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Fungal fruit-bodies are an important economic resource both for recreational pickers and commercial markets. The broad interest in forest fungi and mushrooms has motivated attempts to develop appropriate silvicultural methods able to preserve and improve mushroom yields. Defining best practices for the sustainability and profitability of forest fungal resources is the main aim of 'mycosilviculture'. However, the difficulty of monitoring and studying such a cryptic kingdom (i.e., fungi) under real forest conditions has led to rather scattered scientific knowledge of the effects of forest management regimes and silvicultural operations on wild mushroom resources. Here, we review the current scientific literature regarding the influence of (i) stand characteristics, i.e. stand age, stand density, canopy cover and tree species composition, (ii) silvicultural practices and (iii) other management-related disturbances affecting the yield of wild epigeous mushrooms, with the aim of systematizing existing scientific knowledge and identifying gaps in knowledge in order to suggest future lines of research. Most of the research in the field of mycosilviculture to date has focused on ectomycorrhizal mushrooms, which include the majority of wild commercial mushrooms. The main findings from the literature indicate that forest management practices, by modifying stand characteristics and microclimatic conditions, can influence wild mushroom occurrence and productivity, both positively and negatively, depending on the specific fungal ecological needs, reproductive strategies, forest type and management regime. Further research efforts relating to all aspects of the interaction between forest management, fungal ecology and mushroom production are needed: in particular, additional research efforts should be devoted to understanding mushroom yield dynamics in unevenaged and mixed forests and evaluating the effects of regeneration methods on fungal communities. ; The review was carried out within the COST Action FP1203 (European Non-Wood Forest Products). Sergio de-Miguel was supported by the European Union's Horizon 2020 MultiFUNGtionality Marie Skłodowska-Curie (IF-EF No-655815) and José Antonio Bonet benefited from a Serra-Hunter Fellowship provided by the Generalitat of Catalunya. The study was also partially funded by the Spanish research project AGL2015-66001-C3-1-R (MEC Spain) and by the European project StarTree (No. 311919).

Mushrooms are important non-wood-forest-products in many Mediterranean ecosystems, being highly vulnerable to climate change. However, the ecological scales of variation of mushroom productivity and diversity, and climate dependence has been usually overlooked due to a lack of available data. We determined the spatio-temporal variability of epigeous sporocarps and the climatic factors driving their fruiting to plan future sustainable management of wild mushrooms production. We collected fruiting bodies in Pinus sylvestris stands along an elevation gradient for 8 consecutive years. Overall, sporocarp biomass was mainly dependent on inter-annual variations, whereas richness was more spatial-scale dependent. Elevation was not significant, but there were clear elevational differences in biomass and richness patterns between ectomycorrhizal and saprotrophic guilds. The main driver of variation was late-summer-early-autumn precipitation. Thus, different scale processes (inter-annual vs. spatial-scale) drive sporocarp biomass and diversity patterns; temporal effects for biomass and ectomycorrhizal fungi vs. spatial scale for diversity and saprotrophic fungi. The significant role of precipitation across fungal guilds and spatio-temporal scales indicates that it is a limiting resource controlling sporocarp production and diversity in Mediterranean regions. The high spatial and temporal variability of mushrooms emphasize the need for long-term datasets of multiple spatial points to effectively characterize fungal fruiting patterns. ; This study was funded by the Spanish research projects AGL2012-40035-C03-01 and AGL2015-66001-C3-1-R (MEC Spain) and by the European project StarTree (No. 311919). JGA was supported by Juan de la Cierva-fellowship (IJCI-2014-21393) and DYMECO project and SdM by the European Union's Horizon 2020 MultiFUNGtionality Marie Skłodowska-Curie (IF-EF No. 655815) and JAB benefit from Serra-Hunter Fellow provided by the Generalitat of Catalunya.

