Articles | Volume 18, issue 14
https://doi.org/10.5194/gmd-18-4643-2025
© Author(s) 2025. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/gmd-18-4643-2025
© Author(s) 2025. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Model description paper
| 
30 Jul 2025
Model description paper |
| 30 Jul 2025
| 30 Jul 2025
Simulating the drought response of European tree species with the dynamic vegetation model LPJ-GUESS (v4.1, 97c552c5)
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
João P. Darela-Filho
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Konstantin Gregor
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Allan Buras
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Qiao-Lin Gu
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Andreas Krause
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Daijun Liu
Department of Botany and Biodiversity Research, University of Vienna, Rennweg 14, 1030 Vienna, Austria
Phillip Papastefanou
Department Biogeochemical Signals, Max Planck Institute for Biogeochemistry, Hans-Knoll-Str., 10, 07745 Jena, Thuringia, Germany
Sijeh Asuk
Department of Geography and Environment, School of Social Sciences and Humanities, Loughborough University, Loughborough, LE11 3TU, UK
Thorsten E. E. Grams
Professorship of Land Surface–Atmosphere Interactions, Ecophysiology of Plants, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Christian S. Zang
Professorship of Forests and Climate Change, University of Applied Sciences Weihenstephan-Triesdorf, Freising, Germany
Anja Rammig
Professorship of Land Surface–Atmosphere Interactions, TUM School of Life Sciences, Technical University of Munich, Freising, Germany
Related authors
Lucia S. Layritz, Konstantin Gregor, Andreas Krause, Stefan Kruse, Benjamin F. Meyer, Thomas A. M. Pugh, and Anja Rammig
Biogeosciences, 22, 3635–3660, https://doi.org/10.5194/bg-22-3635-2025, https://doi.org/10.5194/bg-22-3635-2025, 2025
Short summary
Short summary
Disturbances, such as fire, can change which vegetation grows in a forest, affecting water and carbon flows and, thus, the climate. Disturbances are expected to increase with climate change, but it is uncertain by how much. Using a simulation model, we studied how future climate, disturbances, and their combined effect impact northern (high-latitude) forest ecosystems. Our findings highlight the importance of considering these factors and the need to better understand how disturbances will change in the future.
Konstantin Gregor, Benjamin F. Meyer, Tillmann Gaida, Victor Justo Vasquez, Karina Bett-Williams, Matthew Forrest, João P. Darela-Filho, Sam Rabin, Marcos Longo, Joe R. Melton, Johan Nord, Peter Anthoni, Vladislav Bastrikov, Thomas Colligan, Christine Delire, Michael C. Dietze, George Hurtt, Akihiko Ito, Lasse T. Keetz, Jürgen Knauer, Johannes Köster, Tzu-Shun Lin, Lei Ma, Marie Minvielle, Stefan Olin, Sebastian Ostberg, Hao Shi, Reiner Schnur, Urs Schönenberger, Qing Sun, Peter E. Thornton, and Anja Rammig
EGUsphere, https://doi.org/10.5194/egusphere-2025-1733, https://doi.org/10.5194/egusphere-2025-1733, 2025
This preprint is open for discussion and under review for Geoscientific Model Development (GMD).
Short summary
Short summary
Geoscientific models are crucial for understanding Earth’s processes. However, they sometimes do not adhere to highest software quality standards, and scientific results are often hard to reproduce due to the complexity of the workflows. Here we gather the expertise of 20 modeling groups and software engineers to define best practices for making geoscientific models maintainable, usable, and reproducible. We conclude with an open-source example serving as a reference for modeling communities.
Benjamin F. Meyer, Allan Buras, Konstantin Gregor, Lucia S. Layritz, Adriana Principe, Jürgen Kreyling, Anja Rammig, and Christian S. Zang
Biogeosciences, 21, 1355–1370, https://doi.org/10.5194/bg-21-1355-2024, https://doi.org/10.5194/bg-21-1355-2024, 2024
Short summary
Short summary
Late-spring frost (LSF), critically low temperatures when trees have already flushed their leaves, results in freezing damage leaving trees with reduced ability to perform photosynthesis. Forests with a high proportion of susceptible species like European beech are particularly vulnerable. However, this process is rarely included in dynamic vegetation models (DVMs). We show that the effect on simulated productivity and biomass is substantial, warranting more widespread inclusion of LSF in DVMs.
Lucia S. Layritz, Konstantin Gregor, Andreas Krause, Stefan Kruse, Benjamin F. Meyer, Thomas A. M. Pugh, and Anja Rammig
Biogeosciences, 22, 3635–3660, https://doi.org/10.5194/bg-22-3635-2025, https://doi.org/10.5194/bg-22-3635-2025, 2025
Short summary
Short summary
Disturbances, such as fire, can change which vegetation grows in a forest, affecting water and carbon flows and, thus, the climate. Disturbances are expected to increase with climate change, but it is uncertain by how much. Using a simulation model, we studied how future climate, disturbances, and their combined effect impact northern (high-latitude) forest ecosystems. Our findings highlight the importance of considering these factors and the need to better understand how disturbances will change in the future.
Mateus Dantas de Paula, Tatiana Reichert, Laynara F. Lugli, Erica McGale, Kerstin Pierick, João Paulo Darela-Filho, Liam Langan, Jürgen Homeier, Anja Rammig, and Thomas Hickler
Biogeosciences, 22, 2707–2732, https://doi.org/10.5194/bg-22-2707-2025, https://doi.org/10.5194/bg-22-2707-2025, 2025
Short summary
Short summary
This study explores how plant roots with different forms and functions rely on fungal partnerships for nutrient uptake. This relationship was integrated into a vegetation model and was tested in a tropical forest in Ecuador. The model accurately predicted root traits and showed that without fungi, biomass decreased by up to 80 %. The findings highlight the critical role of fungi in ecosystem processes and suggest that root–fungal interactions should be considered in vegetation models.
Konstantin Gregor, Benjamin F. Meyer, Tillmann Gaida, Victor Justo Vasquez, Karina Bett-Williams, Matthew Forrest, João P. Darela-Filho, Sam Rabin, Marcos Longo, Joe R. Melton, Johan Nord, Peter Anthoni, Vladislav Bastrikov, Thomas Colligan, Christine Delire, Michael C. Dietze, George Hurtt, Akihiko Ito, Lasse T. Keetz, Jürgen Knauer, Johannes Köster, Tzu-Shun Lin, Lei Ma, Marie Minvielle, Stefan Olin, Sebastian Ostberg, Hao Shi, Reiner Schnur, Urs Schönenberger, Qing Sun, Peter E. Thornton, and Anja Rammig
EGUsphere, https://doi.org/10.5194/egusphere-2025-1733, https://doi.org/10.5194/egusphere-2025-1733, 2025
This preprint is open for discussion and under review for Geoscientific Model Development (GMD).
Short summary
Short summary
Geoscientific models are crucial for understanding Earth’s processes. However, they sometimes do not adhere to highest software quality standards, and scientific results are often hard to reproduce due to the complexity of the workflows. Here we gather the expertise of 20 modeling groups and software engineers to define best practices for making geoscientific models maintainable, usable, and reproducible. We conclude with an open-source example serving as a reference for modeling communities.
Friedrich J. Bohn, Ana Bastos, Romina Martin, Anja Rammig, Niak Sian Koh, Giles B. Sioen, Bram Buscher, Louise Carver, Fabrice DeClerck, Moritz Drupp, Robert Fletcher, Matthew Forrest, Alexandros Gasparatos, Alex Godoy-Faúndez, Gregor Hagedorn, Martin C. Hänsel, Jessica Hetzer, Thomas Hickler, Cornelia B. Krug, Stasja Koot, Xiuzhen Li, Amy Luers, Shelby Matevich, H. Damon Matthews, Ina C. Meier, Mirco Migliavacca, Awaz Mohamed, Sungmin O, David Obura, Ben Orlove, Rene Orth, Laura Pereira, Markus Reichstein, Lerato Thakholi, Peter H. Verburg, and Yuki Yoshida
Biogeosciences, 22, 2425–2460, https://doi.org/10.5194/bg-22-2425-2025, https://doi.org/10.5194/bg-22-2425-2025, 2025
Short summary
Short summary
An interdisciplinary collaboration of 36 international researchers from 35 institutions highlights recent findings in biosphere research. Within eight themes, they discuss issues arising from climate change and other anthropogenic stressors and highlight the co-benefits of nature-based solutions and ecosystem services. Based on an analysis of these eight topics, we have synthesized four overarching insights.
Mateus Dantas de Paula, Matthew Forrest, David Warlind, João Paulo Darela Filho, Katrin Fleischer, Anja Rammig, and Thomas Hickler
Geosci. Model Dev., 18, 2249–2274, https://doi.org/10.5194/gmd-18-2249-2025, https://doi.org/10.5194/gmd-18-2249-2025, 2025
Short summary
Short summary
Our study maps global nitrogen (N) and phosphorus (P) availability and how they changed from 1901 to 2018. We find that tropical regions are mostly P-limited, while temperate and boreal areas face N limitations. Over time, P limitation increased, especially in the tropics, while N limitation decreased. These shifts are key to understanding global plant growth and carbon storage, highlighting the importance of including P dynamics in ecosystem models.
