Acidic Tundra Research Articles

Soil nutrient availability affects tundra plant community composition and plant–vole interactions

ABSTRACT Climate change in the Arctic is predicted to drastically alter carbon and nutrient pools, plant communities, and plant–animal interactions. We examined how four levels of long-term (16 years) nutrient addition in moist acidic tundra affected plant community structure and the abundance of Eriophorum vaginatum, the preferred forage for tundra voles (Microtus oeconomus). We also explored how simulated press and pulse herbivory for four years affected Eriophorum at these different nutrient levels. Shifts in plant community structure, reductions in species diversity and richness, and decreased Eriophorum abundance were pronounced at the highest levels of nutrient addition. Eriophorum abundance was negatively correlated with deciduous shrub abundance, likely caused by light limitation in fertilized plots. Added nutrients generally increased Eriophorum leaf length, but effects varied among years and simulated herbivory treatments. After being clipped once, Eriophorum leaves regrew in fertilized plots to the same length as leaves that had not been clipped, demonstrating recovery. Our results show how interannual variation in plant growth and animal activity might exacerbate or dampen responses over multiple years. Changes to plant community composition and plant–vole interactions associated with increased soil nutrients resulting from warming could have cascading impacts on arctic ecosystems and carbon cycling.

Mineral organic carbon interactions in dry versus wet tundra soils

Mineral organic carbon interactions (aggregation, organo-mineral associations and organo-metallic complexes) enhance the protection of organic carbon (OC) from microbial degradation in soils. The northern circumpolar permafrost region stores between 1,440 and 1,600 Pg OC of which a significant portion is already thawed or about to thaw in coming years. In the light of this tipping point for climate change, any mechanism that can promote OC stabilization and hence mitigate OC mineralization and greenhouse gas emissions is of crucial interest. Here, we study interactions between metals (Fe, Al, Mn and Ca) and OC in the moist acidic tundra ecosystem of Eight Mile Lake, near Healy, AK, USA. We collected thirteen cores (124 soil samples) in late summer 2019 with shallow and deep active layers (45 to 109 cm deep) and varying water table depths. We find that between 6% and 59% of total OC in Eight Mile Lake tundra soils is mineral-associated (mean 20%), in organo-mineral associations (association between poorly crystalline oxides and OC) and in organo-metallic complexes (associations between Fe, Mn, Al, Ca polyvalent cations and organic acids). We find that total Fe and Mn concentrations can be used as good proxies to assess the reactive pool of these metals able to form associations with OC, i.e., poorly crystalline oxides or metals complexed with OC. We observe that in the active layer, mineral OC interactions are mostly as organo-metallic complexes with Fe cations, with an accumulation at the water table level acting as a soil redox interface. In waterlogged soils with a water table level above surface, no such accumulation of OC-Fe complexes is found due to the absence of a redox interface below soil surface. In the permafrost layer, we find that a combination of complexed metals and poorly crystalline Fe oxides act as reactive phases towards OC. Knowing that upon permafrost thaw tundra soils will become wetter or drier, the assessment of mineral-bound OC in drier or wetter tundra soils is a needed step to better constrain the changes in the proportion of non-protected OC more likely to contribute to C emissions from tundra soils.

NDVI changes in the Arctic: Functional significance in the moist acidic tundra of Northern Alaska.

