{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,30]],"date-time":"2026-06-30T22:42:10Z","timestamp":1782859330158,"version":"3.54.5"},"reference-count":58,"publisher":"The Royal Society","issue":"1","license":[{"start":{"date-parts":[[2026,1,21]],"date-time":"2026-01-21T00:00:00Z","timestamp":1768953600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,1,21]],"date-time":"2026-01-21T00:00:00Z","timestamp":1768953600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004837","name":"Ministry of Science and Innovation of Spain","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,21]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\u2020Otodus megalodon, the largest known macropredatory shark, was globally distributed from the Miocene to Pliocene, yet most ecological and palaeobiological inferences rely heavily on American collections. This geographical bias limits understanding of its population structure and life-history strategies elsewhere. Two explanations have been proposed for its body-size patterns: a Bergmann-type cline, with larger individuals at higher, cooler latitudes and a related hypothesis of Mediterranean dwarfism; and the influence of localized nursery areas that generate assemblages dominated by small juveniles, as documented in several American sites and the Revert\u00e9 quarries in the western Mediterranean. Here, we present the first extensive dataset outside the Americas designed to test these alternatives across two adjacent basins. Body-size estimates show no significant difference between Atlantic and non-Revert\u00e9 Mediterranean populations, whereas the Revert\u00e9 assemblage is smaller. This pattern contradicts basin-wide Mediterranean dwarfism and does not support a Bergmann-type size\u2013latitude relationship. Instead, it strongly indicates a localized nursery at Revert\u00e9. More broadly, the results align with an emerging view of widespread nursery habitats in \u2020O. megalodon, comparable to those of extant sharks. By filling a key regional gap, the Iberian record provides tests of competing hypotheses and refines understanding of \u2020O. megalodon population structure and life history.<\/jats:p>","DOI":"10.1098\/rsbl.2025.0640","type":"journal-article","created":{"date-parts":[[2026,1,21]],"date-time":"2026-01-21T08:00:01Z","timestamp":1768982401000},"source":"Crossref","is-referenced-by-count":1,"title":["The Iberian fossil record of \u2020\n                    <i>Otodus megalodon<\/i>\n                    rejects Mediterranean dwarfism and supports nursery use"],"prefix":"10.1098","volume":"22","author":[{"given":"Jos\u00e9 Luis","family":"Herraiz","sequence":"first","affiliation":[{"name":"University of Valencia 1 , , ,","place":["Valencia, Valencian Community, Spain"],"department":["Cavanilles Institute of Biodiversity and Evolutionary Biology"]},{"name":"Museu Valenci\u00e0 d\u2019Hist\u00f2ria Natural\u2013i\\Biotaxa 2 , , ,","place":["Alginet, Valencia, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2254-8424","authenticated-orcid":false,"given":"Humberto G.","family":"Ferr\u00f3n","sequence":"additional","affiliation":[{"name":"University of Valencia 1 , , ,","place":["Valencia, Valencian Community, Spain"],"department":["Cavanilles Institute of Biodiversity and Evolutionary Biology"]},{"name":"University of Bristol 3 , ,","place":["Bristol, UK"],"department":["School of Earth Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H\u00e9ctor","family":"Botella","sequence":"additional","affiliation":[{"name":"University of Valencia 1 , , ,","place":["Valencia, Valencian Community, Spain"],"department":["Cavanilles Institute of Biodiversity and Evolutionary Biology"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mat\u00edas","family":"Reolid","sequence":"additional","affiliation":[{"name":"University of Jaen 4 , ,","place":["Jaen, Spain"],"department":["Department of Geology and Center for Advanced Studies in Earth and Environmental Sciences (CEACTEMA)"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7795-5997","authenticated-orcid":false,"given":"Carlos","family":"Mart\u00ednez-P\u00e9rez","sequence":"additional","affiliation":[{"name":"University of Valencia 1 , , ,","place":["Valencia, Valencian Community, Spain"],"department":["Cavanilles Institute of Biodiversity and Evolutionary Biology"]},{"name":"University of Bristol 3 , ,","place":["Bristol, UK"],"department":["School of Earth Sciences,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,1,21]]},"reference":[{"key":"2026012102595542600_R1","doi-asserted-by":"crossref","first-page":"1055","DOI":"10.1111\/j.1461-0248.2010.01489.x","article-title":"Patterns and ecosystem consequences of shark declines in the ocean","volume":"13","author":"Ferretti","year":"2010","journal-title":"Ecol. Lett."