Under Stress: Searching for Genes Involved in the Response of Abies pinsapo Boiss to Climate Change.

Currently, Mediterranean forests are experiencing the deleterious effects of global warming, which mainly include increased temperatures and decreased precipitation in the region. Relict Abies pinsapo fir forests, endemic in the southern Iberian Peninsula, are especially sensitive to these recent environmental disturbances, and identifying the genes involved in the response of this endangered tree species to climate-driven stresses is of paramount importance for mitigating their effects. Genomic resources for A. pinsapo allow for the analysis of candidate genes reacting to warming and aridity in their natural habitats. Several members of the complex gene families encoding late embryogenesis abundant proteins (LEAs) and heat shock proteins (HSPs) have been found to exhibit differential expression patterns between wet and dry seasons when samples from distinct geographical locations and dissimilar exposures to the effects of climate change were analyzed. The observed changes were more perceptible in the roots of trees, particularly in declining forests distributed at lower altitudes in the more vulnerable mountains. These findings align with previous studies and lay the groundwork for further research on the molecular level. Molecular and genomic approaches offer valuable insights for mitigating climate stress and safeguarding this endangered conifer.

Ecosystem services provision by Mediterranean forests will be compromised above 2℃ warming.

Forests provide a wide range of provisioning, regulating and cultural services of great value to societies across the Mediterranean basin. In this study, we reviewed the scientific literature of the last 30 years to quantify the magnitude of projected changes in ecosystem services provision by Mediterranean forests under IPCC climate change scenarios. We classified the scenarios according to the temperature threshold of 2℃ set by the Paris Agreement (below or above). The review of 78 studies shows that climate change will lead to a general reduction in the provision of regulating services (e.g. carbon storage, regulation of freshwater quantity and quality) and a general increase in the number of fires, burnt areas and generally, an increase in climate-related forest hazards (median + 62% by 2100). Studies using scenarios above the 2℃ threshold projected significantly more negative changes in regulating services than studies using scenarios below this threshold. Main projected trend changes on material services (e.g. wood products), were less clear and depended on (i) whether or not the studies considered the interaction between the rise in temperatures and other drivers (e.g. forest management, CO2 fertilization) and (ii) differences in productivity responses across the tree species evaluated. Overall, the reviewed studies projected significant reductions in range extent and habitat suitability for the most drought-sensitive forest species (e.g. -88% Fagus sylvatica), while the amount of habitat available for more drought-tolerant species will remain stable or increase; however, the magnitude of projected change for these more xeric species was limited when high-end extreme climatic scenarios were considered (above Paris Agreement). Our review highlights the benefits that climate change mitigation (to keep global mean temperature increase <2℃) can bring in terms of service provision and conservation of Mediterranean forests.

Current and future impacts of drought and ozone stress on Northern Hemisphere forests.

Rising ozone (O3 ) concentrations, coupled with an increase in drought frequency due to climate change, pose a threat to plant growth and productivity which could negatively affect carbon sequestration capacity of Northern Hemisphere (NH) forests. Using long-term observations of O3 mixing ratios and soil water content (SWC), we implemented empirical drought and O3 stress parameterizations in a coupled stomatal conductance-photosynthesis model to assess their impacts on plant gas exchange at three FLUXNET sites: Castelporziano, Blodgett and Hyytiälä. Model performance was evaluated by comparing model estimates of gross primary productivity (GPP) and latent heat fluxes (LE) against present-day observations. CMIP5 GCM model output data were then used to investigate the potential impact of the two stressors on forests by the middle (2041-2050) and end (2091-2100) of the 21st century. We found drought stress was the more significant as it reduced model overestimation of GPP and LE by ~11%-25% compared to 1%-11% from O3 stress. However, the best model fit to observations at all the study sites was obtained with O3 and drought stress combined, such that the two stressors counteract the impact of each other. With the inclusion of drought and O3 stress, GPP at CPZ, BLO and HYY is projected to increase by 7%, 5% and 8%, respectively, by mid-century and by 14%, 11% and 14% by 2091-2100 as atmospheric CO2 increases. Estimates were up to 21% and 4% higher when drought and O3 stress were neglected respectively. Drought stress will have a substantial impact on plant gas exchange and productivity, off-setting and possibly negating CO2 fertilization gains in future, suggesting projected increases in the frequency and severity of droughts in the NH will play a significant role in forest productivity and carbon budgets in future.

