Novel approaches to study climate change effects on terrestrial ecosystems in the field: Drought and passive nighttime warming

Beier C., Emmett B., Gundersen P., Tietema A., Peñuelas J., Estiarte M., Gordon C., Gorissen A., Llorens L., Roda F., Williams D. (2004) Novel approaches to study climate change effects on terrestrial ecosystems in the field: Drought and passive nighttime warming. Ecosystems. 7: 583-597.
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Doi: 10.1007/s10021-004-0178-8

Resum:

This article describes new approaches for manipulation of temperature and water input in the field. Nighttime warming was created by reflection of infrared radiation. Automatically operated reflective curtains covered the vegetation at night to reduce heat loss to the atmosphere. This approach mimicked the way climate change, caused by increased cloudiness and increased greenhouse gas emissions, alters the heat balance of ecosystems. Drought conditions were created by automatically covering the vegetation with transparent curtains during rain events over a 2-5-month period. The experimental approach has been evaluated at four European sites across a climate gradient. All sites were dominated (more than 50%) by shrubs of the ericaceous family. Within each site, replicated 4-m X 5-m plots were established for control, warming, and drought treatments and the effect on climate variables recorded. Results over a two-year period indicate that the warming treatment was successful in achieving an increase of the minimum temperatures by 0.4-1.2°C in the air and soil. The drought treatment resulted in a soil moisture reduction of 33%-82% at the peak of the drought. The data presented demonstrate that the approach minimizes unintended artifacts with respect to water balance, moisture conditions, and light, while causing a small but significant reduction in wind speed by the curtains. Temperature measurements demonstrated that the edge effects associated with the treatments were small. Our method provides a valuable tool for investigating the effects of climate change in remote locations with minimal artifacts.

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The response of soil processes to climate change: Results from manipulation studies of shrublands across an environmental gradient

Emmett B.A., Beier C., Estiarte M., Tietema A., Kristensen H.L., Williams D., Peñuelas J., Schmidt I., Sowerby A. (2004) The response of soil processes to climate change: Results from manipulation studies of shrublands across an environmental gradient. Ecosystems. 7: 625-637.
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Doi: 10.1007/s10021-004-0220-x

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Predicted changes in climate may affect key soil processes such as respiration and net nitrogen (N) mineralization and thus key ecosystem functions such as carbon (C) storage and nutrient availability. To identify the sensitivity of shrubland soils to predicted climate changes, we have carried out experimental manipulations involving ecosystem warming and prolonged summer drought in ericaceous shrublands across a European climate gradient. We used retractable covers to create artificial nighttime warming and prolonged summer drought to 20-m2 experimental plots. Combining the data from across the environmental gradient with the results from the manipulation experiments provides evidence for strong climate controls on soil respiration, net N mineralization and nitrification, and litter decomposition. Trends of 0%-19% increases of soil respiration in response to warming and decreases of 3%-29% in response to drought were observed. Across the environmental gradient and below soil temperatures of 20°C at a depth of 5-10 cm, a mean Q10 of 4.1 in respiration rates was observed although this varied from 2.4 to 7.0 between sites. Highest Q10 values were observed in Spain and the UK and were therefore not correlated with soil temperature. A trend of increased accumulated surface litter mass loss was observed with experimental warming (2%-22%) but there was no consistent response to experimental drought. In contrast to soil respiration and decomposition, variability in net N mineralization was best explained by soil moisture rather than temperature. When water was neither limiting or in excess, a Q10 of 1.5 was observed for net N mineralization rates. These data suggest that key soil processes will be differentially affected by predicted changes in rainfall pattern and temperature and the net effect on ecosystem functioning will be difficult to predict without a greater understanding of the controls underlying the sensitivity of soils to climate variables.

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Reflectance assessment of seasonal and annual changes in biomass and CO2 uptake of a Mediterranean shrubland submitted to experimental warming and drought

Filella I., Peñuelas J., Llorens L., Estiarte M. (2004) Reflectance assessment of seasonal and annual changes in biomass and CO2 uptake of a Mediterranean shrubland submitted to experimental warming and drought. Remote Sensing of Environment. 90: 308-318.
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Doi: 10.1016/j.rse.2004.01.010

