A new approach to derive productivity of tropical forests using radar remote sensing measurements.

Deriving gross & net primary productivity (GPP & NPP) and carbon turnover time of forests from remote sensing remains challenging. This study presents a novel approach to estimate forest productivity by combining radar remote sensing measurements, machine learning and an individual-based forest model. In this study, we analyse the role of different spatial resolutions on predictions in the context of the Radar BIOMASS mission (by ESA). In our analysis, we use the forest gap model FORMIND in combination with a boosted regression tree (BRT) to explore how spatial biomass distributions can be used to predict GPP, NPP and carbon turnover time (τ) at different resolutions. We simulate different spatial biomass resolutions (4 ha, 1 ha and 0.04 ha) in combination with different vertical resolutions (20, 10 and 2 m). Additionally, we analysed the robustness of this approach and applied it to disturbed and mature forests. Disturbed forests have a strong influence on the predictions which leads to high correlations (R2 > 0.8) at the spatial scale of 4 ha and 1 ha. Increased vertical resolution leads generally to better predictions for productivity (GPP & NPP). Increasing spatial resolution leads to better predictions for mature forests and lower correlations for disturbed forests. Our results emphasize the value of the forthcoming BIOMASS satellite mission and highlight the potential of deriving estimates for forest productivity from information on forest structure. If applied to more and larger areas, the approach might ultimately contribute to a better understanding of forest ecosystems.

Accelerated forest fragmentation leads to critical increase in tropical forest edge area.

Large areas of tropical forests have been lost through deforestation, resulting in fragmented forest landscapes. However, the dynamics of forest fragmentation are still unknown, especially the critical forest edge areas, which are sources of carbon emissions due to increased tree mortality. We analyzed the changes in forest fragmentation for the entire tropics using high-resolution forest cover maps. We found that forest edge area increased from 27 to 31% of the total forest area in just 10 years, with the largest increase in Africa. The number of forest fragments increased by 20 million with consequences for connectivity of tropical landscapes. Simulations suggest that ongoing deforestation will further accelerate forest fragmentation. By 2100, 50% of tropical forest area will be at the forest edge, causing additional carbon emissions of up to 500 million MT carbon per year. Thus, efforts to limit fragmentation in the world's tropical forests are important for climate change mitigation.

Tackling unresolved questions in forest ecology: The past and future role of simulation models.

Understanding the processes that shape forest functioning, structure, and diversity remains challenging, although data on forest systems are being collected at a rapid pace and across scales. Forest models have a long history in bridging data with ecological knowledge and can simulate forest dynamics over spatio-temporal scales unreachable by most empirical investigations.We describe the development that different forest modelling communities have followed to underpin the leverage that simulation models offer for advancing our understanding of forest ecosystems.Using three widely applied but contrasting approaches - species distribution models, individual-based forest models, and dynamic global vegetation models - as examples, we show how scientific and technical advances have led models to transgress their initial objectives and limitations. We provide an overview of recent model applications on current important ecological topics and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.Synthesis. This overview shows that forest models, due to their complementarity and mutual enrichment, represent an invaluable toolkit to address a wide range of fundamental and applied ecological questions, hence fostering a deeper understanding of forest dynamics in the context of global change.

Consequences of spatial patterns for coexistence in species-rich plant communities.

Ecology cannot yet fully explain why so many tree species coexist in natural communities such as tropical forests. A major difficulty is linking individual-level processes to community dynamics. We propose a combination of tree spatial data, spatial statistics and dynamical theory to reveal the relationship between spatial patterns and population-level interaction coefficients and their consequences for multispecies dynamics and coexistence. Here we show that the emerging population-level interaction coefficients have, for a broad range of circumstances, a simpler structure than their individual-level counterparts, which allows for an analytical treatment of equilibrium and stability conditions. Mechanisms such as animal seed dispersal, which result in clustering of recruits that is decoupled from parent locations, lead to a rare-species advantage and coexistence of otherwise neutral competitors. Linking spatial statistics with theories of community dynamics offers new avenues for explaining species coexistence and calls for rethinking community ecology through a spatial lens.

General statistical scaling laws for stability in ecological systems.

