Shrub shading moderates the effects of weather on arthropod activity in arctic tundra

1. Rapid warming has facilitated an increase in deciduous shrub cover in arctic tundra. Because shrubs create a cooler microclimate during the growing season, shrub cover could modulate the effects of global warming on the phenology and activity of ectotherms, including arthropods. This possibility was explored here using two dominant arthropod groups (flies and wolf spiders) in Alaskan tundra. 2. We monitored arthropods with pitfall traps over five summers at four sites that differed in shrub abundance, and used generalised additive mixed models (GAMMs) to separate the two underlying components of pitfall trap catch: the seasonal trend in arthropod density and the effects of short‐term weather variation (air temperature, wind speed, rainfall, solar radiation) on arthropod activity. 3. We found that shrub cover significantly altered the seasonal trend in the abundance of flies by reducing early‐season pitfall catch, in line with observed later snowmelt in shrub‐dominated plots at these sites. 4. Additionally, shrub cover modulated the effects of many weather variables on arthropod activity: shrub cover shifted wolf spiders' temperature–activity relationship, dampened the positive effect of solar radiation on the activity of arthropods in total, and ameliorated the negative effect of wind on the activity of flies. 5. Thus, these results indicate that shrub encroachment will probably be accompanied by altered arthropod responses to warming and other key weather variables. Because the rate of key ecological processes – herbivory, decomposition, predation – are controlled by activity at the organismal level, these effects on arthropods will have long‐term ecosystem‐level consequences.     

Variable sensitivity of plant communities in Iceland to experimental warming

Facing an increased threat of rapid climate change in cold‐climate regions, it is important to understand the sensitivity of plant communities both in terms of degree and direction of community change. We studied responses to 3–5 years of moderate experimental warming by open‐top chambers in two widespread but contrasting tundra communities in Iceland. In a species‐poor and nutrient‐deficient moss heath, dominated by Racomitrium lanuginosum, mean daily air temperatures at surface were 1–2°C higher in the warmed plots than the controls whereas soil temperatures tended to be lower in the warmed plots throughout the season. In a species‐rich dwarf shrub heath on relatively rich soils at a cooler site, dominated by Betula nana and R. lanuginosum, temperature changes were in the same direction although more moderate. In the moss heath, there were no detectable community changes while significant changes were detected in the dwarf shrub heath: the abundance of deciduous and evergreen dwarf shrubs significantly increased (>50%), bryophytes decreased (18%) and canopy height increased (100%). Contrary to some other studies of tundra communities, we detected no changes in species richness or other diversity measures in either community and the abundance of lichens did not change. It is concluded that the sensitivity of Icelandic tundra communities to climate warming varies greatly depending on initial conditions in terms of species diversity, dominant species, soil and climatic conditions as well as land‐use history.     

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.     

Shrubs and species identity effects on the distribution and diversity of ground-dwelling arthropods in a Gobi desert

Disentangling the relationship between shrub vegetation and ground-dwelling arthropods at multiple levels of taxonomic resolution is essential to developing a suitable management strategy for the conservation of shrub-associated arthropod biodiversity in shrubland ecosystems. Using Gobi desert (dominated by shrub species Nitraria sphaerocarpa and Reaumuria soongorica) occurring widely in inland arid areas of northwest China as a model system, we sampled ground-dwelling arthropods by pitfall trapping method under canopies of both shrubs and in intershrub bare areas during spring, summer and autumn. Our aim was to determine whether the presence and species identity of shrubs influence the distribution of ground-dwelling arthropods and whether the influence of shrub presence and species identity differs among trophic and taxonomic groups. At the community level, total arthropod abundance and species richness were significantly greater under shrubs than in intershrub bare habitats, whereas more arthropods were captured under N. sphaerocarpa than under R. soongorica. At the trophic group level, the abundance of predator and decomposer arthropods was significantly greater under shrubs than in intershrub bare habitats, whereas herbivore and decomposer arthropods were more abundant under N. sphaerocarpa than under R. soongorica. At the family level, the abundance of Carabidae, Gnaphosidae, Karschiidae, Tenebrionidae and Thomisidae was consistently much greater under shrubs than in intershrub bare habitats, whereas that of Formicidae, Philodromidae and Tettigoniidae did not differ between the shrub microhabitats and the intershrub bare habitats. However, the abundance of Curculionidae under R. soongorica and the abundance of Geotrupidae under N. sphaerocarpa were significantly lower than that in intershrub bare habitats. N. sphaerocarpa was commonly preferred by arthropods in the families of Curculionidae and Tenebrionidae, whereas R. soongorica was commonly preferred by arthropods in the family Geotrupidae, but shrub species identity did not affect the abundance of the remaining families. Our results suggest that shrubs and species identity play important roles in structuring ground-dwelling arthropod communities, but the response of arthropods differed among trophic or taxonomic groups. This study may have important implications for the conservation of invertebrate biodiversity in Gobi desert ecosystems.     

