Divergent spatio-temporal tree growth trends in Pinus pinaster Ait. in South-Western European forests.

Climate change influences forest ecosystems in several ways, such as modifying forest growth or ecosystem functionality. To fully understand the impact of changing climatic conditions on forest growth it is necessary to undertake long-term spatiotemporal analyses. The main purpose of this work is to describe the major trends in tree growth of Pinus pinaster in Spain over the last 70 years, differentiating homogeneous ecological units using an unsupervised classification algorithm and additive modelling techniques. We also aim to relate these growth trends with temporal series for precipitation and temperature, as well as forest variables. We leverage information from a large data set of tree cores (around 2200) extracted during the field campaign of the Fourth Spanish National Forest Inventory. An unsupervised algorithm classified the plots into five classes, which were consistent in ecological terms. We also found a general decline in growth in three of the five ecoregions since the 1970s, concomitant with an increase in temperature and a reduction in precipitation. However, this tree growth decline has not been observed in the Atlantic influenced ecoregion, where the cooler, more humid climatic conditions are more stable. Certain stand features, such as low basal area through forest management practices, may have alleviated the impact of harsh climatic conditions on some areas of inner Spain, while denser stands display a more pronounced decline in tree growth. We concluded that Southern populations show some degrees of growth decline and low growth trends while Northern populations did not exhibit growth decline and have the largest growth rates. Under a forecasted increment of temperatures, the growth decline can be expanded.

Harmonised statistics and maps of forest biomass and increment in Europe.

Forest biomass is an essential resource in relation to the green transition and its assessment is key for the sustainable management of forest resources. Here, we present a forest biomass dataset for Europe based on the best available inventory and satellite data, with a higher level of harmonisation and spatial resolution than other existing data. This database provides statistics and maps of the forest area, biomass stock and their share available for wood supply in the year 2020, and statistics on gross and net volume increment in 2010-2020, for 38 European countries. The statistics of most countries are available at a sub-national scale and are derived from National Forest Inventory data, harmonised using common reference definitions and estimation methodology, and updated to a common year using a modelling approach. For those counties without harmonised statistics, data were derived from the State of Europe's Forest 2020 Report at the national scale. The maps are coherent with the statistics and depict the spatial distribution of the forest variables at 100 m resolution.

Deadwood stocks in south-western European forests: Ecological patterns and large scale assessments.

Forest deadwood is a relevant factor in the provision of ecosystem services (forest biodiversity, carbon sequestration, recreational and aesthetic values), but it also influences the risk and impact of forest perturbations. Hence, reliable estimations are urgently need in the lack of detailed information in Mediterranean forests at large scales. In this study we provide, for the first time, national-level estimations for Spain based on the information from the Spanish National Forest Inventory (38,945 plots). In addition, we compare and validate two approaches for estimating deadwood stocks where data is lacking; the first of these being a modelling approach based on stand, climatic and physiographical variables, and the other considers the ratio between deadwood and living biomass. We also examine the different patterns stock across forest types in four biogeographical regions according to a broad-spectrum of species groups and forests with different degrees of anthropogenic influence. The degrees are based on levels of protection and naturalness categories. The modelling approach provides more robust deadwood estimates and better predictive capacity than the ratio approach. Alpine (6.09 Mg.ha-1) and Atlantic (3.53 Mg.ha-1) bioregion forests store significantly higher mean deadwood biomass stocks than Macaronesian and Mediterranean forests. However, the share of deadwood in relation to the total biomass stock is greater in Mediterranean biogeographical region. As regards species groups, the mean deadwood stock of mixed forests doubled the stocks found in conifer and broadleaved dominated forests. We also found significant differences in deadwood biomass stocks between forests with different levels of anthropogenic protection. However, forest types with intensive forest management had contrasting figures for deadwood stock. The mean values obtained at national level according to forest type, bioregion and degree of anthropogenic influence, provide baseline information for carbon accounting as well as for other forest policy planning and management strategies.

Climate Change Synchronizes Growth and iWUE Across Species in a Temperate-Submediterranean Mixed Oak Forest.

Tree species have good tolerance to a range of environmental conditions, though their ability to respond and persist to environmental changes is dramatically reduced at the rear-edge distribution limits. At those edges, gene flow conferring adaptation is impaired due to lack of populations at lower latitudes. Thus, trees mainly rely on phenotypic changes to buffer against long-term environmental changes. Interspecific hybridization may offer an alternative mechanism in the generation of novel genetic recombinants that could be particularly valuable to ensure persistence in geographically isolated forests. In this paper, we take advantage of the longevity of a temperate-submediterranean mixed-oak forest to explore the long-term impact of environmental changes on two different oak species and their hybrid. Individual trees were genetically characterized and classified into three groups: pure Quercus petraea (Matt.), Liebl, pure Q. pyrenaica Willd, and hybrids. We calculated basal area increment and intrinsic water-use efficiency (iWUE) from tree-ring width and δ13C per genetic group, respectively. Tree-growth drivers were assessed using correlation analyses and generalized linear mixed models for two contrasting climatic periods: (1880-1915, colder with [CO2] < 303 ppm; and 1980-2015, warmer with [CO2] > 338 ppm). The three genetic groups have increased radial growth and iWUE during the last decades, being the least drought-tolerant QuPe the most sensitive species to water stress. However, no significant differences were found among genetic groups neither in mean growth rate nor in mean iWUE. Furthermore, little differences were found in the response to climate among groups. Genetic groups only differed in the relationship between δ13C and temperature and precipitation during the earlier period, but such a difference disappeared during the recent decades. Climate change may have promoted species-level convergence as a response to environment-induced growth limitations, which translated in synchronized growth and response to climate as well as a tighter stomatal control and increased iWUE across coexisting oak species.