},{"key":"2026012102595542600_R2","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1126\/science.1205106","article-title":"Trophic downgrading of planet Earth","volume":"333","author":"Estes","year":"2011","journal-title":"Science"},{"key":"2026012102595542600_R3","doi-asserted-by":"crossref","first-page":"354","DOI":"10.1086\/679735","article-title":"The body size dependence of trophic cascades","volume":"185","author":"DeLong","year":"2015","journal-title":"Am. Nat."},{"key":"2026012102595542600_R4","doi-asserted-by":"crossref","first-page":"1846","DOI":"10.1126\/science.1138657","article-title":"Cascading effects of the loss of apex predatory sharks from a coastal ocean","volume":"315","author":"Myers","year":"2007","journal-title":"Science"},{"key":"2026012102595542600_R5","doi-asserted-by":"crossref","first-page":"1530326","DOI":"10.3389\/fevo.2024.1530326","article-title":"Drivers of diversification in sharks and rays (Chondrichthyes: Elasmobranchii)","volume":"12","author":"Gayford","year":"2025","journal-title":"Front. Ecol. Evol."},{"key":"2026012102595542600_R6","doi-asserted-by":"crossref","first-page":"5138","DOI":"10.1016\/j.cub.2021.09.028","article-title":"Feeding ecology has shaped the evolution of modern sharks","volume":"31","author":"Bazzi","year":"2021","journal-title":"Curr. Biol."},{"key":"2026012102595542600_R7","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0044632","article-title":"An analytical approach for estimating fossil record and diversification events in sharks, skates and rays","volume":"7","author":"Guinot","year":"2012","journal-title":"PLoS One"},{"key":"2026012102595542600_R8","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/B978-012415031-7\/50008-2","volume-title":"Great white sharks: the biology of Carcharodon carcharias","author":"Gottfried","year":"1996"},{"key":"2026012102595542600_R9","article-title":"Body length estimation of Neogene macrophagous lamniform sharks (Carcharodon and Otodus) derived from associated fossil dentitions","volume":"24","author":"P\u00e9rez","year":"2021","journal-title":"Palaeontol. Electron"},{"key":"2026012102595542600_R10","doi-asserted-by":"crossref","first-page":"904","DOI":"10.1080\/08912963.2019.1666840","article-title":"The size of the megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), revisited","volume":"33","author":"Shimada","year":"2021","journal-title":"Hist. Biol."},{"key":"2026012102595542600_R11","article-title":"Reassessment of the possible size, form, weight, cruising speed, and growth parameters of the extinct megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), and new evolutionary insights into its gigantism, life history strategies, ecology, and extinction","volume":"28","author":"Shimada","year":"2025","journal-title":"Palaeontol. Electron."},{"key":"2026012102595542600_R12","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-020-71387-y","article-title":"Body dimensions of the extinct giant shark Otodus megalodon: a 2D reconstruction","volume":"10","author":"Cooper","year":"2020","journal-title":"Sci. Rep."},{"key":"2026012102595542600_R13","article-title":"White shark comparison reveals a slender body for the extinct megatooth shark, Otodus megalodon (Lamniformes: Otodontidae)","volume":"27","author":"Sternes","year":"2024","journal-title":"Palaeontol. Electron."},{"key":"2026012102595542600_R14","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1111\/j.1755-263X.2008.00046.x","article-title":"Reassessing the value of nursery areas to shark conservation and management","volume":"2","author":"Kinney","year":"2009","journal-title":"Conserv. Lett."},{"key":"2026012102595542600_R15","doi-asserted-by":"crossref","first-page":"237","DOI":"10.1071\/MF9920237","article-title":"Preliminary results of a study into the incidental mortality of sharks in gill-nets in two Tasmanian shark nursery areas","volume":"43","author":"Williams","year":"1992","journal-title":"Mar. Freshw. Res."