Jordi Vayreda

Positive biodiversity-productivity relationship predominant in global forests.

Global biodiversity and productivity The relationship between biodiversity and ecosystem productivity has been explored in detail in herbaceous vegetation, but patterns in forests are far less well understood. Liang et al. have amassed a global forest data set from >770,000 sample plots in 44 countries. A positive and consistent relationship can be discerned between tree diversity and ecosystem productivity at landscape, country, and ecoregion scales. On average, a 10% loss in biodiversity leads to a 3% loss in productivity. This means that the economic value of maintaining biodiversity for the sake of global forest productivity is more than fivefold greater than global conservation costs. Science, this issue p. 196 Global forest inventory records suggest that biodiversity loss would result in a decline in forest productivity worldwide. INTRODUCTION The biodiversity-productivity relationship (BPR; the effect of biodiversity on ecosystem productivity) is foundational to our understanding of the global extinction crisis and its impacts on the functioning of natural ecosystems. The BPR has been a prominent research topic within ecology in recent decades, but it is only recently that we have begun to develop a global perspective. RATIONALE Forests are the most important global repositories of terrestrial biodiversity, but deforestation, forest degradation, climate change, and other factors are threatening approximately one half of tree species worldwide. Although there have been substantial efforts to strengthen the preservation and sustainable use of forest biodiversity throughout the globe, the consequences of this diversity loss pose a major uncertainty for ongoing international forest management and conservation efforts. The forest BPR represents a critical missing link for accurate valuation of global biodiversity and successful integration of biological conservation and socioeconomic development. Until now, there have been limited tree-based diversity experiments, and the forest BPR has only been explored within regional-scale observational studies. Thus, the strength and spatial variability of this relationship remains unexplored at a global scale. RESULTS We explored the effect of tree species richness on tree volume productivity at the global scale using repeated forest inventories from 777,126 permanent sample plots in 44 countries containing more than 30 million trees from 8737 species spanning most of the global terrestrial biomes. Our findings reveal a consistent positive concave-down effect of biodiversity on forest productivity across the world, showing that a continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The BPR shows considerable geospatial variation across the world. The same percentage of biodiversity loss would lead to a greater relative (that is, percentage) productivity decline in the boreal forests of North America, Northeastern Europe, Central Siberia, East Asia, and scattered regions of South-central Africa and South-central Asia. In the Amazon, West and Southeastern Africa, Southern China, Myanmar, Nepal, and the Malay Archipelago, however, the same percentage of biodiversity loss would lead to greater absolute productivity decline. CONCLUSION Our findings highlight the negative effect of biodiversity loss on forest productivity and the potential benefits from the transition of monocultures to mixed-species stands in forestry practices. The BPR we discover across forest ecosystems worldwide corresponds well with recent theoretical advances, as well as with experimental and observational studies on forest and nonforest ecosystems. On the basis of this relationship, the ongoing species loss in forest ecosystems worldwide could substantially reduce forest productivity and thereby forest carbon absorption rate to compromise the global forest carbon sink. We further estimate that the economic value of biodiversity in maintaining commercial forest productivity alone is

Disentangling biodiversity and climatic determinants of wood production.

166 billion to

Fire regenerative syndromes of forest woody species across fire and climatic gradients.

490 billion per year. Although representing only a small percentage of the total value of biodiversity, this value is two to six times as much as it would cost to effectively implement conservation globally. These results highlight the necessity to reassess biodiversity valuation and the potential benefits of integrating and promoting biological conservation in forest resource management and forestry practices worldwide. Global effect of tree species diversity on forest productivity. Ground-sourced data from 777,126 global forest biodiversity permanent sample plots (dark blue dots, left), which cover a substantial portion of the global forest extent (white), reveal a consistent positive and concave-down biodiversity-productivity relationship across forests worldwide (red line with pink bands representing 95% confidence interval, right). The biodiversity-productivity relationship (BPR) is foundational to our understanding of the global extinction crisis and its impacts on ecosystem functioning. Understanding BPR is critical for the accurate valuation and effective conservation of biodiversity. Using ground-sourced data from 777,126 permanent plots, spanning 44 countries and most terrestrial biomes, we reveal a globally consistent positive concave-down BPR, showing that continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The value of biodiversity in maintaining commercial forest productivity alone—US

Structural and climatic determinants of demographic rates of Scots pine forests across the Iberian Peninsula

166 billion to 490 billion per year according to our estimation—is more than twice what it would cost to implement effective global conservation. This highlights the need for a worldwide reassessment of biodiversity values, forest management strategies, and conservation priorities.

Determinants of distribution of six Pinus species in Catalonia, Spain.