Geographical adaptation prevails over species-specific determinism in trees' vulnerability to climate change at Mediterranean rear-edge forests.

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short-term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short- and long-term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species-specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species-specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine-scale limitations of the climate data generated by global climate models.

Divergent phenological and leaf gas exchange strategies of two competing tree species drive contrasting responses to drought at their altitudinal boundary.

In Mediterranean mountains, Pinus sylvestris L. is expected to be displaced under a warming climate by more drought-tolerant species such as the sub-Mediterranean Quercus pyrenaica Willd. Understanding how environmental factors drive tree physiology and phenology is, therefore, essential to assess the effect of changing climatic conditions on the performance of these species and, ultimately, their distribution. We compared the cambial and leaf phenology and leaf gas exchange of Q. pyrenaica and P. sylvestris at their altitudinal boundary in Central Spain and assessed the environmental variables involved. Results indicate that P. sylvestris cambial phenology was more sensitive to weather conditions (temperature at the onset and water deficit at the end of the growing season) than Q. pyrenaica. On the other hand, Q. pyrenaica cambial and leaf phenology were synchronized and driven by photoperiod and temperatures. Pinus sylvestris showed lower photosynthetic nitrogen-use efficiency and higher intrinsic water-use efficiency than Q. pyrenaica as a result of a tighter stomatal control in response to summer dry conditions, despite its less negative midday leaf water potentials. These phenological and leaf gas exchange responses evidence a stronger sensitivity to drought of P. sylvestris than that of Q. pyrenaica, which may therefore hold a competitive advantage over P. sylvestris under the predicted increase in recurrence and intensity of drought events. On the other hand, both species could benefit from warmer springs through an advanced phenology, although this effect could be limited in Q. pyrenaica if it maintains a photoperiod control over the onset of xylogenesis.

Xylem and Leaf Functional Adjustments to Drought in Pinus sylvestris and Quercus pyrenaica at Their Elevational Boundary.

Climatic scenarios for the Mediterranean region forecast increasing frequency and intensity of drought events. Consequently, a reduction in Pinus sylvestris L. distribution range is projected within the region, with this species being outcompeted at lower elevations by more drought-tolerant taxa such as Quercus pyrenaica Willd. The functional response of these species to the projected shifts in water availability will partially determine their performance and, thus, their competitive success under these changing climatic conditions. We studied how the cambial and leaf phenology and xylem anatomy of these two species responded to a 3-year rainfall exclusion experiment set at their elevational boundary in Central Spain. Additionally, P. sylvestris leaf gas exchange, water potential and carbon isotope content response to the treatment were measured. Likewise, we assessed inter-annual variability in the studied functional traits under control and rainfall exclusion conditions. Prolonged exposure to drier conditions did not affect the onset of xylogenesis in either of the studied species, whereas xylem formation ceased 1-3 weeks earlier in P. sylvestris. The rainfall exclusion had, however, no effect on leaf phenology on either species, which suggests that cambial phenology is more sensitive to drought than leaf phenology. P. sylvestris formed fewer, but larger tracheids under dry conditions and reduced the proportion of latewood in the tree ring. On the other hand, Q. pyrenaica did not suffer earlywood hydraulic diameter changes under rainfall exclusion, but experienced a cumulative reduction in latewood width, which could ultimately challenge its hydraulic performance. The phenological and anatomical response of the studied species to drought is consistent with a shift in resource allocation under drought stress from xylem to other sinks. Additionally, the tighter stomatal control and higher intrinsic water use efficiency observed in drought-stressed P. sylvestris may eventually limit carbon uptake in this species. Our results suggest that both species are potentially vulnerable to the forecasted increase in drought stress, although P. sylvestris might experience a higher risk of drought-induced decline at its low elevational limit.

Impact of RAV1-engineering on poplar biomass production: a short-rotation coppice field trial.

