Although it is generally agreed that the Arctic flora is among the youngest and least diverse on Earth, the processes that shaped it are poorly understood. Here we present 50 thousand years (kyr) of Arctic vegetation history, derived from the first large-scale ancient DNA metabarcoding study of circumpolar plant diversity. For this interval we also explore nematode diversity as a proxy for modelling vegetation cover and soil quality, and diets of herbivorous megafaunal mammals, many of which became extinct around 10 kyr bp (before present). For much of the period investigated, Arctic vegetation consisted of dry steppe-tundra dominated by forbs (non-graminoid herbaceous vascular plants). During the Last Glacial Maximum (25-15 kyr bp), diversity declined markedly, although forbs remained dominant. Much changed after 10 kyr bp, with the appearance of moist tundra dominated by woody plants and graminoids. Our analyses indicate that both graminoids and forbs would have featured in megafaunal diets. As such, our findings question the predominance of a Late Quaternary graminoid-dominated Arctic mammoth steppe.

The Arctic is an excellent model system for the study of polyploidy. It is one the Earth's most polyploid-rich areas, in particular of high-level and recently evolved polyploids. Here we re-address previous hypotheses on arctic polyploidy based on a new analysis of the circumarctic flora, and review recent molecular, cytological and reproductive studies. The frequency and level of polyploidy strongly increase northwards within the Arctic. We found no clear-cut association between polyploidy and the degree of glaciation for the arctic flora as a whole, which contains many widespread species. However, for 'arctic specialist' taxa with restricted distributions, the frequency of diploids is much higher in the Beringian area, which remained largely unglaciated during the last ice age, than in the heavily glaciated Atlantic area. This result supports the hypothesis that polyploids are more successful than diploids in colonizing after deglaciation. There is abundant molecular evidence for recurrent formation of arctic polyploids at different scales in time and space. Examples are given of low-level polyploids formed after the last glaciation and of repeated and successively more high-level polyploidizations throughout the Quaternary. Recurrent polyploid origins, followed by interbreeding within and across ploidal levels, provide a major explanation for the taxonomic complexity of the arctic flora. In the well-studied, recently deglaciated archipelago of Svalbard, most species are mainly selffertilizing or clonal. All Svalbard polyploids examined so far are genetic allopolyploids with fixed heterozygosity at isozyme loci. The level of heterozygosity in 65 taxa increases dramatically from diploids to high-level polyploids. In the circumarctic area, there is evidence for numerous recently evolved sibling species within diploid taxonomic species. Rapid evolution of crossing barriers at the diploid level promotes further diversification after expansion from different refugia, and may provide new raw materials for allopolyploid formation. We conclude that the evolutionary success of polyploids in the Arctic may be based on their fixed-heterozygous genomes, which buffer against inbreeding and genetic drift through periods of dramatic climate change.

Up to the 1960s, there was nearly complete consensus that disjunctions and endemism in the North Atlantic cannot be explained without in situ survival during the glaciations (the "nunatak hypothesis"). The alternative "tabula rasa hypothesis" of postglacial immigration was regarded to be of merely historical interest. Herein we review recent geological, molecular, taxonomic, and biogeographic data to re‐examine this view. There is now strong geological evidence for some ice‐free North Atlantic areas within the maximum limits of the Late Weichselian/Wisconsian ice sheets, but no fossils have been found to prove continuous in situ existence of life in these areas. Molecular data suggest that many plants and animals have migrated recently across the Atlantic, even if they lack mechanisms promoting long‐distance dispersal. In other species, there are deep trans‐oceanic phylogeographic splits suggesting survival in two or more refugia, but these refugia may have been located outside the ice sheets. For vascular plants, we provide an updated list of 77 north boreal, alpine, and arctic taxa accepted as North Atlantic endemics. The degree of endemism is very low (0.0‐1.9% single‐region endemism). Forty endemics occur in more than one of the isolated Atlantic regions, indicating extensive migration and complicating inferences on the location of refugia. Thirty‐four endemics are probably not hardy enough for nunatak survival and are explained by postglacial immigration (or in situ evolution). Among the 43 "hardy" endemics, there is not a single outcrossing diploid that could suggest long‐term evolution. Most of the hardy endemics are asexual or self‐fertilizing polyploids, some of postglacial hybrid origin. Others are preglacial polyploids which immigrated postglacially or survived in situ. Some ice‐free areas, such as the extensive Greenlandic ones, may have supported survival of some hardy organisms. The evidence accumulated since the 1960s suggests, however, that endemism and disjunctions in the North Atlantic can be explained without invoking in situ glacial survival.

