{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T04:01:41Z","timestamp":1778299301333,"version":"3.51.4"},"reference-count":124,"publisher":"Springer Science and Business Media LLC","issue":"3","license":[{"start":{"date-parts":[[2023,3,6]],"date-time":"2023-03-06T00:00:00Z","timestamp":1678060800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2023,3,6]],"date-time":"2023-03-06T00:00:00Z","timestamp":1678060800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","award":["61408"],"award-info":[{"award-number":["61408"]}],"id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","award":["61408"],"award-info":[{"award-number":["61408"]}],"id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","award":["61408"],"award-info":[{"award-number":["61408"]}],"id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","award":["61408"],"award-info":[{"award-number":["61408"]}],"id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000925","name":"John Templeton Foundation","doi-asserted-by":"publisher","award":["61408"],"award-info":[{"award-number":["61408"]}],"id":[{"id":"10.13039\/100000925","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000270","name":"RCUK | Natural Environment Research Council","doi-asserted-by":"publisher","award":["studentship 2276912"],"award-info":[{"award-number":["studentship 2276912"]}],"id":[{"id":"10.13039\/501100000270","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000270","name":"RCUK | Natural Environment Research Council","doi-asserted-by":"publisher","award":["studentship 2276912"],"award-info":[{"award-number":["studentship 2276912"]}],"id":[{"id":"10.13039\/501100000270","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000268","name":"RCUK | Biotechnology and Biological Sciences Research Council","doi-asserted-by":"publisher","award":["BB\/K015702\/1"],"award-info":[{"award-number":["BB\/K015702\/1"]}],"id":[{"id":"10.13039\/501100000268","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000268","name":"RCUK | Biotechnology and Biological Sciences Research Council","doi-asserted-by":"publisher","award":["BB\/K006754\/1"],"award-info":[{"award-number":["BB\/K006754\/1"]}],"id":[{"id":"10.13039\/501100000268","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Ecol Evol"],"abstract":"<jats:title>Abstract<\/jats:title><jats:p>Complexity, defined as the number of parts and their degree of differentiation, is a poorly explored aspect of macroevolutionary dynamics. The maximum anatomical complexity of organisms has undoubtedly increased through evolutionary time. However, it is unclear whether this increase is a purely diffusive process or whether it is at least partly driven, occurring in parallel in most or many lineages and with increases in the minima as well as the means. Highly differentiated and serially repeated structures, such as vertebrae, are useful systems with which to investigate these patterns. We focus on the serial differentiation of the vertebral column in 1,136 extant mammal species, using two indices that quantify complexity as the numerical richness and proportional distribution of vertebrae across presacral regions and a third expressing the ratio between thoracic and lumbar vertebrae. We address three questions. First, we ask whether the distribution of complexity values in major mammal groups is similar or whether clades have specific signatures associated with their ecology. Second, we ask whether changes in complexity throughout the phylogeny are biased towards increases and whether there is evidence of driven trends. Third, we ask whether evolutionary shifts in complexity depart from a uniform Brownian motion model. Vertebral counts, but not complexity indices, differ significantly between major groups and exhibit greater within-group variation than recognized hitherto. We find strong evidence of a trend towards increasing complexity, where higher values propagate further increases in descendant lineages. Several increases are inferred to have coincided with major ecological or environmental shifts. We find support for multiple-rate models of evolution for all complexity metrics, suggesting that increases in complexity occurred in stepwise shifts, with evidence for widespread episodes of recent rapid divergence. Different subclades evolve more complex vertebral columns in different configurations and probably under different selective pressures and constraints, with widespread convergence on the same formulae. Further work should therefore focus on the ecological relevance of differences in complexity and a more detailed understanding of historical patterns.<\/jats:p>","DOI":"10.1038\/s41559-023-01982-5","type":"journal-article","created":{"date-parts":[[2023,3,6]],"date-time":"2023-03-06T17:06:02Z","timestamp":1678122362000},"page":"367-381","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":22,"title":["Divergent vertebral formulae shape the evolution of axial complexity in mammals"],"prefix":"10.1038","volume":"7","author":[{"given":"Yimeng","family":"Li","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3265-8793","authenticated-orcid":false,"given":"Andrew","family":"Brinkworth","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Emily","family":"Green","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8291-7271","authenticated-orcid":false,"given":"Jack","family":"Oyston","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Matthew","family":"Wills","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6151-0704","authenticated-orcid":false,"given":"Marcello","family":"Ruta","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2023,3,6]]},"reference":[{"key":"1982_CR1","doi-asserted-by":"crossref","unstructured":"McShea, D. W. in Evolutionary Developmental Biology: A Reference Guide (eds de la Rosa, L. N. & M\u00fcller, G.) 169\u2013179 (Springer, 2021).","DOI":"10.1007\/978-3-319-32979-6_123"},{"key":"1982_CR2","doi-asserted-by":"publisher","first-page":"728","DOI":"10.1126\/science.1157719","volume":"323","author":"MJ Benton","year":"2009","unstructured":"Benton, M. J. The Red Queen and the Court Jester: species diversity and the role of biotic and abiotic factors through time. Science 323, 728\u2013732 (2009).","journal-title":"Science"},{"key":"1982_CR3","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1146\/annurev.ecolsys.28.1.129","volume":"28","author":"M Foote","year":"1997","unstructured":"Foote, M. The evolution of morphological diversity. Ann. Rev. Ecol. Syst. 28, 129\u2013152 (1997).","journal-title":"Ann. Rev. Ecol. Syst."