{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,19]],"date-time":"2026-09-19T18:14:09Z","timestamp":1789841649204,"version":"4.0.1"},"reference-count":99,"publisher":"Wiley","issue":"1","license":[{"start":{"date-parts":[[2022,11,11]],"date-time":"2022-11-11T00:00:00Z","timestamp":1668124800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"}],"funder":[{"DOI":"10.13039\/501100003130","name":"Fonds Wetenschappelijk Onderzoek","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100003130","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Morphology"],"published-print":{"date-parts":[[2023,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Given the pervasiveness of gene sharing in evolution and the extent of homology across the tree of life, why is everything not homologous with everything else? The continuity and overlapping genetic contributions to diverse traits across lineages seem to imply that no discrete determination of homology is possible. Although some argue that the widespread overlap in parts and processes should be acknowledged as \u201cpartial\u201d homology, this threatens a broad base of presumed comparative morphological knowledge accepted by most biologists. Following a long scientific tradition, we advocate a strategy of \u201ctheoretical articulation\u201d that introduces further distinctions to existing concepts to produce increased contrastive resolution among the labels used to represent biological phenomena. We pursue this strategy by drawing on successful patterns of reasoning from serial homology at the level of gene sequences to generate an enriched characterization of serial homology as a hierarchical, phylogenetic concept. Specifically, we propose that the concept of serial homology should be applied primarily to repeated but developmentally individualized body parts, such as cell types, differentiated body segments, or epidermal appendages. For these characters, a phylogenetic history can be reconstructed, similar to families of paralogous genes, endowing the notion of serial homology with a hierarchical, phylogenetic interpretation. On this basis, we propose a five\u2010fold theoretical classification that permits a more fine\u2010grained mapping of diverse trait\u2010types. This facilitates answering the question of why everything is not homologous with everything else, as well as how novelty is possible given that any new character possesses evolutionary precursors. We illustrate the fecundity of our account by reference to debates over insect wing serial homologs and vertebrate paired appendages.<\/jats:p>","DOI":"10.1002\/jmor.21531","type":"journal-article","created":{"date-parts":[[2022,11,1]],"date-time":"2022-11-01T06:27:35Z","timestamp":1667284055000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":27,"title":["The hierarchical basis of serial homology and evolutionary novelty"],"prefix":"10.1002","volume":"284","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9600-1013","authenticated-orcid":false,"given":"James","family":"DiFrisco","sequence":"first","affiliation":[{"name":"Institute of Philosophy KU Leuven Leuven Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4422-7779","authenticated-orcid":false,"given":"Alan C.","family":"Love","sequence":"additional","affiliation":[{"name":"Department of Philosophy University of Minnesota Minneapolis Minnesota USA"},{"name":"Minnesota Center for Philosophy of Sciences University of Minnesota Minneapolis Minnesota USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3097-002X","authenticated-orcid":false,"given":"G\u00fcnter P.","family":"Wagner","sequence":"additional","affiliation":[{"name":"Department of Ecology and Evolutionary Biology Yale University New Haven Connecticut USA"},{"name":"Yale Systems Biology Institute Yale University New Haven Connecticut USA"},{"name":"Department of Obstetrics, Gynecology and Reproductive Sciences Yale Medical School New Haven Connecticut USA"},{"name":"Department of Obstetrics and Gynecology Wayne State University Detroit Michigan USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2022,11,11]]},"reference":[{"key":"e_1_2_12_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tree.2021.04.002"},{"key":"e_1_2_12_3_1","first-page":"207","volume-title":"Homology","author":"Abouheif E.","year":"1999"},{"key":"e_1_2_12_4_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg2416"},{"key":"e_1_2_12_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2019.01.022"},{"key":"e_1_2_12_6_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg.2016.127"},{"key":"e_1_2_12_7_1","volume-title":"Homology, an unsolved problem","author":"Beer G.","year":"1971"},{"key":"e_1_2_12_8_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365-2907.1991.tb00290.x"},{"key":"e_1_2_12_9_1","first-page":"171","volume-title":"Pragmatism, Science and Naturalism","author":"Brigandt I.","year":"2011"},{"key":"e_1_2_12_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.b.22461"},{"key":"e_1_2_12_11_1","doi-asserted-by":"publisher","DOI":"10.7551\/mitpress\/12389.001.0001"},{"key":"e_1_2_12_12_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41559-020-01349-0"},{"key":"e_1_2_12_13_1","doi-asserted-by":"publisher","DOI":"10.3389\/fimmu.2014.00459"},{"key":"e_1_2_12_14_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-7091-1868-9_2"},{"key":"e_1_2_12_15_1","volume-title":"Gaining