The Normalized Difference Vegetation Index (NDVI), derived from reflected visible and infrared radiation, has been critical to understanding change across the Arctic, but relatively few ground truthing efforts have directly linked NDVI to structural and functional properties of Arctic tundra ecosystems. To improve the interpretation of changing NDVI within moist acidic tundra (MAT), a common Arctic ecosystem, we coupled measurements of NDVI, vegetation structure, and CO2 flux in seventy MAT plots, chosen to represent the full range of typical MAT vegetation conditions, over two growing seasons. Light-saturated photosynthesis, ecosystem respiration, and net ecosystem CO2 exchange were well predicted by NDVI, but not by vertically-projected leaf area, our nondestructive proxy for leaf area index (LAI). Further, our data indicate that NDVI in this ecosystem is driven primarily by the biochemical properties of the canopy leaves of the dominant plant functional types, rather than purely the amount of leaf area; NDVI was more strongly correlated with top cover and repeated cover of deciduous shrubs than other plant functional types, a finding supported by our data from separate "monotypic" plots. In these pure stands of a plant functional type, deciduous shrubs exhibited higher NDVI than any other plant functional type. Likewise, leaves from the two most common deciduous shrubs, Betula nana and Salix pulchra, exhibited higher leaf-level NDVI than those from the codominant graminoid, Eriophorum vaginatum. Our findings suggest that recent increases in NDVI in MAT in the North American Arctic are largely driven by expanding deciduous shrub canopies, with substantial implications for MAT ecosystem function, especially net carbon uptake.

Quantifying pH buffering capacity in acidic, organic-rich Arctic soils: Measurable proxies and implications for soil carbon degradation

Dynamic pH change promoted by biogeochemical reactions in Arctic tundra soils can be a major control on the production and release of CO2 and CH4, which contribute to rising global temperatures. Large quantities of soil organic matter (SOM) in these soils are susceptible to microbial decomposition, leading to pH changes during permafrost thaw. Soil pH buffering capacity (β) modulates the extent of pH change but has not been thoroughly studied and represented in predictive ecosystem scale biogeochemical models in Arctic tundra soils. In this study, we generated titration curves for 21 acidic tundra soils from three Arctic sites across northern Alaska, United States of America. Geochemical and hydrological soil properties were evaluated, and correlations with β were developed. Strong correlations between β and both gravimetric water content (Θg) (R2 = 0.847, p < 0.001) and soil water retention (SWR) (R2 = 0.849, p = 0.001) indicate that the ability of soil to retain water could be associated with its buffering properties. Correlations between β and soil organic carbon (SOC) and cation exchange capacity (CEC) were also explored, and relationships to SWR are discussed. These correlations were then used with existing soil databases reporting SOC, CEC, and SWR to estimate β across Alaska soils. We further demonstrated the quantitative relationships between β and the simulated rates of biogeochemical reactions and show that lower β leads to higher soil pH and more CH4 production. Our study provides simple proxies for β in Arctic soils and highlights the importance and implications of representing soil buffering in predictive models, thereby enabling quantitative coupling between pH dynamics associated with biogeochemical reactions. Integrating β into predictive models of Arctic biogeochemical cycling may reduce model uncertainty and further our understanding of permafrost SOM degradation accelerated by warming.

Small herbivores with big impacts: Tundra voles (Microtus oeconomus) alter post-fire ecosystem dynamics.

Fire is an important ecological disturbance that can reset ecosystems and initiate changes in plant community composition, ecosystem biogeochemistry, and primary productivity. As herbivores rely on primary producers for food, changes in vegetation may alter plant-herbivore interactions with important-but often unexplored-feedbacks to ecosystems. Here we examined the impact of post-fire changes in plant community composition and structure on habitat suitability and rodent herbivore activity in response to a large, severe, and unprecedented fire in northern Alaskan tundra. In moist acidic tundra where the fire occurred, tundra voles (Microtus oeconomus) are the dominant herbivore and rely on the tussock forming sedge Eriophorum vaginatum for both food and nesting material. Tundra voles were 10 times more abundant at the burned site compared with nearby unburned tundra 7-12 years after the fire. Fire increased the habitat suitability for voles by increasing plant productivity and biomass, food quality, and cover through both taller vegetation and increased microtopography. As a result of elevated vole abundance, Eriophorum mortality caused by vole herbivory was two orders of magnitude higher than natural mortality and approached the magnitude of the mortality rate resulting directly from the fire. These findings suggest that post-fire increases in herbivore pressure on Eriophorum could, in turn, disrupt graminoid recovery and enhance shrub encroachment. Tundra state transitions from graminoid to shrub dominated are also evident following other disturbances and fertilization experiments, suggesting that as Arctic temperatures rise, greater available nutrients and increased frequencies of large-scale disturbances may also alter plant-animal interactions with cascading impacts on plant communities and ecosystem function.