},{"key":"1982_CR4","doi-asserted-by":"publisher","first-page":"20200199","DOI":"10.1098\/rsbl.2020.0199","volume":"16","author":"T Guillerme","year":"2020","unstructured":"Guillerme, T. et al. Disparities in the analysis of morphological disparity. Biol. Lett. 16, 20200199 (2020).","journal-title":"Biol. Lett."},{"key":"1982_CR5","first-page":"13","volume":"57","author":"RA Fortey","year":"1996","unstructured":"Fortey, R. A., Briggs, D. E. G. & Wills, M. A. The Cambrian evolutionary \u2018explosion\u2019: decoupling cladogenesis from morphological disparity. Biol. J. Linn. Soc. 57, 13\u201333 (1996).","journal-title":"Biol. J. Linn. Soc."},{"key":"1982_CR6","first-page":"20131071","volume":"280","author":"M Ruta","year":"2013","unstructured":"Ruta, M., Angielczyk, K. D., Fr\u00f6bisch, J. & Benton, M. J. Decoupling of morphological disparity and taxic diversity during the adaptive radiation of anomodont therapsids. Proc. Biol. Sci. 280, 20131071 (2013).","journal-title":"Proc. Biol. Sci."},{"key":"1982_CR7","doi-asserted-by":"publisher","first-page":"1665","DOI":"10.1111\/jeb.12164","volume":"26","author":"MJ Hopkins","year":"2013","unstructured":"Hopkins, M. J. Decoupling of taxonomic diversity and morphological disparity during decline of the Cambrian trilobite family Pterocephaliidae. J. Evol. Biol. 26, 1665\u20131676 (2013).","journal-title":"J. Evol. Biol."},{"key":"1982_CR8","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1017\/pab.2020.57","volume":"47","author":"J Wan","year":"2021","unstructured":"Wan, J. et al. Decoupling of morphological disparity and taxonomic diversity during the end-Permian mass extinction. Paleobiology 47, 402\u2013417 (2021).","journal-title":"Paleobiology"},{"key":"1982_CR9","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-021-90962-5","volume":"11","author":"L Cavin","year":"2021","unstructured":"Cavin, L., Piuz, A., Ferrante, C. & Guinot, G. Giant Mesozoic coelacanths (Osteichthyes, Actinistia) reveal high body size disparity decoupled from taxic diversity. Sci. Rep. 11, 11812 (2021).","journal-title":"Sci. Rep."},{"key":"1982_CR10","doi-asserted-by":"publisher","first-page":"3428","DOI":"10.1073\/pnas.1113870109","volume":"109","author":"DW Bapst","year":"2012","unstructured":"Bapst, D. W., Bullock, P. C., Melchin, M. J., Sheets, H. D., & Mitchell, C. E. Graptoloid diversity and disparity became decoupled during the Ordovician mass extinction. Proc. Natl Acad. Sci. USA 109, 3428\u20133433 (2012).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR11","doi-asserted-by":"publisher","first-page":"1085","DOI":"10.1002\/bies.10192","volume":"24","author":"C Adami","year":"2002","unstructured":"Adami, C. What is complexity? BioEssays 24, 1085\u20131094 (2002).","journal-title":"BioEssays"},{"key":"1982_CR12","doi-asserted-by":"publisher","first-page":"E8909","DOI":"10.1073\/pnas.1810575115","volume":"115","author":"B Deline","year":"2018","unstructured":"Deline, B. et al. Evolution of metazoan morphological disparity. Proc. Natl Acad. Sci. USA 115, E8909\u2013E8918 (2018).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR13","first-page":"jgs2020","volume":"178","author":"DH Erwin","year":"2021","unstructured":"Erwin, D. H. Developmental capacity and the early evolution of animals. J. Geol. Soc. Soc. 178, jgs2020\u2013jgs2245 (2021).","journal-title":"J. Geol. Soc. Soc."},{"key":"1982_CR14","doi-asserted-by":"publisher","first-page":"1225","DOI":"10.1016\/j.cub.2021.07.071","volume":"31","author":"RB Benson","year":"2021","unstructured":"Benson, R. B., Butler, R., Close, R. A., Saupe, E. & Rabosky, D. L. Biodiversity across space and time in the fossil record. Curr. Biol. 31, 1225\u20131236 (2021).","journal-title":"Curr. Biol."},{"key":"1982_CR15","doi-asserted-by":"publisher","first-page":"2207","DOI":"10.1111\/evo.14078","volume":"74","author":"MN Puttick","year":"2020","unstructured":"Puttick, M. N., Guillerme, T. & Wills, M. A. The complex effects of mass extinctions on morphological disparity. Evolution 74, 2207\u20132220 (2020).","journal-title":"Evolution"},{"key":"1982_CR16","first-page":"20203105","volume":"288","author":"M Wang","year":"2021","unstructured":"Wang, M., Lloyd, G. T., Zhang, C. & Zhou, Z. The patterns and modes of the evolution of disparity in Mesozoic birds. Proc. Biol. Sci. 288, 20203105 (2021).","journal-title":"Proc. Biol. Sci."