ground: The origin and evolution of tetrapods","author":"Clack J. A.","year":"2002"},{"key":"e_1_2_12_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cois.2015.12.005"},{"key":"e_1_2_12_17_1","doi-asserted-by":"publisher","DOI":"10.1007\/s12064-003-0057-4"},{"key":"e_1_2_12_18_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ydbio.2017.03.006"},{"key":"e_1_2_12_19_1","doi-asserted-by":"publisher","DOI":"10.1086\/714959"},{"key":"e_1_2_12_20_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10539-020-09762-2"},{"key":"e_1_2_12_21_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11692-022-09567-z"},{"key":"e_1_2_12_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.semcdb.2022.03.030"},{"key":"e_1_2_12_23_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10539-012-9313-7"},{"key":"e_1_2_12_24_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2021.1808"},{"key":"e_1_2_12_25_1","doi-asserted-by":"publisher","DOI":"10.1007\/s13752-021-00395-6"},{"key":"e_1_2_12_26_1","doi-asserted-by":"publisher","DOI":"10.1101\/gr.5457707"},{"key":"e_1_2_12_27_1","volume-title":"Master control genes in development and evolution: The homeobox story","author":"Gehring W. J.","year":"1998"},{"key":"e_1_2_12_28_1","first-page":"230","volume-title":"Homology: The hierarchical basis of comparative biology","author":"Goodwin B. C.","year":"1994"},{"key":"e_1_2_12_29_1","volume-title":"Evolution of the Insects","author":"Grimaldi D.","year":"2005"},{"key":"e_1_2_12_30_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1004150"},{"key":"e_1_2_12_31_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-319-32979-6_83"},{"key":"e_1_2_12_32_1","volume-title":"Homology: The Hierarchical Basis of Comparative Biology","author":"Hall B. K.","year":"1994"},{"key":"e_1_2_12_33_1","doi-asserted-by":"publisher","DOI":"10.1017\/S1464793102006097"},{"key":"e_1_2_12_34_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00441-019-03096-6"},{"key":"e_1_2_12_35_1","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780198546412.001.0001","volume-title":"The comparative method in evolutionary biology","author":"Harvey P. H.","year":"1991"},{"key":"e_1_2_12_36_1","doi-asserted-by":"publisher","DOI":"10.1242\/dev.129.5.1225"},{"key":"e_1_2_12_37_1","doi-asserted-by":"crossref","DOI":"10.1093\/oso\/9780198540472.001.0001","volume-title":"Early vertebrates","author":"Janvier P.","year":"1996"},{"key":"e_1_2_12_38_1","first-page":"172","volume-title":"Major events in early vertebrate evolution: Palaeontology, phylogeny, genetics and development","author":"Janvier P.","year":"2001"},{"key":"e_1_2_12_39_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2016.05.030"},{"key":"e_1_2_12_40_1","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2021.2682"},{"key":"e_1_2_12_41_1","doi-asserted-by":"publisher","DOI":"10.1139\/z87-352"},{"key":"e_1_2_12_42_1","doi-asserted-by":"publisher","DOI":"10.1080\/00222937008696201"},{"key":"e_1_2_12_43_1","first-page":"799","volume-title":"Encyclopaedia Brittanica, Ninth Edition","author":"Lankester E. R.","year":"1888"},{"key":"e_1_2_12_44_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2100575118"},{"key":"e_1_2_12_45_1","doi-asserted-by":"publisher","DOI":"10.3389\/fcell.2021.695135"},{"key":"e_1_2_12_46_1","doi-asserted-by":"publisher","DOI":"10.1093\/iob\/obac015"},{"key":"e_1_2_12_47_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.thbio.2005.11.006"},{"key":"e_1_2_12_48_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10539-007-9093-7"},{"key":"e_1_2_12_49_1","volume-title":"Principles of systematic zoology","author":"Mayr E.","year":"1969"},{"key":"e_1_2_12_50_1","first-page":"1","volume-title":"Current Topics in Developmental Biology","author":"McKenna K. Z.","year":"2021"},{"key":"e_1_2_12_51_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1525-142X.2005.05021.x"},{"key":"e_1_2_12_52_1","doi-asserted-by":"publisher","DOI":"10.1006\/mpev.1997.0490"},{"key":"e_1_2_12_53_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1525-142x.2000.00054.x"},{"key":"e_1_2_12_54_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1525-142X.2002.01060.x"},{"key":"e_1_2_12_55_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-94-007-6537-5_15"},{"key":"e_1_2_12_56_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1420-9101.1991.4030429.x"},{"key":"e_1_2_12_57_1","doi-asserted-by":"publisher","DOI":"10.3389\/fevo.2016.00071"},{"key":"e_1_2_12_58_1","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:oso\/9780199671427.003.0014"},{"key":"e_1_2_12_59_1","doi-asserted-by":"crossref","first-page":"51","DOI":"10.7551\/mitpress\/5182.001.0001","volume-title":"Origination of organismal form: Beyond the gene in developmental and evolutionary biology","author":"M\u00fcller