Tree carbon allocation explains forest drought-kill and recovery patterns.

The mechanisms governing tree drought mortality and recovery remain a subject of inquiry and active debate given their role in the terrestrial carbon cycle and their concomitant impact on climate change. Counter-intuitively, many trees do not die during the drought itself. Indeed, observations globally have documented that trees often grow for several years after drought before mortality. A combination of meta-analysis and tree physiological models demonstrate that optimal carbon allocation after drought explains observed patterns of delayed tree mortality and provides a predictive recovery framework. Specifically, post-drought, trees attempt to repair water transport tissue and achieve positive carbon balance through regrowing drought-damaged xylem. Furthermore, the number of years of xylem regrowth required to recover function increases with tree size, explaining why drought mortality increases with size. These results indicate that tree resilience to drought-kill may increase in the future, provided that CO2 fertilisation facilitates more rapid xylem regrowth.

Dominance of the suppressed: Power-law size structure in tropical forests.

Tropical tree size distributions are remarkably consistent despite differences in the environments that support them. With data analysis and theory, we found a simple and biologically intuitive hypothesis to explain this property, which is the foundation of forest dynamics modeling and carbon storage estimates. After a disturbance, new individuals in the forest gap grow quickly in full sun until they begin to overtop one another. The two-dimensional space-filling of the growing crowns of the tallest individuals relegates a group of losing, slow-growing individuals to the understory. Those left in the understory follow a power-law size distribution, the scaling of which depends on only the crown area-to-diameter allometry exponent: a well-conserved value across tropical forests.

FOREST ECOLOGY. Pervasive drought legacies in forest ecosystems and their implications for carbon cycle models.

The impacts of climate extremes on terrestrial ecosystems are poorly understood but important for predicting carbon cycle feedbacks to climate change. Coupled climate-carbon cycle models typically assume that vegetation recovery from extreme drought is immediate and complete, which conflicts with the understanding of basic plant physiology. We examined the recovery of stem growth in trees after severe drought at 1338 forest sites across the globe, comprising 49,339 site-years, and compared the results with simulated recovery in climate-vegetation models. We found pervasive and substantial "legacy effects" of reduced growth and incomplete recovery for 1 to 4 years after severe drought. Legacy effects were most prevalent in dry ecosystems, among Pinaceae, and among species with low hydraulic safety margins. In contrast, limited or no legacy effects after drought were simulated by current climate-vegetation models. Our results highlight hysteresis in ecosystem-level carbon cycling and delayed recovery from climate extremes.

Potential role of natural enemies during tree range expansions following climate change.

Recent investigations have shown how chance, long-range dispersal events can allow tree populations to migrate rapidly in response to changes in climate. However, this apparent solution to Reid's paradox applies solely within the context of single species models, while the rapid migration rates seen in pollen records occurred within multispecies communities. Ecologists are therefore presented with a new challenge: reconciling the macroscopic dynamics of spread seen in the pollen record with the rules and interactions governing plant community assembly. A case that highlights this issue is the rapid spread of Beech during the Holocene into a landscape already dominated by a close competitor, Hemlock. In this study, we analyse a simple model of plant community assembly incorporating competition for space and dispersal dynamics, showing how, even when a species is capable of rapid migration into an empty landscape, the presence of an ecologically similar competitor causes Reid's paradox to re-emerge because of the dramatic slowing effect of competitive interactions on a species' rate of spread. We then show how the answer to the question of how tree species dispersed rapidly into occupied landscapes may lie in secondary interactions with host-specific pathogens and parasites. Inclusion of host-specific pathogens into the simple community assembly model illustrates how tree species undergoing range expansions can temporarily outstrip specialist predators, giving rise to a transient Jansen-Connell effect, in which the invader acts as temporary 'super-species' that spreads rapidly into communities already occupied by competitors at rates consistent with those observed in the paleo-record.

Stabilization wedges: solving the climate problem for the next 50 years with current technologies.