},{"key":"2026012102595542600_R16","first-page":"236","article-title":"Management of the sandbar shark, Carcharhinus plumbeus: implications of a stage-based model","volume":"98","author":"Brewster-Geisz","year":"2000","journal-title":"Fish. Bull."},{"key":"2026012102595542600_R17","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1007\/BF00842902","article-title":"The shark nursery of Bulls Bay, South Carolina, with a review of the shark nurseries of the southeastern coast of the United States","volume":"38","author":"Castro","year":"1993","journal-title":"Environ. Biol. Fishes"},{"key":"2026012102595542600_R18","doi-asserted-by":"crossref","first-page":"123","DOI":"10.3354\/meps13483","article-title":"Sensitivity of a shark nursery habitat to a changing climate","volume":"652","author":"Crear","year":"2020","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2026012102595542600_R19","doi-asserted-by":"crossref","first-page":"897","DOI":"10.1071\/MF18081","article-title":"Advances in understanding the roles and benefits of nursery areas for elasmobranch populations","volume":"70","author":"Heupel","year":"2018","journal-title":"Mar. Freshw. Res"},{"key":"2026012102595542600_R20","doi-asserted-by":"crossref","first-page":"287","DOI":"10.3354\/meps337287","article-title":"Shark nursery areas: concepts, definition, characterization and assumptions","volume":"337","author":"Heupel","year":"2007","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2026012102595542600_R21","doi-asserted-by":"crossref","DOI":"10.1098\/rsbl.2020.0746","article-title":"Use of nursery areas by the extinct megatooth shark Otodus megalodon","volume":"16","author":"Herraiz","year":"2020","journal-title":"Biol. Lett"},{"key":"2026012102595542600_R22","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0010552","article-title":"Ancient nursery area for the extinct giant shark Megalodon from the Miocene of Panama","volume":"5","author":"Pimiento","year":"2010","journal-title":"PLoS One"},{"key":"2026012102595542600_R23","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1080\/08912963.2022.2032024","article-title":"Revisiting body size trends and nursery areas of the Neogene megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), reveals Bergmann\u2019s rule possibly enhanced its gigantism in cooler waters","volume":"35","author":"Shimada","year":"2023","journal-title":"Hist. Biol."},{"key":"2026012102595542600_R24","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1046\/j.1365-2699.2003.00837.x","article-title":"On the validity of Bergmann\u2019s rule","volume":"30","author":"Meiri","year":"2003","journal-title":"J. Biogeogr."},{"key":"2026012102595542600_R25","first-page":"595","article-title":"\u00dcber die Verh\u00e4ltnisse der W\u00e4rme\u00f6konomie der Thiere zu ihrer Gr\u00f6sse","volume":"1","author":"Bergmann","year":"1847","journal-title":"G\u00f6ttinger Studien"},{"key":"2026012102595542600_R26","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1111\/j.1466-8238.2010.00577.x","article-title":"Bergmann\u2019s rule\u2014what\u2019s in a name?","volume":"20","author":"Meiri","year":"2011","journal-title":"Glob. Ecol. Biogeogr."},{"key":"2026012102595542600_R27","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1086\/674094","article-title":"The cold-water connection: Bergmann\u2019s rule in North American freshwater fishes","volume":"183","author":"Rypel","year":"2014","journal-title":"Am. Nat."},{"key":"2026012102595542600_R28","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1111\/j.1461-0248.2006.00928.x","article-title":"Ecotypic variation in the context of global climate change: revisiting the rules","volume":"9","author":"Millien","year":"2006","journal-title":"Ecol. Lett."},{"key":"2026012102595542600_R29","first-page":"1","article-title":"List of skeletal material from megatooth sharks (Lamniformes, Otodontidae)","volume":"4","author":"Greenfield","year":"2022","journal-title":"Paleoichthys"},{"key":"2026012102595542600_R30","first-page":"1","article-title":"Phylogenetic affinity of the extinct shark family Otodontidae within Lamniformes remains uncertain\u2014comments on \u2018list of skeletal material from megatooth sharks (Lamniformes, Otodontidae)\u2019 by greenfield","volume":"6","author":"Shimada","year":"2022","journal-title":"Paleoichthys"},{"key":"2026012102595542600_R31","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0011842","article-title":"The