},{"key":"1982_CR17","doi-asserted-by":"publisher","first-page":"717","DOI":"10.1111\/pala.12482","volume":"63","author":"HG Ferron","year":"2020","unstructured":"Ferron, H. G. et al. Categorical versus geometric morphometric approaches to characterizing the evolution of morphological disparity in Osteostraci (Vertebrata, stem Gnathostomata). Palaeontology 63, 717\u2013732 (2020).","journal-title":"Palaeontology"},{"key":"1982_CR18","doi-asserted-by":"crossref","unstructured":"Brandon, R. & McShea, D. W. The Missing Two-Thirds of Evolutionary Theory (Cambridge Univ. Press, 2020).","DOI":"10.1017\/9781108591508"},{"key":"1982_CR19","doi-asserted-by":"publisher","first-page":"1368","DOI":"10.1126\/science.abi6984","volume":"373","author":"AB Leslie","year":"2021","unstructured":"Leslie, A. B., Simpson, C. & Mander, L. Reproductive innovations and pulsed rise in plant complexity. Science 373, 1368\u20131372 (2021).","journal-title":"Science"},{"key":"1982_CR20","doi-asserted-by":"publisher","first-page":"1102","DOI":"10.1038\/35059227","volume":"409","author":"SB Carroll","year":"2001","unstructured":"Carroll, S. B. Chance and necessity: the evolution of morphological complexity and diversity. Nature 409, 1102\u20131109 (2001).","journal-title":"Nature"},{"key":"1982_CR21","doi-asserted-by":"publisher","first-page":"200895","DOI":"10.1098\/rsos.200895","volume":"8","author":"N Rebout","year":"2021","unstructured":"Rebout, N. et al. Measuring complexity in organisms and organizations. R. Soc. Open Sci. 8, 200895 (2021).","journal-title":"R. Soc. Open Sci."},{"key":"1982_CR22","doi-asserted-by":"crossref","unstructured":"McShea, D. W. & Brandon, R. N. Biology\u2019s First Law: The Tendency for Diversity and Complexity to Increase in Evolutionary Systems (Univ. of Chicago Press, 2010).","DOI":"10.7208\/chicago\/9780226562278.001.0001"},{"key":"1982_CR23","doi-asserted-by":"publisher","first-page":"20130249","DOI":"10.1098\/rstb.2013.0249","volume":"369","author":"CP Klingenberg","year":"2014","unstructured":"Klingenberg, C. P. Studying morphological integration and modularity at multiple levels: concepts and analysis. Philos. Trans. R. Soc. Lond. B Biol. Sci. 369, 20130249 (2014).","journal-title":"Philos. Trans. R. Soc. Lond. B Biol. Sci."},{"key":"1982_CR24","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1046\/j.1420-9101.2000.00139.x","volume":"13","author":"G Fusco","year":"2001","unstructured":"Fusco, G. & Minelli, A. Measuring morphological complexity of segmented animals: centipedes as model systems. J. Evol. Biol. 13, 38\u201346 (2001).","journal-title":"J. Evol. Biol."},{"key":"1982_CR25","doi-asserted-by":"publisher","first-page":"4786","DOI":"10.1073\/pnas.0709378105","volume":"105","author":"SJ Adamowicz","year":"2008","unstructured":"Adamowicz, S. J., Purvis, A. & Wills, M. A. Increasing morphological complexity in multiple parallel lineages of the Crustacea. Proc. Natl Acad. Sci. USA 105, 4786\u20134791 (2008).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR26","doi-asserted-by":"publisher","first-page":"8678","DOI":"10.1073\/pnas.1505596112","volume":"112","author":"J Yang","year":"2015","unstructured":"Yang, J. et al. A superarmored lobopodian from the Cambrian of China and early disparity in the evolution of Onychophora. Proc. Natl Acad. Sci. USA 112, 8678\u20138683 (2015).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR27","doi-asserted-by":"publisher","first-page":"6001","DOI":"10.1038\/s41467-021-26285-w","volume":"12","author":"F Lafuma","year":"2021","unstructured":"Lafuma, F., Corfe, I. J., Clavel, J. & Di-Po\u00ef, N. Multiple evolutionary origins and losses of tooth complexity in squamates. Nat. Commun. 12, 6001 (2021).","journal-title":"Nat. Commun."},{"key":"1982_CR28","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1038\/s41559-020-1094-9","volume":"4","author":"KE Jones","year":"2020","unstructured":"Jones, K. E., Gonzalez, S., Angielczyk, K. D. & Pierce, S. E. Regionalization of the axial skeleton predates functional adaptation in the forerunners of mammals. Nat. Ecol. Evol. 4, 470\u2013478 (2020).","journal-title":"Nat. Ecol. Evol."},{"key":"1982_CR29","doi-asserted-by":"crossref","unstructured":"Felice, R. N. & Angielczyk, K. D. in Early Evolutionary History of the Synapsida (eds Kammerer, C. F. et al.) 25\u201351 (Springer, 2014).","DOI":"10.1007\/978-94-007-6841-3_3"},{"key":"1982_CR30","doi-asserted-by":"crossref","unstructured":"Figueirido, B. et al. Body-axis organization in tetrapods: a model-system to disentangle the developmental origins of convergent evolution in deep time. Biol. Lett. 18, 20220047.","DOI":"10.1098\/rsbl.2022.0047"},{"key":"1982_CR31","doi-asserted-by":"publisher","first-page":"23","DOI":"10.1111\/j.1475-4983.2006.00612.x","volume":"50","author":"MJ Benton","year":"2007","unstructured":"Benton, M. J. & Emerson, B. C. How did life become so diverse? The dynamics of diversification according to the fossil record and molecular phylogenetics. Palaeontology 50, 23\u201340 (2007).","journal-title":"Palaeontology"},{"key":"1982_CR32","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1093\/aob\/mcv135","volume":"117","author":"JW Oyston","year":"2016","unstructured":"Oyston, J. W., Hughes, M., Gerber, S. & Wills, M. A. Why should we investigate the morphological disparity of plant clades? Ann. Bot. 117, 859\u2013879 (2016).","journal-title":"Ann. Bot."},{"key":"1982_CR33","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1017\/S0094837300012641","volume":"20","author":"JW Valentine","year":"1994","unstructured":"Valentine, J. W., Collins, A. G. & Meyer, C. P. Morphological complexity increase in metazoans. Paleobiology 20, 131\u2013142 (1994).","journal-title":"Paleobiology"},{"key":"1982_CR34","doi-asserted-by":"publisher","first-page":"1","DOI":"10.2307\/2407115","volume":"27","author":"SM Stanley","year":"1973","unstructured":"Stanley, S. M. An explanation for Copeas rule. Evolution 27, 1\u201326 (1973).","journal-title":"Evolution"},{"key":"1982_CR35","doi-asserted-by":"publisher","first-page":"731","DOI":"10.1126\/science.280.5364.731","volume":"280","author":"J Alroy","year":"1998","unstructured":"Alroy, J. Cope\u2019s rule and the dynamics of body mass evolution in North American fossil mammals. Science 280, 731\u2013734 (1998).","journal-title":"Science"},{"key":"1982_CR36","doi-asserted-by":"publisher","first-page":"98","DOI":"10.1093\/sysbio\/syv075","volume":"65","author":"F Bokma","year":"2016","unstructured":"Bokma, F. et al. Testing for Dep\u00e9ret\u2019s rule (body size increase) in mammals using combined extinct and extant data. Syst. Biol. 65, 98\u2013108 (2016).","journal-title":"Syst. Biol."