Accounting for herbaceous communities in process-based models will advance our understanding of "grassy" ecosystems.

Grassland and other herbaceous communities cover significant portions of Earth's terrestrial surface and provide many critical services, such as carbon sequestration, wildlife habitat, and food production. Forecasts of global change impacts on these services will require predictive tools, such as process-based dynamic vegetation models. Yet, model representation of herbaceous communities and ecosystems lags substantially behind that of tree communities and forests. The limited representation of herbaceous communities within models arises from two important knowledge gaps: first, our empirical understanding of the principles governing herbaceous vegetation dynamics is either incomplete or does not provide mechanistic information necessary to drive herbaceous community processes with models; second, current model structure and parameterization of grass and other herbaceous plant functional types limits the ability of models to predict outcomes of competition and growth for herbaceous vegetation. In this review, we provide direction for addressing these gaps by: (1) presenting a brief history of how vegetation dynamics have been developed and incorporated into earth system models, (2) reporting on a model simulation activity to evaluate current model capability to represent herbaceous vegetation dynamics and ecosystem function, and (3) detailing several ecological properties and phenomena that should be a focus for both empiricists and modelers to improve representation of herbaceous vegetation in models. Together, empiricists and modelers can improve representation of herbaceous ecosystem processes within models. In so doing, we will greatly enhance our ability to forecast future states of the earth system, which is of high importance given the rapid rate of environmental change on our planet.

A Conceptual Framework to Integrate Biodiversity, Ecosystem Function, and Ecosystem Service Models.

Global biodiversity and ecosystem service models typically operate independently. Ecosystem service projections may therefore be overly optimistic because they do not always account for the role of biodiversity in maintaining ecological functions. We review models used in recent global model intercomparison projects and develop a novel model integration framework to more fully account for the role of biodiversity in ecosystem function, a key gap for linking biodiversity changes to ecosystem services. We propose two integration pathways. The first uses empirical data on biodiversity-ecosystem function relationships to bridge biodiversity and ecosystem function models and could currently be implemented globally for systems and taxa with sufficient data. We also propose a trait-based approach involving greater incorporation of biodiversity into ecosystem function models. Pursuing both approaches will provide greater insight into biodiversity and ecosystem services projections. Integrating biodiversity, ecosystem function, and ecosystem service modeling will enhance policy development to meet global sustainability goals.

Growing-season temperature and precipitation are independent drivers of global variation in xylem hydraulic conductivity.

Stem xylem-specific hydraulic conductivity (KS ) represents the potential for plant water transport normalized by xylem cross section, length, and driving force. Variation in KS has implications for plant transpiration and photosynthesis, growth and survival, and also the geographic distribution of species. Clarifying the global-scale patterns of KS and its major drivers is needed to achieve a better understanding of how plants adapt to different environmental conditions, particularly under climate change scenarios. Here, we compiled a xylem hydraulics dataset with 1,186 species-at-site combinations (975 woody species representing 146 families, from 199 sites worldwide), and investigated how KS varied with climatic variables, plant functional types, and biomes. Growing-season temperature and growing-season precipitation drove global variation in KS independently. Both the mean and the variation in KS were highest in the warm and wet tropical regions, and lower in cold and dry regions, such as tundra and desert biomes. Our results suggest that future warming and redistribution of seasonal precipitation may have a significant impact on species functional diversity, and is likely to be particularly important in regions becoming warmer or drier, such as high latitudes. This highlights an important role for KS in predicting shifts in community composition in the face of climate change.

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.

Joint control of terrestrial gross primary productivity by plant phenology and physiology.

Terrestrial gross primary productivity (GPP) varies greatly over time and space. A better understanding of this variability is necessary for more accurate predictions of the future climate-carbon cycle feedback. Recent studies have suggested that variability in GPP is driven by a broad range of biotic and abiotic factors operating mainly through changes in vegetation phenology and physiological processes. However, it is still unclear how plant phenology and physiology can be integrated to explain the spatiotemporal variability of terrestrial GPP. Based on analyses of eddy-covariance and satellite-derived data, we decomposed annual terrestrial GPP into the length of the CO2 uptake period (CUP) and the seasonal maximal capacity of CO2 uptake (GPPmax). The product of CUP and GPPmax explained >90% of the temporal GPP variability in most areas of North America during 2000-2010 and the spatial GPP variation among globally distributed eddy flux tower sites. It also explained GPP response to the European heatwave in 2003 (r(2) = 0.90) and GPP recovery after a fire disturbance in South Dakota (r(2) = 0.88). Additional analysis of the eddy-covariance flux data shows that the interbiome variation in annual GPP is better explained by that in GPPmax than CUP. These findings indicate that terrestrial GPP is jointly controlled by ecosystem-level plant phenology and photosynthetic capacity, and greater understanding of GPPmax and CUP responses to environmental and biological variations will, thus, improve predictions of GPP over time and space.