biodiversity of the Mediterranean Sea: estimates, patterns, and threats","volume":"5","author":"Coll","year":"2010","journal-title":"PLoS One"},{"key":"2026012102595542600_R32","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1038\/267399a0","article-title":"History of the Mediterranean salinity crisis","volume":"267","author":"Hs\u00fc","year":"1977","journal-title":"Nature"},{"key":"2026012102595542600_R33","doi-asserted-by":"crossref","first-page":"240","DOI":"10.1038\/242240a0","article-title":"Late Miocene desiccation of the Mediterranean","volume":"242","author":"Hs\u00fc","year":"1973","journal-title":"Nature"},{"key":"2026012102595542600_R34","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1038\/23231","article-title":"Chronology, causes and progression of the Messinian salinity crisis","volume":"400","author":"Krijgsman","year":"1999","journal-title":"Nature"},{"key":"2026012102595542600_R35","doi-asserted-by":"crossref","first-page":"1259","DOI":"10.1080\/08912963.2023.2211597","article-title":"Tessellated calcified cartilage and placoid scales of the Neogene megatooth shark, Otodus megalodon (Lamniformes: Otodontidae), offer new insights into its biology and the evolution of regional endothermy and gigantism in the otodontid clade","volume":"36","author":"Shimada","year":"2024","journal-title":"Hist. Biol."},{"key":"2026012102595542600_R36","first-page":"181","article-title":"Megalodon, mako shark and planktonic foraminifera from the continental shelf off Portugal and their age","volume":"13","author":"Antunes","year":"2015","journal-title":"Geol. Acta"},{"key":"2026012102595542600_R37","doi-asserted-by":"crossref","first-page":"289","DOI":"10.7203\/sjp.34.2.16118","article-title":"Pre-Messinian ecological diversity of Mediterranean sharks revealed by the study of their dermal denticles","volume":"34","author":"Ferr\u00f3n","year":"2019","journal-title":"Span. J. Palaeont"},{"key":"2026012102595542600_R38","doi-asserted-by":"crossref","first-page":"43395","DOI":"10.11137\/1982-3908_2021_44_43395","article-title":"Fossil chondrichthyes from the Neogene of Portugal: diversity and occurrence","volume":"44","author":"Fialho","year":"2021","journal-title":"Anu. Inst. Geoc"},{"key":"2026012102595542600_R39","doi-asserted-by":"crossref","DOI":"10.1080\/02724634.2024.2307071","article-title":"New data on the Late Miocene chondrichthyans from the Western Mediterranean region (Alcoy Basin, Eastern Spain)","volume":"43","author":"Herraiz","year":"2023","journal-title":"J. Vertebr. Paleontol."},{"key":"2026012102595542600_R40","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1080\/08912963.2017.1326111","article-title":"A Serravallian (Middle Miocene) shark fauna from southeastern Spain and its palaeoenvironment significance","volume":"30","author":"Mart\u00ednez-P\u00e9rez","year":"2018","journal-title":"Hist. Biol."},{"key":"2026012102595542600_R41","doi-asserted-by":"crossref","first-page":"102","DOI":"10.7203\/sjp.28830","article-title":"Analysing trophic competition in \u2020Otodus megalodon and Carcharodon carcharias through 2D-SEM dental microwear","volume":"39","author":"Paredes-Aliaga","year":"2024","journal-title":"Spanish J. Palaeontol."},{"key":"2026012102595542600_R42","article-title":"Record of Carcharocles megalodon in the Eastern Guadalquivir Basin (Upper Miocene, South Spain)","volume":"71","author":"Reolid","year":"2015","journal-title":"Estud. Geol."},{"key":"2026012102595542600_R43","volume-title":"Sel\u00e1ceos do neog\u00e9nico da Ilha de Santa Maria (A\u00e7ores)","author":"de Azevedo","year":"2017"},{"key":"2026012102595542600_R44","first-page":"107","article-title":"First description of a tooth of the extinct giant shark \u2018Carcharocles megalodon\u2019 (Agassiz, 1835) found in the province of Seville (SW Iberian Peninsula) (Otodontidae)","volume":"4","author":"Medina-Gavil\u00e1n","year":"2015","journal-title":"BV news Publ. Cient."},{"key":"2026012102595542600_R45","volume-title":"Handbook of paleoichthyology. Volume 3E. Chondrichthyes, mesozoic and Cenozoic elasmobranchii: teeth","author":"Cappetta","year":"2012"},{"key":"2026012102595542600_R46","doi-asserted-by":"crossref","first-page":"479","DOI":"10.1017\/pab.2015.16","article-title":"Body-size trends of the extinct giant shark Carcharocles megalodon: a deep-time perspective on marine apex predators","volume":"41","author":"Pimiento","year":"2015","journal-title":"Paleobiology"},{"key":"2026012102595542600_R47","first-page":"28","article-title":"The relationship between the tooth size and total body length in the white shark","volume":"35","author":"Shimada","year":"2002","journal-title":"J. Foss. Res."