Speijer, D., Lukeš, J. & Eliáš, M. Sex is a ubiquitous, ancient, and inherent attribute of eukaryotic life. Proc. Natl Acad. Sci. 112, 8827–8834 (2015).
Google Scholar
Bachtrog, D. et al. Sex determination: why so many ways of doing it? PLoS Biol. 12, e1001899 (2014).
Google Scholar
Ah-King, M. & Nylin, S. Sex in an evolutionary perspective: just another reaction norm. Evolut. Biol. 37, 234–246 (2010).
Google Scholar
Leonard, J. L. The evolution of sexual systems in animals. In: Leonard, J.L. (ed.). Transitions between sexual systems: understanding the mechanisms of, and pathways between, dioecy, hermaphroditism and other sexual systems, 1–58 Springer (2019).
Weeks, S. C., Benvenuto, C. & Reed, S. K. When males and hermaphrodites coexist: a review of androdioecy in animals. Integr. Comp. Biol. 46, 449–464 (2006).
Google Scholar
Goldberg, E. E. et al. Macroevolutionary synthesis of flowering plant sexual systems. Evolution 71, 898–912 (2017).
Google Scholar
Waples, R. S., Mariani, S. & Benvenuto, C. Consequences of sex change for effective population size. Proc. R. Soc. B: Biol. Sci. 285, 20181702 (2018).
Google Scholar
Benvenuto, C. & Weeks, S. C. Hermaphroditism and gonochorism. The Natural History of the Crustacea: Reproductive Biology VI, 197–241 (2020).
Mariani, S., Sala-Bozano, M., Chopelet, J. & Benvenuto, C. Spatial and temporal patterns of size-at-sex-change in two exploited coastal fish. Environ. Biol. Fishes 96, 535–541 (2013).
Google Scholar
Käfer, J., Marais, G. A. & Pannell, J. R. On the rarity of dioecy in flowering plants. Mol. Ecol. 26, 1225–1241 (2017).
Google Scholar
Atz, J. Intersexuality in Fishes. In C.N. Amstrong and A.J. Marshall (eds). Intersexuality in vertebrates including man, 145–232 Academic Press, London (1964).
Jarne, P. & Auld, J. R. Animals mix it up too: the distribution of self-fertilization among hermaphroditic animals. Evolution 60, 1816–1824 (2006).
Google Scholar
Leonard, J. L. Williams’ paradox and the role of phenotypic plasticity in sexual systems. Integr. Comp. Biol. 53, 671–688 (2013).
Google Scholar
Weeks, S. C. The role of androdioecy and gynodioecy in mediating evolutionary transitions between dioecy and hermaphroditism in the animalia. Evolution 66, 3670–3686 (2012).
Google Scholar
Renner, S. S. The relative and absolute frequencies of angiosperm sexual systems: dioecy, monoecy, gynodioecy, and an updated online database. Am. J. Bot. 101, 1588–1596 (2014).
Google Scholar
Bawa, K. S. Evolution of dioecy in flowering plants. Annu. Rev. Ecol. Syst. 11, 15–39 (1980).
Google Scholar
Charlesworth, B. & Charlesworth, D. A model for the evolution of dioecy and gynodioecy. Am. Nat. 112, 975–997 (1978).
Google Scholar
Charlesworth, D. Androdioecy and the evolution of dioecy. Biol. J. Linn. Soc. 22, 333–348 (1984).
Google Scholar
Pannell, J. R. The evolution and maintenance of androdioecy. In: Annual Review of Ecology and Systematics 397–425 (2002).
Bull, J. & Charnov, E. On irreversible evolution. Evolution 39, 1149–1155 (1985).
Google Scholar
Barrett, S. C. The evolution of plant reproductive systems: how often are transitions irreversible? Proc. R. Soc. B: Biol. Sci. 280, 20130913 (2013).
Google Scholar
Oyarzún, P. A., Nuñez, J. J., Toro, J. E. & Gardner, J. P. Trioecy in the marine mussel Semimytilus algosus (Mollusca, Bivalvia): stable sex ratios across 22 degrees of a latitudinal gradient. Front. Mar. Sci. 7, 348 (2020).
