The population genetics of sporophytic self-incompatibility in Senecio squalidus L. (Asteraceae): the number, frequency, and dominance interactions of S alleles across its British range

Sporophytic self-incompatibility (SSI) was studied in 11 British Senecio squalidus populations to quantify mating system variation and determine how its recent colonization of the United Kingdom has influenced its mating behavior. S allele number, frequency, and dominance interactions in populations were assessed using full diallels of controlled pollinations. A mean of 5.1 S alleles per population was observed, and no population contained more than six S alleles. Numbers of S alleles within populations of S. squalidus declined with increasing distance from the center of its introduction (Oxford). Cross-classification of S alleles allowed an estimate of approximately seven and no more than 11 S alleles for the entire British S. squalidus population. The low number of S alleles observed in British S. squalidus compared to other SI species is consistent with the population bottleneck associated with S. squalidus' introduction to the Oxford Botanic Garden and subsequent colonization of Britain. Extensive S allele dominance interactions were observed to be a feature of the S. squalidus SSI system and may represent an adaptive response to improve limited mate availability imposed by the presence of so few S alleles. Multilocus allozyme genotypes were also identified for individuals in all populations and geographic patterns of S locus and allozyme loci variation investigated. Less interpopulation structure was observed for the S locus than for allozyme diversity--a finding indicative of the effects of negative frequency-dependent selection at the S locus maintaining equal S phenotypes within populations and enhancing effective migration between populations.     

The population genetics of sporophytic self-incompatibility in Senecio squalidus L. (Asteraceae) I: S allele diversity in a natural population

Twenty-six individuals of the sporophytic self-incompatible (SSI) weed, Senecio squalidus were crossed in a full diallel to determine the number and frequency of S alleles in an Oxford population. Incompatibility phenotypes were determined by fruit-set results and the mating patterns observed fitted a SSI model that allowed us to identify six S alleles. Standard population S allele number estimators were modified to deal with S allele data from a species with SSI. These modified estimators predicted a total number of approximately six S alleles for the entire Oxford population of S. squalidus. This estimate of S allele number is low compared to other estimates of S allele diversity in species with SSI. Low S allele diversity in S. squalidus is expected to have arisen as a consequence of a disturbed population history since its introduction and subsequent colonisation of the British Isles. Other features of the SSI system in S. squalidus were also investigated: (a) the strength of self-incompatibility response; (b) the nature of S allele dominance interactions; and (c) the relative frequencies of S phenotypes. These are discussed in view of the low S allele diversity estimates and the known population history of S. squalidus.     

Population genetics of sporophytic self-incompatibility in Senecio squalidus L. (Asteraceae) II: a spatial autocorrelation approach to determining mating behaviour in the presence of low S allele diversity

We recently estimated that as few as six S alleles represent the extent of S locus diversity in a British population of the self-incompatible (SI) coloniser Senecio squalidus (Oxford Ragwort). Despite the predicted constraints to mating imposed by such a low number of S alleles, S. squalidus maintains a strong sporophytic self-incompatibility (SSI) system and there is no evidence for a breakdown of SSI or any obvious negative reproductive consequences for this highly successful coloniser. The present paper assesses mating behaviour in an Oxford S. squalidus population through observations of its effect on spatial patterns of genetic diversity and thus the extent to which it is responsible for ameliorating the potentially detrimental reproductive consequences of low S allele diversity in British S. squalidus. A spatial autocorrelation (SA) treatment of S locus and allozyme polymorphism data for four loci indicates that mating events regularly occur at all the distance classes examined from 60 to 480 m throughout the entire sample population. Less SA is observed for S locus data than for allozyme data in accordance with the hypothesis that SSI and low diversity at the S locus are driving these large-scale mating events. The limited population structure at small distances of 60 m and less observed for SA analysis of the Me-2 locus and by F-statistics for all the allozyme data, is evidence of some local relatedness due to limited seed and pollen dispersal in S. squalidus. However, the overall impression of mating dynamics in this S. squalidus population is that of ample potential mating opportunities with many individuals at large population scales, indicating that reproductive success is not seriously affected by few S alleles available for mating interactions.     

