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Theory developed to explain the evolution of one such trait, anisogamy, has recently been adapted to explain its superorganismal analogue: large egg-like queens and small sperm-like males. To test this theory with comparative data, we first extended it to determine if changing how multiqueen colonies arise affects the model\u2019s predictions. We then used data from 732 ant species to investigate the effects of colony size, worker caste number, and queen number on queen size (thorax volume) and queen\u2013male dimorphism. Queen size and queen\u2013male size dimorphism both increased with colony size and number of worker castes, consistent with predictions. Contrary to predictions, queen size and queen\u2013male size dimorphism were not associated with queen number. To further understand the commonalities and idiosyncrasies of evolution at different hierarchical levels, future work should consider the adaptive and nonadaptive causes for correlated evolution between queen and male, and egg and sperm sizes.<\/jats:p>","DOI":"10.1093\/evolut\/qpag120","type":"journal-article","created":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T15:46:49Z","timestamp":1782920809000},"page":"1895-1904","source":"Crossref","is-referenced-by-count":0,"title":["Superorganismal anisogamy: a comparative test of an extended theory"],"prefix":"10.1093","volume":"80","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5286-3153","authenticated-orcid":false,"given":"Philip A","family":"Downing","sequence":"first","affiliation":[{"name":"Ecology and Genetics Research Unit, University of Oulu , Oulu ,","place":["Finland"]},{"name":"Animal Ecology, Department of Ecology and Genetics, Uppsala University , Uppsala ,","place":["Sweden"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jussi","family":"Lehtonen","sequence":"additional","affiliation":[{"name":"Department of Biological and Environmental Science, University of Jyv\u00e4skyl\u00e4 , Jyv\u00e4skyl\u00e4 ,","place":["Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Louis","family":"Bell-Roberts","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Oxford , Oxford ,","place":["UK"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Heikki","family":"Helanter\u00e4","sequence":"additional","affiliation":[{"name":"Ecology and Genetics Research Unit, University of Oulu , Oulu ,","place":["Finland"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,7,2]]},"reference":[{"key":"2026093011452696000_bib1","doi-asserted-by":"publisher","first-page":"1959","DOI":"10.1038\/s41559-024-02512-7","article-title":"Larger colony sizes favoured the evolution of more worker castes in ants","volume":"8","author":"Bell-Roberts","year":"2024","journal-title":"Nature Ecology & Evolution"},{"key":"2026093011452696000_bib2","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1111\/evo.13117","article-title":"Defensive traits exhibit an evolutionary trade-off and drive diversification in ants","volume":"71","author":"Blanchard","year":"2017","journal-title":"Evolution"},{"key":"2026093011452696000_bib3","doi-asserted-by":"publisher","DOI":"10.1093\/oso\/9780198746171.001.0001","volume-title":"Domains and major transitions of social evolution","author":"Boomsma","year":"2022"},{"key":"2026093011452696000_bib4","doi-asserted-by":"publisher","first-page":"395","DOI":"10.1146\/annurev.ento.50.071803.130416","article-title":"The evolution of male traits in social insects","volume":"50","author":"Boomsma","year":"2005","journal-title":"Annual Review of Entomology"},{"key":"2026093011452696000_bib5","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/j.anbehav.2014.03.005","article-title":"The evolution of multiqueen breeding in eusocial lineages with permanent physically differentiated castes","volume":"92","author":"Boomsma","year":"2014","journal-title":"Animal Behaviour"},{"key":"2026093011452696000_bib6","doi-asserted-by":"publisher","first-page":"e2026029118","DOI":"10.1073\/pnas.2026029118","article-title":"The evolution of social parasitism in Formica ants revealed by a global phylogeny","volume":"118","author":"Borowiec","year":"2021","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"2026093011452696000_bib7","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1046\/j.1420-9101.1999.00028.x","article-title":"Colony size, social complexity and reproductive conflict in social insects","volume":"12","author":"Bourke","year":"1999","journal-title":"Journal of Evolutionary Biology"},{"key":"2026093011452696000_bib8","doi-asserted-by":"publisher","DOI":"10.1093\/acprof:oso\/9780199231157.001.0001","volume-title":"Principles of social evolution","author":"Bourke","year":"2011"},{"key":"2026093011452696000_bib9","volume-title":"Social evolution in ants","author":"Bourke","year":"1995"},{"key":"2026093011452696000_bib10","doi-asserted-by":"publisher","first-page":"2381","DOI":"10.1098\/rspb.2002.2161","article-title":"The evolution of anisogamy: A game-theoretic approach","volume":"269","author":"Bulmer","year":"2002","journal-title":"Proceedings of the Royal Society of London. 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