Google Scholar
Dani, K. & Kodandaramaiah, U. Plant and animal reproductive strategies: lessons from offspring size and number tradeoffs. Front. Ecol. Evol. 5, 38 (2017).
Google Scholar
Avise, J. & Mank, J. Evolutionary perspectives on hermaphroditism in fishes. Sex. Dev. 3, 152–163 (2009).
Google Scholar
Dornburg, A. & Near, T. J. The Emerging phylogenetic perspective on the evolution of Actinopterygian fishes. Annu. Rev. Ecol. Evol. Syst. 52, 427–452 (2021).
Google Scholar
Costa, W. J., Lima, S. M. & Bartolette, R. Androdioecy in Kryptolebias killifish and the evolution of self-fertilizing hermaphroditism. Biol. J. Linn. Soc. 99, 344–349 (2010).
Google Scholar
Costa, W. Colouration, taxonomy and geographical distribution of mangrove killifishes, the Kryptolebias marmoratus species group, in southern Atlantic coastal plains of Brazil (Cyprinodontiformes: Rivulidae). Ichthyol. Explor. Freshw. 27, 183–192 (2016).
Powell, M. L., Kavanaugh, S. I. & Sower, S. A. Seasonal concentrations of reproductive steroids in the gonads of the Atlantic hagfish, Myxine glutinosa. J. Exp. Zool. Part A Comp. Exp. Biol. 301, 352–360 (2004).
Google Scholar
Pennell, M. W., Mank, J. E. & Peichel, C. L. Transitions in sex determination and sex chromosomes across vertebrate species. Mol. Ecol. 27, 3950–3963 (2018).
Google Scholar
Ghiselin, M. T. The evolution of hermaphroditism among animals. Q. Rev. Biol. 44, 189–208 (1969).
Google Scholar
Eppley, S. M. & Jesson, L. K. Moving to mate: the evolution of separate and combined sexes in multicellular organisms. J. Evol. Biol. 21, 727–736 (2008).
Google Scholar
Warner, R. R. The adaptive significance of sequential hermaphroditism in animals. Am. Nat. 109, 61–82 (1975).
Google Scholar
Warner, R. R., Robertson, D. R. & Leigh, E. G. Sex change and sexual selection. Science 190, 633–638 (1975).
Google Scholar
Charnov, E. L. The Theory of Sex Allocation. Princeton University Press, USA (1982).
Policansky, D. Sex change in plants and animals. Annu. Rev. Ecol. Syst. 13, 471–495 (1982).
Google Scholar
Benvenuto, C., Coscia, I., Chopelet, J., Sala-Bozano, M. & Mariani, S. Ecological and evolutionary consequences of alternative sex-change pathways in fish. Sci. Rep. 7, 9084 (2017).
Google Scholar
Charnov, E. L. Natural selection and sex change in pandalid shrimp: test of a life-history theory. Am. Nat. 113, 715–734 (1979).
Google Scholar
Broquet, T. et al. The size advantage model of sex allocation in the protandrous sex-changer Crepidula fornicata: role of the mating system, sperm storage, and male mobility. Am. Nat. 186, 404–420 (2015).
Google Scholar
Erisman, B. E., Craig, M. T. & Hastings, P. A. A phylogenetic test of the size-advantage model: evolutionary changes in mating behavior influence the loss of sex change in a fish lineage. Am. Nat. 174, E83–E99 (2009).
Google Scholar
Buxton, C. D. & Garratt, P. A. Alternative reproductive styles in seabreams (Pisces: Sparidae). Environ. Biol. Fishes 28, 113–124 (1990).
Google Scholar
Shapiro, D. Y. Social behavior, group structure, and the control of sex reversal in hermaphroditic fish. Adv. Study Behav. 10, 43–102 (1979).
Google Scholar
Stearns, S. C. Life history evolution: successes, limitations, and prospects. Naturwissenschaften 87, 476–486 (2000).
Google Scholar
Waples, R. S., Luikart, G., Faulkner, J. R. & Tallmon, D. A. Simple life-history traits explain key effective population size ratios across diverse taxa. Proc. R. Soc. Lond. B: Biol. Sci. 280, 20131339 (2013).
Martinez, A. S., Willoughby, J. R. & Christie, M. R. Genetic diversity in fishes is influenced by habitat type and life-history variation. Ecol. Evolution 8, 12022–12031 (2018).
