The Terrestrial Biosphere Model Farm.

Model Intercomparison Projects (MIPs) are fundamental to our understanding of how the land surface responds to changes in climate. However, MIPs are challenging to conduct, requiring the organization of multiple, decentralized modeling teams throughout the world running common protocols. We explored centralizing these models on a single supercomputing system. We ran nine offline terrestrial biosphere models through the Terrestrial Biosphere Model Farm: CABLE, CENTURY, HyLand, ISAM, JULES, LPJ-GUESS, ORCHIDEE, SiB-3, and SiB-CASA. All models were wrapped in a software framework driven with common forcing data, spin-up, and run protocols specified by the Multi-scale Synthesis and Terrestrial Model Intercomparison Project (MsTMIP) for years 1901-2100. We ran more than a dozen model experiments. We identify three major benefits and three major challenges. The benefits include: (a) processing multiple models through a MIP is relatively straightforward, (b) MIP protocols are run consistently across models, which may reduce some model output variability, and (c) unique multimodel experiments can provide novel output for analysis. The challenges are: (a) technological demand is large, particularly for data and output storage and transfer; (b) model versions lag those from the core model development teams; and (c) there is still a need for intellectual input from the core model development teams for insight into model results. A merger with the open-source, cloud-based Predictive Ecosystem Analyzer (PEcAn) ecoinformatics system may be a path forward to overcoming these challenges.

Using near-term forecasts and uncertainty partitioning to inform prediction of oligotrophic lake cyanobacterial density.

Near-term ecological forecasts provide resource managers advance notice of changes in ecosystem services, such as fisheries stocks, timber yields, or water quality. Importantly, ecological forecasts can identify where there is uncertainty in the forecasting system, which is necessary to improve forecast skill and guide interpretation of forecast results. Uncertainty partitioning identifies the relative contributions to total forecast variance introduced by different sources, including specification of the model structure, errors in driver data, and estimation of current states (initial conditions). Uncertainty partitioning could be particularly useful in improving forecasts of highly variable cyanobacterial densities, which are difficult to predict and present a persistent challenge for lake managers. As cyanobacteria can produce toxic and unsightly surface scums, advance warning when cyanobacterial densities are increasing could help managers mitigate water quality issues. Here, we fit 13 Bayesian state-space models to evaluate different hypotheses about cyanobacterial densities in a low nutrient lake that experiences sporadic surface scums of the toxin-producing cyanobacterium, Gloeotrichia echinulata. We used data from several summers of weekly cyanobacteria samples to identify dominant sources of uncertainty for near-term (1- to 4-week) forecasts of G. echinulata densities. Water temperature was an important predictor of cyanobacterial densities during model fitting and at the 4-week forecast horizon. However, no physical covariates improved model performance over a simple model including the previous week's densities in 1-week-ahead forecasts. Even the best fit models exhibited large variance in forecasted cyanobacterial densities and did not capture rare peak occurrences, indicating that significant explanatory variables when fitting models to historical data are not always effective for forecasting. Uncertainty partitioning revealed that model process specification and initial conditions dominated forecast uncertainty. These findings indicate that long-term studies of different cyanobacterial life stages and movement in the water column as well as measurements of drivers relevant to different life stages could improve model process representation of cyanobacteria abundance. In addition, improved observation protocols could better define initial conditions and reduce spatial misalignment of environmental data and cyanobacteria observations. Our results emphasize the importance of ecological forecasting principles and uncertainty partitioning to refine and understand predictive capacity across ecosystems.

Alternative stable states of the forest mycobiome are maintained through positive feedbacks.

Most trees on Earth form a symbiosis with either arbuscular mycorrhizal or ectomycorrhizal fungi. By forming common mycorrhizal networks, actively modifying the soil environment and other ecological mechanisms, these contrasting symbioses may generate positive feedbacks that favour their own mycorrhizal strategy (that is, the con-mycorrhizal strategy) at the expense of the alternative strategy. Positive con-mycorrhizal feedbacks set the stage for alternative stable states of forests and their fungi, where the presence of different forest mycorrhizal strategies is determined not only by external environmental conditions but also mycorrhiza-mediated feedbacks embedded within the forest ecosystem. Here, we test this hypothesis using thousands of US forest inventory sites to show that arbuscular and ectomycorrhizal tree recruitment and survival exhibit positive con-mycorrhizal density dependence. Data-driven simulations show that these positive feedbacks are sufficient in magnitude to generate and maintain alternative stable states of the forest mycobiome. Given the links between forest mycorrhizal strategy and carbon sequestration potential, the presence of mycorrhizal-mediated alternative stable states affects how we forecast forest composition, carbon sequestration and terrestrial climate feedbacks.

