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Changes in the distribution of coastal macrophytes in Greenland, and elsewhere in the Arctic are difficult to quantify as the region remains challenging to access and monitor. Satellite imagery, in particular Sentinel-2 (S2), may enable large-scale monitoring of coastal areas in Greenland but its use is impacted by the optically complex environments and the scarcity of supporting data in the region. Additionally, the canopies of the dominant macrophyte species in Greenland do not extend to the sea surface, limiting the use of indices that exploit the reflection of near-infrared radiation by vegetation due to its absorption by seawater. Three hypotheses are tested: I) 10-m S2 imagery and commonly used detection methods can identify intertidal macrophytes that are exposed at low tide in an optically complex fjord system in Greenland impacted by marine and land terminating glaciers; II) detached and floating macrophytes accumulate in patches that are sufficiently large to be detected by 10-m S2 images; III) iceberg scour and/or turbid meltwater runoff shape the spatial distribution of intertidal macroalgae in fjord systems with marine-terminating glaciers. The NDVI produced the best results in optically complex fjord systems in Greenland. 12 km2 of exposed intertidal macrophytes were identified in the study area at low tide. Floating mats of macrophytes ranged in area from 400 m2 to 326,800 m2 and were most common at the mouth of the fjord. Icebergs and turbidity appear to play a role in structuring the distribution of intertidal macrophytes and the retreat of marine terminating glaciers could allow macrophytes cover to expand. The challenges and solutions presented here apply to most fjords in Greenland and, therefore, the methodology may be extended to produce a Greenland-wide estimate of intertidal macrophytes.

Biodiversity studies could strongly benefit from three-dimensional data on ecosystem structure derived from contemporary remote sensing technologies, such as light detection and ranging (lidar). Despite the increasing availability of such data at regional and national scales, the average ecologist has been limited in accessing them due to high requirements on computing power and remote sensing knowledge. We processed Denmark's publicly available national airborne laser scanning (ALS) data set acquired in 2014/15, together with the accompanying elevation model, to compute 70 rasterised descriptors of interest for ecological studies. With a grain size of 10 m, these data products provide a snapshot of high-resolution measures including vegetation height, structure and density, as well as topographic descriptors including elevation, aspect, slope and wetness across more than 40 000 km2 covering almost all of Denmark's terrestrial surface. The resulting data set is comparatively small (∼94 GB, compressed 16.8 GB), and the raster data can be readily integrated into analytical workflows in software familiar to many ecologists (GIS software, R, Python). Source code and documentation for the processing workflow are openly available via a code repository, allowing for transfer to other ALS data sets, as well as modification or re-calculation of future instances of Denmark's national ALS data set. We hope that our high-resolution ecological vegetation and terrain descriptors (EcoDes-DK15) will serve as an inspiration for the publication of further such data sets covering other countries and regions and that our rasterised data set will provide a baseline of the ecosystem structure for current and future studies of biodiversity, within Denmark and beyond. The full data set is available on Zenodo: https://doi.org/10.5281/zenodo.4756556 (Assmann et al., 2021); a 5 MB teaser subset is also available: https://doi.org/10.5281/zenodo.6035188 (Assmann et al., 2022a).

Rapid climate change has been driving changes in Arctic vegetation in recent decades, with increased shrub dominance in many tundra ecosystems. Dendroecological observations of tundra shrubs can provide insight into current and past growth and recruitment patterns, both key components for understanding and predicting ongoing and future Arctic shrub dynamics. However, generalizing these dynamics is challenging as they are highly scale-dependent and vary among sites, species, and individuals. Here, we provide a perspective on how some of these challenges can be overcome. Based on a targeted literature search of dendrochronological studies from 2005 to 2022, we highlight five research gaps that currently limit dendro-based studies from revealing cross-scale ecological insight into shrub dynamics across the Arctic biome. We further discuss the related re- search priorities, suggesting that future studies could consider: 1) increasing focus on intra- and interspecific var- iation, 2) including demographic responses other than radial growth, 3) incorporating drivers, in addition to warming, at different spatial and temporal scales, 4) implementing systematic and unbiased sampling ap- proaches, and 5) investigating the cellular mechanisms behind the observed responses. Focusing on these aspects in dendroecological studies could improve the value of the field for addressing cross-scale and plant community- framed ecological questions. We outline how this could be facilitated through the integration of community- based dendroecology and dendroanatomy with remote sensing approaches. Integrating new technologies and a more multidisciplinary approach in dendroecological research could provide key opportunities to close impor- tant knowledge gaps in our understanding of scale-dependencies, as well as intra- and inter-specific variation, in vegetation community dynamics across the Arctic tundra.