Ecological stability refers to a family of concepts used to describe how systems of interacting species vary through time and respond to disturbances. Because observed ecological stability depends on sampling scales and environmental context, it is notoriously difficult to compare measurements across sites and systems. Here, we apply stochastic dynamical systems theory to derive general statistical scaling relationships across time, space, and ecological level of organisation for three fundamental stability aspects: resilience, resistance, and invariance. These relationships can be calibrated using random or representative samples measured at individual scales, and projected to predict average stability at other scales across a wide range of contexts. Moreover deviations between observed vs. extrapolated scaling relationships can reveal information about unobserved heterogeneity across time, space, or species. We anticipate that these methods will be useful for cross-study synthesis of stability data, extrapolating measurements to unobserved scales, and identifying underlying causes and consequences of heterogeneity.

Lianas in silico, ecological insights from a model of structural parasitism.

Tropical forests are a critical component of the Earth system, storing half of the global forest carbon stocks and accounting for a third of terrestrial photosynthesis. Lianas are structural parasites that can substantially reduce the carbon sequestration capacity of these forests. Simulations of this peculiar growth form have only recently started and a single vegetation model included lianas so far. In this work we present a new liana implementation within the individual based model Formind. Initial tests indicate high structural realism both horizontal and vertical. In particular, we benchmarked the model against empirical observations of size distribution, mean liana cluster size and vertical leaf distribution for the Paracou site in French Guiana. Our model predicted a reduction of above-ground biomass between 10% for mature stands to 45% for secondary plots upon inclusion of lianas in the simulations. The reduced biomass was the result of a lower productivity due to a combination of lower tree photosynthesis and high liana respiration. We evaluated structural metrics (LAI, basal area, mean tree-height) and carbon fluxes (GPP, respiration) by comparing simulations with and without lianas. At the equilibrium, liana productivity was 1.9t C ha - 1 y - 1 , or 23% of the total GPP and the forest carbon stocks were between 5% and 11% lower in simulations with lianas. We also highlight the main strengths and limitations of this new approach and propose new field measurements to further the understanding of liana ecology in a modelling framework.

Network science applied to forest megaplots: tropical tree species coexist in small-world networks.

Network analysis is an important tool to analyze the structure of complex systems such as tropical forests. Here, we infer spatial proximity networks in tropical forests by using network science. First, we focus on tree neighborhoods to derive spatial tree networks from forest inventory data. In a second step, we construct species networks to describe the potential for interactions between species. We find remarkably similar tree and species networks among tropical forests in Panama, Sri Lanka and Taiwan. Across these sites only 32 to 51% of all possible connections between species pairs were realized in the species networks. The species networks show the common small-world property and constant node degree distributions not yet described and explained by network science. Our application of network analysis to forest ecology provides a new approach in biodiversity research to quantify spatial neighborhood structures for better understanding interactions between tree species. Our analyses show that details of tree positions and sizes have no important influence on the detected network structures. This suggests existence of simple principles underlying the complex interactions in tropical forests.

Confronting an individual-based simulation model with empirical community patterns of grasslands.

Grasslands contribute to global biogeochemical cycles and can host a high number of plant species. Both-species dynamics and biogeochemical fluxes-are influenced by abiotic and biotic environmental factors, management and natural disturbances. In order to understand and project grassland dynamics under global change, vegetation models which explicitly capture all relevant processes and drivers are required. However, the parameterization of such models is often challenging. Here, we report on testing an individual- and process-based model for simulating the dynamics and structure of a grassland experiment in temperate Europe. We parameterized the model for three species and confront simulated grassland dynamics with empirical observations of their monocultures and one two-species mixture. The model reproduces general trends of vegetation patterns (vegetation cover and height, aboveground biomass and leaf area index) for the monocultures and two-species community. For example, the model simulates well an average annual grassland cover of 70% in the species mixture (observed cover of 77%), but also shows mismatches with specific observation values (e.g. for aboveground biomass). By a sensitivity analysis of the applied inverse model parameterization method, we demonstrate that multiple vegetation attributes are important for a successful parameterization while leaf area index revealed to be of highest relevance. Results of our study pinpoint to the need of improved grassland measurements (esp. of temporally higher resolution) in close combination with advanced modelling approaches.

