Elevation alters ecosystem properties across temperate treelines globally.

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.

Plant and microbial responses to nitrogen and phosphorus addition across an elevational gradient in subarctic tundra.

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 approximately 3 degrees 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.

Bioavailable soil phosphorus decreases with increasing elevation in a subarctic tundra landscape.

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.