Ecological forecasting of tree growth: Regional fusion of tree-ring and forest inventory data to quantify drivers and characterize uncertainty.

Robust ecological forecasting of tree growth under future climate conditions is critical to anticipate future forest carbon storage and flux. Here, we apply three ingredients of ecological forecasting that are key to improving forecast skill: data fusion, confronting model predictions with new data, and partitioning forecast uncertainty. Specifically, we present the first fusion of tree-ring and forest inventory data within a Bayesian state-space model at a multi-site, regional scale, focusing on Pinus ponderosa var. brachyptera in the southwestern US. Leveraging the complementarity of these two data sources, we parsed the ecological complexity of tree growth into the effects of climate, tree size, stand density, site quality, and their interactions, and quantified uncertainties associated with these effects. New measurements of trees, an ongoing process in forest inventories, were used to confront forecasts of tree diameter with observations, and evaluate alternative tree growth models. We forecasted tree diameter and increment in response to an ensemble of climate change projections, and separated forecast uncertainty into four different causes: initial conditions, parameters, climate drivers, and process error. We found a strong negative effect of fall-spring maximum temperature, and a positive effect of water-year precipitation on tree growth. Furthermore, tree vulnerability to climate stress increases with greater competition, with tree size, and at poor sites. Under future climate scenarios, we forecast increment declines of 22%-117%, while the combined effect of climate and size-related trends results in a 56%-91% decline. Partitioning of forecast uncertainty showed that diameter forecast uncertainty is primarily caused by parameter and initial conditions uncertainty, but increment forecast uncertainty is mostly caused by process error and climate driver uncertainty. This fusion of tree-ring and forest inventory data lays the foundation for robust ecological forecasting of aboveground biomass and carbon accounting at tree, plot, and regional scales, including iterative improvement of model skill.

Liana optical traits increase tropical forest albedo and reduce ecosystem productivity.

Lianas are a key growth form in tropical forests. Their lack of self-supporting tissues and their vertical position on top of the canopy make them strong competitors of resources. A few pioneer studies have shown that liana optical traits differ on average from those of colocated trees. Those trait discrepancies were hypothesized to be responsible for the competitive advantage of lianas over trees. Yet, in the absence of reliable modelling tools, it is impossible to unravel their impact on the forest energy balance, light competition, and on the liana success in Neotropical forests. To bridge this gap, we performed a meta-analysis of the literature to gather all published liana leaf optical spectra, as well as all canopy spectra measured over different levels of liana infestation. We then used a Bayesian data assimilation framework applied to two radiative transfer models (RTMs) covering the leaf and canopy scales to derive tropical tree and liana trait distributions, which finally informed a full dynamic vegetation model. According to the RTMs inversion, lianas grew thinner, more horizontal leaves with lower pigment concentrations. Those traits made the lianas very efficient at light interception and significantly modified the forest energy balance and its carbon cycle. While forest albedo increased by 14% in the shortwave, light availability was reduced in the understorey (-30% of the PAR radiation) and soil temperature decreased by 0.5°C. Those liana-specific traits were also responsible for a significant reduction of tree (-19%) and ecosystem (-7%) gross primary productivity (GPP) while lianas benefited from them (their GPP increased by +27%). This study provides a novel mechanistic explanation to the increase in liana abundance, new evidence of the impact of lianas on forest functioning, and paves the way for the evaluation of the large-scale impacts of lianas on forest biogeochemical cycles.

Effects of the COVID-19 pandemic on noise pollution in three protected areas in metropolitan Boston (USA).