},{"key":"1982_CR37","first-page":"747","volume":"15","author":"P Raia","year":"2013","unstructured":"Raia, P. & Fortelius, M. Cope\u2019s law of the unspecialized, Cope\u2019s rule, and weak directionality in evolution. Evol. Ecol. Res. 15, 747\u2013756 (2013).","journal-title":"Evol. Ecol. Res."},{"key":"1982_CR38","doi-asserted-by":"crossref","unstructured":"Lineweaver, C. H., Davies, P. C. W. & Ruse, M (eds) Complexity and the Arrow of Time (Cambridge Univ. Press, 2013).","DOI":"10.1017\/CBO9781139225700"},{"key":"1982_CR39","doi-asserted-by":"publisher","first-page":"1101","DOI":"10.1111\/evo.13735","volume":"73","author":"DW McShea","year":"2019","unstructured":"McShea, D. W., Wang, S. C. & Brandon, R. N. A quantitative formulation of biology\u2019s first law. Evolution 73, 1101\u20131115 (2019).","journal-title":"Evolution"},{"key":"1982_CR40","doi-asserted-by":"publisher","first-page":"1315","DOI":"10.1126\/science.1060852","volume":"292","author":"E Szathm\u00e1ry","year":"2001","unstructured":"Szathm\u00e1ry, E., Jord\u00e1n, F. & P\u00e1l, C. Can genes explain biological complexity? Science 292, 1315\u20131316 (2001).","journal-title":"Science"},{"key":"1982_CR41","doi-asserted-by":"publisher","first-page":"P1","DOI":"10.1186\/gb-2003-5-1-p1","volume":"5","author":"RJ Taft","year":"2003","unstructured":"Taft, R. J. & Mattick, J. S. Increasing biological complexity is positively correlated with the relative genome-wide expansion of non-protein-coding DNA sequences. Gen. Biol 5, P1 (2003).","journal-title":"Gen. Biol"},{"key":"1982_CR42","doi-asserted-by":"publisher","first-page":"276","DOI":"10.1111\/ede.12301","volume":"21","author":"KK Duclos","year":"2019","unstructured":"Duclos, K. K., Hendrikse, J. L. & Jamniczky, H. A. Investigating the evolution and development of biological complexity under the framework of epigenetics. Evol. Dev. 21, 276\u2013293 (2019).","journal-title":"Evol. Dev."},{"key":"1982_CR43","doi-asserted-by":"publisher","first-page":"e1008570","DOI":"10.1371\/journal.pcbi.1008570","volume":"21","author":"PF Hagolani","year":"2021","unstructured":"Hagolani, P. F., Zimm, R., Vroomans, R. & Salazar-Ciudad, I. On the evolution and development of morphological complexity: a view from gene regulatory networks. PLoS Comput. Biol. 21, e1008570 (2021).","journal-title":"PLoS Comput. Biol."},{"key":"1982_CR44","doi-asserted-by":"publisher","first-page":"1747","DOI":"10.2307\/2410505","volume":"48","author":"DW McShea","year":"1994","unstructured":"McShea, D. W. Mechanisms of large\u2010scale evolutionary trends. Evolution 48, 1747\u20131763 (1994).","journal-title":"Evolution"},{"key":"1982_CR45","doi-asserted-by":"publisher","first-page":"849","DOI":"10.1554\/0014-3820(2001)055[0849:QPADLS]2.0.CO;2","volume":"55","author":"SC Wang","year":"2001","unstructured":"Wang, S. C. Quantifying passive and driven large\u2010scale evolutionary trends. Evolution 55, 849\u2013858 (2001).","journal-title":"Evolution"},{"key":"1982_CR46","doi-asserted-by":"publisher","first-page":"91","DOI":"10.1002\/jez.b.21029","volume":"304","author":"Y Narita","year":"2005","unstructured":"Narita, Y. & Kuratani, S. Evolution of the vertebral formulae in mammals: a perspective on developmental constraints. J. Exp. Zool. B Mol. Dev. Evol 304, 91\u2013106 (2005).","journal-title":"J. Exp. Zool. B Mol. Dev. Evol"},{"key":"1982_CR47","doi-asserted-by":"publisher","first-page":"44","DOI":"10.5603\/FM.a2017.0064","volume":"77","author":"T Kawashima","year":"2018","unstructured":"Kawashima, T., Thorington, R. W. Jr, Bohaska, P. W. & Sato, F. Variability and constraint of vertebral formulae and proportions in colugos, tree shrews, and rodents, with special reference to vertebral modification by aerodynamic adaptation. Folia Morphol. 77, 44\u201356 (2018).","journal-title":"Folia Morphol."},{"key":"1982_CR48","doi-asserted-by":"publisher","first-page":"2118","DOI":"10.1073\/pnas.0912622107","volume":"107","author":"J M\u00fcller","year":"2010","unstructured":"M\u00fcller, J. et al. Homeotic effects, somitogenesis and the evolution of vertebral numbers in recent and fossil amniotes. Proc. Natl Acad. Sci. USA 107, 2118\u20132123 (2010).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR49","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/S1477200006002258","volume":"5","author":"MR S\u00e1nchez\u2010Villagra","year":"2007","unstructured":"S\u00e1nchez\u2010Villagra, M. R., Narita, Y. & Kuratani, S. Thoracolumbar vertebral number: the first skeletal synapomorphy for afrotherian mammals. Syst. Biodiv. 5, 1\u20137 (2007).","journal-title":"Syst. Biodiv."},{"key":"1982_CR50","doi-asserted-by":"publisher","first-page":"18903","DOI":"10.1073\/pnas.1010335107","volume":"107","author":"L Hautier","year":"2010","unstructured":"Hautier, L., Weisbecker, V., S\u00e1nchez-Villagra, M. R., Goswami, A. & Asher, R. J. Skeletal development in sloths and the evolution of mammalian vertebral patterning. Proc. Natl Acad. Sci. USA 107, 18903\u201318908 (2010).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR51","doi-asserted-by":"publisher","first-page":"173","DOI":"10.1007\/s10914-020-09506-9","volume":"28","author":"P Arnold","year":"2021","unstructured":"Arnold, P. Evolution of the mammalian neck from developmental, morpho-functional, and paleontological perspectives. J. Mammal. Evol. 28, 173\u2013183 (2021).","journal-title":"J. Mammal. Evol."