Dynamics of alpine plant litter decomposition in a changing climate

Climatic changes resulting from anthropogenic activities over the passed century are repeatedly reported to alter the functioning of pristine ecosystems worldwide, and especially those in cold biomes. Available literature on the process of plant leaf litter decomposition in the temperate Alpine zone is reviewed here, with emphasis on both direct and indirect effects of climate change phenomena on rates of litter decay. Weighing the impact of biotic and abiotic processes governing litter mass loss, it appears that an immediate intensification of decomposition rates due to temperature rise can be retarded by decreased soil moisture, insufficient snow cover insulation, and shrub expansion in the Alpine zone. This tentative conclusion, remains speculative unless empirically tested, but it has profound implications for understanding the biogeochemical cycling in the Alpine vegetation belt, and its potential role as a buffering mechanism to climate chang

Subalpine Pasture-Woodlands in a Changing Climate:the Role of Land Use Type in Ecosystem Resistance to Simulated Climate Change

Climate change in temperate mountain systems and associated increase in temperature and decrease in precipitation are expected to have strong implications for vegetation productivity, species diversity and carbon turnover in subalpine grasslands. Little is known, however, about the interaction between the effects of climate change and those of local land use management and possible changes in landscape structure. Pasture woodlands in the Swiss Jura Mountains are a traditional landscape, resulting from a long-lived sustainable use of grasslands and woodlands, and as such provide a suite of important ecosystem services to human society. These range from carbon sequestration and biodiversity preservation, to provision of timber and forage for livestock, and last but not least an aesthetic value, much appreciated by tourism. In this thesis various aspects of ecosystem functioning have been studied, investigating the combined effects of experimental climate change and land use on structurally different wooded pastures. An altitudinal gradient method has been used to simulate future climate change conditions, by imposing warmer and drier climate on subalpine turfs transplanted at lower elevation. The resulting gradient in mean annual temperature and precipitation – ranging from cold and wet in the subalpine zone, to warm and dry in the colline zone – has allowed for the detection of tipping points and altered states of ecosystem functioning in response to the treatments. The method employed provided also the possibility for a direct comparison of three land use types: unwooded pastures, sparsely wooded pastures, and densely wooded pastures (the result of pasture management intensity), in their response to climate perturbation. During the four years of experimental work, a series of observations have been made at the plot scale (square metre) in terms of plant performance and biogeochemical cycles, as well as at the landscape scale (hectare) in terms of forage production. A general threshold level for ecosystem resistance to experimental climate change was detected between the moderate IPCC scenario (+2 K mean annual temperature; -20 % annual precipitation) and the intensive IPCC scenario (+4 K mean annual temperature; -40 % annual precipitation). A concomitant gradient in ecosystem response to climate change was observed across the three land use types. The intensively managed unwooded pasture type was consistently more affected by the experimental treatment and rarely exhibited signs of resistance, especially under the intense climate change scenario. A drastic loss of plant species diversity, reduction of herbaceous biomass, impaired litter decomposition and soil microbial metabolic activity have all contributed to the altered state of ecosystem functioning. In contrast, the two extensively managed wooded pasture types showed considerable resistance to climate perturbation in terms of both above and belowground ecosystem processes. The reported inter-annual variation in herbaceous diversity and biomass production within these land use types demonstrated their resilience (recovery) potential too. Using a modelling approach for upscaling these results to the heterogeneous landscape of pasture woodlands in the Swiss Jura Mountains, has proven that extensively used wooded pastures could grant sustainable ecosystem services in terms of forage provision for cattle under climate change. Considering that the two experimental climate change intensities implemented this study are the projected ‘best’ and ‘worst’ case scenarios for the coming decades, the reported resistance of wooded pastures to climate change has to be embraced, and sustainable land use set as a goal in high altitude mountain pastures

