Species mixing effects on forest productivity in the Netherlands

Many monoculture forests (dominated by a single tree species) have been converted to mixed-species forests (dominated by more than one tree species) in Europe over the last decades. The main reason for this conversion was to increase productivity, including timber production, and enhance other ecosystem services, such as conservation of biodiversity and other nature values. In addition, it has been suggested that mixed-species forests are more resistant, resilient and stable to disturbances. In line with the niche complementarity hypothesis, inter-specific differences in crown architecture, leaf phenology, shade tolerance and root distribution may allow tree species to partition resources in mixed forests. Such mechanisms may lead to a higher productivity of mixed forests versus monoculture forests, a phenomenon often referred to as overyielding. Interestingly, the stress-gradient hypothesis and the resource-ratio hypothesis suggests that such inter-specific interactions vary along a soil fertility gradient, but in different ways. The stress-gradient hypothesis emphasizes that more efficient partitioning increases overyielding at low fertility soils, whereas the resource ratio hypothesis considers that the denser packing of crowns on fertile soils allows for partitioning of light and overyielding on high fertility soils. Several studies have been carried out about species mixing effects on forest productivity, but so far their findings are ambiguous. Probably, this ambiguity comes from the sites that they studied, which differ in species, age, management history, and/or environmental conditions. This thesis analyses the mixing effect on productivity in relation to the combination of species, stand age and soil fertility, and discusses possible consequences of forest management, for five two-species mixtures in the Netherlands: Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco)–beech (Fagus sylvatica L.), pine (Pinus sylvestris L.)–oak (Quercus robur L.), oak–beech, oak–birch (Betula pendula Roth) and pine–birch. These mixtures and their corresponding monoculture stands were studied using long-term permanent forest plots over multiple decades, but also using two inventories (around 2003 and 2013) across the entire Netherlands. These forest plots data were used together with empirical models at total stand level (chapter 2), species level (chapter 3) and tree level (chapter 4) to evaluate the mixing effect on forest productivity. In chapter 2, four two-species mixtures and their corresponding monospecific stands were compared for productivity (volume stem wood in m3 ha-1 year-1). It was explored whether mixing species differing in leaf phenology and shade tolerance would lead to overyielding of mixed forest stands, and whether overyielding changes with stand development. In line with the niche complementarity hypothesis, the two evergreen–deciduous species mixtures (Douglas-fir–beech and pine–oak) showed overyielding whereas deciduous–deciduous species mixtures (oak–beech and oak–birch) did not. The overyielding was strongest for the Douglas-fir–beech mixture than the pine–oak mixture, which can be attributed to the greater difference in shade tolerance in the former mixture. Overyielding did not significantly change with stand development. It is argued that the regular thinning maintained the ability of species to partition resources, i.e. the complementary resource use in those mixed stands over all stand ages. In chapter 3, it was analysed which of the two species in these four mixtures contributed to overyielding, and whether this overyielding changed along a soil fertility gradient. It was discovered that both the fast-growing and the slow-growing species could contribute to overyielding. Yet, it was mainly the fast-growing Douglas-fir that contributed to higher productivity in the Douglas-fir–beech mixtures, and the slow-growing oak that did so in the pine–oak mixtures. For both mixtures, the greatest relative productivity gain was achieved by mixtures on the poorer soils. At first sight, these results seem in line with the stress-gradient hypothesis and not the resource-ratio hypothesis. Yet, it was argued that not only complementary use of soil resources, but also use of light, may contribute to the higher productivity of mixed stands on the poorer soils. In chapter 4, it was assessed how the growth of individual trees in mixtures was influenced by inter- and intra-specific competition, and whether this competition was mainly size-symmetric for soil resources or size-asymmetric for light on soils differing in fertility. This chapter focussed on three mixtures, i.e. oak–birch, pine–oak and pine–birch, which were available at sufficient numbers in the Dutch national forest inventory data. It was concluded that intra-specific competition was not necessarily stronger than inter-specific competition and this competitive reduction was less seen at lower soil fertility and dependent on species mixtures, which is not in line with the stress-gradient hypothesis. Moreover, size-asymmetric competition for light was more associated with tree basal area growth than size-symmetric competition for soil resources, suggesting that light is the most limiting resource. Competition for light was generally much stronger at high fertility soils, supporting the resource-ratio hypothesis. These results suggest that light is the most limiting resource for tree basal area growth and that reduced competition for light can be explained to some degree by complementarity in light use to increase tree growth in mixed forests. This thesis thus described the productivity patterns when mixing tree species and explored possible mechanisms of higher productivity in mixed stands compared with monoculture stands in the Netherlands. Complementary use of aboveground and belowground resources probably contributes to the higher productivity in mixed stands, but other factors including pathogens, nutrient cycling and litter decomposition were not addressed but cannot be excluded. Overyielding in Douglas-fir–beech and pine–oak mixtures was maintained over time, probably owing to the intensive thinning in Dutch forests. The results shed new light on the stress-gradient and resource-ratio hypotheses. For mixtures in Dutch forest, the greatest productivity gain in Douglas-fir–beech and pine–oak mixtures was achieved on the poorer soils, and it was argued that this is at least partially driven by complementary use of light, while the role of complementarity in use of soil resources is more obscure. Overall, this thesis suggest a substantial potential of species mixing for increasing productivity, which may run in parallel with enhancing other ecosystem services such as conservation of diversity and other nature values. Yet, more experimental studies on productivity in mixed stands are required to better unravel alternative mechanisms. Such understanding is required to manage the forests effectively in a century of unpreceded human driven changes in environmental conditions.</p