Climate warming is inducing widespread vegetation changes in Arctic tundra ecosystems, with the potential to alter carbon and nutrient dynamics between vegetation and soils. Yet, we lack a detailed understanding of how variation in vegetation and topography influences fine-scale temperatures ("microclimate") that mediate these dynamics, and at what resolution vegetation needs to be sampled to capture these effects. We monitored microclimate at 90 plots across a tundra landscape in western Greenland. Our stratified random study design covered gradients of topography and vegetation, while nested plots (0.8-100 m(2)) enabled comparison across different sampling resolutions. We used Bayesian mixed-effect models to quantify the direct influence of plot-level topography, moisture and vegetation on soil, near-surface and canopy-level temperatures (-6, 2, and 15 cm). During the growing season, colder soils were predicted by shrub cover (-0.24 degrees C per 10% increase), bryophyte cover (-0.35 degrees C per 10% increase), and vegetation height (-0.17 degrees C per 1 cm increase). The same three factors also predicted the magnitude of differences between soil and above-ground temperatures, indicating warmer soils at low cover/height, but colder soils under closed/taller canopies. These findings were consistent across plot sizes, suggesting that spatial predictions of microclimate may be possible at the operational scales of satellite products. During winter, snow cover (+0.75 degrees C per 10 snow-covered days) was the key predictor of soil microclimate. Topography and moisture explained little variation in the measured temperatures. Our results not only underline the close connection of vegetation and snow with microclimate in the Arctic tundra but also point to the need for more studies disentangling their complex interplay across tundra environments and seasons. Future shifts in vegetation cover and height will likely mediate the impact of atmospheric warming on the tundra soil environment, with potential implications for below-ground organisms and ecosystem functioning.

Aim Tundra ecosystems are highly vulnerable to climate change, and climate-growth responses of Arctic shrubs are variable and altered by microsite environmental conditions and biotic factors. With warming and drought during the growing season, insect-driven defoliation is expected to increase in frequency and severity with potential broad-scale impacts on tundra ecosystem functioning. Here we provide the first broad-scale reconstruction of spatio-temporal dynamics of past insect outbreaks by assessing their effects on shrub growth along a typical Greenlandic fjord climate gradient from the inland ice to the sea. Location Nuuk Fjord (64 degrees 30 ' N/51 degrees 23 ' W) and adjacent areas, West Greenland. Taxa Great brocade (Eurois occulta L.) and grey willow (Salix glauca L.). Methods We combined dendro-anatomical and remote sensing analyses. Time series of ring width (RW) and wood-anatomical traits were obtained from chronologies of >40 years established from 153 individuals of S. glauca collected at nine sites. We detected anomalies in satellite-based Normalized Difference Vegetation Index (NDVI) related to defoliation and reconstructed past changes in photosynthetic activity across the region. Results We identified outbreaks as distinctive years with reduced RW, cell-wall thickness and vessel size, without being directly related to climate but matching with years of parallel reduction in NDVI. The two subsequent years after the defoliation showed a significant increase in RW. The reconstructed spatio-temporal dynamics of these events indicate substantial regional variation in outbreak intensity linked to the climate variability across the fjord system. Main conclusions Our results highlight the ability of S. glauca to cope with severe insect defoliation by changing carbon investment and xylem conductivity leading to high resilience and rapid recovery after the disturbance. Our multiproxy approach allows us to pinpoint biotic drivers of narrow ring formation and to provide new broad-scale insight on the C-budget and vegetation productivity of shrub communities in a widespread arctic ecosystem.

Nitrogen availability in Arctic ecosystems is a key driver for biological activity, including plant, growth and thereby directly linked to the greening of the Arctic. Here, we model the redistribution of meltwater following spring snowmelt as well as the accumulation of meltwater and dissolved nitrate at landscape scale. By combining snow mapping with unmanned aerial systems, snow chemistry, and hydrological modelling, we argue that the majority of nitrate in the snowpack is flushed out of the landscape due to the limited storage capacity of meltwater in the early growing season frozen soil. We illustrate how landscape micro-topography is a crucial parameter to quantify storage capacity of meltwater at landscape scale and thereby the associated pool of soluble compounds such as nitrate. This pool will be available for plants and may be important for plant diversity and growth rates in the wettest part of the landscape. This study illustrates that the evenly distributed nitrate input during the Arctic winter may be redistributed during the initial snowmelt and lead to marked differences in biologically available nitrate at the onset of the growing season, but also that the majority of deposited nitrate in snow is lost from the terrestrial to the aquatic environment during snowmelt.

