Reindeer herbivory reduces willow growth and grouse forage in a forest-tundra ecotone

Browsing mammals strongly modify the structure of vegetation of forest-tundra ecotones. We investigated the impact of reindeer browsing on growth and morphology of an arctic willow, Salix glauca, by studying plants inside and outside fenced areas in a tundra habitat at the tree line. We also studied if reindeer feeding has an effect on the forage availability for willow grouse, a herbivore sharing the same food plant. Analyses of 6-year data show that reindeer strongly reduces the growth of tundra willow and changes plant morphology to a stunted growth form. Intense reindeer browsing on willow limited the forage availability for willow grouse and grouse fed less on reindeer-browsed willow than willows protected from reindeer browsing. The results of this study imply that herbivores can counteract the increase of shrubs in forest-tundra ecotones, as has been predicted in some studies discussing the effects of climatic warming on vegetation. Furthermore, trophic interactions should be incorporated in modelling vegetation changes as a response to increased temperatures.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

Ecological concepts and biographical zonation in the North: the need for a generally accepted terminology

The author stresses the need for more clearly defined concepts and terms related to ecological research in the northern biogeographical zones and regions. The concepts ‘arctic’, ‘subarctic’, ‘tundra’, ‘forest-tundra’, ‘open woodland’ (and similar expressions), ‘taiga’, etc., are discussed below. The expressions ‘tree’ and ‘forest’ are briefly mentioned. The main interest below is focused on terms related to the tree- and timber- (forest-) lines and their interrelations.     

Latitudinal response of subarctic tree lines to recent climate change in eastern Canada

Aim The predictions from biogeographical models of poleward expansion of biomes under a warmer 2 × CO2 scenario might not be warranted, given the non‐climatic influences on vegetation dynamics. Milder climatic conditions have occurred in northern Québec, Canada, in the 20th century. The purpose of this study was to document the early signs of a northward expansion of the boreal forest into the subarctic forest‐tundra, a vast heterogeneous ecotone. Colonization of upland tundra sites by black spruce (Picea mariana (Mill.) BSP.) forming local subarctic tree lines was quantified at the biome scale. Because it was previously shown that the regenerative potential of spruce is reduced with increasing latitude, we predicted that tree line advances and recent establishment of seedlings above tree lines will also decrease northwards. Location Black spruce regeneration patterns were surveyed across a > 300‐km latitudinal transect spanning the forest‐tundra of northern Québec, Canada (55°29′–58°27′ N). Methods Elevational transects were positioned at forest–tundra interfaces in two regions from the southern forest‐tundra and two regions from the northern forest‐tundra, including the arctic tree line. The surroundings of stunted black spruce, forming the species limit in the shrub tundra, were also examined. Position, total height and origin (seed or layer) of all black spruce stems established in the elevational transects were determined. Dendrochronological and topographical data allowed recent subarctic tree line advances to be estimated. Age structures of spruce recently established from seed (< 2.5 m high) were constructed and compared between forest‐tundra regions. Five to 20‐year heat sum (growing degree‐days, > 5 °C) and precipitation fluctuations were computed from regional climatic data, and compared with seedling recruitment patterns. Results During the 20th century, all tree lines from the southern forest‐tundra rose slightly through establishment of seed‐origin spruce, while some tree lines in the northern forest‐tundra rose through height growth of stunted spruce already established on the tundra hilltops. However, the rate of rise in tree lines did not slow down with latitude. The density of < 2.5‐m spruce established by seed declined exponentially with latitude. While the majority of < 2.5‐m spruce has established since the late 1970s on the southernmost tundra hilltops, the regeneration pool was mainly composed of old, suppressed individuals in the northern forest‐tundra. Spruce age generally decreased with increasing elevation in the southern forest‐tundra stands, therefore indicating current colonization of tundra hilltops. Although spruce reproductive success has improved over the twentieth century in the southern forest‐tundra, there was hardly any evidence that recruitment of seed‐origin spruce was controlled by 5‐ to 20‐year regional climatic fluctuations, except for winter precipitation. Main conclusions Besides the milder 20th century climate, local topographic factors appear to have influenced the rise in tree lines and recent establishment by seed. The effect of black spruce's semi‐serotinous cones in trapping seeds and the difficulty of establishment on exposed, drought‐prone tundra vegetation are some factors likely to explain the scarcity of significant correlations between tree establishment and climatic variables in the short term. The age data suggest impending reforestation of the southernmost tundra sites, although the development of spruce seedlings into forest might be slowed down by the harsh wind‐exposure conditions.     