Landscape Genomics Provides Evidence of Ecotypic Adaptation and a Barrier to Gene Flow at Treeline for the Arctic Foundation Species Eriophorum vaginatum.

Global climate change has resulted in geographic range shifts of flora and fauna at a global scale. Extreme environments, like the Arctic, are seeing some of the most pronounced changes. This region covers 14% of the Earth’s land area, and while many arctic species are widespread, understanding ecotypic variation at the genomic level will be important for elucidating how range shifts will affect ecological processes. Tussock cottongrass (Eriophorum vaginatum L.) is a foundation species of the moist acidic tundra, whose potential decline due to competition from shrubs may affect ecosystem stability in the Arctic. We used double-digest Restriction Site-Associated DNA sequencing to identify genomic variation in 273 individuals of E. vaginatum from 17 sites along a latitudinal gradient in north central Alaska. These sites have been part of 30 + years of ecological research and are inclusive of a region that was part of the Beringian refugium. The data analyses included genomic population structure, demographic models, and genotype by environment association. Genome-wide SNP investigation revealed environmentally associated variation and population structure across the sampled range of E. vaginatum, including a genetic break between populations north and south of treeline. This structure is likely the result of subrefugial isolation, contemporary isolation by resistance, and adaptation. Forty-five candidate loci were identified with genotype-environment association (GEA) analyses, with most identified genes related to abiotic stress. Our results support a hypothesis of limited gene flow based on spatial and environmental factors for E. vaginatum, which in combination with life history traits could limit range expansion of southern ecotypes northward as the tundra warms. This has implications for lower competitive attributes of northern plants of this foundation species likely resulting in changes in ecosystem productivity.

Shading decreases and delays NDVI and flowering of prostrate Arctic shrubs

Increases in shrub growth and canopy cover are well documented community responses to climate warming in the Arctic. An important consequence of larger deciduous shrubs is shading of prostrate plant species, many of which are important sources of nectar and berries. Here, we present the impact of a shading experiment on two prostrate shrubs, Vaccinium vitis-idaea L. and Arctous alpina L., in northern Alaska over two growing seasons. We implemented three levels of shading (no shade, 40% shade, and 80% shade) in dry heath and moist acidic tundra. Plots were monitored for soil moisture content, surface temperature, normalized difference vegetation index (NDVI), and flowering. Shading was shown to, on average, lower surface temperature (0.7 °C to 5.3 °C) and increase soil moisture content (0.5% to 5.6%) in both communities. Both species- and plot-level NDVI values were delayed in timing of peak values (7 to 13 days) and decreased at the highest shading. Flower abundance of both species was lower in shaded plots and peak flowering was delayed (3 to 8 days) compared with controls. Changes in timing may result in phenological mismatches and can impact other trophic levels in the Arctic as both the flowers and resulting berries are important food sources for animals.

Intraspecific variation in phenology offers resilience to climate change for Eriophorum vaginatum

The phenology of Arctic plants is an important determinant of the pattern of carbon uptake and may be highly sensitive to continued rapid climate change. Eriophorum vaginatum L. (Cyperaceae) has a disproportionate influence over ecosystem processes in moist acidic tundra, but it is unclear whether its growth and phenology will remain competitive in the future. We investigated whether northern tundra ecotypes of E. vaginatum could extend their growing season in response to direct warming and transplanting into southern ecosystems. At the same time, we examined whether southern ecotypes could adjust their growth patterns in order to thrive further north, should they disperse quickly enough. Detailed phenology measurements across three reciprocal transplant gardens over a 2-year period showed that some northern ecotypes were capable of growing for longer when conditions were favourable, but their biomass and growing season length was still shorter than those of the southern ecotype. Southern ecotypes retained large leaf length when transplanted north and mirrored the growing season length better than the others, mainly owing to immediate green-up after snowmelt. All ecotypes retained the same senescence timing, regardless of environment, indicating a strong genetic control. Eriophorum vaginatum may remain competitive in a warming world if southern ecotypes can migrate north.