Humanity already possesses the fundamental scientific, technical, and industrial know-how to solve the carbon and climate problem for the next half-century. A portfolio of technologies now exists to meet the world's energy needs over the next 50 years and limit atmospheric CO2 to a trajectory that avoids a doubling of the preindustrial concentration. Every element in this portfolio has passed beyond the laboratory bench and demonstration project; many are already implemented somewhere at full industrial scale. Although no element is a credible candidate for doing the entire job (or even half the job) by itself, the portfolio as a whole is large enough that not every element has to be used.

Projecting the future of the U.S. carbon sink.

Atmospheric and ground-based methods agree on the presence of a carbon sink in the coterminous United States (the United States minus Alaska and Hawaii), and the primary causes for the sink recently have been identified. Projecting the future behavior of the sink is necessary for projecting future net emissions. Here we use two models, the Ecosystem Demography model and a second simpler empirically based model (Miami Land Use History), to estimate the spatio-temporal patterns of ecosystem carbon stocks and fluxes resulting from land-use changes and fire suppression from 1700 to 2100. Our results are compared with other historical reconstructions of ecosystem carbon fluxes and to a detailed carbon budget for the 1980s. Our projections indicate that the ecosystem recovery processes that are primarily responsible for the contemporary U.S. carbon sink will slow over the next century, resulting in a significant reduction of the sink. The projected rate of decrease depends strongly on scenarios of future land use and the long-term effectiveness of fire suppression.

Deterministic limits to stochastic spatial models of natural enemies.

Stochastic spatial models are becoming an increasingly popular tool for understanding ecological and epidemiological problems. However, due to the complexities inherent in such models, it has been difficult to obtain any analytical insights. Here, we consider individual-based, stochastic models of both the continuous-time Lotka-Volterra system and the discrete-time Nicholson-Bailey model. The stability of these two stochastic models of natural enemies is assessed by constructing moment equations. The inclusion of these moments, which mimic the effects of spatial aggregation, can produce either stabilizing or destabilizing influences on the population dynamics. Throughout, the theoretical results are compared to numerical models for the full distribution of populations, as well as stochastic simulations.

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.

Long-term studies of vegetation dynamics.

By integrating a wide range of experimental, comparative, and theoretical approaches, ecologists are starting to gain a detailed understanding of the long-term dynamics of vegetation. We explore how patterns of variation in demographic traits among species have provided insight into the processes that structure plant communities. We find a common set of mechanisms, derived from ecological and evolutionary principles, that underlie the main forces shaping systems as diverse as annual plant communities and tropical forests. Trait variation between species maintains diversity and has important implications for ecosystem processes. Hence, greater understanding of how Earth's vegetation functions will likely require integration of ecosystem science with ideas from plant evolutionary, population, and community ecology.

Consistent land- and atmosphere-based U.S. carbon sink estimates.

For the period 1980-89, we estimate a carbon sink in the coterminous United States between 0.30 and 0.58 petagrams of carbon per year (petagrams of carbon = 10(15) grams of carbon). The net carbon flux from the atmosphere to the land was higher, 0.37 to 0.71 petagrams of carbon per year, because a net flux of 0.07 to 0.13 petagrams of carbon per year was exported by rivers and commerce and returned to the atmosphere elsewhere. These land-based estimates are larger than those from previous studies (0.08 to 0.35 petagrams of carbon per year) because of the inclusion of additional processes and revised estimates of some component fluxes. Although component estimates are uncertain, about one-half of the total is outside the forest sector. We also estimated the sink using atmospheric models and the atmospheric concentration of carbon dioxide (the tracer-transport inversion method). The range of results from the atmosphere-based inversions contains the land-based estimates. Atmosphere- and land-based estimates are thus consistent, within the large ranges of uncertainty for both methods. Atmosphere-based results for 1980-89 are similar to those for 1985-89 and 1990-94, indicating a relatively stable U.S. sink throughout the period.

Reinterpreting space, time lags, and functional responses in ecological models.

