Sensitivity of carbon cycling in the European Alps to changes of climate and land cover

Assessments of the impacts of global change on carbon stocks in mountain regions have received little attention to date, in spite of the considerable role of these areas for the global carbon cycle. We used the regional hydro-ecological simulation system RHESSys in five case study catchments from different climatic zones in the European Alps to investigate the behavior of the carbon cycle under changing climatic and land cover conditions derived from the SRES scenarios of the IPCC. The focus of this study was on analyzing the differences in carbon cycling across various climatic zones of the Alps, and to explore the differences between the impacts of various SRES scenarios (A1FI, A2, B1, B2), and between several global circulation models (GCMs, i.e., HadCM3, CGCM2, CSIRO2, PCM). The simulation results indicate that the warming trend generally enhances carbon sequestration in these catchments over the first half of the twenty-first century, particularly in forests just below treeline. Thereafter, forests at low elevations increasingly release carbon as a consequence of the changed balance between growth and respiration processes, resulting in a net carbon source at the catchment scale. Land cover changes have a strong modifying effect on these climate-induced patterns. While the simulated temporal pattern of carbon cycling is qualitatively similar across the five catchments, quantitative differences exist due to the regional differences of the climate and land cover scenarios, with land cover exerting a stronger influence. The differences in the simulations with scenarios derived from several GCMs under one SRES scenario are of the same magnitude as the differences between various SRES scenarios derived from one single GCM, suggesting that the uncertainty in climate model projections needs to be narrowed before accurate impact assessments under the various SRES scenarios can be made at the local to regional scale. We conclude that the carbon balance of the European Alps is likely to shift strongly in the future, driven mainly by land cover changes, but also by changes of the climate. We recommend that assessments of carbon cycling at regional to continental scales should make sure to adequately include sub-regional differences of changes in climate and land cover, particularly in areas with a complex topograph

Ungulate browsing in winter reduces the growth of Fraxinus and Acer saplings in subsequent unbrowsed years

Browsing by ungulates has become a hotly debated issue in many European mountain forests in the past century. Acer pseudoplatanus L. and Fraxinus excelsior L. are broadleaved tree species that are preferentially browsed by roe deer (Capreolus capreolus) in central Europe. We analyzed growth data from shaded saplings of both tree species to quantify the extent to which height growth after game browsing is reduced in subsequent, unbrowsed years in forest stands. Sixty saplings of F. excelsior and A. pseudoplatanus from forest stands at Albisriederberg (Switzerland) were available to us that had been dissected into pieces that then were split in the middle for counting tree rings and assessing ungulate damage. We fitted the von Bertalanffy growth equation to these height growth data and included a reduction factor for winter browsing. Both tree species showed significantly reduced height growth in unbrowsed years after one to several browsing events in winter, and this effect increased with the number of browsing events. Saplings with a high growth rate showed a higher growth reduction. After winter browsing, height growth of A. pseudoplatanus saplings was less affected in unbrowsed years than that of F. excelsior saplings. We conclude that browsed saplings of these species in forest stands are not able to compensate browsing-induced height loss, but that height differences between browsed and unbrowsed saplings probably increase over time. A comparison between our analysis and the parameters estimated using equations published by Eiberle for predicting age at 130cm height suggests that our parameter values are rather conservative estimates of the growth reduction effect after winter browsing. Neither F. excelsior nor A. pseudoplatanus show a distinct pattern in browsing-induced growth reduction with respect to soil moisture, nutrient level, and altitude. We thus conclude that our results are likely to be valid for a wide range of forested site

Predicting decay and ground vegetation development in Picea abies snag stands

In a Picea abies (L.) Karst. (Norway spruce) mountain forest on the Gandberg site in the northern Swiss Alps, trees were killed by bark beetles in 1992-1997. A combination of field studies and dynamic modelling was used to project snag decay and future ground vegetation succession in these steep, unharvested stands. In permanent plots, ground vegetation cover and natural tree regeneration have been monitored annually since 1994. To obtain additional information on the abundance of snags, logs, boulders and other microsite types in these stands, the relative frequency of the microsite types was quantified along four strip transects on the montane and subalpine elevational levels. A dynamic model of snag decay and ground vegetation development was constructed (modified matrix model approach). Based on field data and literature values, the model was parameterised and initialised separately for the montane and the subalpine level. For model validation, microsite types were quantified in 2001 with the line-intercept method on both elevational levels. Starting from the conditions in the stands before the bark beetle attacks, it was possible to project short-term succession and to accurately simulate the decay and ground vegetation patterns eight years after tree die-back. Long-term simulations suggest that on the montane level, raspberries (Rubus idaeus L.) will be replaced by Picea abies, while on the subalpine level ferns will dominate for a long tim