Dynamics of alpine plant litter decomposition in a changing climate

Climatic changes resulting from anthropogenic activities over the passed century are repeatedly reported to alter the functioning of pristine ecosystems worldwide, and especially those in cold biomes. Available literature on the process of plant leaf litter decomposition in the temperate Alpine zone is reviewed here, with emphasis on both direct and indirect effects of climate change phenomena on rates of litter decay. Weighing the impact of biotic and abiotic processes governing litter mass loss, it appears that an immediate intensification of decomposition rates due to temperature rise can be retarded by decreased soil moisture, insufficient snow cover insulation, and shrub expansion in the Alpine zone. This tentative conclusion, remains speculative unless empirically tested, but it has profound implications for understanding the biogeochemical cycling in the Alpine vegetation belt, and its potential role as a buffering mechanism to climate change

Transplantation of subalpine wood-pasture turfs along a natural climatic gradient reveals lower resistance of unwooded pastures to climate change compared to wooded ones

Climate change could impact strongly on cold-adapted mountain ecosystems, but little is known about its interaction with traditional land-use practices. We used an altitudinal gradient to simulate a year-round warmer and drier climate for semi-natural subalpine grasslands across a landscape of contrasting land-use management. Turf mesocosms from three pasture-woodland land-use types—unwooded pasture, sparsely wooded pasture, and densely wooded pasture—spanning a gradient from high to low management intensity were transplanted downslope to test their resistance to two intensities of climate change. We found strong overall effects of intensive (+4K) experimental climate change (i.e., warming and reduced precipitation) on plant community structure and function, while moderate (+2K) climate change did not substantially affect the studied land-use types, thus indicating an ecosystem response threshold to moderate climate perturbation. The individual land-use types were affected differently under the +4K scenario, with a 60% decrease in aboveground biomass (AGB) in unwooded pasture turfs, a 40% decrease in sparsely wooded pasture turfs, and none in densely wooded ones. Similarly, unwooded pasture turfs experienced a 30% loss of species, advanced (by 30days) phenological development, and a mid-season senescence due to drought stress, while no such effects were recorded for the other land-use types. The observed contrasting effects of climate change across the pasture-woodland landscape have important implications for future decades. The reduced impact of climate change on wooded pastures as compared to unwooded ones should promote the sustainable land use of wooded pastures by maintaining low management intensity and a sparse forest canopy, which buffer the immediate impacts of climate change on herbaceous vegetation

Isotopic analysis of cyanobacterial nitrogen fixation associated with subarctic lichen and bryophyte species

Dinitrogen fixation by cyanobacteria is of particular importance for the nutrient economy of cold biomes, constituting the main pathway for new N supplies to tundra ecosystems. It is prevalent in cyanobacterial colonies on bryophytes and in obligate associations within cyanolichens. Recent studies, applying interspecific variation in plant functional traits to upscale species effects on ecosystems, have all but neglected cryptogams and their association with cyanobacteria. Here we looked for species-specific patterns that determine cryptogam-mediated rates of N2 fixation in the Subarctic. We hypothesised a contrast in N2 fixation rates (1) between the structurally and physiologically different lichens and bryophytes, and (2) within bryophytes based on their respective plant functional types. Throughout the survey we supplied 15N-labelled N2 gas to quantify fixation rates for monospecific moss, liverwort and lichen turfs. We sampled fifteen species in a design that captures spatial and temporal variations during the growing season in Abisko region, Sweden. We measured N2 fixation potential of each turf in a common environment and in its field sampling site, in order to embrace both comparativeness and realism. Cyanolichens and bryophytes differed significantly in their cyanobacterial N2 fixation capacity, which was not driven by microhabitat characteristics, but rather by morphology and physiology. Cyanolichens were much more prominent fixers than bryophytes per unit dry weight, but not per unit area due to their low specific thallus weight. Mosses did not exhibit consistent differences in N2 fixation rates across species and functional types. Liverworts did not fix detectable amounts of N2. Despite the very high rates of N2 fixation associated with cyanolichens, large cover of mosses per unit area at the landscape scale compensates for their lower fixation rates, thereby probably making them the primary regional atmospheric nitrogen sin

Biotic and Abiotic Constraints on the Decomposition of Fagus sylvatica Leaf Litter Along an Altitudinal Gradient in Contrasting Land-Use Types