Forests synchronize their growth in contrasting Eurasian regions in response to climate warming.

Forests play a key role in the carbon balance of terrestrial ecosystems. One of the main uncertainties in global change predictions lies in how the spatiotemporal dynamics of forest productivity will be affected by climate warming. Here we show an increasing influence of climate on the spatial variability of tree growth during the last 120 y, ultimately leading to unprecedented temporal coherence in ring-width records over wide geographical scales (spatial synchrony). Synchrony in growth patterns across cold-constrained (central Siberia) and drought-constrained (Spain) Eurasian conifer forests have peaked in the early 21st century at subcontinental scales (∼ 1,000 km). Such enhanced synchrony is similar to that observed in trees co-occurring within a stand. In boreal forests, the combined effects of recent warming and increasing intensity of climate extremes are enhancing synchrony through an earlier start of wood formation and a stronger impact of year-to-year fluctuations of growing-season temperatures on growth. In Mediterranean forests, the impact of warming on synchrony is related mainly to an advanced onset of growth and the strengthening of drought-induced growth limitations. Spatial patterns of enhanced synchrony represent early warning signals of climate change impacts on forest ecosystems at subcontinental scales.

Black Truffle Harvesting in Spanish Forests: Trends, Current Policies and Practices, and Implications on its Sustainability.

The European black truffle is a mycorrhizal fungus native to Spanish Mediterranean forests. In most Spanish regions it was originally commercially harvested in the second half of the 20th century. Experts agree that wild truffle yields suffered a sharp decline during the 1970s and 1980s. However, official statistics for Spanish harvest are scarce and seemingly conflicting, and little attention has been paid to the regime for the exploitation of truffle-producing forests and its implications on the sustainability of this resource. Trends in harvest from 1969 to 2013 and current harvesting practices were analyzed as a case study, taking into account that Spain is a major truffle producer worldwide, but at the same time truffles have only recently been exploited. The available statistical sources, which include an increasing proportion of cultivated truffles since the mid-1990s, were explored, with estimates from Truffle Harvesters Federation showing higher consistency. Statistical sources were then compared with proxies for wild harvest (rents from truffle leases in public forests) to corroborate time trends in wild harvesting. Results suggest that black truffle production is recovering in recent years thanks to plantations, whereas wild harvest is still declining. The implications of Spanish legal and institutional framework on sustainability of wild truffle use are reviewed. In the current scenario, the decline of wild harvest is likely to continue and eventually make commercial harvesting economically unattractive, thus aggravating sustainability issues. Strengthening of property rights, rationalization of harvesting pressure, forest planning and involvement of public stakeholders are proposed as corrective measures.

Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests.

Tree populations at the low-altitudinal or -latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree-ring widths and δ13 C and δ18 O chronologies to compare the growth rates and long-term ecophysiological responses to climate in the temperate-Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO2 . Tree-ring δ18 O for both species were mostly correlated with δ18 O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree-ring δ18 O but had a positive effect on Q. ilex tree-ring δ18 O. Furthermore, tree-ring δ18 O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO2 fertilization, in contrast to the water-limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.

Drought legacies in mixed Mediterranean forests: Analysing the effects of structural overshoot, functional traits and site factors.

Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests.

Leaf morpho-anatomical adjustments in a Quercus pubescens forest after 10 years of partial rain exclusion in the field.

In the Mediterranean region, a reduction of annual precipitation and a longer and drier summer season are expected with climate change by the end of the century, eventually endangering forest survival. To cope with such rapid changes, trees may modulate their morpho-anatomical and physiological traits. In the present study, we focused on the variation in leaf gas exchange and different leaf morpho-anatomical functional traits of Quercus pubescens Willd. in summer using a long-term drought experiment in natura consisting of a dynamic rainfall exclusion system where trees have been submitted to amplified drought (AD) (~-30% of annual precipitation) since April 2012 and compared them with trees under natural drought (ND) in a Mediterranean forest. During the study, we analyzed net CO2 assimilation (An), stomatal conductance (gs), transpiration (E), water-use efficiency (WUE), stomatal size and density, density of glandular trichomes and non-glandular trichomes, thickness of the different leaf tissues, specific leaf area and leaf surface. Under AD, tree functioning was slightly impacted, since only An exhibited a 49% drop, while gs, E and WUE remained stable. The decrease in An under AD was regulated by concomitant lower stomatal density and reduced leaf thickness. Trees under AD also featured leaves with a higher non-glandular trichome density and a lower glandular trichome density compared with ND, which simultaneously limits transpiration and production costs. This study points out that Q. pubescens exhibits adjustments of leaf morpho-anatomical traits which can help trees to acclimate to AD scenarios as those expected in the future in the Mediterranean region.

