Harald Bugmann

Ecosystem service supply and vulnerability to global change in Europe

Global change will alter the supply of ecosystem services that are vital for human well-being. To investigate ecosystem service supply during the 21st century, we used a range of ecosystem models and scenarios of climate and land-use change to conduct a Europe-wide assessment. Large changes in climate and land use typically resulted in large changes in ecosystem service supply. Some of these trends may be positive (for example, increases in forest area and productivity) or offer opportunities (for example, “surplus land” for agricultural extensification and bioenergy production). However, many changes increase vulnerability as a result of a decreasing supply of ecosystem services (for example, declining soil fertility, declining water availability, increasing risk of forest fires), especially in the Mediterranean and mountain regions.

A REVIEW OF FOREST GAP MODELS

Forest gap models, initially conceived in 1969 as a special case of individual-tree based models, have become widely popular among forest ecologists for addressing a large number of applied research questions, including the impacts of global change on long-term dynamics of forest structure, biomass, and composition. However, they have been strongly criticized for a number of weaknesses inherent in the original model structure. In this paper, I review the fundamental assumptions underlying forest gap models, the structure of the parent model JABOWA, and examine these criticisms in the context of the many alternative formulations that have been developed over the past 30 years.Four assumptions originally underlie gap models: (1) The forest is abstracted as a composite of many small patches of land, where each can have a different age and successional stage; (2) patches are horizontally homogeneous, i.e., tree position within a patch is not considered; (3) the leaves of each tree are located in an indefinitely thin layer (disk) at the top of the stem; and (4) successional processes are described on each patch separately, i.e., there are no interactions between patches. These simplifications made it possible to consider mixed-species, mixed-age forests, which had been difficult previously mainly because of computing limitations.The structure of JABOWA is analysed in terms of the functional relationships used for formulating the processes of tree establishment, growth, and mortality. It is concluded that JABOWA contains a number of unrealistic assumptions that have not been questioned strongly to date. At the same time, some aspects of JABOWA that were criticized strongly in the past years are internally consistent given the objectives of this specific model.A wide variety of formulations for growth processes, establishment, and mortality factors have been developed in gap models over the past 30 years, and modern gap models include more robust parameterizations of environmental influences on tree growth and population dynamics as compared to JABOWA. Approaches taken in more recent models that led to the relaxation of one or several of the four basic assumptions are discussed. It is found that the original assumptions often have been replaced by alternatives; however, no systematic analysis of the behavioral effects of these conceptual changes has been attempted to date.The feasibility of including more physiological detail (instead of using relatively simple parameterizations) in forest gap models is discussed, and it is concluded that we often lack the data base to implement such approaches for more than a few commercially important tree species. Hence, it is important to find a compromise between using simplistic parameterizations and expanding gap models with physiology-based functions and parameters that are difficult to estimate. While the modeling of tree growth has received a lot of attention over the past years, much less effort has been spent on improving the formulations of tree establishment and mortality, although these processes are likely to be just as sensitive to global change as tree growth itself. Finally, model validation issues are discussed, and it is found that there is no single data source that can reliably be used for evaluating the behavior of forest gap models; instead, I propose a combination of sensitivity analyses, qualitative examinations of process formulations, and quantitative tests of gap models or selected submodels against various kinds of empirical data to evaluate the usefulness of these models for assessing their utility for predicting the impacts of global change on long-term forest dynamics.

Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland

During the 20th century, high mortality rates of Scots pine (Pinus silvestris L.) have been observed over large areas in the Rhône valley (Valais, Switzerland) and in other dry valleys of the European Alps. In this study, we evaluated drought as a possible inciting factor of Scots pine decline in the Valais. Averaged tree-ring widths, standardized tree-ring series, and estimated annual mortality risks were related to a drought index. Correlations between drought indices and standardized tree-ring series from 11 sites showed a moderate association. Several drought years and drought periods could be detected since 1864 that coincided with decreased growth. Although single, extreme drought years had generally a short-term, reversible effect on tree growth, multi-year drought initiated prolonged growth decreases that increased a tree’s long-term risk of death. Tree death occurred generally several years or even decades after the drought. In conclusion, drought has a limiting effect on tree growth and acts as a bottleneck event in triggering Scots pine decline in the Valais.

