SPORE SUPPLY AND HABITAT AVAILABILITY AS SOURCES OF RECRUITMENT LIMITATION IN THE GIANT KELP MACROCYSTIS PYRIFERA (PHAEOPHYCEAE)1

The causes of spatial variation in the recruitment of benthic marine algae are frequently misunderstood because of difficulties in distinguishing among the many factors that influence the supply and establishment of microscopic propagules. We used the recently constructed San Clemente Artificial Reef (SCAR) experiment to examine the roles of dispersal distance, size of spore source, and habitat availability as sources of variation in the recruitment of the giant kelp Macrocystis pyrifera (L.) C. Ag., a species whose recruitment has often been considered to be dispersal limited. Sparse colonization on SCAR by adult Macrocystis occurred within 6 months after reef construction via drifters (i.e. individuals from neighboring kelp beds that became dislodged and set adrift). The abundance of drifters on SCAR declined exponentially with distance from the nearest source population (San Mateo), suggesting that San Mateo was the likely source of drifters. Dense recruitment of small Macrocystis sporophytes was observed within 8 months of reef construction. The density of recruits on SCAR showed an initial increase with distance from San Mateo before declining exponentially. Nonetheless, substantial recruitment was observed at the most distant locations on SCAR located 3.5 km from San Mateo. In contrast to drifters, the density of recruits was positively correlated to the bottom cover of artificial reef substrate. Importantly, no correlation was found between the local density or fecundity of drifters and the local density of kelp recruits suggesting that recruitment on SCAR resulted from widespread spore dispersal rather than from the local dispersal of spores from drifters.     

The influence of self‐fertilization and grouping on fitness attributes in the freshwater snail Physa acuta: population and individual inbreeding depression

The genetic structure, selfing rate and inbreeding depression of the hermaphroditic freshwater snail Physa acuta were jointly analysed in a population near Montpellier, France. Allozymic markers revealed moderate gene diversity (0.138), and no heterozygote deficiency. The mean outcrossing rate, estimated by using progeny arrays, was 0.9, with substantial variation among families. This also suggests that the number of fathers among outcrossed offspring of a given mother is low. Inbreeding depression was estimated over more than one generation using 83 first‐laboratory‐generation (G₁) families. The main parameters measured were parental (G₁) fecundity, offspring (G₂) survival and fecundity. Size and growth were also monitored. Parental fecundity was analysed under several conditions (isolation, pair and quadruplet outcrossing). The self‐fertilization depression, including parental fecundity, offspring survival and fecundity, was about 0.9 at the population level. The genetic data obtained in the same population indicate a value of about 0.3 using Ritland’s (1990) technique, suggesting that the depression over the whole life‐cyle might be even higher than 0.9. Grouping affected neither fecundity nor self‐fertilization depression. Substantial variation in depression for survival was detected among individuals, from no survival in some selfed families to better survival than that of outbred families in others. The overall result (outbred population structure, high outcrossing rate and high self‐fertilization depression) is consistent with what is expected in large outcrossing populations in which inbreeding depression is maintained by mutation‐selection balance.     

Consequences of inbreeding for offspring fitness and gender in Silene vulgaris,a gynodioecious plant

Abstract In gynodioecious plants, hermaphrodite and female plants co‐occur in the same population. In these systems gender typically depends on whether a maternally inherited cytoplasmic male sterility factor (CMS) is counteracted by nuclear restorer alleles. These restorer alleles are often genetically dominant. Although plants of the female morph are obligatorily outcrossing, hermaphrodites may self. This selfing increases homozygosity and may thus have two effects: (1) it may decrease fitness (i.e. result in inbreeding depression) and (ii) it may increase homozygosity of the nuclear restorer alleles and therefore increase the production of females. This, in turn, enhances outcrossing in the following generation. In order to test the latter hypothesis, experimental crosses were conducted using individuals derived from four natural populations of Silene vulgaris, a gynodioecious plant. Treatments included self‐fertilization of hermaphrodites, outcrossing of hermaphrodites and females using pollen derived from the same source population as the pollen recipients, and outcrossing hermaphrodites and females using pollen derived from different source populations. Offspring were scored for seed germination, survivorship to flowering and gender. The products of self‐fertilization had reduced survivorship at both life stages when compared with the offspring of outcrossed hermaphrodites or females. In one population the fitness of offspring produced by within‐population outcrossing of females was significantly less than the fitness of offspring produced by crossing females with hermaphrodites from other populations. Self‐fertilization of hermaphrodites produced a smaller proportion of hermaphroditic offspring than did outcrossing hermaphrodites. Outcrossing females within populations produced a smaller proportion of hermaphrodite offspring than did crossing females with hermaphrodites from other populations. These results are consistent with a cytonuclear system of sex determination with dominant nuclear restorers, and are discussed with regard to how the mating system and the genetics of sex determination interact to influence the evolution of inbreeding depression.     