Background: The prediction of biogeographical patterns from a large number of driving factors with complex interactions, correlations and non-linear dependences require advanced analytical methods and modeling tools. This study compares different statistical and machine learning-based models for predicting fungal productivity biogeographical patterns as a case study for the thorough assessment of the performance of alternative modeling approaches to provide accurate and ecologically-consistent predictions. Methods: We evaluated and compared the performance of two statistical modeling techniques, namely, generalized linear mixed models and geographically weighted regression, and four techniques based on different machine learning algorithms, namely, random forest, extreme gradient boosting, support vector machine and artificial neural network to predict fungal productivity. Model evaluation was conducted using a systematic methodology combining random, spatial and environmental blocking together with the assessment of the ecological consistency of spatially-explicit model predictions according to scientific knowledge. Results: Fungal productivity predictions were sensitive to the modeling approach and the number of predictors used. Moreover, the importance assigned to different predictors varied between machine learning modeling approaches. Decision tree-based models increased prediction accuracy by more than 10% compared to other machine learning approaches, and by more than 20% compared to statistical models, and resulted in higher ecological consistence of the predicted biogeographical patterns of fungal productivity. Conclusions: Decision tree-based models were the best approach for prediction both in sampling-like environments as well as in extrapolation beyond the spatial and climatic range of the modeling data. In this study, we show that proper variable selection is crucial to create robust models for extrapolation in biophysically differentiated areas. This allows for reducing the dimensions of the ecosystem space described by the predictors of the models, resulting in higher similarity between the modeling data and the environmental conditions over the whole study area. When dealing with spatial-temporal data in the analysis of biogeographical patterns, environmental blocking is postulated as a highly informative technique to be used in cross-validation to assess the prediction error over larger scales. ; This work was supported by the Secretariat for Universities and of the Ministry of Business and Knowledge of the Government of Catalonia and the European Social Fund. This work was partially supported by the Spanish Ministry of Science, Innovation and Universities (Grant No. RTI2018–099315-A-I00). J.A.B. benefitted from a Serra-Húnter Fellowship provided by the Government of Catalonia.

Fungi provide important forest ecosystem services worldwide. In Mediterranean pine forests, predicted warmer and drier conditions could lead to a decline in mushroom yields. Climate is a key factor regulating both tree growth and fungal yields, particularly in drought-prone Mediterranean ecosystems. However, the responses of forest growth and mushroom production to climate depend on the differences among tree and fungal species and functional groups (e.g., mycorrhizal vs. saprotrophic), forest types, as well as depending on site conditions. Here we investigate how climatic conditions drive seasonal wood formation (earlywood −EW− and latewood −LW− production) and mycorrhizal mushroom production, to disentangle if growth and fungal yields are related. This assessment was done in Mediterranean forests dominated by four pine species in two areas located in Catalonia (NE Spain) representing mesic and xeric conditions and encompassing wide ecological gradients. The data consisted of 7-year to 13-year long inventories of mushroom production. EW production was favoured by cold and wet climate conditions during the previous fall and winter, and during the current spring and summer. LW production was enhanced by warm and humid conditions from spring to early fall. Mushroom yield was improved by wet late-summer and fall conditions, mainly in the most xeric area. This study confirms the ample differences found in tree growth and fungal production along ecological and climatic gradients. Clear relationships between mycorrhizal fungal yields and tree growth were mostly observed in specific sites characterized by severe summer drought. Specifically, latewood production seems to be the tree-ring variable most tightly linked to mycorrhizal fungal yield in drought-prone areas. ; This study was partially funded by the research projects AGL2012-40035-C03-01 and AGL2015-66001-C3-1-R (Ministerio de Economía y Competitividad of Spain, Secretaría de Estado de Investigación, Desarrollo e Innovación), by the European project 'StarTree—Multipurpose trees and non-wood forest products: a challenge and opportunity' under grant agreement No. 311919. Irantzu Primicia work was supported by a STSM Grant from the COST Action FP1203 (European Non-Wood Forest Products). Sergio de Miguel's work was supported by the European Union's Horizon 2020 research and innovation programme within the framework of the MultiFUNGtionality Marie Skłodowska-Curie Individual Fellowship (IF-EF) under grant agreement No 655815.