Olivier Bouriaud, Ernst-Detlef Schulze, Konstantin Gregor, Issam Bourkhris, Peter Högberg, Roland Irslinger, Phillip Papastefanou, Julia Pongratz, Anja Rammig, Riccardo Valentini, and Christian Körner
EGUsphere, https://doi.org/10.5194/egusphere-2024-3092, https://doi.org/10.5194/egusphere-2024-3092, 2024
Short summary
Short summary
The impact of harvesting on forests' carbon sink capacities is debated. One view is that their sink strength is resilient to harvesting, the other that it disrupts these capacities. Our work shows that leaf area index (LAI) has been overlooked in this discussion. We found that temperate forests' carbon uptake is largely insensitive to variations in LAI beyond about 4 m² m-², but that forests operate at higher levels.
Melanie A. Thurner, Silvia Caldararu, Jan Engel, Anja Rammig, and Sönke Zaehle
Biogeosciences, 21, 1391–1410, https://doi.org/10.5194/bg-21-1391-2024, https://doi.org/10.5194/bg-21-1391-2024, 2024
Short summary
Short summary
Due to their crucial role in terrestrial ecosystems, we implemented mycorrhizal fungi into the QUINCY terrestrial biosphere model. Fungi interact with mineral and organic soil to support plant N uptake and, thus, plant growth. Our results suggest that the effect of mycorrhizal interactions on simulated ecosystem dynamics is minor under constant environmental conditions but necessary to reproduce and understand observed patterns under changing conditions, such as rising atmospheric CO2.
Benjamin F. Meyer, Allan Buras, Konstantin Gregor, Lucia S. Layritz, Adriana Principe, Jürgen Kreyling, Anja Rammig, and Christian S. Zang
Biogeosciences, 21, 1355–1370, https://doi.org/10.5194/bg-21-1355-2024, https://doi.org/10.5194/bg-21-1355-2024, 2024
Short summary
Short summary
Late-spring frost (LSF), critically low temperatures when trees have already flushed their leaves, results in freezing damage leaving trees with reduced ability to perform photosynthesis. Forests with a high proportion of susceptible species like European beech are particularly vulnerable. However, this process is rarely included in dynamic vegetation models (DVMs). We show that the effect on simulated productivity and biomass is substantial, warranting more widespread inclusion of LSF in DVMs.
João Paulo Darela-Filho, Anja Rammig, Katrin Fleischer, Tatiana Reichert, Laynara Figueiredo Lugli, Carlos Alberto Quesada, Luis Carlos Colocho Hurtarte, Mateus Dantas de Paula, and David M. Lapola
Earth Syst. Sci. Data, 16, 715–729, https://doi.org/10.5194/essd-16-715-2024, https://doi.org/10.5194/essd-16-715-2024, 2024
Short summary
Short summary
Phosphorus (P) is crucial for plant growth, and scientists have created models to study how it interacts with carbon cycle in ecosystems. To apply these models, it is important to know the distribution of phosphorus in soil. In this study we estimated the distribution of phosphorus in the Amazon region. The results showed a clear gradient of soil development and P content. These maps can help improve ecosystem models and generate new hypotheses about phosphorus availability in the Amazon.
Jennifer A. Holm, David M. Medvigy, Benjamin Smith, Jeffrey S. Dukes, Claus Beier, Mikhail Mishurov, Xiangtao Xu, Jeremy W. Lichstein, Craig D. Allen, Klaus S. Larsen, Yiqi Luo, Cari Ficken, William T. Pockman, William R. L. Anderegg, and Anja Rammig
Biogeosciences, 20, 2117–2142, https://doi.org/10.5194/bg-20-2117-2023, https://doi.org/10.5194/bg-20-2117-2023, 2023
Short summary
Short summary
Unprecedented climate extremes (UCEs) are expected to have dramatic impacts on ecosystems. We present a road map of how dynamic vegetation models can explore extreme drought and climate change and assess ecological processes to measure and reduce model uncertainties. The models predict strong nonlinear responses to UCEs. Due to different model representations, the models differ in magnitude and trajectory of forest loss. Therefore, we explore specific plant responses that reflect knowledge gaps.
Johannes Oberpriller, Christine Herschlein, Peter Anthoni, Almut Arneth, Andreas Krause, Anja Rammig, Mats Lindeskog, Stefan Olin, and Florian Hartig
Geosci. Model Dev., 15, 6495–6519, https://doi.org/10.5194/gmd-15-6495-2022, https://doi.org/10.5194/gmd-15-6495-2022, 2022
Short summary
Short summary
Understanding uncertainties of projected ecosystem dynamics under environmental change is of immense value for research and climate change policy. Here, we analyzed these across European forests. We find that uncertainties are dominantly induced by parameters related to water, mortality, and climate, with an increasing importance of climate from north to south. These results highlight that climate not only contributes uncertainty but also modifies uncertainties in other ecosystem processes.
Phillip Papastefanou, Christian S. Zang, Zlatan Angelov, Aline Anderson de Castro, Juan Carlos Jimenez, Luiz Felipe Campos De Rezende, Romina C. Ruscica, Boris Sakschewski, Anna A. Sörensson, Kirsten Thonicke, Carolina Vera, Nicolas Viovy, Celso Von Randow, and Anja Rammig
Biogeosciences, 19, 3843–3861, https://doi.org/10.5194/bg-19-3843-2022, https://doi.org/10.5194/bg-19-3843-2022, 2022
Short summary
Short summary
The Amazon rainforest has been hit by multiple severe drought events. In this study, we assess the severity and spatial extent of the extreme drought years 2005, 2010 and 2015/16 in the Amazon. Using nine different precipitation datasets and three drought indicators we find large differences in drought stress across the Amazon region. We conclude that future studies should use multiple rainfall datasets and drought indicators when estimating the impact of drought stress in the Amazon region.
Mats Lindeskog, Benjamin Smith, Fredrik Lagergren, Ekaterina Sycheva, Andrej Ficko, Hans Pretzsch, and Anja Rammig
Geosci. Model Dev., 14, 6071–6112, https://doi.org/10.5194/gmd-14-6071-2021, https://doi.org/10.5194/gmd-14-6071-2021, 2021
Short summary
Short summary
Forests play an important role in the global carbon cycle and for carbon storage. In Europe, forests are intensively managed. To understand how management influences carbon storage in European forests, we implement detailed forest management into the dynamic vegetation model LPJ-GUESS. We test the model by comparing model output to typical forestry measures, such as growing stock and harvest data, for different countries in Europe.
Gilvan Sampaio, Marília H. Shimizu, Carlos A. Guimarães-Júnior, Felipe Alexandre, Marcelo Guatura, Manoel Cardoso, Tomas F. Domingues, Anja Rammig, Celso von Randow, Luiz F. C. Rezende, and David M. Lapola
Biogeosciences, 18, 2511–2525, https://doi.org/10.5194/bg-18-2511-2021, https://doi.org/10.5194/bg-18-2511-2021, 2021
Short summary
Short summary
The impact of large-scale deforestation and the physiological effects of elevated atmospheric CO2 on Amazon rainfall are systematically compared in this study. Our results are remarkable in showing that the two disturbances cause equivalent rainfall decrease, though through different causal mechanisms. These results highlight the importance of not only curbing regional deforestation but also reducing global CO2 emissions to avoid climatic changes in the Amazon.
Anita D. Bayer, Richard Fuchs, Reinhard Mey, Andreas Krause, Peter H. Verburg, Peter Anthoni, and Almut Arneth
Earth Syst. Dynam., 12, 327–351, https://doi.org/10.5194/esd-12-327-2021, https://doi.org/10.5194/esd-12-327-2021, 2021
Short summary
Short summary
Many projections of future land-use/-cover exist. We evaluate a number of these and determine the variability they cause in ecosystems and their services. We found that projections differ a lot in regional patterns, with some patterns being at least questionable in a historical context. Across ecosystem service indicators, resulting variability until 2040 was highest in crop production. Results emphasize that such variability should be acknowledged in assessments of future ecosystem provisions.