Analyzing the carbon dynamics of central European forests: comparison of Biome-BGC simulations with measurements

Biogeochemical models are often used for making projections of future carbon dynamics under scenarios of global change. The aim of this study was to assess the accuracy of the process-based biogeochemical model Biome-BGC for application in central European forests from the lowlands to upper treeline as a pre-requisite for environmental impact assessments. We analyzed model behavior along an altitudinal gradient across the alpine treeline, which provided insights on the sensitivity of simulated average carbon pools to changes in environmental factors. A second set of tests included medium-term (30years) simulations of carbon fluxes, and a third set of tests focused on daily carbon and water fluxes. Model results were compared to aboveground biomass measurements, leaf area index recordings as well as net ecosystem exchange (NEE) and actual evapotranspiration (AET) measurements. The simulated medium-term forest growth agreed well with measured data. Also daily NEE fluxes were simulated adequately in most cases. Problems were detected when simulating ecosystems close to the upper timberline (overestimation of measured growth and pool sizes), and when simulating daily AET fluxes (overestimation of measured fluxes). The results showed that future applications of Biome-BGC could benefit much from an improvement of model algorithms (e.g., the Q10 model for respiration) as well as from a detailed analysis of the ecological significance of crucial parameters (e.g., the canopy water interception coefficient

Timing, light availability and vigour determine the response of Abies alba saplings to leader shoot browsing

Herbivore browsing on tree saplings is a common phenomenon that can cause damage particularly on preferred species. In this study, the combined effects of light availability and timing of browsing on the response of 9-year-old Abies alba saplings were tested experimentally. Leader shoot clipping was applied before budburst, shortly after budburst or in autumn on saplings grown in full light or under artificial shade. Timing of clipping, light availability and tree vigour (expressed as height and tree ring width before clipping) had an effect on the height after clipping. After clipping in autumn or before budburst, fast-growing fir saplings bent up twigs to form new leader shoots and overcompensated height loss; saplings characterised by intermediate growth rates formed new shoots out of regular visible lateral buds; and slow-growing saplings had no new shoot in the first year after clipping, such that the clipping-induced height difference even increased over time. Saplings clipped shortly after budburst elongated the remaining part of the shoot in the first year and developed shoots out of the most distal lateral buds in the second growing season, leading to complete height compensation. Multi-trunking was typical for all clipped trees. We conclude that the microscale conditions under which a tree is growing (i.e. which affect tree vigour) are highly important for determining whether the height reduction imposed by browsing is offset by overcompensation or increases over time relative to unclipped trees. This response can partly be influenced by forest management via enhancing tree vigour via the light regim

The speed of learning instructed stimulus-response association rules in human: experimental data and model.

Humans can learn associations between visual stimuli and motor responses from just a single instruction. This is known to be a fast process, but how fast is it? To answer this question, we asked participants to learn a briefly presented (200ms) stimulus-response rule, which they then had to rapidly apply after a variable delay of between 50 and 1300ms. Participants showed a longer response time with increased variability for short delays. The error rate was low and did not vary with the delay, showing that participants were able to encode the rule correctly in less than 250ms. This time is close to the fastest synaptic learning speed deemed possible by diffusive influx of AMPA receptors. Learning continued at a slower pace in the delay period and was fully completed in average 900ms after rule presentation onset, when response latencies dropped to levels consistent with basic reaction times. A neural model was proposed that explains the reduction of response times and of their variability with the delay by (i) a random synaptic learning process that generates weights of average values increasing with the learning time, followed by (ii) random crossing of the firing threshold by a leaky integrate-and-fire neuron model, and (iii) assuming that the behavioural response is initiated when all neurons in a pool of m neurons have fired their first spike after input onset. Values of m=2 or 3 were consistent with the experimental data. The proposed model is the simplest solution consistent with neurophysiological knowledge. Additional experiments are suggested to test the hypothesis underlying the model and also to explore forgetting effects for which there were indications for the longer delay conditions. This article is part of a Special Issue entitled Neural Coding 2012

Will the CO2 fertilization effect in forests be offset by reduced tree longevity?