Tundra Type Drives Distinct Trajectories of Functional and Taxonomic Composition of Arctic Fungal Communities in Response to Climate Change - Results From Long-Term Experimental Summer Warming and Increased Snow Depth.

The arctic tundra is undergoing climate-driven changes and there are serious concerns related to the future of arctic biodiversity and altered ecological processes under possible climate change scenarios. Arctic land surface temperatures and precipitation are predicted to increase further, likely causing major transformation in terrestrial ecosystems. As a response to increasing temperatures, shifts in vegetation and soil fungal communities have already been observed. Little is known, however, how long-term experimental warming coupled with increased snow depth influence the trajectories of soil fungal communities in different tundra types. We compared edaphic variables and fungal community composition in experimental plots simulating the expected increase in summer warming and winter snow depth, based on DNA metabarcoding data. Fungal communities in the sampled dry and moist acidic tundra communities differed greatly, with tundra type explaining ca. one-third of compositional variation. Furthermore, dry and moist tundra appear to have different trajectories in response to climate change. Specifically, while both warming and increased snow depth had significant effects on fungal community composition and edaphic variables in dry tundra, the effect of increased snow was greater. However, in moist tundra, fungal communities mainly were affected by summer warming, while increased snow depth had a smaller effect and only on some functional groups. In dry tundra, microorganisms generally are limited by moisture in the summer and extremely low temperatures in winter, which is in agreement with the stronger effect of increased snow depth relative to warming. On the contrary, moist tundra soils generally are saturated with water, remain cold year-round and show relatively small seasonal fluctuations in temperature. The greater observed effect of warming on fungi in moist tundra may be explained by the narrower temperature optimum compared to those in dry tundra.

The relationships of present vegetation, bacteria, and soil properties with soil organic matter characteristics in moist acidic tundra in Alaska

Soil organic matter (SOM) is related to vegetation, soil bacteria, and soil properties; however, not many studies link all these parameters simultaneously, particularly in tundra ecosystems vulnerable to climate change. Our aim was to describe the relationships between vegetation, bacteria, soil properties, and SOM composition in moist acidic tundra by integrating physical, chemical, and molecular methods. A total of 70 soil samples were collected at two different depths from 36 spots systematically arranged over an area of about 300 m × 50 m. Pyrolysis-gas chromatography/mass spectrometry and pyrosequencing of the 16S rRNA gene were used to identify the molecular compositions of the SOM and bacterial community, respectively. Vegetation and soil physicochemical properties were also measured. The sampling sites were grouped into three, based on their SOM compositions: Sphagnum moss-derived SOM, lipid-rich materials, and aromatic-rich materials. Our results show that SOM composition is spatially structured and linked to microtopography; however, the vegetation, soil properties, and bacterial community composition did not show overall spatial structuring. Simultaneously, soil properties and bacterial community composition were the main factors explaining SOM compositional variation, while vegetation had a residual effect. Verrucomicrobia and Acidobacteria were related to polysaccharides, and Chloroflexi was linked to aromatic compounds. These relationships were consistent across different hierarchical levels. Our results suggest that SOM composition at a local scale is closely linked with soil factors and the bacterial community. Comprehensive observation of ecosystem components is recommended to understand the in-situ function of bacteria and the fate of SOM in the moist acidic tundra.