Forest bathing and hiking benefits for mental health during the COVID-19 pandemic in Mediterranean regions.

Forest bathing (FB) has evidenced positive effects on individuals' mental health and well-being, but its benefits have mainly been studied in Asian biomes. The present study aimed to evaluate whether its benefits are also generalisable to other forests and biomes of the world, such as the Mediterranean. Eighty-six healthy adults of the general population were assessed before and after a FB near Barcelona (Spain) during the COVID-19 pandemic. A control-hiking group of participants was also analysed to contrast the FB effects on anxiety, affect, mood states and mindfulness. Results show that the guided practice of FB in Mediterranean-Catalan forests increases mindfulness states and positive affect and reduces anxiety and negative affect, with effect sizes being large to very large. Hiking also induced significant changes in all variables tested, but FB showed higher effect sizes. An exploratory analysis also revealed a different profile of the FB participants compared to the hiking practitioners, being highly educated women living in urban areas and with lower basal levels of psychological well-being. Accordingly, it is concluded that both Mediterranean FB and hiking (to a lesser degree) might be cost-effective strategies to promote and restore psychological well-being after the COVID-19 pandemic and to promote sustainable tourism in Mediterranean biomes of the European forested and protected areas.

Simulating diverse forest management options in a changing climate on a Pinus nigra subsp. laricio plantation in Southern Italy.

Mediterranean pine plantations provide several ecosystem services but are vulnerable to climate change. Forest management might play a strategic role in the adaptation of Mediterranean forests, but the joint effect of climate change and diverse management options have seldom been investigated together. Here, we simulated the development of a Laricio pine (Pinus nigra subsp. laricio) stand in the Bonis watershed (southern Italy) from its establishment in 1958 up to 2095 using a state-of-the-science process-based forest model. The model was run under three climate scenarios corresponding to increasing levels of atmospheric CO2 concentration and warming, and six management options with different goals, including wood production and renaturalization. We analysed the effect of climate change on annual carbon fluxes (i.e., gross and net primary production) and stocks (i.e., basal area, standing and harvested carbon woody stocks) of the autotrophic compartment, as well as the impact of different management options compared to a no management baseline. Results show that higher temperatures (+3 to +5 °C) and lower precipitation (-20 % to -22 %) will trigger a decrease in net primary productivity in the second half of the century. Compared to no management, the other options had a moderate effect on carbon fluxes over the whole simulation (between -14 % and +11 %). While standing woody biomass was reduced by thinning interventions and the shelterwood system (between -5 % and -41 %), overall carbon stocks including the harvested wood were maximized (between +41 % and +56 %). Results highlight that management exerts greater effects on the carbon budget of Laricio pine plantations than climate change alone, and that climate change and management are largely independent (i.e., no strong interaction effects). Therefore, appropriate silvicultural strategies might enhance potential carbon stocks and improve forest conditions, with cascading positive effects on the provision of ecosystem services in Mediterranean pine plantations.

Declines in canopy greenness and tree growth are caused by combined climate extremes during drought-induced dieback.

Several dieback episodes triggered by droughts are revealing the high vulnerability of Mediterranean forests, manifested as declines in growth, increased defoliation, and rising mortality rates. Understanding forest responses to such climate extreme events is of high priority for predicting their future vegetation dynamics. We examined how remotely sensed measures of vegetation activity (NDVI, Normalized Difference Vegetation Index) and radial growth (BAI, basal area increment) responded to climate extreme events. We considered tree (Pinus sylvestris, Quercus pubescens, Quercus frainetto) and shrub (Juniperus phoenicea) populations from Italy and Spain showing recent dieback phenomena. Two components of drought, namely elevated atmospheric demand (VPD, vapor pressure deficit) and low soil moisture were analyzed in nearby stands showing or not showing dieback symptoms. Dieback stands exhibited lower NDVI values than non-dieback stands. NDVI and BAI were positively related in all sites except for the dieback stand of Q. frainetto that was negatively related. Such NDVI-BAI linkages were related to specific time windows, which could be useful for identifying when climatic conditions have the greatest influence on vegetation. Growth decline occurred in response to increasing VPD, but responses differed among species. J. phoenicea was the most negatively impacted by higher VPD, whereas oaks responded to soil moisture. A high VPD was related to stronger growth reduction in dieback P. sylvestris trees regardless of soil moisture changes. We highlighted that coupling between proxies of forest productivity (NDVI, BAI) allows better understanding and forecasting of drought-induced dieback phenomena in forests and shrublands. Scaling up from tree to stand levels might be feasible when using the maximum growing season NDVI, which can be applied for retrospective modeling of the impact of drought stress on forest productivity and tree growth.

