Stephen W. Pacala

A Large and Persistent Carbon Sink in the World’s Forests

Net average global annual uptake of atmospheric carbon dioxide by forests was 1.1 petagrams of carbon, roughly one-sixth of fossil fuel emissions. The terrestrial carbon sink has been large in recent decades, but its size and location remain uncertain. Using forest inventory data and long-term ecosystem carbon studies, we estimate a total forest sink of 2.4 ± 0.4 petagrams of carbon per year (Pg C year–1) globally for 1990 to 2007. We also estimate a source of 1.3 ± 0.7 Pg C year–1 from tropical land-use change, consisting of a gross tropical deforestation emission of 2.9 ± 0.5 Pg C year–1 partially compensated by a carbon sink in tropical forest regrowth of 1.6 ± 0.5 Pg C year–1. Together, the fluxes comprise a net global forest sink of 1.1 ± 0.8 Pg C year–1, with tropical estimates having the largest uncertainties. Our total forest sink estimate is equivalent in magnitude to the terrestrial sink deduced from fossil fuel emissions and land-use change sources minus ocean and atmospheric sinks.

Beneficial Biofuels—The Food, Energy, and Environment Trilemma

Exploiting multiple feedstocks, under new policies and accounting rules, to balance biofuel production, food security, and greenhouse-gas reduction. Recent analyses of the energy and greenhouse-gas performance of alternative biofuels have ignited a controversy that may be best resolved by applying two simple principles. In a world seeking solutions to its energy, environmental, and food challenges, society cannot afford to miss out on the global greenhouse-gas emission reductions and the local environmental and societal benefits when biofuels are done right. However, society also cannot accept the undesirable impacts of biofuels done wrong.

Recent patterns and mechanisms of carbon exchange by terrestrial ecosystems

Knowledge of carbon exchange between the atmosphere, land and the oceans is important, given that the terrestrial and marine environments are currently absorbing about half of the carbon dioxide that is emitted by fossil-fuel combustion. This carbon uptake is therefore limiting the extent of atmospheric and climatic change, but its long-term nature remains uncertain. Here we provide an overview of the current state of knowledge of global and regional patterns of carbon exchange by terrestrial ecosystems. Atmospheric carbon dioxide and oxygen data confirm that the terrestrial biosphere was largely neutral with respect to net carbon exchange during the 1980s, but became a net carbon sink in the 1990s. This recent sink can be largely attributed to northern extratropical areas, and is roughly split between North America and Eurasia. Tropical land areas, however, were approximately in balance with respect to carbon exchange, implying a carbon sink that offset emissions due to tropical deforestation. The evolution of the terrestrial carbon sink is largely the result of changes in land use over time, such as regrowth on abandoned agricultural land and fire prevention, in addition to responses to environmental changes, such as longer growing seasons, and fertilization by carbon dioxide and nitrogen. Nevertheless, there remain considerable uncertainties as to the magnitude of the sink in different regions and the contribution of different processes.

Reid's Paradox of Rapid Plant Migration Dispersal theory and interpretation of paleoecological records

James S. Clark, Jason Lynch, and Peter Wyckoff are in the Department of Botany, Duke University, Durham, NC 27708; Chris Fastie and Stephen T. Jackson are in the Department of Botany, University of Wyoming, Laramie, WY 82701; George Hurtt and Stephen Pacala are in the Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544-1003; Carter Johnson is in the Department of Horticulture and Forestry, South Dakota State University, Brookings, SD 57007; George A. King is at Dynamic Corporation, US EPA National Health and Environmental Effects Research Laboratory, Corvallis, OR 97333; Mark Lewis is in the Math Department, University of Utah, Salt Lake City, UT 84112; Colin Prentice is at the School of Ecology, Lund University, Lund, Sweden; Eugene W. Schupp is in the Department of Rangeland Resources, Utah State University, Logan, UT 84322; and Thompson Webb III is in the Department of Geological Sciences, Brown University, Providence, RI 029121846. ? 1998 American Institute of Biological Sciences. A plausible explanation