Summer temperature increase has distinct effects on the ectomycorrhizal fungal communities of moist tussock and dry tundra in Arctic Alaska

Arctic regions are experiencing the greatest rates of climate warming on the planet and marked changes have already been observed in terrestrial arctic ecosystems. While most studies have focused on the effects of warming on arctic vegetation and nutrient cycling, little is known about how belowground communities, such as fungi root‐associated, respond to warming. Here, we investigate how long‐term summer warming affects ectomycorrhizal (ECM) fungal communities. We used Ion Torrent sequencing of the rDNA internal transcribed spacer 2 (ITS2) region to compare ECM fungal communities in plots with and without long‐term experimental warming in both dry and moist tussock tundra. Cortinarius was the most OTU‐rich genus in the moist tundra, while the most diverse genus in the dry tundra was Tomentella. On the diversity level, in the moist tundra we found significant differences in community composition, and a sharp decrease in the richness of ECM fungi due to warming. On the functional level, our results indicate that warming induces shifts in the extramatrical properties of the communities, where the species with medium‐distance exploration type seem to be favored with potential implications for the mobilization of different nutrient pools in the soil. In the dry tundra, neither community richness nor community composition was significantly altered by warming, similar to what had been observed in ECM host plants. There was, however, a marginally significant increase in OTUs identified as ECM fungi with the medium‐distance exploration type in the warmed plots. Linking our findings of decreasing richness with previous results of increasing ECM fungal biomass suggests that certain ECM species are favored by warming and may become more abundant, while many other species may go locally extinct due to direct or indirect effects of warming. Such compositional shifts in the community might affect nutrient cycling and soil organic C storage.     

Impact of the invasive shrub Amur honeysuckle (Lonicera maackii) on shrub-layer insects in a deciduous forest in the eastern United States

Invasive Amur honeysuckle (Lonicera maackii) creates a dense shrub layer in deciduous forests in eastern North America that negatively impacts native herbs and tree seedlings. We predicted that higher vegetative cover caused by this shrub would increase abundance and diversity of insects and alter composition of insect assemblages. We used paired plots, one with and one without honeysuckle, in ten forest locations in southwestern Ohio, USA, to sample insects and measure diversity and vertical cover of vegetation in the shrub layer. Vertical cover of this vegetation was higher on honeysuckle-present plots, but diversity of shrub-layer vegetation did not differ between honeysuckle-present and honeysuckle-absent plots. Species diversity of Hexapoda, Coleoptera, and Psocoptera, richness of Hexapoda, Coleoptera, Diptera, Hymenoptera, and Psocoptera, and abundance of Hexapoda, Diptera, Hymenoptera, and Psocoptera were higher on honeysuckle-present than on honeysuckle-absent plots. Evenness did not differ between honeysuckle treatments. Nonmetric multidimensional scaling distinguished taxonomic composition in honeysuckle-present plots from that in honeysuckle-absent plots. Presence of vertical cover explained higher richness of Hexapoda and Coleoptera, and higher abundance of Hexapoda, Diptera, Hymenoptera, and Psocoptera. Attributes of honeysuckle, independent of its contribution to vertical cover, explained increases in richness of Hexapoda, Coleoptera, and Hymenoptera and abundance of Hexapoda, Hymenopera, and Psocoptera. These attributes of honeysuckle could include a more complex vegetative structure, a greater availability of resources such as food, detritus, or shelter, and/or a more favorable cooler and moister microenvironment. To more fully understand the mechanisms causing increased richness and abundance of insects in honeysuckle-present areas, detailed studies on these attributes of honeysuckle would be necessary.     