Google Scholar
Harvey, P. H. & Pagel, M. D. The comparative method in evolutionary biology. (Oxford University Press, USA, 1991).
Barneche, D. R., Robertson, D. R., White, C. R. & Marshall, D. J. Fish reproductive-energy output increases disproportionately with body size. Science 360, 642–645 (2018).
Google Scholar
Brandl, S. J. & Bellwood, D. R. Pair-formation in coral reef fishes: an ecological perspective. Oceanogr. Mar. Biol.: Annu. Rev. 52, 1–80 (2014).
Fitzpatrick, J. L. Sperm competition and fertilization mode in fishes. Philos. Trans. R. Soc. B: Biol. Sci. 375, 20200074 (2020).
Google Scholar
Parker, G. A. Conceptual developments in sperm competition: a very brief synopsis. Philos. Trans. R. Soc. B: Biol. Sci. 375, 20200061 (2020).
Google Scholar
Warner, R. R. Sex change in fishes: hypotheses, evidence, and objections. Environ. Biol. Fishes 22, 81–90 (1988).
Google Scholar
Molloy, P. P., Goodwin, N. B., Côté, I. M., Reynolds, J. D. & Gage, M. J. Sperm competition and sex change: a comparative analysis across fishes. Evolution 61, 640–652 (2007).
Google Scholar
Erisman, B. E., Petersen, C. W., Hastings, P. A. & Warner, R. R. Phylogenetic perspectives on the evolution of functional hermaphroditism in teleost fishes. Integr. Comp. Biol. 53, 736–754 (2013).
Google Scholar
Sadovy, Y., Colin, P. & Domeier, M. Aggregation and spawning in the tiger grouper, Mycteroperca tigris (Pisces: Serranidae). Copeia 1994, 511–516 (1994).
Google Scholar
Muñoz, R. C. & Warner, R. R. A new version of the size-advantage hypothesis for sex change: incorporating sperm competition and size-fecundity skew. Am. Nat. 161, 749–761 (2003).
Google Scholar
Horne, C. R., Hirst, A. G. & Atkinson, D. Selection for increased male size predicts variation in sexual size dimorphism among fish species. Proc. R. Soc. B: Biol. Sci. 287, 20192640 (2020).
Google Scholar
Parker, G. The evolution of expenditure on testes. J. Zool. 298, 3–19 (2016).
Google Scholar
Stockley, P., Gage, M., Parker, G. & Møller, A. Sperm competition in fishes: the evolution of testis size and ejaculate characteristics. Am. Nat. 149, 933–954 (1997).
Google Scholar
Pla, S., Benvenuto, C., Capellini, I. & Piferrer, F. A phylogenetic comparative analysis on the evolution of sequential hermaphroditism in seabreams (Teleostei: Sparidae). Sci. Rep. 10, 3606 (2020).
Google Scholar
Vrijenhoek, R. C. Unisexual fish: model systems for studying ecology and evolution. Annu. Rev. Ecol. Syst. 25, 71–96 (1994).
Google Scholar
Sadovy de Mitcheson, Y. & Liu, M. Functional hermaphroditism in teleosts. Fish. Fish. 9, 1–43 (2008).
Google Scholar
Rabosky, D. L. et al. An inverse latitudinal gradient in speciation rate for marine fishes. Nature 559, 392 (2018).
Google Scholar
Froese, R., Pauly, D. & Editors. FishBase. World Wide Web electronic publication. www.fishbase.org (2018).
Moore, W. S. Evolutionary ecology of unisexual fishes. In: Evolutionary genetics of fishes, 329–398 (Springer, 1984).
Collin, R. & Miglietta, M. P. Reversing opinions on Dollo’s Law. Trends Ecol. Evol. 23, 602–609 (2008).
Google Scholar
Domes, K., Norton, R. A., Maraun, M. & Scheu, S. Re-evolution of sexuality breaks Dollo’s law. Proc. Natl Acad. Sci. 104, 7139–7144 (2007).
Google Scholar
Dollo, L. Les lois de l’évolution. Bull. Soc. Belge Géol. Paléont. Hydrol. 7, 164–166 (1893).
King, B. & Lee, M. S. Ancestral state reconstruction, rate heterogeneity, and the evolution of reptile viviparity. Syst. Biol. 64, 532–544 (2015).
