Earth system models (ESMs) have been developed to represent the role of terrestrial ecosystems on the energy, water, and carbon cycles. However, many ESMs still lack representation of ...within-ecosystem heterogeneity and diversity. In this paper, we present the Ecosystem Demography model version 2.2 (ED-2.2). In ED-2.2, the biophysical and physiological processes account for the horizontal and vertical heterogeneity of the ecosystem: the energy, water, and carbon cycles are solved separately for a series of vegetation cohorts (groups of individual plants of similar size and plant functional type) distributed across a series of spatially implicit patches (representing collections of micro-environments that have a similar disturbance history). We define the equations that describe the energy, water, and carbon cycles in terms of total energy, water, and carbon, which simplifies the differential equations and guarantees excellent conservation of these quantities in long-term simulation (< 0.1 % error over 50 years). We also show examples of ED-2.2 simulation results at single sites and across tropical South America. These results demonstrate the model's ability to characterize the variability of ecosystem structure, composition, and functioning both at stand and continental scales. A detailed model evaluation was conducted and is presented in a companion paper (Longo et al., 2019a). Finally, we highlight some of the ongoing model developments designed to improve the model's accuracy and performance and to include processes hitherto not represented in the model.
Climatic extreme events are expected to occur more frequently in the future, increasing the likelihood of unprecedented climate extremes (UCEs) or record-breaking events. UCEs, such as extreme ...heatwaves and droughts, substantially affect ecosystem stability and carbon cycling by increasing plant mortality and delaying ecosystem recovery. Quantitative knowledge of such effects is limited due to the paucity of experiments focusing on extreme climatic events beyond the range of historical experience. Here, we present a road map of how dynamic vegetation demographic models (VDMs) can be used to investigate hypotheses surrounding ecosystem responses to one type of UCE: unprecedented droughts. As a result of nonlinear ecosystem responses to UCEs that are qualitatively different from responses to milder extremes, we consider both biomass loss and recovery rates over time by reporting a time-integrated carbon loss as a result of UCE, relative to the absence of drought. Additionally, we explore how unprecedented droughts in combination with increasing atmospheric CO2 and/or temperature may affect ecosystem stability and carbon cycling. We explored these questions using simulations of pre-drought and post-drought conditions at well-studied forest sites using well-tested models (ED2 and LPJ-GUESS). The severity and patterns of biomass losses differed substantially between models. For example, biomass loss could be sensitive to either drought duration or drought intensity depending on the model approach. This is due to the models having different, but also plausible, representations of processes and interactions, highlighting the complicated variability of UCE impacts that still need to be narrowed down in models. Elevated atmospheric CO2 concentrations (eCO2) alone did not completely buffer the ecosystems from carbon losses during UCEs in the majority of our simulations. Our findings highlight the consequences of differences in process formulations and uncertainties in models, most notably related to availability in plant carbohydrate storage and the diversity of plant hydraulic schemes, in projecting potential ecosystem responses to UCEs. We provide a summary of the current state and role of many model processes that give way to different underlying hypotheses of plant responses to UCEs, reflecting knowledge gaps which in future studies could be tested with targeted field experiments and an iterative modeling–experimental conceptual framework.
Methane (CH4) emission by carbon-rich cryosols at the high latitudes in Northern Hemisphere has been studied extensively. In contrast, data on the CH4 emission potential of carbon-poor cryosols is ...limited, despite their spatial predominance. This work employs CH4 flux measurements in the field and under laboratory conditions to show that the mineral cryosols at Axel Heiberg Island in the Canadian high Arctic consistently consume atmospheric CH4. Omics analyses present the first molecular evidence of active atmospheric CH4-oxidizing bacteria (atmMOB) in permafrost-affected cryosols, with the prevalent atmMOB genotype in our acidic mineral cryosols being closely related to Upland Soil Cluster α. The atmospheric (atm) CH4 uptake at the study site increases with ground temperature between 0 °C and 18 °C. Consequently, the atm CH4 sink strength is predicted to increase by a factor of 5-30 as the Arctic warms by 5-15 °C over a century. We demonstrate that acidic mineral cryosols are a previously unrecognized potential of CH4 sink that requires further investigation to determine its potential impact on larger scales. This study also calls attention to the poleward distribution of atmMOB, as well as to the potential influence of microbial atm CH4 oxidation, in the context of regional CH4 flux models and global warming.