Fine-scale spatial genetic structure of a liverwort (<Emphasis Type="Italic">Barbilophozia attenuata</Emphasis>) within a network of ant trails

Fine-scale spatial genetic structure (SGS) of the liverwort, Barbilophozia attenuata, occupying an area characterized by a network of ant trails, was investigated using microsatellite markers. This is the first study investigating SGS in a liverwort. Significant genetic differentiation was detected among colonies along and outside ant trails, and the SGS pattern varied depending on the spatial scale. At short distances, kinship coefficients were significantly positive up to about eight meters, after which they approached zero and turned negative, while at distances greater than 25 m the values were about zero. Thus, nearby individuals are more closely related than expected, at mid-distances less related, and at great distances genotypes show a random distribution. We suggest that the reproductive mode strongly affects SGS in B. attenuata. Asexual propagation by relatively large gemmae allows more effective establishment than sexual reproduction by small-sized spores, and causes an aggregation of similar genotypes, although the inbreeding effect cannot be ruled out. In environments with small-scale disturbances, e.g., ant trails, gemmae are favoured over spores at establishment. Also, the diaspore bank of the forest floor can be activated by disturbances, which may affect SGS. At mid-distances, the isolation by distance effect, presumably related to comparatively ineffective gemma dispersal, is visible, while at greater distances, the role of spores as effective means of dispersal is evident. The Sp statistic values, which quantify the strength of SGS, indicate that outsider colonies possess less SGS than do plant colonies along ant trails, which relates to the more frequent spore production of outsider colonies. Moreover, dispersal from fallen logs or stumps may be more effective than dispersal from ground-level colonies along ant trails. Apparently, ants do not have much role as dispersal vectors, nor do the physical structures of ant trails as dispersal corridors, although they provide open areas for colonization.     

MUTATIONAL MELTDOWN IN SELFING ARABIDOPSIS LYRATA

The majority of plant species and many animals are hermaphrodites, with individuals expressing both female and male function. Although hermaphrodites can potentially reproduce by self‐fertilization, they have a high prevalence of outcrossing. The genetic advantages of outcrossing are described by two hypotheses: avoidance of inbreeding depression because selfing leads to immediate expression of recessive deleterious mutations, and release from drift load because self‐fertilization leads to long‐term accumulation of deleterious mutations due to genetic drift and, eventually, to extinction. I tested both hypotheses by experimentally crossing Arabidopsis lyrata plants (self‐pollinated, cross‐pollinated within the population, or cross‐pollinated between populations) and measuring offspring performance over 3 years. There were 18 source populations, each of which was either predominantly outcrossing, mixed mating, or predominantly selfing. Contrary to predictions, outcrossing populations had low inbreeding depression, which equaled that of selfing populations, challenging the central role of inbreeding depression in mating system shifts. However, plants from selfing populations showed the greatest increase in fitness when crossed with plants from other populations, reflecting higher drift load. The results support the hypothesis that extinction by mutational meltdown is why selfing hermaphroditic taxa are rare, despite their frequent appearance over evolutionary time.     

Costs of selfing prevent the spread of a self‐compatibility mutation that causes reproductive assurance

In flowering plants, shifts from outcrossing to partial or complete self‐fertilization have occurred independently thousands of times, yet the underlying adaptive processes are difficult to discern. Selfing's ability to provide reproductive assurance when pollination is uncertain is an oft‐cited ecological explanation for its evolution, but this benefit may be outweighed by costs diminishing its selective advantage over outcrossing. We directly studied the fitness effects of a self‐compatibility mutation that was backcrossed into a self‐incompatible (SI) population of Leavenworthia alabamica, illuminating the direction and magnitude of selection on the mating‐system modifier. In array experiments conducted in two years, self‐compatible (SC) plants produced 17–26% more seed, but this advantage was counteracted by extensive seed discounting—the replacement of high‐quality outcrossed seeds by selfed seeds. Using a simple model and simulations, we demonstrate that SC mutations with these attributes rarely spread to high frequency in natural populations, unless inbreeding depression falls below a threshold value (0.57 ≤ δ_threshold ≤ 0.70) in SI populations. A combination of heavy seed discounting and inbreeding depression likely explains why outcrossing adaptations such as self‐incompatibility are maintained generally, despite persistent input of selfing mutations, and frequent limits on outcross seed production in nature.     