Integrating the underlying structure of stochasticity into community ecology.

Stochasticity is a core component of ecology, as it underlies key processes that structure and create variability in nature. Despite its fundamental importance in ecological systems, the concept is often treated as synonymous with unpredictability in community ecology, and studies tend to focus on single forms of stochasticity rather than taking a more holistic view. This has led to multiple narratives for how stochasticity mediates community dynamics. Here, we present a framework that describes how different forms of stochasticity (notably demographic and environmental stochasticity) combine to provide underlying and predictable structure in diverse communities. This framework builds on the deep ecological understanding of stochastic processes acting at individual and population levels and in modules of a few interacting species. We support our framework with a mathematical model that we use to synthesize key literature, demonstrating that stochasticity is more than simple uncertainty. Rather, stochasticity has profound and predictable effects on community dynamics that are critical for understanding how diversity is maintained. We propose next steps that ecologists might use to explore the role of stochasticity for structuring communities in theoretical and empirical systems, and thereby enhance our understanding of community dynamics.

From small-scale forest structure to Amazon-wide carbon estimates.

Tropical forests play an important role in the global carbon cycle. High-resolution remote sensing techniques, e.g., spaceborne lidar, can measure complex tropical forest structures, but it remains a challenge how to interpret such information for the assessment of forest biomass and productivity. Here, we develop an approach to estimate basal area, aboveground biomass and productivity within Amazonia by matching 770,000 GLAS lidar (ICESat) profiles with forest simulations considering spatial heterogeneous environmental and ecological conditions. This allows for deriving frequency distributions of key forest attributes for the entire Amazon. This detailed interpretation of remote sensing data improves estimates of forest attributes by 20-43% as compared to (conventional) estimates using mean canopy height. The inclusion of forest modeling has a high potential to close a missing link between remote sensing measurements and the 3D structure of forests, and may thereby improve continent-wide estimates of biomass and productivity.

An Approach to Study Species Persistence in Unconstrained Random Networks.

The connection between structure and stability of ecological networks has been widely studied in the last fifty years. A challenge that scientists continue to face is that in-depth mathematical model analysis is often difficult, unless the considered systems are specifically constrained. This makes it challenging to generalize results. Therefore, methods are needed that relax the required restrictions. Here, we introduce a novel heuristic approach that provides persistence estimates for random systems without limiting the admissible parameter range and system behaviour. We apply our approach to study persistence of species in random generalized Lotka-Volterra systems and present simulation results, which confirm the accuracy of our predictions. Our results suggest that persistence is mainly driven by the linkage density, whereby additional links can both favour and hinder persistence. In particular, we observed "persistence bistability", a rarely studied feature of random networks, leading to a dependency of persistence on initial species densities. Networks with this property exhibit tipping points, in which species loss can lead to a cascade of extinctions. The methods developed in this paper may facilitate the study of more general models and thereby provide a step forward towards a unifying framework of network architecture and stability.

Rarity of monodominance in hyperdiverse Amazonian forests.

Tropical forests are known for their high diversity. Yet, forest patches do occur in the tropics where a single tree species is dominant. Such "monodominant" forests are known from all of the main tropical regions. For Amazonia, we sampled the occurrence of monodominance in a massive, basin-wide database of forest-inventory plots from the Amazon Tree Diversity Network (ATDN). Utilizing a simple defining metric of at least half of the trees ≥ 10 cm diameter belonging to one species, we found only a few occurrences of monodominance in Amazonia, and the phenomenon was not significantly linked to previously hypothesized life history traits such wood density, seed mass, ectomycorrhizal associations, or Rhizobium nodulation. In our analysis, coppicing (the formation of sprouts at the base of the tree or on roots) was the only trait significantly linked to monodominance. While at specific locales coppicing or ectomycorrhizal associations may confer a considerable advantage to a tree species and lead to its monodominance, very few species have these traits. Mining of the ATDN dataset suggests that monodominance is quite rare in Amazonia, and may be linked primarily to edaphic factors.

Tree mortality submodels drive simulated long-term forest dynamics: assessing 15 models from the stand to global scale.

