Evolutionary imbalance, climate and human history jointly shape the global biogeography of alien plants.

Human activities are causing global biotic redistribution, translocating species and providing them with opportunities to establish populations beyond their native ranges. Species originating from certain global regions, however, are disproportionately represented among naturalized aliens. The evolutionary imbalance hypothesis posits that differences in absolute fitness among biogeographic divisions determine outcomes when biotas mix. Here, we compile data from native and alien distributions for nearly the entire global seed plant flora and find that biogeographic conditions predicted to drive evolutionary imbalance act alongside climate and anthropogenic factors to shape flows of successful aliens among regional biotas. Successful aliens tend to originate from large, biodiverse regions that support abundant populations and where species evolve against a diverse backdrop of competitors and enemies. We also reveal that these same native distribution characteristics are shared among the plants that humans select for cultivation and economic use. In addition to influencing species' innate potentials as invaders, we therefore suggest that evolutionary imbalance shapes plants' relationships with humans, impacting which species are translocated beyond their native distributions.

Using genomics to guide seed-sourcing at the right taxonomical level for ecological restoration projects: The complex case of Carex bigelowii s.lat. in Norway.

There is a growing demand for ecological restoration using suitable seeds following international standards or national legal demands for local seed-sourcing. However, before selecting the appropriate geographic origin of seeds, it is vital to explore taxonomic complexity related to the focal taxa. We used ddRAD-seq to screen genomic diversity within Carex bigelowii s.lat. focussing on Norway. This species complex is considered a candidate for seeding, but presents considerable morphological, ecological, and genetic variation. The genetic structure of 132 individuals of C. bigelowii s.lat., including Carex nigra as an outgroup, was explored using ordinations, clustering analyses, and a genetic barrier algorithm. Two highly divergent clusters were evident, supporting the recognition of two taxonomic units "C. dacica" and C. bigelowii "subsp. bigelowii". Previously defined seed-sourcing regions for C. bigelowii s.lat. did not consider the known taxonomic complexity, and therefore interpreted the overall genetic structure as seed-sourcing regions, not taxa. We estimated genetic neighborhood sizes within each taxon to be 100-150 km and 300 km, respectively, indicating species-specific delimitations of local seed-sourcing regions. Frequent hybrids, local genetic distinctiveness, and suggested ecotypes add complexity to the discussed seed-sourcing regions. Our results show how genomic screening of diversity and structure in a species complex can alleviate the taxonomic impediment, inform practical questions, and legal requirements related to seed-sourcing, and together with traditional taxonomic work provide necessary information for a sound management of biodiversity.

Status and trends in Arctic vegetation: Evidence from experimental warming and long-term monitoring.

Changes in Arctic vegetation can have important implications for trophic interactions and ecosystem functioning leading to climate feedbacks. Plot-based vegetation surveys provide detailed insight into vegetation changes at sites around the Arctic and improve our ability to predict the impacts of environmental change on tundra ecosystems. Here, we review studies of changes in plant community composition and phenology from both long-term monitoring and warming experiments in Arctic environments. We find that Arctic plant communities and species are generally sensitive to warming, but trends over a period of time are heterogeneous and complex and do not always mirror expectations based on responses to experimental manipulations. Our findings highlight the need for more geographically widespread, integrated, and comprehensive monitoring efforts that can better resolve the interacting effects of warming and other local and regional ecological factors.

Non-native vascular flora of the Arctic: Taxonomic richness, distribution and pathways.

We present a comprehensive list of non-native vascular plants known from the Arctic, explore their geographic distribution, analyze the extent of naturalization and invasion among 23 subregions of the Arctic, and examine pathways of introductions. The presence of 341 non-native taxa in the Arctic was confirmed, of which 188 are naturalized in at least one of the 23 regions. A small number of taxa (11) are considered invasive; these plants are known from just three regions. In several Arctic regions there are no naturalized non-native taxa recorded and the majority of Arctic regions have a low number of naturalized taxa. Analyses of the non-native vascular plant flora identified two main biogeographic clusters within the Arctic: American and Asiatic. Among all pathways, seed contamination and transport by vehicles have contributed the most to non-native plant introduction in the Arctic.

Population genomic evidence for plant glacial survival in Scandinavia.