Google Scholar
Uller, T. & Helanterä, H. From the origin of sex-determining factors to the evolution of sex-determining systems. Q. Rev. Biol. 86, 163–180 (2011).
Google Scholar
Devlin, R. H. & Nagahama, Y. Sex determination and sex differentiation in fish: an overview of genetic, physiological, and environmental influences. Aquaculture 208, 191–364 (2002).
Google Scholar
Volff, J.-N., Nanda, I., Schmid, M. & Schartl, M. Governing sex determination in fish: regulatory putsches and ephemeral dictators. Sex. Dev. 1, 85–99 (2007).
Google Scholar
Nagahama, Y., Chakraborty, T., Paul-Prasanth, B., Ohta, K. & Nakamura, M. Sex determination, gonadal sex differentiation and plasticity in vertebrate species. Physiol. Rev. 101, 1237–1308 (2020).
Google Scholar
Penman, D. J. & Piferrer, F. Fish gonadogenesis. Part I: genetic and environmental mechanisms of sex determination. Rev. Fish. Sci. 16(S1), 16–34 (2008).
Google Scholar
Mank, J. E., Promislow, D. E. L. & Avise, J. C. Evolution of alternative sex-determining mechanisms in teleost fishes. Biol. J. Linn. Soc. 87, 83–93 (2006).
Google Scholar
Galetti, P. M., Aguilar, C. T. & Molina, W. F. An overview of marine fish cytogenetics. Hydrobiologia 420, 55–62 (2000).
Google Scholar
Yoshida, K. et al. Sex chromosome turnover contributes to genomic divergence between incipient stickleback species. PLoS Genet. 10, e1004223 (2014).
Google Scholar
Ross, J. A., Urton, J. R., Boland, J., Shapiro, M. D. & Peichel, C. L. Turnover of sex chromosomes in the stickleback fishes (Gasterosteidae). PLoS Genet. 5, e1000391 (2009).
Google Scholar
Vicoso, B. Molecular and evolutionary dynamics of animal sex-chromosome turnover. Nature Ecology & Evolution 1–10 (2019).
Gamble, T. et al. Restriction site-associated DNA sequencing (RAD-seq) reveals an extraordinary number of transitions among gecko sex-determining systems. Mol. Biol. Evol. 32, 1296–1309 (2015).
Google Scholar
Pokorná, M. & Kratochvíl, L. Phylogeny of sex-determining mechanisms in squamate reptiles: are sex chromosomes an evolutionary trap? Zool. J. Linn. Soc. 156, 168–183 (2009).
Google Scholar
Furman, B. L. et al. Sex chromosome evolution: sso many exceptions to the rules. Genome Biol. Evol. 12, 750–763 (2020).
Google Scholar
Carvalho, N. D. M. et al. Cytogenetics of Synbranchiformes: a comparative analysis of two Synbranchus Bloch, 1795 species from the Amazon. Genetica 140, 149–158 (2012).
Google Scholar
Piferrer, F. Epigenetic mechanisms in sex determination and in the evolutionary transitions between sexual systems. Philos. Trans. R. Soc. B: Biol. Sci. 376, 20200110 (2021).
Google Scholar
Grant, S. et al. Genetics of sex determination in flowering plants. Dev. Genet. 15, 214–230 (1994).
Google Scholar
Harrington Jr, R. W. How ecological and genetic factors interact to determine when self-fertilizing hermaphrodites of Rivulus marmoratus change into functional secondary males, with a reappraisal of the modes of intersexuality among fishes. Copeia 389–432 (1971).
Adolfi, M. C., Nakajima, R. T., Nóbrega, R. H. & Schartl, M. Intersex, Hermaphroditism, and gonadal plasticity in vertebrates: Evolution of the Müllerian duct and Amh/Amhr2 signalling. Annual Review of Animal Biosciences (2018).
Adkins-Regan, E. Early organizational effects of hormones: an evolutionary perspective. In Adler, N.T. (ed.) Neuroendocrinology of reproduction: physiology and behavior, 159–228 (Springer, USA, 1981).
Navara, K. J. The truth about Nemo’s dad: sex-changing behaviors in fishes. In Choosing Sexes 183–212 (Springer, Cham, 2018).