Natural enemy-victim interactions are of major applied importance and of fundamental interest to ecologists. A key question is what stabilizes these interactions, allowing the long-term coexistence of the two species. Three main theoretical explanations have been proposed: behavioral responses, time-dependent factors such as delayed density dependence, and spatial heterogeneity. Here, using the powerful moment-closure technique, we show a fundamental equivalence between these three elements. Limited movement by organisms is a ubiquitous feature of ecological systems, allowing spatial structure to develop; we show that the effects of this can be naturally described in terms of time lags or within-generation functional responses.

Contributions of land-use history to carbon accumulation in U.S. forests.

Carbon accumulation in forests has been attributed to historical changes in land use and the enhancement of tree growth by CO2 fertilization, N deposition, and climate change. The relative contribution of land use and growth enhancement is estimated by using inventory data from five states spanning a latitudinal gradient in the eastern United States. Land use is the dominant factor governing the rate of carbon accumulation in these states, with growth enhancement contributing far less than previously reported. The estimated fraction of aboveground net ecosystem production due to growth enhancement is 2.0 +/- 4.4%, with the remainder due to land use.

Modeling and analysis of stochastic invasion processes.

In this paper we derive spatially explicit equations to describe a stochastic invasion process. Parents are assumed to produce a random number of offspring which then disperse according to a spatial redistribution kernel. Equations for population moments, such as expected density and covariance averaged over an ensemble of identical stochastic processes, take the form of deterministic integro-difference equations. These equations describe the spatial spread of population moments as the invasion progresses. We use the second order moments to analyse two basic properties of the invasion. The first property is 'permanence of form' in the correlation structure of the wave. Analysis of the asymptotic form of the invasion wave shows that either (i) the covariance in the leading edge of the wave of invasion asymptotically achieves a permanence of form with a characteristic structure described by an unchanging spatial correlation function, or (ii) the leading edge of the wave has no asymptotic permanence of form with the length scales of spatial correlations continually increasing over time. Which of these two outcomes pertains is governed by a single statistic, phi which depends upon the shape of the dispersal kernel and the net reproductive number. The second property of the invasion is its patchy structure. Patchiness, defined in terms of spatial correlations on separate short (within patch) and long (between patch) spatial scales, is linked to the dispersal kernel. Analysis shows how a leptokurtic dispersal kernel gives rise to patchiness in spread of a population.

Limiting Similarity, Species Packing, and System Stability for Hierarchical Competition-Colonization Models.

Hierarchical competition-colonization models have been used to explain limiting similarities among species, successional dynamics, and species loss under habitat destruction. This class of models assumes that there is an inverse relationship between competitive ability and colonization ability and that competitively superior species exclude competitively inferior species when both occupy the same site. This hierarchical model of performance trade-offs, however, exhibits some unusual behaviors in the high-diversity limit, including infinitesimally close species packing, pathologically slow dynamics, and fundamentally important regularities in trait-abundance relationships. In particular, under the condition of constant mortality across species, a 3/2-power-law relationship emerges between abundance and fecundity under infinite packing (abundance of a species with fecundity f is inversely proportional to f to the 3/2 power). In this article, we explore the high-diversity limit of the hierarchical competition-colonization model, with particular emphasis on patterns of species packing, species-abundance relationships, and system stability. Because of the potential for pathologically slow dynamics following perturbations and infinitesimally close species packing in the high-diversity limit for this class of models, the models may need to be modified to include more realistic mechanisms governing the extent and timing of interspecific competitive exclusion in order to effectively capture the structure and dynamics of real-world ecosystems.

A large terrestrial carbon sink in north america implied by atmospheric and oceanic carbon dioxide data and models

Atmospheric carbon dioxide increased at a rate of 2.8 petagrams of carbon per year (Pg C year-1) during 1988 to 1992 (1 Pg = 10(15) grams). Given estimates of fossil carbon dioxide emissions, and net oceanic uptake, this implies a global terrestrial uptake of 1.0 to 2. 2 Pg C year-1. The spatial distribution of the terrestrial carbon dioxide uptake is estimated by means of the observed spatial patterns of the greatly increased atmospheric carbon dioxide data set available from 1988 onward, together with two atmospheric transport models, two estimates of the sea-air flux, and an estimate of the spatial distribution of fossil carbon dioxide emissions. North America is the best constrained continent, with a mean uptake of 1.7 +/- 0.5 Pg C year-1, mostly south of 51 degrees north. Eurasia-North Africa is relatively weakly constrained, with a mean uptake of 0.1 +/- 0.6 Pg C year-1. The rest of the world's land surface is poorly constrained, with a mean source of 0.2 +/- 0.9 Pg C year-1.

