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Squamous or scaly reptilian\u2010like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non\u2010feathered integumentary structures in non\u2010avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non\u2010averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (<jats:italic>Carnotaurus<\/jats:italic>), Allosauroidea (<jats:italic>Allosaurus<\/jats:italic>,<jats:italic>Concavenator<\/jats:italic>,<jats:italic>Lourinhanosaurus<\/jats:italic>), Compsognathidae (<jats:italic>Juravenator<\/jats:italic>), and Tyrannosauroidea (<jats:italic>Santanaraptor<\/jats:italic>,<jats:italic>Albertosaurus<\/jats:italic>,<jats:italic>Daspletosaurus<\/jats:italic>,<jats:italic>Gorgosaurus<\/jats:italic>,<jats:italic>Tarbosaurus<\/jats:italic>,<jats:italic>Tyrannosaurus<\/jats:italic>), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to<jats:italic>Ceratosaurus<\/jats:italic>. Naked, non\u2010scale bearing skin is found in the contentious tetanuran<jats:italic>Sciurumimus<\/jats:italic>, ornithomimosaurians (<jats:italic>Ornithomimus<\/jats:italic>) and possibly tyrannosauroids (<jats:italic>Santanaraptor<\/jats:italic>), and also on the patagia of scansoriopterygids (<jats:italic>Ambopteryx<\/jats:italic>,<jats:italic>Yi<\/jats:italic>). Scales are surprisingly conservative among non\u2010avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and\/or within body regions in<jats:italic>Carnotaurus<\/jats:italic>,<jats:italic>Concavenator<\/jats:italic>and<jats:italic>Juravenator<\/jats:italic>, and include polarised, snake\u2010like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in<jats:italic>Tyrannosaurus<\/jats:italic>, whereas feature scales are present only in<jats:italic>Albertosaurus<\/jats:italic>and<jats:italic>Carnotaurus<\/jats:italic>. Few theropods currently show compelling evidence for the co\u2010occurrence of scales and feathers (e.g.<jats:italic>Juravenator<\/jats:italic>,<jats:italic>Sinornithosaurus<\/jats:italic>), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.<\/jats:p>","DOI":"10.1111\/brv.12829","type":"journal-article","created":{"date-parts":[[2022,1,7]],"date-time":"2022-01-07T00:53:10Z","timestamp":1641516790000},"page":"960-1004","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":45,"title":["Morphology and distribution of scales, dermal ossifications, and other non\u2010feather integumentary structures in non\u2010avialan theropod dinosaurs"],"prefix":"10.1111","volume":"97","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-8500-2405","authenticated-orcid":false,"given":"Christophe","family":"Hendrickx","sequence":"first","affiliation":[{"name":"Unidad Ejecutora Lillo, CONICET\u2010Fundaci\u00f3n Miguel Lillo 251 Miguel Lillo, San Miguel de Tucum\u00e1n Tucum\u00e1n 4000 Argentina"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Phil R.","family":"Bell","sequence":"additional","affiliation":[{"name":"School of Environmental and Rural Science University of New England Armidale NSW 2351 Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6149-3078","authenticated-orcid":false,"given":"Michael","family":"Pittman","sequence":"additional","affiliation":[{"name":"Vertebrate Palaeontology Laboratory, Department of Earth Sciences The University of Hong Kong Pokfulam Hong Kong SAR China"},{"name":"Department of Earth Sciences University College London WC1E 6BT United Kingdom"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4466-3477","authenticated-orcid":false,"given":"Andrew R. 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