Responses of High Arctic wet sedge tundra to climate warming since 1980

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short‐term experimental manipulations, with few studies of long‐term ambient change. We investigated changes in above‐ and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980–1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short‐term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.     

Estimating root biomass in Salix viminalis × Salix schwerinii cultivar “Olof” using the electrical capacitance method

Abstract

Non‐destructive assessment of root systems is important in order to understand and optimise the potential of resource capture and allocation by the plant. We studied the relationships between electrical capacitance (EC) and the below‐ and above‐ground biomass of willows. Cuttings of Salix viminalis × Salix schwerinii cv. Olof were maintained in pots and root development was followed up using a portable capacitance meter over the course of 2.5 months. Pot observations were compared with excavation of two‐year‐old established trees. A strong significant linear relationship (R ² = 0.81, p < 0.001) was obtained between EC and root biomass (dry weight [DW]) for the pot experiment. EC also showed good correlations with stem and leaf biomass, as well as with stem height. In the excavated willow trees, there was a strong logarithmic relationship between EC and root biomass (R ² = 0.66, p < 0.001). These results suggest that EC is a good estimator of below‐ground biomass in willow and may become useful in screening varieties for differences in root biomass traits, especially in distinguishing below‐ground resource allocation at an early stage.     

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.     

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

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

Rising plant‐mediated methane emissions from arctic wetlands

Plant‐mediated CH₄ flux is an important pathway for land–atmosphere CH₄ emissions, but the magnitude, timing, and environmental controls, spanning scales of space and time, remain poorly understood in arctic tundra wetlands, particularly under the long‐term effects of climate change. CH₄ fluxes were measured in situ during peak growing season for the dominant aquatic emergent plants in the Alaskan arctic coastal plain, Carex aquatilis and Arctophila fulva, to assess the magnitude and species‐specific controls on CH₄ flux. Plant biomass was a strong predictor of A. fulva CH₄ flux while water depth and thaw depth were copredictors for C. aquatilis CH₄ flux. We used plant and environmental data from 1971 to 1972 from the historic International Biological Program (IBP) research site near Barrow, Alaska, which we resampled in 2010–2013, to quantify changes in plant biomass and thaw depth, and used these to estimate species‐specific decadal‐scale changes in CH₄ fluxes. A ~60% increase in CH₄ flux was estimated from the observed plant biomass and thaw depth increases in tundra ponds over the past 40 years. Despite covering only ~5% of the landscape, we estimate that aquatic C. aquatilis and A. fulva account for two‐thirds of the total regional CH₄ flux of the Barrow Peninsula. The regionally observed increases in plant biomass and active layer thickening over the past 40 years not only have major implications for energy and water balance, but also have significantly altered land–atmosphere CH₄ emissions for this region, potentially acting as a positive feedback 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.     

Changes in high arctic tundra plant reproduction in response to long‐term experimental warming

We provide new information on changes in tundra plant sexual reproduction in response to long‐term (12 years) experimental warming in the High Arctic. Open‐top chambers (OTCs) were used to increase growing season temperatures by 1–2 °C across a range of vascular plant communities. The warming enhanced reproductive effort and success in most species; shrubs and graminoids appeared to be more responsive than forbs. We found that the measured effects of warming on sexual reproduction were more consistently positive and to a greater degree in polar oasis compared with polar semidesert vascular plant communities. Our findings support predictions that long‐term warming in the High Arctic will likely enhance sexual reproduction in tundra plants, which could lead to an increase in plant cover. Greater abundance of vegetation has implications for primary consumers – via increased forage availability, and the global carbon budget – as a function of changes in permafrost and vegetation acting as a carbon sink. Enhanced sexual reproduction in Arctic vascular plants may lead to increased genetic variability of offspring, and consequently improved chances of survival in a changing environment. Our findings also indicate that with future warming, polar oases may play an important role as a seed source to the surrounding polar desert landscape.     

