Inter-annual variability of NDVI in response to long-term warming and fertilization in wet sedge and tussock tundra

This study explores the relationship between the normalized difference vegetation index (NDVI) and aboveground plant biomass for tussock tundra vegetation and compares it to a previously established NDVI–biomass relationship for wet sedge tundra vegetation. In addition, we explore inter-annual variation in NDVI in both these contrasting vegetation communities. All measurements were taken across long-term experimental treatments in wet sedge and tussock tundra communities at the Toolik Lake Long Term Ecological Research (LTER) site, in northern Alaska. Over 15 years (for wet sedge tundra) and 14 years (for tussock tundra), N and P were applied in factorial experiments (N, P and N+P), air temperature was increased using greenhouses with and without N+P fertilizer, and light intensity was reduced by 50% using shade cloth. during the peak growing seasons of 2001, 2002, and 2003, NDVI measurements were made in both the wet sedge and tussock tundra experimental treatment plots, creating a 3-year time series of inter-annual variation in NDVI. We found that: (1) across all tussock experimental tundra treatments, NDVI is correlated with aboveground plant biomass (r ²=0.59); (2) NDVI–biomass relationships for tussock and wet sedge tundra communities are community specific, and; (3) NDVI values for tussock tundra communities are typically, but not always, greater than for wet sedge tundra communities across all experimental treatments. We suggest that differences between the response of wet sedge and tussock tundra communities in the same experimental treatments result from the contrasting degree of heterogeneity in species and functional types that characterize each of these Arctic tundra vegetation communities.     

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.     

Above‐ and below‐ground plant biomass response to experimental warming in northern Alaska

Question: Does experimental warming, designed to simulate future warming of the Arctic, change the biomass allocation and mycorrhizal infection of tundra plants? Location: High Arctic tundra near Barrow, Alaska, USA (71°18′N 156°40′W). Methods: Above and below ground plant biomass of all species was harvested following 3–4 yr of 1‐2°C of experimental warming. Biomass allocation and arbuscular mycorrhizal infection were also examined in the two dominant species, Salix rotundifolia and Carex aquatilis. Results: Above‐ground biomass of graminoids increased in response to warming but there was no difference in total plant biomass or the ratio of above‐ground to below‐ground biomass for the community as a whole. Carex aquatilis increased above‐ground biomass and proportionally allocated more biomass above ground in response to warming. Salix rotundifolia increased the amount of above‐ and below‐ground biomass allocated per leaf in response to warming. Mycorrhizal infection rates showed no direct response to warming, but total abundance was estimated to have likely increased in response to warming owing to increased root biomass of S. rotundifolia. Conclusions: The community as a whole was resistant to short‐term warming and showed no significant changes in above‐ or below‐ground biomass despite significant increases in above‐ground biomass of graminoids. However, the patterns of biomass allocation for C. aquatilis and S. rotundifolia did change with warming. This suggests that long‐term warming may result in changes in the above‐ground to below‐ground biomass ratio of the community.     

Mammalian herbivores confer resilience of Arctic shrub‐dominated ecosystems to changing climate

Climate change is resulting in a rapid expansion of shrubs in the Arctic. This expansion has been shown to be reinforced by positive feedbacks, and it could thus set the ecosystem on a trajectory toward an alternate, more productive regime. Herbivores, on the other hand, are known to counteract the effects of simultaneous climate warming on shrub biomass. However, little is known about the impact of herbivores on resilience of these ecosystems, that is, the capacity of a system to absorb disturbance and still remain in the same regime, retaining the same function, structure, and feedbacks. Here, we investigated how herbivores affect resilience of shrub‐dominated systems to warming by studying the change of shrub biomass after a cessation of long‐term experimental warming in a forest–tundra ecotone. As predicted, warming increased the biomass of shrubs, and in the absence of herbivores, shrub biomass in tundra continued to increase 4 years after cessation of the artificial warming, indicating that positive effects of warming on plant growth may persist even over a subsequent colder period. Herbivores contributed to the resilience of these systems by returning them back to the original low‐biomass regime in both forest and tundra habitats. These results support the prediction that higher shrub biomass triggers positive feedbacks on soil processes and microclimate, which enable maintaining the rapid shrub growth even in colder climates. Furthermore, the results show that in our system, herbivores facilitate the resilience of shrub‐dominated ecosystems to climate warming.     