Models are pivotal for assessing future forest dynamics under the impacts of changing climate and management practices, incorporating representations of tree growth, mortality, and regeneration. Quantitative studies on the importance of mortality submodels are scarce. We evaluated 15 dynamic vegetation models (DVMs) regarding their sensitivity to different formulations of tree mortality under different degrees of climate change. The set of models comprised eight DVMs at the stand scale, three at the landscape scale, and four typically applied at the continental to global scale. Some incorporate empirically derived mortality models, and others are based on experimental data, whereas still others are based on theoretical reasoning. Each DVM was run with at least two alternative mortality submodels. Model behavior was evaluated against empirical time series data, and then, the models were subjected to different scenarios of climate change. Most DVMs matched empirical data quite well, irrespective of the mortality submodel that was used. However, mortality submodels that performed in a very similar manner against past data often led to sharply different trajectories of forest dynamics under future climate change. Most DVMs featured high sensitivity to the mortality submodel, with deviations of basal area and stem numbers on the order of 10-40% per century under current climate and 20-170% under climate change. The sensitivity of a given DVM to scenarios of climate change, however, was typically lower by a factor of two to three. We conclude that (1) mortality is one of the most uncertain processes when it comes to assessing forest response to climate change, and (2) more data and a better process understanding of tree mortality are needed to improve the robustness of simulated future forest dynamics. Our study highlights that comparing several alternative mortality formulations in DVMs provides valuable insights into the effects of process uncertainties on simulated future forest dynamics.

Global patterns of tropical forest fragmentation.

Remote sensing enables the quantification of tropical deforestation with high spatial resolution. This in-depth mapping has led to substantial advances in the analysis of continent-wide fragmentation of tropical forests. Here we identified approximately 130 million forest fragments in three continents that show surprisingly similar power-law size and perimeter distributions as well as fractal dimensions. Power-law distributions have been observed in many natural phenomena such as wildfires, landslides and earthquakes. The principles of percolation theory provide one explanation for the observed patterns, and suggest that forest fragmentation is close to the critical point of percolation; simulation modelling also supports this hypothesis. The observed patterns emerge not only from random deforestation, which can be described by percolation theory, but also from a wide range of deforestation and forest-recovery regimes. Our models predict that additional forest loss will result in a large increase in the total number of forest fragments-at maximum by a factor of 33 over 50 years-as well as a decrease in their size, and that these consequences could be partly mitigated by reforestation and forest protection.

What drives the spatial distribution and dynamics of local species richness in tropical forest?

Understanding the structure and dynamics of highly diverse tropical forests is challenging. Here we investigate the factors that drive the spatio-temporal variation of local tree numbers and species richness in a tropical forest (including 1250 plots of 20 × 20 m2). To this end, we use a series of dynamic models that are built around the local spatial variation of mortality and recruitment rates, and ask which combination of processes can explain the observed spatial and temporal variation in tree and species numbers. We find that processes not included in classical neutral theory are needed to explain these fundamental patterns of the observed local forest dynamics. We identified a large spatio-temporal variability in the local number of recruits as the main missing mechanism, whereas variability of mortality rates contributed to a lesser extent. We also found that local tree numbers stabilize at typical values which can be explained by a simple analytical model. Our study emphasized the importance of spatio-temporal variability in recruitment beyond demographic stochasticity for explaining the local heterogeneity of tropical forests.

Priorities to Advance Monitoring of Ecosystem Services Using Earth Observation.

Managing ecosystem services in the context of global sustainability policies requires reliable monitoring mechanisms. While satellite Earth observation offers great promise to support this need, significant challenges remain in quantifying connections between ecosystem functions, ecosystem services, and human well-being benefits. Here, we provide a framework showing how Earth observation together with socioeconomic information and model-based analysis can support assessments of ecosystem service supply, demand, and benefit, and illustrate this for three services. We argue that the full potential of Earth observation is not yet realized in ecosystem service studies. To provide guidance for priority setting and to spur research in this area, we propose five priorities to advance the capabilities of Earth observation-based monitoring of ecosystem services.

The importance of forest structure to biodiversity-productivity relationships.

