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Premise of Study In a seminal body of theory, Lloyd showed that the fitness consequences of selfing will depend on its timing in anthesis. Selfing that occurs after opportunities for outcrossing or pollen dispersal can provide reproductive assurance when pollinators are limited and is expected to incur little cost, even when inbreeding depression is high. As a result, delayed selfing is often interpreted as a “best‐of‐both‐worlds” mating system that combines the advantages of selfing and outcrossing. Methods We surveyed 65 empirical studies of delayed selfing, recording floral mechanisms and examining information on inbreeding depression, autofertility, and other parameters to test the support for delayed selfing as a best‐of‐both‐worlds strategy. Key Results Phylogenetic distribution of the diverse floral mechanisms suggests that some basic floral structures may predispose plant taxa to evolve delayed selfing. Delayed selfing appears to serve as a best‐of‐both‐worlds strategy in some but not all species. While the capacity for autonomous selfing is often high, it is lower, in some cases, than in related species with earlier modes of selfing. In other delayed‐selfers, low inbreeding depression and reduced investment in corollas and pollen suggest limited benefits from outcrossing. Conclusions Despite a growing literature on the subject, experimental evidence for delayed selfing is limited and major gaps in knowledge remain, particularly with respect to the stability of delayed selfing and the conditions that may favor transitions between delayed and earlier selfing. Finally, we suggest a potential role of delayed selfing in facilitating transitions from self‐incompatibility to selfing.

Hermaphroditic individuals can produce both selfed and outcrossed progeny, termed mixed mating. General theory predicts that mixed-mating populations should evolve quickly toward high rates of setting, driven by rapid purging of genetic load and loss of inbreeding depression (ID), but the substantial number of mixed-mating species observed in nature calls this prediction into question. Lower average ID reported for selfing than for outcrossing populations is consistent with purging and suggests that mixed-mating taxa in evolutionary transition will have intermediate ID. We compared the magnitude of ID from published estimates for highly selfing (r > 0.8), mixed-mating (0.2 ≤ r ≥ 0.8), and highly outcrossing (r < 0.2) plant populations across 58 species. We found that mixed-mating and outcrossing taxa have equally high average lifetime ID (δ = 0.58 and 0.54, respectively) and similar ID at each of four life-cycle stages. These results are not consistent with evolution toward selfing in most mixed-mating taxa. We suggest that prevention of purging by selective interference could explain stable mixed mating in many natural populations. We identify critical gaps in the empirical data on ID and outline key approaches to filling them.

Mixed mating and variation in outcrossing rate among populations of the same species are common. Outcrossing can be affected by pollinator activity and floral traits that facilitate or impede autonomous self‐fertilization. However, the relative contribution of pollen limitation and evolved differences in the ability to self‐fertilize to variation in the mating system is poorly understood and can only be disentangled using an experimental approach. We placed arrays of plants from eight Campanula americana populations that varied in autonomous selfing ability (hereafter “autonomy”) in sun and shade habitats to create high‐ and low‐pollination environments. Floral visitors were observed and pollen limitation, dichogamy, herkogamy and pollen persistence within the flower were measured. Outcrossing rate was estimated for a subset of the arrays. Pollen limitation was over three times higher in the shade than the sun due to differences in pollinator visitation. Populations with high levels of autonomy displayed greater pollen persistence and dichogamy in the more pollen‐limited environments than in the high‐pollination environments. In these high‐autonomy populations, outcrossing rates were tightly associated with pollen limitation, with reduced outcrossing under strong pollen limitation. In contrast, populations with intermediate autonomy levels had less plastic floral traits and their outcrossing rates changed little with pollen limitation. Mating system was shaped by both the pollination environment and the magnitude of plasticity in floral traits. The experimental approach used here revealed that plasticity in floral traits provides the ability to adjust mating system in response to limited potential for outcrossing. The lack of plasticity in some populations could explain mismatches between pollinator activity and mating system in natural populations. Finally, flexibility in the mating system may help explain the prevalence of mixed mating. A plain language summary is available for this article. Plain Language Summary

