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.     

Rapid, landscape scale responses in riparian tundra vegetation to exclusion of small and large mammalian herbivores

Productive tundra plant communities composed of a variety of fast growing herbaceous and woody plants are likely to attract mammalian herbivores. Such vegetation is likely to respond to different-sized herbivores more rapidly than currently acknowledged from the tundra. Accentuated by currently changing populations of arctic mammals there is a need to understand impacts of different-sized herbivores on the dynamics of productive tundra plant communities. Here we assess the differential effects of ungulate (reindeer) and small rodent herbivores (voles and lemmings) on high productive tundra vegetation. A spatially extensive exclosure experiment was run for three years on river sediment plains along two river catchments in low-arctic Norway. The river catchments were similar in species pools but differed in species abundance composition of both plants and vertebrate herbivores. Biomass of forbs, deciduous shrubs and silica-poor grasses increased by 40–50% in response to release from herbivory, whereas biomass of silica-rich grasses decreased by 50–75%. Hence both additive and compensatory effects of small rodents and reindeer exclusion caused these significant changes in abundance composition of the plant communities. Changes were also rapid, evident after only one growing season, and are among the fastest and strongest ever documented in Arctic vegetation. The rate of changes indicates a tight link between the dynamics of productive tundra vegetation and both small and large herbivores. Responses were however not spatially consistent, being highly different between the catchments. We conclude that despite similar species pools, variation in plant species abundance and herbivore species dynamics give different prerequisites for change.     

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.     

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.     

Herbivores inhibit climate-driven shrub expansion on the tundra

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana , has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (>0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (<0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.     

Open tundra persist,but arctic features decline—Vegetation changes in the warming Fennoscandian tundra

In the forest‐tundra ecotone of the North Fennoscandian inland, summer and winter temperatures have increased by two to three centigrades since 1965, which is expected to result in major vegetation changes. To document the expected expansion of woodlands and scrublands and its impact on the arctic vegetation, we repeated a vegetation transect study conducted in 1976 in the Darju, spanning from woodland to a summit, 200 m above the tree line. Contrary to our expectations, tree line movement was not detected, and there was no increase in willows or shrubby mountain birches, either. Nevertheless, the stability of tundra was apparent. Small‐sized, poorly competing arctic species had declined, lichen cover had decreased, and vascular plants, especially evergreen ericoid dwarf shrubs, had gained ground. The novel climate seems to favour competitive clonal species and species thriving in closed vegetation, creating a community hostile for seedling establishment, but equally hostile for many arctic species, too. Preventing trees and shrubs from invading the tundra is thus not sufficient for conserving arctic biota in the changing climate. The only dependable cure is to stop the global warming.     

Shrubline but not treeline advance matches climate velocity in montane ecosystems of south‐central Alaska

Tall shrubs and trees are advancing into many tundra and wetland ecosystems but at a rate that often falls short of that predicted due to climate change. For forest, tall shrub, and tundra ecosystems in two pristine mountain ranges of Alaska, we apply a Bayesian, error‐propagated calculation of expected elevational rise (climate velocity), observed rise (biotic velocity), and their difference (biotic inertia). We show a sensitive dependence of climate velocity on lapse rate and derive biotic velocity as a rigid elevational shift. Ecosystem presence identified from recent and historic orthophotos ~50 years apart was regressed on elevation. Biotic velocity was estimated as the difference between critical point elevations of recent and historic logistic fits divided by time between imagery. For both mountain ranges, the 95% highest posterior density of climate velocity enclosed the posterior distributions of all biotic velocities. In the Kenai Mountains, mean tall shrub and climate velocities were both 2.8 m y^?1. In the better sampled Chugach Mountains, mean tundra retreat was 1.2 m y^?1 and climate velocity 1.3 m y^?1. In each mountain range, the posterior mode of tall woody vegetation velocity (the complement of tundra) matched climate velocity better than either forest or tall shrub alone, suggesting competitive compensation can be important. Forest velocity was consistently low at 0.1–1.1 m y^?1, indicating treeline is advancing slowly. We hypothesize that the high biotic inertia of forest ecosystems in south‐central Alaska may be due to competition with tall shrubs and/or more complex climate controls on the elevational limits of trees than tall shrubs. Among tall shrubs, those that disperse farthest had lowest inertia. Finally, the rapid upward advance of woody vegetation may be contributing to regional declines in Dall's sheep (Ovis dalli), a poorly dispersing alpine specialist herbivore with substantial biotic inertia due to dispersal reluctance.     