Quaternary glaciations have played a major role in shaping the genetic diversity and distribution of plant species. Strong palaeoecological and genetic evidence supports a postglacial recolonization of most plant species to northern Europe from southern, eastern and even western glacial refugia. Although highly controversial, the existence of small in situ glacial refugia in northern Europe has recently gained molecular support. We used genomic analyses to examine the phylogeography of a species that is critical in this debate. Carex scirpoidea Michx subsp. scirpoidea is a dioecious, amphi-Atlantic arctic-alpine sedge that is widely distributed in North America, but absent from most of Eurasia, apart from three extremely disjunct populations in Norway, all well within the limits of the Weichselian ice sheet. Range-wide population sampling and variation at 5,307 single nucleotide polymorphisms show that the three Norwegian populations comprise unique evolutionary lineages divergent from Greenland with high between-population divergence. The Norwegian populations have low within-population genetic diversity consistent with having experienced genetic bottlenecks in glacial refugia, and host private alleles that probably accumulated in long-term isolated populations. Demographic analyses support a single, pre-Weichselian colonization into Norway from East Greenland, and subsequent divergence of the three populations in separate refugia. Other refugial areas are identified in North-east Greenland, Minnesota/Michigan, Colorado and Alaska. Admixed populations in British Columbia and West Greenland indicate postglacial contact. Taken together, evidence from this study strongly indicates in situ glacial survival in Scandinavia.

Long-distance plant dispersal to North Atlantic islands: colonization routes and founder effect.

Long-distance dispersal (LDD) processes influence the founder effect on islands. We use genetic data for 25 Atlantic species and similarities among regional floras to analyse colonization, and test whether the genetic founder effect on five islands is associated with dispersal distance, island size and species traits. Most species colonized postglacially via multiple dispersal events from several source regions situated 280 to >3000 km away, and often not from the closest ones. A strong founder effect was observed for insect-pollinated mixed maters, and it increased with dispersal distance and decreased with island size in accordance with the theory of island biogeography. Only a minor founder effect was observed for wind-pollinated outcrossing species. Colonization patterns were largely congruent, indicating that despite the importance of stochasticity, LDD is mainly determined by common factors, probably dispersal vectors. Our findings caution against a priori assuming a single, close source region in biogeographic analyses.

Glacial survival may matter after all: nunatak signatures in the rare European populations of two west-arctic species.

Biogeographers claimed for more than a century that arctic plants survived glaciations in ice-free refugia within the limits of the North European ice sheets. Molecular studies have, however, provided overwhelming support for postglacial immigration into northern Europe, even from the west across the Atlantic. For the first time we can here present molecular evidence strongly favouring in situ glacial persistence of two species, the rare arctic-alpine pioneer species Sagina caespitosa and Arenaria humifusa. Both belong to the 'west-arctic element' of amphi-Atlantic disjuncts, having their few and only European occurrences well within the limits of the last glaciation. Sequencing of non-coding regions of chloroplast DNA revealed only limited variation. However, two very distinct and partly diverse genetic groups, one East and one West Atlantic, were detected in each species based on amplified fragment length polymorphisms (AFLPs), excluding postglacial dispersal from North America as explanation for their European occurrences. Patterns of genetic diversity and distinctiveness indicate that glacial populations existed in East Greenland and/or Svalbard (A. humifusa) and in southern Scandinavia (S. caespitosa). Despite their presumed lack of long-distance dispersal adaptations, intermixed populations in several regions indicate postglacial contact zones. Both species are declining in Nordic countries, probably due to climate change-induced habitat loss. Little or no current connectivity between their highly fragmented and partly distinct populations call for conservation of several populations in each geographic region.

Frequent long-distance plant colonization in the changing Arctic.

The ability of species to track their ecological niche after climate change is a major source of uncertainty in predicting their future distribution. By analyzing DNA fingerprinting (amplified fragment-length polymorphism) of nine plant species, we show that long-distance colonization of a remote arctic archipelago, Svalbard, has occurred repeatedly and from several source regions. Propagules are likely carried by wind and drifting sea ice. The genetic effect of restricted colonization was strongly correlated with the temperature requirements of the species, indicating that establishment limits distribution more than dispersal. Thus, it may be appropriate to assume unlimited dispersal when predicting long-term range shifts in the Arctic.