While various relationships between productivity and biodiversity are found in forests, the processes underlying these relationships remain unclear and theory struggles to coherently explain them. In this work, we analyse diversity-productivity relationships through an examination of forest structure (described by basal area and tree height heterogeneity). We use a new modelling approach, called 'forest factory', which generates various forest stands and calculates their annual productivity (above-ground wood increment). Analysing approximately 300 000 forest stands, we find that mean forest productivity does not increase with species diversity. Instead forest structure emerges as the key variable. Similar patterns can be observed by analysing 5054 forest plots of the German National Forest Inventory. Furthermore, we group the forest stands into nine forest structure classes, in which we find increasing, decreasing, invariant and even bell-shaped relationships between productivity and diversity. In addition, we introduce a new index, called optimal species distribution, which describes the ratio of realized to the maximal possible productivity (by shuffling species identities). The optimal species distribution and forest structure indices explain the obtained productivity values quite well (R2 between 0.7 and 0.95), whereby the influence of these attributes varies within the nine forest structure classes.

High resolution analysis of tropical forest fragmentation and its impact on the global carbon cycle.

Deforestation in the tropics is not only responsible for direct carbon emissions but also extends the forest edge wherein trees suffer increased mortality. Here we combine high-resolution (30 m) satellite maps of forest cover with estimates of the edge effect and show that 19% of the remaining area of tropical forests lies within 100 m of a forest edge. The tropics house around 50 million forest fragments and the length of the world's tropical forest edges sums to nearly 50 million km. Edge effects in tropical forests have caused an additional 10.3 Gt (2.1-14.4 Gt) of carbon emissions, which translates into 0.34 Gt per year and represents 31% of the currently estimated annual carbon releases due to tropical deforestation. Fragmentation substantially augments carbon emissions from tropical forests and must be taken into account when analysing the role of vegetation in the global carbon cycle.

Do abundance distributions and species aggregation correctly predict macroecological biodiversity patterns in tropical forests?

AIM: It has been recently suggested that different 'unified theories of biodiversity and biogeography' can be characterized by three common 'minimal sufficient rules': (1) species abundance distributions follow a hollow curve, (2) species show intraspecific aggregation, and (3) species are independently placed with respect to other species. Here, we translate these qualitative rules into a quantitative framework and assess if these minimal rules are indeed sufficient to predict multiple macroecological biodiversity patterns simultaneously. LOCATION: Tropical forest plots in Barro Colorado Island (BCI), Panama, and in Sinharaja, Sri Lanka. METHODS: We assess the predictive power of the three rules using dynamic and spatial simulation models in combination with census data from the two forest plots. We use two different versions of the model: (1) a neutral model and (2) an extended model that allowed for species differences in dispersal distances. In a first step we derive model parameterizations that correctly represent the three minimal rules (i.e. the model quantitatively matches the observed species abundance distribution and the distribution of intraspecific aggregation). In a second step we applied the parameterized models to predict four additional spatial biodiversity patterns. RESULTS: Species-specific dispersal was needed to quantitatively fulfil the three minimal rules. The model with species-specific dispersal correctly predicted the species-area relationship, but failed to predict the distance decay, the relationship between species abundances and aggregations, and the distribution of a spatial co-occurrence index of all abundant species pairs. These results were consistent over the two forest plots. MAIN CONCLUSIONS: The three 'minimal sufficient' rules only provide an incomplete approximation of the stochastic spatial geometry of biodiversity in tropical forests. The assumption of independent interspecific placements is most likely violated in many forests due to shared or distinct habitat preferences. Furthermore, our results highlight missing knowledge about the relationship between species abundances and their aggregation.

Monodominance in tropical forests: modelling reveals emerging clusters and phase transitions.

Tropical forests are highly diverse ecosystems, but within such forests there can be large patches dominated by a single tree species. The myriad presumed mechanisms that lead to the emergence of such monodominant areas is currently the subject of intensive research. We used the most generic of these mechanisms, large seed mass and low dispersal ability of the monodominant species, in a spatially explicit model. The model represents seven identical species with long-distance dispersal of small seeds, competing with one potentially monodominant species with short-distance dispersal of large seeds. Monodominant patches emerged and persisted only for a narrow range of species traits; these results have the characteristic features of phase transitions. Additional mechanisms may explain monodominance in different ecological contexts, but our results suggest that percolation-like phenomena and phase transitions might be pervasive in this type of system.