We review knowledge about the roles of sex chromosomes in vertebrate hybridization and speciation, exploring a gradient of divergences with increasing reproductive isolation (speciation continuum). Under early divergence, well-differentiated sex chromosomes in meiotic hybrids may cause Haldane-effects and introgress less easily than autosomes. Undifferentiated sex chromosomes are more susceptible to introgression and form multiple (or new) sex chromosome systems with hardly predictable dominance hierarchies. Under increased divergence, most vertebrates reach complete intrinsic reproductive isolation. Slightly earlier, some hybrids (linked in 'the extended speciation continuum') exhibit aberrant gametogenesis, leading towards female clonality. This facilitates the evolution of various allodiploid and allopolyploid clonal ('asexual') hybrid vertebrates, where 'asexuality' might be a form of intrinsic reproductive isolation. A comprehensive list of 'asexual' hybrid vertebrates shows that they all evolved from parents with divergences that were greater than at the intraspecific level (K2P-distances of greater than 5-22% based on mtDNA). These 'asexual' taxa inherited genetic sex determination by mostly undifferentiated sex chromosomes. Among the few known sex-determining systems in hybrid 'asexuals', female heterogamety (ZW) occurred about twice as often as male heterogamety (XY). We hypothesize that pre-/meiotic aberrations in all-female ZW-hybrids present Haldane-effects promoting their evolution. Understanding the preconditions to produce various clonal or meiotic allopolyploids appears crucial for insights into the evolution of sex, 'asexuality' and polyploidy. This article is part of the theme issue 'Challenging the paradigm in sex chromosome evolution: empirical and theoretical insights with a focus on vertebrates (Part II)'.

Amphibian Research Center Hiroshima University Higashi Hiroshima 739 8526 Japan

Department of Biology and Ecology Faculty of Science University of Ostrava 701 03 Ostrava Czech Republic

Department of Zoology Faculty of Science Charles University Viničná 7 Prague 2 128 00 Czech Republic

Institute of Animal Physiology and Genetics Laboratory of Fish Genetics The Czech Academy of Sciences 277 21 Libechov Czech Republic

Leibniz Institute of Freshwater Ecology and Inland Fisheries IGB Müggelseedamm 301 12587 Berlin Germany

Research Department for Limnology University of Innsbruck Mondseestrasse 9 A 5310 Mondsee Austria

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Gradual chromosomal lagging drive programmed genome elimination in hemiclonal fishes from the genus Hypseleotris

Unifying framework explaining how parental regulatory divergence can drive gene expression in hybrids and allopolyploids

Inheritance patterns of male asexuality in hybrid males of a water frog Pelophylax esculentus

Premeiotic endoreplication is the mechanism of obligate parthenogenesis in rock lizards of the genus Darevskia

Meiotic deviations and endoreplication lead to diploid oocytes in female hybrids between bighead catfish (Clarias macrocephalus) and North African catfish (Clarias gariepinus)

A cyclical switch of gametogenic pathways in hybrids depends on the ploidy level

Genetic and karyotype divergence between parents affect clonality and sterility in hybrids

Mechanisms of Intrinsic Postzygotic Isolation: From Traditional Genic and Chromosomal Views to Genomic and Epigenetic Perspectives

Sperm-dependent asexual species and their role in ecology and evolution

Achiasmatic meiosis in the unisexual Amazon molly, Poecilia formosa

The high diversity of gametogenic pathways in amphispermic water frog hybrids from Eastern Ukraine

Equilibrated evolution of the mixed auto-/allopolyploid haplotype-resolved genome of the invasive hexaploid Prussian carp

Challenges and Costs of Asexuality: Variation in Premeiotic Genome Duplication in Gynogenetic Hybrids from Cobitis taenia Complex

Sex chromosomes in meiotic, hemiclonal, clonal and polyploid hybrid vertebrates: along the 'extended speciation continuum'

Expanding the classical paradigm: what we have learnt from vertebrates about sex chromosome evolution

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