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Andøya on the NW coast of Norway is a key site for understanding the Last Glacial Maximum (LGM) in northern Europe. Controversy has arisen concerning the local conditions, especially about the timing and extent of local glacial cover, maximum July temperatures and whether pine and/or spruce could have grown there. We reviewed all existing data and add newly analysed ancient sedimentary DNA (sedaDNA), pollen, macrofossils, geochemistry and stable isotopes from three lake sediment cores from Øvre Æråsvatnet. A total of 23 new dates and age-depth modelling suggests the lake has been ice-free since GI2 (Pinus and Picea sedaDNA was found in all three cores but at such low frequencies that it could not be distinguished from background contamination. LGM samples have an exceptionally high organic matter content, with isotopic values indicating that carbon and nitrogen derive from a marine source. Along with finds of bones of the little auk (Alle alle), this indicates that the lake received guano from an adjacent bird colony.SedaDNA, pollen and macrofossil assemblages were dominated by Poaceae, Brassicaceae and Papaver, but scattered occurrence of species currently restricted to the Low Arctic Tundra Zone (July temperature of 8–9 °C) such as Apiaceae (sedaDNA, 8–9 °C), and Alchemilla alpina (macrofossil, 8–9 °C) were also recorded. The review of >14.7 cal ka BP data recorded 94 vascular plant taxa, of which 38% have a northern limit in Shrub Tundra or more southern vegetation zones. This unusual assemblage likely stems from a combination of proximity to ice-free water in summer, geographical isolation linked with stochastic long-distance dispersal events, and the presence of bird-fertilized habitats. The environmental reconstruction based on all records from the area does not preclude local growth of tree species, as the local climate combined with high nutrient input may have led to periodically suitable environmental ‘hotspot’ conditions.

Source at https://doi.org/10.1111/bor.12357 We present a Holocene record of floristic diversity and environmental change for the central Varanger Peninsula, Finnmark, based on ancient DNA extracted from the sediments of a small lake (sedaDNA). The record covers the period c. 10 700 to 3300 cal. a BP and is complemented by pollen data. Measures of species richness, sample evenness and beta diversity were calculated based on sedaDNA sampling intervals and 1000‐year time windows. We identified 101 vascular plant and 17 bryophyte taxa, a high proportion (86%) of which are still growing within the region today. The high species richness (>60 taxa) observed in the Early Holocene, including representatives from all important plant functional groups, shows that modern shrub‐tundra communities, and much of their species complement, were in place as early as c. 10 700 cal. a BP. We infer that postglacial colonization of the area occurred prior to the full Holocene, during the Pleistocene‐Holocene transition, Younger Dryas stadial or earlier. Abundant DNA of the extra‐limital aquatic plant Callitriche hermaphroditica suggests it expanded its range northward between c. 10 200 and 9600 cal. a BP, when summers were warmer than present. High values of Pinus DNA occur throughout the record, but we cannot say with certainty if they represent prior local presence; however, pollen influx values >500 grains cm−2 a−1 between c. 8000 and 7300 cal. a BP strongly suggest the presence of pine woodland during this period. As the site lies beyond the modern tree limit of pine, it is likely that this expansion also reflects a response to warmer Early Holocene summers.

Environmental DNA retrieved from modern soils (eDNA) and late-Quaternary palaeosols and sediments (aDNA and sedaDNA) promises insight into the composition of present and past terrestrial biotic communities, but few studies address the spatial relationship between recovered eDNA and contributing organisms. Svalbard’s vascular plant flora is well known, and a cold climate enhances preservation of eDNA in soils. Thus, Svalbard plant communities are excellent systems for addressing the representation of plant eDNA in soil samples. In two valleys in the inner fjord region of Spitsbergen, we carried out detailed vegetation surveys of circular plots up to a 4-m radius. One or three near-surface soil samples from each plot were used for extraction and metabarcoding of soil-derived eDNA. Use of PCR replicates and appropriate filtering, plus a relevant reference metabarcode catalogue, provided taxon lists that reflected the local flora. There was high concordance between taxa recorded in plot vegetation and those in the eDNA, but floristic diversity was under-sampled, even at the scale of a 1-m radius plot. Most detected taxa grew within < 0.5–1.0 m of the sampling point. Taxa present in vegetation but not in eDNA tended to occur further from the sampling point, and most had above-ground cover of < 5%. Soil-derived eDNA provides a highly local floristic signal, and this spatial constraint should be considered in sampling designs. For palaeoecological or archaeological studies, multiple samples from a given soil horizon that are spatially distributed across the area of interest are likely to provide the most complete picture of species presence.

Source at https://doi.org/10.3389/fevo.2019.00189. The lake sediments of Hässeldala Port in south-east Sweden provide an archive of local and regional environmental conditions ~14.5–9.5 ka BP (thousand years before present) and allow testing DNA sequencing techniques to reconstruct past vegetation changes. We combined shotgun sequencing with plant micro- and macrofossil analyses to investigate sediments dating to the Allerød (14.1–12.7 ka BP), Younger Dryas (12.7–11.7 ka BP), and Preboreal (

International audience

Published version. Source at http://doi.org/10.1111/nph.14199. License CC BY 4.0. Sedimentary DNA (sedDNA) has recently emerged as a new proxy for reconstructing past vegetation, but its taphonomy, source area and representation biases need better assessment. We investigated how sedDNA in recent sediments of two small Scottish lakes reflects a major vegetation change, using well-documented 20th Century plantations of exotic conifers as an experimental system. We used next-generation sequencing to barcode sedDNA retrieved from subrecent lake sediments. For comparison, pollen was analysed from the same samples. The sedDNA record contains 73 taxa (mainly genus or species), all but one of which are present in the study area. Pollen and sedDNA shared 35% of taxa, which partly reflects a difference in source area. More aquatic taxa were recorded in sedDNA, whereas taxa assumed to be of regional rather than local origin were recorded only as pollen. The chronology of the sediments and planting records are well aligned, and sedDNA of exotic conifers appears in high quantities with the establishment of plantations around the lakes. SedDNA recorded other changes in local vegetation that accompanied afforestation. There were no signs of DNA leaching in the sediments or DNA originating from pollen.

Compilation of the species richness data was made possible through the TFI Networks grant to CD, “Effect Studies and Adaptation to Climate Change,” under the Norforsk initiative (2011 – 2014) which supported two CBIONET-AVA workshops held in Denmark during 2013. The genetic studies were funded by the Research Council of Norway (grant nos. 150322/720 and 170952/V40 to CB).

Metabarcoding of lake sediments may reveal current and past biodiversity, but little is known about the degree to which taxa growing in the vegetation are represented in environmental DNA (eDNA) records. We analysed composition of lake and catchment vegetation and vascular plant eDNA at 11 lakes in northern Norway. Out of 489 records of taxa growing within 2 m from the lake shore, 17-49% (mean 31%) of the identifiable taxa recorded were detected with eDNA. Of the 217 eDNA records, 73% and 12% matched taxa recorded in vegetation surveys within 2 m and up to about 50 m away from the lakeshore, respectively, whereas 16% were not recorded in the vegetation surveys of the same lake. The latter include taxa likely overlooked in the vegetation surveys or growing outside the survey area. The percentages detected were 61, 47, 25, and 15 for dominant, common, scattered, and rare taxa, respectively. Similar numbers for aquatic plants were 88, 88, 33 and 62%, respectively. Detection rate and taxonomic resolution varied among plant families and functional groups with good detection of e.g. Ericaceae, Roseaceae, deciduous trees, ferns, club mosses and aquatics. The representation of terrestrial taxa in eDNA depends on both their distance from the sampling site and their abundance and is sufficient for recording vegetation types. For aquatic vegetation, eDNA may be comparable with, or even superior to, in-lake vegetation surveys and therefore be used as an tool for biomonitoring. For reconstruction of terrestrial vegetation, technical improvements and more intensive sampling is needed to detect a higher proportion of rare taxa although DNA of some taxa may never reach the lake sediments due to taphonomical constrains. Nevertheless, eDNA performs similar to conventional methods of pollen and macrofossil analyses and may therefore be an important tool for reconstruction of past vegetation.