Directional change in upland tundra plant communities 20‐30 years after seismic exploration in the Canadian low‐arctic

Question: What is the disturbance response of low‐arctic plant communities two to three decades after seismic exploration. Location: Mackenzie River Delta, low‐arctic, northwestern Canada. Methods: Plant communities in two upland tundra vegetation types were compared between winter seismic lines, created between 1970 and 1986, and adjacent “reference” tundra. Also, we used aerial surveys to quantify the total area impacted by visible linear features. Results: Vascular plant cover was significantly higher, and lichen cover significantly lower, on seismic lines than in reference tundra. The increase in vascular plant cover was attributable to deciduous shrubs and graminoids. There were significant differences in plant community composition between seismic lines and reference tundra but no differences in species diversity or richness. Betula glandulosa and Arctagrostis latifolia were significant indicator species for seismic lines, while Saussurea angustifolia was a significant indicator for reference tundra. Based on the aerial surveys, these effects apply to at least 90% of seismic lines from two‐dimensional programs in these habitat types during the 1970s. Conclusions: Vegetation composition and structure on 20‐30‐year‐old seismic lines differs from reference upland tundra despite no persistent differences in organic layer depth or depth to permafrost. We propose that this reflects: (1) successional redevelopment following changes in soil conditions and nutrient availability arising from the disturbance, and/or (2) disturbance‐initiated succession towards a community reflecting current climatic conditions.     

Tall shrub and tree expansion in Siberian tundra ecotones since the 1960s

Circumpolar expansion of tall shrubs and trees into Arctic tundra is widely thought to be occurring as a result of recent climate warming, but little quantitative evidence exists for northern Siberia, which encompasses the world's largest forest‐tundra ecotonal belt. We quantified changes in tall shrub and tree canopy cover in 11, widely distributed Siberian ecotonal landscapes by comparing very high‐resolution photography from the Cold War‐era ‘Gambit’ and ‘Corona’ satellite surveillance systems (1965–1969) with modern imagery. We also analyzed within‐landscape patterns of vegetation change to evaluate the susceptibility of different landscape components to tall shrub and tree increase. The total cover of tall shrubs and trees increased in nine of 11 ecotones. In northwest Siberia, alder (Alnus) shrubland cover increased 5.3–25.9% in five ecotones. In Taymyr and Yakutia, larch (Larix) cover increased 3.0–6.7% within three ecotones, but declined 16.8% at a fourth ecotone due to thaw of ice‐rich permafrost. In Chukotka, the total cover of alder and dwarf pine (Pinus) increased 6.1% within one ecotone and was little changed at a second ecotone. Within most landscapes, shrub and tree increase was linked to specific geomorphic settings, especially those with active disturbance regimes such as permafrost patterned‐ground, floodplains, and colluvial hillslopes. Mean summer temperatures increased at most ecotones since the mid‐1960s, but rates of shrub and tree canopy cover expansion were not strongly correlated with temperature trends and were better correlated with mean annual precipitation. We conclude that shrub and tree cover is increasing in tundra ecotones across most of northern Siberia, but rates of increase vary widely regionally and at the landscape scale. Our results indicate that extensive changes can occur within decades in moist, shrub‐dominated ecotones, as in northwest Siberia, while changes are likely to occur much more slowly in the highly continental, larch‐dominated ecotones of central and eastern Siberia.     

Pathways of tundra encroachment by trees and tall shrubs in the western Brooks Range of Alaska

