Background and aims

Hybridization is commonly assumed to aid invasions through adaptive introgression. In contrast, a recent theoretical model predicted that there can be non-adaptive demographic advantages from hybridization and that the population consequences will depend on the breeding systems of the species and the extent to which subsequent generations are able to interbreed and reproduce. We examined cross-fertilization success and inheritance of breeding systems of two species in order to better assess the plausibility of the theoretical predictions.

Methods

Reciprocal artificial crosses were made to produce F1, F2 and backcrosses between Cakile maritima (self-incompatible, SI) and Cakile edentula (self-compatible, SC) (Brassicaceae). Flowers were emasculated prior to anther dehiscence and pollen was introduced from donor plants to the recipient's stigma. Breeding system, pollen viability, pollen germination, pollen tube growth and reproductive output were then determined. The results were used to replace the assumptions made in the original population model and new simulations were made.

Key results

The success rate with the SI species as the pollen recipient was lower than when it was the pollen donor, in quantitative agreement with the 'SI × SC rule' of unilateral incompatibility. Similar outcomes were found in subsequent generations where fertile hybrids were produced but lower success rates were observed in crosses of SI pollen donors with SC pollen recipients. Much lower proportions of SC hybrids were produced than expected from a single Mendelian allele. When incorporated into a population model, these results predicted an even faster rate of replacement of the SC species by the SI species than previously reported.

Conclusions

Our study of these two species provides even clearer support for the feasibility of the non-adaptive hybridization hypothesis, whereby the colonization of an SI species can be assisted by transient hybridization with a congener. It also provides novel insight into reproductive biology beyond the F1 generation.

The founding population in most new species introductions, or at the leading edge of an ongoing invasion, is likely to be small. Severe Allee effects-reductions in individual fitness at low population density-may then result in a failure of the species to colonize, even if the habitat could support a much larger population. Using a simulation model for plant populations that incorporates demography, mating systems, quantitative genetics, and pollinators, we show that Allee effects can potentially be overcome by transient hybridization with a resident species or an earlier colonizer. This mechanism does not require the invocation of adaptive changes usually attributed to invasions following hybridization. We verify our result in a case study of sequential invasions by two plant species where the outcrosser Cakile maritima has replaced an earlier, inbreeding, colonizer Cakile edentula (Brassicaceae). Observed historical rates of replacement are consistent with model predictions from hybrid-alleviated Allee effects in outcrossers, although other causes cannot be ruled out.