Long‐term experimental manipulation of climate alters the ectomycorrhizal community of Betula nana in Arctic tundra

Climate warming is leading to shrub expansion in Arctic tundra. Shrubs form ectomycorrhizal (ECM) associations with soil fungi that are central to ecosystem carbon balance as determinants of plant community structure and as decomposers of soil organic matter. To assess potential climate change impacts on ECM communities, we analysed fungal internal transcribed spacer sequences from ECM root tips of the dominant tundra shrub Betula nana growing in treatments plots that had received long‐term warming by greenhouses and/or fertilization as part of the Arctic Long‐Term Ecological Research experiment at Toolik Lake Alaska, USA. We demonstrate opposing effects of long‐term warming and fertilization treatments on ECM fungal diversity; with warming increasing and fertilization reducing the diversity of ECM communities. We show that warming leads to a significant increase in high biomass fungi with proteolytic capacity, especially Cortinarius spp., and a reduction of fungi with high affinities for labile N, especially Russula spp. In contrast, fertilization treatments led to relatively small changes in the composition of the ECM community, but increased the abundance of saprotrophs. Our data suggest that warming profoundly alters nutrient cycling in tundra, and may facilitate the expansion of B. nana through the formation of mycorrhizal networks of larger size.     

Plant–herbivore interactions in Alaskan arctic tundra change with soil nutrient availability

Herbivores in nutrient‐limited systems such as arctic tundra have been suggested to play a minor role in controlling plant growth simply because they are relatively few in number. However, theory predicts that as net primary productivity (NPP) increases because of greater inputs of nutrients or energy, herbivores may have greater effects on plant growth. This prediction has not been tested in the context of climate warming in arctic tundra, which may increase soil nutrient availability and thus NPP. We examined a long‐term experiment that excluded small and large mammalian herbivores and increased soil nutrients in two arctic Alaskan tundra communities: dry heath (DH) and moist acidic tussock (MAT). In the ninth year of manipulations, we measured weekly growth of three plant species of three growth forms: tussock‐forming graminoid, rhizomatous graminoid, and dwarf deciduous shrub, in each community. All species grew better when fertilized. In DH, this increase in growth was exaggerated when plants were protected from herbivores, confirming that herbivory had a negative effect on plant growth under increased nutrient conditions, but was unimportant under ambient soil conditions. However, in MAT, the importance of herbivory differed among species with fertilization. The tussock‐forming sedge at MAT, Eriophorum vaginatum, grew better and flowered more when fenced under both ambient and amended nutrients compared to plants exposed to herbivores. This species decreases in abundance in long‐term fertilized plots when mammals are present, and our results suggest that herbivory may be accounting for at least some of that loss, in addition to shifts in competitive relationships. Although we only focused on individual plants here rather than the entire community, our results suggest that under the increased soil nutrient conditions expected with continued climate warming in the Arctic, herbivores may become more important in affecting several abundant tundra plant populations, and should not be ignored.     

CO2 exchange in three Canadian High Arctic ecosystems: response to long-term experimental warming

