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Resilience of endangered rear edge populations of cold-adapted forests in the Mediterranean basin is increasingly altered by extreme heatwave and drought pressures. It remains unknown, however, whether microclimatic variation in these isolated forests could ultimately result in large intra-population variability in the demographic responses, allowing the coexistence of contrasting declining and resilient trends across small topographic gradients. Multiple key drivers promoting spatial variability in the resilience of rear edge forests remain largely unassessed, including amplified and buffered thermal exposure induced by heatwaves along topographic gradients, and increased herbivory pressure on tree saplings in defaunated areas lacking efficient apex predators. Here we analysed whether indicators of forest resilience to global warming are strongly modulated by local-scale topographic, microclimatic and biotic conditions. We studied a protected rear edge forest of sessile oak Quercus petraea, applying a suite of 20 indicators of resilience of tree secondary growth, including multidecadal and short-term indices. We also analysed sapling recruitment success, recruit/adult ratios and sapling thermal exposure across topographic gradients. We found large within population variation in secondary growth resilience, in recruitment success and in thermal exposure of tree saplings to heatwaves, and this variability was spatially structured along small-scale topographical gradients. Multidecadal resilience indices and curves provide useful descriptors of forest vulnerability to climate warming, complementing assessments based in the analysis of short-term resilience indicators. Species-specific associations of trees with microclimatic variability are reported. Biotic factors are key in determining long-term resilience in climatically stressed rear edge forests, with strong limitation of sapling recruitment by increased roe deer and wild boar herbivory. Our results also support

The Mediterranean region is a hot spot of climate change vulnerable to increased droughts and heat waves. Scaling carbon fluxes from leaf to landscape levels is particularly challenging under drought conditions. We aimed to improve the mechanistic understanding of the seasonal acclimation of photosynthesis and morphology in sunlit and shaded leaves of four Mediterranean trees (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.) under natural conditions. Vc,max and Jmax were not constant, and mesophyll conductance was not infinite, as assumed in most terrestrial biosphere models, but varied significantly between seasons, tree species and leaf position. Favourable conditions in winter led to photosynthetic recovery and growth in the evergreens. Under moderate drought, adjustments in the photo/biochemistry and stomatal/mesophyllic diffusion behaviour effectively protected the photosynthetic machineries. Severe drought, however, induced early leaf senescence mostly in A. unedo and Q. pubescens, and significantly increased leaf mass per area in Q. ilex and P. halepensis. Shaded leaves had lower photosynthetic potentials but cushioned negative effects during stress periods. Species-specificity, seasonal variations and leaf position are key factors to explain vegetation responses to abiotic stress and hold great potential to reduce uncertainties in terrestrial biosphere models especially under drought conditions.

Water availability is the most limiting factor to global plant productivity, yet photosynthetic responses to seasonal drought cycles are poorly understood, with conflicting reports on which limiting process is the most important during drought. We address the problem using a model-data synthesis approach to look at canopy level fluxes, integrating twenty years of half hour data gathered by the FLUXNET network across six Mediterranean sites. The measured canopy level, water and carbon fluxes were used, together with an inverse canopy ecophysiological model, to estimate the bulk canopy conductance, bulk mesophyll conductance, and the canopy scale carbon pools in both the intercellular spaces and at the site of carboxylation in the chloroplasts. Thus the roles of stomatal and mesophyll conductance in the regulation of internal carbon pools and photosynthesis could be separated. A quantitative limitation analysis allowed for the relative seasonal responses of stomatal, mesophyll, and biochemical limitations to be gauged. The concentration of carbon in the chloroplast was shown to be a potentially more reliable estimator of assimilation rates than the intercellular carbon concentration. Both stomatal conductance limitations and mesophyll conductance limitations were observed to regulate the response of photosynthesis to water stress in each of the six species studied. The results suggest that mesophyll conductance could bridge the gap between conflicting reports on plant responses to soil water stress, and that the inclusion of mesophyll conductance in biosphere–atmosphere transfer models may improve their performance, in particular their ability to accurately capture the response of terrestrial vegetation productivity to drought.

Water availability is the most limiting factor to global plant productivity, yet photosynthetic responses to seasonal drought cycles are poorly understood, with conflicting reports on which limiting process is the most important during drought. We address the problem using a model-data synthesis approach to look at canopy level fluxes, integrating twenty years of half hour data gathered by the FLUXNET network across six Mediterranean sites. The measured canopy level, water and carbon fluxes were used, together with an inverse canopy ecophysiological model, to estimate the bulk canopy conductance, bulk mesophyll conductance, and the canopy scale carbon pools in both the intercellular spaces and at the site of carboxylation in the chloroplasts. Thus the roles of stomatal and mesophyll conductance in the regulation of internal carbon pools and photosynthesis could be separated. A quantitative limitation analysis allowed for the relative seasonal responses of stomatal, mesophyll, and biochemical limitations to be gauged. The concentration of carbon in the chloroplast was shown to be a potentially more reliable estimator of assimilation rates than the intercellular carbon concentration. Both stomatal conductance limitations and mesophyll conductance limitations were observed to regulate the response of photosynthesis to water stress in each of the six species studied. The results suggest that mesophyll conductance could bridge the gap between conflicting reports on plant responses to soil water stress, and that the inclusion of mesophyll conductance in biosphere-atmosphere transfer models may improve their performance, in particular their ability to accurately capture the response of terrestrial vegetation productivity to drought.