Recent hybrid origin and invasion of the British Isles by a self-incompatible species, Oxford ragwort (Senecio squalidus L., Asteraceae)

Senecio squalidus is a diploid hybrid species which originated in the British Isles following the introduction of material collected from a hybrid zone on Mount Etna, Sicily, approximately 300 years ago. Introduced hybrid material was cultivated in the Oxford Botanic Garden and gave rise to the stabilized diploid hybrid species, which later spread throughout much of the UK and into some parts of Ireland. Unusually for an invasive species, S. squalidus has a strong system of sporophytic self-incompatibility (SSI) that may have become modified as a result of its recent hybrid origin and spread. First, S. squalidus contains relatively few S alleles (between 2 and 6 S alleles within individual UK populations) compared to other species with SSI (estimates average ~17 S alleles per population). This most probably reflects the population bottleneck experienced by introduced hybrid material. Second, dominance relationships among S. squalidus S alleles are more extensive than those reported in other species with SSI. Third, although pseudo-self-compatibility occurs sporadically in S. squalidus, it is not widespread, indicating that SSI is maintained in the species despite potential mate availability restrictions imposed by low numbers of S alleles. Surveys of other forms of genetic diversity in S. squalidus show that allozyme variation is reduced relative to that within the progenitor species, but Randomly Amplified Polymorphic DNA variation is relatively high. Both types of genetic variation show little or no pattern of isolation-by-distance between populations in keeping with the recent range expansion of the species. During its spread in the British Isles, S. squalidus has hybridized with the native self-compatible (SC) tetraploid species, S. vulgaris, which has led to the origin of three new SC hybrid taxa: a radiate form of S. vulgaris (var. hibernicus), a tetrapoid hybrid species (S. eboracensis) and an allohexaploid (S. cambrensis).     

Sporophytic self-incompatibility in Senecio squalidus L (Asteraceae)--the search for S

Senecio squalidus (Oxford Ragwort) is being used as a model species to study the genetics and molecular genetics of self-incompatibility (SI) in the Asteraceae. S. squalidus has a strong system of sporophytic SI (SSI) and populations within the UK contain very few S alleles probably due to a population bottleneck experienced on its introduction to the UK. The genetic control of SSI in S. squalidus is complex and may involve a second locus epistatic to S. Progress towards identifying the female determinant of SSI in S. squalidus is reviewed here. Research is focused on plants carrying two defined S alleles, S(1) and S(2). S(2) is dominant to S(1) in pollen and stigma. RT-PCR was used to amplify three SRK-like cDNAs from stigmas of S(1)S(2) heterozygotes, but the expression patterns of these cDNAs suggest that they are unlikely to be directly involved in SI or pollen-stigma interactions in contrast to SSI in the Brassicaceae. Stigma-specific proteins associated with the S(1) allele and the S(2) allele have been identified using isoelectric focusing and these proteins have been designated SSP1 (Stigma S-associated Protein 1) and SSP2. SSP1 and SSP2 cDNAs have been cloned by 3' and 5' RACE and shown to be allelic forms of the same gene, SSP. The expression of SSP and its linkage to the S locus are currently being investigated. Initial results show SSP to be expressed exclusively in stigmas and developmentally regulated, with maximal expression occurring at and just before anthesis when SI is fully functional, SSP expression being undetectable in immature buds. Together these data suggest that SSP is a strong candidate for a Senecio S-gene.     

Diversity of S-alleles and mate availability in 3 populations of self-incompatible wild pear (Pyrus pyraster)

Small populations of self-incompatible plants may be expected to be threatened by the limitation of compatible mating partners (i.e., S-Allee effect). However, few empirical studies have explicitly tested the hypothesis of mate limitation in small populations of self-incompatible plants. To do so, we studied wild pear (Pyrus pyraster), which possesses a gametophytic self-incompatibility system. We determined the S-genotypes in complete samplings of all adult trees from 3 populations using a PCR-RFLP approach. We identified a total of 26 different S-alleles, homologous to S-alleles of other woody Rosaceae. The functionality of S-alleles and their Mendelian inheritance were verified in artificial pollination experiments and investigations of pollen tube growth. The smallest population (N = 8) harbored 9 different S-alleles and showed a mate availability of 92.9%, whereas the 2 larger populations harbored 18 and 25 S-alleles and exhibited mate availabilities of 98.4% and 99.2%, respectively. Therefore, we conclude that even small populations of gametophytic self-incompatible plants may exhibit high diversity at the S-locus and are not immediately threatened owing to reduced mate availability.     