G. B.","year":"2003"},{"key":"e_1_2_12_60_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.es.22.110191.001305"},{"key":"e_1_2_12_61_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature19322"},{"key":"e_1_2_12_62_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1525-142X.2010.00402.x"},{"key":"e_1_2_12_63_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-0031.2011.00371.x"},{"key":"e_1_2_12_64_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10539-018-9617-3"},{"key":"e_1_2_12_65_1","doi-asserted-by":"publisher","DOI":"10.1023\/A:1022896515287"},{"key":"e_1_2_12_66_1","doi-asserted-by":"publisher","DOI":"10.1038\/ni.3320"},{"key":"e_1_2_12_67_1","volume-title":"Lectures on the comparative anatomy and physiology of the invertebrate animals","author":"Owen R.","year":"1843"},{"key":"e_1_2_12_68_1","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.118611"},{"key":"e_1_2_12_69_1","first-page":"21","volume-title":"Problems in Phylogenetic Reconstruction","author":"Patterson C.","year":"1982"},{"key":"e_1_2_12_70_1","doi-asserted-by":"publisher","DOI":"10.1093\/icb\/icac026"},{"key":"e_1_2_12_71_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg.2016.167"},{"key":"e_1_2_12_72_1","doi-asserted-by":"publisher","DOI":"10.4159\/9780674042124"},{"key":"e_1_2_12_73_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1096-0031.1991.tb00045.x"},{"key":"e_1_2_12_74_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg928"},{"key":"e_1_2_12_75_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature09977"},{"key":"e_1_2_12_76_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00018-022-04382-3"},{"key":"e_1_2_12_77_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1095-8312.1984.tb00796.x"},{"key":"e_1_2_12_78_1","first-page":"1","volume-title":"Ontogeny and Systematics","author":"Roth V. L.","year":"1988"},{"key":"e_1_2_12_79_1","doi-asserted-by":"publisher","DOI":"10.2307\/2418787"},{"key":"e_1_2_12_80_1","first-page":"423","volume-title":"Homology: The Hierachical Basis of Comparative Biology","author":"Sattler R.","year":"1994"},{"key":"e_1_2_12_81_1","first-page":"317","volume-title":"Challenging the modern synthesis","author":"Schmitt S.","year":"2017"},{"key":"e_1_2_12_82_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)81681-5"},{"key":"e_1_2_12_83_1","doi-asserted-by":"publisher","DOI":"10.1038\/41710"},{"key":"e_1_2_12_84_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature07891"},{"key":"e_1_2_12_85_1","doi-asserted-by":"publisher","DOI":"10.1093\/icb\/36.1.51"},{"key":"e_1_2_12_86_1","doi-asserted-by":"publisher","DOI":"10.7312\/simp92414"},{"key":"e_1_2_12_87_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.21236"},{"key":"e_1_2_12_88_1","first-page":"163","volume-title":"Allgemeine Biologie","author":"Spemann H.","year":"1915"},{"key":"e_1_2_12_89_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.shpsc.2014.03.001"},{"key":"e_1_2_12_90_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-11215-8"},{"key":"e_1_2_12_91_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1525-142X.2012.00561.x"},{"key":"e_1_2_12_92_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg2099"},{"key":"e_1_2_12_93_1","doi-asserted-by":"publisher","DOI":"10.1515\/9781400851461"},{"key":"e_1_2_12_94_1","doi-asserted-by":"publisher","DOI":"10.1002\/jez.1100"},{"key":"e_1_2_12_95_1","volume-title":"Physics avoidance: Essays in conceptual strategy","author":"Wilson M.","year":"2018"},{"key":"e_1_2_12_96_1","first-page":"122","volume-title":"Major Transitions in Vertebrate Evolution","author":"Wilson M. V. H.","year":"2007"},{"key":"e_1_2_12_97_1","doi-asserted-by":"publisher","DOI":"10.2307\/j.ctv1pncnrh"},{"key":"e_1_2_12_98_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrg2681"},{"key":"e_1_2_12_99_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsbl.2011.1033"},{"key":"e_1_2_12_100_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.77.4.2158"}],"container-title":["Journal of Morphology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.21531","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/jmor.21531","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/jmor.21531","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,11,29]],"date-time":"2023-11-29T17:03:48Z","timestamp":1701277428000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/jmor.21531"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,11,11]]},"references-count":99,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2023,1]]}},"alternative-id":["10.1002\/jmor.21531"],"URL":"https:\/\/doi.org\/10.1002\/jmor.21531","archive":["Portico"],"relation":{"has-review":[{"id-type":"doi","id":"10.1002\/JMOR.21531\/v2\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1002\/JMOR.21531\/v1\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1002\/JMOR.21531\/v1\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1002\/JMOR.21531\/v1\/review1","asserted-by":"object"}]},"ISSN":["0362-2525","1097-4687"],"issn-type":[{"value":"0362-2525","type":"print"},{"value":"1097-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,11,11]]},"assertion":[{"value":"2022-08-06","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-10-29","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-11-11","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e21531"}}