Mixed Mediterranean Forest Research Articles

Climate change and exotic pathogens shift carbon allocation in Mediterranean mixed forests

Abstract Forecasting the effects of global change drivers on ecosystems is one of the most pressing challenges for scientists worldwide. Particularly, climate change and exotic pathogens might have a large impact on plant community dynamics and ecosystem functioning through changes in carbon uptake and sinks. Nevertheless, we still have a poor understanding of the combined effects of these two drivers on plant communities. Here, we explored the impact of rainfall reduction and exotic pathogens on the carbon balance of Mediterranean tree species. For this, we performed a 3‐year field experiment taking advantage of rainfall exclusion infrastructures (30% exclusion) installed in the southernmost European oak forests invaded by the aggressive exotic pathogen Phytophthora cinnamomi. We measured a set of 10 variables representative of tree carbon sources (photosynthetic rates) and sinks (primary production, reproduction, defence, and reserves) in adult trees of three species in two forest types: closed forests of Quercus suber and Q. canariensis, and open woodlands of Q. suber and Olea europaea. We found a large variability in the sensitivity of the different carbon sources and sinks to the effects of drought and pathogens, from variables highly sensitive to both factors (carbon fixation and reproduction, root chemistry) to variables only responsive to drought (litter production) or totally unresponsive (tree trunk, leaf chemistry). Although negative effects predominated, positive effects of rainfall exclusion were also detected in wet years, likely due to a reduction of pathogen abundance in drier soil. Trade‐offs between carbon sinks appeared in all tree species, but rainfall exclusion only modified trade‐offs in Q. suber, the species most susceptible to P. cinnamomi. Synthesis. We provide evidence on the complexity of the combined effects of abiotic (drought) and biotic (pathogens) global change drivers on carbon source and sinks of adult trees, including both negative direct effects and positive indirect effects. Our results showed that these effects varied among co‐existing species, particularly for carbon sinks directly related to tree demography (reproduction). Therefore, long‐term changes in the structure of Mediterranean mixed forests might be expected towards the dominance of species highly resistant to both drought and pathogens.

Mixing oak and pine trees in Mediterranean forests increases aboveground hydraulic dysfunctions.

Increasing tree species diversity in Mediterranean forests could reduce drought-induced hydraulic impairments through improved microclimate and reduced competition for water. However, it remains unclear if and how species diversity modulates tree hydraulic functions and how impacts may shift during the growing season. Using unmanaged Mediterranean forest stands composed of one (i.e., monospecific) or four (i.e., multispecific) tree species, we examined the seasonal dynamics of in-situ hydraulic traits (predawn and midday leaf water potential - Ψpd and Ψmd, xylem- and leaf-specific hydraulic conductivity - KS and KL, percentage loss of conductivity - PLC, specific leaf area - SLA, and Huber value - HV) in four co-existing Pinus and Quercus species over two years. We mainly observed adverse impacts of species diversity with lower Ψpd, Ψmd, KS, KL, and higher PLC in multispecific compared to monospecific stands, especially for the two pines. These impacts were observed all along the growing season but were stronger duringthe driest periods of the summer. Beneficial impacts of diversity were rare and only occured for oaks (Q. faginea) after prolonged and intense water stress. Our findings reveal that mixing oaks and pines could mainly enhance hydraulic impairments for all species during the dry season, suggesting a potential decline in mixed Mediterranean forests under future climate.

Different drought-tolerance strategies of tree species to cope with increased water stress under climate change in a mixed forest.