Predicting vegetation type through physiological and environmental interactions with leaf traits: evergreen and deciduous forests in an earth system modeling framework.

Earth system models are incorporating plant trait diversity into their land components to better predict vegetation dynamics in a changing climate. However, extant plant trait distributions will not allow extrapolations to novel community assemblages in future climates, which will require a mechanistic understanding of the trade-offs that determine trait diversity. In this study, we show how physiological trade-offs involving leaf mass per unit area (LMA), leaf lifespan, leaf nitrogen, and leaf respiration may explain the distribution patterns of evergreen and deciduous trees in the temperate and boreal zones based on (1) an evolutionary analysis of a simple mathematical model and (2) simulation experiments of an individual-based dynamic vegetation model (i.e., LM3-PPA). The evolutionary analysis shows that these leaf traits set up a trade-off between carbon- and nitrogen-use efficiency at the scale of individual trees and therefore determine competitively dominant leaf strategies. As soil nitrogen availability increases, the dominant leaf strategy switches from one that is high in nitrogen-use efficiency to one that is high in carbon-use efficiency or, equivalently, from high-LMA/long-lived leaves (i.e., evergreen) to low-LMA/short-lived leaves (i.e., deciduous). In a region of intermediate soil nitrogen availability, the dominant leaf strategy may be either deciduous or evergreen depending on the initial conditions of plant trait abundance (i.e., founder controlled) due to feedbacks of leaf traits on soil nitrogen mineralization through litter quality. Simulated successional patterns by LM3-PPA from the leaf physiological trade-offs are consistent with observed successional dynamics of evergreen and deciduous forests at three sites spanning the temperate to boreal zones.

Where does the carbon go? A model-data intercomparison of vegetation carbon allocation and turnover processes at two temperate forest free-air CO2 enrichment sites.

Elevated atmospheric CO2 concentration (eCO2) has the potential to increase vegetation carbon storage if increased net primary production causes increased long-lived biomass. Model predictions of eCO2 effects on vegetation carbon storage depend on how allocation and turnover processes are represented. We used data from two temperate forest free-air CO2 enrichment (FACE) experiments to evaluate representations of allocation and turnover in 11 ecosystem models. Observed eCO2 effects on allocation were dynamic. Allocation schemes based on functional relationships among biomass fractions that vary with resource availability were best able to capture the general features of the observations. Allocation schemes based on constant fractions or resource limitations performed less well, with some models having unintended outcomes. Few models represent turnover processes mechanistically and there was wide variation in predictions of tissue lifespan. Consequently, models did not perform well at predicting eCO2 effects on vegetation carbon storage. Our recommendations to reduce uncertainty include: use of allocation schemes constrained by biomass fractions; careful testing of allocation schemes; and synthesis of allocation and turnover data in terms of model parameters. Data from intensively studied ecosystem manipulation experiments are invaluable for constraining models and we recommend that such experiments should attempt to fully quantify carbon, water and nutrient budgets.

Evaluation of 11 terrestrial carbon-nitrogen cycle models against observations from two temperate Free-Air CO2 Enrichment studies.

We analysed the responses of 11 ecosystem models to elevated atmospheric [CO2 ] (eCO2 ) at two temperate forest ecosystems (Duke and Oak Ridge National Laboratory (ORNL) Free-Air CO2 Enrichment (FACE) experiments) to test alternative representations of carbon (C)-nitrogen (N) cycle processes. We decomposed the model responses into component processes affecting the response to eCO2 and confronted these with observations from the FACE experiments. Most of the models reproduced the observed initial enhancement of net primary production (NPP) at both sites, but none was able to simulate both the sustained 10-yr enhancement at Duke and the declining response at ORNL: models generally showed signs of progressive N limitation as a result of lower than observed plant N uptake. Nonetheless, many models showed qualitative agreement with observed component processes. The results suggest that improved representation of above-ground-below-ground interactions and better constraints on plant stoichiometry are important for a predictive understanding of eCO2 effects. Improved accuracy of soil organic matter inventories is pivotal to reduce uncertainty in the observed C-N budgets. The two FACE experiments are insufficient to fully constrain terrestrial responses to eCO2 , given the complexity of factors leading to the observed diverging trends, and the consequential inability of the models to explain these trends. Nevertheless, the ecosystem models were able to capture important features of the experiments, lending some support to their projections.

Biodiversity loss reduces global terrestrial carbon storage.