Fungi are responsible for many of the processes that occur in natural ecosystems and largely determine forest ecosystem dynamics, such as the ability of trees to access limiting nutrients and sequester carbon. Understanding and predicting climate change impacts on fungal dynamics over large scales is key in order to gain further insights into the effects of global change on natural ecosystem functioning and related ecosystem services. In this study, we use predictive models based on machine learning algorithms to estimate, in a spatially explicit way, the historical and future (1976–2100) evolution of mycorrhizal and saprotrophic macrofungal productivity in Mediterranean forest areas under climate change scenarios. The greatest changes in total productivity, as well as mycorrhizal fungi, are predicted to occur in subalpine and montane pine forests, where fungal productivity is estimated to decrease, and will be more pronounced under climate change scenarios with higher expected increase in temperature. In contrast to mycorrhizal species, saprotrophic fungi could benefit from pronounced changes in climate and increase their productivity in supra- and mesomediterranean regions at mid-range elevations. Moreover, we estimated that fungal productivity has also changed historically in some scattered areas where changes in climate over the years may have led to a decrease in productivity. This study contributes to raising awareness on the need for anticipating potential global change impacts on this key element of ecosystem functioning, and for deploying possible management policies oriented toward maintaining the important role of fungal productivity in both climate change mitigation and adaptation. ; We thank the "Agencia Estatal de Meteorología (AEMET)" and "Servei Meteorològic de Catalunya (SMC)" for providing daily weather station data. This work was supported by the Secretariat for Universities and of the Ministry of Business and Knowledge of the Government of Catalonia and the European Social Fund. This work was ...

Wild mushrooms contribute to a variety of ecosystem services. The expected warmer and drier conditions for the Mediterranean region as a consequence of climate change, are raising concerns about future mushroom productivity due to potential reduction of soil water availability for fungi. The aim of this study was to increase our understanding of the interaction between climate and soil moisture in relation to their impact on mushroom productivity in Mediterranean forests. Mushroom yield data were obtained from 28 permanent mushroom inventory plots intensively monitored in Maritime pine (Pinus pinaster Ait.) stands of northeastern Iberian Peninsula. Annual productivity of total, edible and marketed mushrooms was obtained from measurements conducted every week during the autumn fruiting season for years 2008–2015. Historical weather conditions were obtained through data interpolation from meteorological stations. Soil moisture data were obtained from continuous plot-level measurements. A process-based soil water balance model was used to predict soil moisture under two climate change scenarios, using the predictions of two different regional climate models. Mixed-effects models using either precipitation or soil moisture as predictors, in combination with other weather variables, were fitted to annual mushroom occurrence and yield data. Mushroom yield was primarily dependent on weather and soil moisture conditions during the same month, with the exception of precipitation, whose effects exhibited a one-month delay. High temperatures limited mushroom yield at the beginning of the fruiting season, but tended to enhance it towards the end. The analysis revealed no apparent negative effect of climate change on long-term mushroom productivity, but rather the opposite (i.e., predicted median productivity of marketed mushrooms for 2016–2100 was 23–93% higher compared to the current yield), mainly due to an elongation of the fruiting season arising from the combined effect of increased precipitation at the beginning of the season and warmer temperatures at the end. ; This work benefited from the Erasmus Mundus Master Program MEDFOR (Mediterranean Forestry and Natural Resources Management) which provided a scholarship to the first author. Sergio de Miguel was supported by the European Union's Horizon 2020 MultiFUNGtionality Marie Skłodowska-Curie (IF-EF No 655815), Miquel De Cáceres benefited from a Ramon y Cajal fellowship (RYC-2012-11109), and José Antonio Bonet benefited from a Serra-Húnter Fellowship provided by the Generalitat of Catalunya. Additional funding came from the research projects MYCOSYTEMS (AGL2015-66001-C3-1-R − MEC Spain) and FORESTCAST (CGL2014- 59742-C2-2-R), and from the European project StarTree (No. 311919). Special thanks to Mr. Antoine Cabon for his assistance with R software.