Carbon dioxide exchange, soil C and N, leaf mineral nutrition and leaf carbon isotope discrimination (LCID‐Δ) were measured in three High Arctic tundra ecosystems over 2 years under ambient and long‐term (9 years) warmed (～2°C) conditions. These ecosystems are located at Alexandra Fiord (79°N) on Ellesmere Island, Nunavut, and span a soil water gradient; dry, mesic, and wet tundra. Growing season CO₂ fluxes (i.e., net ecosystem exchange (NEE), gross ecosystem photosynthesis (GEP), and ecosystem respiration (R_e)) were measured using an infrared gas analyzer and winter C losses were estimated by chemical absorption. All three tundra ecosystems lost CO₂ to the atmosphere during the winter, ranging from 7 to 12 g CO₂‐C m^?2 season^?1 being highest in the wet tundra. The period during the growing season when mesic tundra switch from being a CO₂ source to a CO₂ sink was increased by 2 weeks because of warming and increases in GEP. Warming during the summer stimulated dry tundra GEP more than R_e and thus, NEE was consistently greater under warmed as opposed to ambient temperatures. In mesic tundra, warming stimulated GEP with no effect on R_e increasing NEE by ～10%, especially in the first half of the summer. During the ～70 days growing season (mid‐June–mid‐August), the dry and wet tundra ecosystems were net CO₂‐C sinks (30 and 67 g C m^?2 season^?1, respectively) and the mesic ecosystem was a net C source (58 g C m^?2 season^?1) to the atmosphere under ambient temperature conditions, due in part to unusual glacier melt water flooding that occurred in the mesic tundra. Experimental warming during the growing season increased net C uptake by ～12% in dry tundra, but reduced net C uptake by ～20% in wet tundra primarily because of greater rates of R_e as opposed to lower rates of GEP. Mesic tundra responded to long‐term warming with ～30% increase in GEP with almost no change in R_e reducing this tundra type to a slight C source (17 g C m^?2 season^?1). Warming caused LCID of Dryas integrafolia plants to be higher in dry tundra and lower in Salix arctic plants in mesic and wet tundra. Our findings indicate that: (1) High Arctic ecosystems, which occur in similar mesoclimates, have different net CO₂ exchange rates with the atmosphere; (2) long‐term warming can increase the net CO₂ exchange of High Arctic tundra by stimulating GEP, but it can also reduce net CO₂ exchange in some tundra types during the summer by stimulating R_e to a greater degree than stimulating GEP; (3) after 9 years of experimental warming, increases in soil carbon and nitrogen are detectable, in part, because of increases in deciduous shrub cover, biomass, and leaf litter inputs; (4) dry tundra increases in GEP, in response to long‐term warming, is reflected in D. integrifolia LCID; and (5) the differential carbon exchange responses of dry, mesic, and wet tundra to similar warming magnitudes appear to depend, in part, on the hydrologic (soil water) conditions. Annual net ecosystem CO₂‐C exchange rates ranged from losses of 64 g C m^?2 yr^?1 to gains of 55 g C m^?2 yr^?1. These magnitudes of positive NEE are close to the estimates of NPP for these tundra types in Alexandra Fiord and in other High Arctic locations based on destructive harvests.     

Shifting grassland plant community structure drives positive interactive effects of warming and diversity on aboveground net primary productivity

Ecosystems worldwide are increasingly impacted by multiple drivers of environmental change, including climate warming and loss of biodiversity. We show, using a long‐term factorial experiment, that plant diversity loss alters the effects of warming on productivity. Aboveground primary productivity was increased by both high plant diversity and warming, and, in concert, warming (≈1.5 °C average above and belowground warming over the growing season) and diversity caused a greater than additive increase in aboveground productivity. The aboveground warming effects increased over time, particularly at higher levels of diversity, perhaps because of warming‐induced increases in legume and C4 bunch grass abundances, and facilitative feedbacks of these species on productivity. Moreover, higher plant diversity was associated with the amelioration of warming‐induced environmental conditions. This led to cooler temperatures, decreased vapor pressure deficit, and increased surface soil moisture in higher diversity communities. Root biomass (0–30 cm) was likewise consistently greater at higher plant diversity and was greater with warming in monocultures and at intermediate diversity, but at high diversity warming had no detectable effect. This may be because warming increased the abundance of legumes, which have lower root : shoot ratios than the other types of plants. In addition, legumes increase soil nitrogen (N) supply, which could make N less limiting to other species and potentially decrease their investment in roots. The negative warming × diversity interaction on root mass led to an overall negative interactive effect of these two global change factors on the sum of above and belowground biomass, and thus likely on total plant carbon stores. In total, plant diversity increased the effect of warming on aboveground net productivity and moderated the effect on root mass. These divergent effects suggest that warming and changes in plant diversity are likely to have both interactive and divergent impacts on various aspects of ecosystem functioning.     

