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Although studies have detailed various food\u2010processing strategies and mechanisms across vertebrates, a coherent and comprehensive terminology is lacking. Here, we provide a synthesis, including a unified terminology for the intricate complexity of vertebrate oropharyngeal processing. Among gnathostomes, mandibular food processing predominates, ranging from discrete bites to rhythmic, cyclic chewing facilitated by precise tongue mechanics in aquatic and terrestrial environments alike. By contrast, some taxa have abandoned oropharyngeal processing entirely, relying instead on post\u2010oesophageal strategies such as gastric milling and chemical digestion. Interestingly, teleost (bony) fishes illustrate the evolutionary trade\u2010off between increased jaw protrusion for prey capture and reduced mandibular processing capacity. They compensated for this trade\u2010off by developing derived processing behaviours early in their evolutionary development. Through the re\u2010evolution of mandibular chewing, they succeeded in utilising all three known processing centres. Mastication is a specialised, dimensionally complex form of unignathic mandibular chewing (i.e. chewing restricted to the lower jaw) exclusive to mammals. However, our findings demonstrate that dimensionally complex forms of mandibular chewing have arisen independently multiple times and are widespread among gnathostomes. Notably, diverse taxa, including elasmobranch stingrays, Australian lungfish, sirenid salamanders, various songbirds, herbivorous turtles, and the tuatara, exhibit complex jaw movements combining arcuate, longitudinal, and sometimes transverse components enabled by specialised jaw joints, suspensions, and intracranial motions (\u2018cranial kinesis\u2019). From a comparative, functional\u2013morphological perspective, mammalian mastication may best be characterised as dimensionally complex chewing mediated by the secondary or temporomandibular joint. By contrast, analogous dimensionally complex non\u2010mammalian chewing involving motions confined to the primary or quadrate\u2013articular jaw joint qualifies as pseudomastication. Both mastication and pseudomastication resemble functional masticatory behaviours, while those incorporating intracranial motions and movements of the jaw suspension belong to distinct categories. Our anatomical analysis highlights the convergent evolution of dimensionally complex chewing among gnathostomes and emphasises the importance of comprehensive studies on jaw development and function to deepen our understanding of the evolution of oropharyngeal processing.<\/jats:p>","DOI":"10.1002\/brv.70129","type":"journal-article","created":{"date-parts":[[2026,1,21]],"date-time":"2026-01-21T02:40:42Z","timestamp":1768963242000},"page":"1406-1462","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Beyond mammals: the evolution of chewing and other forms of oropharyngeal food processing in vertebrates"],"prefix":"10.1002","volume":"101","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5845-4027","authenticated-orcid":false,"given":"Daniel","family":"Schwarz","sequence":"first","affiliation":[{"name":"Department of Palaeontology State Museum of Natural History Stuttgart (SMNS)  Rosenstein 1 Stuttgart 70191 Germany"},{"name":"Friedrich Schiller University Jena (FSU), Institute of Zoology and Evolutionary Research  Erbertstra\u00dfe 1 Jena 07743 Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6328-0589","authenticated-orcid":false,"given":"Maja","family":"Mielke","sequence":"additional","affiliation":[{"name":"Laboratory of Functional Morphology, Department of Biology, Faculty of Sciences University of Antwerp  Wilrijk 2610 Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2140-7892","authenticated-orcid":false,"given":"Stephan","family":"Handschuh","sequence":"additional","affiliation":[{"name":"VetCore Facility for Research\/Imaging Unit University of Veterinary Medicine Vienna  Veterinaerplatz 1 Vienna 1210 Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0991-4434","authenticated-orcid":false,"given":"Anthony","family":"Herrel","sequence":"additional","affiliation":[{"name":"D\u00e9partement Adaptations du Vivant UMR 7179 C.N.R.S\/M.N.H.N  B\u00e2timent d'Anatomie Compar\u00e9e, 55 rue Buffon Paris 75005 France"},{"name":"Department of Biology, Evolutionary Morphology of Vertebrates Ghent University  Ghent 9000 Belgium"},{"name":"Department of Biology University of Antwerp  Wilrijk 2610 Belgium"},{"name":"Naturhistorisches Museum Bern  Bern 3005 Switzerland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0262-4047","authenticated-orcid":false,"given":"Patrick","family":"Lemell","sequence":"additional","affiliation":[{"name":"Department of Evolutionary Biology, Unit for Integrative Zoology University of Vienna  Djerassiplatz 1 Vienna 1030 Austria"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-6479-8419","authenticated-orcid":false,"given":"L\u00e9a Da","family":"Cunha","sequence":"additional","affiliation":[{"name":"Institut des Sciences de l'\u00c9volution de Montpellier, Universit\u00e9 de Montpellier, CNRS, IRD  c. c. 64 Montpellier France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3310-9080","authenticated-orcid":false,"given":"Nicolai","family":"Konow","sequence":"additional","affiliation":[{"name":"Department of Biological Sciences University of Massachusetts Lowell  198 Riverside Street Lowell MA 01854 USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,1,20]]},"reference":[{"key":"e_1_2_13_3_1","doi-asserted-by":"publisher","DOI":"10.3354\/meps11188"},{"key":"e_1_2_13_4_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.1995.0027"},{"key":"e_1_2_13_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/0169-5347(96)10048-3"},{"key":"e_1_2_13_6_1","first-page":"189","article-title":"The cranial anatomy of Polypterus, with special reference to Polypterus bichir","volume":"56","author":"Allis E. 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