Sexual reproduction is both extremely costly and extremely common relative to asexuality, indicating that it must confer profound benefits. This in turn points to major disadvantages of asexual reproduction, which is usually given as an explanation for why almost all asexual lineages are apparently quite short-lived. However, a growing body of evidence suggests that some asexual lineages are actually quite old. Insight into why sex is so common may come from understanding why asexual lineages persist in some places or taxa but not others. Here, we review the distribution of asexual lineage ages estimated from a diverse array of taxa, and we discuss our results in light of the main mutational and environmental hypotheses for sex. Along with strengthening the case for wide variation in asexual lineage age and the existence of many old asexual taxa, we also found that the distribution of asexual lineage age estimates follows a surprisingly regular distribution, to the extent that asexual taxa viewed as "scandalously" ancient merely fall on the high end of this distribution. We interpret this result to mean that similar mechanisms may determine asexual lineage age across eukaryotic taxa. We also derive some qualitative predictions for asexual lineage age under different theories for sex and discuss empirical evidence for these predictions. Ultimately, we were limited in the extent to which we could use these data to make inferences about the maintenance of sex by the absence of both clear theoretical expectations and estimates of key parameters.

Understanding the maintenance of sexual reproduction constitutes a difficult problem for evolutionary biologists because of the immediate costs that sex seems to incur. Typically, general benefits to sex and recombination are investigated that might outweigh these costs. However, several factors can strongly influence the complex balance between costs and benefits of sex; these include constraints on the evolution of asexuality, ecological differentiation, and certain lif-history traits. We review these factors and their empirical support for the first time in a unified framework and find that they can reduce the costs of sex, circumvent them, or make them inapplicable. These factors can even tip the scales to a net benefit for sex. The reviewed factors affect species and species groups differently, and we conclude consequently that understanding the maintenance of sex could turn out to be more species-specific than commonly assumed. Interestingly, our study suggests that, in some species, no general benefits to sex and recombination might be needed to understand the maintenance of sex, as in our case study of dandelions.

Why and how sexual reproduction is maintained in natural populations, the so-called "queen of problems," is a key unanswered question in evolutionary biology. Recent efforts to solve the problem of sex have often emphasized results generated from laboratory settings. Here, we use a survey of representative "sex in the wild" literature to review and synthesize the outcomes of empirical studies focused on natural populations. Especially notable results included relatively strong support for mechanisms involving niche differentiation and a near absence of attention to adaptive evolution. Support for a major role of parasites is largely confined to a single study system, and only three systems contribute most of the support for mutation accumulation hypotheses. This evidence for taxon specificity suggests that outcomes of particular studies should not be more broadly extrapolated without extreme caution. We conclude by suggesting steps forward, highlighting tests of niche differentiation mechanisms in both laboratory and nature, and empirical evaluation of adaptive evolution-focused hypotheses in the wild. We also emphasize the value of leveraging the growing body of genomic resources for nonmodel taxa to address whether the clearance of harmful mutations and spread of beneficial variants in natural populations proceeds as expected under various hypotheses for sex.

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