Source at https://doi.org/10.3897/neobiota.37.22088. Biosecurity measures are commonly used to prevent the introduction of non-native species to natural environments globally, yet the efficacy of practices is rarely tested under operational conditions. A voluntary biosecurity measure was trialled in the Norwegian high Arctic following concern that non-native species might be transferred to the region on the footwear of travellers. Passengers aboard an expedition cruise ship disinfected their footwear with the broad spectrum disinfectant Virkon S prior to and in-between landing at sites around the remote Svalbard archipelago. The authors evaluated the efficacy of simply stepping through a disinfectant foot bath, which is the most common practice of footwear disinfection aboard expedition cruise ships in the Arctic. This was compared to a more time consuming and little-used method involving drying disinfected footwear, as proposed by other studies. The two practices were evaluated by measuring microbial growth on paired footwear samples before and after disinfection under both conditions. Step-through disinfection did not substantially reduce microbial growth on the footwear. Allowing disinfected footwear to dry, however, reduced the microbial burden significantly to lower levels. Thus, the currently adopted procedures used aboard ships are ineffective at removing microbial burden and are only effective when footwear is given more time to dry than currently granted under operational conditions. These findings underscore results from empirical research performed elsewhere and suggest the need to better relay this information to practitioners. It is suggested that footwear should minimally be wiped dry after step-through disinfection as a reasonable compromise between biosecurity and practicability.

Aim Previous research on how climatic niches vary across species ranges has focused on a limited number of species, mostly invasive, and has not, to date, been very conclusive. Here we assess the d ...

Birches (Betula spp.) hybridize readily, confounding genetic signatures of refugial isolation and postglacial migration. We aimed to distinguish hybridization from range-shift processes in the two widespread and cold-adapted species B. nana and B. pubescens, previously shown to share a similarly east-west-structured variation in plastid DNA (pDNA). We sampled the two species throughout their ranges and included reference samples of five other Betula species and putative hybrids. We analysed 901 individual plants using mainly nuclear high-resolution markers (amplified fragment length polymorphisms; AFLPs); a subset of 64 plants was also sequenced for two pDNA regions. Whereas the pDNA variation as expected was largely shared between B. nana and B. pubescens, the two species were distinctly differentiated at AFLP loci. In B. nana, both the AFLP and pDNA results corroborated the former pDNA-based hypothesis that it expanded from at least two major refugia in Eurasia, one south of and one east of the North European ice sheets. In contrast, B. pubescens showed a striking lack of geographic structuring of its AFLP variation. We identified a weak but significant increase in nuclear (AFLP) gene flow from B. nana into B. pubescens with increasing latitude, suggesting hybridization has been most frequent at the postglacial expansion front of B. pubescens and that hybrids mainly backcrossed to B. pubescens. Incongruence between pDNA and AFLP variation in B. pubescens can be explained by efficient expansion from a single large refugium combined with leading-edge hybridization and plastid capture from B. nana during colonization of new territory already occupied by this more cold-tolerant species.

Invasive giant hogweeds are infamous in Europe for causing ecological and economic damage, but genetic resources for their study are limited. We used next-generation sequencing to develop a microsatellite library for Heracleum persicum, a widespread invasive in Nordic countries. In addition, these markers were cross amplified with the closely related Heracleum mantegazzianum, Heracleum sosnowskyi, Heracleum sphondylium, and the putative hybrid H. persicum × H. sphondylium, as well as the more distantly related Anthriscus sylvestris. We designed and validated 164 primer pairs. A cost-effective PCR approach with modified forward primer, reverse primer, and fluorescently labeled universal tail was used to test the functionality of each marker. Twenty-five of thirty markers screened on eight geographically distant samples of H. persicum were polymorphic. The number of alleles was 2–4 whereas the expected and observed heterozygosity varied from 0.06 to 0.84 and 0.0 to 1.0 respectively. The cross-species amplification efficiency was 84–100 %, in which 60–76 % of the cross-species amplified markers were polymorphic for Heracleum taxa including H. persicum × sphondylium. Three out of eight of the cross-amplified markers were polymorphic in Anthriscus sylvestris. Ordination revealed a clear genetic structure of Heracleum taxa. Thus, these markers can serve as important genetic resources for understanding taxonomy, population genetics, and phylogeny of giant hogweeds and their hybrids, which in turn, is expected to contribute to invasive species management.

Small and peripheral populations often contain low levels of genetic variation. This may limit their ability to adapt to environmental change, including climate warming. In a recent study published in AoB PLANTS, Birkeland, Skjetne and colleagues show that many rare and threatened plant species in the High Arctic archipelago Svalbard harbour low levels of genetic variation. Most of them are probably relicts from the early Holocene warmer period. They have likely experienced strong genetic founder/bottleneck effects due to climatic limitations. Even though temperatures now are rising, it is highly uncertain whether this will be beneficial for these warmth-demanding species., Small, isolated and/or peripheral populations are expected to harbour low levels of genetic variation and may therefore have reduced adaptability to environmental change, including climate warming. In the Arctic, global warming has already caused vegetation change across the region and is acting as a significant stressor on Arctic biodiversity. Many of the rare plants in the Arctic are relicts from early Holocene warm periods, but their ability to benefit from the current warming is dependent on the viability of their populations. We therefore examined Amplified Fragment Length Polymorphism (AFLP) data from regional red listed vascular plant species in the High Arctic archipelago of Svalbard and reference populations from the main distribution area of: (1) Botrychium lunaria, (2) Carex capillaris ssp. fuscidula, (3) Comastoma tenellum, (4) Kobresia simpliciuscula ssp. subholarctica, (5) Ranunculus wilanderi, (6) Sibbaldia procumbens and (7) Tofieldia pusilla. In addition, we gathered population size data in Svalbard. The Svalbard populations had low genetic diversity and distinctiveness and few or no private markers compared to populations outside the archipelago. This is similar to observations in other rare species in Svalbard and the genetic depletion may be due to an initial founder effect and/or a genetic bottleneck caused by late Holocene cooling. There seems to be limited gene flow from other areas and the Svalbard populations should therefore be considered as demographically independent management units. Overall, these management units have small and/or few populations and are therefore prone to stochastic events which may further increase vulnerability to inbreeding depression, loss of genetic variation, and reduced evolutionary potential. Our results support theory predicting lower levels of genetic diversity in small, isolated and/or peripheral populations and may be of importance for management of other rare plant species in the Arctic.

