The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

The worldwide phenomenon of shrub encroachment in grass-dominated dryland ecosystems is commonly associated with desertification. Studies of the purported desertification effects associated with shrub encroachment are often restricted to relatively few study areas, and document a narrow range of possible impacts upon biota and ecosystem processes. We conducted a study in degraded Mediterranean grasslands dominated by Stipa tenacissima to simultaneously evaluate the effects of shrub encroachment on the structure and composition of multiple biotic community components, and on various indicators of ecosystem function. Shrub encroachment enhanced vascular plant richness, biomass of fungi, actinomycetes and other bacteria, and was linked with greater soil fertility and N mineralization rates. While shrub encroachment may be a widespread phenomenon in drylands, an interpretation that this is an expression of desertification is not universal. Our results suggest that shrub establishment may be an important step in the reversal of desertification processes in the Mediterranean region.

Summary1. Long-term basal area increment (BAI) in Abies pinsapo was studied to investigate the way density-dependent factors modulate the responses of radial growth to climatic stresses in relict stands of a drought-sensitive Mediterranean fir. 2. First, we verified that spatially explicit competition predicts mean A. pinsapo BAI at our study site; i.e. it modulates the degree to which the average climate-driven potential for growth is expressed. Second, we verified that the long-term pattern of temperature predicts the long-term pattern of BAI, estimated as the main trend over a time period of c. 40 years. Finally, we assessed whether the intensity of tree-to-tree competition restrains the potential improvements achieved by our model of BAI when a short-term, high-frequency stressor such as drought (inter-annual precipitation variability) is introduced. 3. We applied Dynamic Factor Analysis (DFA) to characterize regional climatic trends and to test the hypothesis that trees subjected to contrasting competition intensity may differ in their growth pattern. Significant long-term climate trends obtained by DFA were used as predictors of long-term BAI. 4. The mean BAI was mainly determined by competition, whereas growth trends obtained by DFA did not differ among dominant, suppressed and dying trees. Common trends of growth decline were strongly related to long-term, late-winter to summer temperatures, while the residuals were related to total annual precipitation, although with decreasing significance as competition increased. Our results support the contention that the reported patterns of A. pinsapo growth decline and death occur as a result of the interacting effects of both competition and climate stressors acting at longand short-term time scales. 5. Synthesis. Long-term climatic drought stress was the main driving factor of growth decline in A. pinsapo. Moreover, trees already suffering from competition (a long-term stress) were predisposed to decline given an additional short-term stress, such as a severe drought.

Forests play a critical role in the global carbon cycle, being considered an important and continuing carbon sink. However, the response of carbon sequestration in forests to global climate change remains a major uncertainty, with a particularly poor understanding of the origins and environmental responses of soil CO 2 efflux. For example, despite their large biomass, the contribution of ectomycorrhizal (EM) fungi to forest soil CO 2 efflux and responses to changes in environmental drivers has, to date, not been quantified in the field. Their activity is often simplistically included in the 'autotrophic' root respiration term. We set up a multiplexed continuous soil respiration measurement system in a young Lodgepole pine forest, using a mycorrhizal mesh collar design, to monitor the three main soil CO 2 efflux components: root, extraradical mycorrhizal hyphal, and soil heterotrophic respiration.Mycorrhizal hyphal respiration increased during the first month after collar insertion and thereafter remained remarkably stable. During autumn the soil CO 2 flux components could be divided into $ 60% soil heterotrophic, $ 25% EM hyphal, and $15% root fluxes. Thus the extraradical EM mycelium can contribute substantially more to soil CO 2 flux than do roots. While EM hyphal respiration responded strongly to reductions in soil moisture and appeared to be highly dependent on assimilate supply, it did not responded directly to changes in soil temperature. It was mainly the soil heterotrophic flux component that caused the commonly observed exponential relationship with temperature. Our results strongly suggest that accurate modelling of soil respiration, particularly in forest ecosystems, needs to explicitly consider the mycorrhizal mycelium and its dynamic response to specific environmental factors. Moreover, we propose that in forest ecosystems the mycorrhizal CO 2 flux component represents an overflow 'CO 2 tap' through which surplus plant carbon may be returned directly to the atmosphere, thus limiting expected carbon sequestration from trees under elevated CO 2 .

AimsThe combined effects of changes in climate and land use on tree mortality and growth patterns have rarely been addressed. Relict tree species from the Mediterranean Basin serve as appropriate models to investigate these effects, since they grow in climatically stressed areas which have undergone intense cover changes. The aim is to use climate, aerial photographs, stand structure and radial-growth data to explain the mortality and historical patterns of growth of Abies pinsapo in the area where this relict species was first protected.Location Sierra de las Nieves, West Baetic Range, southern Spain. MethodsWe assessed variations of tree cover in A. pinsapo forests through image analyses of aerial photographs spanning the last 50 years. We sampled 31 stands to assess current altitudinal patterns of forest structure and mortality. We evaluated the relationships between radial growth and regional climate using linear models in three sites at different elevations. ResultsRegional warming and a decrease in precipitation were detected. Forest tree cover increased at all elevations from 1957 until 1991, but it afterwards decreased below 1100 m. Currently, the likelihood of tree mortality increases downwards and is associated with dense, closed stands with a low living basal area. In contrast to previous droughts, a sharp synchronized reduction in tree growth, not fully accounted for in linear climate-growth models, occurred at low elevations in 1994-95, but not upwards. It was preceded by a weakening of the negative association between low-elevation growth and water deficit since the late 1970s. ConclusionsThe intense densification of A. pinsapo forests following strict protection measures in the late 1950s enhanced the vulnerability of climate-sensitive A. pinsapo forests to recent drier conditions. Such abrupt land-use changes help to explain recent patterns of mortality and growth decline in low-elevation A. pinsapo forests.