Climate change is expected to increase woody vegetation abundance in the Arctic, yet the magnitude, spatial pattern and pathways of change remain uncertain. We compared historical orthophotos photos (1952 and 1979) with high-resolution satellite imagery (2015) to examine six decades of change in abundance of white spruce Picea glauca and tall shrubs (Salix spp., Alnus spp.) near the Agashashok River in northwest Alaska. We established ~3000 random points within our ~5500 ha study area for classification into nine cover types. To examine physiographic controls on tree abundance, we fit multinomial log-linear models with predictors derived from a digital elevation model and with arctic tundra, alpine tundra and ‘tree’ as levels of a categorical response variable. Between 1952 and 2015, points classified as arctic and alpine tundra decreased by 31% and 15%, respectively. Meanwhile, tall shrubs increased by 86%, trees mixed with tall shrubs increased by 385% and forest increased by 84%. Tundra with tall shrubs rarely transitioned to forest. The best multinomial model explained 71% of variation in cover and included elevation, slope and an interaction between slope and ‘northness’. Treeline was defined as the elevation where the probability of tree presence equaled that of tundra. Mean treeline elevation in 2015 was 202 m, corresponding with a June–August mean air temperature > 11°C, which is > 4°C warmer than the 6–7°C isotherm associated with global treeline elevations. Our results show dramatic increases in the abundance of trees and tall shrubs, question the universality of air temperature as a predictor of treeline elevation and suggest two mutually exclusive pathways of vegetation change, because tundra that gained tall shrubs rarely transitioned to forest. Conversion of tundra to tall shrubs and forest has important and potentially contrasting implications for carbon cycling, surface energy exchange and wildlife habitat in the Arctic.     

Satellite‐based evidence for shrub and graminoid tundra expansion in northern Quebec from 1986 to 2010

Global vegetation models predict rapid poleward migration of tundra and boreal forest vegetation in response to climate warming. Local plot and air‐photo studies have documented recent changes in high‐latitude vegetation composition and structure, consistent with warming trends. To bridge these two scales of inference, we analyzed a 24‐year (1986–2010) Landsat time series in a latitudinal transect across the boreal forest‐tundra biome boundary in northern Quebec province, Canada. This region has experienced rapid warming during both winter and summer months during the last 40 years. Using a per‐pixel (30 m) trend analysis, 30% of the observable (cloud‐free) land area experienced a significant (P < 0.05) positive trend in the Normalized Difference Vegetation Index (NDVI). However, greening trends were not evenly split among cover types. Low shrub and graminoid tundra contributed preferentially to the greening trend, while forested areas were less likely to show significant trends in NDVI. These trends reflect increasing leaf area, rather than an increase in growing season length, because Landsat data were restricted to peak‐summer conditions. The average NDVI trend (0.007 yr^?1) corresponds to a leaf‐area index (LAI) increase of ~0.6 based on the regional relationship between LAI and NDVI from the Moderate Resolution Spectroradiometer. Across the entire transect, the area‐averaged LAI increase was ~0.2 during 1986–2010. A higher area‐averaged LAI change (~0.3) within the shrub‐tundra portion of the transect represents a 20–60% relative increase in LAI during the last two decades. Our Landsat‐based analysis subdivides the overall high‐latitude greening trend into changes in peak‐summer greenness by cover type. Different responses within and among shrub, graminoid, and tree‐dominated cover types in this study indicate important fine‐scale heterogeneity in vegetation growth. Although our findings are consistent with community shifts in low‐biomass vegetation types over multi‐decadal time scales, the response in tundra and forest ecosystems to recent warming was not uniform.     

Recent Shrub Proliferation in the Mackenzie Delta Uplands and Microclimatic Implications

Local observations, repeat photos, and broad-scale remote sensing suggest that tall shrubs are becoming an increasingly dominant component of Low Arctic ecosystems. This shift has the potential to alter the surface energy balance through changes to the surface albedo, snow accumulation and melt, and ground thermal regimes. However, to date there have been few quantitative estimates of the rate of tall shrub expansion. We used soft copy stereo visualization of air photos to map fine-scale changes in tall shrub tundra and green alder density in the upland tundra north of Inuvik, NT between 1972 and 2004. We also used 2004 photos to map tall shrub tundra in areas affected by fires that occurred between 1960 and 1968. To assess the potential impact of vegetation change on microclimate, we used pyranometers to measure albedo and net solar radiation, thermistors attached to data loggers to record ground temperatures, and field surveys to record winter snow conditions in three common vegetation types. Fine-scale mapping shows that green alder stem density has increased by 68% (±24.1) since 1972. Average tall shrub tundra cover has also increased by 15% (±3.6) since 1972. Historical tundra fires had the highest proportion of tall shrub cover of all areas mapped using 2004 photos, ranging from 92 to 99%. Based on these results, we suggest that predicted increases in the size and frequency of tundra fire are likely to drive rapid shrub proliferation in the Low Arctic. Shrub-dominated sites have decreased albedo, increased net solar radiation, deeper snow pack, and elevated near-surface ground temperatures, indicating that continued increases in shrub cover will affect regional climate, hydrology, permafrost temperatures, and terrain stability.     