This work was supported by the Swedish Research Council (grant no. 2013-D0568401), SciLifeLab Stockholm and the Carl Triggers’ Foundation (grant no. 14:371) to L.P., and the Research Council of Norway to I.G.A. (grant no. 213692/F20). Recent advances in sequencing technologies now permit the analyses of plant DNA from fossil samples (ancient plant DNA, plant aDNA), and thus enable the molecular reconstruction of palaeofloras. Hitherto, ancient frozen soils have proved excellent in preserving DNA molecules, and have thus been the most commonly used source of plant aDNA. However, DNA from soil mainly represents taxa growing a few metres from the sampling point. Lakes have larger catchment areas and recent studies have suggested that plant aDNA from lake sediments is a more powerful tool for palaeofloristic reconstruction. Furthermore, lakes can be found globally in nearly all environments, and are therefore not limited to perennially frozen areas. Here, we review the latest approaches and methods for the study of plant aDNA from lake sediments and discuss the progress made up to the present. We argue that aDNA analyses add new and additional perspectives for the study of ancient plant populations and, in time, will provide higher taxonomic resolution and more precise estimation of abundance. Despite this, key questions and challenges remain for such plant aDNA studies. Finally, we provide guidelines on technical issues, including lake selection, and we suggest directions for future research on plant aDNA studies in lake sediments. Postprint

Accepted manuscript version. Published version available at https://doi.org/10.1016/j.ppees.2017.09.008. Accepted manuscript version, licensed CC BY-NC-ND 4.0. In the face of climate change, populations have two survival options − they can remain in situ and tolerate the new climatic conditions (“stay”), or they can move to track their climatic niches (“go”). For sessile and small-stature organisms like alpine plants, staying requires broad climatic tolerances, realized niche shifts due to changing biotic interactions, acclimation through plasticity, or rapid genetic adaptation. Going, in contrast, requires good dispersal and colonization capacities. Neither the magnitude of climate change experienced locally nor the capacities required for staying/going in response to climate change are constant across landscapes, and both aspects may be strongly affected by local microclimatic variation associated with topographic complexity. We combine ideas from population and community ecology to discuss the effects of topographic complexity in the landscape on the immediate “stay” or “go” opportunities of local populations and communities, and on the selective pressures that may have shaped the stay or go capacities of the species occupying contrasting landscapes. We demonstrate, using example landscapes of different topographical complexity, how species’ thermal niches could be distributed across these landscapes, and how these, in turn, may affect many population and community ecological processes that are related to adaptation or dispersal. Focusing on treeless alpine or Arctic landscapes, where temperature is expected to be a strong determinant, our theorethical framework leads to the hypothesis that populations and communities of topographically complex (rough and patchy) landscapes should be both more resistant and more resilient to climate change than those of topographically simple (flat and homogeneous) landscapes. Our theorethical framework further points to how meta-community dynamics such as mass effects in topographically complex landscapes and extinction lags in simple landscapes, may mask and delay the long-term outcomes of these landscape differences under rapidly changing climates.

Source at https://dx.doi.org/10.1002%2Fece3.3055 The impact of invasion on diversity varies widely and remains elusive. Despite the con- siderable attempts to understand mechanisms of biological invasion, it is largely un- known whether some communities’ characteristics promote biological invasion, or whether some inherent characteristics of invaders enable them to invade other com- munities. Our aims were to assess the impact of one of the massive plant invaders of Scandinavia on vascular plant species diversity, disentangle attributes of invasible and noninvasible communities, and evaluate the relationship between invasibility and ge- netic diversity of a dominant invader. We studied 56 pairs of Heracleum persicum Desf. ex Fisch.- invaded and noninvaded plots from 12 locations in northern Norway. There was lower native cover, evenness, taxonomic diversity, native biomass, and species richness in the invaded plots than in the noninvaded plots. The invaded plots had nearly two native species fewer than the noninvaded plots on average. Within the in- vaded plots, cover of H. persicum had a strong negative effect on the native cover, evenness, and native biomass, and a positive association with the height of the native plants. Plant communities containing only native species appeared more invasible than those that included exotic species, particularly H. persicum . Genetic diversity of H. per - sicum was positively correlated with invasibility but not with community diversity. The invasion of a plant community by H. persicum exerts consistent negative pressure on vascular plant diversity. The lack of positive correlation between impacts and genetic diversity of H. persicum indicates that even a small founder population may cause high impact. We highlight community stability or saturation as an important determinant of invasibility. While the invasion by H. persicum may decrease susceptibility of a plant community to further invasion, it severely reduces the abundance of native species and makes them more vulnerable to competitive exclusion.

International audience; 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.

Aim: Anticipated changes in the global ocean climate will affect the vulnerability of marine ecosystems to the negative effects of non-indigenous species (NIS). In the Arctic, there is a need to better characterize present and future marine biological introduction patterns and processes. We use a vector-based assessment to estimate changes in the vulnerability of a high-Arctic archipelago to marine NIS introduction and establishment. Location: Global, with a case study of Svalbard, Norway. Methods: We base our assessment on the level of connectedness to global NIS pools through the regional shipping network and predicted changes in ocean climates. Environmental match of ports connected to Svalbard was evaluated under present and future environmental conditions (2050 and 2100 predicted under the RCP8.5 emissions scenario). Risk of NIS introduction was then estimated based on the potential for known NIS to be transported (in ballast water or as biofouling), environmental match, and a qualitative estimate of propagule pressure. Results: We show that Svalbard will become increasingly vulnerable to marine NIS introduction and establishment. Over the coming century, sea surface warming at high latitudes is estimated to increase the level of environmental match to nearly one-third of ports previously visited by vessels travelling to Svalbard in 2011 (n = 136). The shipping network will then likely connect Svalbard to a much greater pool of known NIS, under conditions more favourable for their establishment. Research and fishing vessels were estimated to pose the highest risk of NIS introduction through biofouling, while ballast water discharge is estimated to pose an increased risk by the end of the century. Main conclusions: In the absence of focused preventative management, the risk of NIS introduction and establishment in Svalbard, and the wider Arctic, will increase over coming decades, prompting a need to respond in policy and action.

Arctic plant species are expected to lose range due to climate change. One approach to preserve the genetic and species diversity for the future is to store propagules in seed vaults. However, germinability of seeds is assumed to be low for Arctic species. We evaluated ex situ storage potential of 113 of the 161 native angiosperms of Svalbard by studying seed ripening and germination. Seeds or bulbils were collected, and germinability was tested after one winter of storage in the Svalbard Global Seed Vault. Twenty-six of the species did not produce ripe propagules, 8 produced bulbils, and 79 produced seeds. Bulbils sprouted to high percentages. Seeds of 10 species did not germinate, 22 had low germination (\20 %), 34 had germination of 21-70 %, and 13 had high germination percentages ((70 %). More than 70 % of the species belonging to Asteraceae, Brassicaceae, Caryophyllaceae, Juncaceae, Rosaceae, and Saxifragaceae germinated. Cold tolerant, common species had higher germination per- centages than relatively thermophilous, rare species. Ger- mination percentages were six times higher than observed in 1969 (n = 51) and 0.7 times that observed in 2008 (n = 22), indicating that recent climate warming improves germination in the Arctic. While in situ conservation is of vital importance, ex situ conservation in seed banks is a potential complementary conservation strategy for the majority of Arctic vascular plant species. For species that did not germinate, other methods for ex situ conservation should be sought, for example, growing in botanical gardens.

