{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T05:01:48Z","timestamp":1785214908080,"version":"3.55.0"},"reference-count":47,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Anatomy"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Enameloid is a type of hard mineralised tissue covering teeth and dermal denticles of extant and extinct chondrichthyans. Understanding the diversity of enameloid microstructures across chondrichthyan clades is key to highlight potential phylogenetic or adaptive signals provided by histological characters. Among chondrichthyans, the enameloid microstructure of teeth and odontodes of batomorphs (skates and rays) has been understudied despite the high extant and past taxonomic and morphological diversities of this clade. Here, we focus on the sclerorhynchoid rays, a speciose extinct (Cretaceous) clade of skates that developed rostra bearing enlarged denticles in most taxa. We explore the enameloid microstructure of both teeth and rostral denticles of 13 species representing a wide range of morphologies. Our results show that the tooth enameloid microstructure of sclerorhynchoid skates achieved a complex arrangement, with a high diversity of bundled components. This microstructure is furthermore complexified by a strong degree of apico\u2010basal variation in the organisation of the enameloid components. This organisation is the most complex known so far among batomorph teeth and probably elasmobranchs in general. Our results further confirm the presence of a similarly complex enameloid microstructure in the dermal denticles covering the elongated rostrum of these extinct skates. Most importantly, we provide evidence for the pervasiveness of this complex enameloid across rostral denticles of all studied taxa, despite the wide range of taxa and morphologies represented. The common enameloid organisation reported in both teeth and rostral denticles is, from base to the apex: a Single\u2010Crystallite Enameloid (SCE) layer passing into Radial Bundled Enameloid (RBE) sometimes through a phase of protobundles as the enameloid thickens slightly higher in the crown, with apico\u2010basal Parallel\u2010Bundled Enameloid (PBE) appearing mixed with RBE bundles higher up at the base of the cusp, followed by the classical three\u2010layered enameloid including SCE, PBE and Tangled\u2010Bundled Enameloid (TBE) in the apex, crossed by rare Thick Radial Bundles (TRB). The presence of protobundles and RBE forming the entire enameloid thickness in the basalmost crown enameloid of teeth and rostral denticles analysed here was so far restricted to placoid scales and teeth of some hybodonts and batomorphs. Our study identified protobundles as transitional organisation between SCE and structured bundles and the RBE as an organisational stage preceding the PBE. We demonstrate that TRB are isolated RBE bundles and that both RBE and PBE bundles can participate in circumferential bundles whose development is linked with the size of the cutting edges. We found no evidence for a clear phylogenetic signal provided by the distribution of enameloid microstructure characters neither within teeth, nor among rostral denticles sampled here. Instead, we propose a possible link with geometric constraints (height, width), with SCE and RBE (with or without protobundles) being restricted to thin enameloid layers. Taken together, these results have strong implications for the understanding of enameloid character distribution in elasmobranch odontodes.<\/jats:p>","DOI":"10.1111\/joa.70215","type":"journal-article","created":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T04:43:55Z","timestamp":1785213835000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Widespread complex enameloid microstructure in teeth and rostral denticles of Cretaceous sclerorhynchoid skates (Rajiformes: \u2020Sclerorhynchoidei)"],"prefix":"10.1111","author":[{"given":"Endy","family":"Chateauneuf\u2010Gautier","sequence":"first","affiliation":[{"name":"ISEM, University of Montpellier, CNRS, IRD  Montpellier France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3363-6972","authenticated-orcid":false,"given":"Guillaume","family":"Guinot","sequence":"additional","affiliation":[{"name":"ISEM, University of Montpellier, CNRS, IRD  Montpellier France"},{"name":"Institut universitaire de France (IUF)  France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,7,27]]},"reference":[{"key":"e_1_2_8_2_1","doi-asserted-by":"crossref","first-page":"24699","DOI":"10.1039\/D5NR03589A","article-title":"Orthogonal design and dual\u2010mode construction of oriented bundles of fluorapatite from organic fibrils in shark