Integrating fungal-based ecosystem services into forest management planning and policy-making requires quantitative knowledge of the yields of fungal sporocarps and their environmental drivers. The aim of this study was to predict edible mushroom yield in Pinus pinaster forests of Central Spain, based on a 17-year data series. Two-stage mixed-effects models were used to examine the effect of predictors on mushroom occurrence and yield separately with the aim of providing further insight into the ecological system. Changes in seasonal precipitation represented the main weather-related driver affecting sporocarp emergence and production, since they were both positively influenced by late summer and early autumn precipitation. Soil acidity positively influenced Lactarius yield. Stand age and sandy soils showed a negative influence on mushroom production. The diversity of drivers became more apparent at the fungal species level. The models can be used for predicting the production of edible fungi under different meteorological and site conditions. ; This study was partially funded by the research projects AGL2012-40035-C03-01 and AGL2012-40035-C03-03 (Ministerio de Economía y Competitividad of Spain, Secretaría de Estado de Investigaci on, Desarrollo e Innovaci on), by the Micosylva þ project (Interreg IVB SUDOE SOE3/P2/E533), and by the Mycological Programme of Castilla y Leon (www.micocyl.es). This study also received funding from the European Union's Horizon 2020 research and innovation programme within the framework of the Multi- FUNGtionality Marie Skłodowska-Curie Individual Fellowship (IFEF) under grant agreement No 655815. This work further benefited from the Erasmus Mundus Master Course MEDFOR (Mediterranean Forestry and Natural Resources Management) (520137-1-2011-1- PT-ERA MUNDUS-EMMC), which provided one scholarship to Mr. Zelalem Mengiste Taye. The inventory and monitoring of the sample plots was funded by Junta de Castilla y Leon and ADEMA from 1997 to 2007, and by Junta de Castilla y Leon and CESEFOR from 2007 to 2014.

Fungi provide important forest ecosystem services worldwide. In Mediterranean pine forests, predicted warmer and drier conditions could lead to a decline in mushroom yields. Climate is a key factor regulating both tree growth and fungal yields, particularly in drought-prone Mediterranean ecosystems. However, the responses of forest growth and mushroom production to climate depend on the differences among tree and fungal species and functional groups (e.g., mycorrhizal vs. saprotrophic), forest types, as well as depending on site conditions. Here we investigate how climatic conditions drive seasonal wood formation (earlywood −EW− and latewood −LW− production) and mycorrhizal mushroom production, to disentangle if growth and fungal yields are related. This assessment was done in Mediterranean forests dominated by four pine species in two areas located in Catalonia (NE Spain) representing mesic and xeric conditions and encompassing wide ecological gradients. The data consisted of 7-year to 13-year long inventories of mushroom production. EW production was favoured by cold and wet climate conditions during the previous fall and winter, and during the current spring and summer. LW production was enhanced by warm and humid conditions from spring to early fall. Mushroom yield was improved by wet late-summer and fall conditions, mainly in the most xeric area. This study confirms the ample differences found in tree growth and fungal production along ecological and climatic gradients. Clear relationships between mycorrhizal fungal yields and tree growth were mostly observed in specific sites characterized by severe summer drought. Specifically, latewood production seems to be the tree-ring variable most tightly linked to mycorrhizal fungal yield in drought-prone areas. ; This study was partially funded by the research projects AGL2012-40035-C03-01 and AGL2015-66001-C3-1-R (Ministerio de Economía y Competitividad of Spain, Secretaría de Estado de Investigación, Desarrollo e Innovación), by the European project "StarTree—Multipurpose trees and non-wood forest products: a challenge and opportunity" under grant agreement No. 311919. Irantzu Primicia work was supported by a STSM Grant from the COST Action FP1203 (European Non-Wood Forest Products). Sergio de Miguel's work was supported by the European Union's Horizon 2020 research and innovation programme within the framework of the MultiFUNGtionality Marie Skłodowska-Curie Individual Fellowship (IF-EF) under grant agreement No 655815.