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.

EXPLAINING FOREST COMPOSITION AND BIOMASS ACROSS MULTIPLE BIOGEOGRAPHICAL REGIONS

Current scientific concerns regarding the impacts of global change include the responses of forest composition and biomass to rapid changes in climate, and forest gap models have often been used to address this issue. These models reflect the concept that forest composition and biomass in the absence of large-scale disturbance are explained by competition among species for light and other resources in canopy gaps formed when dominant trees die. Since their intiation 25 yr ago, a wide variety of gap models have been developed that are applicable to different forest ecosystems all over the world. Few gap models, however, have proved to be equally valid over a wide range of environmental conditions, a problem on which our work is focused. We previously developed a gap model that is capable of simulating forest composition and biomass in temperate forests of Europe and eastern North America based on a single model structure. In the present study, we extend the model to simulate individual tree species response to strong moisture seasonality and low temperature seasonality, and we modify the widespread parabolic temperature response function to mimic nonlinear increases in growth with increased temperature up to species-specific optimal values. The resulting gap model, ForClim V2.9, generates realistic projections of tree species composition and biomass across a complex gradient of temperature and moisture in the Pacific Northwest of the United States. The model is evaluated against measured basal area and stand structure data at three elevations of the H. J. Andrews LTER site, yielding satisfactory results. The very same model also provides improved estimates of species composition and stand biomass in eastern North America and central Europe, where it originated. This suggests that the model modifications we introduced are indeed generic. Temperate forests other than those we studied here are characterized by climates that are quite similar to the ones in the three study regions. Therefore we are confident that it is possible to explain forest composition and biomass of all major temperate forests by means of a single hypothesis as embodied in a forest gap model.

Disentangling biodiversity and climatic determinants of wood production.

Background Despite empirical support for an increase in ecosystem productivity with species diversity in synthetic systems, there is ample evidence that this relationship is dependent on environmental characteristics, especially in structurally more complex natural systems. Empirical support for this relationship in forests is urgently needed, as these ecosystems play an important role in carbon sequestration. Methodology/Principal Findings We tested whether tree wood production is positively related to tree species richness while controlling for climatic factors, by analyzing 55265 forest inventory plots in 11 forest types across five European countries. On average, wood production was 24% higher in mixed than in monospecific forests. Taken alone, wood production was enhanced with increasing tree species richness in almost all forest types. In some forests, wood production was also greater with increasing numbers of tree types. Structural Equation Modeling indicated that the increase in wood production with tree species richness was largely mediated by a positive association between stand basal area and tree species richness. Mean annual temperature and mean annual precipitation affected wood production and species richness directly. However, the direction and magnitude of the influence of climatic variables on wood production and species richness was not consistent, and vary dependent on forest type. Conclusions Our analysis is the first to find a local scale positive relationship between tree species richness and tree wood production occurring across a continent. Our results strongly support incorporating the role of biodiversity in management and policy plans for forest carbon sequestration.

PREDICTING THE TIME OF TREE DEATH USING DENDROCHRONOLOGICAL DATA

Complex interactions of various environmental factors result in high vari- ability of tree mortality in space and time. Tree mortality functions that are implemented in forest succession models have been suggested to play a key role in assessing forest response to climate change. However, these functions are based on theoretical considerations and are likely to be poor predictors of the timing of tree death, since they do not adequately reflect our understanding of tree mortality processes. In addition, these theoretical mortality functions and most empirical mortality functions have not been tested sufficiently with respect to the accuracy of predicting the time of tree death. We introduce a new approach to modeling tree mortality based on different growth patterns of entire tree-ring series. Dendrochronological data from Picea abies (Norway spruce) in the Swiss Alps were used to calibrate mortality models using logistic regression. The autocorrelation of the data was taken into account by a jackknife variance estimator. Model performance was assessed by two criteria for classification accuracy and three criteria for prediction error. The six models with the highest overall performance correctly classified 71-78% of all dead trees and 73-75% of all living trees, and they predicted 44- 56% of all dead trees to die within 0-15 years prior to the actual year of death. For these six models, a maximum of 1.7% of all dead trees and 5% of all living trees were predicted to die .60 years prior to the last measured year. Models including the relative growth rate and a short-term growth trend as explanatory variables were most reliable with respect to inference and prediction. The generality of the mortality models was successfully tested by applying them to two independent P. abies data sets from climatologically and geologically different areas. We conclude that the methods presented improve our understanding of how tree growth and mortality are related, which results in more accurate mortality models that can ultimately be used to increase the reliability of predictions from models of forest dynamics.