Insect defoliations are a major natural disturbance in high-latitude ecosystems and are expected to increase in frequency and severity due to current climatic change. Defoliations cause severe reductions in biomass and carbon investments that affect the functioning and productivity of tundra ecosystems. Here we combined dendro-anatomical analysis with chemical imaging to investigate the direct and lagged effects of insect outbreaks on carbon investment. We analysed the content of lignin vs. holocellulose, i.e. unspecified carbohydrates in xylem samples of Salix glauca L. collected at Iffiartarfik, Nuuk fjord, Greenland, featuring two outbreak events of the moth Eurois occulta L. Cross sections of the growth rings corresponding to both outbreaks ±3 years were analysed using confocal Raman imaging to identify possible chemical signatures related to insect defoliation on fibres, vessels, and ray parenchyma cells and to get insight into species-specific defence responses. Outbreak years with narrower rings and thinner fibre cell walls are accompanied by a change in the content of cell-wall polymers but not their underlying chemistry. Indeed, during the outbreaks the ratio between lignin and carbohydrates significantly increased in fibre but not vessel cell walls due to an increase in lignin content coupled with a reduced content of carbohydrates. Parenchyma cell walls and cell corners did not show any significant changes in the cell-wall biopolymer content. The selective adjustment of the cell-wall composition of fibres but not vessels under stressful conditions could be related to the plants priority to maintain an efficient hydraulic system rather than mechanical support. However, the higher lignin content of fibre cell walls formed during the outbreak events could increase mechanical stiffness to the thin walls by optimizing the available resources. Chemical analysis of xylem traits with Raman imaging is a promising approach to highlight hidden effects of defoliation otherwise overlooked with classical dendroecological methods.

The majority of variation in six traits critical to the growth, survival and reproduction of plant species is thought to be organised along just two dimensions, corresponding to strategies of plant size and resource acquisition. However, it is unknown whether global plant trait relationships extend to climatic extremes, and if these interspecific relationships are confounded by trait variation within species. We test whether trait relationships extend to the cold extremes of life on Earth using the largest database of tundra plant traits yet compiled. We show that tundra plants demonstrate remarkably similar resource economic traits, but not size traits, compared to global distributions, and exhibit the same two dimensions of trait variation. Three quarters of trait variation occurs among species, mirroring global estimates of interspecific trait variation. Plant trait relationships are thus generalizable to the edge of global trait-space, informing prediction of plant community change in a warming world., It is unclear whether plant trait relationships found at the global scale extend to climatic extremes. Here the authors analyse six major aboveground traits to show that known plant trait relationships extend to the tundra biomes and exhibit the same two dimensions of variation detected at the global scale.

Mapping changes in vegetation cover is essential for understanding the consequences of climate change on Arctic ecosystems. Classification of ultra-high spatial-resolution (UHR, < 1 cm) imagery can provide estimates of vegetation cover across space and time. The challenge of this approach is to assure comparability of classification across many images taken at different illumination conditions and locations. With warming, vegetation at higher elevation is expected to resemble current vegetation at lower elevation. To investigate the value of classification of UHR imagery for monitoring vegetation change, we collected visible and near-infrared images from 108 plots with hand-held cameras along an altitudinal gradient in Greenland and examined the classification accuracy of shrub cover on independent images (i.e. classification transferability). We implemented several models to examine if colour calibration improves transferability based on an in-image calibration target. The classifier was trained on different number of images to find the minimum training subset size. With a training set of ~ 20% of the images the overall accuracy levelled off at about 81% and 68% on the non-calibrated training and validation images, respectively. Colour calibration improved the accuracy on training images (1–4%) while it only improved the classifier transferability significantly for training sets < 20%. Linear calibration only based on the target’s grey series improved transferability most. Reasonable transferability of Arctic shrub cover classification can be obtained based only on spectral data and about 20% of all images. This is promising for vegetation monitoring through multi-image classification of UHR imagery acquired with hand-held cameras or Unmanned Aerial Systems.

The disproportionate warming in the Arctic and the resulting adverse ecosystem changes underline the importance of continued monitoring of these ecosystems. Land-cover classification maps of the Arctic regions are essential for monitoring and change detection purposes, as well as upscaling of various ecosystem processes. However, large-scale land cover maps of the Arctic regions are often too coarse to properly capture the heterogeneity of these landscapes. In this study, we bridge this gap through incorporating multi temporal Landsat-8 OLI data in a large-scale land cover classification, and subsequently produce a tundra classification map for the entire Greenland. An algorithm is developed that allows for the extraction of vegetation phenology from single-year time series of 4169 OLI scenes at 30 m resolution despite the low revisit frequency of the satellite and persistent cloud cover. The phenological metrics, satellite-derived wetness, and terrain information are then used to separate land surface classes using a random forest classifier. The optimal algorithm parameters and input layers are identified, ultimately yielding a cross-validation accuracy of 89.25% across the studied area. Finally, we have conducted a comprehensive analysis on the resulting land-cover map and for the first time presented the geographical distribution, latitudinal gradients, and climate linkages of the various tundra vegetation classes across the ice-free part of Greenland. With a resolution of 30 m and Greenland-wide spatial coverage, the produced land-cover map can support various applications at scales ranging from the landscape to regional level.