Phenology and species determine growing‐season albedo increase at the altitudinal limit of shrub growth in the sub‐Arctic

Arctic warming is resulting in reduced snow cover and increased shrub growth, both of which have been associated with altered land surface–atmospheric feedback processes involving sensible heat flux, ground heat flux and biogeochemical cycling. Using field measurements, we show that two common Arctic shrub species (Betula glandulosa and Salix pulchra), which are largely responsible for shrub encroachment in tundra, differed markedly in albedo and that albedo of both species increased as growing season progressed when measured at their altitudinal limit. A moveable apparatus was used to repeatedly measure albedo at six precise spots during the summer of 2012, and resampled in 2013. Contrary to the generally accepted view of shrub‐covered areas having low albedo in tundra, full‐canopy prostrate B. glandulosa had almost the highest albedo of all surfaces measured during the peak of the growing season. The higher midsummer albedo is also evident in localized MODIS albedo aggregated from 2000 to 2013, which displays a similar increase in growing‐season albedo. Using our field measurements, we show the ensemble summer increase in tundra albedo counteracts the generalized effect of earlier spring snow melt on surface energy balance by approximately 40%. This summer increase in albedo, when viewed in absolute values, is as large as the difference between the forest and tundra transition. These results indicate that near future (<50 years) changes in growing‐season albedo related to Arctic vegetation change are unlikely to be particularly large and might constitute a negative feedback to climate warming in certain circumstances. Future efforts to calculate energy budgets and a sensible heating feedback in the Arctic will require more detailed information about the relative abundance of different ground cover types, particularly shrub species and their respective growth forms and phenology.     

Effect of defoliation upon root growth,phosphate absorption and respiration in nutrient-limited tundra graminoids

Summary Moderate experimental defoliation stimulated root respiration and phosphate absorption in two tundra graminoids, Eriophorum vaginatum and Carex aquatilis, growing under nutrient-limited field conditions in northern Alaska. The increase in phosphate absorption rate following defoliation of Eriophorum was associated with a decrease in root phosphate and available carbohydrate contents per unit root length but a constant root nitrogen content. Only after four repeated defoliations did phosphate absorption rate decrease below control levels. We suggest that the stimulation of root respiration and phosphate absorption immediately following defoliation resulted from lowered root phosphorus status as nutrient reserves were reallocated to support shoot regrowth. Root growth was affected more severely by defoliation than was root activity. Two or more defoliations reduced root elongation, initiation and weight per unit length, but root mortality increased only after four defoliations. Carex aquatilis, a species with large belowground biomass, was less sensitive to defoliation than Eriophorum. Phosphate absorption rate increased only after four defoliations in this species, and root elongation, initiation and mortality were affected only by the most severe clipping regimes. Responses of plants to repeated defoliation over two growing seasons were consistent with results of short-term studies.     

Effects of a nematode Meloidogyne incognita and its interaction with above‐ground herbivory on an invasive wetland plant,alligator weed (Alternanthera philoxeroides)

Invasive plants may be attacked both above ground and below ground. Few studies have, however, investigated the simultaneous effects of above‐ground and below‐ground herbivory. In the present study, we report the effects of beetle herbivory and nematode infection on alligator weed, Alternanthera philoxeroides, an invasive plant in China. We found that the root‐knot nematode Meloidogyne incognita widely occurred on the plant in south China. To examine its effect on the plant in conjunction with above‐ground herbivory, we conducted a field common garden experiment with a local insect defoliator, Cassida piperata. We also included the native congener Alternanthera sessilis in our experiments for a comparison of the response of invasive and native species. We found no significant effects on plant biomass of the nematode infection in conjunction with the above‐ground herbivory. Further chemical analysis, however, showed that the water‐soluble carbohydrate content in roots of A. philoxeroides was significantly increased in plants attacked by both the nematode and the herbivore compared with the water‐soluble carbohydrate content in plants attacked by only the nematode or herbivore alone. We found no such change in the native congener A. sessilis. Together these results may suggest that A. philoxeroides tolerates joint above‐ground and below‐ground damage by allocating more resources to below‐ground material.     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.     