Background Despite empirical support for an increase in ecosystem productivity with species diversity in synthetic systems, there is ample evidence that this relationship is dependent on environmental characteristics, especially in structurally more complex natural systems. Empirical support for this relationship in forests is urgently needed, as these ecosystems play an important role in carbon sequestration. Methodology/Principal Findings We tested whether tree wood production is positively related to tree species richness while controlling for climatic factors, by analyzing 55265 forest inventory plots in 11 forest types across five European countries. On average, wood production was 24% higher in mixed than in monospecific forests. Taken alone, wood production was enhanced with increasing tree species richness in almost all forest types. In some forests, wood production was also greater with increasing numbers of tree types. Structural Equation Modeling indicated that the increase in wood production with tree species richness was largely mediated by a positive association between stand basal area and tree species richness. Mean annual temperature and mean annual precipitation affected wood production and species richness directly. However, the direction and magnitude of the influence of climatic variables on wood production and species richness was not consistent, and vary dependent on forest type. Conclusions Our analysis is the first to find a local scale positive relationship between tree species richness and tree wood production occurring across a continent. Our results strongly support incorporating the role of biodiversity in management and policy plans for forest carbon sequestration.

WOODY PLANT RICHNESS AND NDVI RESPONSE TO DROUGHT EVENTS IN CATALONIAN (NORTHEASTERN SPAIN) FORESTS

Community resilience after fire is determined by species’ ability to regenerate through two main mechanisms growth of new sprouts (resprouter species) and germination from surviving seed banks or from seeds arriving from neighbouring populations (seeder species). Both the mechanisms are present in Mediterranean communities. The occurrence of both the types in a community depends on fire history and the bio-geographical history determining the available species pool. Regenerative traits also covary with other functional attributes associated with resource acquisition and stress tolerance. As post-fire regenerative responses can be related to various ecological factors other than fire, we tested the hypothesis of a different proportional representation of post-fire regenerative syndromes in forest woody species along a climatic gradient in Catalonia (NE Spain) ranging from Mediterranean to temperate-boreal climates. Specifically, we expected seeder species to become less common with colder and moister conditions while resprouters would not be so influenced by the climatic gradient. We also tested the hypothesis of change in the relative abundance of regenerative syndromes in relation to recent fire history. We analysed a large database obtained from extensive forestry surveys and remote sensing fire records. After correction for spatial autocorrelation, we found an increase in the proportion of seeder species under more Mediterranean conditions and a decrease in fire-sensitive species (with no efficient mechanisms of post-fire recovery) in moister conditions. Resprouter species were similarly present across the whole gradient. A similar pattern was observed after excluding recently burnt plots. Therefore, post-fire regenerative syndromes segregate along the climatic gradient. Recent fires reduced the occurrence of fire-sensitive species and increased the proportion of seeder species. No significant effect was observed on resprouter species. Fire has a sorting effect, shaping the occurrence of species with different regenerative traits. Overall, fire seems to explain better the variability of the proportion of fire-sensitive species and climate the variability of seeder species. In addition, other factors (forestry practices and the covariation between regenerative and functional attributes) are likely to contribute to the regional pattern of regenerative syndromes.

Biodiversity correlates with regional patterns of forest litter pools

The demographic rates of tree species typically show large spatial variation across their range. Understanding the environmental factors underlying this variation is a key topic in forest ecology, with far-reaching management implications. Scots pine (Pinus sylvestris L.) covers large areas of the Northern Hemisphere, the Iberian Peninsula being its southwestern distribution limit. In recent decades, an increase in severe droughts and a densification of forests as a result of changes in forest uses have occurred in this region. Our aim was to use climate and stand structure data to explain mortality and growth patterns of Scots pine forests across the Iberian Peninsula. We used data from 2392 plots dominated by Scots pine, sampled for the National Forest Inventory of Spain. Plots were sampled from 1986 to 1996 (IFN2) and were resampled from 1997 to 2007 (IFN3), allowing for the calculation of growth and mortality rates. We fitted linear models to assess the response of growth and mortality rates to the spatial variability of climate, climatic anomalies, and forest structure. Over the period of approximately 10 years between the IFN2 and IFN3, the amount of standing dead trees increased 11-fold. Higher mortality rates were related to dryness, and growth was reduced with increasing dryness and temperature, but results also suggested that effects of climatic stressors were not restricted to dry sites only. Forest structure was strongly related to demographic rates, suggesting that stand development and competition are the main factors associated with demography. In the case of mortality, forest structure interacted with climate, suggesting that competition for water resources induces tree mortality in dry sites. A slight negative relationship was found between mortality and growth, indicating that both rates are likely to be affected by the same stress factors. Additionally, regeneration tended to be lower in plots with higher mortality. Taken together, our results suggest a large-scale self-thinning related to the recent densification of Scots pine forests. This process appears to be enhanced by dry conditions and may lead to a mismatch in forest turnover. Forest management may be an essential adaptive tool under the drier conditions predicted by most climate models.