Orban, L., Sreenivasan, R. & Olsson, P. E. Long and winding roads: testis differentiation in zebrafish. Mol. Cell. Endocrinol. 312, 35–41 (2009).
Google Scholar
Zohar, Y., Abraham, M. & Gordin, H. The gonadal cycle of the captivity-reared hermaphroditic teleost Sparus aurata (L.) during the first two years of life. Annales de. Biologie Anim. Biochim. Biophys. 18, 877–882 (1978).
Google Scholar
Chang, C.-F. & Yueh, W.-S. Annual cycle of gonadal histology and steroid profiles in the juvenile males and adult females of the protandrous black porgy, Acanthopagrus schlegelii. Aquaculture 91, 179–196 (1990).
Google Scholar
Miura, S., Nakamura, S., Kobayashi, Y., Piferrer, F. & Nakamura, M. Differentiation of ambisexual gonads and immunohistochemical localization of P450 cholesterol side-chain cleavage enzyme during gonadal sex differentiation in the protandrous anemonefish, Amphiprion clarkii. Comp. Biochem. Physiol. Part B: Biochem. Mol. Biol. 149, 29–37 (2008).
Google Scholar
Yamaguchi, S. & Iwasa, Y. Advantage for the sex changer who retains the gonad of the nonfunctional sex. Behav. Ecol. Sociobiol. 71, 39 (2017).
Google Scholar
Munday, P. L., Kuwamura, T. & Kroon, F. J. Bi-directional sex change in marine fishes. In: Cole, K.S. (ed.) Reproduction and sexuality in marine fishes: Patterns and processes. 241–271 (University of California Press, Berkeley, USA, 2010).
Uller, T., Feiner, N., Radersma, R., Jackson, I. S. & Rago, A. Developmental plasticity and evolutionary explanations. Evol. Dev. 22, 47–55 (2020).
Google Scholar
Pla, S., Maynou, F. & Piferrer, F. Hermaphroditism in fish: incidence, distribution and associations with abiotic environmental factors. Rev. Fish. Biol. Fish. 31, 935–955 (2021).
Google Scholar
Boettiger, C., Lang, D. T. & Wainwright, P. C. rfishbase: exploring, manipulating and visualizing FishBase data from R. J. Fish. Biol. 81, 2030–2039 (2012).
Google Scholar
Pagel, M., Meade, A. & Barker, D. Bayesian estimation of ancestral character states on phylogenies. Syst. Biol. 53, 673–684 (2004).
Google Scholar
Pagel, M. & Meade, A. Bayesian analysis of correlated evolution of discrete characters by reversible-jump Markov chain Monte Carlo. Am. Nat. 167, 808–825 (2006).
Google Scholar
Pagel, M. Detecting correlated evolution on phylogenies: a general method for the comparative analysis of discrete. Proc. R. Soc. B: Biol. Sci. 255, 37–45 (1994).
Google Scholar
Currie, T. E. & Meade, A. In Modern phylogenetic comparative methods and their application in evolutionary biology, 263–286 (Springer, 2014).
Furness, A. I. & Capellini, I. The evolution of parental care diversity in amphibians. Nat. Commun. 10, 1–12 (2019).
Google Scholar
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 (2018).
Google Scholar
Paradis, E. & Schliep, K. ape 5.0: an environment for modern phylogenetics and evolutionary analyses in R. Bioinformatics 35, 526–528 (2019).
Google Scholar
Freckleton, R. P., Harvey, P. H. & Pagel, M. Phylogenetic analysis and comparative data: a test and review of evidence. Am. Nat. 160, 712–726 (2002).
Google Scholar
Pagel, M. Inferring evolutionary processes from phylogenies. Zool. Scr. 26, 331–348 (1997).
Google Scholar
Pagel, M. Inferring the historical patterns of biological evolution. Nature 401, 877–884 (1999).
Google Scholar
Orme, D. The caper package: comparative analysis of phylogenetics and evolution in R. https://cran.r-project.org/web/packages/caper/vignettes/caper.pdf (2018).
Schiettekatte, N., Brandl, S. & Casey, J. Fishualize: Color palettes based on fish species. R package v0.2.2 (2021).
Source: Ecology - nature.com
