Community structure and ecosystem processes often vary along elevational gradients. Their responses to elevation are commonly driven by changes in temperature, and many community-and ecosystem-level ...variables therefore frequently respond similarly to elevation across contrasting gradients. There are also many exceptions, sometimes because other factors such as precipitation can also vary with elevation. Given this complexity, our capacity to predict when and why the same variable responds differently among disparate elevational gradients is often limited. Furthermore, there is utility in using elevational gradients for understanding community and ecosystem responses to global climate change at much larger spatial and temporal scales than is possible through conventional ecological experiments. However, future studies that integrate elevational gradient approaches with experimental manipulations will provide powerful information that can improve predictions of climate change impacts within and across ecosystems.
Temperature and nutrients are major limiting factors in subarctic tundra. Experimental manipulation of nutrient availability along elevational gradients (and thus temperature) can improve our ...understanding of ecological responses to climate change. However, no study to date has explored impacts of nutrient addition along a tundra elevational gradient, or across contrasting vegetation types along any elevational gradient. We set up a full factorial nitrogen (N) and phosphorus (P) fertilization experiment in each of two vegetation types (heath and meadow) at 500 m, 800 m, and 1000 m elevation in northern Swedish tundra. We predicted that plant and microbial communities in heath or at lower elevations would be more responsive to N addition while communities in meadow or at higher elevations would be more responsive to P addition, and that fertilizer effects would vary more with elevation for the heath than for the meadow. Although our results provided little support for these predictions, the relationship between nutrient limitation and elevation differed between vegetation types. Most plant and microbial properties were responsive to N and/or P fertilization, but responses often varied with elevation and/or vegetation type. For instance, vegetation density significantly increased with N + P fertilization relative to the other fertilizer treatments, and this increase was greatest at the lowest elevation for the heath but at the highest elevation for the meadow. Arbuscular mycorrhizae decreased with P fertilization at 500 m for the meadow, but with all fertilizer treatments in both vegetation types at 800 m. Fungal to bacterial ratios were enhanced by N + P fertilization for the two highest elevations in the meadow only. Additionally, microbial responses to fertilization were primarily direct rather than indirect via plant responses, pointing to a decoupled response of plant and microbial communities to nutrient addition and elevation. Because our study shows how two community types differ in their responses to fertilization and elevation, and because the temperature range across this gradient is ∼3°C, our study is informative about how nutrient limitation in tundra may be influenced by temperature shifts that are comparable to those expected under climate change during this century.
Arctic plant growth is predominantly nitrogen (N) limited. This limitation is generally attributed to slow soil microbial processes due to low temperatures. Here, we show that arctic plant-soil N ...cycling is also substantially constrained by the lack of larger detritivores (earthworms) able to mineralize and physically translocate litter and soil organic matter. These new functions provided by earthworms increased shrub and grass N concentration in our common garden experiment. Earthworm activity also increased either the height or number of floral shoots, while enhancing fine root production and vegetation greenness in heath and meadow communities to a level that exceeded the inherent differences between these two common arctic plant communities. Moreover, these worming effects on plant N and greening exceeded reported effects of warming, herbivory and nutrient addition, suggesting that human spreading of earthworms may lead to substantial changes in the structure and function of arctic ecosystems.
Elevational gradients are increasingly recognized as a valuable tool for understanding how community and ecosystem properties respond to climatic factors, but little is known about how plant traits ...and their effects on ecosystem processes respond to elevation. We studied the response of plant leaf and litter traits, and litter decomposability across a gradient of elevation, and thus temperature, in subarctic tundra in northern Sweden for each of two contrasting vegetation types, heath and meadow, dominated by dwarf shrubs and herbaceous plants respectively. This was done at each of three levels; across species, within individual species, and the plant community using a community weighted average approach. Several leaf and litter traits shifted with increasing elevation in a manner consistent with greater conservation of nutrients at all three levels, and the most consistent response was an increase in tissue N to P ratio. However, litter decomposition was less directly responsive to elevation because the leaf and litter traits which were most responsive to elevation were not necessarily those responsible for driving decomposition. At the community level, the response to elevation of foliar and litter traits, and decomposability, varied greatly among the two vegetation types, highlighting the importance of vegetation type in determining ecological responses to climatic factors such as temperature. Finally our results highlight how understanding the responses of leaf and litter characteristics of functionally distinct vegetation types, and the processes that they drive, to temperature helps provide insights about how future climate change could affect tundra ecosystems.