Altres ajuts europeus: P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586)P.A.W. was additionally supported by the European Union Fourth Environment and Climate Framework Programme (Project Number ENV4-CT970586). The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

Recent changes in arctic vegetation might not be driven by climate change alone. Legacies of human activities have received little attention as a contributing factor. We examine the extent to which traditional human activities (hunting, herding, fire, wood extraction, and agriculture) have had lasting effects on arctic woody plant communities and therefore might continue to affect biome-wide responses to climate change. Evidence suggests that legacies are likely to be evident across meters to hundreds of kilometers and for decades, centuries, and millennia. The evidence, however, is currently sparse, and we highlight the potential to develop systematic assessments through a circumarctic collaboratory consisting of a network of interdisciplinary field sites, standardized protocols, participatory research, and new approaches. We suggest that human activities should be brought into consideration to increase our understanding of arctic vegetation dynamics in general and to assess woody plant responses to climate change in particular.

Climate change has been reported to affect shrub growth positively at several sites at high northern latitudes, including several arctic environments. The observed growth rates are, however, not uniform in space and time, and the mechanistic drivers of these patterns remain poorly understood. Here we investigated spatio-temporal interactions between climatic conditions, xylem anatomical traits, and annual growth of 21 Betula nana L. individuals from western Greenland for the period 2001–2011. Structural equation modeling showed that summer precipitation and winter temperature are affecting annual growth positively. Furthermore, vessel lumen area and vessel grouping, which are related to water conductivity and hydraulic connectivity of the xylem, respectively, positively influenced annual growth. To optimize growth B. nana was thus able to adjust its water transporting system. Annual variation in vessel lumen area seemed to be driven mostly by spring and summer temperatures, whereas annual variati...

The Arctic is warming at twice the rate of the rest of the world. This impacts Arctic species both directly, through increased temperatures, and indirectly, through structural changes in their habitats. Species are expected to exhibit idiosyncratic responses to structural change, which calls for detailed investigations at the species and community level. Here, we investigate how arthropod assemblages of spiders and beetles respond to variation in habitat structure at small spatial scales. We sampled transitions in shrub dominance and soil moisture between three different habitats (fen, dwarf shrub heath, and tall shrub tundra) at three different sites along a fjord gradient in southwest Greenland, using yellow pitfall cups. We identified 2,547 individuals belonging to 47 species. We used species richness estimation, indicator species analysis and latent variable modeling to examine differences in arthropod community structure in response to habitat variation at local (within site) and regional scales (between sites). We estimated species responses to the environment by fitting species-specific generalized linear models with environmental covariates. Species assemblages were segregated at the habitat and site level. Each habitat hosted significant indicator species, and species richness and diversity were significantly lower in fen habitats. Assemblage patterns were significantly linked to changes in soil moisture and vegetation height, as well as geographic location. We show that meter-scale variation among habitats affects arthropod community structure, supporting the notion that the Arctic tundra is a heterogeneous environment. To gain sufficient insight into temporal biodiversity change, we require studies of species distributions detailing species habitat preferences.

Aim Today’s genetic population structure and diversity of species can be understood as the result of range expansion from the area of origin, past climatic oscillations and contemporary processes. We examined the relative importance of these factors in Veratrum album L., a toxic weed of mountain grasslands. Location Continental Europe. Methods Forty populations from the Asian border (Urals and Caucasus) to Portugal were studied using amplified fragment length polymorphisms (AFLPs) combined with selected plant and population measures. The data were analysed with phylogenetic, population genetic and regression methods inferring both genetic structure and diversity from geographic and ecological factors. Results Fragment frequency clines together with genetic distance clustering and principal coordinates analysis indicated an east–west direction in the genetic structure of V. album, suggesting ancient migration into Europe from a proposed Asian origin. However, the strong geographic pattern in the genetic structure, pronounced isolation by distance (R2 = 0.74) and moderate overall population differentiation (FST = 0.13) suggests high historical gene flow, possibly during glacials, and vicariance into mountainous regions during interglacials. Occurrence of V. album during the last glaciation in several areas along the periphery of the Alps and recolonization of this mountain range from both eastern and central–western areas was indicated. Genetic diversity was highest in central Europe, a pattern that did not agree with the expectations from east–west migration into Europe. Furthermore, managed habitats showed higher levels of genetic diversity compared to unmanaged habitats. Stepwise linear regression determined shoot density and soil phosphorus as the main predictors of within-population genetic diversity (R2 = 0.40). Main conclusions Our results showed that V. album retained genetic imprints of historical range expansion into Europe, although this was alleviated by the influence of climatic oscillations and contemporary processes. For example, genetic population structure was strongly affected by post-glacial vicariance while patterns of genetic diversity seemed mainly to be influenced by human land use. Our findings highlight the importance of applying a synthetic approach, testing the influence of both historical and contemporary processes on genetic structure and diversity in order to understand complex phylogeographic patterns. This may especially apply to widespread species, such as weeds. Implications of our findings for biological control are briefly discussed.