Responses of moist non-acidic arctic tundra to altered environment: productivity, biomass, and species richness

In arctic Alaska, researchers have manipulated air temperature, light availability, and soil nutrient availability in several tundra communities over the past two decades. These communities responded quite differently to the same manipulations, and species responded individualistically within communities and among sites. For example, moist acidic tundra is primarily nitrogen (N)‐limited, whereas wet sedge tundra is primarily phosphorus (P)‐limited, and the magnitude of growth responses varies across sites within communities. Here we report results of four years of manipulated nutrients (N and/or P) and/or air temperature in an understudied, diverse plant community, moist non‐acidic tussock tundra, in northern Alaska. Our goals were to determine which factors limit above‐ground net primary productivity (ANPP) and biomass, how community composition changes may affect ecosystem attributes, and to compare these results with those from other communities to determine their generality. Although relative abundance of functional groups shifted in several treatments, the only significant change in community‐level ANPP and biomass occurred in plots that received both N and P, driven by an increase in graminoid biomass and production resulting from a positive effect of adding N. There was no difference in community biomass among any other treatments; however, some growth forms and individual species did respond. After four years no one species has come to dominate the treatment plots and species richness has not changed. These results are similar to studies in dry heath, wet sedge, and moist acidic tundra where community biomass had the greatest response to both N and P and warming results were more subtle. Unlike in moist acidic tundra where shrub biomass increased markedly with fertilization, our results suggest that in non‐acidic tundra carbon sequestration in plant biomass will not increase substantially under increased soil nutrient conditions because of the lack of overstory shrub species.     

The response of Alaskan arctic tundra to experimental warming: differences between short- and long-term responses

Global climate models predict continued rapid warming for most of the Arctic throughout the next century. To further understand the response of arctic tundra to climate warming, four sites in northern Alaska were warmed for five to seven consecutive growing seasons using open‐top chambers. Sites were located in dry heath and wet meadow communities near Barrow (71°18′N, 156°40′W) and Atqasuk (70°29′N, 157°25′W). Change in plant community composition was measured using a point frame method. During the period of observation, species richness declined in control plots by up to 2.7 species plot^?1. Responses to warming varied by site but similar trends included increased canopy height (?0.1 to 2.3 cm) and relative cover of standing dead plant matter (1.5–6.0%) and graminoids (1.8–5.8%) and decreased species diversity (0.1–1.7 species plot^?1) and relative cover of lichens (0.2–9.1%) and bryophytes (1.4–4.6%) (parentheses enclose the range of average values for the sites). The response to warming was separated into an initial short‐term response assessed after two growing seasons of warming and a secondary longer‐term response assessed after an additional three to five growing seasons of warming. The initial responses to warming were similar in the four sites, while the secondary responses varied by site. The response to warming was greater at Barrow than Atqasuk because of a greater initial response at Barrow. However, the long‐term response to warming was projected to be greater at Atqasuk because of a greater secondary response at Atqasuk. These findings show that predictions of vegetation change due to climate warming based on manipulative experiments will differ depending on both the duration and plant community on which the study focuses.     

