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Plant mating systems play a key role in structuring genetic variation both within and between species. In hybrid zones, the outcomes and dynamics of hybridization are usually interpreted as the balance between gene flow and selection against hybrids. Yet, mating systems can introduce selective forces that alter these expectations; with diverse outcomes for the level and direction of gene flow depending on variation in outcrossing and whether the mating systems of the species pair are the same or divergent. We present a survey of hybridization in 133 species pairs from 41 plant families and examine how patterns of hybridization vary with mating system. We examine if hybrid zone mode, level of gene flow, asymmetries in gene flow and the frequency of reproductive isolating barriers vary in relation to mating system/s of the species pair. We combine these results with a simulation model and examples from the literature to address two general themes: (1) the two-way interaction between introgression and the evolution of reproductive systems, and (2) how mating system can facilitate or restrict interspecific gene flow. We conclude that examining mating system with hybridization provides unique opportunities to understand divergence and the processes underlying reproductive isolation.

In most mammals, both sexes display different survial patterns, often involving faster senescence in males. Being under intense sexual competition to secure mating opportunities, males of polygynous species allocate resources to costly behaviors and conspicuous sexual traits, which might explain these observed differences in longevity and senescence patterns. However, comparative studies performed to date have led to conflicting results. We aimed to resolve this problem by first reviewing case studies of the relationship between the strength of sexual selection and age-specific survival metrics. Then, we performed a comprehensive comparative analysis to test whether such relationships exist among species of captive ruminants. We found that the strength of sexual selection negatively influenced the onset of actuarial senescence in males, with males senescing earlier in polygynous than in monogamous species, which led to reduced male longevity in polygynous species. Moreover, males of territorial species senesced earlier but slower, and have a shorter longevity than males of species displaying other mating tactics. We detected little influence of the strength of sexual selection on the rate of actuarial senescence. Our findings demonstrate that the onset of actuarial senescence, rather than its rate, is a side effect of physiological mechanisms linked to sexual selection, and potentially accounts for observed differences in longevity.

A compound hypothesis positing that self-fertilization is an evolutionary dead end conflates two distinct claims: the transition from outcrossing to selfing is unidirectional; and the diversification rate, or the balance of the speciation and extinction rate, is negative for selfing species. Both claims have enjoyed widespread informal support for decades, but have recently come under suspicion. Sources of data that apparently contradict strongly asymmetric mating system transitions often rely on statistical phylogenetic tests plagued by profound flaws. Although recently developed models mend preceding approaches, they have been employed sparingly, and many problems remain. Theoretical investigations, genetic data and applications of new phylogenetic methods provide indirect support for an association of selfing with negative diversification rates. We lack direct tests of reversals from selfing to outcrossing, and require data concerning the genetic basis and complexity of independently evolved outcrossing adaptations. The identification of the mechanisms that limit the longevity of selfing lineages has been difficult. Limitations may include brief and variable durations of selfing lineages, as well as ongoing difficulties in relating additive genetic and nucleotide variation. Furthermore, a common line of evidence for the stability of mixed mating – based simply on its frequent occurrence – is misleading. We make specific suggestions for research programs that aim to provide a richer understanding of mating system evolution and seriously challenge Stebbins' venerable hypothesis.

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

Most spermatophytes need conspecific pollen in order to produce seed. This need for specialization seems to conflict with the generalized nature of most plant–pollinator interactions. Specialization and generalization are dynamic – not fixed – and exist simultaneously in multiple states at different levels of biological hierarchy. Over the short term, specialization ensures conspecific pollen transfer, whereas over the long term, generalization improves system-level robustness. The balance between specialization and generalization at different scales is critical for different kinds of ecological functioning and is an important factor in plant speciation and the evolution of plant mating systems. Community context, including diversity and interaction network structure at different levels of aggregation, is a key driver of specialization dynamics.

The evolution and maintenance of secondary sexual characteristics and behavior are heavily influenced by the variance in mating success among individuals in a population. The operational sex ratio (OSR) is often used as a predictor of the intensity of competition for mates, as it describes the relative number of males and females who are ready to mate. We investigate changes in aggression, courtship, mate guarding, and sperm release as a function of changes in the OSR using meta-analytic techniques. As the OSR becomes increasingly biased, aggression increases as competitors attempt to defend mates, but this aggression begins to decrease at an OSR of 1.99, presumably due to the increased costs of competition as rivals become more numerous. Sperm release follows a similar but not significant trend. By contrast, courtship rate decreases as the OSR becomes increasingly biased, whereas mate guarding and copulation duration increase. Overall, predictable behavioral changes occur in response to OSR, although the nature of the change is dependent on the type of mating behavior. These results suggest considerable flexibility of mating system structure within species, which can be predicted by OSR and likely results in variation in the strength of sexual selection.