Models suggesting field experiments to test two hypotheses explaining successional diversity.

A simple mathematical model of competition is developed that includes two alternative mechanisms promoting successional diversity. The first underpins the competition-colonization hypothesis in which early successional species are able to persist because they colonize disturbed habitats before the arrival of late successional dominant competitors. The second underpins the niche hypothesis, in which early successional species are able to persist, even with unlimited colonization by late successional dominants, because they specialize on the resource-rich conditions typical of recently disturbed sites. We modify the widely studied competition-colonization model so that it also includes the mechanism behind the niche hypothesis. Analysis of this model suggests simple experiments that determine whether the successional diversity of a field system is maintained primarily by the competition-colonization mechanism, primarily by the niche mechanism, by neither, or by both. We develop quantitative metrics of the relative importance of the two mechanisms. We also discuss the implications for the management of biodiversity in communities structured by the two mechanisms.

An analytical treatment of root-to-shoot ratio and plant competition for soil nutrient and light.

By restricting biomass allocation to two compartments only-roots and leaves-we are able to treat plant resource competition analytically. We derive A*i, the optimum allocation strategy for invasion into a bare habitat, and A*f, the allocation strategy of the dominant species in an equilibrium habitat. Furthermore, we prove that in a given habitat succession proceeds from species A*i to species A*f, as long as enough intermediate species are present. Analysis of the relationship between A*i and A*f and the soil nutrient supply and disturbance rates of a habitat predicts that a dichotomy exists in succession. Low soil nutrient supply and disturbance rates favor successions from rooty to leafy species, and high soil nutrient supply and disturbance rates favor the converse. Experimental research is needed to determine whether this pattern exists in nature. Light "consumption" by plants is fundamentally different from soil nutrient consumption in our model. This difference in light and nutrient consumption makes all possible two-species equilibria unstable and leads to founder control. In our model founder control can result in arrested succession and contributes to community diversity.

Herbivores and plant diversity.

We study spatial lottery models of competition between two plant species in which competitive ability is affected by levels of herbivory. Herbivory may enhance plant diversity in two qualitatively different ways. The first is global frequency dependence; the level of herbivory suffered by a plant decreases as the species becomes rare. Second, spatial variability in levels of herbivory can create ephemeral, local refuges for herbivory for each plant species. Both of these mechanisms operate only if there is not a negative correlation between a plant's palatability and its competitive ability. Both mechanisms also require that herbivores have sufficiently strong diet preferences (or, equivalently, that the plants have sufficiently different grazing tolerances). If there is no relationship between palatability and competitive ability, then plant diversity is a monotonically increasing function of the herbivore's degree of monophagy. In contrast, if there is a positive correlation between palatability and competitive ability, then the diversity/degree-of-monophagy relationship may be either monotonically increasing or humped.

Aggregation and the population dynamics of parasitoids and predators.

Aggregation by parasitoids and predators has long been considered an important factor helping to stabilize host-parasitoid and predator-prey interactions. This consensus has recently been challenged by W. W. Murdoch and A. Stewart-Oaten, who conclude, from an analysis of a model formulated in a novel way, that aggregation independent of host density has no effect on stability, whereas aggregation to host or prey patches of high density is normally destabilizing. It is argued here that part of the discrepancy in conclusions results from the use of different concepts of aggregation: behavioral versus statistical. It is also argued that the conclusions of the Murdoch and Stewart-Oaten model should be treated with caution, because unrealistic assumptions have to be made to derive the model, and it is not clear that the conclusions are robust to the relaxation of these assumptions.