Polar bears (Ursus maritimus) are heavily dependent on marine prey, in particular ice-associated seals, which they hunt on landfast sea ice or free-floating pack ice. Dramatic current (and predicted) losses of sea ice habitat make it increasingly important to gain more knowledge of the relative use by bears of all types of prey from the marine food web as well as from terrestrial sources. This study uses frequency of occurrence of food items in 119 polar bear scats sampled on the sea ice as well as on shore in coastal areas in the Svalbard Archipelago, mainly in spring, between 2003 and 2010 to explore the diet of bears in the region. Ringed seals (Pusa hispida) occurred in 62.2 % (CI 52.8–70.9 %) of the scat samples examined. Various terrestrial plants (32.8 %, CI 24.4–42.0 %) and marine algae (21.8 %, CI 14.8–30.4 %) also occurred frequently in the scats; the significance of this high occurrence of plants and algae is not clear. Bearded seals (Erignathus barbatus) and various bird species constituted only minor components of the diet, while Svalbard reindeer (Rangifer tarandus platyrhynchus) occurred in 9.2 % (CI 4.7–15.9 %) of the scats, indicating that this species may play a more important role than previously reported. The novel combination of genetic analyses of material in the fecal samples along with detailed exploration of the physical–structural properties of prey hairs and plant parts provided a much fuller picture of the diet of polar bears than would have been possible from observational studies of polar bear predation behavior alone. This approach may provide an important tool for monitoring the responses of polar bears to ongoing ecosystem changes that will result from continued warming in the Arctic.

Source: doi: 10.1088/1748-9326/11/5/055005 Satellite-derived remote-sensing products are providing a modern circumpolar perspective of Arctic vegetation and its changes, but this new view is dependent on a long heritage of ground-based observations in the Arctic. Several products of the Conservation of Arctic Flora and Fauna are key to our current understanding.Wereview aspects of the PanArctic Flora, the Circumpolar Arctic Vegetation Map, the Arctic Biodiversity Assessment, and the Arctic Vegetation Archive (AVA) as they relate to efforts to describe and map the vegetation, plant biomass, and biodiversity of the Arctic at circumpolar, regional, landscape and plot scales. Cornerstones for all these tools are ground-based plant-species and plant-community surveys. TheAVA is in progress and will store plot-based vegetation observations in a public-accessible database for vegetation classification, modeling, diversity studies, and other applications.Wepresent the current status of the Alaska Arctic Vegetation Archive (AVA-AK), as a regional example for the panarctic archive, and with a roadmap for a coordinated international approach to survey, archive and classify Arctic vegetation.Wenote the need for more consistent standards of plot-based observations, and make several recommendations to improve the linkage between plot-based observations biodiversity studies and satellite-based observations of Arctic vegetation.

Aim High intra specific genetic diversity is necessary for species adaptation to novel environments under climate change but species tracking suitable conditions are losing alleles through successive founder events during range shift. Here we investigated the relationship between range shift since the Last Glacial Maximum (LGM) and extant population genetic diversity across multiple plant species to understand variability in species responses. Location The circumpolar Arctic and northern temperate alpine ranges. Methods We estimated the climatic niches of 30 cold adapted plant species using range maps coupled with species distribution models and hindcasted species suitable areas to reconstructions of the mid Holocene and LGM climates. We computed the species specific migration distances from the species glacial refugia to their current distribution and correlated distances to extant genetic diversity in 1295 populations. Differential responses among species were related to life history traits. Results We found a negative association between inferred migration distances from refugia and genetic diversities in 25 species but only 11 had statistically significant negative slopes. The relationships between inferred distance and population genetic diversity were steeper for insect pollinated species than wind pollinated species but the difference among pollination system was marginally independent from phylogenetic autocorrelation. Main conclusion The relationships between inferred migration distances and genetic diversities in 11 species independent from current isolation indicate that past range shifts were associated with a genetic bottleneck effect with an average of 21 loss of genetic diversity per 1000 km-1. In contrast the absence of relationship in many species also indicates that the response is species specific and may be modulated by plant pollination strategies or result from more complex historical contingencies than those modelled here.

High amounts of driftwood sail across the oceans and provide habitat for organisms tolerating the rough and saline environment. Fungi have adapted to the extremely cold and saline conditions which driftwood faces in the high north. For the first time, we applied high-throughput sequencing to fungi residing in driftwood to reveal their taxonomic richness, community composition, and ecology in the North Atlantic. Using pyrosequencing of ITS2 amplicons obtained from 49 marine logs, we found 807 fungal operational taxonomic units (OTUs) based on clustering at 97 % sequence similarity cut-off level. The phylum Ascomycota comprised 74 % of the OTUs and 20 % belonged to Basidiomycota. The richness of basidiomycetes decreased with prolonged submersion in the sea, supporting the general view of ascomycetes being more extremotolerant. However, more than one fourth of the fungal OTUs remained unassigned to any fungal class, emphasising the need for better DNA reference data from the marine habitat. Different fungal communities were detected in coniferous and deciduous logs. Our results highlight that driftwood hosts a considerably higher fungal diversity than currently known. The driftwood fungal community is not a terrestrial relic but a speciose assemblage of fungi adapted to the stressful marine environment and different kinds of wooden substrates found in it.

Published version. Source at http://dx.doi.org/10.1098/rsbl.2016.0264 Sea ice has been suggested to be an important factor for dispersal of vascular plants in the Arctic. To assess its role for postglacial colonization in the North Atlantic region, we compiled data on the first Late Glacial to Holocene occurrence of vascular plant species in East Greenland, Iceland, the Faroe Islands and Svalbard. For each record, we reconstructed likely past dispersal events using data on species distributions and genetics. We compared these data to sea-ice reconstructions to evaluate the potential role of sea ice in these past colonization events and finally evaluated these results using a compilation of driftwood records as an independent source of evidence that sea ice can disperse biological material. Our results show that sea ice was, in general, more prevalent along the most likely dispersal routes at times of assumed first colonization than along other possible routes. Also, driftwood is frequently dispersed in regions that have sea ice today. Thus, sea ice may act as an important dispersal agent. Melting sea ice may hamper future dispersal of Arctic plants and thereby cause more genetic differentiation. It may also limit the northwards expansion of competing boreal species, and hence favour the persistence of Arctic species.

PREMISE OF THE STUDY Climate change forces many species to migrate. Empirical small-scale data on migration and colonization in the Arctic are scarce. Retreating glaciers provide new territory for cold-adapted plant species, but the genetic consequences depend on dispersal distances and frequencies. We estimated local, regional, and long-distance dispersal frequencies, as well as their effect on levels of genetic diversity, in diploid and tetraploid individuals of Saxifraga oppositifolia. METHODS Samples were collected in four aged moraines in each of three glacier forelands, in surrounding areas and reference populations in the Arctic archipelago Svalbard. These samples were analyzed for neutral amplified fragment length polymorphisms (AFLPs, n = 707) and ploidy levels (n = 30). KEY RESULTS Genetic clustering and ploidy analyses revealed two distinct genetic groups representing diploids and tetraploids, with few intermediate triploids. The groups were intermixed in most sampled populations. No differences in genetic diversity were found between tetraploids and diploids, or between established and glacier foreland populations. Seeds were dispersed over local, regional, and long distances, with the highest proportions of seeds originating from close sources. A minimum of 4-15 founding individuals from several source populations had initially established in each glacier foreland. CONCLUSIONS Our data suggest that S. oppositifolia can rapidly colonize new deglaciated areas without losing genetic diversity. Thus, glacier forelands can be alternative habitats for cold-adapted vascular plants tracking their climatic niche. Our data show no difference in colonization success between diploid and tetraploid individuals.