Noise pollution can reduce the ability of urban protected areas to provide a refuge for people and habitat for wildlife. Amidst an unprecedented global pandemic, it is unknown if the changes in human activity have significantly impacted noise pollution in metropolitan parks. We tested the hypothesis that reduced human activity associated with the COVID-19 pandemic lockdowns would lead to reduced sound levels in protected areas compared with non-pandemic times. We measured sound levels in three urban protected areas in metropolitan Boston, MA (USA) at three time periods: in the fall and summer before the pandemic, immediately after the government-imposed lockdown in March 2020 when the trees were leafless, and during the beginning of reopening in early June 2020 when the trees had leaves. At all time periods, sound levels were highest near major roads and demonstrated a logarithmic decrease further from roads. At the two protected areas closest to the city center, sound levels averaged 1-3 dB lower during the time of the pandemic lockdown. In contrast, at the third protected area, which is transected by a major highway, sound levels were 4-6 dB higher during the time of the pandemic, likely because reduced traffic allowed vehicles to travel faster and create more noise. This study demonstrates that altered human levels of activity, in this case associated with the COVID-19 pandemic, can have major, and in some cases unexpected, effects on the levels of noise pollution in protected areas.

Soil microbiome predictability increases with spatial and taxonomic scale.

Soil microorganisms shape ecosystem function, yet it remains an open question whether we can predict the composition of the soil microbiome in places before observing it. Furthermore, it is unclear whether the predictability of microbial life exhibits taxonomic- and spatial-scale dependence, as it does for macrobiological communities. Here, we leverage multiple large-scale soil microbiome surveys to develop predictive models of bacterial and fungal community composition in soil, then test these models against independent soil microbial community surveys from across the continental United States. We find remarkable scale dependence in community predictability. The predictability of bacterial and fungal communities increases with the spatial scale of observation, and fungal predictability increases with taxonomic scale. These patterns suggest that there is an increasing importance of deterministic versus stochastic processes with scale, consistent with findings in plant and animal communities, suggesting a general scaling relationship across biology. Biogeochemical functional groups and high-level taxonomic groups of microorganisms were equally predictable, indicating that traits and taxonomy are both powerful lenses for understanding soil communities. By focusing on out-of-sample prediction, these findings suggest an emerging generality in our understanding of the soil microbiome, and that this understanding is fundamentally scale dependent.

Towards robust statistical inference for complex computer models.

Ecologists increasingly rely on complex computer simulations to forecast ecological systems. To make such forecasts precise, uncertainties in model parameters and structure must be reduced and correctly propagated to model outputs. Naively using standard statistical techniques for this task, however, can lead to bias and underestimation of uncertainties in parameters and predictions. Here, we explain why these problems occur and propose a framework for robust inference with complex computer simulations. After having identified that model error is more consequential in complex computer simulations, due to their more pronounced nonlinearity and interconnectedness, we discuss as possible solutions data rebalancing and adding bias corrections on model outputs or processes during or after the calibration procedure. We illustrate the methods in a case study, using a dynamic vegetation model. We conclude that developing better methods for robust inference of complex computer simulations is vital for generating reliable predictions of ecosystem responses.

Forest responses to last-millennium hydroclimate variability are governed by spatial variations in ecosystem sensitivity.

Forecasts of future forest change are governed by ecosystem sensitivity to climate change, but ecosystem model projections are under-constrained by data at multidecadal and longer timescales. Here, we quantify ecosystem sensitivity to centennial-scale hydroclimate variability, by comparing dendroclimatic and pollen-inferred reconstructions of drought, forest composition and biomass for the last millennium with five ecosystem model simulations. In both observations and models, spatial patterns in ecosystem responses to hydroclimate variability are strongly governed by ecosystem sensitivity rather than climate exposure. Ecosystem sensitivity was higher in models than observations and highest in simpler models. Model-data comparisons suggest that interactions among biodiversity, demography and ecophysiology processes dampen the sensitivity of forest composition and biomass to climate variability and change. Integrating ecosystem models with observations from timescales extending beyond the instrumental record can better understand and forecast the mechanisms regulating forest sensitivity to climate variability in a complex and changing world.

The National Ecological Observatory Network's soil metagenomes: assembly and basic analysis.