• Self-fertilisation has consequences for variation across the genome as it reduces effective population size, effect recombination rates and pollen flow, with implications for local adaptation. • We conducted simulations of divergent stabilising selection on a quantitative trait with drift, pollen flow, mutation, recombination and different outcrossing rates. We quantified trait divergence and the genetic architecture of adaptation. We conducted an FST outlier analysis to identify candidate loci and quantified the impact of mating system on detectability. • Selfing promoted trait divergence mainly through reductions in pollen flow. Moreover, trait architecture became more diffuse with selfing. Average effect size of trait loci was lower, while the number of loci, and their clustering distance increased. The genetic architecture of selfers was also more diffuse than outcrossers for equivalent migration rates. However, when deleterious alleles were included, architectures became more concentrated in selfers, likely to be because of reductions in population size caused by mutational meltdown and impacts of background selection on Nₑ. • Our simulations demonstrate that mating system has important impacts on adaptive divergence of traits and the genetic landscape underlying that divergence. Selfing has a significant effect on detectability of regions of the genome important for adaptation because of neutral divergence and diffuse trait architecture.

Male–female coevolution has taken different paths among closely related species, but our understanding of the factors that govern its direction is limited. While it is clear that ecological factors, life history, and the economics of reproduction are connected, the divergent links are often obscure. We propose that a complete understanding requires the conceptual integration of metabolic phenotypes. Metabolic rate, a nexus of life history evolution, is constrained by ecological factors and may exert important direct and indirect effects on the evolution of sexual dimorphism. We performed standardized experiments in 12 seed beetle species to gain a rich set of sex-specific measures of metabolic phenotypes, life history traits, and the economics of mating and analyzed our multivariate data using phylogenetic comparative methods. Resting metabolic rate (RMR) showed extensive evolution and evolved more rapidly in males than in females. The evolution of RMR was tightly coupled with a suite of life history traits, describing a pace-of-life syndrome (POLS), with indirect effects on the economics of mating. As predicted, high resource competition was associated with a low RMR and a slow POLS. The cost of mating showed sexually antagonistic coevolution, a hallmark of sexual conflict. The sex-specific costs and benefits of mating were predictably related to ecology, primarily through the evolution of male ejaculate size. Overall, our results support the tenet that resource competition affects metabolic processes that, in turn, have predictable effects on both life history evolution and reproduction, such that ecology shows both direct and indirect effects on male–female coevolution.

Dispersal affects species' ability to move or adapt in response to environmental change. Successful long‐distance dispersal also requires reproduction in areas with few mates, thus mating systems, especially the capacity for self‐fertilization, may influence the speed and success of range shifts. Here, we review: the theoretical predictions regarding dispersal and mating‐system evolution at equilibrium, expanding and contracting range limits; the empirical support for these predictions; and how these geographic patterns may influence future range evolution. Equilibrium range limits can arise from environmental gradients in habitat quality, temporal variation or habitat heterogeneity. Dispersal has been predicted to increase or decrease towards range edges, depending on which life‐history traits respond to the ecological gradient(s). In general, spatial habitat isolation selects against dispersal, whereas temporal stochasticity favours dispersal. At expanding range fronts, dispersal should increase due to spatial sorting for dispersive individuals and the benefits of colonizing vacant habitat. Dispersal evolution is likely more constrained during native range shifts than invasions. Models of expansion across environmental gradients and during climate‐tracking range shifts are lacking. Little theory considers evolution at contracting range margins. We suggest that increased dispersal should be selected if there is local adaptation to climate, as dispersers from warmer areas will out‐compete nondispersers no longer adapted to new climatic conditions. Dispersal increases should be more pronounced in regions where local adaptation is stronger. Self fertilization may be favoured at equilibrium, expanding or contracting range margins by providing reproductive assurance. However, this benefit depends on how inbreeding depression is influenced by genetic load, the severity of the abiotic environment, and the competitive milieu in edge populations. Models for the joint evolution of mating and dispersal in plants suggest that although selfing may evolve at range limits, it will not necessarily be associated with high dispersal. Empirical evidence to test these predictions is scarce. Geographic surveys of dispersal traits, mating‐system traits and relevant selective factors are needed, especially studies of: (i) stable range limits that identify underlying environmental gradients; (ii) moving range limits that compare traits across space and time; and (iii) contracting limits that assess variation in local adaptation towards the range edge.