Aim Previous research on how climatic niches vary across species ranges has focused on a limited number of species, mostly invasive, and has not, to date, been very conclusive. Here we assess the degree of niche conservatism between distant populations of native alpine plant species that have been separated for thousands of years.LocationEuropean Alps and Fennoscandia. Methods Of the studied pool of 888 terrestrial vascular plant species occurring in both the Alps and Fennoscandia, we used two complementary approaches to test and quantify climatic-niche shifts for 31 species having strictly disjunct populations and 358 species having either a contiguous or a patchy distribution with distant populations. First, we used species distribution modelling to test for a region effect on each species' climatic niche. Second, we quantified niche overlap and shifts in niche width (i.e. ecological amplitude) and position (i.e. ecological optimum) within a bi-dimensional climatic space. Results Only one species (3%) of the 31 species with strictly disjunct populations and 58 species (16%) of the 358 species with distant populations showed a region effect on their climatic niche. Niche overlap was higher for species with strictly disjunct populations than for species with distant populations and highest for arctic–alpine species. Climatic niches were, on average, wider and located towards warmer and wetter conditions in the Alps. Main conclusion Climatic niches seem to be generally conserved between populations that are separated between the Alps and Fennoscandia and have probably been so for 10,000–15,000 years. Therefore, the basic assumption of species distribution models that a species' climatic niche is constant in space and time – at least on time scales 104 years or less – seems to be largely valid for arctic–alpine plants.

Expanding visitation to Polar regions combined with climate warming increases the potential for alien species introduction and establishment. We quantified vascular plant propagule pressure associated with different groups of travelers to the high-Arctic archipelago of Svalbard, and evaluated the potential of introduced seeds to germinate under the most favorable average Svalbard soil temperature (10°C). We sampled the footwear of 259 travelers arriving by air to Svalbard during the summer of 2008, recording 1,019 seeds: a mean of 3.9 (±0.8) seeds per traveler. Assuming the seed influx is representative for the whole year, we estimate a yearly seed load of around 270,000 by this vector alone. Seeds of 53 species were identified from 17 families, with Poaceae having both highest diversity and number of seeds. Eight of the families identified are among those most invasive worldwide, while the majority of the species identified were non-native to Svalbard. The number of seeds was highest on footwear that had been used in forested and alpine areas in the 3 months prior to traveling to Svalbard, and increased with the amount of soil affixed to footwear. In total, 26% of the collected seeds germinated under simulated Svalbard conditions. Our results demonstrate high propagule transport through aviation to highly visited cold-climate regions and isolated islands is occurring. Alien species establishment is expected to increase with climate change, particularly in high latitude regions, making the need for regional management considerations a priority.

Sexual reproduction is crucial for plant populations to track and adapt to climate change, but it is uncertain to what degree arctic vascular plants reproduce by seed. Several studies on arctic species show low germination. To re-examine seed germination and evaluate factors limiting sexual reproduction, seeds of 6–22 arctic species were germinated in five different, increasingly more realistic, conditions. Thirteen out of 15 species that were tested in an earlier study in Svalbard, Norway, germinated better in our study. Compared with perceived optimal conditions in a growth chamber, average germination per species was 6%–52% lower in five out of six species germinating at a colder temperature in soil, 36%–64% lower when germinating outdoors in soil, 49%–91% lower when germinating in a moss covered moraine, and 55%–91% lower when germinating in open soil on a moraine. Germination outdoors was below 5% in 10 out of 13 species and not correlated to germination in perceived optimal conditions. The high germination compared with earlier studies suggests that climate warming has already increased seed viability. However, caution should be taken when evaluating species-recruitment potential based on laboratory studies, as germination in the field was limited by species-specific responses to low temperatures, moisture, predation, and safe-site availability.

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 noncoding 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.

Aim The oceanic Saxifraga rivularis L. presents one of the most extreme disjunctions known in the arctic flora: it has a small amphi-Beringian range and a larger amphi-Atlantic one. It was recently suggested to have had a single allopolyploid origin in Beringia at least one glacial cycle ago, followed by gradual expansion in a more humid period and differentiation into two allopatric subspecies (the Atlantic ssp. rivularis and the Beringian ssp. arctolitoralis). Here we explore the history of its extreme disjunction. Location The amphi-Beringian and northern amphi-Atlantic regions. Methods We obtained amplified fragment length polymorphisms (AFLPs) and chloroplast DNA sequences from 36 populations (287 individuals) and 13 populations (15 individuals), respectively. The data were analysed using principal coordinates analyses, Bayesian clustering methods, and analyses of molecular variance. Results Two distinctly divergent AFLP groups were observed, corresponding to the two described subspecies, but, surprisingly, four of the West Atlantic populations belonged to the supposedly Beringian endemic ssp. arctolitoralis. This was confirmed by re-examination of their morphological characteristics. The overall AFLP diversity in the species was low (26.4% polymorphic markers), and there was no variation in the five investigated chloroplast DNA (cpDNA) regions. There was little geographic structuring of the AFLP diversity within each subspecies, even across the extreme disjunction in ssp. arctolitoralis, across the Bering Sea, and across the Atlantic Ocean, except that most plants from the arctic Svalbard archipelago formed a separate genetic group with relatively high diversity. Main conclusions The extreme disjunction in S. rivularis has evidently formed at least twice. The first expansion from Beringia was followed by allopatric differentiation into one Beringian and one Atlantic subspecies, which are distinctly divergent at AFLP loci but still harbour identical cpDNA haplotypes, suggesting that the expansion was quite recent but before the last glaciation. The next expansion from Beringia probably occurred by means of several long-distance dispersals in the current interglacial, resulting in the colonization of the western Atlantic region by ssp. arctolitoralis. The poor geographic structuring within each subspecies suggests frequent long-distance dispersals from two main Weichselian refugia, one Beringian and one western-central European, but it is possible that the genetic group in Svalbard originates from an additional refugium.

A major drawback of Amplified Fragment Length Polymorphisms (AFLP) as genetic makers for phylogeographic studies is their lack of a temporal dimension. In a recent publication in Molecular Ecology, Kropf et al. (2009) proposed a molecular clock for AFLP. In this comment we evaluate the proposed approach both theoretically and empirically. A linear increase with time is a prerequisite to use a genetic distance as molecular clock. Testing the relationship between genetic distance and time in the data of Kropf et al. (2009) for linearity revealed that the relationship was in fact not linear for their pooled data, as well as for one of the three species analyzed. Also, the relationship was not linear in two new species, where divergence times could be inferred from macrofossils. When applying the proposed molecular clock to data from eight species, dates obtained were plausible in some cases, but very improbable in others. The suggested genetic distance was also influenced by intrapopulation genetic diversity, leading to a potential bias. In the future, investigations of AFLP mutation rates combined with phylogeographic modelling may contribute to adding a time scale to the understanding of AFLP data.

