{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,9]],"date-time":"2026-06-09T04:37:10Z","timestamp":1780979830077,"version":"3.54.1"},"reference-count":71,"publisher":"Geological Society of London","issue":"5","license":[{"start":{"date-parts":[[2021,3,22]],"date-time":"2021-03-22T00:00:00Z","timestamp":1616371200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-002"}],"funder":[{"name":"Strategic Priority Research Program (B) of the Chinese Academy of Sciences","award":["XDB26000000"],"award-info":[{"award-number":["XDB26000000"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["41921002"],"award-info":[{"award-number":["41921002"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]},{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology","award":["20181105"],"award-info":[{"award-number":["20181105"]}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["42072006"],"award-info":[{"award-number":["42072006"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["www.lyellcollection.org"],"crossmark-restriction":true},"short-container-title":["JGS"],"published-print":{"date-parts":[[2021,9]]},"abstract":"<jats:p>\n            In spite of their unrivalled ecological success, the origins of terrestrial mandibulates have long remained virtually unknown. In recent years, claims have been made based on phylogenetic results that fuxianhuiids, iconic fossils of the Chengjiang biota and purported basal euarthropods, were in fact mandibulates, allied with the problematic euthycarcinoids as the closest sister-group to Mandibulata or even stem myriapods. A re-examination of euthycarcinoid mouthparts has since brought additional support to this view. Here, we reinvestigated the anterior anatomy of the Cambrian euarthropod\n            <jats:italic>Fuxianhuia<\/jats:italic>\n            and its relatives. We demonstrate that the fuxianhuiid head is that of a mandibulate, sharing similarities with hymenocarines, including a limbless intercalary segment. The hypostome is a sub-triangular sclerite partially fused but anatomically independent from a large, bilobed labral plate, as observed in many extant mandibulate taxa as well as euthycarcinoids. The so-called \u2018specialized post-antenn(ular) appendages' are therefore the mandibles, with a tripartite palp and large dented gnathal lobe partially covered by the hypostome-labrum complex. Under a \u2018deep split' phylogenetic scenario,\n            <jats:italic>Fuxianhuia<\/jats:italic>\n            and its relatives are here resolved as sister-group to crown Mandibulata.\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:\/\/protect-us.mimecast.com\/s\/avp5CQWRnRt6JoBRTxMIBz?domain=lyellcollection.org\">https:\/\/www.lyellcollection.org\/cc\/advances-cambrian-explosion<\/jats:ext-link>\n          <\/jats:p>\n          <jats:p content-type=\"supplementary-material\">\n            <jats:bold>Supplementary material<\/jats:bold>\n            : Additional remarks on terminology, morphological interpretations and a list of modified\/new characters used for the phylogenetic analysis 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.5305042\">https:\/\/doi.org\/10.6084\/m9.figshare.c.5305042<\/jats:ext-link>\n          <\/jats:p>","DOI":"10.1144\/jgs2020-246","type":"journal-article","created":{"date-parts":[[2021,2,17]],"date-time":"2021-02-17T15:55:25Z","timestamp":1613577325000},"update-policy":"https:\/\/doi.org\/10.1144\/crossmark-policy","source":"Crossref","is-referenced-by-count":30,"title":["Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda"],"prefix":"10.1144","volume":"178","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0567-1730","authenticated-orcid":false,"given":"C\u00e9dric","family":"Aria","sequence":"first","affiliation":[{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology &amp; Center for Excellence in Life and Palaeoenvironment, Chinese Academy of Sciences, Nanjing 210008, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8473-4439","authenticated-orcid":false,"given":"Fangchen","family":"Zhao","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology &amp; Center for Excellence in Life and Palaeoenvironment, Chinese Academy of Sciences, Nanjing 210008, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7327-9856","authenticated-orcid":false,"given":"Maoyan","family":"Zhu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology &amp; Center for Excellence in Life and Palaeoenvironment, Chinese Academy of Sciences, Nanjing 210008, China"},{"name":"College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, Beijing 100049, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1881","published-online":{"date-parts":[[2021,3,22]]},"reference":[{"key":"e_1_3_5_2_1","doi-asserted-by":"publisher","DOI":"10.1111\/1475-4983.00308"},{"key":"e_1_3_5_3_1","doi-asserted-by":"publisher","DOI":"10.1017\/pab.2020.36"},{"key":"e_1_3_5_4_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0124979"},{"key":"e_1_3_5_5_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature22080"},{"key":"e_1_3_5_6_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-017-1088-7"},{"key":"e_1_3_5_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/pala.12161"},{"key":"e_1_3_5_8_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12862-019-1560-7"},{"key":"e_1_3_5_9_1","doi-asserted-by":"publisher","DOI":"10.1080\/11035890809452772"},{"key":"e_1_3_5_10_1","first-page":"1","article-title":"The arthropod Branchiocaris n.gen., Middle Cambrian, Burgess Shale, British Columbia","volume":"264","author":"Briggs D.E.G","year":"1976","unstructured":"Briggs, D.E.G. 1976. The arthropod Branchiocaris n.gen., Middle Cambrian, Burgess Shale, British Columbia. Geological Survey of Canada, Bulletin, 264, 1\u201329.","journal-title":"Geological Survey of Canada, Bulletin"},{"key":"e_1_3_5_11_1","first-page":"631","article-title":"Anomalocaris, the largest known Cambrian arthropod","volume":"22","author":"Briggs D.E.G","year":"1979","unstructured":"Briggs, D.E.G. 1979. Anomalocaris, the largest known Cambrian arthropod. 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Proceedings of an International Symposium Held at the University of Camerino 27\u201331 March 1989, 189\u2013244."},{"key":"e_1_3_5_17_1","first-page":"191","article-title":"The organization of the subesophageal nervous system in tardigrades: Insights into the evolution of the arthropod hypostome and tritocerebrum","volume":"238","author":"Dewel R.A.","year":"1999","unstructured":"Dewel, R.A., Budd, G.E., Castano, D.F. and Dewel, W.C. 1999. The organization of the subesophageal nervous system in tardigrades: Insights into the evolution of the arthropod hypostome and tritocerebrum. 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