Summary
Fine roots, and their functional traits, influence associated rhizosphere microorganisms via root exudation and root litter quality. However, little information is known about their ...relationship with rhizosphere microbial taxa and functional guilds.
We investigated the relationships of 11 fine root traits of 20 sub‐arctic tundra meadow plant species and soil microbial community composition, using phospholipid fatty acids (PLFAs) and high‐throughput sequencing. We primarily focused on the root economics spectrum, as it provides a useful framework to examine plant strategies by integrating the co‐ordination of belowground root traits along a resource acquisition–conservation trade‐off axis.
We found that the chemical axis of the fine root economics spectrum was positively related to fungal to bacterial ratios, but negatively to Gram‐positive to Gram‐negative bacterial ratios. However, this spectrum was unrelated to the relative abundance of functional guilds of soil fungi. Nevertheless, the relative abundance of arbuscular mycorrhizal fungi was positively correlated to root carbon content, but negatively to the numbers of root forks per root length.
Our results suggest that the fine root economics spectrum is important for predicting broader groups of soil microorganisms (i.e. fungi and bacteria), while individual root traits may be more important for predicting soil microbial taxa and functional guilds.
Litter decomposition is an important driver of soil carbon and nutrient cycling in nutrient-limited Arctic ecosystems. However, climate change is expected to induce changes that directly or ...indirectly affect decomposition. We examined the direct effects of long-term warming relative to differences in soil abiotic properties associated with vegetation type on litter decomposition across six subarctic vegetation types.
In six vegetation types, rooibos and green tea bags were buried for 70–75 days at 8 cm depth inside warmed (by open-top chambers) and control plots that had been in place for 20–25 years. Standardized initial decomposition rate and stabilization of the labile material fraction of tea (into less decomposable material) were calculated from tea mass losses. Soil moisture and temperature were measured bi-weekly during summer and plant-available nutrients were measured with resin probes.
Initial decomposition rate was decreased by the warming treatment. Stabilization was less affected by warming and determined by vegetation type and soil moisture. Soil metal concentrations impeded both initial decomposition rate and stabilization.
While a warmer Arctic climate will likely have direct effects on initial litter decomposition rates in tundra, stabilization of organic matter was more affected by vegetation type and soil parameters and less prone to be affected by direct effects of warming.
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•Warming with open top chambers decreased initial decomposition rate but not stabilization during decomposition in tundras.•Soil moisture, temperature and metal concentrations explained differences in the tea bag index between vegetation types.•In dry locations, initial decomposition rate and stabilization were not affected by soil temperature.•In wetter locations, initial decomposition rate decreased and stabilization increased with increasing soil temperature.•Increased soil metal concentrations impeded decomposition, which may affect decomposition irrespective of climate change.
Temperature is a primary driver of the distribution of biodiversity as well as of ecosystem boundaries. Declining temperature with increasing elevation in montane systems has long been recognized as ...a major factor shaping plant community biodiversity, metabolic processes, and ecosystem dynamics. Elevational gradients, as thermoclines, also enable prediction of long-term ecological responses to climate warming. One of the most striking manifestations of increasing elevation is the abrupt transitions from forest to treeless alpine tundra. However, whether there are globally consistent above- and belowground responses to these transitions remains an open question. To disentangle the direct and indirect effects of temperature on ecosystem properties, here we evaluate replicate treeline ecotones in seven temperate regions of the world. We find that declining temperatures with increasing elevation did not affect tree leaf nutrient concentrations, but did reduce ground-layer community-weighted plant nitrogen, leading to the strong stoichiometric convergence of ground-layer plant community nitrogen to phosphorus ratios across all regions. Further, elevation-driven changes in plant nutrients were associated with changes in soil organic matter content and quality (carbon to nitrogen ratios) and microbial properties. Combined, our identification of direct and indirect temperature controls over plant communities and soil properties in seven contrasting regions suggests that future warming may disrupt the functional properties of montane ecosystems, particularly where plant community reorganization outpaces treeline advance.