The Ecosystem Demography model version 2.2 (ED-2.2) is a terrestrial biosphere model that simulates the biophysical, ecological, and biogeochemical dynamics of vertically and horizontally ...heterogeneous terrestrial ecosystems. In a companion paper (Longo et al., 2019a), we described how the model solves the energy, water, and carbon cycles, and verified the high degree of conservation of these properties in long-term simulations that include long-term (multi-decadal) vegetation dynamics. Here, we present a detailed assessment of the model's ability to represent multiple processes associated with the biophysical and biogeochemical cycles in Amazon forests. We use multiple measurements from eddy covariance towers, forest inventory plots, and regional remote-sensing products to assess the model's ability to represent biophysical, physiological, and ecological processes at multiple timescales, ranging from subdaily to century long. The ED-2.2 model accurately describes the vertical distribution of light, water fluxes, and the storage of water, energy, and carbon in the canopy air space, the regional distribution of biomass in tropical South America, and the variability of biomass as a function of environmental drivers. In addition, ED-2.2 qualitatively captures several emergent properties of the ecosystem found in observations, specifically observed relationships between aboveground biomass, mortality rates, and wood density; however, the slopes of these relationships were not accurately captured. We also identified several limitations, including the model's tendency to overestimate the magnitude and seasonality of heterotrophic respiration and to overestimate growth rates in a nutrient-poor tropical site. The evaluation presented here highlights the potential of incorporating structural and functional heterogeneity within biomes in Earth system models (ESMs) and to realistically represent their impacts on energy, water, and carbon cycles. We also identify several priorities for further model development.
We assessed whether diversity in plant hydraulic traits can explain the observed diversity in plant responses to water stress in seasonally dry tropical forests (SDTFs).
The Ecosystem Demography ...model 2 (ED2) was updated with a trait-driven mechanistic plant hydraulic module, as well as novel drought-phenology and plant water stress schemes. Four plant functional types were parameterized on the basis of meta-analysis of plant hydraulic traits. Simulations from both the original and the updated ED2 were evaluated against 5 yr of field data from a Costa Rican SDTF site and remote-sensing data over Central America.
The updated model generated realistic plant hydraulic dynamics, such as leaf water potential and stem sap flow. Compared with the original ED2, predictions from our novel traitdriven model matched better with observed growth, phenology and their variations among functional groups. Most notably, the original ED2 produced unrealistically small leaf area index (LAI) and underestimated cumulative leaf litter. Both of these biases were corrected by the updated model. The updated model was also better able to simulate spatial patterns of LAI dynamics in Central America.
Plant hydraulic traits are intercorrelated in SDTFs. Mechanistic incorporation of plant hydraulic traits is necessary for the simulation of spatiotemporal patterns of vegetation dynamics in SDTFs in vegetation models.
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.
In this review, we summarize in detail the development of vegetation demographics models as components of Earth System Models. We particularly highlight the ecological uncertainties around the strength of growth‐resource acquisition feedbacks that are common across model developments, and illustrate the myriad new opportunities for ecological‐scale data streams to inform and validate these new model structures.