Trees' functional strategies to cope with extreme drought are essential under climate change. In a mixed Mediterranean forest, we analyzed the functional strategy in response to drought of four co-occurring species (Pinus pinea, Pinus pinaster, Juniperus oxycedrus, and Quercus ilex) during two years. Specifically, we assessed functional traits related to tree water status, leaf water relations, and gas exchange. Different trait-syndrome metrics and the functional strategies under water stress observed suggested a species drought-tolerance differentiation, with the more anysohidric Q. ilex and J. oxycedrus showing a much higher drought tolerance than the more isohydric P. pinea and P. pinaster. All species recovered from negative leaf turgor reached during peak water stress in summer. Q. ilex and J. oxycedrus kept lower leaf osmotic potentials and lower sensitivity of leaf gas exchange and leaf photochemistry to water stress. In contrast, the pine species exhibited more drought-avoidant and water-conservative strategies, yet this behavior was less effective in mitigating water stress's impact on their physiology. The pine species were the most affected by drought, with prolonged near-zero net photosynthesis during summer. P. pinaster was more isohydric than P. pinea and exhibited a lower capacity to maintain leaf turgor. Physiological processes regulating leaf turgor under drought constitute a key functional strategy involved in the carbon and water-related mechanisms, ultimately inducing mortality under hot drought. The currently observed mortality dynamics for P. pinaster, and to a lower extent in P. pinea, may be exacerbated by loss of functional homeostasis.

Common field measures and geometric assumptions of tree shape produce consistently biased estimates of tree and canopy structure in mixed Mediterranean forests

Crown shape and canopy structure are crucial determinants of tree and forest performance, and forest-climate interactions, but methods of quantifying these from field measurements are based on untested assumptions. Historically, measurements of crown size, shape and interactions have been limited to what can be quantified using simple tools such as a tape measure or clinometer, which are then combined with simple geometric assumptions to calculate metrics including crown area, crown volume and canopy cover and overlap. Terrestrial Laser Scanning (TLS) captures the three-dimensional structure of trees and can derive such metrics directly, bypassing the need for geometric assumptions. Here we use TLS data from c. 2500 trees in mixed Mediterranean forests to present a first test of the validity of common assumptions of tree shape. We simulate field measurements from TLS, use these to estimate crown and canopy properties based on geometric assumptions and compare with direct TLS estimates. We find that commonly used methods produce consistently biased estimates of individual tree crown area and crown volume, and whole-plot crown projected area and crown overlap. Although TLS is often not the practical choice for forest measurement, we show that it can guide best practice and here present recommendations of ground measurements that minimise errors.

Wild ungulate effects on soil fungal diversity in Mediterranean mixed forests

The effect of wild ungulate density on ecosystems varies according to different factors: climate and physiography conditions, forest type, management history, and herbivore identity. In this study, we evaluated the effect of historically high densities of red deer (Cervus elaphus L.) on the soil fungal communities in Mediterranean ecosystems using 30 paired plots, open on the one hand and exclosure plots on the other one. Plots were established at the end of 2020 in a perimeter-fenced hunting estate of 6600 ha in Toledo, Spain. Three months after plots were established, we analysed fungal communities in 60 soil samples using Illumina 250-bp paired-end sequencing. We estimated changes in total fungal richness and in the richness of trophic groups through Linear Mixed Effects models using fenced/unfenced type and deer habitat use as fixed variables and the location of the plots and the main tree host species as random variables. Fungal composition was analysed using non-metric multidimensional scaling and permutational multivariate ANOVA; edaphic characteristics were incorporated to explain differences. Soil fungal communities were not differentially affected by excluding ungulates for three months. Areas with high deer densities had a richer saprotrophic community and where lowland environments were dominated by the main tree hosts Quercus faginea and Quercus ilex. Arbutus unedo was found in mountain areas where there was less herbivore pressure, a greater richness of ectomycorrhizal and lichenized fungi and soils positively associated to nitrogen, phosphorus, potassium and organic matter levels.

The alien conifer Cupressus arizonica can outcompete native pines in Mediterranean mixed forests under climate change

The alien conifer Cupressus arizonica can outcompete native pines in Mediterranean mixed forests under climate change

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.

Effects of plant leaf traits, abundance and phylogeny on differentiation of herbivorous insect assemblages in Mediterranean mixed forest