Aim Climate change may cause loss of genetic diversity. Here we explore how a multidisciplinary approach can be used to infer effects of past climate change on species distribution and genetic diversity and also to predict loss of diversity due to future climate change. We use the arctic-alpine plant Salix herbacea L. as a model.Location Europe, Greenland and eastern North America.Methods We analysed 399 samples from 41 populations for amplified fragment length polymorphism (AFLP) to identify current patterns of genetic structure and diversity and likely historical dispersal routes. Macrofossil records were compiled to infer past distribution, and species distribution models were used to predict the Last Glacial Maximum (LGM) and future distribution of climatically suitable areas.Results We found strong genetic differentiation between the populations from Europe/East Greenland and those from Canada/West Greenland, indicating a split probably predating the LGM. Much less differentiation was observed among the four genetic groups identified in Europe, and diversity was high in the Scandinavian as well as in southern alpine populations. Continuous distribution in Central Europe during the last glaciation was inferred based on the fossil records and distribution modelling. A 46-57% reduction in suitable areas was predicted in 2080 compared to present. However, mainly southern alpine populations may go extinct, causing a loss of about 5% of the genetic diversity in the species.Main conclusions From a continuous range in Central Europe during the last glaciation, northward colonization probably occurred as a broad front maintaining diversity as the climate warmed. This explains why potential extinction of southern populations by 2080 will cause a comparatively low loss of the genetic diversity in S. herbacea. For other species with different glacial histories, however, the expected climate-change induced regional extinction may cause a more severe loss of genetic diversity. We conclude that our multidisciplinary approach may be a useful tool for assessing impact of climate change on loss of genetic diversity.

Background: Many alpine plants are rare in Scotland. Their persistence depends on their ability to withstand habitat fragmentation and loss due to changes in land use, increased grazing pressure, and climate change. Aims: We use a phylogeographic approach to address the origin and genetic diversity of Scottish populations, which is relevant for their future management and protection. Methods: We review phylogeographic studies that include Scottish populations of common and rare alpine plants, and present unpublished data for eleven more species. We use results from amplified fragment length polymorphism (AFLP) studies to address the genetic distinctiveness of the Scottish populations and their levels of genetic diversity. Results: A positive correlation was found between relative rarity of each species in Scotland and the level of genetic diversity in Scottish populations compared with the species' total. Scottish populations of all species analysed except Saxifraga cernua belong to larger European/Eurasi...

Aim During the last ice age large parts of the north boreal and subarctic zones were covered by ice, while the climate in ice-free regions of northern Asia was extremely cold and dry. The extensive peatlands of these zones with their characteristic vegetation developed at the beginning of the Holocene. We combine a phylogeographical approach with maps of pollen records to identify regions where Rubus chamaemorus, a plant of moist, peaty soils, was likely to grow during this period. Location Circumarctic/circumboreal. Methods Samples were collected from 45 locations throughout much of the range of R. chamaemorus and 398 plants were analysed with amplified fragment length polymorphism (AFLP). Estimates of diversity and differentiation, principal coordinates analysis and Bayesian clustering methods were used for the analysis of genetic data. Dated pollen records were retrieved from the European and the Global Pollen Databases. Results The plants from Sakhalin are highly divergent from the rest of the material and represent the previously described var. pseudochamaemorus. The main genetic division in R. chamaemorus sensu stricto is found in the Taymyr region in central Eurasia. Genetic diversity and the relative number of rare markers are highest in central Siberia and eastern Asia and decrease towards Europe and to a lesser extent eastwards through North America. Pollen dating back to the last ice age is found in central and eastern Siberia and Alaska. The maximum observed clone size is about 250 m, and more than one clone is found in nearly every local population. Main conclusions The genetic data are consistent with the pollen records and indicate that R. chamaemorus was growing in several areas of northern central Siberia and Beringia during the last glaciation. This finding suggests that sufficient humidity for this and other species of peaty soils was present locally in different parts of the generally dry ice-free areas of northern Asia, as had been previously documented for Beringia. The AFLP data show that var. pseudochamaemorus, which is also morphologically quite divergent, clearly represents a distinct genetic entity.

To fully understand the contemporary genetic structure of plants, both nuclear and plastid markers are needed. Three chloroplast DNA (cpDNA) lineages, which probably diverged before the major Pleistocene glaciations, have been identified in the circumpolar/circumboreal Vaccinium uliginosum. Here we investigate its nuclear DNA variation using nuclear ribosomal internal transcribed spacer (ITS) sequences, DNA ploidy level measurements and amplified fragment length polymorphisms (AFLPs). We also extend the cpDNA dataset. Two ITS lineages, corresponding to diploids and tetraploids, respectively, were identified. However, both main sequence types apparently occurred in most individual plants but showed ploidy-biased homogenization and possibly reflect paralogy predating the origin of V. uliginosum. The ploidy levels were largely consistent with the cpDNA lineages, suggesting that the initial cpDNA divergence followed early polyploidizations. Five main AFLP groups were identified, consistent with recent glacial refugia in Beringia, western Siberia, the southern European mountains and areas south/east of the Scandinavian and Laurentide ice sheets. Except from the southern European mountains, there has been extensive expansion from all refugia, resulting in several contact zones. Surprisingly, the presumably older ploidy and cpDNA patterns were partly inconsistent with the main AFLP groups and more consistent with AFLP subgroups. A likely major driver causing the inconsistencies is recent nuclear gene flow via unreduced pollen from diploids to tetraploids. This may prevent cytoplasmic introgression and result in overlayed patterns formed by processes dominating at different time scales. The data also suggest more recent polyploidizations, as well as several chloroplast capture events, further complicating this scenario. This study highlights the importance of combining different marker systems to unravel intraspecific histories.

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., 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.

Several decades of research on invasive marine species have yielded a broad understanding of the nature of species invasion mechanisms and associated threats globally. However, this is not true of the Arctic, a region where ongoing climatic changes may promote species invasion. Here, we evaluated risks associated with non-indigenous propagule loads discharged with ships' ballast water to the high-Arctic archipelago, Svalbard, as a case study for the wider Arctic. We sampled and identified transferred propagules using traditional and DNA barcoding techniques. We then assessed the suitability of the Svalbard coast for non-indigenous species under contemporary and future climate scenarios using ecophysiological models based on critical temperature and salinity reproductive thresholds. Ships discharging ballast water in Svalbard carried high densities of zooplankton (mean 1522335 SE individuals m(-3)), predominately comprised of indigenous species. Ballast water exchange did not prevent non-indigenous species introduction. Non-indigenous coastal species were present in all except one of 16 ballast water samples (mean 14467 SE individuals m(-3)), despite five of the eight ships exchanging ballast water en route. Of a total of 73 taxa, 36 species including 23 non-indigenous species were identified. Of those 23, sufficient data permitted evaluation of the current and future colonization potential for eight widely known invaders. With the exception of one of these species, modelled suitability indicated that the coast of Svalbard is unsuitable presently; under the 2100 Representative Concentration Pathway (RCP) 85 climate scenario, however, modelled suitability will favour colonization for six species.Synthesis and applications. We show that current ballast water management practices do not prevent non-indigenous species from being transferred to the Arctic. Consequences of these shortcomings will be shipping-route dependent, but will likely magnify over time: our models indicate future conditions will favour the colonization of non-indigenous species Arctic-wide. Invasion threats will be greatest where shipping transfers organisms across biogeographic realms, and for these shipping routes ballast water treatment technologies may be required to prevent impacts. Our results also highlight critical gaps in our understanding of ballast water management efficacy and prioritization. Thereby, our study provides an agenda for research and policy development. We show that current ballast water management practices do not prevent non-indigenous species from being transferred to the Arctic. Consequences of these shortcomings will be shipping-route dependent, but will likely magnify over time: our models indicate future conditions will favour the colonization of non-indigenous species Arctic-wide. Invasion threats will be greatest where shipping transfers organisms across biogeographic realms, and for these shipping routes ballast water treatment technologies may be required to prevent impacts. Our results also highlight critical gaps in our understanding of ballast water management efficacy and prioritization. Thereby, our study provides an agenda for research and policy development.