},{"key":"1982_CR52","doi-asserted-by":"publisher","first-page":"949","DOI":"10.1038\/s41559-019-0894-2","volume":"3","author":"SA Williams","year":"2019","unstructured":"Williams, S. A. et al. Increased variation in numbers of presacral vertebrae in suspensory mammals. Nat. Ecol. Evol. 3, 949\u2013956 (2019).","journal-title":"Nat. Ecol. Evol."},{"key":"1982_CR53","doi-asserted-by":"publisher","first-page":"853","DOI":"10.1002\/bies.200900053","volume":"31","author":"RJ Asher","year":"2009","unstructured":"Asher, R. J., Bennett, N. & Lehmann, T. The new framework for understanding placental mammal evolution. Bioessays 31, 853\u2013864 (2009).","journal-title":"Bioessays"},{"key":"1982_CR54","doi-asserted-by":"crossref","unstructured":"Jones, K. E., Angielczyk, K. D. & Pierce, S. E. Stepwise shifts underlie evolutionary trends in morphological complexity of the mammalian vertebral column. Nat. Comm. 10, 5071 (2019).","DOI":"10.1038\/s41467-019-13026-3"},{"key":"1982_CR55","doi-asserted-by":"crossref","unstructured":"Figueirido, B. et al. Serial disparity in the carnivoran backbone unveils a complex adaptive role in metameric evolution. Comm. Biol. 4, 863 (2021).","DOI":"10.1038\/s42003-021-02346-0"},{"key":"1982_CR56","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1002\/jez.b.21316","volume":"314","author":"MA McCollum","year":"2010","unstructured":"McCollum, M. A., Rosenman, B. A., Suwa, G., Meindl, R. S. & Lovejoy, C. O. The vertebral formula of the last common ancestor of African apes and humans. J. Exp. Zool. B Mol. Dev. Evol 314, 123\u2013134 (2010).","journal-title":"J. Exp. Zool. B Mol. Dev. Evol"},{"key":"1982_CR57","doi-asserted-by":"publisher","first-page":"175","DOI":"10.1017\/S0952836901000164","volume":"253","author":"EA Buchholtz","year":"2001","unstructured":"Buchholtz, E. A. Vertebral osteology and swimming style in living and fossil whales (Order: Cetacea). J. Zool. 253, 175\u2013190 (2001).","journal-title":"J. Zool."},{"key":"1982_CR58","doi-asserted-by":"crossref","unstructured":"Jones, K. E., Benitez, L., Angielczyk, K. D. & Pierce, S. E. Adaptation and constraint in the evolution of the mammalian backbone. BMC Evol. Biol. 18, 172 (2018).","DOI":"10.1186\/s12862-018-1282-2"},{"key":"1982_CR59","doi-asserted-by":"publisher","first-page":"1249","DOI":"10.1126\/science.aar3126","volume":"361","author":"KE Jones","year":"2018","unstructured":"Jones, K. E. et al. Fossils reveal the complex evolutionary history of the mammalian regionalized spine. Science 361, 1249\u20131252 (2018).","journal-title":"Science"},{"key":"1982_CR60","unstructured":"Owen, R. Descriptive Catalogue of the Osteological Series Contained in the Museum of the Royal College of Surgeons of England Vol. 2 (Taylor and Francis, 1853)."},{"key":"1982_CR61","doi-asserted-by":"publisher","first-page":"11401","DOI":"10.1073\/pnas.1401392111","volume":"111","author":"F Galis","year":"2014","unstructured":"Galis, F. et al. Fast running restricts evolutionary change of the vertebral column in mammals. Proc. Natl Acad. Sci. USA 111, 11401\u201311406 (2014).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR62","doi-asserted-by":"publisher","first-page":"278","DOI":"10.1111\/j.1525-142X.2007.00160.x","volume":"9","author":"EA Buchholtz","year":"2007","unstructured":"Buchholtz, E. A. Modular evolution of the cetacean vertebral column. Evol. Dev. 9, 278\u2013289 (2007).","journal-title":"Evol. Dev."},{"key":"1982_CR63","first-page":"20191771","volume":"286","author":"A Gillet","year":"2019","unstructured":"Gillet, A., Fr\u00e9d\u00e9rich, B. & Parmentier, E. Divergent evolutionary morphology of the axial skeleton as a potential key innovation in modern cetaceans. Proc. Biol. Sci. 286, 20191771 (2019).","journal-title":"Proc. Biol. Sci."},{"key":"1982_CR64","doi-asserted-by":"publisher","first-page":"2341","DOI":"10.1590\/0001-3765201720170076","volume":"89","author":"PJ Gaudioso","year":"2017","unstructured":"Gaudioso, P. J., D\u00edaz, M. M. & Barquez, R. M. Morphology of the axial skeleton of seven bat genera (Chiroptera: Phyllostomidae). An. Acad. Bras. Cienc. 89, 2341\u20132358 (2017).","journal-title":"An. Acad. Bras. Cienc."},{"key":"1982_CR65","unstructured":"Walton, D. W. & Walton, G. M. in About Bats (eds Slaughter, B. H. & Walton, D. W.) 93\u2013126 (Southern Methodist Univ. Press, 1970)."},{"key":"1982_CR66","doi-asserted-by":"publisher","first-page":"730","DOI":"10.2307\/2410179","volume":"47","author":"DW McShea","year":"1993","unstructured":"McShea, D. W. Evolutionary change in the morphological complexity of the mammalian vertebral column. Evolution 47, 730\u2013740 (1993).","journal-title":"Evolution"},{"key":"1982_CR67","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1038\/s41586-021-03433-2","volume":"592","author":"F Mao","year":"2021","unstructured":"Mao, F., Zhang, C., Liu, C. & Meng, J. Fossoriality and evolutionary development in two Cretaceous mammaliamorphs. Nature 592, 577\u2013582 (2021).","journal-title":"Nature"},{"key":"1982_CR68","doi-asserted-by":"publisher","first-page":"e3000494","DOI":"10.1371\/journal.pbio.3000494","volume":"17","author":"NS Upham","year":"2019","unstructured":"Upham, N. S., Esselstyn, J. A. & Jetz, W. Inferring the mammal tree: species-level sets of phylogenies for questions in ecology, evolution, and conservation. PLoS Biol. 17, e3000494 (2019).","journal-title":"PLoS Biol."