moulted cuticle that predisposes them to exceptional preservation. Ecdysozoans dominate the oldest exceptionally preserved bilaterian animal biotas in the early to mid-Cambrian (\n            <jats:italic>c.<\/jats:italic>\n            520\u2013508\u2005Ma), with possible trace fossils in the latest Ediacaran (&lt;556\u2005Ma). The fossil record of Ecdysozoa is among the best understood of major animal clades and is believed to document their origins and evolutionary history well. Strikingly, however, molecular clock analyses have implied a considerably deeper Precambrian origin for Ecdysozoa, much older than their earliest fossils. Here, using an improved set of fossil calibrations, we performed Bayesian analyses to estimate an evolutionary time-tree for Ecdysozoa, sampling all eight phyla for the first time. Our results recover Scalidophora as the sister group to Nematoida\u2009+\u2009Panarthropoda (= Cryptovermes nov.) and suggest that the Ediacaran divergence of Ecdysozoa occurred at least 23\u2005myr before the first potential ecdysozoan trace fossils. This finding is impervious to the use of all plausible phylogenies, fossil prior distributions, evolutionary rate models and matrix partitioning strategies. Arthropods exhibit more precision and less incongruence between fossil- and clock-based estimates of clade ages than other ecdysozoan phyla.\n          <\/jats:p>\n          <jats:p content-type=\"supplementary-material\">\n            <jats:bold>Supplementary material:<\/jats:bold>\n            Full methodologies used and an appendix of fossil calibration points are available at\n            <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" specific-use=\"dataset is-supplemented-by\" xlink:href=\"https:\/\/doi.org\/10.6084\/m9.figshare.c.5811381\">https:\/\/doi.org\/10.6084\/m9.figshare.c.5811381<\/jats:ext-link>\n          <\/jats:p>\n          <jats:p content-type=\"thematic-collection\">\n            <jats:bold>Thematic collection:<\/jats:bold>\n            This article is part of the Advances in the Cambrian Explosion collection available at:\n            <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" ext-link-type=\"uri\" xlink:href=\"https:\/\/www.lyellcollection.org\/cc\/advances-cambrian-explosion\">https:\/\/www.lyellcollection.org\/cc\/advances-cambrian-explosion<\/jats:ext-link>\n          <\/jats:p>","DOI":"10.1144\/jgs2021-107","type":"journal-article","created":{"date-parts":[[2022,2,1]],"date-time":"2022-02-01T11:15:40Z","timestamp":1643714140000},"update-policy":"https:\/\/doi.org\/10.1144\/crossmark-policy","source":"Crossref","is-referenced-by-count":67,"title":["The Ediacaran origin of Ecdysozoa: integrating fossil and phylogenomic data"],"prefix":"10.1144","volume":"179","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1360-5696","authenticated-orcid":false,"given":"Richard J.","family":"Howard","sequence":"first","affiliation":[{"name":"Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Penryn TR10 9FE, UK"},{"name":"Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK"},{"name":"MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Yunnan University, Chenggong Campus, Kunming 650504, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Mattia","family":"Giacomelli","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Bristol, Bristol BS8 1TQ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3597-1221","authenticated-orcid":false,"given":"Jesus","family":"Lozano-Fernandez","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Bristol, Bristol BS8 1TQ, UK"},{"name":"Institute of Evolutionary Biology (CSIC-UPF), Barcelona 08003, Spain"},{"name":"Department of Genetics, Microbiology and Statistics, Biodiversity Research Institute (IRBio), University of Barcelona, Barcelona 08028, Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9591-8011","authenticated-orcid":false,"given":"Gregory D.","family":"Edgecombe","sequence":"additional","affiliation":[{"name":"Department of Earth Sciences, The Natural History Museum, London SW7 5BD, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9987-3984","authenticated-orcid":false,"given":"James