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas-fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ(13) C and δ(18) O in early- and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ(13) C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long-term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.

Forest dynamics and ungulate herbivory: from leaf to landscape

Due primarily to anthropogenic influences such as land use change, exotic species introductions, livestock grazing, altered hunting regimes, and predator control, wild ungulate populations have undergone tremendous shifts in recent decades. Although the result has been endangerment or extirpation in some regions, ungulate numbers have increased to locally or regionally high levels in other areas, causing shifts in plant species composition, problems for forest regeneration, and conflict with humans and domestic livestock. Our ability to find solutions to these problems is limited because we lack sufficient understanding of how ungulate species interact with predators, habitat, forage, competing species, and humans at multiple scales from small foraging patches to large regions. To bring together new findings in this area, encompassing various scales and foci of investigation, an international conference on ‘‘Forest Dynamics and Ungulate Herbivory’’ was held from 3 to 6 October 2001 in Davos, Switzerland. We provide a synthetic overview of the papers contained in this Special Issue, arising from that conference. A companion Special Issue of the Journal for Nature Conservation is devoted to the more management-oriented aspects of forest‐ungulate interactions, entitled ‘‘Forest‐Ungulate Interactions: Monitoring, Modeling and Management.’’ From the papers in this issue, a number of important generalizations emerge to guide our understanding and further research. Our predictive understanding of foraging ecology needs to be generalized and scaled up, if it is to become useful for predicting the consequences of herbivory for broad-scale vegetation dynamics. Further, it is important to move beyond single-factor studies, to embrace the complexity of ungulate‐vegetation interactions. The direction and magnitude of ungulate influences can be difficult to gauge because of complex interactions among species, structural units of vegetation or landscapes, ecosystem processes, and natural disturbances. Researchers need to look beyond the simplistic concept of ‘‘game damage’’, and comprehensively address the direct and indirect effects of ungulates on communities, ecosystems and landscapes. This may require a more systems-oriented, and less species-oriented approach. Ungulate‐vegetation interactions need to be better understood over multiple scales, and particularly at the coarser scales that are of interest to managers and policy-makers. Finally, a long-term view of ungulate‐vegetation interactions, where past, present, and future effects are considered in their appropriate temporal context, is absolutely essential. # 2003 Elsevier Science B.V. All rights reserved.

Radial growth responses to drought of Pinus sylvestris and Quercus pubescens in an inner-Alpine dry valley

Abstract Question: Lower montane treeline ecotones such as the inner Alpine dry valleys are regarded as sensitive to climate change. In the dry Valais valley (Switzerland) the composition of the widespread, low altitude Pinus forests is shifting towards a mixed deciduous state. The sub-boreal P. sylvestris shows high mortality rates, whereas the deciduous sub-mediterranean Quercus pubescens is spreading. These species may act as early indicators of climate change. We evaluate this hypothesis by focusing on their differences in drought tolerance, which are hardly known, but are likely to be crucial in the current forest shift and also for future forest development. Methods: We used dendroecological methods to detect species-specific patterns in the growth response to drought. The relationship between radial growth of 401 trees from 15 mixed stands and drought was analysed by calculating response functions using yearly tree-ring indices and monthly drought indices. PCA was applied to the response ratios to discover spatial patterns of drought response. Results: A species-specific response to moisture as well as a sub-regional differentiation of the response patterns were found. While Quercus showed a response mainly to the conditions of the previous autumn and those of current spring, Pinus did not start responding before May, but showed responses throughout the whole summer. Quercus may restrict physiological activity to moist periods; growth of Pinus was much more dependent on prior growth. Conclusions: Given that the climate is changing towards (1) longer summer drought periods, (2) higher mean temperatures and (3) shifted seasonality of moisture availability, Quercus may benefit from adapting better to drier conditions. Pinus may increasingly face problems related to drought stress as it depends on summer moisture and has a smaller adaptive capacity due to its long-lived photosynthetic tissue.