Comparative transcriptomics of an arctic foundation species, tussock cottongrass (Eriophorum vaginatum), during an extreme heat event

Tussock cottongrass (Eriophorum vaginatum) is a foundation species for much of the arctic moist acidic tundra, which is currently experiencing extreme effects of climate change. The Arctic is facing higher summer temperatures and extreme weather events are becoming more common. We used Illumina RNA-Seq to analyse cDNA libraries for differential expression of genes from leaves of ecologically well-characterized ecotypes of tussock cottongrass found along a latitudinal gradient in the Alaskan Arctic and transplanted into a common garden. Plant sampling was performed on a typical summer day and during an extreme heat event. We obtained a de novo assembly that contained 423,353 unigenes. There were 363 unigenes up-regulated and 1,117 down-regulated among all ecotypes examined during the extreme heat event. Of these, 26 HSP unigenes had >log2-fold up-regulation. Several TFs associated with heat stress in previous studies were identified that had >log2-fold up- or down-regulation during the extreme heat event (e.g., DREB, NAC). There was consistent variation in DEGs among ecotypes, but not specifically related to whether plants originated from taiga or tundra ecosystems. As the climate changes it is essential to determine ecotypic diversity at the genomic level, especially for widespread species that impact ecosystem function.

Deeper snow increases the net soil organic carbon accrual rate in moist acidic tussock tundra: 210Pb evidence from Arctic Alaska

ABSTRACT The net change in the carbon inventory of arctic tundra remains uncertain as global warming leads to shifts in arctic water and carbon cycles. To better understand the response of arctic tundra carbon to changes in winter precipitation amount, we investigated soil depth profiles of carbon concentration and radionuclide activities (7Be, 137Cs, 210Pb, and 241Am) in the active layer of a twenty-two-year winter snow depth manipulation experiment in moist acidic tussock tundra at Toolik Lake, Alaska. Depth correlations of cumulative carbon dry mass (g cm−2) vs. unsupported 210Pb activity (mBq g−1) were examined using a modified constant rate of supply (CRS) model. Results were best fit by two-slope CRS models indicating an apparent step temporal increase in the accumulation rate of soil organic carbon. Most of the best-fit model chronologies indicated that the increase in carbon accumulation rate apparently began and persisted after snow fence construction in 1994. The inhomogeneous nature of permafrost soils and their relatively low net carbon accumulation rates make it challenging to establish robust chronologic records. Nonetheless, the data obtained in this study support a decadal-scale increase in net soil organic carbon accumulation rate in the active layer of arctic moist acidic tussock tundra under conditions of increased winter precipitation.

Above- and belowground responses to long-term herbivore exclusion

ABSTRACT Herbivores can play an important role in determining arctic ecosystem function with effects determined in part by herbivore identity. We examined the impact of long-term (twenty-two years) small and large mammal herbivore exclusion in two arctic plant communities in northern Alaska: dry heath (DH) and moist acidic tundra (MAT). Our aims were to examine how herbivore exclusion influences (1) plant communities and (2) soil nutrient pools and microbial processes. Though herbivore absence increased moss and decreased evergreen shrub cover in MAT, there were few other significant effects on vegetation in either community. We also observed no influence of exclusion on most soil properties. However, in DH, phosphatase activity was greater in areas where small mammals alone were present, suggesting that they are altering phosphorus (P) availability, perhaps through herbivores’ influence on the plant community and subsequently on competition for P with the microbial community. We conclude that herbivore impacts in the Arctic are dependent on both the plant community and herbivore identity (size). We show the importance of understanding the roles of herbivores in the Arctic and contribute to a growing number of herbivore studies in a biome likely to experience future changes in herbivore communities and ecosystem function.

Decoupled above‐ and belowground responses to multi‐decadal nitrogen and phosphorus amendments in two tundra ecosystems