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.     

Do herbivores cause habitat degradation or vegetation state transition? Evidence from the tundra

Range expansion and increasing densities of large herbivores are held responsible for large-scale habitat degradation in a wide range of natural and semi-natural ecosystems. Herbivore-driven ecosystem changes frequently represent predictable transitions from one vegetation state to another. Whether such predictable changes justify the value judgement 'habitat degradation' may be debatable as this strongly depends on individual perspective.
To further the debate on herbivore-driven habitat degradation, I apply the concept of alternative stable states to arctic tundra as a framework to capture predictable stepwise vegetation transitions in which the productivity and hence herbivore-carrying capacity increases with grazing pressure. Specifically, evidence is provided that large parts of the tundra biome can be in either of three relatively discrete vegetation states and that changes in reindeer/caribou density are responsible for sudden, predictable but often reversible state transitions. From this, it appears that the relatively rapidly emerging vegetation changes do not necessarily equate to habitat degradation, but in many cases reflect predictable vegetation change. Acknowledgement of the existence of predictable state transitions in tundra ecosystems may help to evaluate the observed radical vegetation changes occurring throughout the reindeer/caribou range.     

Reindeer grazing and soil microbial processes in two suboceanic and two subcontinental tundra heaths

In oceanic, nutrient-rich Fennoscandian arctic-alpine tundra heaths, grazing by reindeer has been found to increase herbs and graminoids in relation to dwarf shrubs. In continental lichen heaths in the inland with nutrient-poor conditions, however, slowly decomposable dwarf shrubs are favoured by grazing. According to a hypothesis, by favouring easily decomposing plants in nutrient-rich conditions and slowly decomposing plants in nutrient-poor conditions, herbivory enhances soil nutrient cycling in nutrient-rich and retards it in nutrient-poor areas. We tested this hypothesis by comparing the impact of reindeer grazing on soil C and N mineralization between two oceanic and two continental arctic-alpine tundra heaths.
Although soil respiration and microbial metabolic activity were enhanced by grazing in the suboceanic but not in the subcontinental tundra heaths, gross N mineralization rates were higher in the grazed areas in soils from all study sites, indicating that reindeer grazing leads to increased rates of nutrient cycling in both nutrient-poor and nutrient-rich tundra heaths. Thus, in the subcontinental tundra heaths, the increase in soil N concentrations due to mammalian waste products enhances N mineralization rates, even though the organic C quality is not improved by reindeer grazing. There was some site-specific variation in the strength of the reindeer effects on various microbial processes and soil properties, which can be related to spatial variation in grazing intensity and timing, as these factors in turn affect the nutrient sink strength of the vegetation.     

Plant carbon allocation drives turnover of old soil organic matter in permafrost tundra soils

Carbon cycle feedbacks from permafrost ecosystems are expected to accelerate global climate change. Shifts in vegetation productivity and composition in permafrost regions could influence soil organic carbon (SOC) turnover rates via rhizosphere (root zone) priming effects (RPEs), but these processes are not currently accounted for in model predictions. We use a radiocarbon (bomb‐¹⁴C) approach to test for RPEs in two Arctic tall shrubs, alder (Alnus viridis (Chaix) DC.) and birch (Betula glandulosa Michx.), and in ericaceous heath tundra vegetation. We compare surface CO₂ efflux rates and ¹⁴C content between intact vegetation and plots in which below‐ground allocation of recent photosynthate was prevented by trenching and removal of above‐ground biomass. We show, for the first time, that recent photosynthate drives mineralization of older (>50 years old) SOC under birch shrubs and ericaceous heath tundra. By contrast, we find no evidence of RPEs in soils under alder. This is the first direct evidence from permafrost systems that vegetation influences SOC turnover through below‐ground C allocation. The vulnerability of SOC to decomposition in permafrost systems may therefore be directly linked to vegetation change, such that expansion of birch shrubs across the Arctic could increase decomposition of older SOC. Our results suggest that carbon cycle models that do not include RPEs risk underestimating the carbon cycle feedbacks associated with changing conditions in tundra regions.     