},{"key":"1982_CR69","doi-asserted-by":"publisher","first-page":"2216","DOI":"10.1093\/bioinformatics\/btu181","volume":"30","author":"M Pennell","year":"2014","unstructured":"Pennell, M. et al. geiger v2.0: an expanded suite of methods for fitting macroevolutionary models to phylogenetic trees. Bioinformatics 30, 2216\u20132218 (2014).","journal-title":"Bioinformatics"},{"key":"1982_CR70","doi-asserted-by":"publisher","first-page":"3578","DOI":"10.1111\/j.1558-5646.2011.01401.x","volume":"65","author":"JM Eastman","year":"2011","unstructured":"Eastman, J. M., Alfaro, M. E., Joyce, P., Hipp, A. L. & Harmon, L. J. A novel comparative method for identifying shifts in the rate of character evolution on trees. Evolution 65, 3578\u20133589 (2011).","journal-title":"Evolution"},{"key":"1982_CR71","doi-asserted-by":"crossref","unstructured":"Brillouin, L. Science and Information Theory (Academic Press, 1956).","DOI":"10.1063\/1.3059856"},{"key":"1982_CR72","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1016\/0022-5193(78)90302-8","volume":"70","author":"RR Laxton","year":"1978","unstructured":"Laxton, R. R. The measure of diversity. J. Theor. Biol. 70, 51\u201367 (1978).","journal-title":"J. Theor. Biol."},{"key":"1982_CR73","doi-asserted-by":"publisher","first-page":"496","DOI":"10.2307\/1934984","volume":"56","author":"RK Peet","year":"1975","unstructured":"Peet, R. K. Relative diversity indices. Ecology 56, 496\u2013498 (1975).","journal-title":"Ecology"},{"key":"1982_CR74","unstructured":"Pielou, E. C. Ecological Diversity (Wiley, 1975)."},{"key":"1982_CR75","unstructured":"Magurran, A. E. Measuring Biological Diversity (Blackwell Publishing, 2004)."},{"key":"1982_CR76","doi-asserted-by":"publisher","first-page":"1164","DOI":"10.1111\/evo.13217","volume":"71","author":"LC Soul","year":"2017","unstructured":"Soul, L. C. & Benson, R. J. Developmental mechanisms of macroevolutionary change in the tetrapod axis: a case study of Sauropterygia. Evolution 71, 1164\u20131177 (2017).","journal-title":"Evolution"},{"key":"1982_CR77","unstructured":"Mangiafico, S. S. An R Companion for the Handbook of Biological Statistics Version 1.3.2 (2015); rcompanion.org\/documents\/RCompanionBioStatistics.pdf"},{"key":"1982_CR78","doi-asserted-by":"publisher","first-page":"265","DOI":"10.1093\/sysbio\/42.3.265","volume":"42","author":"T Garland Jr.","year":"1993","unstructured":"Garland, T. Jr., Dickerman, A. W., Janis, C. M. & Jones, J. A. Phylogenetic analysis of covariance by computer simulation. Syst. Biol. 42, 265\u2013292 (1993).","journal-title":"Syst. Biol."},{"key":"1982_CR79","doi-asserted-by":"publisher","first-page":"70","DOI":"10.1214\/aos\/1176346577","volume":"13","author":"JA Hartigan","year":"1985","unstructured":"Hartigan, J. A. & Hartigan, P. M. The dip test of unimodality.Ann. Stat 13, 70\u201384 (1985).","journal-title":"Ann. Stat"},{"key":"1982_CR80","doi-asserted-by":"publisher","first-page":"754","DOI":"10.1111\/2041-210X.12066","volume":"4","author":"LJ Revell","year":"2013","unstructured":"Revell, L. J. Two new graphical methods for mapping trait evolution on phylogenies. Methods Ecol. Evol. 4, 754\u2013759 (2013).","journal-title":"Methods Ecol. Evol."},{"key":"1982_CR81","first-page":"601","volume":"89","author":"AH Schultz","year":"1945","unstructured":"Schultz, A. H. & Straus, W. L. Jr. The numbers of vertebrae in primates. Proc. Am. Philos. Soc. 89, 601\u2013626 (1945).","journal-title":"Proc. Am. Philos. Soc."},{"key":"1982_CR82","first-page":"33","volume":"6","author":"A Burt","year":"1989","unstructured":"Burt, A. Comparative methods using phylogenetically independent contrasts. Oxf. Surv. Evol. Biol. 6, 33\u201353 (1989).","journal-title":"Oxf. Surv. Evol. Biol."},{"key":"1982_CR83","unstructured":"Orme, D. et al. The caper package: comparative analysis of phylogenetics and evolution in R. R package version 5 https:\/\/mirror.rcg.sfu.ca\/mirror\/CRAN\/web\/packages\/caper\/ (2013)."},{"key":"1982_CR84","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1017\/S1089332600001893","volume":"16","author":"G Hunt","year":"2010","unstructured":"Hunt, G. & Carrano, M. T. Models and methods for analyzing phenotypic evolution in lineages and clades. Paleontol. Soc. Pap. 16, 245\u2013269 (2010).","journal-title":"Paleontol. Soc. Pap."},{"key":"1982_CR85","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1666\/0094-8373(2000)026<0319:UTDOTW>2.0.CO;2","volume":"26","author":"J Alroy","year":"2000","unstructured":"Alroy, J. Understanding the dynamics of trends within evolving lineages. Paleobiology 26, 319\u2013329 (2000).","journal-title":"Paleobiology"},{"key":"1982_CR86","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1666\/06028.1","volume":"33","author":"JD Marcot","year":"2007","unstructured":"Marcot, J. D. & McShea, D. W. Increasing hierarchical complexity throughout the history of life: phylogenetic tests of trend mechanisms. Paleobiology 33, 182\u2013200 (2007).","journal-title":"Paleobiology"},{"key":"1982_CR87","first-page":"477","volume":"50","author":"DW McShea","year":"1996","unstructured":"McShea, D. W. Metazoan complexity and evolution: is there a trend? Evolution 50, 477\u2013492 (1996).","journal-title":"Evolution"},{"key":"1982_CR88","first-page":"43","volume":"274","author":"JAF Diniz-Filho","year":"2007","unstructured":"Diniz-Filho, J. A. F., Rangel, T. F. L., Bini, L. M. & Hawkins, B. A. Macroevolutionary dynamics in environmental space and the latitudinal diversity gradient in New World birds. Proc. Biol. Sci. 274, 43\u201352 (2007).","journal-title":"Proc. Biol. Sci."},{"key":"1982_CR89","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1111\/j.1420-9101.2005.00998.x","volume":"19","author":"M Verd\u00fa","year":"2006","unstructured":"Verd\u00fa, M. Tempo, mode and phylogenetic associations of relative embryo size evolution in angiosperms. J. Evol. Biol. 19, 625\u2013634 (2006).","journal-title":"J. Evol. Biol."