Quantitative trait locus (QTL) mapping is a first step toward understanding the genetic basis of adaptive evolution and may also reveal reproductive incompatibilities unique to hybrids. In plants, the shift from outcrossing to self‐pollination is common, providing the opportunity for comparisons of QTL architecture among parallel evolutionary transitions. We used QTL mapping in hybrids between the bee‐pollinated monkeyflower Mimulus lewisii and the closely related selfer Mimulus parishii to determine the genetic basis of divergence in floral traits and flowering time associated with mating‐system evolution, and to characterize hybrid anther sterility. We found a moderately polygenic and highly directional basis for floral size evolution, suggesting adaptation from standing variation or in pursuit of a moving optimum, whereas only a few major loci accounted for substantial flowering‐time divergence. Cytonuclear incompatibilities caused hybrid anther sterility, confounding estimation of reproductive organ QTLs. The genetic architecture of floral traits associated with selfing in M. parishii was primarily polygenic, as in other QTL studies of this transition, but in contrast to the previously characterized oligogenic basis of a pollinator shift in close relatives. Hybrid anther sterility appeared parallel at the molecular level to previously characterized incompatibilities, but also raised new questions about cytonuclear co‐evolution in plants.

G. Ledyard Stebbins suggested that self-fertilization (selfing) may be an evolutionary dead end because it may result in the loss of genetic diversity and consequently preclude adaptation to changing environments. While the basic premise of selfing as a dead end is widely accepted, there have been few rigorous evaluations of the hypothesis. We examine the foundations of the dead-end hypothesis by considering theoretical advances in the study of mating-system evolution. We discuss theories predicting the irreversibility of self-fertilization and the extinction of selfing lineages through the loss of adaptive potential and genetic degradation. In the second portion of the review, focusing on the irreversibility of selfing, we summarize the contribution of phylogenetic studies of mating-system evolution to determine if evolutionary history supports this well-established hypothesis. Most studies are in accord with the hypothesis; no single study unequivocally demonstrates the transition from highly selfing to outcrossing lineages. Finally, we discuss the problems encountered when phylogenetic studies rely on reconstruction of ancestral mating systems. To avoid some of these problems, we applied likelihood ratio tests of irreversibility of mating-system evolution to several data sets and found that current data sets are probably too small for this test.

Inbreeding depression is a key factor influencing mating system evolution in plants, but current understanding of its relationship with selfing rate is limited by a sampling bias with few estimates for self-incompatible species. We quantified inbreeding depression (δ) over two growing seasons in two populations of the self-incompatible perennial herb Arabidopsis lyrata ssp. petraea in Scandinavia. Inbreeding depression was strong and of similar magnitude in both populations. Inbreeding depression for overall fitness across two seasons (the product of number of seeds, offspring viability, and offspring biomass) was 81% and 78% in the two populations. Chlorophyll deficiency accounted for 81% of seedling mortality in the selfing treatment, and was not observed among offspring resulting from outcrossing. The strong reduction in both early viability and late quantitative traits suggests that inbreeding depression is due to deleterious alleles of both large and small effect, and that both populations experience strong selection against the loss of self-incompatibility. A review of available estimates suggested that inbreeding depression tends to be stronger in self-incompatible than in self-compatible highly outcrossing species, implying that undersampling of self-incompatible taxa may bias estimates of the relationship between mating system and inbreeding depression.