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We aimed to evaluate how the remote sensing vegetation indices NDVI and PRI responded to seasonal and annual changes in an early successional stage Mediterranean coastal shrubland canopy that was submitted to experimental warming and drought simulating predicted climate change for the next decades. These conditions were obtained by using a new non-intrusive methodological approach that increases the temperature and prolongs the drought period by using roofs that automatically cover the vegetation after the sunset or when it rains. On average, warming increased air temperature by 0.7°C and soil temperature by 1.6°C, and the drought treatment reduced soil moisture by 22%. We measured spectral reflectance at the canopy level and at the individual plant level seasonally during 4 years. Shrubland NDVI tracked the community development and activity. In control and warming treatments, NDVI increased with the years while it did not change in the drought treatment. There was a good relationship between NDVI and both community and individual plant biomass. NDVI also decreased in summer seasons when some species dry or decolour. The NDVI of E. multiflora plant individuals was lower in autumn and winter than in the other seasons, likely because of flowering. Shrubland PRI decreased only in winter, similarly to the PRI of the most dominant species, G. alypum. At this community scale, NDVI was better related than PRI to photosynthetic activity, probably because photosynthetic fluxes followed canopy seasonal greening in this complex canopy, which includes brevideciduous, annual and evergreen species and variable morphologies and canopy coverage. PRI followed the seasonal variations in photosynthetic rates in E. multiflora and detected the decreased photosynthetic rates of drought treatment. However, PRI did not track the photosynthetic rates of G. alypum plants which have lower LAIs than E. multiflora. In this community, which is in its early successional stages, NDVI was able to track biomass, and indirectly, CO2 uptake changes, likely because LAI values did not saturate NDVI. Thus, NDVI appears as a valid tool for remote tracking of this community development. PRI was less adequate for photosynthetic assessment of this community especially for its lower LAI canopies. PRI usefulness was also species-dependent and could also be affected by flowering. These results will help to improve the interpretation of remote sensing information on the structure and physiological status of these Mediterranean shrublands, and to gain better insight on ecological and environmental controls on their ecosystem carbon dioxide exchange. They also show the possibility of assessing the impacts of climate change on shrubland communities. © 2004 Elsevier Inc. All rights reserved.

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Experimental evidences of climate change effects on plant recruitment in the Western Mediterranean Basin.

Lloret F, Peñuelas J, Estiarte M, Ogaya R (2004) Experimental evidences of climate change effects on plant recruitment in the Western Mediterranean Basin. In Arianoutsou M, Papanastasis VP (eds) Ecology, conservation and management of Mediterranean climate ecosystems. Millpress, Rotterdam. Edició en CD-ROM (ISBN-90-5966-016-1).

Climate change affects carbon allocation to the soil in shrublands

Gorissen A., Tietema A., Joosten N.N., Estiarte M., Peñuelas J., Sowerby A., Emmett B.A., Beier C. (2004) Climate change affects carbon allocation to the soil in shrublands. Ecosystems. 7: 650-661.
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Doi: 10.1007/s10021-004-0218-4

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Climate change may affect ecosystem functioning through increased temperatures or changes in precipitation patterns. Temperature and water availability are important drivers for ecosystem processes such as photosynthesis, carbon translocation, and organic matter decomposition. These climate changes may affect the supply of carbon and energy to the soil microbial population and subsequently alter decomposition and mineralization, important ecosystem processes in carbon and nutrient cycling. In this study, carried out within the cross-European research project CLIMOOR, the effect of climate change, resulting from imposed manipulations, on carbon dynamics in shrubland ecosystems was examined. We performed a 14C-labeling experiment to probe changes in net carbon uptake and allocation to the roots and soil compartments as affected by a higher temperature during the year and a drought period in the growing season. Differences in climate, soil, and plant characteristics resulted in a gradient in the severity of the drought effects on net carbon uptake by plants with the impact being most severe in Spain, followed by Denmark, with the UK showing few negative effects at significance levels of p ≤ 0.10. Drought clearly reduced carbon flow from the roots to the soil compartments. The fraction of the 14C fixed by the plants and allocated into the soluble carbon fraction in the soil and to soil microbial biomass in Denmark and the UK decreased by more than 60%. The effects of warming were not significant, but, as with the drought treatment, a negative effect on carbon allocation to soil microbial biomass was found. The changes in carbon allocation to soil microbial biomass at the northern sites in this study indicate that soil microbial biomass is a sensitive, early indicator of drought- or temperature-initiated changes in these shrubland ecosystems. The reduced supply of substrate to the soil and the response of the soil microbial biomass may help to explain the observed acclimation of CO2 exchange in other ecosystems.