Cited articles
Adams, H. D., Zeppel, M. J. B., Anderegg, W. R. L., Hartmann, H., Landhäusser, S. M., Tissue, D. T., Huxman, T. E., Hudson, P. J., Franz, T. E., Allen, C. D., Anderegg, L. D. L., Barron-Gafford, G. A., Beerling, D. J., Breshears, D. D., Brodribb, T. J., Bugmann, H., Cobb, R. C., Collins, A. D., Dickman, L. T., Duan, H., Ewers, B. E., Galiano, L., Galvez, D. A., Garcia-Forner, N., Gaylord, M. L., Germino, M. J., Gessler, A., Hacke, U. G., Hakamada, R., Hector, A., Jenkins, M. W., Kane, J. M., Kolb, T. E., Law, D. J., Lewis, J. D., Limousin, J.-M., Love, D. M., Macalady, A. K., Martínez-Vilalta, J., Mencuccini, M., Mitchell, P. J., Muss, J. D., O'Brien, M. J., O'Grady, A. P., Pangle, R. E., Pinkard, E. A., Piper, F. I., Plaut, J. A., Pockman, W. T., Quirk, J., Reinhardt, K., Ripullone, F., Ryan, M. G., Sala, A., Sevanto, S., Sperry, J. S., Vargas, R., Vennetier, M., Way, D. A., Xu, C., Yepez, E. A., and Mcdowell, N. G.: A Multi-Species Synthesis of Physiological Mechanisms in Drought-Induced Tree Mortality, Nat. Ecol. Evol., 1, 1285–1291, https://doi.org/10.1038/s41559-017-0248-x, 2017. a
Ahlström, A., Schurgers, G., Arneth, A., and Smith, B.: Robustness and Uncertainty in Terrestrial Ecosystem Carbon Response to CMIP5 Climate Change Projections, Environ. Res. Lett., 7, 044008, https://doi.org/10.1088/1748-9326/7/4/044008, 2012. a
Allen, C. D., Macalady, A. K., Chenchouni, H., Bachelet, D., McDowell, N., Vennetier, M., Kitzberger, T., Rigling, A., Breshears, D. D., Hogg, E. H. T., Gonzalez, P., Fensham, R., Zhang, Z., Castro, J., Demidova, N., Lim, J.-H., Allard, G., Running, S. W., Semerci, A., and Cobb, N.: A Global Overview of Drought and Heat-Induced Tree Mortality Reveals Emerging Climate Change Risks for Forests, Forest Ecol. Manage., 259, 660–684, https://doi.org/10.1016/j.foreco.2009.09.001, 2010. a, b
Allen, C. D., Breshears, D. D., and McDowell, N. G.: On Underestimation of Global Vulnerability to Tree Mortality and Forest Die-off from Hotter Drought in the Anthropocene, Ecosphere, 6, art129, https://doi.org/10.1890/ES15-00203.1, 2015. a
Allen, R. G., Pereira, L. S., Raes, D., and Smith, M.: Crop evapotranspiration – guidelines for computing crop water requirements, Food and Agriculture Organization of the United Nations, https://www.fao.org/3/x0490e/x0490e00.htm (last access: 16 October 2024), 1998. a
Anderegg, W. R. L., Berry, J. A., Smith, D. D., Sperry, J. S., Anderegg, L. D. L., and Field, C. B.: The Roles of Hydraulic and Carbon Stress in a Widespread Climate-Induced Forest Die-Off, P. Natl. Acad. Sci. USA, 109, 233–237, https://doi.org/10.1073/pnas.1107891109, 2012. a
Anderegg, W. R. L., Hicke, J. A., Fisher, R. A., Allen, C. D., Aukema, J., Bentz, B., Hood, S., Lichstein, J. W., Macalady, A. K., McDowell, N., Pan, Y., Raffa, K., Sala, A., Shaw, J. D., Stephenson, N. L., Tague, C., and Zeppel, M.: Tree Mortality from Drought, Insects, and Their Interactions in a Changing Climate, New Phytol., 208, 674–683, https://doi.org/10.1111/nph.13477, 2015. a, b, c
Anderegg, W. R. L., Klein, T., Bartlett, M., Sack, L., Pellegrini, A. F. A., Choat, B., and Jansen, S.: Meta-Analysis Reveals That Hydraulic Traits Explain Cross-Species Patterns of Drought-Induced Tree Mortality across the Globe, P. Natl. Acad. Sci. USA, 113, 5024–5029, https://doi.org/10.1073/pnas.1525678113, 2016. a, b, c, d
Arend, M., Link, R. M., Patthey, R., Hoch, G., Schuldt, B., and Kahmen, A.: Rapid Hydraulic Collapse as Cause of Drought-Induced Mortality in Conifers, P. Natl. Acad. Sci. USA, 118, e2025251118, https://doi.org/10.1073/pnas.2025251118, 2021. a
Arend, M., Link, R. M., Zahnd, C., Hoch, G., Schuldt, B., and Kahmen, A.: Lack of Hydraulic Recovery as a Cause of Post-drought Foliage Reduction and Canopy Decline in European Beech, New Phytol., 234, 1195–1205, https://doi.org/10.1111/nph.18065, 2022. a, b
Bigler, C. and Vitasse, Y.: Premature Leaf Discoloration of European Deciduous Trees Is Caused by Drought and Heat in Late Spring and Cold Spells in Early Fall, Agr. Forest Meteorol., 307, 108492, https://doi.org/10.1016/j.agrformet.2021.108492, 2021. a
Bigler, C., Bräker, O. U., Bugmann, H., Dobbertin, M., and Rigling, A.: Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland, Ecosystems, 9, 330–343, https://doi.org/10.1007/s10021-005-0126-2, 2006. a, b
Blackman, C. J., Brodribb, T. J., and Jordan, G. J.: Leaf Hydraulic Vulnerability Is Related to Conduit Dimensions and Drought Resistance across a Diverse Range of Woody Angiosperms, New Phytol., 188, 1113–1123, https://doi.org/10.1111/j.1469-8137.2010.03439.x, 2010. a
Bonan, G. B.: Forests and Climate Change: Forcings, Feedbacks, and the Climate Benefits of Forests, Science, 320, 1444–1449, https://doi.org/10.1126/science.1155121, 2008. a
Brodribb, T. J., Powers, J., Cochard, H., and Choat, B.: Hanging by a Thread? Forests and Drought, Science, 368, 261–266, https://doi.org/10.1126/science.aat7631, 2020. a
Buras, A., Meyer, B., and Rammig, A.: Record reduction in European forest canopy greenness during the 2022 drought, EGU General Assembly 2023, Vienna, Austria, 24–28 April 2023, EGU23-8927, https://doi.org/10.5194/egusphere-egu23-8927, 2023. a
Cabon, A. and Anderegg, W. R. L.: Turgor-Driven Tree Growth: Scaling-up Sink Limitations from the Cell to the Forest, Tree Physiol., 42, 225–228, https://doi.org/10.1093/treephys/tpab146, 2022. a
Cabon, A., Peters, R. L., Fonti, P., Martínez-Vilalta, J., and De Cáceres, M.: Temperature and Water Potential Co-Limit Stem Cambial Activity along a Steep Elevational Gradient, New Phytol., 226, 1325–1340, https://doi.org/10.1111/nph.16456, 2020. a
Carminati, A. and Javaux, M.: Soil Rather Than Xylem Vulnerability Controls Stomatal Response to Drought, Trends Plant Sci., 25, 868–880, https://doi.org/10.1016/j.tplants.2020.04.003, 2020. a
Carnicer, J., Coll, M., Ninyerola, M., Pons, X., Sánchez, G., and Peñuelas, J.: Widespread Crown Condition Decline, Food Web Disruption, and Amplified Tree Mortality with Increased Climate Change-Type Drought, P. Natl. Acad. Sci. USA, 108, 1474–1478, https://doi.org/10.1073/pnas.1010070108, 2011. a
Choat, B., Jansen, S., Brodribb, T. J., Cochard, H., Delzon, S., Bhaskar, R., Bucci, S. J., Feild, T. S., Gleason, S. M., Hacke, U. G., Jacobsen, A. L., Lens, F., Maherali, H., Martínez-Vilalta, J., Mayr, S., Mencuccini, M., Mitchell, P. J., Nardini, A., Pittermann, J., Pratt, R. B., Sperry, J. S., Westoby, M., Wright, I. J., and Zanne, A. E.: Global Convergence in the Vulnerability of Forests to Drought, Nature, 491, 752–755, https://doi.org/10.1038/nature11688, 2012. a, b, c, d, e, f
Christoffersen, B. O., Gloor, M., Fauset, S., Fyllas, N. M., Galbraith, D. R., Baker, T. R., Kruijt, B., Rowland, L., Fisher, R. A., Binks, O. J., Sevanto, S., Xu, C., Jansen, S., Choat, B., Mencuccini, M., McDowell, N. G., and Meir, P.: Linking Hydraulic Traits to Tropical Forest Function in a Size-Structured and Trait-Driven Model (TFS v.1-Hydro), Geosci. Model Dev., 9, 4227–4255, https://doi.org/10.5194/gmd-9-4227-2016, 2016. a, b