Males of Caenorhabditis elegans provide a crucial practical tool in the laboratory, but, as the rarer and more finicky sex, have not enjoyed the same depth of research attention as hermaphrodites. Males, however, have attracted the attention of evolutionary biologists who are exploiting the C. elegans system to test longstanding hypotheses about sexual selection, sexual conflict, transitions in reproductive mode, and genome evolution, as well as to make new discoveries about Caenorhabditis organismal biology. Here, we review the evolutionary concepts and data informed by study of males of C. elegans and other Caenorhabditis. We give special attention to the important role of sperm cells as a mediator of inter-male competition and male–female conflict that has led to drastic trait divergence across species, despite exceptional phenotypic conservation in many other morphological features. We discuss the evolutionary forces important in the origins of reproductive mode transitions from males being common (gonochorism: females and males) to rare (androdioecy: hermaphrodites and males) and the factors that modulate male frequency in extant androdioecious populations, including the potential influence of selective interference, host–pathogen coevolution, and mutation accumulation. Further, we summarize the consequences of males being common vs rare for adaptation and for trait divergence, trait degradation, and trait dimorphism between the sexes, as well as for molecular evolution of the genome, at both micro-evolutionary and macro-evolutionary timescales. We conclude that C. elegans male biology remains underexploited and that future studies leveraging its extensive experimental resources are poised to discover novel biology and to inform profound questions about animal function and evolution.

Mating in seed plants arises from interactions between plant traits and the environmental and demographic context in which individuals reside. These interactions commonly cause nonrandom mating, including selfing and promiscuous outcrossing within local neighborhoods. Shared features of seed plants, specifically immobility, hermaphroditism, and modularity, shape the essential character of mating mediated by animals, wind, and water. In addition, diverse floral strategies promote cross- and self-mating, depending on environmental circumstances. Extrinsic ecological factors influence all stages of the mating process-pollination, pollen-tube growth, ovule fertilization-as well as seed development, determining offspring quantity and quality. Traditionally, measures of plant mating systems have focused on a single axis of variation, the maternal outcrossing rate. Instead, we argue for an expanded perspective encompassing mating portfolios, which include all offspring to which individuals contribute genetically as maternal or paternal parents. This approach should expose key ecological determinants of mating-system variation and their evolutionary consequences.

We hypothesize that floral features promoting pollen competition in angiosperms may have evolved, in some cases, in response to selection generated by the negative effects of inbreeding, at least in plants with mixed-mating systems. Screening of haploid genotypes through pollen competition may purge recessive (or additive) deleterious alleles that are expressed in haploid pollen and hence may reduce the fitness cost of self-pollination, geitonogamy, or biparental inbreeding. We tested one prediction of this hypothesis, that offspring produced by more intense competition among self-pollen have higher fitness than offspring produced by less intense competition. Dalechampia scandens (Euphorbiaceae) flowers were pollinated with pollen from other flowers on the same plant (geitonogamous self-fertilization). Those flowers experiencing more intense pollen competition as a result of low pollen dispersion (positional variance) on the stigma produced heavier seeds and seedlings with faster-growing radicles than flowers experiencing less intense pollen competition (high pollen dispersion), as predicted by our hypothesis.

.— Inbreeding depression is a general phenomenon that is well documented in many plants and animals. Furthermore, it is generally considered to be the driving force behind mating‐system evolution. Traditionally, the focus has been on the mean level of inbreeding depression in populations. However, more recently, the variation in inbreeding depression among individuals within populations has been shown to be influential in mating‐system evolution. One set of theories predicts that genetic associations will develop between a mating‐system locus and loci causing inbreeding depression, whereas another suggests either that no such association will occur or that it will be difficult to detect empirically. Here, we focus on variation in inbreeding depression among individuals and present empirical evidence of a genetic association between genes causing inbreeding depression and a floral trait influencing the mating system (i.e., selfing rate). We found a positive association between inbreeding depression and herkogamy (the degree to which the stigma and anthers are separated) in an annual plant, Gilia achilleifolia. These results are consistent with theory predicting that an individual's history of inbreeding will affect its level of inbreeding depression and highlight the potential importance of genetic associations between selfing‐modifier traits and viability in mating‐system evolution.