The taxonomy of the Centaurea stoebe complex is controversial. Diploid and tetraploid plants occur in its native European range, but to date only tetraploids have been recorded from its introduced range in North America. We examined morphological differentiation of C. stoebe using multivariate and univariate approaches to clarify the taxonomic status of the known cytotypes. We measured more than 40 morphological traits on plants originating from 78 populations, grown from seed under uniform glasshouse conditions. The ploidy of almost 300 plants from 2 native and 20 introduced populations from Canada was assessed to test for the absence of diploids from North America. Finally, we explored whether postintroduction processes have resulted in phenotypic changes in introduced plants which may have contributed to the invasion success of C. stoebe. Morphometric analyses showed a clear separation of 2x and 4x plants and thus supported recognition of both cytotypes as separate taxa. Differences in the life cycle, the number of florets, the shape of capitula, and the shape of young rosette leaves were the best discriminant characters. Only minor differences were found between native and introduced tetraploids. All plants from the introduced range except for one hexaploid were found to be tetraploid. Rare diploids from Canada were identified as Centaurea diffusa or Centaurea psamogenna.

Summary 1. Introduced plants have the potential to rapidly evolve traits of ecological importance that may add to their innate potential to become invasive. During invasions, selection may favour genotypes that are already pre-adapted to conditions in the new habitat and, over time, alter the characteristics of subsequent generations. 2. Spotted knapweed (Centaurea stoebe) occurs in two predominantly spatially separated cytotypes in its native range (Europe–Western Asia), but currently only the tetraploid form has been confirmed in the introduced range (North America), where it is invasive. We used several common garden experiments to examine, across multiple populations, whether tetraploids and diploids from the native range differ in life cycle, leaf traits and reproductive capacity and if such differences would explain the predominance of tetraploids and their advance into new habitats in the introduced range. We also compared the same traits in tetraploids from the native and introduced range to determine whether any rapid adaptive changes had occurred since introduction that may have enhanced invasive potential of the species in North America. 3. We found tetraploids had lower specific leaf area, less lamina dissection and fewer, narrower leaves than diploids. Diploids exhibited a monocarpic life cycle and produced few if any accessory rosettes. Diploids produced significantly more seeds per capitulum and had more capitula per plant than tetraploids. In contrast, the vast majority of European tetraploids continued to flower in both seasons by regenerating from multiple secondary rosettes, demonstrating a predominantly polycarpic life cycle. 4. During early growth tetraploids from North America achieved greater biomass than both tetraploids and diploids from the native range but this did not manifest as larger above-ground biomass at maturity. In North American tetraploids there was also evidence of a shift towards a more strictly polycarpic life cycle, less leaf dissection, greater carbon investment per leaf, and greater seed production per capitulum. 5. Synthesis. Our results suggest that the characteristics of tetraploid C. stoebe pre-adapted them (compared to diploid conspecifics) for spread and persistence of the species into habitats in North America characterized by a more continental climate. After the species’ introduction, small but potentially important shifts in tetraploid biology have occurred that may have contributed significantly to successful invasion.

Polyploidy is often assumed to increase the spread and thus the success of alien plant species, but few empirical studies exist. We tested this hypothesis with Centaurea maculosa Lam., a species native to Europe and introduced into North America approximately 120 years ago where it became highly invasive. We analyzed the ploidy level of more than 2000 plants from 93 native and 48 invasive C. maculosa populations and found a pronounced shift in the relative frequency of diploid and tetraploid cytotypes. In Europe diploid populations occur in higher frequencies than tetraploids and only four populations had both cytotypes, while in North America diploid plants were found in only one mixed population and thus tetraploids clearly dominated. Our results showed a pronounced shift in the climatic niche between tetraploid populations in the native and introduced range toward drier climate in North America and a similar albeit smaller shift between diploids and tetraploids in the native range. The field data indicate that diploids have a predominately monocarpic life cycle, while tetraploids are often polycarpic. Additionally, the polycarpic life-form seems to be more prevalent among tetraploids in the introduced range than among tetraploids in the native range. Our study suggests that both ploidy types of C. maculosa were introduced into North America, but tetraploids became the dominant cytotype with invasion. We suggest that the invasive success of C. maculosa is partly due to preadaptation of the tetraploid cytotype in Europe to drier climate and possibly further adaptation to these conditions in the introduced range. The potential for earlier and longer seed production associated with the polycarpic life cycle constitutes an additional factor that may have led to the dominance of tetraploids over diploids in the introduced range.