Ciais, P., Reichstein, M., Viovy, N., Granier, A., Ogée, J., Allard, V., Aubinet, M., Buchmann, N., Bernhofer, C., Carrara, A., Chevallier, F., Noblet, N. D., Friend, A. D., Friedlingstein, P., Grünwald, T., Heinesch, B., Keronen, P., Knohl, A., Krinner, G., Loustau, D., Manca, G., Matteucci, G., Miglietta, F., Ourcival, J. M., Papale, D., Pilegaard, K., Rambal, S., Seufert, G., Soussana, J. F., Sanz, M. J., Schulze, E. D., Vesala, T., and Valentini, R.: Europe-Wide Reduction in Primary Productivity Caused by the Heat and Drought in 2003, Nature, 437, 529–533, https://doi.org/10.1038/nature03972, 2005. a, b, c
Cochard, H., Badel, E., Herbette, S., Delzon, S., Choat, B., and Jansen, S.: Methods for Measuring Plant Vulnerability to Cavitation: A Critical Review, J. Exp. Bot., 64, 4779–4791, https://doi.org/10.1093/jxb/ert193, 2013. a
Cochard, H., Pimont, F., Ruffault, J., and Martin-StPaul, N.: SurEau: A Mechanistic Model of Plant Water Relations under Extreme Drought, Ann. Forest Sci., 78, 55, https://doi.org/10.1007/s13595-021-01067-y, 2021. a
Collatz, G. J., Ball, J. T., Grivet, C., and Berry, J. A.: Physiological and environmental regulation of stomatal conductance, photosynthesis and transpiration: a model that includes a laminar boundary layer, Agr. Forest Meteorol., 54, 107–136, https://doi.org/10.1016/0168-1923(91)90002-8, 1991. a
Collier, N., Hoffman, F. M., Lawrence, D. M., Keppel-Aleks, G., Koven, C. D., Riley, W. J., Mu, M., and Randerson, J. T.: The International Land Model Benchmarking (ILAMB) System: Design, Theory, and Implementation, J. Adv. Model. Earth Syst., 10, 2731–2754, https://doi.org/10.1029/2018MS001354, 2018. a
Comins, H. N. and McMurtrie, R. E.: Long-Term Response of Nutrient-Limited Forests to CO2 Enrichment; Equilibrium Behavior of Plant-Soil Models, Ecol. Appl., 3, 666–681, https://doi.org/10.2307/1942099, 1993. a
Cook, B. I., Mankin, J. S., Marvel, K., Williams, A. P., Smerdon, J. E., and Anchukaitis, K. J.: Twenty-First Century Drought Projections in the CMIP6 Forcing Scenarios, Earth's Future, 8, e2019EF001461, https://doi.org/10.1029/2019EF001461, 2020. a
Da Sois, L., Mencuccini, M., Castells, E., Sanchez-Martinez, P., and Martínez-Vilalta, J.: How Are Physiological Responses to Drought Modulated by Water Relations and Leaf Economics' Traits in Woody Plants?, Agr. Water Manage., 291, 108613, https://doi.org/10.1016/j.agwat.2023.108613, 2024. a
De Kauwe, M. G., Medlyn, B. E., Ukkola, A. M., Mu, M., Sabot, M. E. B., Pitman, A. J., Meir, P., Cernusak, L. A., Rifai, S. W., Choat, B., Tissue, D. T., Blackman, C. J., Li, X., Roderick, M., and Briggs, P. R.: Identifying Areas at Risk of Drought-Induced Tree Mortality across South-Eastern Australia, Global Change Biol., 26, 5716–5733, https://doi.org/10.1111/gcb.15215, 2020. a
Desprez-Loustau, M.-L., Marçais, B., Nageleisen, L.-M., Piou, D., and Vannini, A.: Interactive Effects of Drought and Pathogens in Forest Trees, Ann. Forest Sci., 63, 597–612, https://doi.org/10.1051/forest:2006040, 2006. a
Dietze, M. C., Fox, A., Beck-Johnson, L. M., Betancourt, J. L., Hooten, M. B., Jarnevich, C. S., Keitt, T. H., Kenney, M. A., Laney, C. M., Larsen, L. G., Loescher, H. W., Lunch, C. K., Pijanowski, B. C., Randerson, J. T., Read, E. K., Tredennick, A. T., Vargas, R., Weathers, K. C., and White, E. P.: Iterative Near-Term Ecological Forecasting: Needs, Opportunities, and Challenges, P. Natl. Acad. Sci. USA, 115, 1424–1432, https://doi.org/10.1073/pnas.1710231115, 2018. a
Eckes-Shephard, A. H., Tiavlovsky, E., Chen, Y., Fonti, P., and Friend, A. D.: Direct Response of Tree Growth to Soil Water and Its Implications for Terrestrial Carbon Cycle Modelling, Global Change Biol., 27, 121–135, https://doi.org/10.1111/gcb.15397, 2021. a
Eller, C. B., Rowland, L., Oliveira, R. S., Bittencourt, P. R. L., Barros, F. V., da Costa, A. C. L., Meir, P., Friend, A. D., Mencuccini, M., Sitch, S., and Cox, P.: Modelling Tropical Forest Responses to Drought and El Niño with a Stomatal Optimization Model Based on Xylem Hydraulics, Philos. T. Roy. Soc. B, 373, 20170315, https://doi.org/10.1098/rstb.2017.0315, 2018. a, b
Eller, C. B., Rowland, L., Mencuccini, M., Rosas, T., Williams, K., Harper, A., Medlyn, B. E., Wagner, Y., Klein, T., Teodoro, G. S., Oliveira, R. S., Matos, I. S., Rosado, B. H. P., Fuchs, K., Wohlfahrt, G., Montagnani, L., Meir, P., Sitch, S., and Cox, P. M.: Stomatal Optimization Based on Xylem Hydraulics (SOx) Improves Land Surface Model Simulation of Vegetation Responses to Climate, New Phytol., 226, 1622–1637, https://doi.org/10.1111/nph.16419, 2020. a, b
European Environment Agency: Global and European Temperature, https://www.eea.europa.eu/data-and-maps/indicators/global-and-european-temperature-9/assessment (last access: 19 August 2024), 2019. a
Fink, A. H., Brücher, T., Krüger, A., Leckebusch, G. C., Pinto, J. G., and Ulbrich, U.: The 2003 European Summer Heatwaves and Drought – Synoptic Diagnosis and Impacts, Weather, 59, 209–216, https://doi.org/10.1256/wea.73.04, 2004. a
Fisher, R. A., Williams, M., Do Vale, R. L., Da Costa, A. L., and Meir, P.: Evidence from Amazonian Forests Is Consistent with Isohydric Control of Leaf Water Potential, Plant Cell Environ., 29, 151–165, https://doi.org/10.1111/j.1365-3040.2005.01407.x, 2006. a
Flexas, J., Scoffoni, C., Gago, J., and Sack, L.: Leaf Mesophyll Conductance and Leaf Hydraulic Conductance: An Introduction to Their Measurement and Coordination, J. Exp. Bot., 64, 3965–3981, https://doi.org/10.1093/jxb/ert319, 2013. a
Flo, V., Martínez-Vilalta, J., Granda, V., Mencuccini, M., and Poyatos, R.: Vapour Pressure Deficit Is the Main Driver of Tree Canopy Conductance across Biomes, Agr. Forest Meteorol., 322, 109029, https://doi.org/10.1016/j.agrformet.2022.109029, 2022. a
Frei, E. R., Gossner, M. M., Vitasse, Y., Queloz, V., Dubach, V., Gessler, A., Ginzler, C., Hagedorn, F., Meusburger, K., Moor, M., Samblás Vives, E., Rigling, A., Uitentuis, I., von Arx, G., and Wohlgemuth, T.: European Beech Dieback after Premature Leaf Senescence during the 2018 Drought in Northern Switzerland, Plant Biol., 24, 1132–1145, https://doi.org/10.1111/plb.13467, 2022. a
Gampe, D., Zscheischler, J., Reichstein, M., O'Sullivan, M., Smith, W. K., Sitch, S., and Buermann, W.: Increasing Impact of Warm Droughts on Northern Ecosystem Productivity over Recent Decades, Nat. Clim. Change, 11, 772–779, https://doi.org/10.1038/s41558-021-01112-8, 2021. a
Génard, M., Fishman, S., Vercambre, G., Huguet, J.-G., Bussi, C., Besset, J., and Habib, R.: A Biophysical Analysis of Stem and Root Diameter Variations in Woody Plants, Plant Physiol., 126, 188–202, 2001. a
Gerten, D., Schaphoff, S., Haberlandt, U., Lucht, W., and Sitch, S.: Terrestrial Vegetation and Water Balance – Hydrological Evaluation of a Dynamic Global Vegetation Model, J. Hydrol., 286, 249–270, https://doi.org/10.1016/j.jhydrol.2003.09.029, 2004. a
Guada, G., Camarero, J. J., Sánchez-Salguero, R., and Cerrillo, R. M. N.: Limited Growth Recovery after Drought-Induced Forest Dieback in Very Defoliated Trees of Two Pine Species, Front. Plant Sci., 7, 418, https://doi.org/10.3389/fpls.2016.00418, 2016. a
Hajek, P., Link, R. M., Nock, C. A., Bauhus, J., Gebauer, T., Gessler, A., Kovach, K., Messier, C., Paquette, A., Saurer, M., Scherer-Lorenzen, M., Rose, L., and Schuldt, B.: Mutually Inclusive Mechanisms of Drought-induced Tree Mortality, Global Change Biol., 28, 3365–3378, https://doi.org/10.1111/gcb.16146, 2022. a, b, c