Seasonal Distribution and Diversity of Ground Arthropods in Microhabitats Following a Shrub Plantation Age Sequence in Desertified Steppe

In desertified regions, shrub-dominated patches are important microhabitats for ground arthropod assemblages. As shrub age increases, soil, vegetation and microbiological properties can change remarkably and spontaneously across seasons. However, relatively few studies have analyzed how ground arthropods respond to the microhabitats created by shrubs of different plantation ages across seasons. Using 6, 15, 24 and 36 year-old plantations of re-vegetated shrubs (Caragana koushinskii) in the desert steppe of northwestern China as a model system, we sampled ground arthropod communities using a pitfall trapping method in the microhabitats under shrubs and in the open areas between shrubs, during the spring, summer and autumn. The total ground arthropod assemblage was dominated by Carabidae, Melolonthidae, Curculionidae, Tenebrionidae and Formicidae that were affected by plantation age, seasonal changes, or the interaction between these factors, with the later two groups also influenced by microhabitat. Overall, a facilitative effect was observed, with more arthropods and a greater diversity found under shrubs as compared to open areas, but this was markedly affected by seasonal changes. There was a high degree of similarity in arthropod assemblages and diversity between microhabitats in summer and autumn. Shrub plantation age significantly influenced the distribution of the most abundant groups, and also the diversity indices of the ground arthropods. However, there was not an overall positive relationship between shrub age and arthropod abundance, richness or diversity index. The influence of plantation age on arthropod communities was also affected by seasonal changes. From spring through summer to autumn, community indices of ground arthropods tended to decline, and a high degree of similarity in these indices (with fluctuation) was observed among different ages of shrub plantation in autumn. Altogether the recovery of arthropod communities was markedly affected by seasonal variability, and they demonstrated distinctive communal fingerprints in different microhabitats for each plantation age stage.     

Long‐term nutrient addition alters arthropod community composition but does not increase total biomass or abundance

A simple bottom–up hypothesis predicts that plant responses to nutrient addition should determine the response of consumers: more productive and less diverse plant communities, the usual result of long‐term nutrient addition, should support greater consumer abundances and biomass and less consumer diversity. We tested this hypothesis for the response of an aboveground arthropod community to an uncommonly long‐term (24‐year) nutrient addition experiment in moist acidic tundra in arctic Alaska. This experiment altered plant community composition, decreased plant diversity and increased plant production and biomass as a deciduous shrub, Betula nana, became dominant. Consistent with strong effects on the plant community, nutrient addition altered arthropod community composition, primarily through changes to herbivore taxa in the canopy‐dwelling arthropod assemblage and detritivore taxa in the ground assemblage. Surprisingly, however, the loss of more than half of plant species was accompanied by negligible changes to diversity (rarefied richness) of arthropod taxa (which were primarily identified to family). Similarly, although long‐term nutrient addition in this system roughly doubles plant production and biomass, arthropod abundance was either unchanged or decreased by nutrient addition, and total arthropod biomass was unaffected. Our findings differ markedly from the handful of terrestrial studies that have found bottom‐up diversity cascades and productivity responses by consumers to nutrient addition. This is probably because unlike grasslands and salt marshes (where such studies have historically been conducted), this arctic tundra community becomes less palatable, rather than more so, after many years of nutrient addition due to increased dominance of B. nana. Additionally, by displacing insulating mosses and increasing the cover of shrubs that cool and shade the canopy microenvironment, fertilization may displace arthropods keenly attuned to microclimate. These results indicate that terrestrial arthropod assemblages may be more constrained by producer traits (i.e. palatability, structure) than they are by total primary production or producer diversity.     