THE EVOLUTION OF DOMINANCE IN SPOROPHYTIC SELF-INCOMPATIBILITY SYSTEMS. II. MATE AVAILABILITY AND RECOMBINATION

Sporophytic self-incompatibility (SSI) is a self-pollen recognition system that enforces outcrossing in plants. Recognition in SSI systems is typically controlled by a complex locus ( S -locus) with separate genes that determine pollen and stigma specificity. Experimental studies show that S -alleles can be dominant, recessive, or codominant, and that the dominance level of a given S -allele can depend upon whether pollen or stigma specificity is examined. Here and in the companion paper by Llaurens and colleagues, the evolution of dominance in single-locus SSI is explored using numerical models and simulation. Particular attention is directed at factors that can cause S -allele dominance to differ in pollen versus stigma. The effect of recombination between the S -locus and modifier locus is also examined. The models predict that limitation in the number of compatible mates is required for the evolution of S -allele dominance in the stigma but not in the pollen. Tight linkage between the S -locus and modifier promotes the evolution of S -allele dominance hierarchies. Model results are interpreted with respect to published information on the molecular basis of dominance in SSI systems, and reported S -allele dominance relationships in a variety of species. These studies show that dominant S -alleles are more common in the pollen than in the stigma, a pattern that when interpreted in light of model predictions, suggests that mate limitation may be relatively infrequent in natural populations with SSI.     

Estimating the number,frequency, and dominance of S-alleles in a natural population of Arabidopsis lyrata(Brassicaceae) with sporophytic control of self-incompatibility

In homomorphic plant self-incompatibility (SI) systems, large numbers of alleles may be maintained at a single Mendelian locus. Most estimators of the number of alleles present in natural populations are designed for gametophytic self-incompatibility systems (GSI) in which the recognition phenotype of the pollen is determined by its own haploid genotype. In sporophytic systems (SSI), the recognition phenotype of the pollen is determined by the diploid genotype of its parent, and dominance differs among alleles. We describe research aimed at estimates of S-allele numbers in a natural population of Arabidopsis lyrata (Brassicaceae), whose SSI system has recently been described. Using a combination of pollination studies and PCR-based identification of alleles at a locus equivalent to the Brassica SRK gene, we identified and sequenced 11 putative alleles in a sample of 20 individuals from different maternal seed sets. The pollination results indicate that we have not amplified all alleles that must be present. Extensive partial incompatibility, nonreciprocal compatibility differences, and evidence of weakened expression of SI in some genotypes, prevent us from determining the exact number of missing alleles based only on cross-pollination data. Although we show that none of the theoretical models currently proposed is completely appropriate for estimating the number of alleles in this system, we estimate that there are between 13 and 16 different S-alleles in our sample, probably between 16 and 25 alleles in the population, and discuss the relative frequency of alleles in relation to dominance.     

S-allele diversity suggests no mate limitation in small populations of a self-incompatible plant

Small populations of self-incompatible plants are assumed to be threatened by a limitation of compatible mating partners due to low genetic diversity at the self-incompatibility (S) locus. In contrast, we show by using a PCR-RFLP approach for S-genotype identification that 15 small populations (N = 8-88) of the rare wild pear (Pyrus pyraster) displayed no mate limitation. S-allele diversity within populations was high (N = 9-21) as was mate availability (92.9-100%). Although population size and S-allele diversity were strongly related, no relationship was found between population size and mate availability, gene diversity (He), or fixation index (F(IS)), based on five neutral microsatellite loci. As we determined the principal mate availability within populations based on the S-genotypes observed, the realized mate availability under natural conditions may differ from our estimates, for example, due to spatially limited pollen dispersal. We therefore urge studies on self-incompatible plants to proceed from the simple assessment of principal mate availability to the determination of realized mate availability in natural populations.     

EVOLUTION OF DOMINANCE IN SPOROPHYTIC SELF-INCOMPATIBILITY SYSTEMS: I. GENETIC LOAD AND COEVOLUTION OF LEVELS OF DOMINANCE IN POLLEN AND PISTIL

Recent theoretical advances have suggested that various forms of balancing selection may promote the evolution of dominance through an increase of the proportion of heterozygote genotypes. We test whether dominance can evolve in the sporophytic self-incompatibility (SSI) system in plants. SSI prevents mating between individuals expressing identical SI phenotypes by recognition of pollen by pistils, which avoids selfing and inbreeding depression. SI phenotypes depend on a complex network of dominance relationships between alleles at the self-incompatibility locus ( S -locus). Empirical studies suggest that these relationships are not random, but the exact evolutionary processes shaping these relationships remain unclear. We investigate the expected patterns of dominance under the hypothesis that dominance is a direct target of natural selection. We follow the fate of a mutant allele at the S -locus whose dominance relationships are changed but whose specificity remains unaltered. We show that strict codominance is not evolutionarily stable in SSI, and that inbreeding depression due to deleterious mutations linked or unlinked to the S -locus exerts strong constraints on changes in relative levels of dominance in pollen and pistil. Our results provide a general adaptive explanation for most patterns of dominance relationships empirically observed in natural plant populations.     