Spatially explicit fire-climate history of the boreal forest-tundra (Eastern Canada) over the last 2000 years

Across the boreal forest, fire is the main disturbance factor and driver of ecosystem changes. In this study, we reconstructed a long-term, spatially explicit fire history of a forest-tundra region in northeastern Canada. We hypothesized that current occupation of similar topographic and edaphic sites by tundra and forest was the consequence of cumulative regression with time of forest cover due to compounding fire and climate disturbances. All fires were mapped and dated per 100 year intervals over the last 2,000 years using several fire dating techniques. Past fire occurrences and post-fire regeneration at the northern forest limit indicate 70% reduction of forest cover since 1800 yr BP and nearly complete cessation of forest regeneration since 900 yr BP. Regression of forest cover was particularly important between 1500s-1700s and possibly since 900 yr BP. Although fire frequency was very low over the last 100 years, each fire event was followed by drastic removal of spruce cover. Contrary to widespread belief of northward boreal forest expansion due to recent warming, lack of post-fire recovery during the last centuries, in comparison with active tree regeneration more than 1,000 years ago, indicates that the current climate does not favour such expansion.     

Long-term Impacts of Contrasting Management of Large Ungulates in the Arctic Tundra-Forest Ecotone: Ecosystem Structure and Climate Feedback

The arctic forest-tundra ecotone (FTE) represents a major transition zone between contrasting ecosystems, which can be strongly affected by climatic and biotic factors. Expected northward expansion and encroachment on arctic tundra in response to climate warming may be counteracted by natural and anthropogenic processes such as defoliating insect outbreaks and grazing/browsing regimes. Such natural and anthropogenic changes in land cover can substantially affect FTE dynamics, alter ground albedo (index of the amount of solar energy reflected back into the atmosphere) and provide important feedbacks into the climate system. We took advantage of a naturally occurring contrast between reindeer grazing regimes in a border region between northern Finland and Norway which was recently defoliated by an outbreak of the geometrid moth. We examined ecosystem-wide contrasts between potentially year-round (but mainly summer) grazed (YRG) regions in Finland and mainly winter grazed (WG) regions in Norway. We also used a remotely sensed vegetation index and albedo to quantify effects on local energy balance and potential climate feedbacks. Although differences in soil characteristics and ground vegetation cover were small, we found dramatic differences in the tree layer component of the ecosystem. Regeneration of mountain birch stands appears to have been severely hampered in the YRG regime, by limiting regeneration from basal shoots and reestablishment of individual trees from saplings. This has led to a more open forest structure and a significant 5% increase in spring albedo in the summer grazed compared to the winter grazed regions. This supports recent suggestions that ecosystem processes in the Arctic can significantly influence the climate system, and that such processes must be taken into account when developing climate change scenarios and adaptation strategies.     

Zooplankton community composition of lakes in the Yukon and Northwest Territories (Canada): relationship to physical and chemical limnology

We analysed associations between zooplankton species composition and local abiotic factors in 30 lakes located along a 900 km south-north transect from Whitehorse (Yukon Territory) to Inuvik (Northwest Territories). The lakes were situated in three broadly defined vegetation zones: (i) Boreal forest (between Whitehorse and Dawson City), (ii) alpine tundra (Ogilvie mountains north of Dawson City) and (iii) subarctic forest-tundra (near Inuvik). Lakes in the alpine tundra were characterised by lower conductivity, temperature, chlorophyll a and nutrients than those in the other two zones. Those in the forest-tundra were generally small and shallow, and had higher chlorophyll a concentrations than lakes further south. Lakes in forested catchments spanned a larger latitudinal range and exhibited a greater variety of physical and chemical characteristics. However, they were generally deeper, with higher conductivity, temperature and ionic concentrations. Forty-one zooplankton taxa were identified from the 30 lakes, of which the most frequently occurring were the rotifers Conochilus unicornis, Kellicottia longispina, Keratella cochlearis and Polyarthra vulgaris, the cladocerans Daphnia middendorffiana and Bosmina longirostris, and the copepods Leptodiaptomus pribilofensis, Heterocope septentrionalis and Cyclops spp. The lakes contained between two and fifteen species (mean = 6.9). Alpine tundra lakes contained slightly less species (mean = 5.8) than those at lower elevations; in particular the cladoceran fauna was depauperate or absent. Relationships among the lakes, species and environmental factors were examined using canonical correspondence analysis, with forward selection and associated Monte Carlo permutation tests. Chloride, silica and temperature showed statistically significant relationships with species distribution, and together these abiotic factors explained 25% of the variation in zooplankton communities within Yukon and Northwest Territories lakes.     