Soil–plant N processes in a High Arctic ecosystem,NW Greenland are altered by long‐term experimental warming and higher rainfall

Rapid temperature and precipitation changes in High Arctic tundra ecosystems are altering the biogeochemical cycles of carbon (C) and nitrogen (N), but in ways that are difficult to predict. The challenge grows from the uncertainty of N cycle responses and the extent to which shifts in soil N are coupled with the C cycle and productivity of tundra systems. We used a long‐term (since 2003) experiment of summer warming and supplemental summer water additions to a High Arctic ecosystem in NW Greenland, and applied a combination of discrete sampling and in situ soil core incubations to measure C and N pools and seasonal microbial processes that might control plant‐available N. We hypothesized that elevated temperature and increased precipitation would stimulate microbial activity and net inorganic N mineralization, thereby increasing plant N‐availability through the growing season. While we did find increased N mineralization rates under both global change scenarios, water addition also significantly increased net nitrification rates, loss of NO₃^?‐N via leaching, and lowered rates of labile organic N production. We also expected the chronic warming and watering would lead to long‐term changes in soil N‐cycling that would be reflected in soil δ¹⁵N values. We found that soil δ¹⁵N decreased under the different climate change scenarios. Our results suggest that temperature accelerates biological processes and existing C and N transformations, but moisture increases soil hydraulic connectivity and so alters the pathways, and changes the fate of the products of C and N transformations. In addition, our findings indicate that warmer, wetter High Arctic tundra will be cycling N and C in ways that may transform these landscapes in part leading to greater C sequestration, but simultaneously, N losses from the upper soil profile that may be transported to depth dissolved in water and or transported off site in lateral flow.     

Soil aeration in relation to soil physical properties, nitrogen availability, and root characteristics within an arctic watershed

The seasonal change in soil oxygen availability was determined in several habitats along a topographic moisture gradient in an arctic watershed. The effect of changes in soil aeration on soil chemical and plant properties was examined by comparison of the driest (tussocks) and wettest (wet sedge tundra) sites along this gradient. Spatial variability and seasonal change in soil oxygen availability was closely linked to the hydrologic regime and the thickness of the organic soil horizon. The greatest extension of the aerobic soil layer was found beneath well-drained tussocks, while less than 10% of the unfrozen soil layer is aerated in flooded wet sedge tundra. Intertussock areas and watertracks (channels of water drainage) have intermediate levels of aeration. In tussock tundra, soil oxygen diffusion is restricted in the mineral soil layer below the organic horizon due to reduced pore space. Organic matter constituents and their change with depth were similar beneath tussocks and in wet sedge tundra, indicating that factors other than soil aeration (e.g. low soil temperatures, short growing season) are the primary controls on decomposition in these two arctic tundra systems. NH₄ ⁺, the dominant form of inorganic N, was more available in wet sedge tundra than in tussock tundra. At both sites, extractable and soil solution NO₃ ^- concentrations increased 4 to 8 fold in the second part of the growing season, indicating increased nitrifier activity with improved soil oxygen availability. Although soils thawed as deep as 60 cm, approx. 90% of the root biomass was concentrated within 20 cm of the surface. Despite the anaerobic soil environment in wet sedge tundra, the dominant species there, Eriophorum angustifolium, reached slightly greater rooting depths than E. vaginatum, whose roots grow in the elevated, aerobic portion of tussocks. E. angustifolium had a root porosity of 31%, within the range found for wetland species, while roots of E. vaginatum had a porosity close to 12%. Rhizome porosity were low in both species (11%).     

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.     

The response of Arctic vegetation and soils following an unusually severe tundra fire

Fire causes dramatic short-term changes in vegetation and ecosystem function, and may promote rapid vegetation change by creating recruitment opportunities. Climate warming likely will increase the frequency of wildfire in the Arctic, where it is not common now. In 2007, the unusually severe Anaktuvuk River fire burned 1039 km² of tundra on Alaska''s North Slope. Four years later, we harvested plant biomass and soils across a gradient of burn severity, to assess recovery. In burned areas, above-ground net primary productivity of vascular plants equalled that in unburned areas, though total live biomass was less. Graminoid biomass had recovered to unburned levels, but shrubs had not. Virtually all vascular plant biomass had resprouted from surviving underground parts; no non-native species were seen. However, bryophytes were mostly disturbance-adapted species, and non-vascular biomass had recovered less than vascular plant biomass. Soil nitrogen availability did not differ between burned and unburned sites. Graminoids showed allocation changes consistent with nitrogen stress. These patterns are similar to those seen following other, smaller tundra fires. Soil nitrogen limitation and the persistence of resprouters will likely lead to recovery of mixed shrub–sedge tussock tundra, unless permafrost thaws, as climate warms, more extensively than has yet occurred.     

