Unlike seed plants where global biogeographical patterns typically involve interspecific phylogenetic history, spore-producing bryophyte species often have intercontinental distributions that are best understood from a population genetic perspective. We sought to understand how reproductive processes, especially dispersal, have contributed to the intercontinental 'Pacific Rim' distribution of Sphagnum miyabeanum. In total, 295 gametophyte plants from western North America (California, Oregon, British Columbia, Alaska), Russia, Japan, and China were genotyped at 12 microsatellite loci. Nucleotide sequences were obtained for seven anonymous nuclear loci plus two plastid regions from 21 plants of S. miyabeanum and two outgroup species. We detected weak but significant genetic differentiation among plants from China, Japan, Alaska, British Columbia, and the western USA. Alaskan plants are genetically most similar to Asian plants, and British Columbian plants are most similar to those in the western USA. There is detectable migration between regions, with especially high levels between Alaska and Asia (China and Japan). Migration appears to be recent and/or ongoing, and more or less equivalent in both directions. There is weak (but significant) isolation-by-distance within geographical regions, and the slope of the regression of genetic on geographical distance differs for Asian versus North American plants. A distinctive Vancouver Island morphotype is very weakly differentiated, and does not appear to be reproductively isolated from plants of the normal morphotype. The intercontinental geographical range of S. miyabeanum reflects recent and probably ongoing migration, facilitated by the production of tiny spores capable of effective long distance dispersal. The results of the present study are consistent with Pleistocene survival of S. miyabeanum in unglaciated Beringia, although we cannot eliminate the possibility that the species recolonized Alaska from Asia more recently.

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Aim Arctic plant species are often characterized by a complex genetic structure because of changes in their population size, the fragmentation of metapopulation systems, extensive hybridization and allopolyploidization, and survival in disjunct refugia, historical features associated with Pleistocene glaciation. We assessed the biogeographical and genetic patterns in three closely related northern species of peat-moss (Sphagnum inexspectatum, S. orientale and S. miyabeanum), especially interspecific hybridization, infraspecific geographical differentiation and Pleistocene survival in one or more refugial areas. We tested alternative hypotheses of refugial survival in three widely disjunct regions: eastern Asia, Alaska and Greenland.Location North America (Canada, western USA and Greenland), China, Japan and Russia.Methods Four hundred and forty-three plants were genotyped at 12 microsatellite loci. Nucleotide sequences from 130 accessions for two plastid and two nuclear loci were used to reconstruct haplotype relationships. Population genetic analyses produced estimates of genetic diversity, levels of interspecific gene flow and rates of infraspecific intercontinental migration. Approximate Bayesian computation was used to test alternative biogeographical scenarios. ResultsWe found evidence of hybridization between two of the three species, but phylogenetic patterns are predominantly divergent rather than reticulate. Disjunct populations of one species, S. orientale, in Greenland, Alaska and China are genetically differentiated, but migration has occurred among all three metapopulation systems. Divergence-time analyses strongly support the hypothesis that S. orientale survived the Last Glacial Maximum in Beringia and also in Greenland. Main conclusionsOur results indicate that Beringia served as a refugium for peat-mosses, and therefore peatlands, which are currently extensive at high latitudes of the Northern Hemisphere. Sphagnum orientale also appears to have survived the Last Glacial Maximum in Greenland, indicating that the species persisted in multiple Pleistocene refugia. Phylogenetic patterns are relatively simple in these mosses compared with those commonly encountered in Arctic angiosperms. S. inexspectatum: 18 plants Alaska S. inexspectatum: 5 plants AlaskaS. inexpectatum: 7 plants Alaska S. orientale: 5 plants Alaska, British Columbia, China S. orientale: 3 plants Alaska, Greenland S. orientale: 2 plants Alaska, NWT S. orientale: 8 plants Alaska, China S. orientale: 2 plants Alaska S. orientale: 2 plants Alaska S. miyabeanum: 2 plants Idaho S. orientale: 3 plants Alaska S. inexspectatum: 1 plant Alaska S. orientale: 1 plant Alaska S. inexspectatum: 4 plants Alaska S. orientale: 1 plant China S. miyabeanum: 13 plants British Columbia, California, Russia S. miyabeanum: 5 plants California, Wyoming, China S. miyabeanum: 1 plant Alaska S. miyabeanum: 23 plants Alaska, British Columbia, California S. miyabeanum: 1 plant Oregon S. miyabeanum: 17 plants Alaska, Japan, Russia S. miyabeanum: 1 plant Russia Rapd...

Origin of earliest land plants from ancestral algae dramatically accelerated the evolution of Earth’s terrestrial ecosystems, in which microbial symbioses have played key roles. Recent molecular diversification analyses identify the rare, geographically-limited moss Takakia as Earth’s most archaic modern land plant. Despite occupying a phylogenetic position pivotal for understanding earliest plants, Takakia microbial associations are poorly known. Here, we describe symbiosis-related structural features and contig-based metagenomic data that illuminate the evolutionary transition from streptophyte algae to early embryophytes. We observed that T. lepidozioides shares with streptophyte algae secretion of microbe-harboring mucilage and bacterial taxa such as Rhizobium and genes indicating nitrogen fixation. We find that Takakia root-analogs produce lateral mucilage organs that are more complex than generally understood, having structural analogies to angiosperm lateral roots adapted for N-fixation symbioses, including presence of intracellular microbes. We also find structural and metagenomic evidence for mycorrhiza-like species of glomalean fungi (including Rhizophagus irregularis) not previously known for mosses, as well as ascomycete fungi (e.g. Rhizoscyphus ericae) that associate with other early-diverging plants. Because Takakia is the oldest known modern plant genus, this study of plants of a remote locale not strongly influenced by human activities may indicate microbiome features of early land plants.