Modes and rates of selfing and associated inbreeding depression in the self-incompatible plant Senecio squalidus (Asteraceae): a successful colonizing species in the British Isles

The strength of the self-incompatibility (SI) response in Senecio squalidus was measured across its British range. Geographic variation in SI was investigated and the extent and inheritance of pseudo-self-compatibility (PSC) and inbreeding depression were determined. Mean self-fruit-set per capitulum was calculated for individuals and sample populations. The heritability of PSC and the magnitude of inbreeding depression were assessed by comparing selfing rates and fitness trait values between SI and PSC parent-progeny lines. SI was found to be strongly expressed in S. squalidus throughout its British range, with only 3.1% of the individuals sampled showing PSC. This PSC had relatively low heritability with stronger expression of SI in selfed progeny relative to PSC parents. Inbreeding depression was shown to be great in S. squalidus, with mean life history stage values ranging from 0.18 to 0.25. The strength of SI in S. squalidus appears not to have weakened in response to its rapid colonization of Britain. The avoidance of inbreeding depression is likely to be the primary factor maintaining strong SI in this successful colonizing species.     

Incest versus abstinence: reproductive trade‐offs between mate limitation and progeny fitness in a self‐incompatible invasive plant

Plant mating systems represent an evolutionary and ecological trade‐off between reproductive assurance through selfing and maximizing progeny fitness through outbreeding. However, many plants with sporophytic self‐incompatibility systems exhibit dominance interactions at the S‐locus that allow biparental inbreeding, thereby facilitating mating between individuals that share alleles at the S‐locus. We investigated this trade‐off by estimating mate availability and biparental inbreeding depression in wild radish from five different populations across Australia. We found dominance interactions among S‐alleles increased mate availability relative to estimates based on individuals that did not share S‐alleles. Twelve of the sixteen fitness variables were significantly reduced by inbreeding. For all the three life‐history phases evaluated, self‐fertilized offspring suffered a greater than 50% reduction in fitness, while full‐sib and half‐sib offspring suffered a less than 50% reduction in fitness. Theory indicates that fitness costs greater than 50% can result in an evolutionary trajectory toward a stable state of self‐incompatibility (SI). This study suggests that dominance interactions at the S‐locus provide a possible third stable state between SI and SC where biparental inbreeding increases mate availability with relatively minor fitness costs. This strategy allows weeds to establish in new environments while maintaining a functional SI system.     

The evolution of self-incompatibility when mates are limiting

Self-incompatibility (SI) is a genetic barrier to inbreeding that is broadly distributed in angiosperms. In finite populations of SI plants, the loss of S-allele diversity can limit plant reproduction by reducing the availability of compatible mates. Many studies have shown that small or fragmented plant populations suffer from mate limitation. The advent of molecular typing of S-alleles in many species has paved the way to address quantitatively the importance of mate limitation, and to provide greater insight into why and how SI systems breakdown frequently in nature. In this review, we highlight the ecological factors that contribute to mate limitation in SI taxa, discuss their consequences for the evolution and functioning of SI, and propose new empirical research directions.     

Distribution of S-alleles in island populations of flowering cherry, Prunus lannesiana var. speciosa

We surveyed the distribution of S-alleles in natural island populations of Prunus lannesiana var. speciosa sampled from seven sites on the Izu Peninsula and six Izu islands, Japan. The S-genotypes of sampled individuals were determined by Southern analysis of RFLPs generated by restriction enzyme digestion of genomic DNA, using cDNA of the S-RNase gene as a probe. All individuals were heterozygous, as expected under gametophytic self-incompatibility (GSI). Sixty-three S-alleles were observed in the species, but 12 private to the Izu Peninsula population seemed to be derived from related species, giving a total of 75. The estimated number of S-alleles in each population ranged from 26 to 62, and was inversely correlated with the respective population's distance from the Izu Peninsula, the closest point in the mainland to the islands. This geographical cline in the estimated numbers of S-alleles suggests that gene flow to and from the distant island populations was less frequent, and that the studied species has migrated from the mainland to the Izu islands. The genetic relationship at the S-locus among populations also gave an "isolation by distance" pattern. The genetic differentiation at the S-locus among the populations was very low (F(ST) = 0.014, p < 0.001). The number of S-alleles in the species did not seem to depend on genetic differences associated with population subdivisions. This might be due to the greater effective migration rates of S-alleles, as expected under balancing selection in GSI.     