},{"key":"1982_CR90","doi-asserted-by":"publisher","first-page":"925","DOI":"10.1023\/A:1006512121434","volume":"12","author":"BA Maurer","year":"1998","unstructured":"Maurer, B. A. The evolution of body size in birds. I. Evidence of non-random diversification. Evol. Ecol. 12, 925\u2013934 (1998).","journal-title":"Evol. Ecol."},{"key":"1982_CR91","unstructured":"Fox, J. Applied Regression Analysis, Linear Models, and Related Models (Sage, 1997)."},{"key":"1982_CR92","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1016\/j.jmva.2016.01.004","volume":"147","author":"A Ghosh","year":"2016","unstructured":"Ghosh, A., Mandal, A., Mart\u00edn, N. & Pardo, L. Influence analysis of robust Wald-type tests. J. Multivar. Anal. 147, 102\u2013126 (2016).","journal-title":"J. Multivar. Anal."},{"key":"1982_CR93","doi-asserted-by":"publisher","first-page":"5093","DOI":"10.1073\/pnas.1419823112","volume":"112","author":"J Baker","year":"2015","unstructured":"Baker, J., Meade, A., Pagel, M. & Venditti, C. Adaptive evolution toward larger size in mammals. Proc. Natl Acad. Sci. USA 112, 5093\u20135098 (2015).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"1982_CR94","first-page":"20152947","volume":"283","author":"KE Jones","year":"2016","unstructured":"Jones, K. E. New insights on equid locomotory evolution from the lumbar region of fossil horses. Proc. Biol. Sci. 283, 20152947 (2016).","journal-title":"Proc. Biol. Sci."},{"key":"1982_CR95","doi-asserted-by":"publisher","first-page":"624","DOI":"10.1002\/ar.20534","volume":"290","author":"EA Buchholtz","year":"2007","unstructured":"Buchholtz, E. A., Booth, A. C. & Webbink, K. E. Vertebral anatomy in the Florida manatee, Trichechus manatus latirostris: a developmental and evolutionary analysis. Anat. Rec. 290, 624\u2013637 (2007).","journal-title":"Anat. Rec."},{"key":"1982_CR96","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1111\/j.1469-7998.1986.tb04706.x","volume":"208","author":"JP Gasc","year":"1986","unstructured":"Gasc, J. P., Jouffroy, F. K., Renous, S. & Blottnitz, F. V. Morphofunctional study of the digging system of the Namib Desert golden mole (Eremitalpa granti namibensis): cinefluorographical and anatomical analysis. J. Zool. 208, 9\u201335 (1986).","journal-title":"J. Zool."},{"key":"1982_CR97","doi-asserted-by":"publisher","first-page":"501","DOI":"10.1006\/jhev.1994.1030","volume":"26","author":"MS Fischer","year":"1994","unstructured":"Fischer, M. S. Crouched posture and high fulcrum, a principle in the locomotion of small mammals: the example of the rock hyrax (Procavia capensis) (Mammalia: Hyracoidea). J. Hum. Evol. 26, 501\u2013524 (1994).","journal-title":"J. Hum. Evol."},{"key":"1982_CR98","unstructured":"Vizca\u00edno, S. F. & Loughry, W. J. Biology of the Xenarthra (Univ. Press of Florida, 2008)."},{"key":"1982_CR99","doi-asserted-by":"crossref","unstructured":"Goswami, A. & Friscia, A. (eds) Carnivoran Evolution: New Views on Phylogeny, Form and Function Vol. 1 (Cambridge Univ. Press, 2010).","DOI":"10.1017\/CBO9781139193436"},{"key":"1982_CR100","doi-asserted-by":"crossref","unstructured":"Pierce, S. E., Clack, J. A. & Hutchinson, J. R. Comparative axial morphology in pinnipeds and its correlation with aquatic locomotory behaviour. J. Anat. 219, 502\u2013514.","DOI":"10.1111\/j.1469-7580.2011.01406.x"},{"key":"1982_CR101","doi-asserted-by":"crossref","unstructured":"Larkey, D. J., Datwyler, S. L. & Lancaster, W. C. in Evolutionary History of Bats (eds Gunnell, G. F. & Simmons, N. B.) 500\u2013529 (Cambridge Univ. Press, 2012).","DOI":"10.1017\/CBO9781139045599.016"},{"key":"1982_CR102","doi-asserted-by":"crossref","unstructured":"L\u00f3pez-Aguirre, C., Hand, S. J., Koyabu, D., Son, N. T. & Wilson, L. A. Postcranial heterochrony, modularity, integration and disparity in the prenatal ossification in bats (Chiroptera). BMC Evol. Biol. 19, 75 (2019).","DOI":"10.1186\/s12862-019-1396-1"},{"key":"1982_CR103","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1111\/mam.12211","volume":"50","author":"SC Anderson","year":"2020","unstructured":"Anderson, S. C. & Ruxton, G. D. The evolution of flight in bats: a novel hypothesis. Mamm. Rev. 50, 426\u2013439 (2020).","journal-title":"Mamm. Rev."},{"key":"1982_CR104","doi-asserted-by":"publisher","first-page":"818","DOI":"10.1038\/nature06549","volume":"451","author":"NB Simmons","year":"2008","unstructured":"Simmons, N. B., Seymour, K. L., Habersetzer, J. & Gunnell, G. F. Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation. Nature 451, 818\u2013821 (2008).","journal-title":"Nature"},{"key":"1982_CR105","doi-asserted-by":"publisher","first-page":"592","DOI":"10.1093\/bioinformatics\/btq706","volume":"27","author":"K Schliep","year":"2011","unstructured":"Schliep, K. phangorn: phylogenetic analysis in R. Bioinformatics 27, 592\u2013593 (2011).","journal-title":"Bioinformatics"},{"key":"1982_CR106","first-page":"459","volume":"11","author":"MK Kuhner","year":"1994","unstructured":"Kuhner, M. K. & Felsenstein, J. A simulation comparison of phylogeny algorithms under equal and unequal evolutionary rates. Mol. Biol. Evol. 11, 459\u2013468 (1994).","journal-title":"Mol. Biol. Evol."},{"key":"1982_CR107","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1016\/0025-5564(81)90043-2","volume":"53","author":"DF Robinson","year":"1981","unstructured":"Robinson, D. F. & Foulds, L. R. Comparison of phylogenetic trees. Math. Biosci. 53, 131\u2013147 (1981).","journal-title":"Math. Biosci."},{"key":"1982_CR108","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1109\/TCBB.2011.157","volume":"9","author":"L Yu","year":"2011","unstructured":"Yu, L., Rajan, V. & Moret, B. M. E. A metric for phylogenetic trees based on matching. IEEE\/ACM Trans. Comput. Biol. Bioinf. 9, 1014\u20131022 (2011).","journal-title":"IEEE\/ACM Trans. Comput. Biol. Bioinf."