BACKGROUND: Early branching or syllepsis has been positively correlated with high biomass yields in short-rotation coppice (SRC) poplar plantations, which could represent an important lignocellulosic feedstock for the production of second-generation bioenergy. In prior work, we generated hybrid poplars overexpressing the chestnut gene RELATED TO ABI3/VP1 1 (CsRAV1), which featured c. 80% more sylleptic branches than non-modified trees in growth chambers. Given the high plasticity of syllepsis, we established a field trial to monitor the performance of these trees under outdoor conditions and a SRC management. RESULTS: We examined two CsRAV1-overexpression poplar events for their ability to maintain syllepsis and their potential to enhance biomass production. Two poplar events with reduced expression of the CsRAV1 homologous poplar genes PtaRAV1 and PtaRAV2 were also included in the trial. Under our culture conditions, CsRAV1-overexpression poplars continued developing syllepsis over two cultivation cycles. Biomass production increased on completion of the first cycle for one of the overexpression events, showing unaltered structural, chemical, or combustion wood properties. On completion of the second cycle, aerial growth and biomass yields of both overexpression events were reduced as compared to the control. CONCLUSIONS: These findings support the potential application of CsRAV1-overexpression to increase syllepsis in commercial elite trees without changing their wood quality. However, the syllepsis triggered by the introduction of this genetic modification appeared not to be sufficient to sustain and enhance biomass production.

Towards complete and harmonized assessment of soil carbon stocks and balance in forests: The ability of the Yasso07 model across a wide gradient of climatic and forest conditions in Europe.

Accurate carbon-balance accounting in forest soils is necessary for the development of climate change policy. However, changes in soil organic carbon (SOC) occur slowly and these changes may not be captured through repeated soil inventories. Simulation models may be used as alternatives to SOC measurement. The Yasso07 model presents a suitable alternative because most of the data required for the application are readily available in countries with common forest surveys. In this study, we test the suitability of Yasso07 for simulating SOC stocks and stock changes in a variety of European forests affected by different climatic, land use and forest management conditions and we address country-specific cases with differing resources and data availability. The simulated SOC stocks differed only slightly from measured data, providing realistic, reasonable mean SOC estimations per region or forest type. The change in the soil carbon pool over time, which is the target parameter for SOC reporting, was generally found to be plausible although not in the case of Mediterranean forest soils. As expected under stable forest management conditions, both land cover and climate play major roles in determining the SOC stock in forest soils. Greater mean SOC stocks were observed in northern latitudes (or at higher altitude) than in southern latitudes (or plains) and conifer forests were found to store a notably higher amount of SOC than broadleaf forests. Furthermore, as regards change in SOC, an inter-annual sink effect was identified for most of the European forest types studied. Our findings corroborate the suitability of Yasso07 to assess the impact of forest management and land use change on the SOC balance of forests soils, as well as to accurately simulate SOC in dead organic matter (DOM) and mineral soil pools separately. The obstacles encountered when applying the Yasso07 model reflect a lack of available input data. Future research should focus on improving our knowledge of C inputs from compartments such as shrubs, herbs, coarse woody debris and fine roots. This should include turnover rates and quality of the litter in all forest compartments from a wider variety of tree species and sites. Despite the limitations identified, the SOC balance estimations provided by the Yasso07 model are sufficiently complete, accurate and transparent to make it suitable for reporting purposes such as those required under the UNFCCC (United Nations Framework Convention on Climate Change) and KP (Kyoto Protocol) for a wide range of forest conditions in Europe.

Stand competition determines how different tree species will cope with a warming climate.

Plant-plant interactions influence how forests cope with climate and contribute to modulate species response to future climate scenarios. We analysed the functional relationships between growth, climate and competition for Pinus sylvestris, Quercus pyrenaica and Quercus faginea to investigate how stand competition modifies forest sensitivity to climate and simulated how annual growth rates of these species with different drought tolerance would change throughout the 21st century. Dendroecological data from stands subjected to thinning were modelled using a novel multiplicative nonlinear approach to overcome biases related to the general assumption of a linear relationship between covariates and to better mimic the biological relationships involved. Growth always decreased exponentially with increasing competition, which explained more growth variability than climate in Q. faginea and P. sylvestris. The effect of precipitation was asymptotic in all cases, while the relationship between growth and temperature reached an optimum after which growth declined with warmer temperatures. Our growth projections indicate that the less drought-tolerant P. sylvestris would be more negatively affected by climate change than the studied sub-Mediterranean oaks. Q. faginea and P. sylvestris mean growth would decrease under all the climate change scenarios assessed. However, P. sylvestris growth would decline regardless of the competition level, whereas this decrease would be offset by reduced competition in Q. faginea. Conversely, Q. pyrenaica growth would remain similar to current rates, except for the warmest scenario. Our models shed light on the nature of the species-specific interaction between climate and competition and yield important implications for management. Assuming that individual growth is directly related to tree performance, trees under low competition would better withstand the warmer conditions predicted under climate change scenarios but in a variable manner depending on the species. Thinning following an exponential rule may be desirable to ensure long-term conservation of high-density Mediterranean woodlands, particularly in drought-limited sites.