The different mechanisms of sporophytic self-incompatibility

Flowering plants have evolved a multitude of mechanisms to avoid self-fertilization and promote outbreeding. Self-incompatibility (SI) is by far the most common of these, and is found in ca. 60% of flowering plants. SI is a genetically controlled pollen-pistil recognition system that provides a barrier to fertilization by self and self-related pollen in hermaphrodite (usually co-sexual) flowering plants. Two genetically distinct forms of SI can be recognized: gametophytic SI (GSI) and sporophytic SI (SSI), distinguished by how the incompatibility phenotype of the pollen is determined. GSI appears to be the most common mode of SI and can operate through at least three different mechanisms, two of which have been characterized extensively at a molecular level in the Solanaceae and Papaveraceae. Because molecular studies of SSI have been largely confined to species from the Brassicaceae, predominantly Brassica species, it is not yet known whether SSI, like GSI, can operate through different molecular mechanisms. Molecular studies of SSI are now being carried out on Ipomoea trifida (Convolvulaceae) and Senecio squalidus (Asteraceae) and are providing important preliminary data suggesting that SSI in these two families does not share the same molecular mechanism as that of the Brassicaceae. Here, what is currently known about the molecular regulation of SSI in the Brassicaceae is briefly reviewed, and the emerging data on SSI in I. trifida, and more especially in S. squalidus, are discussed.     

Routes of origin of two recently evolved hybrid taxa: Senecio vulgaris var. hibernicus and York radiate groundsel (Asteraceae)

The possible pathways of origin of two recently arisen introgressant forms of Senecio vulgaris (i.e., var. hibernicus and York radiate groundsel) were investigated in experimental crosses between tetraploid S. vulgaris var. vulgaris and the normally diploid S. squalidus. Comparison of the morphology of synthesized hybrid progeny with established taxa, by discriminant function analysis, revealed that fertile hybrid offspring similar in morphology to S. vulgaris var. hibernicus and York radiate groundsel could be synthesized: (1) following formation of genomically stable diploid gametes by the triploid hybrid; (2) through the production of unreduced gametes by diploid S. squalidus; and (3) when a tetraploid form of S. squalidus acted as one of the parents. It was evident that hybrid offspring similar in morphology to the two introgressant taxa were more often produced in backcrosses to S. vulgaris than in segregating F2 or F3 generations (53% as opposed to 36%), and that fertile hybrid progeny were formed within two generations. Because hybridization between S. vulgaris and S. squalidus occurs regularly, although at very low frequency, in natural mixed populations in the British Isles, there is the potential for multiple origins to occur in the wild of both S. vulgaris var. hibernicus and York radiate groundsel.     

Segregation analyses of partial self-incompatibility in self and cross progeny of Solanum carolinense reveal a leaky S-allele

Natural populations of self-incompatible species often exhibit marked phenotypic variation among individuals in the strength of self-incompatibility (SI). In previous studies, we found that the strength of the SI response in Solanum carolinense, a weedy invasive with RNase-mediated SI, is a plastic trait. Selfing can be particularly important for weeds and other successional species that typically undergo repeated colonization and local extinction events and whose population sizes are often small. We applied a PCR-based protocol to identify the S-alleles present in 16 maternal genotypes and their offspring and performed a two-generation greenhouse study to determine whether variation in the strength of SI is due to the existence of weak and strong S-alleles differing in their ability to recognize and reject self-pollen. We found that allele S9 sets significantly more self seed than the other S-alleles in the population we sampled and that its ability to self is not dependent on interactions with other S-alleles. Our data suggest that the observed variations in self-fertility are likely due to factors that directly influence the expression of SI by altering the translation, turnover, or activity of the S-RNase. The variability in the strength of SI among individuals that we have observed in this and our previous studies raises the possibility that plasticity in the strength of SI in S. carolinense may play a role in the colonization and establishment of this weedy species.     