Evenness mediates the global relationship between forest productivity and richness

1. Biodiversity is an important component of natural ecosystems, with higher species richness often correlating with an increase in ecosystem productivity. Yet, this relationship varies substantially across environments, typically becoming less pronounced at high levels of species richness. However, species richness alone cannot reflect all important properties of a community, including community evenness, which may mediate the relationship between biodiversity and productivity. If the evenness of a community correlates negatively with richness across forests globally, then a greater number of species may not always increase overall diversity and productivity of the system. Theoretical work and local empirical studies have shown that the effect of evenness on ecosystem functioning may be especially strong at high richness levels, yet the consistency of this remains untested at a global scale. 2. Here, we used a dataset of forests from across the globe, which includes composition, biomass accumulation and net primary productivity, to explore whether productivity correlates with community evenness and richness in a way that evenness appears to buffer the effect of richness. Specifically, we evaluated whether low levels of evenness in speciose communities correlate with the attenuation of the richness–productivity relationship. 3. We found that tree species richness and evenness are negatively correlated across forests globally, with highly speciose forests typically comprising a few dominant and many rare species. Furthermore, we found that the correlation between diversity and productivity changes with evenness: at low richness, uneven communities are more productive, while at high richness, even communities are more productive. 4. Synthesis. Collectively, these results demonstrate that evenness is an integral component of the relationship between biodiversity and productivity, and that the attenuating effect of richness on forest productivity might be partly explained by low evenness in speciose communities. Productivity generally increases with species richness, until reduced evenness limits the overall increases in community diversity. Our research suggests that evenness is a fundamental component of biodiversity–ecosystem function relationships, and is of critical importance for guiding conservation and sustainable ecosystem management decisions

Native diversity buffers against severity of non-native tree invasions

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species$^{1,2}$. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies$^{3,4}$. Here, leveraging global tree databases$^{5,6,7}$, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions

Native diversity buffers against severity of non-native tree invasions

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions

Native diversity buffers against severity of non-native tree invasions.

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions

The global biogeography of tree leaf form and habit

Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17-34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cycling

The global biogeography of tree leaf form and habit.

Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17-34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cycling

The global biogeography of tree leaf form and habit

Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17–34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cycling

The global biogeography of tree leaf form and habit

Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17-34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cyclin