AbstractHerbivorous insect assemblages are functionally diverse, with each species exploiting plant tissues in different ways. Availability and palatability of plant tissues influence the diversity and composition of herbivorous insect assemblages. However, few studies have compared herbivorous insect assemblages and their ecological correlates across multiple plant species within the same plant community. Here, we sampled insect assemblages from the canopies of 1060 plants belonging to 36 woody species in two mixed Mediterranean forest stands. 401 insect species were classified as herbivores and grouped into sucker or chewer guilds. We explored differences in the diversity and composition of each insect guild across plant species, and tested their relationships with plant leaf traits, abundance or phylogeny, and explored whether the structures of plant-herbivorous insect networks depended on any of the studied plant traits. Plant identity accounted for the highest proportion of variation in the composition of each insect guild. Plant species abundance showed a positive effect on both insect guilds’ diversity. Suckers’ diversity was higher in plant species with deciduous leaves and low SLA, while the composition was more similar between phylogenetically closer plant species. Chewers diversity increased with the leaf area, while plants with similar LA, leaf nitrogen, SLA and distinct leaf habit showed more similar assemblages. Similarly, closely related angiosperms showed similar chewer assemblages. Plant–insect interaction networks present a modular structure, in which plants belonging to the same module tend to be related and share more sucker species. We add to the evidence supporting the role of plant species features as filters for structuring their associated herbivore insect assemblages.

Increased belowground tree carbon allocation in a mature mixed forest in a dry versus a wet year.

Tree species differ in their carbon (C) allocation strategies during environmental change. Disentangling species-specific strategies and contribution to the C balance of mixed forests requires observations at the individual tree level. We measured a complete set of C pools and fluxes at the tree level in five tree species, conifers and broadleaves, co-existing in a mature evergreen mixed Mediterranean forest. Our study period included a drought year followed by an above-average wet year, offering an opportunity to test the effect of water availability on tree C allocation. We found that in comparison to the wet year, C uptake was lower in the dry year, C use was the same, and allocation to belowground sinks was higher. Among the five major C sinks, respiration was the largest (ca. 60%), while root exudation (ca. 10%) and reproduction (ca. 2%) were those that increased the most in the dry year. Most trees relied on stored starch for maintaining a stable soluble sugars balance, but no significant differences were detected in aboveground storage between dry and wet years. The detailed tree-level analysis of nonstructural carbohydrates and δ13 C dynamics suggest interspecific differences in C allocation among fluxes and tissues, specifically in response to the varying water availability. Overall, our findings shed light on mixed forest physiological responses to drought, an increasing phenomenon under the ongoing climate change.

Short-term effect of thinning on inter- and intra-annual radial increment in Mediterranean Scots pine-oak mixed forests

Thinning treatments along with the establishment of mixed forest stands have been put forward as possible adaptation strategies to cope with climate change, although the effectiveness of combining these two measures has scarcely been studied and may vary depending on stand conditions and the thinning regime employed. The aim of this study was to better understand the effect of commercial thinning and of the different growth behavior of two coexisting species on their inter- and intra-annual cumulative radial increment patterns. For this purpose, we studied radial increment in a Scots pine-Pyrenean oak (Pinus sylvestris L.-Quercus pyrenaica Willd.) Mediterranean mixed forest in north-west Spain over two climatically contrasting years (2016–2017). The data came from a thinning trial consisting of a randomized latin square design with a control and two commercial thinning treatments from below; one moderate and the other heavy, removing 25% and 50 % of initial basal area, respectively, of both species. The radial increment was analyzed based on bi-weekly readings from band dendrometers installed on 90 oak and pine trees. A non-linear mixed model based on double-Richards curve was fitted to explore the differences between thinning treatments and species response in the intra-annual cumulative radial increment patterns. Inter-annual basal area increments for each species at stand level were quantified by aggregating the tree estimates obtained from the model fitted in the first step. Tree and stand level growth were greater in Scots pine, which also showed a greater growth response to early spring droughts than the Pyrenean oak. Heavy thinning increased radial increment in trees of both species at the expense of decreased total stand basal area. At species level, basal area growth in Scots pine decreased through thinning, whereas for Pyrenean oak, the heavy thinning intensity resulted in the same basal area growth as the control. Thus, heavy thinning induced a trade-off between total stand growth and tree-level response to climatic conditions for Scots pine but with no loss in productivity in the case of the Pyrenean oak. Hence, heavy thinning may be an appropriate measure to attain productive stability of the oak coppice in the studied mixed forest as well as to adapt tree growth to future droughts associated with climate change.