LIGHT BOUNDARIES AND THE COUPLED EFFECTS OF SURFACE HYDROPHOBICITY AND LIGHT ON SPORE SETTLEMENT IN THE BROWN ALGA HINCKSIA IRREGULARIS (PHAEOPHYCEAE) 1

We examined the effects of light and surface hydrophobicity individually and in tandem on Hincksia irregularis (Kützing) Amsler spore settlement. Hincksia irregularis spores were determined to be negatively phototactic by the use of computer‐assisted motion analysis. Spore settlement was significantly influenced by surface hydrophobicity and by light, individually and in tandem. Experiments conducted using culture plates modified to reduce well edge artifacts revealed significantly higher settlement on hydrophobic surfaces and in dark environments when compared with negatively charged surfaces and lighted environments. Experimental light/dark boundaries elicited distinct spore settlement responses, with spores displaying dissimilar settlement patterns on plates with different surface hydrophobicities. The results of this study indicate H. irregularis spores possess the capacity for complex responses to their environment. These complex responses may influence dispersal and aid spores in the detection of suitable settlement locations in marine microenvironments.     

PARASITES AND THE EVOLUTION OF SELF-FERTILIZATION

Abstract.— Assuming all else is equal, an allele for selfing should spread when rare in an outcrossing population and rapidly reach fixation. Such an allele will not spread, however, if self‐fertilization results in inbreeding depression so severe that the fitness of selfed offspring is less that half that of outcrossed offspring. Here we consider an ecological force that may also counter the spread of a selfing allele: coevolution with parasites. Computer simulations were conducted for four different genetic models governing the details of infection. Within each of these models, we varied both the level of selfing in the parasite and the level of male‐gamete discounting in the host (i.e., the reduction in outcrossing fitness through male function due to the selfing allele). We then sought the equilibrium level of host selfing under the different conditions. The results show that, over a wide range of conditions, parasites can select for host reproductive strategies in which both selfed and outcrossed progeny are produced (mixed mating). In addition, mixed mating, where it exits, tends to be biased toward selfing.     

Spatial and temporal patterns of larval dispersal in a coral‐reef fish metapopulation: evidence of variable reproductive success

Many marine organisms can be transported hundreds of kilometres during their pelagic larval stage, yet little is known about spatial and temporal patterns of larval dispersal. Although traditional population‐genetic tools can be applied to infer movement of larvae on an evolutionary timescale, large effective population sizes and high rates of gene flow present serious challenges to documenting dispersal patterns over shorter, ecologically relevant, timescales. Here, we address these challenges by combining direct parentage analysis and indirect genetic analyses over a 4‐year period to document spatial and temporal patterns of larval dispersal in a common coral‐reef fish: the bicolour damselfish (Stegastes partitus). At four island locations surrounding Exuma Sound, Bahamas, including a long‐established marine reserve, we collected 3278 individuals and genotyped them at 10 microsatellite loci. Using Bayesian parentage analysis, we identified eight parent‐offspring pairs, thereby directly documenting dispersal distances ranging from 0 km (i.e., self‐recruitment) to 129 km (i.e., larval connectivity). Despite documenting substantial dispersal and gene flow between islands, we observed more self‐recruitment events than expected if the larvae were drawn from a common, well‐mixed pool (i.e., a completely open population). Additionally, we detected both spatial and temporal variation in signatures of sweepstakes and Wahlund effects. The high variance in reproductive success (i.e., ‘sweepstakes’) we observed may be influenced by seasonal mesoscale gyres present in the Exuma Sound, which play a prominent role in shaping local oceanographic patterns. This study documents the complex nature of larval dispersal in a coral‐reef fish, and highlights the importance of sampling multiple cohorts and coupling both direct and indirect genetic methods in order disentangle patterns of dispersal, gene flow and variable reproductive success.     