Hammond, W. M., Yu, K., Wilson, L. A., Will, R. E., Anderegg, W. R. L., and Adams, H. D.: Dead or Dying? Quantifying the Point of No Return from Hydraulic Failure in Drought-induced Tree Mortality, New Phytol., 223, 1834–1843, https://doi.org/10.1111/nph.15922, 2019. a, b, c
Hartmann, H., Bastos, A., Das, A. J., Esquivel-Muelbert, A., Hammond, W. M., Martínez-Vilalta, J., McDowell, N. G., Powers, J. S., Pugh, T. A. M., Ruthrof, K. X., and Allen, C. D.: Climate Change Risks to Global Forest Health: Emergence of Unexpected Events of Elevated Tree Mortality Worldwide, Annu. Rev. Plant Biol., 73, 673–702, https://doi.org/10.1146/annurev-arplant-102820-012804, 2022. a
Haxeltine, A. and Prentice, I. C.: BIOME3: An Equilibrium Terrestrial Biosphere Model Based on Ecophysiological Constraints, Resource Availability, and Competition among Plant Functional Types, Global Biogeochem. Cy., 10, 693–709, https://doi.org/10.1029/96GB02344, 1996. a, b, c, d
Helton, J. C. and Davis, F. J.: Latin Hypercube Sampling and the Propagation of Uncertainty in Analyses of Complex Systems, Reliabil. Eng. Syst. Safe., 81, 23–69, https://doi.org/10.1016/S0951-8320(03)00058-9, 2003. a
Hickler, T., Prentice, I. C., Smith, B., Sykes, M. T., and Zaehle, S.: Implementing plant hydraulic architecture within the LPJ Dynamic Global Vegetation Model, Global Ecol. Biogeogr., 15, 567–577, https://doi.org/10.1111/j.1466-8238.2006.00254.x, 2006. a, b, c
Hickler, T., Vohland, K., Feehan, J., Miller, P. A., Smith, B., Costa, L., Giesecke, T., Fronzek, S., Carter, T. R., Cramer, W., Kühn, I., and Sykes, M. T.: Projecting the Future Distribution of European Potential Natural Vegetation Zones with a Generalized, Tree Species-Based Dynamic Vegetation Model: Future Changes in European Vegetation Zones, Global Ecol. Biogeogr., 21, 50–63, https://doi.org/10.1111/j.1466-8238.2010.00613.x, 2012. a, b
IIASA – International Institute for Applied Systems Analysis: Harmonized World Soil Database Version 2.0, FAO,, ISBN 978-92-5-137499-3, https://doi.org/10.4060/cc3823en, 2023. a
Jansen, M. J.: Analysis of Variance Designs for Model Output, Comput. Phys. Commun., 117, 35–43, https://doi.org/10.1016/S0010-4655(98)00154-4, 1999. a
Johnson, D., Woodruff, D., McCulloh, K., and Meinzer, F.: Leaf Hydraulic Conductance, Measured in Situ, Declines and Recovers Daily: Leaf Hydraulics, Water Potential and Stomatal Conductance in Four Temperate and Three Tropical Tree Species, Tree Physiol., 29, 879–887, https://doi.org/10.1093/treephys/tpp031, 2009. a
Johnson, D. M., Wortemann, R., McCulloh, K. A., Jordan-Meille, L., Ward, E., Warren, J. M., Palmroth, S., and Domec, J.-C.: A Test of the Hydraulic Vulnerability Segmentation Hypothesis in Angiosperm and Conifer Tree Species, Tree Physiol., 36, 983–993, https://doi.org/10.1093/treephys/tpw031, 2016. a
Jones, H. G. and Sutherland, R. A.: Stomatal Control of Xylem Embolism, Plant Cell Environ., 14, 607–612, https://doi.org/10.1111/j.1365-3040.1991.tb01532.x, 1991. a
Jönsson, A. M., Schroeder, L. M., Lagergren, F., Anderbrant, O., and Smith, B.: Guess the Impact of Ips Typographus – An Ecosystem Modelling Approach for Simulating Spruce Bark Beetle Outbreaks, Agr. Forest Meteorol., 166–167, 188–200, https://doi.org/10.1016/j.agrformet.2012.07.012, 2012. a
Joshi, J., Stocker, B. D., Hofhansl, F., Zhou, S., Dieckmann, U., and Prentice, I. C.: Towards a Unified Theory of Plant Photosynthesis and Hydraulics, Nat. Plants, 8, 1304–1316, https://doi.org/10.1038/s41477-022-01244-5, 2022. a
Kannenberg, S. A., Driscoll, A. W., Malesky, D., and Anderegg, W. R.: Rapid and Surprising Dieback of Utah Juniper in the Southwestern USA Due to Acute Drought Stress, Forest Ecol. Manage., 480, 118639, https://doi.org/10.1016/j.foreco.2020.118639, 2021. a
Kennedy, D., Swenson, S., Oleson, K. W., Lawrence, D. M., Fisher, R., Lola da Costa, A. C., and Gentine, P.: Implementing Plant Hydraulics in the Community Land Model, Version 5, J. Adv. Model. Earth Syst., 11, 485–513, https://doi.org/10.1029/2018MS001500, 2019. a, b
Kirschbaum, M. U. F. and Paul, K. I.: Modelling C and N Dynamics in Forest Soils with a Modified Version of the CENTURY Model, Soil Biol. Biochem., 34, 341–354, https://doi.org/10.1016/S0038-0717(01)00189-4, 2002. a
Klein, T.: The Variability of Stomatal Sensitivity to Leaf Water Potential across Tree Species Indicates a Continuum between Isohydric and Anisohydric Behaviours, Funct. Ecol., 28, 1313–1320, https://doi.org/10.1111/1365-2435.12289, 2014. a, b
Körner, C.: Paradigm Shift in Plant Growth Control, Curr. Opin. Plant Biol., 25, 107–114, https://doi.org/10.1016/j.pbi.2015.05.003, 2015. a, b
Körner, C.: No Need for Pipes When the Well Is Dr – a Comment on Hydraulic Failure in Trees, Tree Physiol., 39, 695–700, https://doi.org/10.1093/treephys/tpz030, 2019. a, b
Köstner, B. M. M., Schulze, E. D., Kelliher, F. M., Hollinger, D. Y., Byers, J. N., Hunt, J. E., McSeveny, T. M., Meserth, R., and Weir, P. L.: Transpiration and Canopy Conductance in a Pristine Broad-Leaved Forest of Nothofagus: An Analysis of Xylem Sap Flow and Eddy Correlation Measurements, Oecologia, 91, 350–359, https://doi.org/10.1007/BF00317623, 1992. a
Lagergren, F., Jönsson, A. M., Blennow, K., and Smith, B.: Implementing Storm Damage in a Dynamic Vegetation Model for Regional Applications in Sweden, Ecol. Model., 247, 71–82, https://doi.org/10.1016/j.ecolmodel.2012.08.011, 2012. a
Lamarque, J.-F., Kyle, G. P., Meinshausen, M., Riahi, K., Smith, S. J., van Vuuren, D. P., Conley, A. J., and Vitt, F.: Global and Regional Evolution of Short-Lived Radiatively-Active Gases and Aerosols in the Representative Concentration Pathways, Climatic Change, 109, 191, https://doi.org/10.1007/s10584-011-0155-0, 2011. a
Lambers, H. and Oliveira, R. S.: Plant Water Relations, Springer International Publishing, Cham, 187–263, ISBN 978-3-030-29638-4, https://doi.org/10.1007/978-3-030-29639-1_5, 2019. a, b, c
Lan, X., Tans, P., Thoning, K., and NOAA Global Monitoring Laboratory: Trends in globally-averaged CO2 determined from NOAA Global Monitoring Laboratory measurements, GML NOAA, https://doi.org/10.15138/9N0H-ZH07, 2023. a
Lindeskog, M., Smith, B., Lagergren, F., Sycheva, E., Ficko, A., Pretzsch, H., and Rammig, A.: Accounting for Forest Management in the Estimation of Forest Carbon Balance Using the Dynamic Vegetation Model LPJ-GUESS (v4.0, R9710): Implementation and Evaluation of Simulations for Europe, Geosci. Model Dev., 14, 6071–6112, https://doi.org/10.5194/gmd-14-6071-2021, 2021. a, b, c
Lloret, F., Siscart, D., and Dalmases, C.: Canopy Recovery after Drought Dieback in Holm-Oak Mediterranean Forests of Catalonia (NE Spain), Global Change Biol., 10, 2092–2099, https://doi.org/10.1111/j.1365-2486.2004.00870.x, 2004. a
Martínez-Vilalta, J. and Garcia-Forner, N.: Water Potential Regulation, Stomatal Behaviour and Hydraulic Transport under Drought: Deconstructing the Iso/Anisohydric Concept, Plant Cell Environ., 40, 962–976, https://doi.org/10.1111/pce.12846, 2017. a
Martínez-Vilalta, J., Poyatos, R., Aguadé, D., Retana, J., and Mencuccini, M.: A New Look at Water Transport Regulation in Plants, New Phytol., 204, 105–115, https://doi.org/10.1111/nph.12912, 2014. a