Biomass and leaf‐level gas exchange characteristics of three African savanna C4 grass species under optimum growth conditions

C₄ savanna grass species, Digitaria eriantha, Eragrostis lehmanniana and Panicum repens, were grown under optimum growth conditions with the aim of characterizing their above‐ and below‐ground biomass allocation and the response of their gas exchange to changes in light intensity, CO₂ concentration and leaf‐to‐air vapour pressure deficit gradient (D_l). Digitaria eriantha showed the largest above‐ and below‐ground biomass, high efficiency in carbon gain under light‐limiting conditions, high water use efficiency (WUE) and strong stomatal sensitivity to D_l (P = 0.002; r² = 0.5). Panicum repens had a high aboveground biomass and attained high light saturated photosynthetic rates (A_sat, 47 μmol m^?2 s^?1), stomatal conductance, (g_sat, 0.25 mol m^?2 s^?1) at relatively high WUE. Eragrostis lehmanniana had almost half the biomass of other species, and had similar A_sat and g_sat but were attained at lower WUE than the other species. This species also showed the weakest stomatal response to D_l (P = 0.19, r² = 0. 1). The potential ecological significance of the contrasting patterns of biomass allocation and variations in gas exchange parameters among the species are discussed.     

Large‐scale pattern of biomass partitioning across China's grasslands

Aim To investigate large‐scale patterns of above‐ground and below‐ground biomass partitioning in grassland ecosystems and to test the isometric theory at the community level. Location Northern China, in diverse grassland types spanning temperate grasslands in arid and semi‐arid regions to alpine grasslands on the Tibetan Plateau. Methods We investigated above‐ground and below‐ground biomass in China's grasslands by conducting five consecutive sampling campaigns across the northern part of the country during 2001–05. We then documented the root : shoot ratio (R/S) and its relationship with climatic factors for China's grasslands. We further explored relationships between above‐ground and below‐ground biomass across different grassland types. Results Our results indicated that the overall R/S of China's grasslands was larger than the global average (6.3 vs. 3.7). The R/S for China's grasslands did not show any significant trend with either mean annual temperature or mean annual precipitation. Above‐ground biomass was nearly proportional to below‐ground biomass with a scaling exponent (the slope of log–log linear relationship between above‐ground and below‐ground biomass) of 1.02 across various grassland types. The slope did not differ significantly between temperate and alpine grasslands or between steppe and meadow. Main conclusions Our findings support the isometric theory of above‐ground and below‐ground biomass partitioning, and suggest that above‐ground biomass scales isometrically with below‐ground biomass at the community level.     

Structural complexity and land‐surface energy exchange along a gradient from arctic tundra to boreal forest

Question: Current climate changes in the Alaskan Arctic, which are characterized by increases in temperature and length of growing season, could alter vegetation structure, especially through increases in shrub cover or the movement of treeline. These changes in vegetation structure have consequences for the climate system. What is the relationship between structural complexity and partitioning of surface energy along a gradient from tundra through shrub tundra to closed canopy forest? Location: Arctic tundra‐boreal forest transition in the Alaskan Arctic. Methods: Along this gradient of increasing canopy complexity, we measured key vegetation characteristics, including community composition, biomass, cover, height, leaf area index and stem area index. We relate these vegetation characteristics to albedo and the partitioning of net radiation into ground, latent, and sensible heating fluxes. Results: Canopy complexity increased along the sequence from tundra to forest due to the addition of new plant functional types. This led to non‐linear changes in biomass, cover, and height in the understory. The increased canopy complexity resulted in reduced ground heat fluxes, relatively conserved latent heat fluxes and increased sensible heat fluxes. The localized warming associated with increased sensible heating over more complex canopies may amplify regional warming, causing further vegetation change in the Alaskan Arctic.     