Assessing carbon and water fluxes in a mixed Mediterranean protected forest under climate change: An integrated bottom – up and top – down approach

This paper proposes a framework based on the integration of bottom-up and top-down approaches to assess and quantify the effect of local climate change on carbon and water fluxes in a protected Mediterranean mixed forest. The forest, with an extent of 50 ha, is in a coastal zone and is in decline because of severe summer drought stress. Since 2018, it has been involved in the LIFE PRIMED ecological restoration project.The bottom-up approach used gas exchange measurements and the MOCA model (MOdelling for Carbon Assessment) to quantify photosynthesis and transpiration of the main species in the forest (P. latifolia, P. lentiscus, Q. cerris, and F. angustifolia sub. oxycarpa). The top-down approach used the satellite models C-fix and NDVI-CWS, to quantify and spatialise gross primary production (GPP), net primary production (NPP), and evapotranspiration (ETA) at the forest stand scale. The up-scaling process from leaf to forest stand was made possible by calibrating two key parameters: light use efficiency (ε) and vegetation transpiration coefficient (Kcveg).The results indicate a decline in GPP, NPP, and ETA during the study period (2018–2022) due to the summer drought conditions caused by rising temperatures (16.60 °C in 2018 vs. 16.98 °C in 2022), increased vapor pressure deficit, reduced soil water content, and rainfall (1328 mm in 2018 vs. 468 mm in 2022). In 2018, the annual GPP was 1186.83 ± 351.40 gC/m2y, which decreased to 1128.67 ± 331.21 gC/m2y in 2022. During the dry summer period, GPP ranged from 401.76 ± 119.54 gC/m2y in 2018 to 347.39 ± 101.62 gC/m2y in 2022. Annual ETA rates varied between 1184.86 ± 351.86 mmH₂O/m2y (2018) and 1304.92 ± 384.07 mmH₂O/m2y (2022). The summer ETA rates were significantly influenced by drought conditions, decreasing from 638.73 ± 190.67 mmH₂O/m2y in 2018 to 390.35 ± 108.83 mmH₂O/m2y in 2022.Based on the observed trend in other Mediterranean areas, species-specific responsiveness to local climate change suggests that P. latifolia and P. lentiscus, which are better adapted to water-limited environments, could gradually replace Q. cerris and F. angustifolia subs. oxycarpa.This study provides useful quantitative indications for the management and restoration goals of the area and suggests a methodology that can be applicable in several management practices, particularly for small heterogeneous and fragmented forested areas such as the Mediterranean ones.

Close-to-nature management effects on tree growth and soil moisture in Mediterranean mixed forests

Given the rising frequency of drought events, close-to-nature principles appear to be a suitable management option for promoting more adapted forests by increasing diversity and vitality (e.g., enhancing tree growth), while preserving soil moisture. In this study, we evaluate individual tree growth from selected trees (including future crop trees) and soil moisture responses to close-to-nature silvicultural treatments in sub-humid Mediterranean mixed forests in Catalonia (Northeastern Spain). We relied on 18 permanent sample plots (i.e., 9 paired-plots: managed / control) representing a competition intensity gradient and different forest types (dominated by either Quercus ilex, Castanea sativa, Quercus pubescens/petraea, Pinus pinea and Pinus sylvestris). From them, 8 plots were subject to intense monitoring in terms of intra-annual tree radial growth, monitored by manual band dendrometers, and hourly soil water content. Basal area increment, and annual and seasonal (May, June, July) radial increment rates were computed as a proxy of tree growth for all plots and for those plots with dendrometers, respectively. We conducted Spearman correlation analyses between tree growth, competition intensity, soil water content and climatic variables to detect significant relations between forest management and variables of interest (e.g., basal area increment, radial increment rates, soil water content). To investigate further into the effect of this silviculture on tree growth, we used mixed-effects models using as predictors variables related to competition intensity and forest management (dummy variable) in combination with monthly soil water content variables. We observed that basal area increment and radial increment rates had a positive (albeit weak) correlation with forest management, while competition intensity had a negative correlation with both variables. The models were not capable of detecting clear species-specific tree growth responses to competition. Compared to control plots, on average, the treatments boosted soil water content by about a 10% in managed plots in Q. ilex, P. sylvestris and P. pinea stands, and this increase was particularly significant during the growing season. Moreover, the best-fitted model predicted seasonal radial increment rates using not only forest management and competition variables, but also monthly soil water content factors. Our results suggest that close-to-nature-forest management may reduce the vulnerability to drought by increasing tree vitality and soil moisture, at least in the short term (<3 years).