Beyond width and density: stable carbon and oxygen isotopes in cork-rings provide insights of physiological responses to water stress in Quercus suber L.

As climate change increasingly affects forest ecosystems, detailed understanding of major effects is important to anticipate their consequences under future climate scenarios. The Mediterranean region is a prominent climate change hotspot, and evergreen cork oak (Quercus suber L.) woodlands are particularly climatically sensitive due to cork (bark) harvesting. Cork oak's drought avoidance strategy is well-known and includes structural and physiological adaptations that maximise soil water uptake and transport and limit water use, potentially leading to reduced stem and cork growth. Trees' responses to cope with water-limited conditions have been extensively described based on cork-rings width and, more recently, on cork-rings density, in dendroecological studies. However, so far, tree functional attributes and physiological strategies, namely photosynthetic metabolism adjustments affecting cork formation, have never been addressed and/or integrated on these previous cork-rings-based studies. In this study, we address the relation between carbon and oxygen stable isotopes of cork rings and precipitation and temperature, in two distinct locations of southwestern Portugal-the (wetter) Tagus basin peneplain and the (drier) Grândola mountains. We aimed at assessing whether the two climatic factors affect cork-ring isotopic composition under contrasting conditions of water availability, and, therefore, if carbon and oxygen signatures in cork can reflect tree functional (physiological and structural) responses to stressful conditions, which might be aggravated by climate change. Our results indicate differences between the study areas. At the drier site, the stronger statistically significant negative cork δ 13C correlations were found with mean temperature, whereas strong positive cork δ 18O correlations were fewer and found only with precipitation. Moreover, at the wetter site, cork rings are enriched in 18O and depleted in 13C, indicating, respectively, shallow groundwater as the water source for physiological processes related with biosynthesis of non-photosynthetic secondary tissues, such as suberin, and a weak stomatal regulation under high water availability, consistent with non-existent water availability constrains. In contrast, at the drier site, trees use water from deeper ground layers, depleted in 18O, and strongly regulate stomatal conductance under water stress, thus reducing photosynthetic carbon uptake and probably relying on stored carbon reserves for cork ring formation. These results suggest that although stable isotopes signatures in cork rings are not proxies for net growth, they may be (fairly) robust indicators of trees' physiological and structural adjustments to climate and environmental changes in Mediterranean environments.

The Impact of Climate Change on Forest Development: A Sustainable Approach to Management Models Applied to Mediterranean-Type Climate Regions.

Forest ecosystems are divided into three major groups: boreal, temperate, and tropical. These can be subdivided according to the particularities of each type due to its relative location (littoral, mountain, etc.), climatic conditions, or even geological substrate. Climate change affects each type of forest ecosystem differently. However, it seems to affect temperate forests in Mediterranean-type climate regions more intensely. These regions are located over several continents, with major impacts of increased temperature during summer and decreased precipitation during winter. This situation affects Mediterranean forest ecosystems by increasing the risk of fires, which arise more frequently and are more severe. In addition, the emergence of pests and the spread of invasive species are well-known problems affecting these ecosystems. All of these conditions contribute to losses of productivity and biodiversity. To avoid the destruction of forest resources, and since Mediterranean-type climate regions are considered climate change hot spots with increased vulnerability to disturbances, the implementation of adaptive forest management models could contribute to increasing the resilience of such forests, which could also contribute to mitigating climate change.

Seasonal variations of electrical signals of Pinus halepensis Mill. in Mediterranean forests in dependence on climatic conditions.