This is the submitted manuscript version. Published version can be found at http://doi.org/10.1111/mec.13411. Sparse, incomplete and inappropriate historical records of invasive species often hamper invasive species management interventions. Population genetic analyses of invaders might provide a suitable context for the identification of their source populations and possible introduction routes. Here, we describe the population genetics of Heracleum persicum Desf. ex Fisch and trace its route of introduction into Europe. Microsatellite markers revealed a significantly higher genetic diversity of H. persicum in its native range, and the loss of diversity in the introduced range may be attributed to a recent genetic bottleneck. Bayesian cluster analysis on regional levels identified three and two genetic clusters in the native and the introduced ranges, respectively. A global structure analysis revealed two worldwide distinct genetic groups: one primarily in Iran and Denmark, the other primarily in Norway. There were also varying degrees of admixture in England, Sweden, Finland and Latvia. Approximate Bayesian computation indicated two independent introductions of H. persicum from Iran to Europe: the first one in Denmark and the second one in England. Finland was subsequently colonized by English populations. In contrast to the contemporary hypothesis of English origin of Norwegian populations, we found Finland to be a more likely source for Norwegian populations, a scenario supported by higher estimated histor-ical migration from Finland to Norway. Genetic diversity per se is not a primary determinant of invasiveness in H. persicum. Our results indicate that, due to either pre-adaptations or rapid local adaptations, introduced populations may have acqu-ired invasiveness after subsequent introductions, once a suitable environment was encountered.

Reconstructing past vegetation and species diversity from arctic lake sediments can be challenging because of low pollen and plant macrofossil concentrations. Information may be enhanced by metabarcoding of sedimentary ancient DNA ( sedaDNA). We developed a Holocene record from Lake Skartjørna, Svalbard, using sedaDNA, plant macrofossils and sediment properties, and compared it with published records. All but two genera of vascular plants identified as macrofossils in this or a previous study were identified with sedaDNA. Six additional vascular taxa were found, plus two algal and 12 bryophyte taxa, by sedaDNA analysis, which also detected more species per sample than macrofossil analysis. A shift from Salix polaris-dominated vegetation, with Koenigia islandica, Ranunculaceae and the relatively thermophilic species Arabis alpina and Betula, to Dryas octopetala-dominated vegetation ~6600–5500 cal. BP suggests a transition from moist conditions 1–2°C warmer than today to colder/drier conditions. This coincides with a decrease in runoff, inferred from core lithology, and an independent record of declining lacustrine productivity. This mid-Holocene change in terrestrial vegetation is broadly coincident with changes in records from marine sediments off the west coast of Svalbard. Over the Holocene sedaDNA records little floristic change, and it clearly shows species persisted near the lake during time intervals when they are not detected as macrofossils. The flora has shown resilience in the presence of a changing climate, and, if future warming is limited to 2°C or less, we might expect only minor floristic changes in this region. However, the Holocene record provides no analogues for greater warming.

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 self-fertilizing 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. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82, 521–536.

An aim for conservation in Norway is preserving the Svalbard archipelago as one of the least disturbed areas in the Arctic. Information on local distribution, population sizes and ecology is summarized for 20 thermophilous vascular plant species. The need for conservation of northern, marginal populations in Svalbard is reviewed, using World Conservation Union categories and criteria at a regional scale. Thirteen species reach their northernmost distribution in Svalbard, the remaining seven in the western Arctic. Nine species have 1-8 populations in Svalbard and are assigned to Red List categories endangered or critically endangered: Campanula rotundifolia, Euphrasia frigida, Juncus castaneus, Kobresia simpliciuscula, Rubus chamaemorus, Alchemilla glomerulans, Ranunculus wilanderi, Salix lanata and Vaccinium uliginosum, the last four species needing immediate protective measures. Five species are classified as vulnerable: Betula nana, Carex marina ssp. pseudolagopina, Luzula wahlenbergii, Ranunculus arcticus and Ranunculus pallasii. Six species are considered at lower risk: Calamagrostis stricta, Empetrum nigrum ssp. hermaphroditum, Hippuris vulgaris (only occurring on Bjørnøya), Juncus triglumis, Ranunculus lapponicus and Rhodiola rosea. The warmer Inner Arctic Fjord Zone of Spitsbergen supports most of the 20 target species and is of particular importance for conservation. Endangered or vulnerable species were found in a variety of edaphic conditions; thus, several kinds of habitats need protection.

The rare thermophilous species in the arctic archipelago of Svalbard are probably relicts from previous warmer periods and may be unable to reproduce sexually under the present climatic conditions. Germination of seeds, seed banks, and vegetative sprouts were studied in one Betula nana L., one Vaccinium uliginosum L., and two Campanula rotundifolia L. habitats in Svalbard. For comparison, one islet and three alpine habitats in northern Norway were studied. In each habitat, 50 soil cores (9.6 cm2) were collected and placed in a phytotron for germination tests. In Svalbard, no germinable seeds or seed bank of the three species were observed, whereas vegetative sprout densities were 83, 1060, and 21–1060/m2 for B. nana, V. uliginosum, and C. rotundifolia, respectively. In northern Norway, germination percentages of the collected seeds were 59, 10–27, and 15–40%, seed bank densities of seedlings 21–2765, 21–187, and 21–374/m2, and vegetative sprouts densities 94–206, 56–674, and 94–711/m2 for B. nana, V. uliginosum, and C. rotundifolia, respectively. Less thermophilous plant species produced germinable seeds in Svalbard, as 1247–5405 seedlings/m2 of 26 species germinated from the soil cores. In comparison, 5322–9626 seedlings/m2 of 43 species germinated in soil cores from the alpine habitats and 3534 seedlings/m2 of 26 species from the islet habitat. The results suggest that the most thermophilous species in Svalbard rarely produce germinable seeds under the present climatic conditions.Key words: arctic–alpine, Betula nana, Campanula rotundifolia, seed bank, sexual reproduction, thermophily, Vaccinium uliginosum.

Plant species defined as strongly thermophilous in Svalbard are rare and may be threatened by extinction due to suppressed sexual reproduction, small population sizes, genetic depauperation, and human activity. We examined isozyme diversity in three of these species-Betula nana, Vaccinium uliginosum, and Campanula rotundifolia-in Svalbard. Twenty-five to 60 ramets from each of two to three (sub)populations of each species were analyzed for eight enzyme systems. In B. nana, genetic diversity was low within sampling plots (D: 0.00-0.49) and intermediate within populations (mean D = 0.70, mean Hep = 0.119). Genetic diversity at the species level (Hes) was 0.141, and the differentiation among populations (FsT = 0.193) was higher than the average for other species with similar life traits. In V. uliginosum, no variation was observed within plots, and genetic diversity within populations was low (mean D = 0.29, mean Hep = 0.082). Genetic diversity at the species level was also low (Hes = 0.152), and the differentiation among populations (FsT = 0.493) was much higher than in most other species studied. The only Svalbard population of C. rotundifolia has 2n = 34 and is traditionally considered as diploid, but the observed enzyme banding patterns suggest polyploidy. In this species, genotypic diversity was low within plots (D: 0.00-0.33), intermediate within subpopulations (mean D = 0.74), and all multilocus phenotypes were local. The present populations of all three species in Svalbard are probably fragmented relics of larger populations in the postglacial warm period. On the basis of the observed distribution of genetic diversity and population sizes, we propose to conserve all Svalbard populations of V. uliginosum and C. rotundifolia, and at least one population of B. nana.