Warming-induced expansion of trees and shrubs into tundra vegetation will strongly impact Arctic ecosystems. Today, a small subset of the boreal woody flora found during certain Plio-Pleistocene warm periods inhabits Greenland. Whether the twenty-first century warming will induce a re-colonization of a rich woody flora depends on the roles of climate and migration limitations in shaping species ranges. Using potential treeline and climatic niche modelling, we project shifts in areas climatically suitable for tree growth and 56 Greenlandic, North American and European tree and shrub species from the Last Glacial Maximum through the present and into the future. In combination with observed tree plantings, our modelling highlights that a majority of the non-native species find climatically suitable conditions in certain parts of Greenland today, even in areas harbouring no native trees. Analyses of analogous climates indicate that these conditions are widespread outside Greenland, thus increasing the likelihood of woody invasions. Nonetheless, we find a substantial migration lag for Greenland's current and future woody flora. In conclusion, the projected climatic scope for future expansions is strongly limited by dispersal, soil development and other disequilibrium dynamics, with plantings and unintentional seed dispersal by humans having potentially large impacts on spread rates., Philosophical Transactions of the Royal Society B: Biological Sciences, 368 (1624), ISSN:0962-8436, ISSN:1471-2970, ISSN:0080-4622

news and update ISSN 1948‐6596 commentary Tree refugia and slow forest development in response to post‐ LGM warming in North‐Eastern European Russia Though the Postglacial period counts its thousands of years, it was not indefinitely long, and few plants that merely scatter their seed could advance more than a yard in a year […]. The oak, to gain its present most northerly position in North Britain after being driven out by the cold, probably had to travel fully six hundred miles, and this without external aid would take some‐ thing like a million years. (Clement Reid 1899, p. 25) Driven by an urge to understand how the compo‐ sition of the British flora arose, Clement Reid was puzzled by the discrepancy between the apparent low dispersal capacity of big seeded trees and the high postglacial migration rates deduced from fos‐ sil evidence. Possible explanations resolving Reid’s paradox, as it was later termed, were fast migra‐ tion by long‐distance dispersal events or glacial survival of small scattered populations at shel‐ tered sites further north than evidenced by the fossil record (Reid 1899, Skellam 1951). More than 100 years have passed since the early writings of Reid, but two questions leading from his thoughts are still debated: How far north did tree species survive the Last Glacial Maximum (LGM, ca. 21 kya)? How fast did tree species spread and thus current forest structure develop during post‐LGM warming? A new paleoecological study by Valiranta et al. (2011, Journal of Biogeography) provides, in addition to several other recent studies (e.g., Bin‐ ney et al. 2009, Heikkila et al. 2009), insight into these unresolved questions. Valiranta et al. (2011) summarized macrofossil and stomata records from five sites at the late‐glacial–early Holocene transition at the European Russian arctic tree line; an area east of the margin of the Scandinavian ice sheet and west of the Ural Mountains (Fig. 1). Their data evidenced the presence of Norway spruce (Picea abies) and tree birch (Betula pen‐ dula/pubescens) during the early Holocene (11,500–10,000 cal. yr BP) at all five sites. At one site, two radiocarbon dates even suggested the presence of these taxa up to 1000 yrs prior to the beginning of the Holocene. Historical biogeogra‐ phy, however, depends on reliable chronologies of fossil records. Ideally, each site should have a well established chronology based on an adequate number of radiocarbon dated fragile and precisely identified terrestrial plant remains that show no signs of redeposition. In this respect, the chro‐ nologies of Valiranta et al. (2011) were not opti‐ mal and the late‐glacial presence of P. abies and tree Betula can be questioned, but their early Holocene records were well documented. Hence, the smoking gun, i.e., direct palaeoecological proof of LGM‐populations, might be evasive, but scattered glacial and widespread early Holocene fossil records indeed reflect consistent patterns of early expansions in North‐Eastern European Rus‐ sia. Furthermore, independent climate reconstruc‐ tions are crucial when evaluating the possible LGM survival of boreal tree populations so far north. Glacier‐based climate models do indeed indicate relatively modest LGM temperature de‐ pressions in the arctic Ural Mountains (Allen et al. 2008). This might have facilitated survival of trees. Besides, the ability to produce krummholz forms (Oberg & Kullman 2011) or other strategies for non‐reproductive persistence through millennia (Bhagwat & Willis 2008) may have been decisive. Nevertheless, as concluded by Valiranta et al. (2011), it certainly seems likely that postglacial development of high‐latitude forests occurred through expansion of local tree populations rather than through re‐colonisation from distant south‐ ern refugia. Using molecular markers, recent phy‐ logeographic studies for both P. abies and Betula support glacial survival in north‐eastern Europe (Maliouchenko et al. 2007, Tollefsrud et al. 2008). For example, P. abies shows distinct gene pools of similar within‐population genetic diversity for northern and central‐southern Europe (Tollefsrud et al. 2008): a pattern that supports west expan‐ sion from such north‐eastern refugia and not re‐ colonisation of northern Europe from the south. For decades, pollen data have suggested early frontiers of biogeography 2.4, 2011 — © 2011 the authors; journal compilation © 2011 The International Biogeography Society