THE SECRET LIFE OF KELPS: PLANKTONIC PROCESSES AND POPULATION DYNAMICS

Understanding spatio‐temporal variability in recruitment is vital to studies of kelp population dynamics. Research on settlement and post‐settlement processes has suggested that arrival of kelp zoospores to suitable substrate is important in limiting kelp recruitment, yet the role of planktonic processes in kelp population dynamics has not been studied due to difficulties in sampling and identifying zoospores. I developed a method to estimate kelp zoospore abundance from in situ plankton samples and used it to study various processes regulating the availability of giant kelp (Macrocystis pyrifera) zoospores for settlement. My studies focused on (1) identifying temporal scales over which zoospore abundance is most variable, (2) describing physical and biological processes that regulate this variability, and (3) determining the relationship between zoospore abundance and settlement. I found that short‐term variability in zoospore abundance (<24 hrs) was not due to changes in supply but rather dispersion, caused by oscillating hydrodynamic forces (e.g. waves). Long‐term variability in zoospore abundance, however, was best explained by the size and density of reproductive adult plants, with zoospore abundance being most variable at the scale of days to months. Changes in adult reproductive condition caused rapid changes in zoospore abundance suggesting that the supply of kelp zoospores is sensitive to environmental regulation of adult physiology. Thus, unlike with marine animals, these results indicate that variability in kelp propagule supply, over scales most likely to affect subsequent settlement and recruitment, is more tightly coupled to demographic and reproductive mechanisms than to physical transport processes.     

Individual and Temporal Variation in Outcrossing Rates and Pollen Flow Patterns in Ceiba pentandra (Malvaceae: Bombacoidea)

Ceiba pentandra is a tropical tree with high rates of selfing in some populations. In mixed‐mating species, variation in selfing is due to changes in adult density or variability of incompatibility systems. The effect of spatial isolation and phenology on selfing rates and pollen flow distances was analyzed using microsatellites in a fragmented population of Ceiba pentandra, in southern Costa Rica. Adult trees within a heterogeneous landscape were classified as grouped or isolated. We compared selfing rates at the individual level, between isolation conditions and 2 yr (2007, 2009), which differed in the number of flowering individuals. Mixed mating was estimated in both years (t_m = 0.624–0.759). Trees mated predominantly by outcrossing, while only a few trees reproduced through selfing. Spatial isolation did not significantly affect outcrossing rates. The progeny of grouped trees was mostly sired by near‐neighbors (<1 km) and by long‐distance pollen flow events in isolated trees. A reduction in the number of flowering individuals in 2009 reduced near‐neighbor matings, increased selfing in grouped trees, and decreased the number of unsampled sires in the progeny. Comparing selfing rates on individuals that flowered in both reproductive periods suggests a flexible mating system. Variation in self‐fertilization rates in this population appears to depend on variation of individual traits, such as genetic variability in self‐incompatibility genes, but it is independent of landscape heterogeneity. In contrast, pollen flow distances depend on local tree density as bats concentrate their foraging between near individuals to maximize energy efficiency.     

PRESENCE OF SPOROPHYLLS IN FLOATING KELP RAFTS OF MACROCYSTIS SPP. (PHAEOPHYCEAE) ALONG THE CHILEAN PACIFIC COAST1

Some species of macroalgae continue to live for extended periods of time after detachment and may even maintain reproductive structures, yet very little is known about this process. Here, we describe the presence of sporophylls (with sporogenous tissues) on floating kelp rafts of Macrocystis spp. along the coast of Chile. Surveys were conducted at nine sites (18–50° S) during austral summer 2002, and floating kelp rafts were seen and collected at seven of these nine sites (between 22 and 50° S). Fifteen (26.8%) of the 56 samples had sporophylls, indicating maintenance of sporophylls after detachment. Some of the kelp sporophytes with reproductive blades showed signs of having been afloat for long periods (indicated by the large size of attached stalked barnacles). Additionally, experiments showed that floating kelps released viable zoospores. To understand the reproductive dynamics of floating kelps, we compared these results with information from attached populations of Macrocystis spp. at nearby coastal sites. In general, attached kelp had higher proportions of sporophylls than floating rafts, suggesting that detachment may negatively affect reproductive status. Nevertheless, floating kelps remained functionally reproductive, suggesting that zoospores may be dispersed via floating rafts. Published reports on other macroalgae indicate that some species (Lessoniaceae, Fucaceae, and Sargassaceae) are fertile and probably release zoospores or zygotes while floating or drifting in ocean currents. Because dispersal distances achieved by spores of most macroalgae are relatively short, release of spores from floating algae may be an alternative mechanism of long‐distance dispersal.     