Key message. No temporal change was recorded during summer in fuel availability in Pinus pinaster stands, contrary to predictions from the Forest Fire Weather Index. Also, thinning had no mid-term effect on fuel moisture or canopy structure. Context. Forest fires are a major problem in Mediterranean countries. Management actions, such as fuel reductions, are one of the main tools to diminish fire risk, but the midterm efficacy of such tools remains largely untested with empirical data. Aims. Here, we test for midterm effects of thinning on fuel moisture and crown bulk density in P. pinaster stands and whether temporal variations in fuel moisture correlated with predictions from the Fire Weather Index, a commonly used index on fire risk, and its components. Methods. We compared fuel moisture over a fire season and crown bulk density in nine pairs of thinned/unthinned plots 7 years after treatments were applied. Results. We observed that fuel moisture remained stable during a fire season, as a likely result of drought-induced physiological adjustments, including stomatal regulation and others, which allow leaves to maintain a large humidity even during drought, and that thinning had no midterm effect on fuel moisture or crown bulk density. Moreover, the Fire Weather Index and its components displayed different temporal dynamics than those observed in fuel moisture. Conclusion. These results are important as they indicate that thinning may only have a limited, short-term impact towards diminishing the potential for crown fire spread in these stands and that current indices to evaluate fire risk may require a re-evaluation. ; This study is funded by the Spanish Government (RYC-2012-10970, AGL2015-69151-R, AGL2015-68274-C3-3-R).

Drying soil samples before DNA extraction is commonly used for specific fungal DNA quantification and metabarcoding studies, but the impact of different drying procedures on both the specific fungal DNA quantity and the fungal community composition has not been analyzed. We tested three different drying procedures (freeze-drying, oven-drying, and room temperature) on 12 different soil samples to determine (a) the soil mycelium biomass of the ectomycorrhizal species Lactarius vinosus using qPCR with a specifically designed TaqMan® probe and (b) the fungal community composition and diversity using the PacBio® RS II sequencing platform. Mycelium biomass of L. vinosus was significantly greater in the freeze-dried soil samples than in samples dried at oven and room temperature. However, drying procedures had no effect on fungal community composition or on fungal diversity. In addition, there were no significant differences in the proportions of fungi according to their functional roles (moulds vs. mycorrhizal species) in response to drying procedures. Only six out of 1139 operational taxonomic units (OTUs) had increased their relative proportions after soil drying at room temperature, with five of these OTUs classified as mould or yeast species. However, the magnitude of these changes was small, with an overall increase in relative abundance of these OTUs of approximately 2 %. These results suggest that DNA degradation may occur especially after drying soil samples at room temperature, but affecting equally nearly all fungi and therefore causing no significant differences in diversity and community composition. Despite the minimal effects caused by the drying procedures at the fungal community composition, freeze-drying resulted in higher concentrations of L. vinosus DNA and prevented potential colonization from opportunistic species. ; This work was supported by a STSM Grant from COST Action FP1203 and by the Spanish Ministry of Economy and Competitivity (MINECO) through the project AGL 2012-40035-C03. Carles Castaño received support from the Secretaria d'Universitats i Recerca del Departament d'Economia i Coneixement de la Generalitat de Catalunya through the program of Doctorats Industrials, funded by the European Union and the European Social Fund. Josu G. Alday was supported by Juan de la Cierva fellowships (IJCI-2014-21393).