Can Reindeer Overabundance Cause a Trophic Cascade?

Abstract The region Finnmark, in northernmost Europe, harbors dense populations of semi-domestic reindeer of which some exhibit characteristics of overabundance. Whereas overabundance is evident in terms of density-dependent reductions in reindeer body mass, population growth and abundance of forage plants, claims have been made that this reindeer overabundance also has caused a trophic cascade. These claims are based on the main premise that reindeer overgrazing negatively impacts small-sized, keystone tundra herbivores. We tested this premise by a large-scale study in which the abundance of small rodents, hares and ptarmigans was indexed across reindeer management districts with strong differences in stocking densities. We examined the scale-dependent relations between reindeer, vegetation and these small-sized herbivores by employing a spatially hierarchical sampling design within the management districts. A negative impact of reindeer on ptarmigan, probably as a result of browsing reducing tall Salix, was indicated. However, small rodents (voles and lemmings), which are usually the keystone herbivores in the plant-based tundra food web, were not negatively impacted. On the contrary, there was a strong positive relationship between small rodents and reindeer, both at the scale of landscape areas and local patches, with characteristics of snow-bed vegetation, suggesting facilitation between Norwegian lemmings and reindeer. We conclude that the recent dampening of the vole and lemming population cycle with concurrent declines of rodent predators in northernmost Europe were not caused by large herbivore overgrazing.     

Greater deciduous shrub abundance extends tundra peak season and increases modeled net CO2 uptake

Satellite studies of the terrestrial Arctic report increased summer greening and longer overall growing and peak seasons since the 1980s, which increases productivity and the period of carbon uptake. These trends are attributed to increasing air temperatures and reduced snow cover duration in spring and fall. Concurrently, deciduous shrubs are becoming increasingly abundant in tundra landscapes, which may also impact canopy phenology and productivity. Our aim was to determine the influence of greater deciduous shrub abundance on tundra canopy phenology and subsequent impacts on net ecosystem carbon exchange (NEE) during the growing and peak seasons in the arctic foothills region of Alaska. We compared deciduous shrub‐dominated and evergreen/graminoid‐dominated community‐level canopy phenology throughout the growing season using the normalized difference vegetation index (NDVI). We used a tundra plant‐community‐specific leaf area index (LAI) model to estimate LAI throughout the green season and a tundra‐specific NEE model to estimate the impact of greater deciduous shrub abundance and associated shifts in both leaf area and canopy phenology on tundra carbon flux. We found that deciduous shrub canopies reached the onset of peak greenness 13 days earlier and the onset of senescence 3 days earlier compared to evergreen/graminoid canopies, resulting in a 10‐day extension of the peak season. The combined effect of the longer peak season and greater leaf area of deciduous shrub canopies almost tripled the modeled net carbon uptake of deciduous shrub communities compared to evergreen/graminoid communities, while the longer peak season alone resulted in 84% greater carbon uptake in deciduous shrub communities. These results suggest that greater deciduous shrub abundance increases carbon uptake not only due to greater leaf area, but also due to an extension of the period of peak greenness, which extends the period of maximum carbon uptake.     