The fate of 15N added to high Arctic tundra to mimic increased inputs of atmospheric nitrogen released from a melting snowpack

Increases in the long‐range aerial transport of reactive N species from low to high latitudes will lead to increased accumulation in the Arctic snowpack, followed by release during the early summer thaw. We followed the release of simulated snowpack N, and its subsequent fate over three growing seasons, on two contrasting high Arctic tundra types on Spitsbergen (79°N). Applications of ¹⁵N (99 atom%) at 0.1 and 0.5 g N m^?2 were made immediately after snowmelt in 2001 as either Na¹⁵NO₃ or ¹⁵NH₄Cl. These applications are approximately 1 × and 5 × the yearly atmospheric deposition rates. The vegetation at the principal experimental site was dominated by bryophytes and Salix polaris while at the second site, vegetation included bryophytes, graminoids and lichens. Audits of the applied ¹⁵N were undertaken, over two or three growing seasons, by determining the amounts of labeled N in the soil (0–3 and 3–10 cm), soil microbial biomass and different vegetation fractions. Initial partitioning of the ¹⁵N at the first sampling time showed that ～60% of the applied ¹⁵N was recovered in soil, litter and plants, regardless of N form or application rate, indicating that rapid immobilization into organic forms had occurred at both sites. Substantial incorporation of the ¹⁵N was found in the microbial biomass in the humus layer and in the bryophyte and lichen fractions. After initial partitioning there appeared to be little change in the total ¹⁵N recovered over the following two or three seasons in each of the sampled fractions, indicating highly conservative N retention. The most obvious transfer of ¹⁵N, following assimilation, was from the microbial biomass into stable forms of humus, with an apparent half‐life of just over 1 year. At the principal site the microbial biomass and vascular plants were found to immobilize the greatest proportion of ¹⁵N compared with their total N concentration. In the more diverse tundra of the second site, lichen species and graminoids competed effectively for ¹⁵NH₄‐N and ¹⁵NO₃‐N, respectively. Results suggest that Arctic tundra habitats have a considerable capacity to immobilize additional inorganic N released from the snow pack. However, with 40% of the applied ¹⁵N apparently lost there is potential for N enrichment in the surrounding fjordal systems during the spring thaw.     

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.     

Variation in plant performance in a grassland: Species‐specific and neighbouring root mass effects

Question: What characteristics of local biotic neighbourhood is the best proxy of competitive effects experienced by plants in a herbaceous community: (1) total above‐ground biomass, (2) root mass or (3) relative above‐ground abundance of selected species? Location: Grassland at ca. 1100 m a.s.l. in the Krkono?e Mts., northern Czech Republic. Methods: We implanted two phytometer species, Antho‐xanthum alpinum and Festuca rubra, into a mountain grassland, and examined their response to local variation in (1) total above‐ground biomass, (2) root mass at three soil depths, and (3) relative abundance of individual species above‐ground. Results : Performance of both phytometer species was determined much more consistently by the mass of neighbouring roots and by species composition of neighbours than by the total above‐ground biomass. The two phytometer species showed different responses to these parameters. The most important relationships were (1) negative relationship between performance of Anthoxanthum and mass of neighbouring roots at 0–3 cm, (2) positive relationship between performance of Festuca and mass of neighbouring roots at 3–6 cm, and (3) negative relationship between performance of Festuca and relative abundance of Festuca in the neighbourhood. Conclusions: Neighbouring root mass and above‐ground species composition are better determinants of biotic interactions than total above‐ground biomass of neighbours in the studied mountain grassland. However, the relationships found are not necessarily due to variation in competitive intensity but can be due to other hidden factors as well, e.g. local availability of resources.     

Demography of three dominant sedges under contrasting grazing regimes in the High Arctic