Natural ecosystems store large amounts of carbon globally, as organisms absorb carbon from the atmosphere to build large, long-lasting, or slow-decaying structures such as tree bark or root systems. An ecosystem's carbon sequestration potential is tightly linked to its biological diversity. Yet when considering future projections, many carbon sequestration models fail to account for the role biodiversity plays in carbon storage. Here, we assess the consequences of plant biodiversity loss for carbon storage under multiple climate and land-use change scenarios. We link a macroecological model projecting changes in vascular plant richness under different scenarios with empirical data on relationships between biodiversity and biomass. We find that biodiversity declines from climate and land use change could lead to a global loss of between 7.44-103.14 PgC (global sustainability scenario) and 10.87-145.95 PgC (fossil-fueled development scenario). This indicates a self-reinforcing feedback loop, where higher levels of climate change lead to greater biodiversity loss, which in turn leads to greater carbon emissions and ultimately more climate change. Conversely, biodiversity conservation and restoration can help achieve climate change mitigation goals.

Relative information contributions of model vs. data to short- and long-term forecasts of forest carbon dynamics.

Biogeochemical models have been used to evaluate long-term ecosystem responses to global change on decadal and century time scales. Recently, data assimilation has been applied to improve these models for ecological forecasting. It is not clear what the relative information contributions of model (structure and parameters) vs. data are to constraints of short- and long-term forecasting. In this study, we assimilated eight sets of 10-year data (foliage, woody, and fine root biomass, litter fall, forest floor carbon [C], microbial C, soil C, and soil respiration) collected from Duke Forest into a Terrestrial Ecosystem model (TECO). The relative information contribution was measured by Shannon information index calculated from probability density functions (PDFs) of carbon pool sizes. The null knowledge without a model or data was defined by the uniform PDF within a prior range. The relative model contribution was information content in the PDF of modeled carbon pools minus that in the uniform PDF, while the relative data contribution was the information content in the PDF of modeled carbon pools after data was assimilated minus that before data assimilation. Our results showed that the information contribution of the model to constrain carbon dynamics increased with time whereas the data contribution declined. The eight data sets contributed more than the model to constrain C dynamics in foliage and fine root pools over the 100-year forecasts. The model, however, contributed more than the data sets to constrain the litter, fast soil organic matter (SOM), and passive SOM pools. For the two major C pools, woody biomass and slow SOM, the model contributed less information in the first few decades and then more in the following decades than the data. Knowledge of relative information contributions of model vs. data is useful for model development, uncertainty analysis, future data collection, and evaluation of ecological forecasting.

Assimilation of multiple data sets with the ensemble Kalman filter to improve forecasts of forest carbon dynamics.

The ensemble Kalman filter (EnKF) has been used in weather forecasting to assimilate observations into weather models. In this study, we examine how effectively forecasts of a forest carbon cycle can be improved by assimilating observations with the EnKF. We used the EnKF to assimilate into the terrestrial ecosystem (TECO) model eight data sets collected at the Duke Forest between 1996 and 2004 (foliage biomass, fine root biomass, woody biomass, litterfall, microbial biomass, forest floor carbon, soil carbon, and soil respiration). We then used the trained model to forecast changes in carbon pools from 2004 to 2012. Our daily analysis of parameters indicated that all the exit rates were well constrained by the EnKF, with the exception of the exit rates controlling the loss of metabolic litter and passive soil organic matter. The poor constraint of these two parameters resulted from the low sensitivity of TECO predictions to their values and the poor correlation between these parameters and the observed variables. Using the estimated parameters, the model predictions and observations were in agreement. Model forecasts indicate 15 380-15 660 g C/ m2 stored in Duke Forest by 2012 (a 27% increase since 2004). Parameter uncertainties decreased as data were sequentially assimilated into the model using the EnKF. Uncertainties in forecast carbon sinks increased over time for the long-term carbon pools (woody biomass, structure litter, slow and passive SOM) but remained constant over time for the short-term carbon pools (foliage, fine root, metabolic litter, and microbial carbon). Overall, EnKF can effectively assimilate multiple data sets into an ecosystem model to constrain parameters, forecast dynamics of state variables, and evaluate uncertainty.

Robust leaf trait relationships across species under global environmental changes.

Recent studies show coordinated relationships between plant leaf traits and their capacity to predict ecosystem functions. However, how leaf traits will change within species and whether interspecific trait relationships will shift under future environmental changes both remain unclear. Here, we examine the bivariate correlations between leaf economic traits of 515 species in 210 experiments which mimic climate warming, drought, elevated CO2, and nitrogen deposition. We find divergent directions of changes in trait-pairs between species, and the directions mostly do not follow the interspecific trait relationships. However, the slopes in the logarithmic transformed interspecific trait relationships hold stable under environmental changes, while only their elevations vary. The elevation changes of trait relationship are mainly driven by asymmetrically interspecific responses contrary to the direction of the leaf economic spectrum. These findings suggest robust interspecific trait relationships under global changes, and call for linking within-species responses to interspecific coordination of plant traits.