Martin-StPaul, N., Delzon, S., and Cochard, H.: Sureau Database: A Database Of Hydraulic And Stomatal Traits For Modelling Drought Resistance In Plants, Zenodo [data set], https://doi.org/10.5281/ZENODO.854700, 2017. a, b, c
Mcdowell, N., Pockman, W. T., Allen, C. D., Breshears, D. D., Cobb, N., Kolb, T., Plaut, J., Sperry, J., West, A., Williams, D. G., and Yepez, E. A.: Mechanisms of Plant Survival and Mortality during Drought: Why Do Some Plants Survive While Others Succumb to Drought?, New Phytol., 178, 719–739, https://doi.org/10.1111/j.1469-8137.2008.02436.x, 2008. a, b, c
Mckay, M. D., Beckman, F. J., and Conover, W. J.: A Comparison of Three Methods for Selecting Values of Input Variables in the Analysis of Output From a Computer Code, Technometrics, 42, 55–61, https://doi.org/10.1080/00401706.2000.10485979, 2000. a
Meyer, B. F.: Data analysis pipeline for “Simulating the drought response of European tree species with the dynamic vegetation model LPJ-GUESS”, Zenodo [data set], https://doi.org/10.5281/ZENODO.14001089, 2024. a
Meyer, B. F., Darela-Filho, J. P., Gregor, K., Buras, A., Gu, Q.-L., Krause, A., Liu, D., Papastefanou, P., Asuk, S., Grams, T. E. E., Zang, C. S., and Rammig, A.: LPJ-GUESS model code used in “Simulating the drought response of European tree species with the dynamic vegetation model LPJ-GUESS”, Zenodo [code], https://doi.org/10.5281/ZENODO.14000805, 2024. a
Méndez-Alonzo, R., Ewers, F. W., Jacobsen, A. L., Pratt, R. B., Scoffoni, C., Bartlett, M. K., and Sack, L.: Covariation between Leaf Hydraulics and Biomechanics Is Driven by Leaf Density in Mediterranean Shrubs, Trees, 33, 507–519, https://doi.org/10.1007/s00468-018-1796-7, 2019. a
Nadal-Sala, D., Ruehr, N. K., and Sabaté, S.: Overcoming drought: life traits driving tree strategies to confront drought stress, J. Exp. Bot., 75, 3758–3761, https://doi.org/10.1093/jxb/erae219, 2024. a
Neycken, A., Scheggia, M., Bigler, C., and Lévesque, M.: Long-Term Growth Decline Precedes Sudden Crown Dieback of European Beech, Agr. Forest Meteorol., 324, 109103, https://doi.org/10.1016/j.agrformet.2022.109103, 2022. a
Nolan, R. H., Gauthey, A., Losso, A., Medlyn, B. E., Smith, R., Chhajed, S. S., Fuller, K., Song, M., Li, X., Beaumont, L. J., Boer, M. M., Wright, I. J., and Choat, B.: Hydraulic Failure and Tree Size Linked with Canopy Die-back in Eucalypt Forest during Extreme Drought, New Phytol., 230, 1354–1365, https://doi.org/10.1111/nph.17298, 2021. a
Nolf, M., Creek, D., Duursma, R., Holtum, J., Mayr, S., and Choat, B.: Stem and Leaf Hydraulic Properties Are Finely Coordinated in Three Tropical Rain Forest Tree Species, Plant Cell Environ., 38, 2652–2661, https://doi.org/10.1111/pce.12581, 2015. a
Nord, J., Anthoni, P., Gregor, K., Gustafson, A., Hantson, S., Lindeskog, M., Meyer, B., Miller, P., Nieradzik, L., Olin, S., Papastefanou, P., Smith, B., Tang, J., Wårlind, D., and Past LPJ-GUESS Contributors: LPJ-GUESS Release v4.1.1 model code, Zenodo [code], https://doi.org/10.5281/ZENODO.8065736, 2021. a
Oberpriller, J., Herschlein, C., Anthoni, P., Arneth, A., Krause, A., Rammig, A., Lindeskog, M., Olin, S., and Hartig, F.: Climate and Parameter Sensitivity and Induced Uncertainties in Carbon Stock Projections for European Forests (Using LPJ-GUESS 4.0), Geosci. Model Dev., 15, 6495–6519, https://doi.org/10.5194/gmd-15-6495-2022, 2022. a, b, c, d
Pammenter, N. W. and Van Der Willigen, C.: A Mathematical and Statistical Analysis of the Curves Illustrating Vulnerability of Xylem to Cavitation, Tree Physiol., 18, 589–593, https://doi.org/10.1093/treephys/18.8-9.589, 1998. a
Pan, Y., Birdsey, R. A., Fang, J., Houghton, R., Kauppi, P. E., Kurz, W. A., Phillips, O. L., Shvidenko, A., Lewis, S. L., Canadell, J. G., Ciais, P., Jackson, R. B., Pacala, S. W., Mcguire, A. D., Piao, S., Rautiainen, A., Sitch, S., and Hayes, D.: A Large and Persistent Carbon Sink in the World's Forests, Science, 333, 988–993, https://doi.org/10.1126/science.1201609, 2011. a, b
Pan, Y., Birdsey, R. A., Phillips, O. L., Houghton, R. A., Fang, J., Kauppi, P. E., Keith, H., Kurz, W. A., Ito, A., Lewis, S. L., Nabuurs, G.-J., Shvidenko, A., Hashimoto, S., Lerink, B., Schepaschenko, D., Castanho, A., and Murdiyarso, D.: The Enduring World Forest Carbon Sink, Nature, 631, 563–569, https://doi.org/10.1038/s41586-024-07602-x, 2024. a
Papastefanou, P., Zang, C. S., Pugh, T. A. M., Liu, D., Grams, T. E. E., Hickler, T., and Rammig, A.: A Dynamic Model for Strategies and Dynamics of Plant Water-Potential Regulation Under Drought Conditions, Front. Plant Sci., 11, 373, https://doi.org/10.3389/fpls.2020.00373, 2020. a, b, c, d, e, f, g
Papastefanou, P., Pugh, T. A. M., Buras, A., Fleischer, K., Grams, T. E. E., Hickler, T., Lapola, D., Liu, D., Zang, C. S., and Rammig, A.: Simulated sensitivity of the Amazon rainforest to extreme drought, Environ. Res. Lett., 19, 124072, https://doi.org/10.1088/1748-9326/ad8f48, 2024. a, b, c, d, e
Pappas, C., Fatichi, S., Leuzinger, S., Wolf, A., and Burlando, P.: Sensitivity Analysis of a Process-Based Ecosystem Model: Pinpointing Parameterization and Structural Issues, J. Geophys. Res.-Biogeo., 118, 505–528, https://doi.org/10.1002/jgrg.20035, 2013. a
Parton, W. J., Scurlock, J. M. O., Ojima, D. S., Gilmanov, T. G., Scholes, R. J., Schimel, D. S., Kirchner, T., Menaut, J.-C., Seastedt, T., Garcia Moya, E., Kamnalrut, A., and Kinyamario, J. I.: Observations and Modeling of Biomass and Soil Organic Matter Dynamics for the Grassland Biome Worldwide, Global Biogeochem. Cy., 7, 785–809, https://doi.org/10.1029/93GB02042, 1993. a
Parton, W. J., Hanson, P. J., Swanston, C., Torn, M., Trumbore, S. E., Riley, W., and Kelly, R.: ForCent Model Development and Testing Using the Enriched Background Isotope Study Experiment, J. Geophys. Res.-Biogeo., 115, G04001, https://doi.org/10.1029/2009JG001193, 2010. a
Pastorello, G., Trotta, C., Canfora, E., Chu, H., Christianson, D., Cheah, Y.-W., Poindexter, C., Chen, J., Elbashandy, A., Humphrey, M., Isaac, P., Polidori, D., Reichstein, M., Ribeca, A., Van Ingen, C., Vuichard, N., Zhang, L., Amiro, B., Ammann, C., Arain, M. A., Ardö, J., Arkebauer, T., Arndt, S. K., Arriga, N., Aubinet, M., Aurela, M., Baldocchi, D., Barr, A., Beamesderfer, E., Marchesini, L. B., Bergeron, O., Beringer, J., Bernhofer, C., Berveiller, D., Billesbach, D., Black, T. A., Blanken, P. D., Bohrer, G., Boike, J., Bolstad, P. V., Bonal, D., Bonnefond, J.-M., Bowling, D. R., Bracho, R., Brodeur, J., Brümmer, C., Buchmann, N., Burban, B., Burns, S. P., Buysse, P., Cale, P., Cavagna, M., Cellier, P., Chen, S., Chini, I., Christensen, T. R., Cleverly, J., Collalti, A., Consalvo, C., Cook, B. D., Cook, D., Coursolle, C., Cremonese, E., Curtis, P. S., D'Andrea, E., Da Rocha, H., Dai, X., Davis, K. J., Cinti, B. D., Grandcourt, A. D., Ligne, A. D., De Oliveira, R. C., Delpierre, N., Desai, A. R., Di Bella, C. M., Tommasi, P. D., Dolman, H., Domingo, F., Dong, G., Dore, S., Duce, P., Dufrêne, E., Dunn, A., Dušek, J., Eamus, D., Eichelmann, U., Elkhidir, H. A. M., Eugster, W., Ewenz, C. M., Ewers, B., Famulari, D., Fares, S., Feigenwinter, I., Feitz, A., Fensholt, R., Filippa, G., Fischer, M., Frank, J., Galvagno, M., Gharun, M., Gianelle, D., Gielen, B., Gioli, B., Gitelson, A., Goded, I., Goeckede, M., Goldstein, A. H., Gough, C. M., Goulden, M. L., Graf, A., Griebel, A., Gruening, C., Grünwald, T., Hammerle, A., Han, S., Han, X., Hansen, B. U., Hanson, C., Hatakka, J., He, Y., Hehn, M., Heinesch, B., Hinko-Najera, N., Hörtnagl, L., Hutley, L., Ibrom, A., Ikawa, H., Jackowicz-Korczynski, M., Janouš, D., Jans, W., Jassal, R., Jiang, S., Kato, T., Khomik, M., Klatt, J., Knohl, A., Knox, S., Kobayashi, H., Koerber, G., Kolle, O., Kosugi, Y., Kotani, A., Kowalski, A., Kruijt, B., Kurbatova, J., Kutsch, W. L., Kwon, H., Launiainen, S., Laurila, T., Law, B., Leuning, R., Li, Y., Liddell, M., Limousin, J.