Abstract. Tiller demography of Carex aquatilis ssp. stans, Carex membranacea, and Eriophorum angustifolium ssp. triste was investigated in ungrazed and grazed high arctic vegetation on central Ellesmere Island, Canada. Tiller birth, growth, flowering and death were studied from excavated clonal fragments, and tiller density and biomass were studied from excavated turfs. Five life‐cycle stages were determined: dormant buds, juvenile, mature, flowering and dead tillers. A stage‐based transition matrix model was developed to estimate the long‐term dynamics of the sedge populations and to compare life‐history strategies between ungrazed and grazed populations. Short‐term and retrospective models, based on the growth during the sampling year and during the lifetime of the clonal fragments, respectively, were compared to see how well the short‐term model can describe demography of long‐lived plants. According to the short‐term model, tiller populations were decreasing (λ < 1 except for C. membranacea), whereas the retrospective model indicated that the tiller populations were increasing. Tiller population growth rates did not differ between ungrazed and grazed habitats. Nevertheless, the similar growth rates may be obtained by balanced differences in the vital rates between plants of the two habitats. The plants in the ungrazed habitat tended to remain in their current life‐cycle stage, whereas plants in the grazed habitat moved quickly to the next stage and died earlier. C. aquatilis ssp. stans appears to gain a competitive advantage over the other species under intensive grazing, as indicated by the higher tiller density and greater below‐ground biomass in grazed vegetation. The greater amount of below‐ground biomass apparently buffers C. aquatilis ssp. stans against grazing better than the other species.     

Depth‐based differentiation in nitrogen uptake between graminoids and shrubs in an Arctic tundra plant community

Questions

The rapid climate warming in tundra ecosystems can increase nutrient availability in the soil, which may initiate shifts in vegetation composition. The direction in which the vegetation shifts will co‐determine whether Arctic warming is mitigated or accelerated, making the understanding of successional trajectories urgent. One of the key factors influencing the competitive relationships between plant species is their access to nutrients, depending on the depth where they take up most nutrients. However, nutrient uptake at different soil depths by tundra plant species that differ in rooting depth is unclear.

Location

Kytalyk Nature Reserve, northeast Siberia, Russia.

Methods

We injected ¹⁵N to 5 cm, 15 cm and the thaw front of the soil in a moist tussock tundra. The absorption of ¹⁵N by grasses, sedges, deciduous shrubs and evergreen shrubs from the three depths was compared.

Results

The results clearly show a vertical differentiation of N uptake by these plant functional types, corresponding to their rooting strategy. Shallow‐rooting dwarf shrubs were more capable of absorbing nutrients from the upper soil than from deeper soil. Deep‐rooting grasses and sedges were more capable of absorbing nutrients from deeper soil than the dwarf shrubs. The natural ¹⁵N abundances in control plants also indicate that graminoids can absorb more nutrients from the deeper soil than dwarf shrubs.

Conclusions

Our results show that graminoids and shrubs in the Arctic differ in their N uptake strategies, with graminoids profiting from nutrients released at the thaw front, while shrubs mainly forage in upper soil layers. Our results suggest that tundra vegetation will become graminoid‐dominated as permafrost thaw progresses and nutrient availability increases in the deep soil.     

Allometric scaling relationship between above‐ and below‐ground biomass within and across five woody seedlings

Allometric biomass allocation theory predicts that leaf biomass (M_L) scaled isometrically with stem (M_S) and root (M_R) biomass, and thus above‐ground biomass (leaf and stem) (M_A) and root (M_R) scaled nearly isometrically with below‐ground biomass (root) for tree seedlings across a wide diversity of taxa. Furthermore, prior studies also imply that scaling constant should vary with species. However, litter is known about whether such invariant isometric scaling exponents hold for intraspecific biomass allocation, and how variation in scaling constants influences the interspecific scaling relationship between above‐ and below‐ground biomass. Biomass data of seedlings from five evergreen species were examined to test scaling relationships among biomass components across and within species. Model Type II regression was used to compare the numerical values of scaling exponents and constants among leaf, stem, root, and above‐ to below‐ground biomass. The results indicated that M_L and M_S scaled in an isometric or a nearly isometric manner with M_R, as well as M_A to M_R for five woody species. Significant variation was observed in the Y‐intercepts of the biomass scaling curves, resulting in the divergence for intraspecific scaling and interspecific scaling relationships for M_L versus M_S and M_L versus M_R, but not for M_S versus M_R and M_A versus M_R. We conclude, therefore, that a nearly isometric scaling relationship of M_A versus M_R holds true within each of the studied woody species and across them irrespective the negative scaling relationship between leaf and stem.