Compositional differentiation,vegetation‐environment relationships and classification of willow‐characterised vegetation in the western Eurasian Arctic

Question: How does willow‐characterised tundra vegetation of western Eurasia vary, and what are the main vegetation types? What are the ecological gradients and climatic regimes underlying vegetation differentiation? Location: The dataset was collected across a wide spectrum of tundra habitats at 12 sites in subarctic and arctic areas spanning from NW Fennoscandia to West Siberia. Methods: The dataset, including 758 vegetation sample plots (relevés), was analysed using a TWINSPAN classification and NMDS ordination that also included analyses of vegetation‐environment correlations. Results: Based on the TWINSPAN classification, eight vegetation types characterised by willow (cover of upright willows >10%) were discerned: (1) Salix glauca‐Carex aquatilis type, (2) Aulacomnium‐Tomentypnum type, (3) Salix‐Betula‐Hylocomium type, (4) Salix lanata‐Brachythecium mildeanum type, (5) Salix‐Pachypleurum type, (6) S. lanata‐Myosotis nemorosa type, (7) Salix‐Trollius‐Geranium type and (8) Salix‐Comarum palustre‐Filipendula ulmaria type. Willow‐characterised vegetation types were compositionally differentiated from other tundra vegetation and were confined to relatively moist valley and sloping tundra sites, from mire to mineral soils. These vegetation types were encountered across a broad latitudinal zone in which July mean temperature ranged from 6 to 10°C. Conclusions: Willow‐characterised tundra vegetation forms a broad category of ecologically and geographically differentiated vegetation types that are linked to dwarf shrub tundra, shrub tundra or mire. Because of complex ecological gradients underlying compositional differentiation, predicting the responses of willow‐characterised tundra vegetation to a warming climate may be complicated.     

Long- and short-term effects of reindeer grazing on tundra wetland vegetation

We studied long-term (50 years) and short-term (4 years) effects of summer grazing of reindeer on subarctic tundra wetland vegetation. The long-term effects of summer grazing were studied by comparing vegetation on Finnish and Norwegian sides of the fence line separating reindeer grazing regimes. The Finnish side was intensively grazed and trampled throughout the year, whereas the Norwegian side was grazed in winter. Experimental fences were erected to examine short-term effects of grazing exclusion. Both in the long- and short-term, summer grazing decreased the height of Salix lapponum whereas the short-term effects on willow cover were less clear than the long-term effects. In contrast, Carex spp. benefited from grazing. Long-term grazing had little effect on total bryophyte cover. Grazing had negligible effects on the nutrient content of leaves of S. lapponum and Eriophorum angustifolium. We conclude that tundra wetlands can withstand moderately high grazing pressure sustained over several decades.     

Modern pollen and stomate deposition in lake surface sediments from across the treeline on the Kola Peninsula, Russia

We sampled and analyzed surface sediments from 31 lakes along a latitudinal transect crossing the coniferous treeline on the Kola Peninsula, Russia. The major vegetation zones along the transect were tundra, birch-forest tundra, pine-forest tundra, and forest. The results indicate that the major vegetation types in our study area have distinct pollen spectra. Sum-of-squares cluster analysis and principal components analysis (PCA) groupings of pollen sites correspond to the major vegetation zones. PCA ordination of taxa indicates that the first axis separates taxa typical of the forest zone (Pinus, Picea) from taxa typical of tundra and forest-tundra zones (Polypodiaceae, Ericaceae, and Betula). The current position of the coniferous treeline, defined in our region by Pinus sylvestris, occurs roughly where Pinus pollen values reach 35% or greater. Arboreal pollen (AP)/non-arboreal pollen (NAP) ratios were calculated for each site and plotted against geographic distance along the transect. AP/NAP ratios of 7 or greater are found within pine-forest tundra and forest vegetation zones. Pinus stomates (dispersed stomatal guard cells) are absent from sites north of the coniferous treeline and all but two samples from the forested sites contain stomates. Stomate concentrations among the samples are highly variable and range from 10 to 458 per ml and positively correlate with the changing Pinus pollen values.     