Radial growth, wood anatomical traits and remote sensing indexes reflect different impacts of drought on Mediterranean forests

Drought reduces canopy cover, productivity and tree growth in forests. However, there is still little knowledge on how drought affects coupling between canopy greenness assessed by remote sensing and hydraulic conductivity detected by wood anatomy. This combination could improve the understanding of forest response to climate change. Thus, we investigated the impacts of a hot drought, which occurred in summer 2017, on radial growth, earlywood hydraulic diameter (Dh), a proxy of conductivity, and several remote-sensing indices in mixed Mediterranean hardwood forests (Quercus pubescens Willd. – Fraxinus ornus L.). In general, growth showed a higher coherence among trees and a higher responsiveness to climate. Growth decreased during the drought year, particularly for Q. pubescens, which showed high defoliation and dieback intensity. Both species showed a decline of Dh in 2018 after the drought and subsequent warm winter conditions. We found positive relationships between Dh and remote-sensing data for Q. pubescens in some of these vulnerable sites, where (i) growth was constrained by dry spring-summer conditions and (ii) Dh and growth covaried. These findings indicate a high variability among sites and tree species in their responses to drought considering earlywood anatomy, growth canopy cover and water content. However, some common patterns emerge such as links between potential hydraulic conductivity (Dh), tree cover and Dh-growth covariation in the most impacted sites. Further, F. ornus seem to perform better in terms of growth under drought conditions, showing less mortality and dieback than Q. pubescens. Future studies could explore how water transport and changes in canopy cover respond to dry and warm conditions and if that covariation indicates vulnerability to drought.

Temporal Variations in Temperature and Moisture Soil Profiles in a Mediterranean Maquis Forest in Greece

Soil moisture (SM) and temperature (ST) are critical factors in forest eco-hydrological research. In this study, we investigated the inter- and intra-annual changes in SM and ST profiles in a mixed Mediterranean maquis forest stand together with soil and meteorological parameters. Hourly data from three field measurements points at four depths (−5, −20, −40 and −70 cm) for 6 years were interpolated using the kriging method to produce annual SM and ST profiles. The results indicate that air temperature highly affects the upper 5 cm of the mineral soil. In general, it increases with depth in winter at an average rate of 0.036 °C/cm and decreases in summer (0.035 °C/cm), presenting higher values compared to air temperature from April to August and lower ones during the rest of the period. Precipitation is the main factor driving SM variations up to a superficial soil depth of 40 cm. The upper soil layer (0–40 cm) infiltrates water faster and presents high SM variability, especially in monthly and seasonal (year to year) time steps. The maquis forest stands are likely to be strongly affected by climate change, therefore the results of this study could be useful in hydrological and climate change studies focused on maquis vegetation water management.

Plant Traits and Phylogeny Predict Soil Carbon and Nutrient Cycling in Mediterranean Mixed Forests

Soil functioning is closely linked to the interactions between biological communities with the physical environment. Yet, the impact of plant community attributes on metabolic processes promoting soil nutrient cycling remains largely unknown. We hypothesized that the plant community acts as a regulating agent of nutrient mobilization in soils according to the phylogenetic and morpho-functional traits of plant species of which it is composed. Rhizosphere soils were collected in autumn and spring under 32 tree and shrub species in two Mediterranean mixed forests (four plots in each) located in southern Spain, and nine soil enzymatic activities related to C, N and P mobilization were assessed. Phylogeny and morpho-functional traits of plant species were recorded and their imprint in soil enzymatic activities across forests was determined. The results showed a plant phylogenetic signal for N mobilization in both forests, while it varied across forests for non-labile C and P mobilization. The plant phylogenetic signals were primarily driven by lineages that diversified through the Miocene, about 25 Myr ago. In addition, leaf traits and plant’s mycorrhizal type explained soil enzymatic activities independently from phylogeny. C and P mobilization increased under ectomycorrhizal plants, whilst enhanced N mobilization did occur under arbuscular mycorrhizal ones. The plant community composition led to a different carbon and nutrient mobilization degree, which in turn was mediated by distinct microbial communities mirroring differentiated resource-acquisition strategies of plants. Our results highlight the role of plant traits and mycorrhizal interactions in modulating carbon and nutrient cycling in Mediterranean mixed forest soils.