Changes in the structure and function of northern Alaskan ecosystems when considering variable leaf‐out times across groupings of species in a dynamic vegetation model

The phenology of arctic ecosystems is driven primarily by abiotic forces, with temperature acting as the main determinant of growing season onset and leaf budburst in the spring. However, while the plant species in arctic ecosystems require differing amounts of accumulated heat for leaf‐out, dynamic vegetation models simulated over regional to global scales typically assume some average leaf‐out for all of the species within an ecosystem. Here, we make use of air temperature records and observations of spring leaf phenology collected across dominant groupings of species (dwarf birch shrubs, willow shrubs, other deciduous shrubs, grasses, sedges, and forbs) in arctic and boreal ecosystems in Alaska. We then parameterize a dynamic vegetation model based on these data for four types of tundra ecosystems (heath tundra, shrub tundra, wet sedge tundra, and tussock tundra), as well as ecotonal boreal white spruce forest, and perform model simulations for the years 1970–2100. Over the course of the model simulations, we found changes in ecosystem composition under this new phenology algorithm compared with simulations with the previous phenology algorithm. These changes were the result of the differential timing of leaf‐out, as well as the ability for the groupings of species to compete for nitrogen and light availability. Regionally, there were differences in the trends of the carbon pools and fluxes between the new phenology algorithm and the previous phenology algorithm, although these differences depended on the future climate scenario. These findings indicate the importance of leaf phenology data collection by species and across the various ecosystem types within the highly heterogeneous Arctic landscape, and that dynamic vegetation models should consider variation in leaf‐out by groupings of species within these ecosystems to make more accurate projections of future plant distributions and carbon cycling in Arctic regions.     

Density-dependent switches in diet: a likely mechanism for negative feedbacks on goose population increase?

Goose grazing on arctic tundra vegetation has shown both positive and negative effects on subsequent foraging conditions. To understand the potential of a density-dependent feedback on herbivore population size, the relation between grazing pressure and future foraging conditions is essential. We studied the effect of increasing grazing pressure of barnacle geese (Branta leucopsis) on Spitsbergen. During the establishment of a breeding colony in the period 1992–2004, the proportion of graminoids decreased in the diet of wild geese, while the percentage of mosses increased. Grazing trials with captive geese in an unexploited area showed a similar shift in diet composition. High-quality food plants were depleted within years and over years. Intake rate declined too and as consequence, metabolisable energy intake rate (MEIR) decreased rapidly with increasing grazing pressure. During three successive years of experimental grazing, MEIR decreased at all levels of grazing pressure and declined below minimal energetic requirements when grazing exceeded natural levels of grazing pressure. This suggests that foraging conditions rapidly decline with increasing grazing pressure in these low-productive habitats. The potential for density-dependent feedbacks on local population increase is discussed.     

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.     

The effects of reindeer grazing on the composition and species richness of vegetation in forest–tundra ecotone

We conducted an 8-year exclosure experiment (1999–2006) in a forest–tundra ecotonal area in northwestern Finnish Lapland to study the effects of reindeer grazing on vegetation in habitats of variable productivity and microhabitat structure. The experimental sites included tundra heath, frost heath and riparian habitats, and the two latter habitats were characterized by hummock-hollow ground forms. The total cover of vegetation, cover of willow (Salix spp.), dwarf birch (Betula nana), dwarf shrubs, forbs and grasses (Poaceae spp.) increased in exclosures in all habitats. The increase in the total cover of vegetation and in the covers of willow and dwarf birch tended to be greatest in the least productive tundra heath. Opposing to the increase in the dominant vascular plant groups, the cover and species number of bryophytes decreased in exclosures. We conclude that the effects of reindeer grazing on vegetation composition depend on environmental heterogeneity and the responses vary among plant groups. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Arctic arthropod assemblages in habitats of differing shrub dominance

Recent climate warming in the Arctic has caused advancement in the timing of snowmelt and expansion of shrubs into open tundra. Such an altered climate may directly and indirectly (via effects on vegetation) affect arctic arthropod abundance, diversity and assemblage taxonomic composition. To allow better predictions about how climate changes may affect these organisms, we compared arthropod assemblages between open and shrub‐dominated tundra at three field sites in northern Alaska that encompass a range of shrub communities. Over ten weeks of sampling in 2011, pitfall traps captured significantly more arthropods in shrub plots than open tundra plots at two of the three sites. Furthermore, taxonomic richness and diversity were significantly greater in shrub plots than open tundra plots, although this pattern was site‐specific as well. Patterns of abundance within the five most abundant arthropod orders differed, with spiders (Order: Araneae) more abundant in open tundra habitats and true bugs (Order: Hemiptera), flies (Order: Diptera), and wasps and bees (Order: Hymenoptera) more abundant in shrub‐dominated habitats. Few strong relationships were found between vegetation and environmental variables and arthropod abundance; however, lichen cover seemed to be important for the overall abundance of arthropods. Some arthropod orders showed significant relationships with other vegetation variables, including maximum shrub height (Coleoptera) and foliar canopy cover (Diptera). As climate warming continues over the coming decades, and with further shrub expansion likely to occur, changes in arthropod abundance, richness, and diversity associated with shrub‐dominated habitat may have important ecological effects on arctic food webs since arthropods play important ecological roles in the tundra, including in decomposition and trophic interactions.     