The temporal evolution of the electrical signal generated by Pinus halepensis was measured in a sample of 15 trees. Weekly experiments were carried out during a long-term campaign lasting over a year, while trials with a high frequency of measurements were also performed during several days. In the latter case, day-night oscillations of the electrical magnitudes were observed. Additionally, punctual meteorological events such as rainfall and electrical storms affect the electrical signal as well.The measured electrical intensity grows exponentially with the voltage. In fact, no electrical intensity that exceeds the threshold of 0.01 µA is gathered when voltage values are lower than 0.6 V. In general, higher electrical signals were gathered during the rainy seasons with moderate temperatures; while very low signals, including few measures of zero intensity, were obtained during the most stressful periods over the year, mainly by mid-summer.There is a strong correlation between the rainfall and the electrical signal. The rain-intensity correlation, together with sustained intensity values during the reproductive period in spring, suggests that this electrical magnitude could be an indicator of the physiological state of the tree and thus used for in situ and minimally invasive forest monitoring.

Additive or non-additive effect of mixing oak in pine stands on soil properties depends on the tree species in Mediterranean forests.

This study investigated how oak abundance in pine stands (using relative Oak Basal Area %, OBA%) may modulate soil microbial functioning. Forests were composed of sclerophyllous species i.e. Quercus ilex mixed with Pinus halepensis Miller or of Q. pubescens mixed with P. sylvestris. We used a series of plots with OBA% ranging from 0 to 100% in the two types of stand (n=60) and both OLF and A-horizon compartments were analysed. Relations between OBA% and either soil chemical (C and N contents, quality of organic matter via solid-state NMR, pH, CaCO3) or microbial (enzyme activities, basal respiration, biomass and catabolic diversity via BIOLOG) characteristics were described. OBA% increase led to a decrease in the recalcitrant fraction of organic matter (OM) in OLF and promoted microbial growth. Catabolic profiles of microbial communities from A-horizon were significantly modulated in Q. ilex and P. halepensis stand by OBA% and alkyl C to carboxyl C ratio (characteristic of cutin from Q. ilex tissues) and in Q. pubescens and P. sylvestris stands, by OBA% and pH. In A-horizon under Q. ilex and P. halepensis stands, linear regressions were found between catabolic diversity, microbial biomass and OBA% suggesting an additive effect. Conversely, in A-horizon Q. pubescens and P. sylvestris stands, the relationship between OBA% and either cellulase activities, polysaccharides or ammonium contents, suggested a non-additive effect of Q. pubescens and P. sylvestris, enhancing mineralization of the OM labile fraction for plots characterized by an OBA% ranging from 40% to 60%. Mixing oak with pine thus favored microbial dynamics in both type of stands though OBA% print varied with tree species and consequently sustainable soil functioning depend strongly on the composition of mixed stands. Our study indeed revealed that, when evaluating the benefits of forest mixed stand on soil microbial functioning and OM turnover, the identity of tree species has to be considered.

Beyond ectomycorrhizal bipartite networks: projected networks demonstrate contrasted patterns between early- and late-successional plants in Corsica.

The ectomycorrhizal (ECM) symbiosis connects mutualistic plants and fungal species into bipartite networks. While links between one focal ECM plant and its fungal symbionts have been widely documented, systemic views of ECM networks are lacking, in particular, concerning the ability of fungal species to mediate indirect ecological interactions between ECM plant species (projected-ECM networks). We assembled a large dataset of plant-fungi associations at the species level and at the scale of Corsica using molecular data and unambiguously host-assigned records to: (i) examine the correlation between the number of fungal symbionts of a plant species and the average specialization of these fungal species, (ii) explore the structure of the plant-plant projected network and (iii) compare plant association patterns in regard to their position along the ecological succession. Our analysis reveals no trade-off between specialization of plants and specialization of their partners and a saturation of the plant projected network. Moreover, there is a significantly lower-than-expected sharing of partners between early- and late-successional plant species, with fewer fungal partners for early-successional ones and similar average specialization of symbionts of early- and late-successional plants. Our work paves the way for ecological readings of Mediterranean landscapes that include the astonishing diversity of below-ground interactions.

A Minimal Model for Forest Fire Regimes.

We show in this article how the characteristics of fire recurrence in forests can be theoretically derived from simple information concerning forest morphology. The task is accomplished by means of a minimal model encapsulating a few assumptions on the interactions between overstory and understory species and on the mechanisms of fire development and transmission. The main difference with other models for fire prediction and simulation is that, here, fire is an endogenous variable with purely deterministic dynamics. Nevertheless, the analysis shows that fire recurrence can be chaotic for parameter values corresponding to Mediterranean forests. By contrast, the model shows that boreal forests and savannas have the tendency to experience periodic fires. These general results are in agreement with the studies carried out on many different forests in this century.