enameloid","volume":"17","author":"Akimoto S.","year":"2025","journal-title":"Nanoscale"},{"key":"e_1_2_8_3_1","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/S0016-6995(95)80003-4","article-title":"\u00c9tude micropal\u00e9ontologique du S\u00e9nonien du synclinal de Tighboula, Moyen Atlas, Maroc: Biostratigraphie et pal\u00c9oenvironnements","volume":"28","author":"Andreu B.","year":"1995","journal-title":"Geobios"},{"key":"e_1_2_8_4_1","first-page":"504","article-title":"Selachians of the Carlile Shale (Turonian) of South Dakota","volume":"47","author":"Cappetta H.","year":"1973","journal-title":"Journal of Paleontology"},{"key":"e_1_2_8_5_1","first-page":"225","article-title":"Sclerorhynchidae nov. fam., Pristidae et Pristiophoridae: un exemple de parall\u00e9lisme chez les S\u00e9laciens","volume":"278","author":"Cappetta H.","year":"1974","journal-title":"Comptes Rendus de l'Acad\u00e9mie des Sciences de Paris"},{"key":"e_1_2_8_6_1","first-page":"149","article-title":"Les s\u00e9laciens du Cr\u00e9tac\u00e9 sup\u00e9rieur du Liban. II: Bato\u00efdes","volume":"168","author":"Cappetta H.","year":"1980","journal-title":"Palaeontographica Abteilung A Palaozoologie Stratigraphie"},{"key":"e_1_2_8_7_1","first-page":"17","article-title":"D\u00e9couverte de nouvelles faunes de s\u00e9laciens (Neoselachii) dans les phosphates maastrichtiens de la Mer Rouge, Egypte","volume":"19","author":"Cappetta H.","year":"1991","journal-title":"M\u00fcnchner Geowissenschaftliche Abhandlungen"},{"key":"e_1_2_8_8_1","first-page":"435","article-title":"Late Cretaceous selachian faunas from Bolivia","volume":"12","author":"Cappetta H.","year":"1991","journal-title":"Revista T\u00e9cnica de Yacimientos Petrol\u00edferos Fiscales Bolivianos"},{"key":"e_1_2_8_9_1","volume-title":"Chondrichthyes\u2014Mesozoic and Cenozoic Elasmobranchii: teeth","author":"Cappetta H.","year":"2012"},{"key":"e_1_2_8_10_1","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1016\/j.palaeo.2014.04.020","article-title":"Marine vertebrate faunas from the 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Anatomy"},{"key":"e_1_2_8_17_1","volume-title":"Evolution of dental tissues and paleobiology in selachians","author":"Cuny G.","year":"2017"},{"key":"e_1_2_8_18_1","first-page":"97","volume-title":"Late Palaeozoic and Mesozoic ecosystems in SE Asia","author":"Cuny G.","year":"2009"},{"key":"e_1_2_8_19_1","doi-asserted-by":"crossref","first-page":"144","DOI":"10.1111\/zoj.12059","article-title":"Simplification of the enameloid microstructure of large stingrays (Chondrichthyes: Myliobatiformes): a functional approach","volume":"169","author":"Enault S.","year":"2013","journal-title":"Zoological Journal of the Linnean Society"},{"key":"e_1_2_8_20_1","doi-asserted-by":"crossref","first-page":"549","DOI":"10.1111\/zoj.12244","article-title":"Chondrichthyan tooth enameloid: past, present, and future","volume":"174","author":"Enault S.","year":"2015","journal-title":"Zoological Journal of the Linnean 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Biostratigraphic and paleoenvironmental implications","volume":"1989","author":"Lewy Z.","year":"1989","journal-title":"Neues Jahrbuch f\u00fcr Geologie und Pal\u00e4ontologie (Monatsheft)"},{"key":"e_1_2_8_32_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s41513-018-0049-3","article-title":"On the enameloid microstructure of Archaeobatidae (Neoselachii, Chondrichthyes)","volume":"44","author":"Manzanares E.","year":"2018","journal-title":"Journal of Iberian Geology"},{"key":"e_1_2_8_33_1","doi-asserted-by":"crossref","first-page":"1060","DOI":"10.1134\/S0031030114100062","article-title":"The enameloid microstructure of euselachian (Chondrichthyes) scales","volume":"48","author":"Manzanares E.","year":"2014","journal-title":"Paleontological Journal"},{"key":"e_1_2_8_34_1","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1111\/zoj.12377","article-title":"New insights into the enameloid microstructure of batoid fishes (Chondrichthyes)","volume":"177","author":"Manzanares E.","year":"2016","journal-title":"Zoological Journal of the Linnean Society"},{"key":"e_1_2_8_35_1","first-page":"321","article-title":"Histologic studies of placoderms and fossil elasmobranchs 1. The endoskeleton, with remarks on the hard tissues of lower vertebrates in general","volume":"2","author":"\u00d8rvig T.","year":"1951","journal-title":"Arkiv F\u00f6r Zoologi"},{"key":"e_1_2_8_36_1","first-page":"65","volume-title":"Dental morphology and evolution","author":"Poole D.F.G.","year":"1971"},{"key":"e_1_2_8_37_1","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1111\/j.1463-6409.1974.tb00753.x","article-title":"Morphologie und Ultrastuktur des Hai\u2010\u201cSchmelzes\u201d","volume":"2","author":"Reif W.