Circumpolar arctic tundra biomass and productivity dynamics in response to projected climate change and herbivory

Satellite remote sensing data have indicated a general ‘greening’ trend in the arctic tundra biome. However, the observed changes based on remote sensing are the result of multiple environmental drivers, and the effects of individual controls such as warming, herbivory, and other disturbances on changes in vegetation biomass, community structure, and ecosystem function remain unclear. We apply ArcVeg, an arctic tundra vegetation dynamics model, to estimate potential changes in vegetation biomass and net primary production (NPP) at the plant community and functional type levels. ArcVeg is driven by soil nitrogen output from the Terrestrial Ecosystem Model, existing densities of Rangifer populations, and projected summer temperature changes by the NCAR CCSM4.0 general circulation model across the Arctic. We quantified the changes in aboveground biomass and NPP resulting from (i) observed herbivory only; (ii) projected climate change only; and (iii) coupled effects of projected climate change and herbivory. We evaluated model outputs of the absolute and relative differences in biomass and NPP by country, bioclimate subzone, and floristic province. Estimated potential biomass increases resulting from temperature increase only are approximately 5% greater than the biomass modeled due to coupled warming and herbivory. Such potential increases are greater in areas currently occupied by large or dense Rangifer herds such as the Nenets‐occupied regions in Russia (27% greater vegetation increase without herbivores). In addition, herbivory modulates shifts in plant community structure caused by warming. Plant functional types such as shrubs and mosses were affected to a greater degree than other functional types by either warming or herbivory or coupled effects of the two.     

The effect of nutrient level on plant structure and production in a wet grassland: a field study

Since the 1950s, agricultural intensification has affected the structure and functioning of ecological systems including wet grasslands. Our study site, a wet grassland near Třeboň, Czech Republic, was historically a sedge meadow, but increased nutrient additions, a long-lasting flood in 2002 and changed mowing patterns resulted in domination by Phalaris arundinacea. The aim of the study was to determine how different nutrient conditions may affect plant structure and production in a wet grassland used for hay production. Species composition and percent cover were determined from line intercepts. Aboveground biomass was harvested six times each in 2007 and 2008 and primary production then calculated. Ingrowth core bags were used to determine belowground production. Aboveground production was two times greater in the high nutrient versus the low nutrient area in both years, while belowground production was very similar. The high nutrient area was still dominated by P. arundinacea, but Carex gracilis was now a co-dominant in the low nutrient area. However, other factors, such as water level and mowing regime, may interact with nutrient level to govern wet grassland structure and function.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.     

Enhanced summer warming reduces fungal decomposer diversity and litter mass loss more strongly in dry than in wet tundra

Many Arctic regions are currently experiencing substantial summer and winter climate changes. Litter decomposition is a fundamental component of ecosystem carbon and nutrient cycles, with fungi being among the primary decomposers. To assess the impacts of seasonal climatic changes on litter fungal communities and their functioning, Betula glandulosa leaf litter was surface‐incubated in two adjacent low Arctic sites with contrasting soil moisture regimes: dry shrub heath and wet sedge tundra at Disko Island, Greenland. At both sites, we investigated the impacts of factorial combinations of enhanced summer warming (using open‐top chambers; OTCs) and deepened snow (using snow fences) on surface litter mass loss, chemistry and fungal decomposer communities after approximately 1 year. Enhanced summer warming significantly restricted litter mass loss by 32% in the dry and 17% in the wet site. Litter moisture content was significantly reduced by summer warming in the dry, but not in the wet site. Likewise, fungal total abundance and diversity were reduced by OTC warming at the dry site, while comparatively modest warming effects were observed in the wet site. These results suggest that increased evapotranspiration in the OTC plots lowered litter moisture content to the point where fungal decomposition activities became inhibited. In contrast, snow addition enhanced fungal abundance in both sites but did not significantly affect litter mass loss rates. Across sites, control plots only shared 15% of their fungal phylotypes, suggesting strong local controls on fungal decomposer community composition. Nevertheless, fungal community functioning (litter decomposition) was negatively affected by warming in both sites. We conclude that although buried soil organic matter decomposition is widely expected to increase with future summer warming, surface litter decay and nutrient turnover rates in both xeric and relatively moist tundra are likely to be significantly restricted by the evaporative drying associated with warmer air temperatures.     