GISS-E2.1: Configurations and Climatology.

This paper describes the GISS-E2.1 contribution to the Coupled Model Intercomparison Project, Phase 6 (CMIP6). This model version differs from the predecessor model (GISS-E2) chiefly due to parameterization improvements to the atmospheric and ocean model components, while keeping atmospheric resolution the same. Model skill when compared to modern era climatologies is significantly higher than in previous versions. Additionally, updates in forcings have a material impact on the results. In particular, there have been specific improvements in representations of modes of variability (such as the Madden-Julian Oscillation and other modes in the Pacific) and significant improvements in the simulation of the climate of the Southern Oceans, including sea ice. The effective climate sensitivity to 2 × CO2 is slightly higher than previously at 2.7-3.1°C (depending on version) and is a result of lower CO2 radiative forcing and stronger positive feedbacks.

Modeling patterns of nonlinearity in ecosystem responses to temperature, CO2, and precipitation changes.

It is commonly acknowledged that ecosystem responses to global climate change are nonlinear. However, patterns of the nonlinearity have not been well characterized on ecosystem carbon and water processes. We used a terrestrial ecosystem (TECO) model to examine nonlinear patterns of ecosystem responses to changes in temperature, CO2, and precipitation individually or in combination. The TECO model was calibrated against experimental data obtained from a grassland ecosystem in the central United States and ran for 100 years with gradual change at 252 different scenarios. We primarily used the 100th-year results to explore nonlinearity of ecosystem responses. Variables examined in this study are net primary production (NPP), heterotrophic respiration (R(h)), net ecosystem carbon exchange (NEE), runoff, and evapotranspiration (ET). Our modeling results show that nonlinear patterns were parabolic, asymptotic, and threshold-like in response to temperature, CO2, and precipitation anomalies, respectively, for NPP, NEE, and R(h). Runoff and ET exhibited threshold-like pattern in response to both temperature and precipitation anomalies but were less sensitive to CO2 changes. Ecosystem responses to combined temperature, CO2, and precipitation anomalies differed considerably from the responses to individual factors in terms of response patterns and/or critical points of nonlinearity. Our results suggest that nonlinear patterns in response to multiple global-change factors were diverse and were considerably affected by combined climate anomalies on ecosystem carbon and water processes. The diverse response patterns in nonlinearity have profound implications for both experimental design and theoretical development.

Forest water use and water use efficiency at elevated CO2 : a model-data intercomparison at two contrasting temperate forest FACE sites.

Predicted responses of transpiration to elevated atmospheric CO2 concentration (eCO2 ) are highly variable amongst process-based models. To better understand and constrain this variability amongst models, we conducted an intercomparison of 11 ecosystem models applied to data from two forest free-air CO2 enrichment (FACE) experiments at Duke University and Oak Ridge National Laboratory. We analysed model structures to identify the key underlying assumptions causing differences in model predictions of transpiration and canopy water use efficiency. We then compared the models against data to identify model assumptions that are incorrect or are large sources of uncertainty. We found that model-to-model and model-to-observations differences resulted from four key sets of assumptions, namely (i) the nature of the stomatal response to elevated CO2 (coupling between photosynthesis and stomata was supported by the data); (ii) the roles of the leaf and atmospheric boundary layer (models which assumed multiple conductance terms in series predicted more decoupled fluxes than observed at the broadleaf site); (iii) the treatment of canopy interception (large intermodel variability, 2-15%); and (iv) the impact of soil moisture stress (process uncertainty in how models limit carbon and water fluxes during moisture stress). Overall, model predictions of the CO2 effect on WUE were reasonable (intermodel µ = approximately 28% ± 10%) compared to the observations (µ = approximately 30% ± 13%) at the well-coupled coniferous site (Duke), but poor (intermodel µ = approximately 24% ± 6%; observations µ = approximately 38% ± 7%) at the broadleaf site (Oak Ridge). The study yields a framework for analysing and interpreting model predictions of transpiration responses to eCO2 , and highlights key improvements to these types of models.

Dynamic disequilibrium of the terrestrial carbon cycle under global change.

In this review, we propose a new framework, dynamic disequilibrium of the carbon cycles, to assess future land carbon-sink dynamics. The framework recognizes internal ecosystem processes that drive the carbon cycle toward equilibrium, such as donor pool-dominated transfer; and external forces that create disequilibrium, such as disturbances and global change. Dynamic disequilibrium within one disturbance-recovery episode causes temporal changes in the carbon source and sink at yearly and decadal scales, but has no impacts on longer-term carbon sequestration unless disturbance regimes shift. Such shifts can result in long-term regional carbon loss or gain and be quantified by stochastic statistics for use in prognostic modeling. If the regime shifts result in ecosystem state changes in regions with large carbon reserves at risk, the global carbon cycle might be destabilized.