Spatial heterogeneity of tundra vegetation response to recent temperature changes

The spatial heterogeneity of recent decadal dynamics in vegetation greenness and biomass in response to changes in summer warmth index (SWI) was investigated along spatial gradients on the Arctic Slope of Alaska. Image spatial analysis was used to examine the spatial pattern of greenness dynamics from 1991 to 2000 as indicated by variations of the maximum normalized difference vegetation index (Peak NDVI) and time‐integrated NDVI (TI‐NDVI) along latitudinal gradients. Spatial gradients for both the means and temporal variances of the NDVI indices for 0.1° latitude intervals crossing three bioclimate subzones were analyzed along two north–south Arctic transects. NDVI indices were generally highly variable over the decade, with great heterogeneity across the transects. The greatest variance in TI‐NDVI was found in low shrub vegetation to the south (68.7–68.8°N) and corresponded to high fractional cover of shrub tundra and moist acidic tundra (MAT), while the greatest variance in Peak‐NDVI predominately occurred in areas dominated by wet tundra (WT) and moist nonacidic tundra (MNT). Relatively high NDVI temporal variances were also related to specific transitional areas between dominant vegetation types. The regional temporal variances of NDVI from 1991 to 2000 were largely driven by meso‐scale climate dynamics. The spatial heterogeneity of the NDVI variance was mostly explained by the fractional land cover composition, different responses of each vegetation type to climate change, and patterned ground features. Aboveground plant biomass exhibited similar spatial heterogeneity as TI‐NDVI; however, spatial patterns are slightly different from NDVI because of their nonlinear relationship.     

Nitrogen‐dependent recovery of subarctic tundra vegetation after simulation of extreme winter warming damage to Empetrum hermaphroditum

Vast areas of (sub)arctic tundra are dominated by the ericoid dwarf shrub Empetrum hermaphroditum. Recent experimental and observational data have shown that Empetrum can be damaged heavily by recurrent extreme winter warming. In addition, summer warming leads to increased soil N availability in tundra ecosystems. In a 7‐year experiment, I investigated the recovery of subarctic Empetrum‐dominated tundra vegetation using a factorial combination of various degrees of aboveground Empetrum removal (simulating the damaging effects of extreme winter warming) and N addition (simulating one of the effects of summer warming). After 7 years no new species had established in the plots. The growth of planted Betula nana seedlings was stimulated by Empetrum removal and reduced by N addition. This Empetrum‐dominated tundra ecosystem was resilient against severe disturbances. Only when Empetrum was 100% removed did it fail to recover, and only in combination with high N supply the subordinate species (notably Eriophorum vaginatum and Rubus chamaemorus, a graminoid and a forb) could benefit. In the 50% removal treatment Empetrum recovered in 7 years when no N was supplied and the cover of the subordinate species did not change. However, when N was added Empetrum recovered faster (in 4 years) and the subordinates decreased. When Empetrum was not removed and N was added, Empetrum even increased in abundance at the expense of the subordinate species. Thus, profound changes in tundra ecosystems can only be expected when Empetrum is very heavily damaged as a result of recurrent extreme winter warming and when soil N availability is increased as a result of summer warming. These changes in species composition upon extreme disturbance events may lead to a wide variety of ecosystem feedbacks and cascade processes as this tundra system is relatively species‐poor, and can be hypothesized to have low functional redundancy.     

Decomposition of mountain birch leaf litter at the forest-tundra ecotone in the Fennoscandian mountains in relation to climate and soil conditions

