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

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

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

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

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

Key factors affecting the future provision of tree-based forest ecosystem goods and services

The continuous provisioning of forest ecosystem goods and services (EGS) is of considerable interest to society. To provide insights on how much EGS provision will change with a changing climate and which factors will influence this change the most, we simulated forest stands on six climatically different sites in Central Europe under several scenarios of species diversity, management, and climate change. We evaluated the influence of these factors on the provision of a range of tree-based EGS, represented by harvested basal area, total biomass, stand diversity, and productivity. The most influential factor was species diversity, with diverse forest stands showing a lower sensitivity to climate change than monocultures. Management mainly influenced biomass, with the most intensively managed stands retaining more of their original biomass than others. All three climate-change scenarios yielded very similar results. We showed that (1) only few factor combinations perform worse under climate-change conditions than others, (2) diversity aspects are important for adaptive management measures, but for some indicators, management may be more important than diversity, and (3) at locations subject to increasing drought, the future provision of EGS may decrease regardless of the factor combination. This quantitative evaluation of the influence of different factors on changes in the provision of forest EGS with climate change represents an important step towards the design of more focused adaptation strategies and highlights key factors that should be considered in simulation studies under climate chang

The relative importance of land use and climatic change in Alpine catchments

Carbon storage and catchment hydrology are influenced both by land use changes and climatic changes, but there are few studies addressing both responses under both driving forces. We investigated the relative importance of climate change vs. land use change for four Alpine catchments using the LPJ-GUESS model. Two scenarios of grassland management were calibrated based on the more detailed model PROGRASS. The simulations until 2100 show that only reforestation could lead to an increase of carbon storage under climatic change, whereby a cessation of carbon accumulation occurred in all catchments after 2050. The initial increase in carbon storage was attributable mainly to forest re-growth on abandoned land, whereas the stagnation and decline in the second half of the century was mainly driven by climate change. If land was used more intensively, i.e. as grassland, litter input to the soil decreased due to harvesting, resulting in a decline of soil carbon storage (1.2−2.9kgC m-2) that was larger than the climate-induced change (0.8-1.4kgC m−2). Land use change influenced transpiration both directly and in interaction with climate change. The response of forested catchments diverged with climatic change (11-40mm increase in AET), reflecting the differences in forest age, topography and water holding capacity within and between catchments. For grass-dominated catchments, however, transpiration responded in a similar manner to climate change (light management: 23-32mm AET decrease, heavy management: 29-44mm AET decrease), likely because grassroots are concentrated in the uppermost soil layers. Both the water and the carbon cycle were more strongly influenced by land use compared to climatic changes, as land use had not only a direct effect on carbon storage and transpiration, but also an indirect effect by modifying the climate change response of transpiration and carbon flux in the catchments. For the carbon cycle, climate change led to a cessation of the catchment response (sink/source strength is limited), whereas for the water cycle, the effect of land use change remains evident throughout the simulation period (changes in evapotranspiration do not attenuate). Thus we conclude that management will have a large potential to influence the carbon and water cycle, which needs to be considered in management planning as well as in climate and hydrological modellin

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

Evaluation of the forest growth model SILVA along an elevational gradient in Switzerland

The semi-empirical single-tree model SILVA 2.2 has been developed and parameterised using forest research and inventory data from Germany that range from the colline to the montane zone. The focus of the model evaluation presented in this study was to test the applicability of the model for the main Swiss forest types and at elevations ranging from the colline to the upper subalpine zone. To this end, SILVA was initialized using data from long-term forest yield research plots. The results at the end of the 30-year simulation were compared with observed data. The analysis of the results at each test site showed that there were no significant differences in model performance between forest types. However, the deviation between simulated and observed growth depended strongly on the elevational zone, i.e., on climate. As expected, the best results were found in the colline zone, for which the model had been calibrated, whereas the upper subalpine sites revealed the strongest differences. The quality of the data regarding forest structure that were available for model initialization had a strong impact on the simulation results, mainly at high-elevation zones (i.e., supalpine and upper subalpine). We conclude that SILVA 2.2 is a suitable tool to estimate the development of single trees and standing volume for a large fraction of the forests in Switzerland. However, extreme climate conditions should be avoided with the model, and the availability of detailed stand structure information is a key priority that has a strong effect on the quality of the simulation result