Mixing of complementary tree species may increase stand productivity, mitigate the effects of drought and other risks, and pave the way to forest production systems which may be more resource-use efficient and stable in the face of climate change. However, systematic empirical studies on mixing effects are still missing for many commercially important and widespread species combinations. Here we studied the growth of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) in mixed versus pure stands on 32 triplets located along a productivity gradient through Europe, reaching from Sweden to Bulgaria and from Spain to the Ukraine. Stand inventory and taking increment cores on the mainly 60-80 year-old trees and 0.02-1.55 ha sized, fully stocked plots provided insight how species mixing modifies the structure, dynamics and productivity compared with neighbouring pure stands. In mixture standing volume (?12 %), stand density (?20 %), basal area growth (?12 %), and stand volume growth (?8 %) were higher Communicated by Peter Biber. than the weighted mean of the neighbouring pure stands. Scots pine and European beech contributed rather equally to the overyielding and overdensity. In mixed stands mean diameter (?20 %) and height (?6 %) of Scots pine was ahead, while both diameter and height growth of European beech were behind (-8 %). The overyielding and overdensity were independent of the site index, the stand growth and yield, and climatic variables despite the wide variation in precipitation (520-1175 mm year -1 ), mean annual temperature (6-10.5°C), and the drought index by de Martonne (28-61 mm°C -1 ) on the sites. Therefore, this species combination is potentially useful for increasing productivity across a wide range of site and climatic conditions. Given the significant overyielding of stand basal area growth but the absence of any relationship with site index and climatic variables, we hypothesize that the overyielding and overdensity results from several different types of interactions (light-, water-, and nutrient-related) that are all important in different circumstances. We discuss the relevance of the results for ecological theory and for the ongoing silvicultural transition from pure to mixed stands and their adaptation to climate change. Electronic supplementary material
The area of forest plantations is increasing worldwide helping to meet timber demand and protect natural forests. However, with global change, monospecific plantations are increasingly vulnerable to abiotic and biotic disturbances. As an adaption measure we need to move to plantations that are more diverse in genotypes, species, and structure, with a design underpinned by science. TreeDivNet, a global network of tree diversity experiments, responds to this need by assessing the advantages and disadvantages of mixed species plantations. The network currently consists of 18 experiments, distributed over 36 sites and five ecoregions. With plantations 1–15 years old, TreeDivNet can already provide relevant data for forest policy and management. In this paper, we highlight some early results on the carbon sequestration and pest resistance potential of more diverse plantations. Finally, suggestions are made for new, innovative experiments in understudied regions to complement the existing network.
Despite considerable research demonstrating that biodiversity increases productivity in forests and regulates herbivory and pathogen damage, there remain gaps in our understanding of the shape, magnitude, and generality of these biodiversity-ecosystem functioning (BEF) relationships. Here, we review findings from TreeDivNet, a global network of 25 tree diversity experiments, on relationships between levels of biodiversity and (a) tree growth and survival and (b) damage to trees from pests and pathogens. Tree diversity often improved the survival and above-and belowground growth of young trees. The mechanistic bases of the diversity effects on tree growth and survival include both selection effects (i.e., an increasing impact of particular species in more species-rich communities) and complementary effects (e.g. related to resource differentiation and facilitation). Plant traits and abiotic stressors may mediate these relationships. Studies of the responses of invertebrate and vertebrate herbivory and pathogen damage have demonstrated that trees in more diverse experimental plots may experience more, less, or similar damage compared to conspecific trees in less diverse plots. Documented mechanisms producing these patterns include changes in concentration, frequency, and apparency of hosts; herbivore and pathogen diet breadth; the spatial scale of interactions; and herbivore and pathogen regulation by natural enemies. Our review of findings from TreeDivNet indicates that tree diversity experiments are extending BEF research across systems and scales, complementing previous BEF work in grasslands by providing opportunities to use remote sensing and spectral approaches to study BEF dynamics, integrate belowground and aboveground approaches, and trace the consequences of tree physiology for ecosystem functioning. This extension of BEF research into tree-dominated systems is improving ecologists' capacity to understand the mechanistic bases behind BEF relationships. Tree diversity experiments also present opportunities for novel research. Since experimental tree diversity plantations enable measurements at tree, neighbourhood and plot level, they allow for explicit consideration of temporal and spatial scales in BEF dynamics. Presently, most TreeDivNet experiments have run for less than ten years. Given the longevity of trees, exciting results on BEF relationships are expected in the future.
TreeDivNet is the largest network of biodiversity experiments worldwide, but needs to expand. We encourage colleagues to establish new experiments on the relation between tree species diversity and forest ecosystem functioning, and to make use of the platform for collaborative research.
We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900-2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4% increase in plant height with every degree northward shift (almost a two-fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three-factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring-flowering Anemone was affected more by temperature, whereas early-summer flowering Milium was affected more by local and other latitude-related factors. Finally, previously reported cross-species correlations between LHS traits and latitude were generally unsupported by our within-species approach
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