Experimental studies suggest that tree growth is stimulated in a greenhouse atmosphere, leading to faster carbon accumulation (i.e., a higher rate of gap filling). However, higher growth may be coupled with reduced longevity, thus leading to faster carbon release (i.e., a higher rate of gap creation). The net effect of these two counteracting processes is not known. We quantify this net effect on aboveground carbon stocks using a novel combination of data sets and modeling. Data on maximum growth rate and maximum longevity of 141 temperate tree species are used to derive a relationship between growth stimulation and changes in longevity. We employ this relationship to modify the respective parameter values of tree species in a forest succession model and study aboveground biomass in a factorial design of growth stimulation×reduced maximum longevity at multiple sites along a climate gradient from the cold to the dry treeline. The results show that (1) any growth stimulation at the tree level leads to a disproportionately small increase of stand biomass due to negative feedback effects, even in the absence of reduced longevity; (2) a reduction of tree longevity tends to offset the growth-related biomass increase; at the most likely value of reduced longevity, the net effect is very close to zero in most multi- and single-species simulations; and (3) when averaging the response across all sites to mimic a "landscape-level” response, the net effect is close to zero. Thus, it is important to consider ecophysiological responses with their linkage to demographic processes in forest trees if one wishes to avoid erroneous inference at the ecosystem level. We conclude that any CO2 fertilization effect is quite likely to be offset by an associated reduction in the longevity of forest trees, thus strongly reducing the carbon mitigation potential of temperate forest

Ungulate herbivory modifies the effects of climate change on mountain forests

Recent temperature observations suggest a general warming trend that may be causing the range of tree species to shift to higher latitudes and altitudes. Since biotic interactions such as herbivory can change tree species composition, it is important to understand their contribution to vegetation changes triggered by climate change. To investigate the response of forests to climate change and herbivory by wild ungulates, we used the forest gap model ForClim v2.9.6 and simulated forest development in three climatically different valleys in the Swiss Alps. We used altitudinal transects on contrasting slopes covering a wide range of forest types from the cold (upper) to the dry (lower) treeline. This allowed us to investigate (1) altitudinal range shifts in response to climate change, (2) the consequences for tree species composition, and (3) the combined effect of climate change and ungulate herbivory. We found that ungulate herbivory changed species composition and that both basal area and stem numbers decreased with increasing herbivory intensity. Tree species responded differently to the change in climate, and their ranges did not change concurrently, thus causing a succession to new stand types. While climate change partially compensated for the reductions in basal area caused by ungulate herbivory, the combined effect of these two agents on the mix of the dominant species and forest type was non-compensatory, as browsing selectively excluded species from establishing or reaching dominance and altered competition patterns, particularly for light. We conclude that there is an urgent need for adaptive forest management strategies that address the joint effects of climate change and ungulate herbivor

Linking Forest Fire Regimes and Climate—A Historical Analysis in a Dry Inner Alpine Valley

Forest fire regimes are likely to experience considerable changes in the European Alps due to climatic changes. However, little is known about the recent regional fire history and the impact of local climate on the fire regime during the 20th century. We therefore reconstructed the fire history in a dry continental valley of the Swiss Alps (Valais) over the past 100years based on documentary evidence, and investigated the relationship between the reconstructed fire regime and the local climatic variability. We compared the impact of temperature, precipitation, drought and dry foehn winds on fire frequency, extent of burnt area, and fire seasonality on various spatial and temporal scales. In the subalpine zone, the fire regime appears to have been mainly driven by temperature and precipitation, whereas these variables seem to have played only a secondary role in the colline-montane zones. Here, foehn winds and, probably, non-climatic factors seem to have been more important. Temperature and precipitation played a major role in shaping fire frequency and burnt area in the first half of the 20th century, but lost their importance during the second half. Our case study illustrates the occurrence of different fire regime patterns and their driving forces on small spatial scales (a few hundred square kilometers). We conclude that the strong rise in temperature over the past century has not profoundly changed the fire regime in Valais, but in the second half of the 20th century temperature was no longer a strong determinant for forest fires as compared to human activities or biomass availability in forest