Positive feedbacks and alternative stable states in forest leaf types.

The emergence of alternative stable states in forest systems has significant implications for the functioning and structure of the terrestrial biosphere, yet empirical evidence remains scarce. Here, we combine global forest biodiversity observations and simulations to test for alternative stable states in the presence of evergreen and deciduous forest types. We reveal a bimodal distribution of forest leaf types across temperate regions of the Northern Hemisphere that cannot be explained by the environment alone, suggesting signatures of alternative forest states. Moreover, we empirically demonstrate the existence of positive feedbacks in tree growth, recruitment and mortality, with trees having 4-43% higher growth rates, 14-17% higher survival rates and 4-7 times higher recruitment rates when they are surrounded by trees of their own leaf type. Simulations show that the observed positive feedbacks are necessary and sufficient to generate alternative forest states, which also lead to dependency on history (hysteresis) during ecosystem transition from evergreen to deciduous forests and vice versa. We identify hotspots of bistable forest types in evergreen-deciduous ecotones, which are likely driven by soil-related positive feedbacks. These findings are integral to predicting the distribution of forest biomes, and aid to our understanding of biodiversity, carbon turnover, and terrestrial climate feedbacks.

How do trees respond to species mixing in experimental compared to observational studies?

For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species-specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.

Erratum: Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe.

This corrects the article DOI: 10.1038/srep32233.

Climate- and successional-related changes in functional composition of European forests are strongly driven by tree mortality.

Intense droughts combined with increased temperatures are one of the major threats to forest persistence in the 21st century. Despite the direct impact of climate change on forest growth and shifts in species abundance, the effect of altered demography on changes in the composition of functional traits is not well known. We sought to (1) quantify the recent changes in functional composition of European forests; (2) identify the relative importance of climate change, mean climate and forest development for changes in functional composition; and (3) analyse the roles of tree mortality and growth underlying any functional changes in different forest types. We quantified changes in functional composition from the 1980s to the 2000s across Europe by two dimensions of functional trait variation: the first dimension was mainly related to changes in leaf mass per area and wood density (partially related to the trait differences between angiosperms and gymnosperms), and the second dimension was related to changes in maximum tree height. Our results indicate that climate change and mean climatic effects strongly interacted with forest development and it was not possible to completely disentangle their effects. Where recent climate change was not too extreme, the patterns of functional change generally followed the expected patterns under secondary succession (e.g. towards late-successional short-statured hardwoods in Mediterranean forests and taller gymnosperms in boreal forests) and latitudinal gradients (e.g. larger proportion of gymnosperm-like strategies at low water availability in forests formerly dominated by broad-leaved deciduous species). Recent climate change generally favoured the dominance of angiosperm-like related traits under increased temperature and intense droughts. Our results show functional composition changes over relatively short time scales in European forests. These changes are largely determined by tree mortality, which should be further investigated and modelled to adequately predict the impacts of climate change on forest function.

Complementarity effects on tree growth are contingent on tree size and climatic conditions across Europe.

Neglecting tree size and stand structure dynamics might bias the interpretation of the diversity-productivity relationship in forests. Here we show evidence that complementarity is contingent on tree size across large-scale climatic gradients in Europe. We compiled growth data of the 14 most dominant tree species in 32,628 permanent plots covering boreal, temperate and Mediterranean forest biomes. Niche complementarity is expected to result in significant growth increments of trees surrounded by a larger proportion of functionally dissimilar neighbours. Functional dissimilarity at the tree level was assessed using four functional types: i.e. broad-leaved deciduous, broad-leaved evergreen, needle-leaved deciduous and needle-leaved evergreen. Using Linear Mixed Models we show that, complementarity effects depend on tree size along an energy availability gradient across Europe. Specifically: (i) complementarity effects at low and intermediate positions of the gradient (coldest-temperate areas) were stronger for small than for large trees; (ii) in contrast, at the upper end of the gradient (warmer regions), complementarity is more widespread in larger than smaller trees, which in turn showed negative growth responses to increased functional dissimilarity. Our findings suggest that the outcome of species mixing on stand productivity might critically depend on individual size distribution structure along gradients of environmental variation.

Comparison of calculation methods for estimating annual carbon stock change in German forests under forest management in the German greenhouse gas inventory.

BACKGROUND: The German greenhouse gas inventory in the land use change sector strongly depends on national forest inventory data. As these data were collected periodically 1987, 2002, 2008 and 2012, the time series on emissions show several "jumps" due to biomass stock change, especially between 2001 and 2002 and between 2007 and 2008 while within the periods the emissions seem to be constant due to the application of periodical average emission factors. This does not reflect inter-annual variability in the time series, which would be assumed as the drivers for the carbon stock changes fluctuate between the years. Therefore additional data, which is available on annual basis, should be introduced into the calculations of the emissions inventories in order to get more plausible time series. RESULTS: This article explores the possibility of introducing an annual rather than periodical approach to calculating emission factors with the given data and thus smoothing the trajectory of time series for emissions from forest biomass. Two approaches are introduced to estimate annual changes derived from periodic data: the so-called logging factor method and the growth factor method. The logging factor method incorporates annual logging data to project annual values from periodic values. This is less complex to implement than the growth factor method, which additionally adds growth data into the calculations. CONCLUSION: Calculation of the input variables is based on sound statistical methodologies and periodically collected data that cannot be altered. Thus a discontinuous trajectory of the emissions over time remains, even after the adjustments. It is intended to adopt this approach in the German greenhouse gas reporting in order to meet the request for annually adjusted values.

Converting probabilistic tree species range shift projections into meaningful classes for management.

The paper deals with the management problem how to decide on tree species suitability under changing environmental conditions. It presents an algorithm that classifies the output of a range shift model for major tree species in Europe into multiple classes that can be linked to qualities characterizing the ecological niche of the species. The classes: i) Core distribution area, ii) Extended distribution area, iii) Occasional occurrence area, and iv) No occurrence area are first theoretically developed and then statistically described. The classes are interpreted from an ecological point of view using criteria like population structure, competitive strength, site spectrum and vulnerability to biotic hazards. The functioning of the algorithm is demonstrated using the example of a generalized linear model that was fitted to a pan-European dataset of presence/absence of major tree species with downscaled climate data from a General Circulation Model (GCM). Applications of the algorithm to tree species suitability classification on a European and regional level are shown. The thresholds that are used by the algorithm are precision-based and include Cohen's Kappa. A validation of the algorithm using an independent dataset of the German National Forest Inventory shows good accordance of the statistically derived classes with ecological traits for Norway spruce, while the differentiation especially between core and extended distribution for European beech that is in the centre of its natural range in this area is less accurate. We hypothesize that for species in the core of their range regional factors like forest history superimpose climatic factors. Problems of uncertainty issued from potentially applying a multitude of modelling approaches and/or climate realizations within the range shift model are discussed and a way to deal with the uncertainty by revealing the underlying attitude towards risk of the decision maker is proposed.