Litter decomposition is a key process in terrestrial ecosystems, releasing nutrients, returning CO₂ to the atmosphere, and contributing to the formation of humus. Litter decomposition is strongly controlled both by climate and by litter quality: global warming scenarios involving shifts in vegetation communities are therefore of particular interest in this context. The objective of the present study was to quantify the role of climatic environment and underlying substrate chemistry for the decomposition of standard mountain birch (Betula pubescens Ehrh. spp. czerepanovii) leaf litter at four sites, spanning the forest-tundra ecotone, in the Fennoscandian mountain range. Litter quality effects were thus held constant, but the study incorporated systematic changes in (i) latitude/altitude, (ii) `continentality', and (iii) vegetation community at each site, together with (iv) experimental manipulation of temperature using passive warming systems. The study was undertaken during a 3 year period, and forms part of a broader investigation of forest-tundra ecotone dynamics in the Fennoscandian mountains. Our results showed (1) higher decomposition rates in forest sites compared to tundra, (2) that the difference between the two vegetation communities was most pronounced at the more maritime sites, and (3) that chemistry of litter remaining after the three years experiment varied according to site and vegetation community (e.g. at the most southerly site, more lignin had decomposed at tundra communities compared with the forest). (4) Surface temperature explained 58% of the variation in mass loss at forest sites; at tundra sites, however, we hypothesise that litter moisture content was the more important factor. (5) Experimental warming lent weight to this hypothesis by reducing rates of mass loss: this reduction was likely the result of surface soil drying, an artefact of the warming treatment. We conclude that a replacement of tundra by forest would likely accelerate litter decomposition both via changes in surface and near-surface temperature and moisture regimes, although the strength of this response will vary between maritime and continental parts of the mountain range.     

Predicted changes in vegetation structure affect the susceptibility to invasion of bryophyte-dominated subarctic heath

Background and Aims

A meta-analysis of global change experiments in arctic tundra sites suggests that plant productivity and the cover of shrubs, grasses and dead plant material (i.e. litter) will increase and the cover of bryophytes will decrease in response to higher air temperatures. However, little is known about which effects these changes in vegetation structure will have on seedling recruitment of species and invasibility of arctic ecosystems.

Methods

A field experiment was done in a bryophyte-dominated, species-rich subarctic heath by manipulating the cover of bryophytes and litter in a factorial design. Three phases of seedling recruitment (seedling emergence, summer seedling survival, first-year recruitment) of the grass Anthoxanthum alpinum and the shrub Betula nana were analysed after they were sown into the experimental plots.

Key Results

Bryophyte and litter removal significantly increased seedling emergence of both species but the effects of manipulations of vegetation structure varied strongly for the later phases of recruitment. Summer survival and first-year recruitment were significantly higher in Anthoxanthum. Although bryophyte removal generally increased summer survival and recruitment, seedlings of Betula showed high mortality in early August on plots where bryophytes had been removed.

Conclusions

Large species-specific variation and significant effects of experimental manipulations on seedling recruitment suggest that changes in vegetation structure as a consequence of global warming will affect the abundance of grasses and shrubs, the species composition and the susceptibility to invasion of subarctic heath vegetation.     

The Circumpolar Arctic vegetation map

Abstract. Question: What are the major vegetation units in the Arctic, what is their composition, and how are they distributed among major bioclimate subzones and countries? Location: The Arctic tundra region, north of the tree line. Methods: A photo‐interpretive approach was used to delineate the vegetation onto an Advanced Very High Resolution Radiometer (AVHRR) base image. Mapping experts within nine Arctic regions prepared draft maps using geographic information technology (ArcInfo) of their portion of the Arctic, and these were later synthesized to make the final map. Area analysis of the map was done according to bioclimate subzones, and country. The integrated mapping procedures resulted in other maps of vegetation, topography, soils, landscapes, lake cover, substrate pH, and above‐ground biomass. Results: The final map was published at 1:7 500 000 scale map. Within the Arctic (total area = 7.11 × 106 km²), about 5.05 × 10⁶ km² is vegetated. The remainder is ice covered. The map legend generally portrays the zonal vegetation within each map polygon. About 26% of the vegetated area is erect shrublands, 18% peaty graminoid tundras, 13% mountain complexes, 12% barrens, 11% mineral graminoid tundras, 11% prostrate‐shrub tundras, and 7% wetlands. Canada has by far the most terrain in the High Arctic mostly associated with abundant barren types and prostrate dwarf‐shrub tundra, whereas Russia has the largest area in the Low Arctic, predominantly low‐shrub tundra. Conclusions: The CAVM is the first vegetation map of an entire global biome at a comparable resolution. The consistent treatment of the vegetation across the circumpolar Arctic, abundant ancillary material, and digital database should promote the application to numerous land‐use, and climate‐change applications and will make updating the map relatively easy.