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This is because ribosomal gene (rDNA) copy number and activity mediate protein biosynthesis, stress response, ageing, disease, dosage compensation and genome stability. Here, we found contrasting dosage compensation of sex\u2010linked NORs in turtles with male and female heterogamety. Most taxa examined exhibit homomorphic rRNA gene clusters in a single autosome pair (determined by 28S rDNA fluorescence in situ hybridization), whereas NORs are sex\u2010linked in <jats:italic>Apalone spinifera<\/jats:italic>, <jats:italic>Pelodiscus sinensis<\/jats:italic> and <jats:italic>Staurotypus triporcatus<\/jats:italic>. Full\u2010dosage compensation upregulates the male X\u2010NOR (determined via silver staining\u2014AgNOR) in <jats:italic>Staurotypus<\/jats:italic> (who lacks Y\u2010NOR) compared with female X\u2010AgNORs. In softshell <jats:italic>Apalone<\/jats:italic> and <jats:italic>Pelodiscus<\/jats:italic>, who share homologous ZZ\/ZW micro\u2010chromosomes, their enlarged W\u2010NOR is partially active (due to 28S rDNA invasion by R2 retroelements), whereas their smaller Z\u2010NOR is silent in females but active in both male\u2010Zs (presumably because the W\u2010NOR meets cellular demands and excessive NOR activity is costly). We hypothesize that R2 disruption favoured W enlargement to add intact 28S\u2010units, perhaps facilitated by reduced recombination during sex chromosome evolution. The molecular basis of the potentially adaptive female Z\u2010silencing is likely intricate and perhaps epigenetic, as non\u2010ribosomal Z genes are active in <jats:italic>Apalone<\/jats:italic> females. Yet, <jats:italic>Emydura maquarii<\/jats:italic> exhibit identical heteromorphism in their autosomal NOR (R2 invaded 28S\u2010units and the small\u2010autosome NOR is silent), suggesting that the softshell turtle pattern can evolve independent of sex chromosome evolution. Our study illuminates the complex sex chromosome evolution and dosage compensation of non\u2010model systems that challenges classic paradigms.<\/jats:p>","DOI":"10.1111\/jeb.14064","type":"journal-article","created":{"date-parts":[[2022,7,25]],"date-time":"2022-07-25T16:51:06Z","timestamp":1658767866000},"page":"1709-1720","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":10,"title":["Evolution and dosage compensation of nucleolar organizing regions (NORs) mediated by mobile elements in turtles with female (<scp>ZZ<\/scp>\/<scp>ZW<\/scp>) but not with male (<scp>XX<\/scp>\/<scp>XY<\/scp>) heterogamety"],"prefix":"10.1093","volume":"35","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2461-9268","authenticated-orcid":false,"given":"Eugenia E.","family":"Montiel","sequence":"first","affiliation":[{"name":"Department of Ecology, Evolution, and Organismal Biology Iowa State University  Ames Iowa USA"},{"name":"Department of Experimental Biology (Genetics Area) University of Ja\u00e9n  Ja\u00e9n Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0956-5011","authenticated-orcid":false,"given":"Daleen","family":"Badenhorst","sequence":"additional","affiliation":[{"name":"Department of Ecology, Evolution, and Organismal Biology Iowa State University  Ames Iowa USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7129-9500","authenticated-orcid":false,"given":"LingSze","family":"Lee","sequence":"additional","affiliation":[{"name":"Department of Ecology, Evolution, and Organismal Biology Iowa State University  Ames Iowa USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1148-631X","authenticated-orcid":false,"given":"Nicole","family":"Valenzuela","sequence":"additional","affiliation":[{"name":"Department of Ecology, Evolution, and Organismal Biology Iowa State University  Ames Iowa USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"286","published-online":{"date-parts":[[2022,7,25]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10577-009-9095-1"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tig.2011.05.005"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10577-013-9343-2"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1590\/S1415-47572013000100010"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/0014-4827(63)90173-3"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF02424488"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.3390\/genes11040416"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2020.0101"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1023\/A:1009238402051"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/hdy.2011.105"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/nyas.13307"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0960-9822(02)00448-7"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2010.04.042"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2148-10-271"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.1093\/genetics\/138.3.787"},{"key":"e_1_2_10_17_1","first-page":"29","article-title":"Ag\u2010NOR staining in the Bennett wallaby, Macropus rufogriseus \u2013 Evidence for dosage compensation","volume":"56","author":"Dhaliwal M. 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