AbstractGlobal change in the Arctic promotes deeper soil thaw and enhanced soil microbial activity, increasing nitrogen (N) and phosphorus (P) availability to plants and microbes in strongly nutrient‐limited ecosystems. This critical, positive climate feedback has been examined through fertilization experiments that describe short‐term (&lt;10 yr) above‐ or belowground responses to combined NP additions, with evidence of enhanced shrub growth, nutrient availability, and soil organic matter decomposition. There has been less opportunity for long‐term comparisons of both above‐ and belowground responses with factorial N and P additions in different systems, despite broad awareness that ecosystem response can shift with time, and the potential for decoupled above‐ vs. belowground or N vs. P responses, currently and with further predicted global change. We examined the response of the plants, soil microbes, and soil nutrients, to factorial N and P additions in the moist acidic tundra (MAT; 26 yr of nutrient additions) and moist non‐acidic tundra (MNT; 16 yr). Aboveground, the MAT plant community continues to change as predicted by earlier studies: Functional groups responded independently to N and P, but NDVI‐biomass, especially of Betula nana, only increased with N addition. Unlike shorter‐term MNT studies, the MNT vegetation, which does not include B. nana, shows few new fertilization responses. Belowground responses were not predicted by aboveground responses in either MAT or MNT. In contrast to the N response aboveground, MAT microbial biomass responded positively and microbial phosphatase activity negatively to P additions, implying possible release from microbial P limitation. Critically, earlier published results of declines in soil total carbon (C) with combined NP addition in the MAT are not present in the long term. We make two conclusions: (1) Arctic ecosystems are not universally N‐limited but also exhibit complex responses to P alone or in combination with N; and (2) the presence or absence of key vegetation species can cascade from aboveground to belowground and restrict the extrapolation of responses of nutrient addition in a single arctic ecosystem to other arctic ecosystems, the short‐term to the long term, or aboveground to belowground.

Complete Genome Sequence of the Aerobic Facultative Methanotroph Methylocella tundrae Strain T4.

Methylocella tundrae T4T is a facultative aerobic methanotroph which was isolated from an acidic tundra wetland and possesses only a soluble methane monooxygenase. The complete genome, which includes two megaplasmids, was sequenced using a combination of Illumina and Nanopore technologies. One of the megaplasmids carries a propane monooxygenase gene cluster.

Predicting the decomposability of arctic tundra soil organic matter with mid infrared spectroscopy

Vast amounts of soil organic matter (SOM) have been preserved in arctic soils over millennia time scales due to the limiting effects of cold and wet environments on decomposer activity. With the increase in high latitude warming due to climate change, the potential decomposability of this SOM needs to be assessed. In this study, we investigated the capability of mid infrared (MIR) spectroscopy to quickly predict soil carbon and nitrogen concentrations and carbon (C) mineralized during short-term incubations of tundra soils. Active layer and upper permafrost soils collected from four tundra sites on the North Slope of Alaska were incubated at 1, 4, 8 and 16 °C for 60 days. All incubated soils were scanned to obtain the MIR spectra and analyzed for total organic carbon (TOC) and total nitrogen (TN) concentrations, and salt-extractable organic matter carbon (SEOM). Partial least square regression (PLSR) models, constructed using the MIR spectral data for all soils, were excellent predictors of soil TOC and TN concentrations and good predictors of mineralized C for these tundra soils. We explored whether we could improve the prediction of mineralized C by splitting the soils into the groups defined by the influential factors and thresholds identified in a principal components analysis: (1) TOC >10%, (2) TOC < 10%, (3) TN < 0.6%, (4) TN > 0.6%, (5) acidic tundra, and (6) non-acidic tundra. The best PLSR mineralization models were found for soils with TOC < 10% and TN < 0.6%. Analysis of the PLSR loadings and beta coefficients from these models indicated a small number of influential spectral bands. These bands were associated with clay content, phenolics, aliphatics, silicates, carboxylic acids, and amides. Our results suggest that MIR could serve as a useful tool for quickly and reasonably estimating the initial decomposability of tundra soils, particularly for mineral soils and the mixed organic-mineral horizons of cryoturbated soils.