\u2010E.","year":"1973","journal-title":"Zoologica Scripta"},{"key":"e_1_2_8_38_1","first-page":"565","article-title":"Tooth enameloid as a taxonomic criterion: 1. A new euselachian shark from the Rhaeic\u2010Liassic boundary","volume":"1977","author":"Reif W.\u2010E.","year":"1977","journal-title":"Neues Jahrbuch f\u00fcr Geologie und Pal\u00e4ontologie (Monatsheft)"},{"key":"e_1_2_8_39_1","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1017\/S1464793102006073","article-title":"Formation of dermal skeletal and dental tissues in fish: a comparative and evolutionary approach","volume":"78","author":"Sire J.\u2010Y.","year":"2003","journal-title":"Biological Reviews"},{"key":"e_1_2_8_40_1","first-page":"1","article-title":"Jurazeittiche Neoselachier aus Deutschland und S\u00fcd\u2010England","volume":"58","author":"Thies D.","year":"1983","journal-title":"Courier Forschungsinstitut Senckenberg"},{"key":"e_1_2_8_41_1","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1666\/04069.1","article-title":"Diversification of the Neoselachii (Chondrichthyes) during the Jurassic and Cretaceous","volume":"32","author":"Underwood C.J.","year":"2006","journal-title":"Paleobiology"},{"key":"e_1_2_8_42_1","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1144\/SP430.7","article-title":"Sclerorhynchus atavus and the convergent evolution of rostrum\u2010bearing chondrichthyans","volume":"430","author":"Underwood C.J.","year":"2016","journal-title":"Geological Society, London, Special Publications"},{"key":"e_1_2_8_43_1","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1093\/zoolinnean\/zlaa166","article-title":"The skeletal remains of the euryhaline sclerorhynchoid \u2020Onchopristis (Elasmobranchii) from the \u2018mid\u2019\u2010Cretaceous and their palaeontological implications","volume":"193","author":"Villalobos\u2010Segura E.","year":"2021","journal-title":"Zoological Journal of the Linnean Society"},{"key":"e_1_2_8_44_1","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1002\/spp2.1287","article-title":"The first skeletal record of the enigmatic Cretaceous sawfish genus Ptychotrygon (Chondrichthyes, Batoidea) from the Turonian of Morocco","volume":"7","author":"Villalobos\u2010Segura E.","year":"2021","journal-title":"Papers in Palaeontology"},{"key":"e_1_2_8_45_1","doi-asserted-by":"crossref","first-page":"1847","DOI":"10.1080\/14772019.2019.1578832","article-title":"The first three\u2010dimensional fossils of Cretaceous sclerorhynchid sawfish: Asflapristis cristadentis gen. et sp. nov., and implications for the phylogenetic relations of the Sclerorhynchoidei (Chondrichthyes)","volume":"17","author":"Villalobos\u2010Segura E.","year":"2019","journal-title":"Journal of Systematic Palaeontology"},{"key":"e_1_2_8_46_1","first-page":"99","article-title":"New sharks and rays from the Cenomanian and Turonian of Charentes, France","volume":"52","author":"Vullo R.","year":"2007","journal-title":"Acta Palaeontologica Polonica"},{"key":"e_1_2_8_47_1","doi-asserted-by":"crossref","DOI":"10.1098\/rsos.150189","article-title":"Evolutionary origins and development of saw\u2010teeth on the sawfish and sawshark rostrum (Elasmobranchii; Chondrichthyes)","volume":"2","author":"Welten M.","year":"2015","journal-title":"Royal Society Open Science"},{"key":"e_1_2_8_48_1","doi-asserted-by":"crossref","first-page":"445","DOI":"10.1007\/s11160-009-9112-7","article-title":"The biology of extinct and extant sawfish (Batoidea: Sclerorhynchidae and Pristidae)","volume":"19","author":"Wueringer B.E.","year":"2009","journal-title":"Reviews in Fish Biology and Fisheries"}],"container-title":["Journal of Anatomy"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/joa.70215","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/joa.70215","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/joa.70215","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T04:44:11Z","timestamp":1785213851000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/joa.70215"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,27]]},"references-count":47,"alternative-id":["10.1111\/joa.70215"],"URL":"https:\/\/doi.org\/10.1111\/joa.70215","archive":["Portico"],"relation":{},"ISSN":["0021-8782","1469-7580"],"issn-type":[{"value":"0021-8782","type":"print"},{"value":"1469-7580","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,27]]},"assertion":[{"value":"2026-05-05","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-06","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-27","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"joa.70215"}}