A method for scaling vegetation dynamics: The ecosystem demography model (ED)

The problem of scale has been a critical impediment to incorporating im- portant fine-scale processes into global ecosystem models. Our knowledge of fine-scale physiological and ecological processes comes from a variety of measurements, ranging from forest plot inventories to remote sensing, made at spatial resolutions considerably smaller than the large scale at which global ecosystem models are defined. In this paper, we describe a new individual-based, terrestrial biosphere model, which we label the eco- system demography model (ED). We then introduce a general method for scaling stochastic individual-based models of ecosystem dynamics (gap models) such as ED to large scales. The method accounts for the fine-scale spatial heterogeneity within an ecosystem caused by stochastic disturbance events, operating at scales down to individual canopy-tree-sized gaps. By conditioning appropriately on the occurrence of these events, we derive a size- and age-structured (SAS) approximation for the first moment of the stochastic ecosystem model. With this approximation, it is possible to make predictions about the large scales of interest from a description of the fine-scale physiological and population-dynamic pro- cesses without simulating the fate of every plant individually. We use the SAS approxi- mation to implement our individual-based biosphere model over South America from 15 8 Nt o1 5 8S, showing that the SAS equations are accurate across a range of environmental conditions and resulting ecosystem types. We then compare the predictions of the biosphere model to regional data and to intensive data at specific sites. Analysis of the model at these sites illustrates the importance of fine-scale heterogeneity in governing large-scale eco- system function, showing how population and community-level processes influence eco- system composition and structure, patterns of aboveground carbon accumulation, and net ecosystem production.

Mechanisms of long-distance dispersal of seeds by wind

Long-distance dispersal (LDD) is central to species expansion following climate change, re-colonization of disturbed areas and control of pests. The current paradigm is that the frequency and spatial extent of LDD events are extremely difficult to predict. Here we show that mechanistic models coupling seed release and aerodynamics with turbulent transport processes provide accurate probabilistic descriptions of LDD of seeds by wind. The proposed model reliably predicts the vertical distribution of dispersed seeds of five tree species observed along a 45-m high tower in an eastern US deciduous forest. Simulations show that uplifting above the forest canopy is necessary and sufficient for LDD, hence, they provide the means to define LDD quantitatively rather than arbitrarily. Seed uplifting probability thus sets an upper bound on the probability of long-distance colonization. Uplifted yellow poplar seeds are on average lighter than seeds at the forest floor, but also include the heaviest seeds. Because uplifting probabilities are appreciable (as much as 1–5%), and tree seed crops are commonly massive, some LDD events will establish individuals that can critically affect plant dynamics on large scales.

Seedling Recruitment in Forests: Calibrating Models to Predict Patterns of Tree Seedling Dispersion

Recruitment, the addition of new individuals into a community, is an im- portant factor that can substantially affect community composition and dynamics. We present a method for calibrating spatial models of plant recruitment that does not require identifying the specific parent of each recruit. This method calibrates seedling recruitment functions by comparing tree seedling distributions with adult distributions via a maximum likelihood analysis. The models obtained from this method can then be used to predict the spatial distributions of seedlings from adult distributions. We calibrated recruitment functions for 10 tree species characteristic of transition oak- northern hardwood forests. Significant differences were found in recruit abundances and spatial distributions. Predicted seedling recruitment limitation for test stands varied sub- stantially between species, with little recruitment limitation for some species and strong recruitment limitation for others. Recruitment was limited due to low overall recruit production or to restricted recruit dispersion. When these seedling recruitment parameters were incorporated into a spatial, individual-based model of forest dynamics, called SOR- TIE, alterations of recruitment parameters produced substantial changes in species abun- dance, providing additional support for the potential importance of seedling recruitment processes in community structure and dynamics.

