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.
Ocean climate change strongly affects organisms and ecosystems, and the causes, consequences, and underlying mechanisms need to be documented. In the Baltic Sea, a marginal sea under severe eutrophication stress, a longer productive season, and changes in the phytoplankton community over the last few decades have likely impacted diet and condition of keystone species, from individual to population level. This study uses stable isotopes (δ 13 C, δ 15 N, and derived isotope niche metrics) to trace energy and nutrient flows in archived samples of blue mussel (Mytilus edulis trossulus) spanning 24 yr (1993-2016). We test if long-term changes in isotope and elemental composition in mussels, as well as population abundance and biomass, can be explained by changes in abiotic and biotic variables, using partial least square regressions and structural equation modeling. We found decreasing trends in δ 13 C and δ 15 N as well as in mean size and total biomass of mussels, but no unidirectional changes in their stoichiometry or condition index. Changes in isotope composition were best explained by nitrogen-fixing cyanobacteria, by increased terrestrial organic carbon from land runoff (reflecting precipitation) and by decreases in dissolved inorganic nitrogen (indicative of successful eutrophication mitigation) and in biomass of a mixotrophic ciliate species. The trophic niche (assessed from isotope niche) was included as the best predictor for both mussel body condition and the observed decline in their total biomass. This study reveals that altered trophic relationships from climate-induced changes in the productivity base may strongly impact keystone species, with potential knock-on effects on ecosystem functions.
Genetic characteristics of populations can have substantial impacts on the adaptive potential of a species. Species are heterogeneous, often defined by variability at a range of scales including at the genetic, individual and population level. Using microsatellite genotyping, we characterize patterns underlying the genetic heterogeneity in marine macroalga Fucus vesiculosus, with a particular focus on two forms: attached and free‐living. Here we demonstrate that sympatric populations representing the two forms display marked differences in characteristics of reproduction and genetic diversity. Asexual reproduction was ubiquitous in the free‐living form despite being almost entirely absent in the attached form, while signals of polyploidy were common in both forms despite the distinct reproductive modes. Gene flow within and between the forms differed, with barriers to gene flow occurring between forms at various spatial scales due to the reproductive modes employed by individuals of each form. The divergent genetic characteristics of F. vesiculosus demonstrate that intraspecific differences can influence the properties of populations with consequential effects on the whole ecosystem. The differing genetic patterns and habitat requirements of the two forms define separate but closely associated ecological entities that will likely display divergent responses to future changes in environmental conditions.
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