The effects of mysid grazing on kelp zoospore survival and settlement

Recent studies have indicated that long‐distance dispersal by kelp zoospores may play an important role in the colonization of newly exposed rocky habitats and in the recovery of recently disturbed kelp forests. This may be facilitated by the vertical transport of zoospores into the shallower portions of the water column where they are exposed to greater alongshore currents that increase their dispersal potential. However, this vertical transport can also expose them to elevated irradiances and enhanced grazing by zooplankton, both of which negatively impact zoospore survival and settlement. In this study, we used plankton tows to show that zooplankton (mysids) were at least seven times more abundant in the surface waters than near the benthos along the edge of a large kelp forest at the time of our spring sampling. We then used feeding experiments and epifluorescence microscopy to verify that these mysids grazed on kelp zoospores. Finally, we conducted laboratory experiments to show that grazing by these mysids over a 12 h period reduced kelp zoospore settlement by at least 50% relative to treatments without grazing. Together with previous studies that have revealed the impacts of high irradiance on zoospore survival and settlement, our study indicates that the vertical transport of kelp zoospores into the shallower portions of the water can also expose them to significantly increased mortality from mysid grazing. Thus, if these patterns are consistent over broader temporal and geographic scales, vertical transport may not be a viable method for sustained long‐distance zoospore dispersal.     

Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales

We used species‐specific spore traps to measure airborne dispersal of the wood decay fungus Phlebia centrifuga (spore size 6.5–9 × 2.5–3 μm) up to 1000 m distance from a point source. We fitted two simple dispersal models, an empirical power law model and a semi‐mechanistic diffusion model to the data using the Bayesian approach. The diffusion model provided a better fit than the power law model which underestimated deposition at 3–55 m and overestimated deposition at longer and shorter distances. Model fit improved by allowing overdispersion, suggesting that spores are not dispersed independently but wind can transport spores in groups inside discrete air packages up to considerable distances. Using the fitted diffusion model and available information on the establishment rates of wood‐decay fungi, we examine the distance up to which colonisation from a single fruit body is likely to occur. We conclude that the diluting effect of distance and low establishment success make the occurrence of P. centrifuga dispersal limited possibly already at the distance of tens of metres and very probably at a few hundred metres from the nearest fruit body, despite the fact that under favourable conditions a high proportion of the spores can disperse considerably further. This conclusion is likely to hold generally for those fungal species that inhabit fragmented landscapes, have specialised resource and habitat requirements, and have similar spore size and other dispersal traits as P. centrifuga.     

Dispersal potential of spores and asexual propagules in the epixylic hepatic Anastrophyllum hellerianum

Dispersal ability is of great importance for plants, which commonly occupy spatially and temporally limited substrate patches. Mixed reproductive strategies with abundant diaspore production are favoured in a heterogeneous landscape to ensure successful colonisation at different distances. In bryophytes, long-distance dispersal has been thought to take place primarily by spores, while asexual propagules are important in local dispersal and in the maintenance of colonies. In the present study, we investigated the dispersal potential of two equally sized propagules, sexually formed spores and asexually produced gemmae in the dioecious, epixylic hepatic, Anastrophyllum hellerianum, which inhabits spatially and temporally limited substrate patches. We trapped propagules at different distances (0–10 m) and directions from the source colonies in two experiments: one in a natural habitat within a forest and another involving an artificial set-up in an open habitat. Spore dispersal showed only slight distance dependence both in the open and the forest habitats, presumably as a consequence of wind affecting the dispersal pattern. Gemma dispersal was more strongly distance-dependent in the open habitat than in the forest sites. Considerably more gemmae were deposited during rainy than dry periods, possibly because of the effect of rain drops on gemma release. However, weather conditions had no effect on the dispersal patterns of spores or gemmae. In A. hellerianum, the combination of occasional spore production and practically continuous, massive gemma production facilitates dispersal both on local scale and over long distances. Unlike previously assumed, not only spores but also the asexual propagules may contribute to long-distance dispersal, thus allowing considerable gene flow at the landscape level.     

Coevolutionary interactions with parasites constrain the spread of self‐fertilization into outcrossing host populations

Given the cost of sex, outcrossing populations should be susceptible to invasion and replacement by self‐fertilization or parthenogenesis. However, biparental sex is common in nature, suggesting that cross‐fertilization has substantial short‐term benefits. The Red Queen hypothesis (RQH) suggests that coevolution with parasites can generate persistent selection favoring both recombination and outcrossing in host populations. We tested the prediction that coevolving parasites can constrain the spread of self‐fertilization relative to outcrossing. We introduced wild‐type Caenorhabditis elegans hermaphrodites, capable of both self‐fertilization, and outcrossing, into C. elegans populations that were fixed for a mutant allele conferring obligate outcrossing. Replicate C. elegans populations were exposed to the parasite Serratia marcescens for 33 generations under three treatments: a control (avirulent) parasite treatment, a fixed (nonevolving) parasite treatment, and a copassaged (potentially coevolving) parasite treatment. Self‐fertilization rapidly invaded C. elegans host populations in the control and the fixed‐parasite treatments, but remained rare throughout the entire experiment in the copassaged treatment. Further, the frequency of the wild‐type allele (which permits selfing) was strongly positively correlated with the frequency of self‐fertilization across host populations at the end of the experiment. Hence, consistent with the RQH, coevolving parasites can limit the spread of self‐fertilization in outcrossing populations.     