Global change is altering species distributions and thus interactions among organisms. Organisms live in concert with thousands of other species, some beneficial, some pathogenic, some which have ...little to no effect in complex communities. Since natural communities are composed of organisms with very different life history traits and dispersal ability it is unlikely they will all respond to climatic change in a similar way. Disjuncts in plant-pollinator and plant-herbivore interactions under global change have been relatively well described, but plant-soil microorganism and soil microbe-microbe relationships have received less attention. Since soil microorganisms regulate nutrient transformations, provide plants with nutrients, allow co-existence among neighbors, and control plant populations, changes in soil microorganism-plant interactions could have significant ramifications for plant community composition and ecosystem function. In this paper we explore how climatic change affects soil microbes and soil microbe-plant interactions directly and indirectly, discuss what we see as emerging and exciting questions and areas for future research, and discuss what ramifications changes in these interactions may have on the composition and function of ecosystems.
Elevational gradients are useful for predicting how plant communities respond to global warming, because communities at lower elevations experience warmer temperatures. Fine root traits and root ...trait variation could play an important role in determining plant community responses to warming in cold‐climate ecosystems where a large proportion of plant biomass is allocated belowground. Here, we investigated the effects of elevation‐associated temperature change on twelve chemical and morphological fine root traits of plant species and plant communities in a Swedish subarctic tundra. We also assessed the relative contributions of plant species turnover and intraspecific variation to the total amount of community‐level root trait variation explained by elevation. Several root traits, both at the species and whole community levels, had significant linear or quadratic relationships with elevation, but the direction and strength of these relationships varied among traits and plant species. Further, we found no support for a unidirectional change from more acquisitive root trait values at the lower elevations towards trait values associated with greater nutrient conservation at the higher elevations, either at the species or community level. On the other hand, root trait coefficients of variation at the community level increased with elevation for several root traits. Further, for a large proportion of the community‐level traits we found that intraspecific variation was relatively more important than species turnover, meaning that trait plasticity is important for driving community‐level trait responses to environmental factors in this tundra system. Our findings indicate that with progressing global warming, intraspecific trait variation may drive plant community composition but this may not necessarily lead to shifts in root resource–acquisition strategy for all species.
Phosphorus (P) is an important macronutrient in arctic and subarctic tundra and its bioavailability is regulated by the mineralization of organic P. Temperature is likely to be an important control ...on P bioavailability, although effects may differ across contrasting plant communities with different soil properties. We used an elevational gradient in northern Sweden that included both heath and meadow vegetation types at all elevations to study the effects of temperature, soil P sorption capacity and oxalate-extractable aluminium (Alox) and iron (Feox) on the concentration of different soil P fractions. We hypothesized that the concentration of labile P fractions would decrease with increasing elevation (and thus declining temperature), but would be lower in meadow than in heath, given that N to P ratios in meadow foliage are higher. As expected, labile P in the form of Resin-P declined sharply with elevation for both vegetation types. Meadow soils did not have lower concentrations of Resin-P than heath soils, but they did have 2-fold and 1.5-fold higher concentrations of NaOH-extractable organic P and Residual P, respectively. Further, meadow soils had 3-fold higher concentrations of Alox + Feox and a 20% higher P sorption index than did heath soils. Additionally, Resin-P expressed as a proportion of total soil P for the meadow was on average half that in the heath. Declining Resin-P concentrations with elevation were best explained by an associated 2.5-3.0 °C decline in temperature. In contrast, the lower P availability in meadow relative to heath soils may be associated with impaired organic P mineralization, as indicated by a higher accumulation of organic P and P sorption capacity. Our results indicate that predicted temperature increases in the arctic over the next century may influence P availability and biogeochemistry, with consequences for key ecosystem processes limited by P, such as primary productivity.
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