Positive effects of warming do not compensate growth reduction due to increased aridity in Mediterranean mixed forests

AbstractIncreased winter and early spring temperatures due to climate change can enhance forest productivity due to earlier growth onset in temperate regions. However, drought‐prone forests can be highly vulnerable to the combined effect of high summer temperatures and water stress. Understanding how water stress and rising temperatures along the growing season control tree growth in co‐occurring species with contrasting drought tolerances is key to project climate change effects on forest dynamics. Here, we evaluated the interactive effect of seasonal temperature and water availability on annual radial growth and climate–growth instability over six decades (1951–2015) in three pine species with contrasting drought tolerances (Pinus pinaster, Pinus nigra, and Pinus sylvestris). For this, we retrospectively observed radial tree growth using dendroecological methods and evaluated seasonal temperature and water availability effects by using linear mixed models. Early growing season temperature and water availability had a positive effect on tree growth, but the positive effect of late season temperature was modulated by water availability. Moving time‐window analyses revealed temporal instability in climate–growth relationships. Since the 1980s, pine species showed a higher growth sensitivity to both seasonal temperatures and annual water availability. Furthermore, growth reductions were more pronounced due to increased summer temperatures and reduced precipitation. Our results were similar for the three studied pine species despite their contrasting tolerance to drought. Overall, climate warming effects on pine growth are contingent upon water availability in Mediterranean continental forests. Synchronization among species, climate–growth instability, and negative growth trends suggests an increased vulnerability to drought of Mediterranean pine species in response to ongoing climate change.

Spatial phylogenetic and phenotypic patterns reveal ontogenetic shifts in ecological processes of plant community assembly

The analysis of the spatial phylogenetic and phenotypic structure of plant communities can provide insight into the underlying processes and interactions governing their assembly, and how these may change during plant ontogeny. We used point pattern analysis to find out if saplings and adult plants are surrounded by phylogenetically and phenotypically more similar or more dissimilar neighbours than expected by chance, and whether these associations change from the sapling to the adult stage. To this end, we combined information on the phylogenetic structure and eight phenotypic traits of 15 woody plant species in two Mediterranean mixed forests of southeastern Spain. At the community level, we found that the sapling bank at both sites did not show phylogenetic or phenotypic spatial patterns, but adults showed phylogenetic clustering (i.e. heterospecific neighbours were more similar than expected). At the species level, we found frequently repulsive patterns in the sapling bank of less abundant species (i.e. heterospecific sapling or adult neighbours were more dissimilar than expected) in both, phylogenetic and phenotypic analyses. For the adult stage, we found phylogenetic attraction (i.e. more similar neighbours) in just one species and phenotypic clustering in four species. The processes driving the assembly of the communities of saplings and adults leave detectable signals in the spatial phylogenetic and phenotypic structure of our two forest communities. Our findings reinforce the existence of ontogenetic shifts in the mechanisms involved in plant community assembly. Facilitation between phylogenetically distant and phenotypically divergent species favours the recruitment of less abundant species. However, processes acting later in the ontogeny ameliorate the competition between close relatives and determine the spatial structure of adult plants. Nevertheless, the role of phenotype in shaping the interactions between adult plants was context‐ and trait‐dependent. The use of spatial point pattern analysis allowed a nuanced interpretation of the phylogenetic and phenotypic structures of plant communities.

Forest type, microhabitat conditions and human presence predict occurrence of two sympatric Columbidae species in the Mediterranean’s largest mixed forest: Implications for management and conservation