Palaeoenvironments and former climates are typically inferred from pollen and macrofossil records. This approach is time-consuming and suffers from low taxonomic resolution and biased taxon sampling. Here, we test an alternative DNA-based approach utilizing the P6 loop in the chloroplast trnL (UAA) intron; a short (13-158 bp) and variable region with highly conserved flanking sequences. For taxonomic reference, a whole trnL intron sequence database was constructed from recently collected material of 842 species, representing all widespread and/or ecologically important taxa of the species-poor arctic flora. The P6 loop alone allowed identification of all families, most genera (>75%) and one-third of the species, thus providing much higher taxonomic resolution than pollen records. The suitability of the P6 loop for analysis of samples containing degraded ancient DNA from a mixture of species is demonstrated by high-throughput parallel pyrosequencing of permafrost-preserved DNA and reconstruction of two plant communities from the last glacial period. Our approach opens new possibilities for DNA-based assessment of ancient as well as modern biodiversity of many groups of organisms using environmental samples.

Our study provides new knowledge of two processes that are important for plant adaptation in a changing environment: 1) long-distance dispersal patterns, and 2) genetic founder effect on islands. Although the theoretical framework for the genetic founder effect on islands was proposed in 1973, we are the first to quantify it in relation to island size, dispersal distance, and plant traits. In addition, our genetic results are mainly coherent with post-glacial colonisation rather than in situ glacial survival, and should therefore bring a final end to the 140-year-long glacial survival-tabula rasa debate among northern biologists.

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Paradoxically, several of the ecologically most important plant groups in the Arctic are little understood in terms of taxonomy and biogeographic history. The circumpolar Carex bigelowii s. l. (Cyperaceae) is abundant in the Arctic and is one of the most complicated arctic plant groups. While its ecology and population genetics have been extensively studied, its taxonomy is largely unexplored. We analyzed the large-scale geographical structuring of amplified fragment length polymorphisms (AFLPs) covering most of the distribution range. We detected high levels of genetic variation, most (66%) within populations, and a fairly weak genetic structure. Only the Central Asian populations, referred to as C. orbicularis, were strongly divergent. For the remaining populations, Bayesian clustering separated three distinct clusters (one European, one amphi-Atlantic, and one broadly amphi-Beringian), probably reflecting different major glacial refugia and recent transoceanic dispersal. The isolated central European populations were most closely related to those from a larger distribution area in northern Europe. Differences in genetic diversity suggest that the Alpine and Tatra populations have experienced strong bottlenecks, whereas the Krkonoše population may have been part of a continuous distribution area during the cold stages of the Pleistocene. Finally, we discuss the relevance of our results for a uniform, range-wide taxonomic concept.

Phylogenetic relationships and biogeography of the genus Cerastium were studied using sequences of three noncoding plastid DNA regions (trnL intron, trnL-trnF spacer, and psbA-trnH spacer). A total of 57 Cerastium taxa was analyzed using two species of the putative sister genus Stellaria as outgroups. Maximum parsimony analyses identified four clades that largely corresponded to previously recognized infrageneric groups. The results suggest an Old World origin and at least two migration events into North America from the Old World. The first event possibly took place across the Bering land bridge during the Miocene. Subsequent colonization of South America occurred after the North and South American continents joined during the Pliocene. A more recent migration event into North America probably across the northern Atlantic took place during the Quaternary, resulting in the current circumpolar distribution of the Arctic species. Molecular clock dating of major biogeographic events was internally consistent on the phylogenetic trees. The arctic high-polyploid species form a polytomy together with some boreal and temperate species of the C. tomentosum group and the C. arvense group. Lack of genetic variation among the arctic species probably indicates a recent origin. The annual life form is shown to be of polyphyletic origin.