Chloroplast DNA sequences are of great interest for population genetics and phylogenetic studies. However, only a small set of markers are commonly used. Most of them have been designed for amplification in a large range of Angiosperms and are located in the Large Single Copy (LSC). Here we developed a new set of 100 primer pairs optimized for amplification in Monocotyledons. Primer pairs amplify coding (exon) and non-coding regions (intron and intergenic spacer). They span the different chloroplast regions: 72 are located in the LSC, 13 in the Small Single Copy (SSC) and 15 in the Inverted Repeat region (IR). Amplification and sequencing were tested in 13 species of Monocotyledons: Dioscorea abyssinica, D. praehensilis, D. rotundata, D. dumetorum, D. bulbifera, Trichopus sempervirens (Dioscoreaceae), Phoenix canariensis, P. dactylifera, Astrocaryum scopatum, A. murumuru, Ceroxylon echinulatum (Arecaceae), Digitaria excilis and Pennisetum glaucum (Poaceae). The diversity found in Dioscorea, Digitaria and Pennisetum mainly corresponded to Single Nucleotide Polymorphism (SNP) while the diversity found in Arecaceae also comprises Variable Number Tandem Repeat (VNTR). We observed that the most variable loci (rps15-ycf1, rpl32-ccsA, ndhF-rpl32, ndhG-ndhI and ccsA) are located in the SSC. Through the analysis of the genetic structure of a wild-cultivated species complex in Dioscorea, we demonstrated that this new set of primers is of great interest for population genetics and we anticipate that it will also be useful for phylogeny and bar-coding studies.

Aim We examined whether species occurrences are primarily limited by physiological tolerance in the abiotically more stressful end of climatic gradients (the asymmetric abiotic stress limitation (AASL) hypothesis) and the geographical predictions of this hypothesis: abiotic stress mainly determines upper-latitudinal and upper-altitudinal species range limits, and the importance of abiotic stress for these range limits increases the further northwards and upwards a species occurs. Location Europe and the Swiss Alps. Methods The AASL hypothesis predicts that species have skewed responses to climatic gradients, with a steep decline towards the more stressful conditions. Based on presence-absence data we examined the shape of plant species responses (measured as probability of occurrence) along three climatic gradients across latitudes in Europe (1577 species) and altitudes in the Swiss Alps (284 species) using Huisman-Olff-Fresco, generalized linear and generalized additive models. Results We found that almost half of the species from Europe and one-third from the Swiss Alps showed responses consistent with the predictions of the AASL hypothesis. Cold temperatures and a short growing season seemed to determine the upper-latitudinal and upper-altitudinal range limits of up to one-third of the species, while drought provided an important constraint at lower-latitudinal range limits for up to one-fifth of the species. We found a biome-dependent influence of abiotic stress and no clear support for abiotic stress as a stronger upper range-limit determinant for species with higher latitudinal and altitudinal distributions. However, the overall influence of climate as a range-limit determinant increased with latitude. Main conclusions Our results support the AASL hypothesis for almost half of the studied species, and suggest that temperature-related stress controls the upper-latitudinal and upper-altitudinal range limits of a large proportion of these species, while other factors including drought stress may be important at the lower range limits.

The “evolution of increased competitive ability” (EICA) hypothesis proposes that escape from natural enemies, e.g., after transcontinental introductions, alters the selection regime because costly defenses no longer enhance fitness. Such an evolutionary loss of defenses enables resources to be directed toward growth or other traits improving performance. We tested the EICA hypothesis in a novel framework in which the natural enemy is the traveler that follows its widespread host by accidental or deliberate (biocontrol) introductions. In a greenhouse experiment we used populations of Senecio vulgaris from North America, Europe, and Australia that differ in the history of exposure to the rust fungus Puccinia lagenophorae. Contrary to what is predicted by EICA, we found no evidence for increased levels of resistance to the rust fungus in plant populations with a longer history of rust exposure (Australia). Similarly, there was no evidence for reduced fecundity in these populations, although vegetative vigor, measured as secondary branching and growth rate, was lower. The maintenance of high rust resistance in populations with no (North America) or only a short history (Europe) of rust exposure is surprising given that resistance seems to incur considerable fitness costs, as indicated by the negative association between family mean resistance and fitness in the absence of disease observed for all three continents. The comparison of population differentiation in quantitative traits with estimates of differentiation in amplified fragment length polymorphic (AFLP) markers suggests that a number of fitness-related traits are under divergent selection among the studied populations. The proposed framework to test changes in the evolutionary trajectory underlying EICA can be employed in an expanded range of systems. These may include investigations on a cosmopolitan weed or crop when an antagonist is expanding its geographic range (such as our study), studies along a chronosequence of introduction time with expected increasing accumulation of natural enemies over time, or comparisons between introduced plant populations that differ in exposure time to biocontrol organisms.