Soil microclimate is a potentially important regulator of the composition of plant‐associated fungal communities in climates with significant drought periods. Here, we investigated the spatio‐temporal dynamics of soil fungal communities in a Mediterranean Pinus pinaster forest in relation to soil moisture and temperature. Fungal communities in 336 soil samples collected monthly over 1 year from 28 long‐term experimental plots were assessed by PacBio sequencing of ITS2 amplicons. Total fungal biomass was estimated by analysing ergosterol. Community changes were analysed in the context of functional traits. Soil fungal biomass was lowest during summer and late winter and highest during autumn, concurrent with a greater relative abundance of mycorrhizal species. Intra‐annual spatio‐temporal changes in community composition correlated significantly with soil moisture and temperature. Mycorrhizal fungi were less affected by summer drought than free‐living fungi. In particular, mycorrhizal species of the short‐distance exploration type increased in relative abundance under dry conditions, whereas species of the long‐distance exploration type were more abundant under wetter conditions. Our observations demonstrate a potential for compositional and functional shifts in fungal communities in response to changing climatic conditions. Free‐living fungi and mycorrhizal species with extensive mycelia may be negatively affected by increasing drought periods in Mediterranean forest ecosystems. ; This work was supported by the Spanish Ministry of Economy and Competitivity (MINECO) (grant number AGL2015‐66001‐C3). C.C. received support from the Secretaria d'Universitats i Recerca del Departament d'Economia i Coneixement de la Generalitat de Catalunya through the program of Doctorats Industrials, funded by the European Union and the European Social Fund. J.G.A. was supported by Juan de la Cierva (Grant number IJCI‐2014‐21393) and Ramon y Cajal fellowships (RYC‐2016‐20528) and J.A.B. benefits from the Serra‐Hunter Fellowship.

Despite the assumption that mushroom fruiting is dependent on climate conditions, recent changes in temperature and precipitation regimes in Mediterranean-type ecosystems have opened new questions about how climate changes impact epigeous sporocarps yields. Here, we described the epigeous fungal sporocarp yield anomalies together with temperature and precipitation fluctuations that have triggered atypical fungal production peaks during the last 20 years in different forest ecosystems of Mediterranean Spain. We used the largest standardized, spatio-temporal epigeous sporocarp collection dataset available in Mediterranean-type forest ecosystems and climates. Two clear positive yield anomalies were found in 2006 and 2014, representing more than 270% and 210% increase, respectively, in comparison with timeseries mean annual fungal yields. Late-summer-early-autumn precipitation was the most influential variable triggering these extreme mushroom production peaks in comparison with temperature. This suggests that fungal fruiting is sensitive to late-summer-early-autumn precipitation pulses, producing discrete yield pulses when conditions are optimal, which are interspersed among periods of limited water resource availability and lower yields. ; This study was partially funded by the research project AGL2012-40035-C03 (MCI Spain), by VA206U13 (Junta-CyL) and European project StarTree-Multipurpose-trees (No-311919). JGA was supported by Juan de la Cierva-fellowship (IJCI-2014-21393), SdM by the European Union's Horizon 2020 MultiFUNGtionality Marie Skłodowska-Curie (IF-EF No-655815) and JAB benefit from Serra-Hunter Fellow provided by the Generalitat of Catalunya.