SMOS and climate data applicability for analyzing forest decline and forest fires

This study explores the determinants of distribu- tion, abundance and regeneration of six Pinus species (P. halepensis, P. nigra, P. pinaster, P. pinea, P. sylvestris and P. uncinata) that occur naturally in Catalonia, northeastern Spain. The aim of this study was to generate accurate predictions of the distribution of each species using simple and readily available environmental variables. We used recursive parti- tioning and GIS analyses to relate the data base of 10 600 field plots from the Forestry Inventory of Catalonia with abiotic and biotic characteristics of each plot. We present general patterns of distribution, dominance and regeneration for the six species and then focus on P. halepensis, the most abundant pine species in the western Mediterranean Basin. For all six spe- cies, the models correctly classified more than 80% of the distribution using abiotic factors, mainly altitude and rainfall variability. Biotic factors such as the basal area of other pine species were necessary to accurately predict patterns of pine species dominance. Biotic factors, especially the basal area of evergreen species (mainly Quercus ilex), were of overriding importance when predicting patterns of seedling occurrence. Potentially important factors such as land use and fire history were of little significance for predicting distribution at the scale of our study. Our models failed to predict accurately which species (and in which numbers) co-occur with P. halepensis. Factors not included in this study, such as stand age, disturbance (cutting, clearing) and other human-induced factors, are probably the main determinants of co-existence patterns.

Climate Change Could Negate Positive Tree Diversity Effects on Forest Productivity: A Study Across Five Climate Types in Spain and Canada

The role of species diversity on ecosystem resistance in the face of strong environmental fluctuations has been addressed from both theoretical and experimental viewpoints to reveal a variety of positive and negative relationships. Here we explore empirically the relationship between the richness of forest woody species and canopy resistance to extreme drought episodes. We compare richness data from an extensive forest inventory to a temporal series of satellite imagery that estimated drought impact on forest canopy as NDVI (normalized difference vegetation index) anomalies of the dry summer in 2003 in relation to records of previous years. We considered five different types of forests that are representative of the main climatic and altitudinal gradients of the region, ranging from lowland Mediterranean to mountain boreal-temperate climates. The observed relationship differed among forest types and interacted with the climate, summarised by the Thorntwaite index. In Mediterranean Pinus halepensis forests, NDVI decreased during the drought. This decrease was stronger in forests with lower richness. In Mediterranean evergreen forests of Quercus ilex, drought did not result in an overall NDVI loss, but lower NDVI values were observed in drier localities with lower richness, and in more moist localities with higher number of species. In mountain Pinus sylvestris forests NDVI decreased, mostly due to the drought impact on drier localities, while no relation to species richness was observed. In moist Fagus sylvatica forests, NDVI only decreased in plots with high richness. No effect of drought was observed in the high mountain Pinus uncinata forests. Our results show that a shift on the diversity-stability relationship appears across the regional, climatic gradient. A positive relationship appears in drier localities, supporting a null model where the probability of finding a species able to cope with drier conditions increases with the number of species. However, in more moist localities we hypothesize that the proportion of drought-sensitive species would increase in richer localities, due to a higher likelihood of co-occurrence of species that share moist climatic requirements. The study points to the convenience of considering the causes of disturbance in relation to current environmental gradients and historical environmental constraints on the community.

The spatial level of analysis affects the patterns of forest ecosystem services supply and their relationships

We compared litter pools of more than 1,000 forests differing in tree species’ diversity over a large scale in Catalonia (NE Spain). Monospecific forests always had smaller litter pools than mixed (from 2 to 5 tree species) forests. Whether there was a positive effect beyond two species mixtures depended on the species and functional identity of the dominant tree species. In sclerophyllous forests the positive effect of diversity was a step-function from one to more species. However, in conifers, litter pools increased constantly with tree diversity. The identity of the dominant tree species and functional type had also a significant effect on litter pools. For instance, forests dominated by sclerophyllous tree species had larger litter pools than forests dominated by deciduous and conifer tree species. When other forest structure parameters (i.e. tree basal area, wood production, successional stage, shrub cover and leaf area index) and environmental factors (i.e. mean annual temperature, mean annual precipitation, annual evapotranspiration and hillside position) where included in the analysis only leaf area index, basal area, wood production and mean temperature influenced litter pools positively. Our analysis emphasizes that at the regional scale, the litter compartment can be as influenced by biodiversity components as by other forest structure and climate components. In mixed forests, species and functional identity of the trees determine whether litter pools increase with tree diversity.