Semiarid environments throughout the world have lost a major part of their woody vegetation and biodiversity due to the effects of wood cutting, cattle grazing and subsistence agriculture. The resulting state is typically used for cattle production, but the productivity of these systems is often very low, and erosion of the unprotected soil is a common problem. Such dry-land degradation is of great international concern, not only because the resulting state is hardly productive but also because it paves the way to desertification. The natural distribution of the genus Prosopis includes arid and semiarid zones of the Americas, Africa and Asia, but the majority of the Prosopis species are, however, native to the Americas. In order to assess a likely gradient in the response of tree species to precipitation, temperature and their connection to El Niño southern oscillation (ENSO) events, two Prosopis species were chosen along a latitudinal gradient in Latin America, from northern Peru to central Chile: Prosopis pallida from a semi-arid land in northern and southern Peru and P. chilensis from a semiarid land in central Chile. Growth rings of each species were crossdated at each sampling site using classical dendrochronological techniques. Chronologies were related with instrumental climatic records in each site, as well as with SOI and N34 series. Cross-correlation, spectral and wavelet analysis techniques were used to assess the relation of growth with precipitation and temperature. Despite the long distance among sites, the two Prosopis species presented similar responses. Thus, the two species' growth is positively correlated to precipitation, while with temperature it is not. In northern Peru, precipitation and growth of P. pallida present a similar cyclic pattern, with a period of around 3 years. On the other hand, P. pallida in southern Peru, and P. chilensis also present this cyclic pattern, but also another one with lower frequency, coinciding with the pattern of

Semiarid environments throughout the world have lost a major part of their woody vegetation and biodiversity due to the effects of wood cutting, cattle grazing and subsistence agriculture. The resulting state is typically used for cattle production, but the productivity of these systems is often very low, and erosion of the unprotected soil is a common problem. Such dry-land degradation is of great international concern, not only because the resulting state is hardly productive but also because it paves the way to desertification. The natural distribution of the genus Prosopis includes arid and semiarid zones of the Americas, Africa and Asia, but the majority of the Prosopis species are, however, native to the Americas. In order to assess a likely gradient in the response of tree species to precipitation, temperature and their connection to El Niño southern oscillation (ENSO) events, two Prosopis species were chosen along a latitudinal gradient in Latin America, from northern Peru to central Chile: Prosopis pallida from a semi-arid land in northern and southern Peru and P. chilensis from a semiarid land in central Chile. Growth rings of each species were crossdated at each sampling site using classical dendrochronological techniques. Chronologies were related with instrumental climatic records in each site, as well as with SOI and N34 series. Cross-correlation, spectral and wavelet analysis techniques were used to assess the relation of growth with precipitation and temperature. Despite the long distance among sites, the two Prosopis species presented similar responses. Thus, the two species' growth is positively correlated to precipitation, while with temperature it is not. In northern Peru, precipitation and growth of P. pallida present a similar cyclic pattern, with a period of around 3 years. On the other hand, P. pallida in southern Peru, and P. chilensis also present this cyclic pattern, but also another one with lower frequency, coinciding with the pattern of

Semiarid environments throughout the world have lost a major part of their woody vegetation and biodiversity due to the effects of wood cutting, cattle grazing and subsistence agriculture. The resulting state is typically used for cattle production, but the productivity of these systems is often very low, and erosion of the unprotected soil is a common problem. Such dry-land degradation is of great international concern, not only because the resulting state is hardly productive but also because it paves the way to desertification. The natural distribution of the genus Prosopis includes arid and semiarid zones of the Americas, Africa and Asia, but the majority of the Prosopis species are, however, native to the Americas. In order to assess a likely gradient in the response of tree species to precipitation, temperature and their connection to El Niño southern oscillation (ENSO) events, two Prosopis species were chosen along a latitudinal gradient in Latin America, from northern Peru to central Chile: Prosopis pallida from a semi-arid land in northern and southern Peru and P. chilensis from a semiarid land in central Chile. Growth rings of each species were crossdated at each sampling site using classical dendrochronological techniques. Chronologies were related with instrumental climatic records in each site, as well as with SOI and N34 series. Cross-correlation, spectral and wavelet analysis techniques were used to assess the relation of growth with precipitation and temperature. Despite the long distance among sites, the two Prosopis species presented similar responses. Thus, the two species' growth is positively correlated to precipitation, while with temperature it is not. In northern Peru, precipitation and growth of P. pallida present a similar cyclic pattern, with a period of around 3 years. On the other hand, P. pallida in southern Peru, and P. chilensis also present this cyclic pattern, but also another one with lower frequency, coinciding with the pattern of