-M., Lion, M., Liska, A. J., Lohila, A., López-Ballesteros, A., López-Blanco, E., Loubet, B., Loustau, D., Lucas-Moffat, A., Lüers, J., Ma, S., Macfarlane, C., Magliulo, V., Maier, R., Mammarella, I., Manca, G., Marcolla, B., Margolis, H. A., Marras, S., Massman, W., Mastepanov, M., Matamala, R., Matthes, J. H., Mazzenga, F., Mccaughey, H., Mchugh, I., Mcmillan, A. M. S., Merbold, L., Meyer, W., Meyers, T., Miller, S. D., Minerbi, S., Moderow, U., Monson, R. K., Montagnani, L., Moore, C. E., Moors, E., Moreaux, V., Moureaux, C., Munger, J. W., Nakai, T., Neirynck, J., Nesic, Z., Nicolini, G., Noormets, A., Northwood, M., Nosetto, M., Nouvellon, Y., Novick, K., Oechel, W., Olesen, J. E., Ourcival, J.-M., Papuga, S. A., Parmentier, F.-J., Paul-Limoges, E., Pavelka, M., Peichl, M., Pendall, E., Phillips, R. P., Pilegaard, K., Pirk, N., Posse, G., Powell, T., Prasse, H., Prober, S. M., Rambal, S., Rannik, Ü., Raz-Yaseef, N., Rebmann, C., Reed, D., Dios, V. R. D., Restrepo-Coupe, N., Reverter, B. R., Roland, M., Sabbatini, S., Sachs, T., Saleska, S. R., Sánchez-Cañete, E. P., Sanchez-Mejia, Z. M., Schmid, H. P., Schmidt, M., Schneider, K., Schrader, F., Schroder, I., Scott, R. L., Sedlák, P., Serrano-Ortíz, P., Shao, C., Shi, P., Shironya, I., Siebicke, L., Šigut, L., Silberstein, R., Sirca, C., Spano, D., Steinbrecher, R., Stevens, R. M., Sturtevant, C., Suyker, A., Tagesson, T., Takanashi, S., Tang, Y., Tapper, N., Thom, J., Tomassucci, M., Tuovinen, J.-P., Urbanski, S., Valentini, R., Van Der Molen, M., Van Gorsel, E., Van Huissteden, K., Varlagin, A., Verfaillie, J., Vesala, T., Vincke, C., Vitale, D., Vygodskaya, N., Walker, J. P., Walter-Shea, E., Wang, H., Weber, R., Westermann, S., Wille, C., Wofsy, S., Wohlfahrt, G., Wolf, S., Woodgate, W., Li, Y., Zampedri, R., Zhang, J., Zhou, G., Zona, D., Agarwal, D., Biraud, S., Torn, M., and Papale, D.: The FLUXNET2015 Dataset and the ONEFlux Processing Pipeline for Eddy Covariance Data, Sci. Data, 7, 225, https://doi.org/10.1038/s41597-020-0534-3, 2020. a, b
Peters, R. L., Steppe, K., Cuny, H. E., De Pauw, D. J., Frank, D. C., Schaub, M., Rathgeber, C. B., Cabon, A., and Fonti, P.: Turgor – a Limiting Factor for Radial Growth in Mature Conifers along an Elevational Gradient, New hytol., 229, 213–229, https://doi.org/10.1111/nph.16872, 2021. a, b
Potkay, A., Hölttä, T., Trugman, A. T., and Fan, Y.: Turgor-Limited Predictions of Tree Growth, Height and Metabolic Scaling over Tree Lifespans, Tree Physiol., 42, 229–252, https://doi.org/10.1093/treephys/tpab094, 2022. a
Powell, T. L., Galbraith, D. R., Christoffersen, B. O., Harper, A., Imbuzeiro, H. M. A., Rowland, L., Almeida, S., Brando, P. M., Da Costa, A. C. L., Costa, M. H., Levine, N. M., Malhi, Y., Saleska, S. R., Sotta, E., Williams, M., Meir, P., and Moorcroft, P. R.: Confronting Model Predictions of Carbon Fluxes with Measurements of Amazon Forests Subjected to Experimental Drought, New Phytol., 200, 350–365, https://doi.org/10.1111/nph.12390, 2013. a
Pugh, T. A. M., Arneth, A., Kautz, M., Poulter, B., and Smith, B.: Important Role of Forest Disturbances in the Global Biomass Turnover and Carbon Sinks, Nat. Geosci., 12, 730–735, https://doi.org/10.1038/s41561-019-0427-2, 2019. a
Puy, A., Piano, S. L., Saltelli, A., and Levin, S. A.: sensobol: An R Package to Compute Variance-Based Sensitivity Indices, J. Stat. Soft., 102, 1–37, https://doi.org/10.18637/jss.v102.i05, 2022. a
Rouault, G., Candau, J.-N., Lieutier, F., Nageleisen, L.-M., Martin, J.-C., and Warzée, N.: Effects of Drought and Heat on Forest Insect Populations in Relation to the 2003 Drought in Western Europe, Ann. Forest Sci., 63, 613–624, https://doi.org/10.1051/forest:2006044, 2006. a
Ruffault, J., Pimont, F., Cochard, H., Dupuy, J.-L., and Martin-StPaul, N.: SurEau-Ecos v2.0: A Trait-Based Plant Hydraulics Model for Simulations of Plant Water Status and Drought-Induced Mortality at the Ecosystem Level, Geosci. Model Dev., 15, 5593–5626, https://doi.org/10.5194/gmd-15-5593-2022, 2022. a, b
Saltelli, A., Annoni, P., Azzini, I., Campolongo, F., Ratto, M., and Tarantola, S.: Variance Based Sensitivity Analysis of Model Output. Design and Estimator for the Total Sensitivity Index, Comput. Phys. Commun., 181, 259–270, https://doi.org/10.1016/j.cpc.2009.09.018, 2010. a, b
Saxton, K. E., Rawls, W. J., Romberger, J. S., and Papendick, R. I.: Estimating Generalized Soil-water Characteristics from Texture, Soil Sci. Soc. Am. J., 50, 1031–1036, https://doi.org/10.2136/sssaj1986.03615995005000040039x, 1986. a, b, c
Schönbeck, L. C., Schuler, P., Lehmann, M. M., Mas, E., Mekarni, L., Pivovaroff, A. L., Turberg, P., and Grossiord, C.: Increasing Temperature and Vapour Pressure Deficit Lead to Hydraulic Damages in the Absence of Soil Drought, Plant Cell Environ., 45, 3275–3289, https://doi.org/10.1111/pce.14425, 2022. a, b
Schuldt, B., Buras, A., Arend, M., Vitasse, Y., Beierkuhnlein, C., Damm, A., Gharun, M., Grams, T. E., Hauck, M., Hajek, P., Hartmann, H., Hiltbrunner, E., Hoch, G., Holloway-Phillips, M., Körner, C., Larysch, E., Lübbe, T., Nelson, D. B., Rammig, A., Rigling, A., Rose, L., Ruehr, N. K., Schumann, K., Weiser, F., Werner, C., Wohlgemuth, T., Zang, C. S., and Kahmen, A.: A First Assessment of the Impact of the Extreme 2018 Summer Drought on Central European Forests, Basic Appl. Ecol., 45, 86–103, https://doi.org/10.1016/j.baae.2020.04.003, 2020. a, b
Scoffoni, C., Rawls, M., McKown, A., Cochard, H., and Sack, L.: Decline of Leaf Hydraulic Conductance with Dehydration: Relationship to Leaf Size and Venation Architecture, Plant Physiol., 156, 832–843, https://doi.org/10.1104/pp.111.173856, 2011. a
Seiler, C., Melton, J. R., Arora, V. K., Sitch, S., Friedlingstein, P., Anthoni, P., Goll, D., Jain, A. K., Joetzjer, E., Lienert, S., Lombardozzi, D., Luyssaert, S., Nabel, J. E. M. S., Tian, H., Vuichard, N., Walker, A. P., Yuan, W., and Zaehle, S.: Are Terrestrial Biosphere Models Fit for Simulating the Global Land Carbon Sink?, J. Adv. Model. Earth Syst., 14, e2021MS002946, https://doi.org/10.1029/2021ms002946, 2022. a, b
Senf, C., Buras, A., Zang, C. S., Rammig, A., and Seidl, R.: Excess Forest Mortality Is Consistently Linked to Drought across Europe, Nat. Commun., 11, 6200, https://doi.org/10.1038/s41467-020-19924-1, 2020. a
Sitch, S., Smith, B., Prentice, I. C., Arneth, A., Bondeau, A., Cramer, W., Kaplan, J. O., Levis, S., Lucht, W., Sykes, M. T., Thonicke, K., and Venevsky, S.: Evaluation of Ecosystem Dynamics, Plant Geography and Terrestrial Carbon Cycling in the LPJ Dynamic Global Vegetation Model, Global Change Biol., 9, 161–185, https://doi.org/10.1046/j.1365-2486.2003.00569.x, 2003. a, b