},{"key":"1982_CR109","doi-asserted-by":"publisher","first-page":"526","DOI":"10.1093\/bioinformatics\/bty633","volume":"35","author":"E Paradis","year":"2019","unstructured":"Paradis, E. & Schliep, K. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics 35, 526\u2013528 (2019).","journal-title":"Bioinformatics"},{"key":"1982_CR110","doi-asserted-by":"publisher","first-page":"1463","DOI":"10.1093\/bioinformatics\/btq166","volume":"26","author":"S Kembel","year":"2010","unstructured":"Kembel, S. et al. Picante: R tools for integrating phylogenies and ecology. Bioinformatics 26, 1463\u20131464 (2010).","journal-title":"Bioinformatics"},{"key":"1982_CR111","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1111\/2041-210X.12628","volume":"8","author":"G Yu","year":"2017","unstructured":"Yu, G., Smith, D., Zhu, H., Guan, Y. & Lam, T. T. ggtree: an R package for visualization and annotation of phylogenetic trees with their covariates and other associated data. Meth. Ecol. Evol. 8, 28\u201336 (2017).","journal-title":"Meth. Ecol. Evol."},{"key":"1982_CR112","unstructured":"Fox, J. & Weisberg, S. An R Companion to Applied Regression 3rd edn (Sage, 2019)."},{"key":"1982_CR113","doi-asserted-by":"publisher","first-page":"1821","DOI":"10.21105\/joss.01821","volume":"4","author":"N Frerebeau","year":"2019","unstructured":"Frerebeau, N. tabula: an R package for analysis, seriation, and visualization of archaeological count data. J. Open Source Soft. 4, 1821 (2019).","journal-title":"J. Open Source Soft."},{"key":"1982_CR114","unstructured":"Adler, D. & Kelly, S. T. vioplot: violin plot. R package version 0.3.6 https:\/\/cran.r-project.org\/web\/packages\/vioplot\/index.html (2021)."},{"key":"1982_CR115","doi-asserted-by":"crossref","unstructured":"Wickham, H. ggplot2: Elegant Graphics for Data Analysis (Springer-Verlag, 2016).","DOI":"10.1007\/978-3-319-24277-4"},{"key":"1982_CR116","doi-asserted-by":"crossref","unstructured":"Venables, W. N. & Ripley, B. D. Modern Applied Statistics with S 4th edn (Springer, 2002).","DOI":"10.1007\/978-0-387-21706-2"},{"key":"1982_CR117","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1080\/00031305.1980.10483031","volume":"34","author":"SR Searle","year":"1980","unstructured":"Searle, S. R., Speed, F. M. & Milliken, G. A. Population marginal means in the linear model: an alternative to least squares means. Am. Stat. 34, 216\u2013221 (1980).","journal-title":"Am. Stat."},{"key":"1982_CR118","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1111\/j.2041-210X.2011.00169.x","volume":"3","author":"LJ Revell","year":"2012","unstructured":"Revell, L. J. phytools: an R package for phylogenetic comparative biology (and other things). Meth. Ecol. Evol. 3, 217\u2013223 (2012).","journal-title":"Meth. Ecol. Evol."},{"key":"1982_CR119","doi-asserted-by":"publisher","first-page":"1243","DOI":"10.1038\/s41559-021-01516-x","volume":"5","author":"N Brocklehurst","year":"2021","unstructured":"Brocklehurst, N. & Benson, R. J. Multiple paths to morphological diversification during the origin of amniotes. Nat. Ecol. Evol. 5, 1243\u20131249 (2021).","journal-title":"Nat. Ecol. Evol."},{"key":"1982_CR120","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1086\/284325","volume":"125","author":"J Felsenstein","year":"1985","unstructured":"Felsenstein, J. Phylogenies and the comparative method. Am. Nat. 125, 1\u201315 (1985).","journal-title":"Am. Nat."},{"key":"1982_CR121","first-page":"246","volume":"15","author":"F Galton","year":"1886","unstructured":"Galton, F. Regression towards mediocrity in hereditary stature. J. Anthropol. Inst. Gt Brit. Irel. 15, 246\u2013263 (1886).","journal-title":"J. Anthropol. Inst. Gt Brit. Irel."},{"key":"1982_CR122","doi-asserted-by":"publisher","first-page":"700","DOI":"10.1086\/497402","volume":"166","author":"C Kelly","year":"2005","unstructured":"Kelly, C. & Price, T. D. Correcting for regression to the mean in behavior and ecology. Am. Nat. 166, 700\u2013707 (2005).","journal-title":"Am. Nat."},{"key":"1982_CR123","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1111\/j.2041-210X.2012.00223.x","volume":"3","author":"DW Bapst","year":"2012","unstructured":"Bapst, D. W. paleotree: an R package for paleontological and phylogenetic analyses of evolution. Meth. Ecol. Evol. 3, 803\u2013807 (2012).","journal-title":"Meth. Ecol. Evol."},{"key":"1982_CR124","doi-asserted-by":"publisher","first-page":"901","DOI":"10.1093\/sysbio\/syy032","volume":"67","author":"A Rambaut","year":"2018","unstructured":"Rambaut, A., Drummond, A. J., Xie, D., Baele, G. & Suchard, M. A. Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Syst. Biol. 67, 901\u2013904 (2018).","journal-title":"Syst. Biol."}],"container-title":["Nature Ecology &amp; Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41559-023-01982-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-023-01982-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-023-01982-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,10,15]],"date-time":"2024-10-15T21:29:44Z","timestamp":1729027784000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41559-023-01982-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,3,6]]},"references-count":124,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2023,3]]}},"alternative-id":["1982"],"URL":"https:\/\/doi.org\/10.1038\/s41559-023-01982-5","relation":{},"ISSN":["2397-334X"],"issn-type":[{"value":"2397-334X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,3,6]]},"assertion":[{"value":"31 January 2022","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"3 January 2023","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 March 2023","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}