Climate change can affect the process of carbon cycling and leaf litter decomposition in multiple ways, both directly and indirectly, though the strength and direction of this relationship is often context dependent. In this experiment, we followed decomposition of a standard litter type—senescent leaves of Fagus sylvatica collected from a single location—along a 1000m altitudinal gradient of four sites over 2.5years. To control the edaphic conditions, we transplanted intact turf mesocosms from three different land-use types [densely wooded, sparsely wooded, and unwooded (UW) pastures] from the highest altitude site into UW pastures along the altitudinal gradient from the moist, cool high-elevation site to the dry, warm low-elevation site, using shade cloth to mimic the light conditions in the original habitats. Decomposition in the drier UW pasture mesocosms increased with altitude, likely because of higher moisture at the highest sites. Decomposition in the more mesic mesocosms from sparsely and densely wooded sites was insensitive to altitude, suggesting an overriding moisture, rather than temperature, constraint on decomposition across these sites. The functional composition of decomposer microbial communities (fungal/bacterial ratio) was similarly insensitive to altitude. Our findings bring substantial evidence for the controlling role of soil moisture on litter decomposition, as well as for the indirect effects of climate through changes in the decomposer community

Soil organic carbon stocks did not change after 130 years of afforestation on a former Swiss Alpine pasture

Soil organic matter (SOM) plays an important role in the global carbon cycle, especially in alpine ecosystems. However, ongoing forest expansion in high-elevation systems potentially alters SOM storage through changes in organic matter (OM) inputs and microclimate. In this study, we investigated the effects of an Picea abies L. afforestation chrono-sequence (0 to 130 years) of a former subalpine pasture in Switzerland on soil organic carbon (SOC) stocks and SOM dynamics. We found that SOC stocks remained constant throughout the chrono-sequence, with comparable SOC stocks in the mineral soils after afforestation and previous pasture (SOC forest40 = 11.6 ± 1.1 kg m−2, SOC forest130 = 11.0 ± 0.3 kg m−2 and SOC pasture = 11.5 ± 0.5 kg m−2). However, including the additional carbon of the organic horizons in the forest, reaching up to 1.7 kg m−2 in the 55-year old forest, resulted in an increase in the overall SOC stocks following afforestation. We found that the soil C:N ratio in the mineral soil increased in the topsoil (0–5 cm) with increasing forest stand age, from 11.9 ± 1.3 in the pasture to 14.3 ± 1.8 in the 130-year old forest. In turn, we observed a decrease in the soil C:N ratio with increasing depth in all forest stand ages. This suggests that litter-derived organic matter (C:N from 35.1 ± 1.9 to 42.4 ± 10.8) is likely to be incorporated and translocated from the organic horizon to the mineral topsoil (0–10 cm) of the profiles. Due to the high root C:N ratio (pasture 63.5 ± 2.8 and forests between 54.7 ± 3.9 and 61.2 ± 2.9), particulate root-derived organic matter seems to have a rather small effect on forest soil C:N ratios, as well as on SOC accumulation in the mineral soil. These results suggest that, although afforestation does not change the SOC stock in the mineral soil, there is an apparent alteration in the SOM dynamics through changes in the litter composition caused by the vegetation shift. We conclude that, at our study site, spruce afforestation on a former subalpine pasture does not change the total SOC stock and that, consequently, there is no additional SOC sequestration on a decadal to centennial scale

Plant-microbial linkages underpin carbon sequestration in contrasting mountain tundra vegetation types