Drought‐related tree mortality is now a widespread phenomenon predicted to increase in magnitude with climate change. However, the patterns of which species and trees are most vulnerable to drought, ...and the underlying mechanisms have remained elusive, in part due to the lack of relevant data and difficulty of predicting the location of catastrophic drought years in advance. We used long‐term demographic records and extensive databases of functional traits and distribution patterns to understand the responses of 20–53 species to an extreme drought in a seasonally dry tropical forest in Costa Rica, which occurred during the 2015 El Niño Southern Oscillation event. Overall, species‐specific mortality rates during the drought ranged from 0% to 34%, and varied little as a function of tree size. By contrast, hydraulic safety margins correlated well with probability of mortality among species, while morphological or leaf economics spectrum traits did not. This firmly suggests hydraulic traits as targets for future research.
We took advantage of an exceptionally strong El Niño‐related drought in 2015 to understand the effects on tree mortality in a diverse, seasonally dry tropical forest. Tree mortality rates differed widely among species, ranging from 0% to 34%, and varied little as a function of tree size. We correlated the mean mortality rates by species with a large number of potential explanatory variables and found that hydraulic safety margins correlated well with probability of mortality among species, while soft traits such as wood density did not. This firmly suggests hydraulic traits as targets for future research.
Seasonally dry tropical forests (SDTF) are located in regions with alternating wet and dry seasons, with dry seasons that last several months or more. By the end of the 21st century, climate models ...predict substantial changes in rainfall regimes across these regions, but little is known about how individuals, species, and communities in SDTF will cope with the hotter, drier conditions predicted by climate models. In this review, we explore different rainfall scenarios that may result in ecological drought in SDTF through the lens of two alternative hypotheses: 1) these forests will be sensitive to drought because they are already limited by water and close to climatic thresholds, or 2) they will be resistant/resilient to intra- and inter-annual changes in rainfall because they are adapted to predictable, seasonal drought. In our review of literature that spans microbial to ecosystem processes, a majority of the available studies suggests that increasing frequency and intensity of droughts in SDTF will likely alter species distributions and ecosystem processes. Though we conclude that SDTF will be sensitive to altered rainfall regimes, many gaps in the literature remain. Future research should focus on geographically comparative studies and well-replicated drought experiments that can provide empirical evidence to improve simulation models used to forecast SDTF responses to future climate change at coarser spatial and temporal scales.
Lianas, or woody vines, and trees dominate the canopy of tropical forests and comprise the majority of tropical aboveground carbon storage. These growth forms respond differently to contemporary ...variation in climate and resource availability, but their responses to future climate change are poorly understood because there are very few predictive ecosystem models representing lianas. We compile a database of liana functional traits (846 species) and use it to parameterize a mechanistic model of liana-tree competition. The substantial difference between liana and tree hydraulic conductivity represents a critical source of inter-growth form variation. Here, we show that lianas are many times more sensitive to drying atmospheric conditions than trees as a result of this trait difference. Further, we use our competition model and projections of tropical hydroclimate based on Representative Concentration Pathway 4.5 to show that lianas are more susceptible to reaching a hydraulic threshold for viability by 2100.
• There are two theories about how allocation of metabolic products occurs. The allometric biomass partitioning theory (APT) suggests that all plants follow common allometric scaling rules. The ...optimal partitioning theory (OPT) predicts that plants allocate more biomass to the organ capturing the most limiting resource.
• Whole-plant harvests of mature and juvenile tropical deciduous trees, evergreen trees, and lianas and model simulations were used to address the following knowledge gaps: (1) Do mature lianas comply with the APT scaling laws or do they invest less biomass in stems compared to trees? (2) Do juveniles follow the same allocation patterns as mature individuals? (3) Is either leaf phenology or life form a predictor of rooting depth?
• It was found that: (1) mature lianas followed the same allometric scaling laws as trees; (2) juveniles and mature individuals do not follow the same allocation patterns; and (3) mature lianas had shallowest coarse roots and evergreen trees had the deepest.
• It was demonstrated that: (1) mature lianas invested proportionally similar biomass to stems as trees and not less, as expected; (2) lianas were not deeper-rooted than trees as had been previously proposed; and (3) evergreen trees had the deepest roots, which is necessary to maintain canopy during simulated dry seasons.
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