Productivity–diversity patterns in arctic tundra vegetation

Productivity has long been argued to be a major driver of species richness patterns. In the present study we test alternative productivity–diversity hypotheses using vegetation data from the vast Eurasian tundra. The productivity–species pool hypothesis predicts positive relationships at both fine and coarse grain sizes, whereas the productivity–interaction hypothesis predicts unimodal patterns at fine grain size, and monotonic positive patterns at coarse grain size. We furthermore expect to find flatter positive (productivity–species pool hypothesis) or more strongly negative (productivity–interaction hypothesis) relationships for lichens and bryophytes than for vascular plants, because as a group, lichens and bryophytes are better adapted to extreme arctic conditions and more vulnerable to competition for light than the taller‐growing vascular plants. The normalised difference vegetation index (NDVI) was used as a proxy of productivity. The generally unimodal productivity–diversity patterns were most consistent with the productivity–interaction hypothesis. There was a general trend of decreasing species richness from moderately to maximally productive tundra, in agreement with an increasing importance of competitive interactions. High richness of vascular plants and lichens occurred in moderately low productive tundra areas, whereas that of bryophytes occurred in the least productive tundra habitats covered by this study. The fine and coarse grain richness trends were surprisingly uniform and no variation in beta diversity along the productivity gradient was seen for vascular plants or bryophytes. However, lichen beta diversity varied along the productivity gradient, probably reflecting their sensitivity to habitat conditions and biotic interactions. Overall, the results show evidence that productivity–diversity gradients exist in tundra and that these appear to be largely driven by competitive interactions. Our results also imply that climate warming‐driven increases in productivity will strongly affect arctic plant diversity patterns.     

The nature of spatial transitions in the Arctic

Aim Describe the spatial and temporal properties of transitions in the Arctic and develop a conceptual understanding of the nature of these spatial transitions in the face of directional environmental change. Location Arctic tundra ecosystems of the North Slope of Alaska and the tundra‐forest region of the Seward Peninsula, Alaska Methods We synthesize information from numerous studies on tundra and treeline ecosystems in an effort to document the spatial changes that occur across four arctic transitions. These transitions are: (i) the transition between High‐Arctic and Low‐Arctic systems, (ii) the transition between moist non‐acidic tundra (MNT) and moist acidic tundra (MAT, also referred to as tussock tundra), (iii) the transition between tussock tundra and shrub tundra, (iv) the transition between tundra and forested systems. By documenting the nature of these spatial transitions, in terms of their environmental controls and vegetation patterns, we develop a conceptual model of temporal dynamics of arctic ecotones in response to environmental change. Results Our observations suggest that each transition is sensitive to a unique combination of controlling factors. The transition between High and Low Arctic is sensitive primarily to climate, whereas the MNT/MAT transition is also controlled by soil parent material, permafrost and hydrology. The tussock/shrub tundra transition appears to be responsive to several factors, including climate, topography and hydrology. Finally, the tundra/forest boundary responds primarily to climate and to climatically associated changes in permafrost. There were also important differences in the demography and distribution of the dominant plant species across the four vegetation transitions. The shrubs that characterize the tussock/shrub transition can achieve dominance potentially within a decade, whereas spruce trees often require several decades to centuries to achieve dominance within tundra, and Sphagnum moss colonization of non‐acidic sites at the MNT/MAT boundary may require centuries to millennia of soil development. Main conclusions We suggest that vegetation will respond most rapidly to climatic change when (i) the vegetation transition correlates more strongly with climate than with other environmental variables, (ii) dominant species exhibit gradual changes in abundance across spatial transitions, and/or (iii) the dominant species have demographic properties that allow rapid increases in abundance following climatic shifts. All three of these properties characterize the transition between tussock tundra and low shrub tundra. It is therefore not surprising that of the four transitions studied this is the one that appears to be responding most rapidly to climatic warming.     

Long‐term experimental warming alters community composition of ascomycetes in Alaskan moist and dry arctic tundra

Arctic tundra regions have been responding to global warming with visible changes in plant community composition, including expansion of shrubs and declines in lichens and bryophytes. Even though it is well known that the majority of arctic plants are associated with their symbiotic fungi, how fungal community composition will be different with climate warming remains largely unknown. In this study, we addressed the effects of long‐term (18 years) experimental warming on the community composition and taxonomic richness of soil ascomycetes in dry and moist tundra types. Using deep Ion Torrent sequencing, we quantified how OTU assemblage and richness of different orders of Ascomycota changed in response to summer warming. Experimental warming significantly altered ascomycete communities with stronger responses observed in the moist tundra compared with dry tundra. The proportion of several lichenized and moss‐associated fungi decreased with warming, while the proportion of several plant and insect pathogens and saprotrophic species was higher in the warming treatment. The observed alterations in both taxonomic and ecological groups of ascomycetes are discussed in relation to previously reported warming‐induced shifts in arctic plant communities, including decline in lichens and bryophytes and increase in coverage and biomass of shrubs.     