The presence of natural forests in association with the planted forests has become a common quite landscape in the Mediterranean basin and even elsewhere. Understanding how native game bird use the different types of forest habitats is a major issue aiming at an effective assessment of their management and conservation. In this study, we investigated this issue with regards to the common Woodpigeon (WD) (Columba palumbus) and the vulnerable European Turtle-Dove (TD) (Streptopelia turtur) in Mamora forest, the largest mixed Mediterranean forest. We used data from three sets of variables, namely forest type, microhabitat conditions and human disturbance, to determine the best predictors that cause variation in the WD and TD occurrence by means of generalized linear mixed models. Our results showed that pine and eucalypt plantations were the most attractive for TD and natural cork oak forests for WD. In pine plantations, TD occurrence was influenced positively by the extent of the herbaceous layer and negatively by the nearest distance to the shepherd’s house with enclosure, whereas in eucalyptus plantations, it was influenced positively by the cover of trees and shrubs. In natural cork oak forests, WD was exclusively affected by the tree cover. Variation partitioning analysis revealed that space was the most robust in explaining the TD occurrence in pine (adj. R2 = 0.20, p < 0.001) and eucalyptus (adj. R2 = 0.20, p < 0.001) plantations, whereas in cork oak natural forests, human presence (adj. R2 = 0.15, p = 0.002) was the most relevant in explaining WD occurrence. Specific management actions are suggested to improve the attractiveness of gamebirds at natural and artificial forests of the Mamora. It is imperative to test the geographical generality of our results in other mixed Mediterranean forests.

Scale dependency of ectomycorrhizal fungal community assembly processes in Mediterranean mixed forests

The assembly of biological communities depends on deterministic and stochastic processes whose influence varies across spatial and temporal scales. Although ectomycorrhizal (ECM) fungi play a key role in forest ecosystems, our knowledge on ECM community assembly processes and their dependency on spatial scales is still scarce. We analysed the assembly processes operating on ECM fungal communities associated with Cistus albidus L. and Quercus spp. in Mediterranean mixed forests (Southern Spain), for which root tip ECM fungi were characterized by high-throughput sequencing. The relative contribution of deterministic and stochastic processes that govern the ECM fungal community assembly was inferred by using phylogenetic and compositional turnover descriptors across spatial scales. Our results revealed that stochastic processes had a significantly higher contribution than selection on root tip ECM fungal community assembly. The strength of selection decreased at the smallest scale and it was linked to the plant host identity and the environment. Dispersal limitation increased at finer scales, whilst drift showed the opposite pattern likely suggesting a main influence of priority effects on ECM fungal community assembly. This study highlights the potential of phylogeny to infer ECM fungal community responses and brings new insights into the ecological processes affecting the structure and dynamics of Mediterranean forests.

Competition and climate influence in the basal area increment models for Mediterranean mixed forests

Competition plays a key role controlling tree growth in mixed forests. Contrary to monocultures, quantifying species mixing influence on tree growth suppose a challenge since the presence of two or more species requires to estimate the degree of intra- and inter-specific competition among trees. Moreover, it is well known that aridity can also influence tree growth, especially in the Mediterranean Basin. In the present context of climate change, it is essential to take into account species mixing and aridity uncertainty in the design of sustainable management guidelines for Mediterranean mixed forests. To achieve that, data from Spanish National Forest Inventory was used in this study to fit new mixed-effects basal area increment (BAI) models for 29 two-species compositions in Spain. A wide range of different competition structures (intra-specific, inter-specific, size-symmetric and size-asymmetric) and aridity conditions (in terms of the De Martonne Index) were included and tested into the BAI models. Parameter estimations were obtained for all possible species, mixtures and combinations by Maximum Likelihood (ML). Models with all the coefficients being significant (p < 0.05) were first selected. Among these models, we used Akaike Evidence Ratios for selecting the best one by species for each mixture. The best model for each species and mixture was used to analyze the competition and climatic influence on tree growth. Regarding competition influence, a common trend among mixtures was found with higher productivity in mixed than pure stands, suggesting that BAI values may increase with the increment of species diversity. Based on intra and inter-specific competition indexes, competition seemed to be the most representative biological interaction in conifer-conifer mixtures, since neutralism and facilitation may occur more frequently in conifer-broadleaved and broadleaved-broadleaved mixtures. Our findings also suggested that tree growth may be significantly limited by arid conditions, excepting for Pinus halepensis and Pinus pinea. Our rigorous modelling approach successfully uncovered not only possible mixing effect among various species but also help us to understand the effect of aridity on tree growth. Thus, models presented in this study can be used in the design and implementation of management and adaptation guidelines under future climate change scenarios.