Extreme changes to gene expression associated with homoploid hybrid speciation

Hybridization is an important cause of abrupt speciation. Hybrid speciation without a change in ploidy (homoploid hybrid speciation) is well-established in plants but has also been reported in animals and fungi. A notable example of recent homoploid hybrid speciation is Senecio squalidus (Oxford ragwort), which originated in the UK in the 18th Century following introduction of hybrid material from a hybrid zone between S. chrysanthemifolius and S. aethnensis on Mount Etna, Sicily. To investigate genetic divergence between these taxa, we used complementary DNA microarrays to compare patterns of floral gene expression. These analyses revealed major differences in gene expression between the parent species and wild and resynthesized S. squalidus . Comparisons of gene expression between S. aethnensis , S. chrysanthemifolius and natural S. squalidus identified genes potentially involved in local environmental adaptation. The analysis also revealed non-additive patterns of gene expression in the hybrid relative to its progenitors. These expression changes were more dramatic and widespread in resynthesized hybrids than in natural S. squalidus , suggesting that a unique expression pattern may have been fixed during the allopatric divergence of British S. squalidus . We speculate that hybridization-induced gene-expression change may provide an immediate source of novel phenotypic variation upon which selection can act to facilitate homoploid hybrid speciation in plants.     

Recent, allopatric, homoploid hybrid speciation: the origin of Senecio squalidus (Asteraceae) in the British Isles from a hybrid zone on Mount Etna, Sicily

Homoploid hybrid speciation occurs through stabilization of a hybrid segregate (or segregates) isolated by premating and/or postmating barriers from parent taxa. Theory predicts that ecological and spatial isolation are of critical importance during homoploid hybrid speciation, and all confirmed homoploid hybrid species are ecologically isolated from their parents. Until recently, such species have been identified long after they originated, and consequently it has not been possible to determine the relative importance of spatial and ecological isolation during their origin. Here we present evidence for the recent origin (within the past 300 years) of a new homoploid hybrid species, Senecio squalidus (Asteraceae), in the British Isles, following long-distance dispersal of hybrid material from a hybrid zone between S. aethnensis and S. chrysanthemifolius on Mount Etna, Sicily, Italy. Historical records show that such hybrid material from Sicily was introduced to the Oxford Botanic Garden in Britain in the early part of the 18th century and that S. squalidus began to spread from there after approximately 90 years. A survey of randomly amplified polymorphic DNA/intersimple sequence repeats (RAPD/ISSR) marker variation demonstrated that S. squalidus is a diploid hybrid derivative of S. aethnensis and S. chrysanthemifolius that grow at high and low altitudes, respectively, on Mount Etna and that form a hybrid zone at intermediate altitudes. Senecio squalidus contained 11 of 13 RAPD/ISSR markers that were recorded at high frequency in S. chrysanthemifolius but were absent or occurred at low frequency in S. aethnensis, and 10 of 13 markers for which the reverse was true. Bayesian admixture analysis showed that all individuals of S. squalidus surveyed were of mixed ancestry with relatively high mean proportions of ancestry derived from both S. chrysanthemifolius and S. aethnensis (0.644 and 0.356, respectively). We argue that long-distance isolation of hybrid material from its parents on Mount Etna would have helped favor the origin and establishment of S. squalidus in the British Isles, regardless of whether the initial hybrid material introduced to Britain was preadapted to local conditions.     

Molecular genetic and quantitative trait divergence associated with recent homoploid hybrid speciation: a study of Senecio squalidus (Asteraceae)

Hybridization is increasingly seen as a trigger for rapid evolution and speciation. To quantify and qualify divergence associated with recent homoploid hybrid speciation, we compared quantitative trait (QT) and molecular genetic variation between the homoploid hybrid species Senecio squalidus and its parental species, S. aethnensis and S. chrysanthemifolius, and also their naturally occurring Sicilian hybrids. S. squalidus originated and became invasive in the United Kingdom following the introduction of hybrid plants from Mount Etna, Sicily, about 300 years ago. We recorded considerable molecular genetic differentiation between S. squalidus and its parents and their Sicilian hybrids in terms of both reduced genetic diversity and altered allele frequencies, potentially due to the genetic bottleneck associated with introduction to the United Kingdom. S. squalidus is also distinct from its parents and Sicilian hybrids for QTs, but less so than for molecular genetic markers. We suggest that this is due to resilience of polygenic QTs to changes in allele frequency or lack of selection for hybrid niche divergence in geographic isolation. While S. squalidus is intermediate or parental-like for most QTs, some trangressively distinct traits were observed, which might indicate emerging local adaptation in its invasive range. This study emphasizes the important contribution of founder events and geographic isolation to successful homoploid hybrid speciation.