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Effects of an experimental increase of temperature and drought on the photosynthetic performance of two ericaceous shrub species along a north-south European gradient

Llorens L., Peñuelas J., Beier C., Emmett B., Estiarte M., Tietema A. (2004) Effects of an experimental increase of temperature and drought on the photosynthetic performance of two ericaceous shrub species along a north-south European gradient. Ecosystems. 7: 613-624.
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Doi: 10.1007/s10021-004-0180-1

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Plant ecophysiological changes in response to climatic change may be different in northern and southern European countries because different abiotic factors constrain plant physiological activity. We studied the effects of experimental warming and drought on the photosynthetic performance of two ericaceous shrubs (Erica multiflora and Calluna vulgaris) along a European gradient of temperature and precipitation (UK, Denmark, The Netherlands, and Spain). At each site, a passive warming treatment was applied during the night throughout the whole year, whereas the drought treatment excluded rain events over 6-10 weeks during the growing season. We measured leaf gas exchange, chlorophyll a fluorescence, and leaf carbon isotope ratio (δ 13C) during the growing seasons of 1999 and 2000. Leaf net photosynthetic rates clearly followed a gradient from northern to southern countries in agreement with the geographical gradient in water availability. Accordingly, there was a strong correlation between net photosynthetic rates and the accumulated rainfall over the growing season. Droughted plants showed lower leaf gas exchange rates than control plants in the four sites. Interestingly, although leaf photosynthetic rates decreased along the precipitation gradient and in response to drought treatment, droughted plants were able to maintain higher leaf photosynthetic rates than control plants in relation to the accumulated rainfall over the months previous to the measurements. Droughted plants also showed higher values of potential photochemical efficiency (F v/Fm) in relation to controls, mainly at midday. The warming treatment did not affect significantly any of the studied instantaneous ecophysiological variables.

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Contrasting growth changes in two dominant species of a mediterranean shrubland submitted to experimental drought and warming

Llorens L., Peñuelas J., Estiarte M., Bruna P. (2004) Contrasting growth changes in two dominant species of a mediterranean shrubland submitted to experimental drought and warming. Annals of Botany. 94: 843-853.
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Doi: 10.1093/aob/mch211

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• Background and Aims: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. • Methods: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. • Key Results: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6°C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. • Conclusions: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term. © 2004 Annals of Botany Company.

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Experimental evidence of reduced diversity of seedlings due to climate modification in a Mediterranean-type community

Lloret F., Penuelas J., Estiarte M. (2004) Experimental evidence of reduced diversity of seedlings due to climate modification in a Mediterranean-type community. Global Change Biology. 10: 248-258.
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Doi: 10.1111/j.1365-2486.2004.00725.x

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We are still lacking in experimental evidence of the effects of climate change on the richness of plant species under field conditions. We report a decrease in the species richness of recruited seedlings in a Mediterranean shrubland in experimentally induced drought and warming over 4 consecutive years. Drought decreased the number of emerging seedlings and their respective species richness. Warming also decreased seedling species richness, but it did not affect the number of emerging seedlings. Species that produce fewer recruits are more likely to disappear in drier or warmer scenarios. However, when the effect of induced climate treatment was greatest, the more abundant species in control stands were not necessarily the ones least affected by treatment; in other words, species-idiosyncratic responses may occur. These results show that demographic processes are sensitive to minor climate changes, with probable consequences on the diversity and structure of the future plant communities. © 2004 Blackwell Publishing Ltd.

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Efectos del cambio climático.

Llorens L, Estiarte M, Peñuelas J (2004) Efectos del cambio climático. Quercus 218:40-45.

Complex spatiotemporal phenological shifts as a response to rainfall changes

Peñuelas J., Filella I., Zhang X., Llorens L., Ogaya R., Lloret F., Comas P., Estiarte M., Terradas J. (2004) Complex spatiotemporal phenological shifts as a response to rainfall changes. New Phytologist. 161: 837-846.
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Doi: 10.1111/j.1469-8137.2004.01003.x

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• Climatic warming produces significant gradual alterations in the timing of life-cycle events, and here we study the phenological effects of rainfall-pattern changes. • We conducted ecosystem field experiments that partially excluded rain and runoff during the growing season in a Mediterranean forest and in a mediterranean shrubland. Studies of time-series of leaf-unfolding, flowering and fruiting over the last 50 yr in central Catalonia were carried out, and greenup onset in the Iberian Peninsula was monitored by satellite images. • Experimental, historical and geographical changes in rainfall produced significant, complex and strongly species-specific, as well as spatially and temporally variable, phenological effects. Among these changes, it was found that in the Iberian Peninsula, greenup onset changes from spring (triggered by rising temperatures) in the northern cool-wet regions to autumn (triggered by the arrival of autumn rainfalls) in the southern warm-dry regions. Even in the mesic Mediterranean central Catalonia (NE of the peninsula) rainfall had a stronger relative influence than temperature on fruiting phenology. • The results show that changes in rainfall and water availability, an important driver of climate change, can cause complex phenological changes with likely far-reaching consequences for ecosystem and biosphere functioning and structure. The seasonal shift in the Iberian Peninsula further highlights this importance and indicates that vegetation may respond to climate change not only with gradual, but also with abrupt temporal and spatial, changes in the timing of greenup onset.

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