Background: Phylogenetic and phylogeographic investigations have been previously performed to study the evolution of the olive tree complex (Olea europaea). A particularly high genomic diversity has been found in north-west Africa. However, to date no exhaustive study has been addressed to infer putative polyploidization events and their evolutionary significance in the diversification of the olive tree and its relatives. Methods: Representatives of the six olive subspecies were investigated using (a) flow cytometry to estimate genome content, and (b) six highly variable nuclear microsatellites to assess the presence of multiple alleles at co-dominant loci. In addition, nine individuals from a controlled cross between two individuals of O. europaea subsp. maroccana were characterized with microsatellites to check for chromosome inheritance. Key Results: Based on flow cytometry and genetic analyses, strong evidence for polyploidy was obtained in subspp. cerasiformis (tetraploid) and maroccana (hexaploid), whereas the other subspecies appeared to be diploids. Agreement between flow cytometry and genetic analyses gives an alternative approach to chromosome counting to determine ploidy level of trees. Lastly, abnormalities in chromosomes inheritance leading to aneuploid formation were revealed using microsatellite analyses in the offspring from the controlled cross in subsp. maroccana. Conclusions: This study constitutes the first report for multiple polyploidy in olive tree relatives. Formation of tetraploids and hexaploids may have played a major role in the diversification of the olive complex in north-west Africa. The fact that polyploidy is found in narrow endemic subspecies from Madeira (subsp. cerasiformis) and the Agadir Mountains (subsp. maroccana) suggests that polyploidization has been favoured to overcome inbreeding depression. Lastly, based on previous phylogenetic analyses, we hypothesize that subsp. cerasiformis resulted from hybridization between ancestors of subspp. guanchica and europaea.

International audience; Niche-based models calibrated in the native range by relating species observations to climatic variables are commonly used to predict the potential spatial extent of species' invasion. This climate matching approach relies on the assumption that invasive species conserve their climatic niche in the invaded ranges. We test this assumption by analysing the climatic niche spaces of Spotted Knapweed in western North America and Europe. We show with robust cross-continental data that a shift of the observed climatic niche occurred between native and non-native ranges, providing the first empirical evidence that an invasive species can occupy climatically distinct niche spaces following its introduction into a new area. The models fail to predict the current invaded distribution, but correctly predict areas of introduction. Climate matching is thus a useful approach to identify areas at risk of introduction and establishment of newly or not-yet-introduced neophytes, but may not predict the full extent of invasions.

We examined whether nitrogen (N) and carbohydrates reserves allow Veratrum album, an alpine forb, to start spring growth earlier than the neighbouring vegetation and to survive unpredictable disturbances resulting in loss of above-ground biomass.Seasonal dynamics of plant reserves, soil N availability and vegetation growth were monitored. Veratrum album shoots were experimentally removed when carbohydrate reserves were at a seasonal minimum and the subsequent changes in biomass and reserves were compared with those in control plants. Reserves did not give V. album a competitive advantage in spring; however, they did function as a buffer against the impact of calamities. Shoot removal resulted in significantly lower root dry weight, higher N concentration in rhizome and roots and lower starch concentrations in rhizome and roots but no plant mortality was observed.Veratrum album used stored N reserves to supplement N uptake and establish high leaf N concentrations, which facilitated a rapid refilling of depleted carbohydrate reserves. The primary function of N reserves appears to be to allow V. album to complete the growing cycle in as short a period as possible, thus minimizing exposure to above-ground risks.

The combined effects of climate change and nutrient availability on Arctic vegetation growth are poorly understood. Archaeological sites in the Arctic could represent unique nutrient hotspots for studying the long-term effect of nutrient enrichment. In this study, we analysed a time-series of ring widths of Salix glauca L. collected at nine archaeological sites and in their natural surroundings along a climate gradient in the Nuuk fjord region, Southwest Greenland, stretching from the edge of the Greenlandic Ice Sheet in the east to the open sea in the west. We assessed the temperature-growth relationship for the last four decades distinguishing between soils with past anthropogenic nutrient enrichment (PANE) and without (controls). Along the East–West gradient, the inner fjord sites showed a stronger temperature signal compared to the outermost ones. Individuals growing in PANE soils had wider ring widths than individuals growing in the control soils and a stronger climate-growth relation, especially in the inner fjord sites. Thereby, the individuals growing on the archaeological sites seem to have benefited more from the climate warming in recent decades. Our results suggest that higher nutrient availability due to past human activities plays a role in Arctic vegetation growth and should be considered when assessing both the future impact of plants on archaeological sites and the general greening in landscapes with contrasting nutrient availability.