Predicted changes in global climate might negatively affect the soil microbiome and associated ecosystem processes in Mediterranean forests. Forest treatments, such as forest thinning, have been suggested to mitigate climate change impacts on vegetation by reducing competition between trees, thus increasing water availability. Studies addressing the combined effects of climate and forest thinning on belowground fungal communities are still scarce, being fundamental to elaborate adaptive strategies to global warming. The aim of this study was to evaluate the tree density reduction effects on soil fungal communities and their response to inter-annual changes in weather conditions. The temporal dynamics of soil fungal communities in relation to these two drivers (i.e., forest management and weather conditions) were studied from 2009 until 2014 in a set of 12 pairs of thinned and un-thinned plots dominated by Pinus pinaster Ait. Thinning (from 30% up to 70% reduction in stand basal area) was conducted in 2009 and soil fungal community composition was studied during 4 years. Here, we used autumn precipitation and temperature to describe the impact of inter-annual weather changes. We used Pacific Biosciences sequencing of fungal ITS2 amplicons to study fungal communities in soil samples. Forest thinning did not significantly affect fungal community composition nor fungal species richness and diversity, indicating that the soil fungal community is resistant to forest thinning regardless of its intensity. However, fungal species composition changed progressively across years, both at the species level and with regards to functional guilds. These changes in community composition were partly driven by inter-annual variation in precipitation and temperature, with free-living fungi increasing in abundance under wetter conditions, and symbiotic fungi being more prominent under drier and colder conditions. The results indicate that mycorrhizal communities in Mediterranean forest ecosystems can resist forest thinning, if enough trees and functional roots from thinned trees are retained. ; This work was supported by the Spanish Ministry of Economy and Competitivity (MINECO) [grant number AGL2015-66001-C3] and by the Collaborative European project ERANET-INFORMED (PCIN-2014-050). Carles Castaño received the support of the Doctorats Industrials program, funded by the European Union and the European Social Fund. Sergio de-Miguel was supported by the European Union's Horizon 2020 MultiFUNGtionality Marie Skłodowska-Curie [Grant number IF-EF No-655815]. Josu G. Alday was supported by Ramon y Cajal fellowship [Grant number RYC-2016-20528] and José Antonio Bonet benefited from a Serra-Húnter Fellowship provided by the Generalitat of Catalunya. The authors are very grateful to the PNIN of Poblet for its considerable help with the process of installing and maintaining the experimental plots. We thank Liu Bing, Daniel Oliach, Francesc Bolaño, Jordi Margalef, Josep Miró and Jewel Yurkewich for their assistance with sampling the plots and processing the samples. The constructive comments and suggestions of two anonymous reviewers were improved substantially this manuscript.

Forest ecosystems provide a wide range of goods and services to society and host high levels of biodiversity. Nevertheless, forest ecosystem services (ES) are often quantified and assessed using simplified methodologies (e.g., proxy methods based exclusively on Land Use Land Cover maps) that introduce substantial uncertainty in the analysis by ignoring, for instance, the species composition and spatial configuration of the ecosystems studied. In this work we defined and calculated a set of 12 indicators of several ES for the forests of the highly populated region of Catalonia (North-eastern Iberian Peninsula). The indicators combined different sources of information such as forest surveys, ecological model predictions and official statistics, but also included additional land cover information. All ES indicators were aggregated at the municipality level to compare their values and distribution patterns. We assessed spatial trade-offs and synergies among ES, as well as their relationships with a set of socioeconomic, climatic and biodiversity variables using correlation analyses and mixed-effects models. The results suggest a clustering of provisioning and regulating ES in mountainous zones towards the North of the study area. These two types of services showed a high degree of spatial similarity and presented high positive correlations. In contrast, cultural ES showed a more scattered pattern, which included lower elevation areas in the South of the study region. Climatic conditions were the main determinants of the spatial variability in the supply of the different ES, with most indicators being positively associated with precipitation and negatively associated with temperature. In addition, biodiversity (particularly woody species richness) showed positive relations with most of these ES, while socioeconomic variables (such as population density and the percentage employment in agriculture) showed negative associations with most of them. The combination of information from different data sources (including primary data) allowed for a detailed analysis of forest ES, likely removing some of the problems derived from approaches based only on proxy methods. In addition, the use of municipalities as study unit makes results directly relevant to management and planning strategies operating at this scale (e.g., forest management and planning). ; Funding was obtained from the Catalan Office for Climate Change (OCCC) through project ForESMap, from EU FORESTERRA program (INFORMED project) and from the Spanish government (CGL2013-46808-R, AGL2015-66001-C3-1-R and CGL2014-59742). We also thank the ECOMETAS (CGL2014-53840-REDT) network for support. This study also received funding from the European Union's Horizon 2020 research and innovation programme within the framework of the MultiFUNGtionality Marie Skłodowska-Curie Individual Fellowship (IF-EF) under grant agreement No655815 and from the Generalitat de Catalunya (Serra-Hunter Fellow). JVRD was supported by the Government of Asturias and the FP7-Marie Curie-COFUND program of the European Commission (Grant 'Clarín' ACA17-02).