Smith, B., Prentice, I. C., and Sykes, M. T.: Representation of Vegetation Dynamics in the Modelling of Terrestrial Ecosystems: Comparing Two Contrasting Approaches within European Climate Space: Vegetation Dynamics in Ecosystem Models, Global Ecol. Biogeogr., 10, 621–637, https://doi.org/10.1046/j.1466-822X.2001.t01-1-00256.x, 2001. a, b
Smith, B., Wårlind, D., Arneth, A., Hickler, T., Leadley, P., Siltberg, J., and Zaehle, S.: Implications of Incorporating N Cycling and N Limitations on Primary Production in an Individual-Based Dynamic Vegetation Model, Biogeosciences, 11, 2027–2054, https://doi.org/10.5194/bg-11-2027-2014, 2014. a
Sperry, J. S., Adler, F. R., Campbell, G. S., and Comstock, J. P.: Limitation of Plant Water Use by Rhizosphere and Xylem Conductance: Results from a Model, Plant Cell Environ., 21, 347–359, https://doi.org/10.1046/j.1365-3040.1998.00287.x, 1998. a
Steppe, K., De Pauw, D. J. W., Lemeur, R., and Vanrolleghem, P. A.: A Mathematical Model Linking Tree Sap Flow Dynamics to Daily Stem Diameter Fluctuations and Radial Stem Growth, Tree Physiol., 26, 257–273, https://doi.org/10.1093/treephys/26.3.257, 2006. a
Sulman, B. N., Roman, D. T., Yi, K., Wang, L., Phillips, R. P., and Novick, K. A.: High Atmospheric Demand for Water Can Limit Forest Carbon Uptake and Transpiration as Severely as Dry Soil, Geophys. Res. Lett., 43, 9686–9695, https://doi.org/10.1002/2016GL069416, 2016. a
Sykes, M. T. and Prentice, I.: Climate Change, Tree Species Distributions and Forest Dynamics: A Case Study in the Mixed Conifer/Northern Hardwoods Zone of Northern Europe, Climatic Change, 34, 161–177, https://doi.org/10.1007/bf00224628, 1996. a
Tardieu, F., Simonneau, T., and Parent, B.: Modelling the Coordination of the Controls of Stomatal Aperture, Transpiration, Leaf Growth, and Abscisic Acid: Update and Extension of the Tardieu–Davies Model, J. Exp. Bot., 66, 2227–2237, https://doi.org/10.1093/jxb/erv039, 2015. a
Torres-Ruiz, J. M., Cochard, H., Delzon, S., Boivin, T., Burlett, R., Cailleret, M., Corso, D., Delmas, C. E. L., De Caceres, M., Diaz-Espejo, A., Fernández-Conradi, P., Guillemot, J., Lamarque, L. J., Limousin, J.-M., Mantova, M., Mencuccini, M., Morin, X., Pimont, F., De Dios, V. R., Ruffault, J., Trueba, S., and Martin-Stpaul, N. K.: Plant Hydraulics at the Heart of Plant, Crops and Ecosystem Functions in the Face of Climate Change, New Phytol., 241, 984–999, https://doi.org/10.1111/nph.19463, 2024. a
Tschumi, E., Lienert, S., Bastos, A., Ciais, P., Gregor, K., Joos, F., Knauer, J., Papastefanou, P., Rammig, A., van der Wiel, K., Williams, K., Xu, Y., Zaehle, S., and Zscheischler, J.: Large Variability in Simulated Response of Vegetation Composition and Carbon Dynamics to Variations in Drought-Heat Occurrence, J. Geophys. Res.-Biogeo., 128, e2022JG007332, https://doi.org/10.1029/2022JG007332, 2023. a
Tyree, M. T. and Sperry, J. S.: Vulnerability of Xylem to Cavitation and Embolism, Annu. Rev. Plant Physiol. Plant Molec. Biol., 40, 19–36, https://doi.org/10.1146/annurev.pp.40.060189.000315, 1989. a
Tyree, M. T., Davis, S. D., and Cochard, H.: Biophysical Perspectives of Xylem Evolution: Is There a Tradeoff of Hydraulic Efficiency for Vulnerability to Dysfunction?, IAWA J., 15, 335–360, 1994. a
van der Woude, A. M., Peters, W., Joetzjer, E., Lafont, S., Koren, G., Ciais, P., Ramonet, M., Xu, Y., Bastos, A., Botía, S., Sitch, S., de Kok, R., Kneuer, T., Kubistin, D., Jacotot, A., Loubet, B., Herig-Coimbra, P.-H., Loustau, D., and Luijkx, I. T.: Temperature Extremes of 2022 Reduced Carbon Uptake by Forests in Europe, Nature Commun., 14, 6218, https://doi.org/10.1038/s41467-023-41851-0, 2023. a, b, c
Wagner, Y., Volkov, M., Nadal-Sala, D., Ruehr, N. K., Hochberg, U., and Klein, T.: Relationships between Xylem Embolism and Tree Functioning during Drought, Recovery, and Recurring Drought in Aleppo Pine, Physiol. Plant., 175, e13995, https://doi.org/10.1111/ppl.13995, 2023. a
Walthert, L., Ganthaler, A., Mayr, S., Saurer, M., Waldner, P., Walser, M., Zweifel, R., and von Arx, G.: From the Comfort Zone to Crown Dieback: Sequence of Physiological Stress Thresholds in Mature European Beech Trees across Progressive Drought, Sci. Total Environ., 753, 141792, https://doi.org/10.1016/j.scitotenv.2020.141792, 2021. a, b
Warm Winter 2020 Team and ICOS Ecosystem Thematic Centre: Warm Winter 2020 Ecosystem Eddy Covariance Flux Product for 73 Stations in FLUXNET-Archive Format – Release 2022-1 (Version 1.0), https://doi.org/10.18160/2G60-ZHAK, 2022. a, b, c
Whitehead, D.: Regulation of Stomatal Conductance and Transpiration in Forest Canopies, Tree Physiol., 18, 633–644, https://doi.org/10.1093/treephys/18.8-9.633, 1998. a
Xu, C., Christoffersen, B., Robbins, Z., Knox, R., Fisher, R. A., Chitra-Tarak, R., Slot, M., Solander, K., Kueppers, L., Koven, C., and McDowell, N.: Quantification of Hydraulic Trait Control on Plant Hydrodynamics and Risk of Hydraulic Failure within a Demographic Structured Vegetation Model in a Tropical Forest (FATES–HYDRO V1.0), Geosci. Model Dev., 16, 6267–6283, https://doi.org/10.5194/gmd-16-6267-2023, 2023. a, b
Xu, X., Medvigy, D., Powers, J. S., Becknell, J. M., and Guan, K.: Diversity in Plant Hydraulic Traits Explains Seasonal and Inter-Annual Variations of Vegetation Dynamics in Seasonally Dry Tropical Forests, New Phytol., 212, 80–95, https://doi.org/10.1111/nph.14009, 2016. a, b
Yao, Y., Joetzjer, E., Ciais, P., Viovy, N., Cresto Aleina, F., Chave, J., Sack, L., Bartlett, M., Meir, P., Fisher, R., and Luyssaert, S.: Forest Fluxes and Mortality Response to Drought: Model Description (ORCHIDEE-CAN-NHA R7236) and Evaluation at the Caxiuanã Drought Experiment, Geosci. Model Dev., 15, 7809–7833, https://doi.org/10.5194/gmd-15-7809-2022, 2022. a, b
Zaehle, S., Sitch, S., Smith, B., and Hatterman, F.: Effects of Parameter Uncertainties on the Modeling of Terrestrial Biosphere Dynamics, Global Biogeochem. Cy., 19, 2004GB002395, https://doi.org/10.1029/2004GB002395, 2005. a, b, c
Zhou, H., Tang, J., Olin, S., and Miller, P. A.: A Comprehensive Evaluation of Hydrological Processes in a Second-Generation Dynamic Vegetation Model, Hydrol. Process., 38, e15152, https://doi.org/10.1002/hyp.15152, 2024. a
Zhou, S., Duursma, R. A., Medlyn, B. E., Kelly, J. W. G., and Prentice, I. C.: How Should We Model Plant Responses to Drought? An Analysis of Stomatal and Non-Stomatal Responses to Water Stress, Agr. Forest Meteorol., 182–183, 204–214, https://doi.org/10.1016/j.agrformet.2013.05.009, 2013. a
Zweifel, R., Haeni, M., Buchmann, N., and Eugster, W.: Are Trees Able to Grow in Periods of Stem Shrinkage?, New Phytol., 211, 839–849, https://doi.org/10.1111/nph.13995, 2016. a
Short summary
Climate change has increased the likelihood of drought events across Europe, potentially threatening the European forest carbon sink. Dynamic vegetation models with mechanistic plant hydraulic architecture are needed to model these developments. We evaluate the plant hydraulic architecture version of LPJ-GUESS and show its ability to capture species-specific evapotranspiration responses to drought and to reproduce flux observations of both gross primary production and evapotranspiration.
Climate change has increased the likelihood of drought events across Europe, potentially...