},{"key":"2026012102595542600_R48","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1017\/pab.2022.40","article-title":"Diversification trajectories and paleobiogeography of Neogene chondrichthyans from Europe","volume":"49","author":"Villafa\u00f1a","year":"2023","journal-title":"Paleobiology"},{"key":"2026012102595542600_R49","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v028.c01","article-title":"Beanplot: a boxplot alternative for visual comparison of distributions","volume":"28","author":"Kampstra","year":"2008","journal-title":"J. Stat. Softw."},{"key":"2026012102595542600_R50","unstructured":"R Core Team\n          . 2025R: a language and environment for statistical computing. R Foundation for Statistical Computing. Seehttps:\/\/www.R-project.org."},{"key":"2026012102595542600_R51","unstructured":"Wickham\n              H\n            , BrianJ. 2025readxl: read Excel files. (doi: 10.32614\/CRAN.package.readxl)"},{"key":"2026012102595542600_R52","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","volume-title":"Ggplot2: elegant graphics for data analysis","author":"Wickham","year":"2016"},{"key":"2026012102595542600_R53","volume-title":"Atlas of Paleogene paleogeographic maps (Mollweide projection), maps 8\u201315, volume 1: the Cenozoic. PALEOMAP Atlas for ArcGIS","author":"Scotese","year":"2014"},{"key":"2026012102595542600_R54","doi-asserted-by":"crossref","DOI":"10.1093\/biolinnean\/blae097","article-title":"The largest to the south? Assessment of the latitudinal variation in Mustelus schmitti (Chondrichthyes: Carcharhiniforms: Triakidae), an endemic coastal shark of the Southwest Atlantic Ocean","volume":"143","author":"Leyton","year":"2024","journal-title":"Biol. J. Linn. Soc."},{"key":"2026012102595542600_R55","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0067596","article-title":"Decadal changes in the world\u2019s coastal latitudinal temperature gradients","volume":"8","author":"Baumann","year":"2013","journal-title":"PLoS One"},{"key":"2026012102595542600_R56","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1146\/annurev-marine-010814-015656","article-title":"The role of the Gulf Stream in European climate","volume":"7","author":"Palter","year":"2015","journal-title":"Ann. Rev. Mar. Sci."},{"key":"2026012102595542600_R57","doi-asserted-by":"crossref","DOI":"10.1029\/2021GL096683","article-title":"Understanding differences in North Atlantic poleward ocean heat transport and its variability in global climate models","volume":"49","author":"Madonna","year":"2022","journal-title":"Geophys. Res. Lett."},{"key":"2026012102595542600_R58","article-title":"Supplementary material from: The Iberian fossil record of \u2020Otodus megalodon rejects Mediterranean dwarfism and supports nursery use. Figshare.","author":"Herraiz","year":"2025"}],"container-title":["Biology Letters"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article-pdf\/doi\/10.1098\/rsbl.2025.0640\/5655236\/rsbl.2025.0640.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article-pdf\/doi\/10.1098\/rsbl.2025.0640\/5655236\/rsbl.2025.0640.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,21]],"date-time":"2026-01-21T08:00:14Z","timestamp":1768982414000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rsbl\/article\/22\/1\/20250640\/479650\/The-Iberian-fossil-record-of-Otodus-megalodon"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,21]]},"references-count":58,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2026,1,21]]}},"URL":"https:\/\/doi.org\/10.1098\/rsbl.2025.0640","relation":{"has-review":[{"id-type":"doi","id":"10.1098\/RSBL.2025.0640\/v1\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1098\/RSBL.2025.0640\/v2\/response1","asserted-by":"object"},{"id-type":"doi","id":"10.1098\/RSBL.2025.0640\/v1\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1098\/RSBL.2025.0640\/v1\/review1","asserted-by":"object"},{"id-type":"doi","id":"10.1098\/RSBL.2025.0640\/v2\/decision1","asserted-by":"object"}]},"ISSN":["1744-957X"],"issn-type":[{"value":"1744-957X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,21]]},"article-number":"20250640"}}