The largest dataset of soil metagenomes has recently been released by the National Ecological Observatory Network (NEON), which performs annual shotgun sequencing of soils at 47 sites across the United States. NEON serves as a valuable educational resource, thanks to its open data and programming tutorials, but there is currently no introductory tutorial for accessing and analyzing the soil shotgun metagenomic dataset. Here, we describe methods for processing raw soil metagenome sequencing reads using a bioinformatics pipeline tailored to the high complexity and diversity of the soil microbiome. We describe the rationale, necessary resources, and implementation of steps such as cleaning raw reads, taxonomic classification, assembly into contigs or genomes, annotation of predicted genes using custom protein databases, and exporting data for downstream analysis. The workflow presented here aims to increase the accessibility of NEON's shotgun metagenome data, which can provide important clues about soil microbial communities and their ecological roles.

Beyond ecosystem modeling: A roadmap to community cyberinfrastructure for ecological data-model integration.

In an era of rapid global change, our ability to understand and predict Earth's natural systems is lagging behind our ability to monitor and measure changes in the biosphere. Bottlenecks to informing models with observations have reduced our capacity to fully exploit the growing volume and variety of available data. Here, we take a critical look at the information infrastructure that connects ecosystem modeling and measurement efforts, and propose a roadmap to community cyberinfrastructure development that can reduce the divisions between empirical research and modeling and accelerate the pace of discovery. A new era of data-model integration requires investment in accessible, scalable, and transparent tools that integrate the expertise of the whole community, including both modelers and empiricists. This roadmap focuses on five key opportunities for community tools: the underlying foundations of community cyberinfrastructure; data ingest; calibration of models to data; model-data benchmarking; and data assimilation and ecological forecasting. This community-driven approach is a key to meeting the pressing needs of science and society in the 21st century.

Does the leaf economic spectrum hold within plant functional types? A Bayesian multivariate trait meta-analysis.

The leaf economic spectrum is a widely studied axis of plant trait variability that defines a trade-off between leaf longevity and productivity. While this has been investigated at the global scale, where it is robust, and at local scales, where deviations from it are common, it has received less attention at the intermediate scale of plant functional types (PFTs). We investigated whether global leaf economic relationships are also present within the scale of plant functional types (PFTs) commonly used by Earth System models, and the extent to which this global-PFT hierarchy can be used to constrain trait estimates. We developed a hierarchical multivariate Bayesian model that assumes separate means and covariance structures within and across PFTs and fit this model to seven leaf traits from the TRY database related to leaf longevity, morphology, biochemistry, and photosynthetic metabolism. Although patterns of trait covariation were generally consistent with the leaf economic spectrum, we found three approximate tiers to this consistency. Relationships among morphological and biochemical traits (specific leaf area [SLA], N, P) were the most robust within and across PFTs, suggesting that covariation in these traits is driven by universal leaf construction trade-offs and stoichiometry. Relationships among metabolic traits (dark respiration [Rd ], maximum RuBisCo carboxylation rate [Vc,max ], maximum electron transport rate [Jmax ]) were slightly less consistent, reflecting in part their much sparser sampling (especially for high-latitude PFTs), but also pointing to more flexible plasticity in plant metabolistm. Finally, relationships involving leaf lifespan were the least consistent, indicating that leaf economic relationships related to leaf lifespan are dominated by across-PFT differences and that within-PFT variation in leaf lifespan is more complex and idiosyncratic. Across all traits, this covariance was an important source of information, as evidenced by the improved imputation accuracy and reduced predictive uncertainty in multivariate models compared to univariate models. Ultimately, our study reaffirms the value of studying not just individual traits but the multivariate trait space and the utility of hierarchical modeling for studying the scale dependence of trait relationships.

Global imprint of mycorrhizal fungi on whole-plant nutrient economics.

Mycorrhizal fungi are critical members of the plant microbiome, forming a symbiosis with the roots of most plants on Earth. Most plant species partner with either arbuscular or ectomycorrhizal fungi, and these symbioses are thought to represent plant adaptations to fast and slow soil nutrient cycling rates. This generates a second hypothesis, that arbuscular and ectomycorrhizal plant species traits complement and reinforce these fungal strategies, resulting in nutrient acquisitive vs. conservative plant trait profiles. Here we analyzed 17,764 species level trait observations from 2,940 woody plant species to show that mycorrhizal plants differ systematically in nitrogen and phosphorus economic traits. Differences were clearest in temperate latitudes, where ectomycorrhizal plant species are more nitrogen use- and phosphorus use-conservative than arbuscular mycorrhizal species. This difference is reflected in both aboveground and belowground plant traits and is robust to controlling for evolutionary history, nitrogen fixation ability, deciduousness, latitude, and species climate niche. Furthermore, mycorrhizal effects are large and frequently similar to or greater in magnitude than the influence of plant nitrogen fixation ability or deciduous vs. evergreen leaf habit. Ectomycorrhizal plants are also more nitrogen conservative than arbuscular plants in boreal and tropical ecosystems, although differences in phosphorus use are less apparent outside temperate latitudes. Our findings bolster current theories of ecosystems rooted in mycorrhizal ecology and support the hypothesis that plant mycorrhizal association is linked to the evolution of plant nutrient economic strategies.

