In dioecious species with both sexual and asexual reproduction, the spatial distribution of individual clones affects the potential for sexual reproduction and local adaptation. The seaweed Fucus radicans, endemic to the Baltic Sea, has separate sexes, but new attached thalli may also form asexually. We mapped the spatial distribution of clones (multilocus genotypes, MLGs) over macrogeographic (>500 km) and microgeographic (<100 m) scales in the Baltic Sea to assess the relationship between clonal spatial structure, sexual recruitment, and the potential for natural selection. Sexual recruitment was predominant in some areas, while in others asexual recruitment dominated. Where clones of both sexes were locally intermingled, sexual recruitment was nevertheless low. In some highly clonal populations, the sex ratio was strongly skewed due to dominance of one or a few clones of the same sex. The two largest clones (one female and one male) were distributed over 100–550 km of coast and accompanied by small and local MLGs formed by somatic mutations and differing by 1–2 mutations from the large clones. Rare sexual events, occasional long‐distance migration, and somatic mutations contribute new genotypic variation potentially available to natural selection. However, dominance of a few very large (and presumably old) clones over extensive spatial and temporal scales suggested that either these have superior traits or natural selection has only been marginally involved in the structuring of genotypes.

Closely related taxa provide significant case studies for understanding evolution of new species but may simultaneously challenge species identification and definition. In the Baltic Sea, two dominant and perennial brown algae share a very recent ancestry. Fucus vesiculosus invaded this recently formed postglacial sea 8000 years ago and shortly thereafter Fucus radicans diverged from this lineage as an endemic species. In the Baltic Sea both species reproduce sexually but also recruit fully fertile new individuals by asexual fragmentation. Earlier studies have shown local differences in morphology and genetics between the two taxa in the northern and western Bothnian Sea, and around the island of Saaremaa in Estonia, but geographic patterns seem in conflict with a single origin of F. radicans. To investigate the relationship between northern and Estonian distributions, we analysed the genetic variation using 9 microsatellite loci in populations from eastern Bothnian Sea, Archipelago Sea and the Gulf of Finland. These populations are located in between earlier studied populations. However, instead of bridging the disparate genetic gap between N-W Bothnian Sea and Estonia, as expected from a simple isolation-by-distance model, the new populations substantially increased overall genetic diversity and showed to be strongly divergent from the two earlier analysed regions, showing signs of additional distinct populations. Contrasting earlier findings of increased asexual recruitment in low salinity in the Bothnian Sea, we found high levels of sexual reproduction in some of the Gulf of Finland populations that inhabit extremely low salinity. The new data generated in this study supports the earlier conclusion of two reproductively isolated but very closely related species. However, the new results also add considerable genetic and morphological complexity within species. This makes species separation at geographic scales more demanding and suggests a need for more comprehensive approaches to further disentangle the intriguing relationship and history of the Baltic Sea fucoids.

In species reproducing both sexually and asexually clones are often more common in recently established populations. Earlier studies have suggested that this pattern arises due to natural selection favouring generally or locally successful genotypes in new environments. Alternatively, as we show here, this pattern may result from neutral processes during species' range expansions. We model a dioecious species expanding into a new area in which all individuals are capable of both sexual and asexual reproduction, and all individuals have equal survival rates and dispersal distances. Even under conditions that favour sexual recruitment in the long run, colonization starts with an asexual wave. After colonization is completed, a sexual wave erodes clonal dominance. If individuals reproduce more than one season, and with only local dispersal, a few large clones typically dominate for thousands of reproductive seasons. Adding occasional long-distance dispersal, more dominant clones emerge, but they persist for a shorter period of time. The general mechanism involved is simple: edge effects at the expansion front favour asexual (uniparental) recruitment where potential mates are rare. Specifically, our model shows that neutral processes (with respect to genotype fitness) during the population expansion, such as random dispersal and demographic stochasticity, produce genotype patterns that differ from the patterns arising in a selection model. The comparison with empirical data from a post-glacially established seaweed species (Fucus radicans) shows that in this case, a neutral mechanism is strongly supported.

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