F.","family":"Fleming","sequence":"additional","affiliation":[{"name":"University of Oslo Natural History Museum, Oslo 0562, Norway"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Reinhardt M.","family":"Kristensen","sequence":"additional","affiliation":[{"name":"Natural History Museum of Denmark, University of Copenhagen, Copenhagen 1123, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1172-6611","authenticated-orcid":false,"given":"Xiaoya","family":"Ma","sequence":"additional","affiliation":[{"name":"Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Penryn TR10 9FE, UK"},{"name":"MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, Yunnan University, Chenggong Campus, Kunming 650504, China"},{"name":"Yunnan Key Laboratory for Palaeobiology, Institute of Palaeontology, Yunnan University, Chenggong Campus, Kunming 650504, China"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9582-7083","authenticated-orcid":false,"given":"J\u00f8rgen","family":"Olesen","sequence":"additional","affiliation":[{"name":"Natural History Museum of Denmark, University of Copenhagen, Copenhagen 1123, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0377-0276","authenticated-orcid":false,"given":"Martin V.","family":"S\u00f8rensen","sequence":"additional","affiliation":[{"name":"Natural History Museum of Denmark, University of Copenhagen, Copenhagen 1123, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Philip F.","family":"Thomsen","sequence":"additional","affiliation":[{"name":"Department of Biology, Aarhus University, Aarhus 8000, Denmark"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Matthew A.","family":"Wills","sequence":"additional","affiliation":[{"name":"Milner Centre for Evolution, Department of Biology &amp; Biochemistry, University of Bath, Bath BA2 7AZ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Philip C. J.","family":"Donoghue","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Bristol, Bristol BS8 1TQ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0949-6682","authenticated-orcid":false,"given":"Davide","family":"Pisani","sequence":"additional","affiliation":[{"name":"School of Biological Sciences, University of Bristol, Bristol BS8 1TQ, UK"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1881","published-online":{"date-parts":[[2022,3,10]]},"reference":[{"key":"e_1_3_6_2_1","volume-title":"Ultrastruktur und Phylogenie von Seison Nebaliae (Grube 1859) und Seison Annulatus (Claus 1876): Hypothesen zu Phylogenetischen Verwandschaftsverh\u00e4ltnissen Innerhalb der Bilateria","author":"Ahlrichs W.H.","year":"1995","unstructured":"Ahlrichs, W.H. 1995. Ultrastruktur und Phylogenie von Seison Nebaliae (Grube 1859) und Seison Annulatus (Claus 1876): Hypothesen zu Phylogenetischen Verwandschaftsverh\u00e4ltnissen Innerhalb der Bilateria. Cuvillier, G\u00f6ttingen."},{"key":"e_1_3_6_3_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-662-08874-6_5"},{"key":"e_1_3_6_4_1","doi-asserted-by":"publisher","DOI":"10.1093\/molbev\/msv075"},{"key":"e_1_3_6_5_1","doi-asserted-by":"publisher","DOI":"10.1093\/gbe\/evy118"},{"key":"e_1_3_6_6_1","article-title":"Constraints on the timescale of animal evolutionary history","volume":"18","author":"Benton M.J.","year":"2015","unstructured":"Benton, M.J., Donoghue, P.C.J., Asher, R.J., Friedman, M., Near, T.J. and Vinther, J. 2015. Constraints on the timescale of animal evolutionary history. 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Backhuys, Leiden, 103\u2013110."},{"key":"e_1_3_6_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41559-018-0644-x"},{"key":"e_1_3_6_9_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1744-7410.2002.tb00136.x"},{"key":"e_1_3_6_10_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ympev.2014.08.001"},{"key":"e_1_3_6_11_1","first-page":"1","article-title":"The Cambrian substrate revolution","volume":"10","author":"Bottjer D.J.","year":"2000","unstructured":"Bottjer, D.J., Hagadorn, J.W. and Dornbos, S.Q. 2000. The Cambrian substrate revolution. 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