This is the peer reviewed version of the following article: Eidesen, P.B., Ehrich, D., Bakkestuen, V., Alsos, I.G., Gilg, O., Taberlet, P. & Brochmann, C. (2013). Genetic roadmap of the Arctic: plant dispersal highways, traffic barriers and capitals of diversity. New Phytologist, 200(3), 898-910. https://doi.org/10.1111/nph.12412, which has been published in final form at https://doi.org/10.1111/nph.12412. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. We provide the first comparative multispecies analysis of spatial genetic structure and diversity in the circumpolar Arctic using a common strategy for sampling and genetic analyses. We aimed to identify and explain potential general patterns of genetic discontinuity/connectivity and diversity, and to compare our findings with previously published hypotheses. We collected and analyzed 7707 samples of 17 widespread arctic–alpine plant species for amplified fragment length polymorphisms (AFLPs). Genetic structure, diversity and distinctiveness were analyzed for each species, and extrapolated to cover the geographic range of each species. The resulting maps were overlaid to produce metamaps. The Arctic and Atlantic Oceans, the Greenlandic ice cap, the Urals, and lowland areas between southern mountain ranges and the Arctic were the strongest barriers against gene flow. Diversity was highest in Beringia and gradually decreased into formerly glaciated areas. The highest degrees of distinctiveness were observed in Siberia. We conclude that large‐scale general patterns exist in the Arctic, shaped by the Pleistocene glaciations combined with long‐standing physical barriers against gene flow. Beringia served as both refugium and source for interglacial (re)colonization, whereas areas further west in Siberia served as refugia, but less as sources for (re)clonization.

Saxifraga oppositifolia L. is a common circumpolar plant species that displays considerable morphological and genetic variation throughout its range. It is mainly diploid, but tetraploids are reported from several regions. The growth form varies from prostate to cushion-shaped, and the plant thrives in wet snow beds as well as on dry ridges. This variation has triggered the curiosity of many researchers, but as yet, no one has explained the observed morphological variation using ecological and/or genetic factors. However, the ploidy level has rarely been taken into account. This is the first study that demonstrates a significant correlation between ploidy level, ecology and growth form in S. oppositifolia . We successfully analysed 193 individuals of S. oppositifolia from 15 locations in Svalbard to investigate possible relationships among growth forms (prostrate, intermediate and cushion), ecological factors (vegetation and soil characteristics) and ploidy level. Results from flow cytometry reported 106 diploids, eight triploids and 79 tetraploids. Tetraploids almost exclusively showed prostrate growth, while the diploids displayed all three growth forms, evidence that growth form is at least partly genetically determined. Our analyses of environmental and vegetation data in relation to ploidy level indicated overlapping niches, but the tetraploids showed a narrower niche, and one shifted towards more benign habitats characterized by higher pH, higher soil temperatures and higher cover of vascular plants. The latter may suggest that tetraploids are slightly better competitors, but less hardy. Thus, autopolyploidy in S. oppositifolia has expanded the ecological amplitude of this species complex. Keywords: Autopolyploidy; flow cytometry; morphotypes; habitat segregation; purple saxifrage (Published: 6 June 2013) To access the supplementary material for this article, please see Supplementary files in the column to the right (under Article Tools) Citation: Polar Research 2013, 32 , 20071, http://dx.doi.org/10.3402/polar.v32i0.20071

Recent studies from mountainous areas of small spatial extent (2500 km(2) ) suggest that fine-grained thermal variability over tens or hundreds of metres exceeds much of the climate warming expected for the coming decades. Such variability in temperature provides buffering to mitigate climate-change impacts. Is this local spatial buffering restricted to topographically complex terrains? To answer this, we here study fine-grained thermal variability across a 2500-km wide latitudinal gradient in Northern Europe encompassing a large array of topographic complexities. We first combined plant community data, Ellenberg temperature indicator values, locally measured temperatures (LmT) and globally interpolated temperatures (GiT) in a modelling framework to infer biologically relevant temperature conditions from plant assemblages within1000-m(2) units (community-inferred temperatures: CiT). We then assessed: (1) CiT range (thermal variability) within 1-km(2) units; (2) the relationship between CiT range and topographically and geographically derived predictors at 1-km resolution; and (3) whether spatial turnover in CiT is greater than spatial turnover in GiT within 100-km(2) units. Ellenberg temperature indicator values in combination with plant assemblages explained 46-72% of variation in LmT and 92-96% of variation in GiT during the growing season (June, July, August). Growing-season CiT range within 1-km(2) units peaked at 60-65°N and increased with terrain roughness, averaging 1.97 °C (SD = 0.84 °C) and 2.68 °C (SD = 1.26 °C) within the flattest and roughest units respectively. Complex interactions between topography-related variables and latitude explained 35% of variation in growing-season CiT range when accounting for sampling effort and residual spatial autocorrelation. Spatial turnover in growing-season CiT within 100-km(2) units was, on average, 1.8 times greater (0.32 °C km(-1) ) than spatial turnover in growing-season GiT (0.18 °C km(-1) ). We conclude that thermal variability within 1-km(2) units strongly increases local spatial buffering of future climate warming across Northern Europe, even in the flattest terrains.

The dwarf shrub Cassiope tetragona (Arctic bell-heather) is increasingly used for arctic climate reconstructions, the reliability of which depends on the existence of a linear climategrowth relationship. This relationship was examined over a high-arctic to sub-arctic temperature gradient and under multi-year artificial warming at a high-arctic site.Growth chronologies of annual shoot length, as well as total leaf length, number of leaves and average leaf length per year, were constructed for three sites. Cassiope tetragona was sampled near its cold tolerance limit at Ny-lesund, Svalbard, at its assumed climatic optimum in Endalen, Svalbard, and near its European southern limit at Abisko, Sweden. Together these sites represent the entire temperature gradient of this species. Leaf life span was also determined. Each growing season from 2004 to 2010, 17 open top chambers (OTCs) were placed near Ny-lesund, thus increasing the daily mean temperatures by 123C. At the end of the 2010 growing season, shoots were harvested from OTCs and control plots, and growth parameters were measured.All growth parameters, except average leaf length, exhibited a linear positive response (R-2 between 063 and 091) to mean July temperature over the temperature gradient. Average leaf life span was 14 years shorter in sub-arctic Sweden compared with arctic Svalbard. All growth parameters increased in response to the experimental warming; the leaf life span was, however, not significantly affected by OTC warming.The linear July temperaturegrowth relationships, as well as the 7 year effect of experimental warming, confirm that the growth parameters annual shoot length, total leaf length and number of leaves per year can reliably be used for monitoring and reconstructing temperature changes. Furthermore, reconstructing July temperature from these parameters is not hampered by divergence.