A genetic approach to estimating natal dispersal distances and self‐recruitment in resident rainforest birds

Natal dispersal is a fundamental component of the ecology and evolutionary history of birds, yet is often prohibitively difficult to study. We characterized natal dispersal for the first time in a bird using molecular genetic parentage analyses in a tropical rainforest understory species, the chestnut‐backed antbird (Thamnophilidae: Myrmeciza exsul). Median natal dispersal distance was ～800 m (mean = 931 ± 84 (SE) m, n = 48), with ～90% of all distances < 1500 m. We found no evidence of sex‐biased dispersal. An index of self‐recruitment (i.e. individuals establishing a territory within the population of origin) was higher in sites largely or entirely surrounded by non‐forest, suggesting birds are reluctant to disperse out of preferred forest habitat. Via simulations, we confirmed that the genetic data had sufficient resolution to correctly identify parent‐offspring dyads, but lacked resolution to identify other relationships (full‐sib and half‐sib) with confidence. Chestnut‐backed antbirds have measurable self‐recruitment rates caused by short natal dispersal distances, and self‐recruitment may be amplified by reluctance to disperse out of sites bordered by non‐forest. Some tropical forest understory birds have naturally short dispersal distances, and our results have implications for understanding how species will be affected by fragmented landscapes and for the design of reserves.     

Repeated evolution and reversibility of self‐fertilization in the volvocine green algae*

Outcrossing and self‐fertilization are fundamental strategies of sexual reproduction, each with different evolutionary costs and benefits. Self‐fertilization is thought to be an evolutionary “dead‐end” strategy, beneficial in the short term but costly in the long term, resulting in self‐fertilizing species that occupy only the tips of phylogenetic trees. Here, we use volvocine green algae to investigate the evolution of self‐fertilization. We use ancestral‐state reconstructions to show that self‐fertilization has repeatedly evolved from outcrossing ancestors and that multiple reversals from selfing to outcrossing have occurred. We use three phylogenetic metrics to show that self‐fertilization is not restricted to the tips of the phylogenetic tree, a finding inconsistent with the view of self‐fertilization as a dead‐end strategy. We also find no evidence for higher extinction rates or lower speciation rates in selfing lineages. We find that self‐fertilizing species have significantly larger colonies than outcrossing species, suggesting the benefits of selfing may counteract the costs of increased size. We speculate that our macroevolutionary results on self‐fertilization (i.e., non‐tippy distribution, no decreased diversification rates) may be explained by the haploid‐dominant life cycle that occurs in volvocine algae, which may alter the costs and benefits of selfing.     

Reduced mate availability leads to evolution of self‐fertilization and purging of inbreeding depression in a hermaphrodite

Basic models of mating‐system evolution predict that hermaphroditic organisms should mostly either cross‐fertilize, or self‐fertilize, due to self‐reinforcing coevolution of inbreeding depression and outcrossing rates. However transitions between mating systems occur. A plausible scenario for such transitions assumes that a decrease in pollinator or mate availability temporarily constrains outcrossing populations to self‐fertilize as a reproductive assurance strategy. This should trigger a purge of inbreeding depression, which in turn encourages individuals to self‐fertilize more often and finally to reduce male allocation. We tested the predictions of this scenario using the freshwater snail Physa acuta, a self‐compatible hermaphrodite that preferentially outcrosses and exhibits high inbreeding depression in natural populations. From an outbred population, we built two types of experimental evolution lines, controls (outcrossing every generation) and constrained lines (in which mates were often unavailable, forcing individuals to self‐fertilize). After ca. 20 generations, individuals from constrained lines initiated self‐fertilization earlier in life and had purged most of their inbreeding depression compared to controls. However, their male allocation remained unchanged. Our study suggests that the mating system can rapidly evolve as a response to reduced mating opportunities, supporting the reproductive assurance scenario of transitions from outcrossing to selfing.