The influence of canopy radiation parameter uncertainty on model projections of terrestrial carbon and energy cycling.

Reducing uncertainties in Earth System Model predictions requires carefully evaluating core model processes. Here we examined how canopy radiative transfer model (RTM) parameter uncertainties, in combination with canopy structure, affect terrestrial carbon and energy projections in a demographic land-surface model, the Ecosystem Demography model (ED2). Our analyses focused on temperate deciduous forests and tested canopies of varying structural complexity. The results showed a strong sensitivity of tree productivity, albedo, and energy balance projections to RTM parameters. Impacts of radiative parameter uncertainty on stand-level canopy net primary productivity ranged from ~2 to > 20% and was most sensitive to canopy clumping and leaf reflectance/transmittance in the visible spectrum (~400-750 nm). ED2 canopy albedo varied by ~1 to ~10% and was most sensitive to near-infrared reflectance (~800-1200 nm). Bowen ratio, in turn, was most sensitive to wood optical properties parameterization; this was much larger than expected based on literature, suggesting model instabilities. In vertically and spatially complex canopies the model response to RTM parameterization may show an apparent reduced sensitivity when compared to simpler canopies, masking much larger changes occurring within the canopy. Our findings highlight both the importance of constraining canopy RTM parameters in models and valuating how the canopy structure responds to those parameter values. Finally, we advocate for more model evaluation, similar to this study, to highlight possible issues with model behavior or process representations, particularly models with demographic representations, and identify potential ways to inform and constrain model predictions.

Spatial vs. temporal controls over soil fungal community similarity at continental and global scales.

Large-scale environmental sequencing efforts have transformed our understanding of the spatial controls over soil microbial community composition and turnover. Yet, our knowledge of temporal controls is comparatively limited. This is a major uncertainty in microbial ecology, as there is increasing evidence that microbial community composition is important for predicting microbial community function in the future. Here, we use continental- and global-scale soil fungal community surveys, focused within northern temperate latitudes, to estimate the relative contribution of time and space to soil fungal community turnover. We detected large intra-annual temporal differences in soil fungal community similarity, where fungal communities differed most among seasons, equivalent to the community turnover observed over thousands of kilometers in space. inter-annual community turnover was comparatively smaller than intra-annual turnover. Certain environmental covariates, particularly climate covariates, explained some spatial-temporal effects, though it is unlikely the same mechanisms drive spatial vs. temporal turnover. However, these commonly measured environmental covariates could not fully explain relationships between space, time and community composition. These baseline estimates of fungal community turnover in time provide a starting point to estimate the potential duration of legacies in microbial community composition and function.

Continental-scale nitrogen pollution is shifting forest mycorrhizal associations and soil carbon stocks.

Most tree roots on Earth form a symbiosis with either ecto- or arbuscular mycorrhizal fungi. Nitrogen fertilization is hypothesized to favor arbuscular mycorrhizal tree species at the expense of ectomycorrhizal species due to differences in fungal nitrogen acquisition strategies, and this may alter soil carbon balance, as differences in forest mycorrhizal associations are linked to differences in soil carbon pools. Combining nitrogen deposition data with continental-scale US forest data, we show that nitrogen pollution is spatially associated with a decline in ectomycorrhizal vs. arbuscular mycorrhizal trees. Furthermore, nitrogen deposition has contrasting effects on arbuscular vs. ectomycorrhizal demographic processes, favoring arbuscular mycorrhizal trees at the expense of ectomycorrhizal trees, and is spatially correlated with reduced soil carbon stocks. This implies future changes in nitrogen deposition may alter the capacity of forests to sequester carbon and offset climate change via interactions with the forest microbiome.