Spatial Moment Equations for Plant Competition: Understanding Spatial Strategies and the Advantages of Short Dispersal

A plant lineage can compete for resources in a spatially variable environment by colonizing new areas, exploiting resources in those areas quickly before other plants arrive to compete with it, or tolerating competition once other plants do arrive. These specializations are ubiquitous in plant communities, but all three have never been derived from a spatial model of community dynamics—instead, the possibility of rapid exploitation has been either overlooked or confounded with colonization. We use moment equations, equations for the mean densities and spatial covariance of competing plant populations, to characterize these strategies in a fully spatial stochastic model. The moment equations predict endogenous spatial pattern formation and the efficacy of spatial strategies under different conditions. The model shows that specializations for colonization, exploitation, and tolerance are all possible, and these are the only possible spatial strategies; among them, they partition all of the endogenous spatial structure in the environment. The model predicts two distinct short‐dispersal specializations where parents disperse their offspring locally, either to exploit empty patches quickly or to fill patches to exclude competitors.

Greater focus needed on methane leakage from natural gas infrastructure

Natural gas is seen by many as the future of American energy: a fuel that can provide energy independence and reduce greenhouse gas emissions in the process. However, there has also been confusion about the climate implications of increased use of natural gas for electric power and transportation. We propose and illustrate the use of technology warming potentials as a robust and transparent way to compare the cumulative radiative forcing created by alternative technologies fueled by natural gas and oil or coal by using the best available estimates of greenhouse gas emissions from each fuel cycle (i.e., production, transportation and use). We find that a shift to compressed natural gas vehicles from gasoline or diesel vehicles leads to greater radiative forcing of the climate for 80 or 280 yr, respectively, before beginning to produce benefits. Compressed natural gas vehicles could produce climate benefits on all time frames if the well-to-wheels CH4 leakage were capped at a level 45–70% below current estimates. By contrast, using natural gas instead of coal for electric power plants can reduce radiative forcing immediately, and reducing CH4 losses from the production and transportation of natural gas would produce even greater benefits. There is a need for the natural gas industry and science community to help obtain better emissions data and for increased efforts to reduce methane leakage in order to minimize the climate footprint of natural gas.

POPULATION REGULATION: HISTORICAL CONTEXT AND CONTEMPORARY CHALLENGES OF OPEN VS. CLOSED SYSTEMS

By definition, a population is regulated if it persists for many generations with fluctuations bounded above zero with high probability. Regulation thus requires den- sity-dependent negative feedback whereby the population has a propensity to increase when small and decrease when large. Ultimately, extinction occurs due to regulating mechanisms becoming weaker than various disruptive events and stochastic variation. Population reg- ulation is one of the foundational concepts of ecology, yet this paradigm has often been challenged, during the first half of the 20th century when the concept was not clearly defined, and more recently by some who study demographically open populations. The history of ecology reveals that earlier manifestations of the concept focused mostly on competition as the mechanism of population regulation. Because competition is often not evident in nature, it was sometimes concluded that population regulation was therefore also absent. However, predation in the broadest sense can also cause density dependence. By the 1950s, the idea that demographic density dependence was essential (but not suffi- cient) for population regulation was well established, and since then, challenges to the general concept have been short lived. However, some now believe that metapopulations composed of demographically open local populations can persist without density depen- dence. In particular, some recent manifestations of the Recruitment Limitation Hypothesis all but preclude the possibility of regulation. The theory of locally open populations indicates that persistence always relies on direct demographic density dependence at some spatial and temporal scale, even in models re- portedly demonstrating the contrary. There is also increasing empirical evidence, especially in marine systems where competition for space is not self evident, that local density de- pendence is more pervasive than previously assumed and is often caused by predation. However, there are currently insufficient data to test unequivocally whether or not any persistent metapopulation is regulated. The challenge for more complete understanding of regulation of metapopulations lies in combined empirical and theoretical studies that bridge the gap between smaller scale field experiments and larger scale phenomena that can pres- ently be explored solely by theory.