Greater shrub dominance alters breeding habitat and food resources for migratory songbirds in Alaskan arctic tundra

Climate warming is affecting the Arctic in multiple ways, including via increased dominance of deciduous shrubs. Although many studies have focused on how this vegetation shift is altering nutrient cycling and energy balance, few have explicitly considered effects on tundra fauna, such as the millions of migratory songbirds that breed in northern regions every year. To understand how increasing deciduous shrub dominance may alter breeding songbird habitat, we quantified vegetation and arthropod community characteristics in both graminoid and shrub dominated tundra. We combined measurements of preferred nest site characteristics for Lapland longspurs (Calcarius lapponicus) and Gambel's White‐crowned sparrows (Zonotrichia leucophrys gambelii) with modeled predictions for the distribution of plant community types in the Alaskan arctic foothills region for the year 2050. Lapland longspur nests were found in sedge‐dominated tussock tundra where shrub height does not exceed 20 cm, whereas White‐crowned sparrows nested only under shrubs between 20 cm and 1 m in height, with no preference for shrub species. Shrub canopies had higher canopy‐dwelling arthropod availability (i.e. small flies and spiders) but lower ground‐dwelling arthropod availability (i.e. large spiders and beetles). Since flies are the birds' preferred prey, increasing shrubs may result in a net enhancement in preferred prey availability. Acknowledging the coarse resolution of existing tundra vegetation models, we predict that by 2050 there will be a northward shift in current White‐crowned sparrow habitat range and a 20–60% increase in their preferred habitat extent, while Lapland longspur habitat extent will be equivalently reduced. Our findings can be used to make first approximations of future habitat change for species with similar nesting requirements. However, we contend that as exemplified by this study's findings, existing tundra modeling tools cannot yet simulate the fine‐scale habitat characteristics that are critical to accurately predicting future habitat extent for many wildlife species.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Does Coastal Foredune Stabilization with Ammophila arenaria Restore Plant and Arthropod Communities in Southeastern Australia?

In this study we examine whether stabilization of denuded coastal foredunes in southeastern Australia with the exotic grass species Ammophila arenaria (marram grass) restores plant and ground‐active arthropod assemblages characteristic of undisturbed foredunes. Vascular plants and arthropods were sampled from foredunes that had been stabilized with marram grass in 1982, and from foredunes with no obvious anthropogenic disturbance (control dunes). All arthropods collected were sorted to Order, and ants (81.5% of all specimens) were further sorted to morphospecies. Abundance within arthropod Orders, as well as richness, composition, and structure of the plant and ant assemblages from control and stabilized dunes, were compared. The abundance of Diptera was significantly greater on stabilized dunes, while the abundance of Isopoda was significantly greater on control dunes. There were no significant differences in morphospecies richness or composition of ant assemblages on the two dunes types, although some differences in the abundances of individual morphospecies were observed. By contrast, stabilized dunes exhibited lower plant species richness and highly significant differences in plant species composition, due mainly to the large projected foliage cover of marram grass. The study revealed that after 12 years, the vegetation composition and structure of stabilized dunes was still dominated by marram grass and, as a result, invertebrate assemblages had not been restored to those characteristic of undisturbed foredunes.     

Avian taxonomic and functional diversity in early stage of longleaf pine (Pinus palustris) stands restored at agricultural lands: variations in scale dependency