Bird Communities of the Arctic Shrub Tundra of Yamal: Habitat Specialists and Generalists

Background

The ratio of habitat generalists to specialists in birds has been suggested as a good indicator of ecosystem changes due to e.g. climate change and other anthropogenic perturbations. Most studies focusing on this functional component of biodiversity originate, however, from temperate regions. The Eurasian Arctic tundra is currently experiencing an unprecedented combination of climate change, change in grazing pressure by domestic reindeer and growing human activity.

Methodology/Principal Findings

Here we monitored bird communities in a tundra landscape harbouring shrub and open habitats in order to analyse bird habitat relationships and quantify habitat specialization. We used ordination methods to analyse habitat associations and estimated the proportions of specialists in each of the main habitats. Correspondence Analysis identified three main bird communities, inhabiting upland, lowland and dense willow shrubs. We documented a stable structure of communities despite large multiannual variations of bird density (from 90 to 175 pairs/km²). Willow shrub thickets were a hotspot for bird density, but not for species richness. The thickets hosted many specialized species whose main distribution area was south of the tundra.

Conclusion/Significance

If current arctic changes result in a shrubification of the landscape as many studies suggested, we would expect an increase in the overall bird abundance together with an increase of local specialists, since they are associated with willow thickets. The majority of these species have a southern origin and their increase in abundance would represent a strengthening of the boreal component in the southern tundra, perhaps at the expense of species typical of the subarctic zone, which appear to be generalists within this zone.     

Where do the treeless tundra areas of northern highlands fit in the global biome system: toward an ecologically natural subdivision of the tundra biome

According to some treatises, arctic and alpine sub‐biomes are ecologically similar, whereas others find them highly dissimilar. Most peculiarly, large areas of northern tundra highlands fall outside of the two recent subdivisions of the tundra biome. We seek an ecologically natural resolution to this long‐standing and far‐reaching problem. We studied broad‐scale patterns in climate and vegetation along the gradient from Siberian tundra via northernmost Fennoscandia to the alpine habitats of European middle‐latitude mountains, as well as explored those patterns within Fennoscandian tundra based on climate–vegetation patterns obtained from a fine‐scale vegetation map. Our analyses reveal that ecologically meaningful January–February snow and thermal conditions differ between different types of tundra. High precipitation and mild winter temperatures prevail on middle‐latitude mountains, low precipitation and usually cold winters prevail on high‐latitude tundra, and Scandinavian mountains show intermediate conditions. Similarly, heath‐like plant communities differ clearly between middle latitude mountains (alpine) and high‐latitude tundra vegetation, including its altitudinal extension on Scandinavian mountains. Conversely, high abundance of snowbeds and large differences in the composition of dwarf shrub heaths distinguish the Scandinavian mountain tundra from its counterparts in Russia and the north Fennoscandian inland. The European tundra areas fall into three ecologically rather homogeneous categories: the arctic tundra, the oroarctic tundra of northern heights and mountains, and the genuinely alpine tundra of middle‐latitude mountains. Attempts to divide the tundra into two sub‐biomes have resulted in major discrepancies and confusions, as the oroarctic areas are included in the arctic tundra in some biogeographic maps and in the alpine tundra in others. Our analyses based on climate and vegetation criteria thus seem to resolve the long‐standing biome delimitation problem, help in consistent characterization of research sites, and create a basis for further biogeographic and ecological research in global tundra environments.