Iterative near-term ecological forecasting: Needs, opportunities, and challenges.

Two foundational questions about sustainability are "How are ecosystems and the services they provide going to change in the future?" and "How do human decisions affect these trajectories?" Answering these questions requires an ability to forecast ecological processes. Unfortunately, most ecological forecasts focus on centennial-scale climate responses, therefore neither meeting the needs of near-term (daily to decadal) environmental decision-making nor allowing comparison of specific, quantitative predictions to new observational data, one of the strongest tests of scientific theory. Near-term forecasts provide the opportunity to iteratively cycle between performing analyses and updating predictions in light of new evidence. This iterative process of gaining feedback, building experience, and correcting models and methods is critical for improving forecasts. Iterative, near-term forecasting will accelerate ecological research, make it more relevant to society, and inform sustainable decision-making under high uncertainty and adaptive management. Here, we identify the immediate scientific and societal needs, opportunities, and challenges for iterative near-term ecological forecasting. Over the past decade, data volume, variety, and accessibility have greatly increased, but challenges remain in interoperability, latency, and uncertainty quantification. Similarly, ecologists have made considerable advances in applying computational, informatic, and statistical methods, but opportunities exist for improving forecast-specific theory, methods, and cyberinfrastructure. Effective forecasting will also require changes in scientific training, culture, and institutions. The need to start forecasting is now; the time for making ecology more predictive is here, and learning by doing is the fastest route to drive the science forward.

Vegetation demographics in Earth System Models: A review of progress and priorities.

Numerous current efforts seek to improve the representation of ecosystem ecology and vegetation demographic processes within Earth System Models (ESMs). These developments are widely viewed as an important step in developing greater realism in predictions of future ecosystem states and fluxes. Increased realism, however, leads to increased model complexity, with new features raising a suite of ecological questions that require empirical constraints. Here, we review the developments that permit the representation of plant demographics in ESMs, and identify issues raised by these developments that highlight important gaps in ecological understanding. These issues inevitably translate into uncertainty in model projections but also allow models to be applied to new processes and questions concerning the dynamics of real-world ecosystems. We argue that stronger and more innovative connections to data, across the range of scales considered, are required to address these gaps in understanding. The development of first-generation land surface models as a unifying framework for ecophysiological understanding stimulated much research into plant physiological traits and gas exchange. Constraining predictions at ecologically relevant spatial and temporal scales will require a similar investment of effort and intensified inter-disciplinary communication.

Probing the limits of predictability: data assimilation of chaotic dynamics in complex food webs.

The daunting complexity of ecosystems has led ecologists to use mathematical modelling to gain understanding of ecological relationships, processes and dynamics. In pursuit of mathematical tractability, these models use simplified descriptions of key patterns, processes and relationships observed in nature. In contrast, ecological data are often complex, scale-dependent, space-time correlated, and governed by nonlinear relations between organisms and their environment. This disparity in complexity between ecosystem models and data has created a large gap in ecology between model and data-driven approaches. Here, we explore data assimilation (DA) with the Ensemble Kalman filter to fuse a two-predator-two-prey model with abundance data from a 2600+ day experiment of a plankton community. We analyse how frequently we must assimilate measured abundances to predict accurately population dynamics, and benchmark our population model's forecast horizon against a simple null model. Results demonstrate that DA enhances the predictability and forecast horizon of complex community dynamics.

Prediction in ecology: a first-principles framework.

Quantitative predictions are ubiquitous in ecology, yet there is limited discussion on the nature of prediction in this field. Herein I derive a general quantitative framework for analyzing and partitioning the sources of uncertainty that control predictability. The implications of this framework are assessed conceptually and linked to classic questions in ecology, such as the relative importance of endogenous (density-dependent) vs. exogenous factors, stability vs. drift, and the spatial scaling of processes. The framework is used to make a number of novel predictions and reframe approaches to experimental design, model selection, and hypothesis testing. Next, the quantitative application of the framework to partitioning uncertainties is illustrated using a short-term forecast of net ecosystem exchange. Finally, I advocate for a new comparative approach to studying predictability across different ecological systems and processes and lay out a number of hypotheses about what limits predictability and how these limits should scale in space and time.