Potential contributions of root decomposition to the nitrogen cycle in arctic forest and tundra

Plant contributions to the nitrogen (N) cycle from decomposition are likely to be altered by vegetation shifts associated with climate change. Roots account for the majority of soil organic matter input from vegetation, but little is known about differences between vegetation types in their root contributions to nutrient cycling. Here, we examine the potential contribution of fine roots to the N cycle in forest and tundra to gain insight into belowground consequences of the widely observed increase in woody vegetation that accompanies climate change in the Arctic. We combined measurements of root production from minirhizotron images with tissue analysis of roots from differing root diameter and color classes to obtain potential N input following decomposition. In addition, we tested for changes in N concentration of roots during early stages of decomposition, and investigated whether vegetation type (forest or tundra) affected changes in tissue N concentration during decomposition. For completeness, we also present respective measurements of leaves. The potential N input from roots was twofold greater in forest than in tundra, mainly due to greater root production in forest. Potential N input varied with root diameter and color, but this variation tended to be similar in forest and tundra. As for roots, the potential N input from leaves was significantly greater in forest than in tundra. Vegetation type had no effect on changes in root or leaf N concentration after 1 year of decomposition. Our results suggest that shifts in vegetation that accompany climate change in the Arctic will likely increase plant‐associated potential N input both belowground and aboveground. In contrast, shifts in vegetation might not alter changes in tissue N concentration during early stages of decomposition. Overall, differences between forest and tundra in potential contribution of decomposing roots to the N cycle reinforce differences between habitats that occur for leaves.     

Responses of phenology and biomass production of boreal fens to climate warming under different water‐table level regimes

Climate change affects peatlands directly through increased air temperatures and indirectly through changes in water‐table level (WL). The interactions of these two still remain poorly known. We determined experimentally the separate and interactive effects of temperature and WL regime on factors of relevance for the inputs to the carbon cycle: plant community composition, phenology, biomass production, and shoot:root allocation in two wet boreal sedge‐dominated fens, “southern” at 62°N and “northern” at 68°Ν. Warming (1.5°C higher average daily air temperature) was induced with open‐top chambers and WL drawdown (WLD; 3–7 cm on average) by shallow ditches. Total biomass production varied from 250 to 520 g/m², with belowground production comprising 25%–63%. Warming was associated with minor effects on phenology and negligible effects on community composition, biomass production, and allocation. WLD clearly affected the contribution of different plant functional types (PFTs) in the community and the biomass they produced: shrubs benefited while forbs and mosses suffered. These responses did not depend on the warming treatment. Following WLD, aboveground biomass production decreased mainly due to reduced growth of mosses in the southern fen. Aboveground vascular plant biomass production remained unchanged but the contribution of different PFTs changed. The observed changes were also reflected in plant phenology, with different PFTs showing different responses. Belowground production increased following WLD in the northern fen only, but an increase in the contributions of shrubs and forbs was observed in both sites, while sedge contribution decreased. Moderate warming alone seems not able to drive significant changes in plant productivity or community composition in these wet ecosystems. However, if warming is accompanied by even modest WL drawdown, changes should be expected in the relative contribution of PFTs, which could lead to profound changes in the function of fens. Consequently, hydrological scenarios are of utmost importance when estimating their future function.