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192 S. Schumacher et al. / Ecological Modelling 180 (2004) 175–194 after such events. Studying transitions from weakly to strongly disturbed landscapes is an important issue, as global environmental change is likely to bring about changes in the distribution of extreme climatic events, which may trigger changes in the large-scale disturbance regimes in many landscapes. Because the modified model explicitly incorporates the effects of a range of climatic factors on tree regeneration and growth, it is more suitable for addressing the direct impacts of climatic changes on forested landscapes. Through a simple set of scenario calculations (which should not be mistaken as predictions), we were able to demonstrate that the model is sensitive to changing climatic parameters. Notably, the effects of climatic changes on forest landscape dynamics are emergent properties of the modified model, rather than being prescribed through site-specific or speciesspecific parameters, which was the case in earlier versions of LANDIS. The modified model can also serve as a tool to help with decision-making in the context of longterm forest management. It provides variables—such as biomass, stand density, species dominance, or vertical structure—that can be used for assessing, for example, the habitat requirements of wildlife, and indices that are relevant to a range of other ecological questions. Also, variables, such as stand density and vertical stand structure are crucial for predicting the long-term consequences of forest management decisions, e.g. with respect to the ability of mountain forests to avert natural hazards, such as avalanches, rockfall or landslides. Finally, the ability of the model to simulate landscape-scale aboveground biomass storage allows us to determine the effect of Global Change scenarios on aspects of the carbon balance of mountain ecosystems. Hence, the modified LANDIS model as described in this study has the potential to provide an integrated picture of the impact of both direct and indirect effects of climate change on forest landscape dynamics. Acknowledgment Thanks are due to Hong He, University of Missouri—Columbia, who supplied the LANDIS model and helped us to get acquainted with it. Dean Urban and an anonymous reviewer provided constructive criticisms of an earlier version of this manuscript. References Anonymous, 1983. Ertragstafeln für Buche, Lärche, Fichte und Tanne, third ed., Eidg. Anstalt für das forstliche Versuchswesen, CH-8903 Birmensdorf, Switzerland. Assmann, E., 1961. Waldertragskunde: organische Produktion, Struktur, Zuwachs und Ertag von Waldbeständen. BLV, München a.o. Baker, W.L., 1989. A review of models of landscape change. Landsc. Ecol. 2 (2), 111–133. Bassett, J.R., 1964. Tree growth as affected by soil moisture availability. Soil Sci. Proc. 28, 436–438. Begon, M., Mortimer, M., Thompson, D.J., 1996. Population Ecology: A Unified Study of Animals and Plants. Oxford University Press, Oxford. Botkin, D.B., 1993. Forest dynamics: An ecological model. Oxford University Press, Oxford/New York. Bugmann, H., 1994. On the ecology of mountainous forests in a changing climate: a simulation study. Ph.D. Thesis no. 10638, Swiss Federal Institute of Technology. Zurich, Switzerland. Bugmann, H., 1996. A simplified forest model to study species composition along climate gradients. Ecology 77 (7), 2055–2074. Bugmann, H., 1999. Anthropogene Klimaveränderung, Sukzessionsprozesse und forstwirtschaftliche Optionen. Schweiz. Z. Forstwes. 150, 275–287. Bugmann, H., Cramer, W., 1998. Improving the behaviour of forest gap models along drought gradients. Forest Ecol. Manage. 103 (2–3), 247–263. Bugmann, H., Pfister, C., 2000. Impacts of interannual climate variability on past and future forest composition. Reg. Environ. Change 1, 112–125. Bugmann, H.K.M., Solomon, A.M., 2000. Explaining forest composition and biomass across multiple biogeographical regions. Ecol. Appl. 10 (1), 95–114. Bugmann, H.K.M., Yan, X., Sykes, M.T., Martin, P., Lindner, M., Desanker, P.V., Cumming, S.G., 1996. A comparison of forest gap models: model structure and behaviour. Clim. Change 34, 289–313. Burger, H., 1945. Holz Blattmenge und Zuwachs. Part VII. Die Lärche: Mitt. Schweiz. Anst. forstl. Versuchswes 24, 7–103. Burger, H., 1947. Holz Blattmenge und Zuwachs. Part VIII. Die Eiche: Mitt. Schweiz. Anst. forstl. Versuchswes 25, 211–279. Burger, H., 1948. Holz Blattmenge und Zuwachs. Part IX. Die Föhre: Mitt. Schweiz. Anst. forstl. Versuchswes 25, 435–493. Burger, H., 1950a. Forstliche Versuchsflächen im schweizerischen Nationalpark: Mitt. Schweiz. Anst. forstl. Versuchswes 26, 583–634. Burger, H., 1950b. Holz Blattmenge und Zuwachs. Part X. Die Buche: Mitt. Schweiz. Anst. forstl. Versuchswes 26, 419– 468. Burger, H., 1951. Holz Blattmenge und Zuwachs. Part XI. Die Tanne: Mitt. Schweiz. Anst. forstl. Versuchswes 27, 247–286.
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