Tundra ecosystems hold large stocks of soil organic matter (SOM), likely due to low temperatures limiting rates of microbial SOM decomposition more than those of SOM accumulation from plant primary productivity and microbial necromass inputs. Here we test the hypotheses that distinct tundra vegetation types and their carbon supply to characteristic rhizosphere microbes determine SOM cycling independent of temperature. In the subarctic Scandes, we used a three-way factorial design with paired heath and meadow vegetation at each of two elevations, and with each combination of vegetation type and elevation subjected during one growing season to either ambient light (i.e., ambient plant productivity), or 95% shading (i.e., reduced plant productivity). We assessed potential above-and belowground ecosystem linkages by uni-and multivariate analyses of variance, and structural equation modelling. We observed direct coupling between tundra vegetation type and microbial community composition and function, which underpinned the ecosystem's potential for SOM storage. Greater primary productivity at low elevation and ambient light supported higher microbial biomass and nitrogen immobilisation, with lower microbial mass-specific enzymatic activity and SOM humification. Congruently, larger SOM at lower elevation and in heath sustained fungal-dominated microbial communities, which were less substrate-limited, and invested less into enzymatic SOM mineralisation, owing to a greater carbon-use efficiency (CUE). Our results highlight the importance of tundra plant community characteristics (i.e., productivity and vegetation type), via their effects on soil microbial community size, structure and physiology, as essential drivers of SOM turnover. The here documented concerted patterns in above-and belowground ecosystem functioning is strongly supportive of using plant community characteristics as surrogates for assessing tundra carbon storage potential and its evolution under climate and vegetation changes

Climate change effects on the stability and chemistry of soil organic carbon pools in a subalpine grassland

Mountain soils stock large quantities of carbon as particulate organic matter that may be highly vulnerable to climate change. To explore potential shifts in soil organic matter (SOM) form and stability under climate change (warming and reduced precipitations), we studied the dynamics of SOM pools of a mountain grassland in the Swiss Jura as part of a climate manipulation experiment. The climate manipulation (elevational soil transplantation) was set up in October 2009 and simulated two realistic climate change scenarios. After 4 years of manipulation, we performed SOM physical fractionation to extract SOM fractions corresponding to specific turnover rates, in winter and in summer. Soil organic matter fraction chemistry was studied with ultraviolet, 3D fluorescence, and mid-infrared spectroscopies. The most labile SOM fractions showed high intra-annual dynamics (amounts and chemistry) mediated via the seasonal changes of fresh plant debris inputs and confirming their high contribution to the microbial loop. Our climate change manipulation modified the chemical differences between free and intra-aggregate organic matter, suggesting a modification of soil macro-aggregates dynamics. Interestingly, the 4-year climate manipulation affected directly the SOM dynamics, with a decrease in organic C bulk soil content, resulting from significant C-losses in the mineral-associated SOM fraction (MAOM), the most stable form of SOM. This SOC decrease was associated with a decrease in clay content, above- and belowground plants biomass, soil microbial biomass and activity. The combination of these climate changes effects on the plant–soil system could have led to increase C-losses from the MAOM fraction through clay-SOM washing out and DOC leaching in this subalpine grassland

Isotopic analysis of cyanobacterial nitrogen fixation associated with subarctic lichen and bryophyte species.

Dinitrogen fixation by cyanobacteria is of particular importance for the nutrient economy of cold biomes, constituting the main pathway for new N supplies to tundra ecosystems. It is prevalent in cyanobacterial colonies on bryophytes and in obligate associations within cyanolichens. Recent studies, applying interspecific variation in plant functional traits to upscale species effects on ecosystems, have all but neglected cryptogams and their association with cyanobacteria. Here we looked for species-specific patterns that determine cryptogam-mediated rates of N-2 fixation in the Subarctic. We hypothesised a contrast in N-2 fixation rates (1) between the structurally and physiologically different lichens and bryophytes, and (2) within bryophytes based on their respective plant functional types. Throughout the survey we supplied N-15-labelled N-2 gas to quantify fixation rates for monospecific moss, liverwort and lichen turfs. We sampled fifteen species in a design that captures spatial and temporal variations during the growing season in Abisko region, Sweden. We measured N-2 fixation potential of each turf in a common environment and in its field sampling site, in order to embrace both comparativeness and realism. Cyanolichens and bryophytes differed significantly in their cyanobacterial N-2 fixation capacity, which was not driven by microhabitat characteristics, but rather by morphology and physiology. Cyanolichens were much more prominent fixers than bryophytes per unit dry weight, but not per unit area due to their low specific thallus weight. Mosses did not exhibit consistent differences in N-2 fixation rates across species and functional types. Liverworts did not fix detectable amounts of N-2. Despite the very high rates of N-2 fixation associated with cyanolichens, large cover of mosses per unit area at the landscape scale compensates for their lower fixation rates, thereby probably making them the primary regional atmospheric nitrogen sink