Glucose addition increases the magnitude and decreases the age of soil respired carbon in a long-term permafrost incubation study

Higher temperatures in northern latitudes will increase permafrost thaw and stimulate above- and belowground plant biomass growth in tundra ecosystems. Higher plant productivity increases the input of easily decomposable carbon (C) to soil, which can stimulate microbial activity and increase soil organic matter decomposition rates. This phenomenon, known as the priming effect, is particularly interesting in permafrost because an increase in C supply to deep, previously frozen soil may accelerate decomposition of C stored for hundreds to thousands of years. The sensitivity of old permafrost C to priming is not well known; most incubation studies last less than one year, and so focus on fast-cycling C pools. Furthermore, the age of respired soil C is rarely measured, even though old C may be vulnerable to labile C inputs. We incubated soil from a moist acidic tundra site in Eight Mile Lake, Alaska for 409 days at 15 °C. Soil from surface (0–25 cm), transition (45–55 cm), and permafrost (65–85 cm) layers were amended with three pulses of uniformly 13C-labeled glucose or cellulose every 152 days. Glucose addition resulted in positive priming in the permafrost layer 7 days after each substrate addition, eliciting a two-fold increase in cumulative soil C loss relative to unamended soils with consistent effects across all three pulses. In the transition and permafrost layers, glucose addition significantly decreased the age of soil-respired CO2-C with Δ14C values that were 115‰ higher. Previous field studies that measured the age of respired C in permafrost regions have attributed younger Δ14C ecosystem respiration values to higher plant contributions. However, the results from this study suggest that positive priming, due to an increase in fresh C supply to deeply thawed soil layers, can also explain the respiration of younger C observed at the ecosystem scale. We must consider priming effects to fully understand permafrost C dynamics, or we risk underestimating the contribution of soil C to ecosystem respiration.

Divergent patterns of experimental and model-derived permafrost ecosystem carbon dynamics in response to Arctic warming

In the last few decades, temperatures in the Arctic have increased twice as much as the rest of the globe. As permafrost thaws in response to this warming, large amounts of soil organic matter may become vulnerable to decomposition. Microbial decomposition will release carbon (C) from permafrost soils, however, warmer conditions could also lead to enhanced plant growth and C uptake. Field and modeling studies show high uncertainty in soil and plant responses to climate change but there have been few studies that reconcile field and model data to understand differences and reduce uncertainty. Here, we evaluate gross primary productivity (GPP), ecosystem respiration (Reco), and net ecosystem C exchange (NEE) from eight years of experimental soil warming in moist acidic tundra against equivalent fluxes from the Community Land Model during simulations parameterized to reflect the field conditions associated with this manipulative field experiment. Over the eight-year experimental period, soil temperatures and thaw depths increased with warming in field observations and model simulations. However, the field and model results do not agree on warming effects on water table depth; warming created wetter soils in the field and drier soils in the models. In the field, initial increases in growing season GPP, Reco, and NEE to experimentally-induced permafrost thaw created a higher C sink capacity in the first years followed by a stronger C source in years six through eight. In contrast, both models predicted linear increases in GPP, Reco, and NEE with warming. The divergence of model results from field experiments reveals the role subsidence, hydrology, and nutrient cycling play in influencing the C flux responses to permafrost thaw, a complexity that the models are not structurally able to predict, and highlight challenges associated with projecting C cycle dynamics across the Arctic.

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.

Mammalian herbivory exacerbates plant community responses to long-term increased soil nutrients in two Alaskan tundra plant communities

The interaction between bottom-up and top-down forces in regulating plant communities is a long-standing topic of interest in ecology. Factorial field experiments examining these factors have been relatively few, but recent meta-analyses provide predictions that can be tested in a range of ecosystems. We tested the prediction that added nutrients would reduce species richness and evenness, while herbivore activity would offset those changes in two tundra plant communities after 11 years. In moist acidic tundra (MAT), herbivores reduced richness more in fertilized plots when mammals were present compared with fertilized plots without herbivores. In dry heath (DH), evenness was significantly reduced in fertilized plots only when herbivores were present, also providing evidence that herbivores enhanced community changes caused by greater nutrient availability. The difference in response between MAT and the meta-analysis predictions appears to be driven by Betula nana, the species that dramatically increased with added nutrients in MAT. Unlike in similar studies and in DH, B. nana is not as palatable as most of the species in the community and is generally avoided by herbivores. These results highlight how the effects of herbivory and nutrients differ across communities and can be affected by the traits of the species present.