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Climate Change Impacts on the Mediterranean Forest and Shrubland, Their Ecophysiology, Demography and Community Composition

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Climate Change Impacts on the Mediterranean Forest and Shrubland, Their Ecophysiology, Demography and Community Composition ADVERTIMENT. Lʼaccés als continguts dʼaquesta tesi queda condicionat a lʼacceptació de les condicions dʼús establertes per la següent llicència Creative Commons: http://cat.creativecommons.org/?page_id=184 ADVERTENCIA. El acceso a los contenidos de esta tesis queda condicionado a la aceptación de las condiciones de uso establecidas por la siguiente licencia Creative Commons: http://es.creativecommons.org/blog/licencias/ WARNING. The access to the contents of this doctoral thesis it is limited to the acceptance of the use conditions set by the following Creative Commons license: https://creativecommons.org/licenses/?lang=en Tesis doctoral Centre de Recerca Ecològica i Aplicacions Forestals Climate change impacts on the mediterranean forest and shrubland, their ecophysiology, demography and community composition Daijun Liu Josep Peñuelas Reixach Roma Ogaya Inurrigarro Director Table of contents Abstract ........................................................................................................................... 3 General introduction ...................................................................................................... 5 Block 1. Ecophysiological changes (leaf photosynthetic activities) and water relations ......................................................................................................................... 17 Chapter 1 Foliar CO2 in a holm oak forest subjected to 15 years of climate change simulation .................................................................................................................... 18 Chapter 2. Physiological adjustments of a Mediterranean shrub to long-term experimental warming and drought treatments .......................................................... 37 Block 2. The changes in population demography (growth, mortality and recruitment) of Mediterranean holm-oak forest in response to long-term experimental drought ................................................................................................... 63 Chapter 3. Contrasting impacts of continuous moderate drought and episodic severe droughts on the aboveground-biomass increment and litterfall of three coexisting Mediterranean woody species ..................................................................................... 64 Chapter 4. Long-term experimental drought combined with natural extremes accelerate forest degradation and vegetation shifts in a Mediterranean holm oak forest .................................................................................................................................... 98 Block 3. The community structure and compositional shifts of the Mediterranean shrubland in response to long-term experimental warming and drought. ........... 139 Chapter 5. Species selection under long-term experimental warming and drought explained by distribution range ................................................................................. 140 1 Chapter 6. Shift in community structure in an early-successional Mediterranean shrubland driven by long-term experimental warming and drought and natural extreme droughts ....................................................................................................... 178 General conclusions .................................................................................................... 213 Supplementary material ............................................................................................ 216 Acknowledgement ....................................................................................................... 246 2 Abstract Anthropogenic disturbances induce rapid global warming and precipitation changes with a great impact on Mediterranean ecosystems, leading to changes in plant physiological processes (e.g. photosynthesis), demography (e.g. stem growth, mortality and seedling recruitment) and community structure and composition (e.g. species loss, dominance shifts). Moreover, great changes for future climate scenarios are expected to be unabated by most climatic models, and the potential threats of these climatic changes are still dramatic even in the context of under 2 °C global warming target. However, to date, the effective method to accurately assess species growth, community structure and composition shifts in response to future climate change are not well-established. Here, we conducted long-term (≥15 years) climate manipulation experiments in a Mediterranean holm-oak forest and in a Mediterranean shrubland to study the changes in plant ecophysiology, demography and community structure. In the Mediterranean forest, long-term experimental drought decreased photosynthetic rates of the dominant species of Q. ilex and P. latifolia, but the latter was more resistant to drought, which could lead it to gain a competitive advantage relative to Q. ilex in the future. Moreover, the long-term experimental drought significantly decreased the biomass increment and increased stem mortality, branch and leaf litterfall, and these responses were also strongly correlated with natural droughts. However, these responses were associated with natural droughts, but were also species-specific because Q. ilex resulted more vulnerable to drought than P. latifolia. Moreover, there was a dampening effect of these changes in aboveground net primary productivity (ANPP) during the long-term experimental drought. The long-term experimental drought also increased annual rates of stem mortality and decreased stem growth at the community level in plots dominated by tall trees (high canopy plots (H-)) and low trees (low canopy 3 plots (L-)) over the study period. The changes in mean stem mortality, recruitment, stem density and growth indicated that Q. ilex was more sensitive to drought than P. latifolia. The long-term experimental drought combined with natural droughts accelerated the decreases in percentage of abundance and basal area of Q. ilex, but promoted the increases for P. latifolia in low canopy plots (L-), indicating a vegetation shift with drier conditions. In the Mediterranean shrubland, long-term experimental warming moderately influenced photosynthetic rates, shoot water potential and stomatal conductance in E. multiflora, whereas drought treatment strongly influenced these variables, especially during summer. However, E. multiflora also improved the acclimation to drier and hotter conditions by the adjustments of stomatal conductance and water-use efficiency. And yet, we found that warming and drought imposed the decrease in species abundance associated with the species distributions. Long-term experimental warming caused a progressive decrease in the abundance of those species distributed in cooler regions. Correspondingly, reduced precipitation resulted in abrupt decreases in wet-distributed species, followed by a delayed increase in dry-distributed species. Moreover, we observed that the decreases in species richness in both warming and drought treatments, and the strongest decreases in community diversity and evenness were found under the drought treatment. The responses in community structure and compositional shifts were also strongly correlated with natural droughts. Thus, our results suggest that rapid warming and drought have imposed profound influences on plant physiological processes, population demography and community composition. More attention should also be paid on these changes at regional- and global-scales. 4 General introduction Anthropogenic disturbances have accelerated global climate changes with notable warming and changes in the distribution of precipitations (drier conditions in some regions) since the middle of last century, which subsequent effects in plant growth, population demography and community structure and composition (Myers et al., 2000; Walther et al., 2002; Hector & Bagchi et al., 2007; Hooper et al., 2012; Peñuelas et al., 2013; 2017; Scheffers et al., 2016). Mediterranean ecosystems are one of the hotpots suffering profound impacts induced by climate change, including loss of endemic species (Myers et al., 2000; Schröter et al., 2005; Peñuelas et al., 2007; Gottfried et al., 2012), decrease in plant photosynthetic activities (Llorens et al., 2003; Prieto et al., 2009a; Ogaya et al., 2014) and biomass growth (Ogaya & Peñuelas, 2007; Barbeta et al., 2013) and shifts in community structure and composition (Prieto et al., 2009b; Peñuelas et al., 2013; 2017; Doblas-Miranda et al., 2015). Moreover, future climate changes are expected to be unabated in the Mediterranean ecosystems, especially combined with climatic extremes as projected by most climatic models, and the impacts of these climate changes are expected to be strong for the immediate coming decades, leading to profound influences for multiple ecosystem functions and climatic feedbacks (Myers et al., 2000; Sala et al., 2000; Doblas-Miranda et al., 2015; 2016). Effective methods for accurately assessing the impacts of ongoing and future climate change on plant communities, remain poorly established. Over the last two decades, observations of examining biodiversity losses and community changes, are conducted in different time periods, but the responses are usually confused by other covarying factors (e.g. human disturbances) (Schröter et al., 2005; Gottfried et al., 2012). Performing climate manipulation experiments in natural ecosystems offers an opportunity to study plant photosynthetic activities, biomass growth, community structure and functioning 5 changes under manipulated future climatic conditions
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