In agricultural landscapes, the longleaf pine initiative (LLPI) and the Bobwhite Quail Initiative (BQI) aim to restore longleaf pine forests and early successional habitats, respectively. The early stage of longleaf pine stands and grass and forb vegetation produced by a combination of both restoration programs (LLPI‐BQI) may form habitat conditions favorable to early successional bird species and other birds, increasing avian diversity. We investigated how the LLPI and BQI programs affected taxonomic and functional diversity of birds and abundance of early successional birds (grassland and scrub/shrub species), and what environmental characteristics were associated with the diversity and abundance of birds. Our study was performed at 41 fields in Georgia, United States, during 2001–2002 by considering environmental characteristics at two spatial scales: local‐scale vegetation features and restoration program type (LLPI or LLPI‐BQI) and landscape‐scale vegetation features and landscape heterogeneity. Functional evenness, species richness, and abundance of grassland and scrub/shrub species did not show a clear association with local‐ or landscape‐scale variables. Shannon‐Wiener diversity was slightly influenced by restoration program type (local‐scale variable) with higher value at LLPI‐BQI stands than at LLPI stands despite no significant differences in local vegetation features between those stands. Functional divergence was strongly positively associated with landscape‐scale variables. That is, niche differentiation increased with increasing shrub coverage within a landscape, reducing competition between abundant bird species and others. Our results suggest that although a combination of BQI and LLPI program may have a positive effect on avian taxonomic diversity, it is important to consider shrub vegetation cover within a landscape to improve functional diversity.     

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.     

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.     

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.     

Elevation and experimental snowmelt manipulation affect emergence phenology and abundance of soil‐hibernating arthropods

1. Effects of climate change, such as higher average temperatures and earlier snowmelt, are already apparent, especially in alpine regions. However, community responses of functionally important arthropod taxa to changing climatic conditions are mostly unknown. 2. In this study, an earlier snowmelt was simulated at 15 plots along an elevational gradient in the German Alps. At each study site, soil emergence traps were established for sampling soil‐hibernating arthropods on earlier and control snowmelt treatments during the growing season. The abundance and emergence phenology of the five most common arthropod orders (Araneae, Coleoptera, Diptera, Hemiptera, Hymenoptera) were analysed, as well as the species richness of Coleoptera. 3. There was increasing abundance and species richness of Coleoptera along the elevational gradient, indicating that at higher altitudes more individuals and species hibernate in the soil. Abundances of Diptera also increased with elevation. By contrast, abundances of Hemiptera declined with increasing elevation, while abundances of Araneae and Hymenoptera did not show significant elevational patterns. Arthropods at higher elevations emerged, on average, 5 weeks later than arthropods at lower elevations, because of a longer‐lasting snow cover. The earlier snowmelt treatment resulted in higher abundances of Araneae and Hymenoptera compared with the control plots, indicating that the time of snowmelt influenced the abundance of predators, such as spiders or parasitic wasps, more than that of herbivores. 4. An earlier emergence of certain arthropod guilds and a change in relative abundance of guilds might desynchronise species interactions, leading to a possible loss of biodiversity.     

Arctic warming on two continents has consistent negative effects on lichen diversity and mixed effects on bryophyte diversity

Little is known about the impact of changing temperature regimes on composition and diversity of cryptogam communities in the Arctic and Subarctic, despite the well‐known importance of lichens and bryophytes to the functioning and climate feedbacks of northern ecosystems. We investigated changes in diversity and abundance of lichens and bryophytes within long‐term (9–16 years) warming experiments and along natural climatic gradients, ranging from Swedish subarctic birch forest and subarctic/subalpine tundra to Alaskan arctic tussock tundra. In both Sweden and Alaska, lichen diversity responded negatively to experimental warming (with the exception of a birch forest) and to higher temperatures along climatic gradients. Bryophytes were less sensitive to experimental warming than lichens, but depending on the length of the gradient, bryophyte diversity decreased both with increasing temperatures and at extremely low temperatures. Among bryophytes, Sphagnum mosses were particularly resistant to experimental warming in terms of both abundance and diversity. Temperature, on both continents, was the main driver of species composition within experiments and along gradients, with the exception of the Swedish subarctic birch forest where amount of litter constituted the best explanatory variable. In a warming experiment in moist acidic tussock tundra in Alaska, temperature together with soil ammonium availability were the most important factors influencing species composition. Overall, dwarf shrub abundance (deciduous and evergreen) was positively related to warming but so were the bryophytes Sphagnum girgensohnii, Hylocomium splendens and Pleurozium schreberi; the majority of other cryptogams showed a negative relationship to warming. This unique combination of intercontinental comparison, natural gradient studies and experimental studies shows that cryptogam diversity and abundance, especially within lichens, is likely to decrease under arctic climate warming. Given the many ecosystem processes affected by cryptogams in high latitudes (e.g. carbon sequestration, N₂‐fixation, trophic interactions), these changes will have important feedback consequences for ecosystem functions and climate.