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.

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.

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.

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.

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.

The parasites of Anolis lizards the northern Lesser Antilles : II. The structure of the parasite community.

The communities of parasitic helminths from ten species of lizards on seven islands in the Caribbean were examined to ascertain the relative importance of predictable deterministic factors and unpredicatable colonization or extinction events in determining the structure of the parasite community. A simple graphical model of community structure is used as a "null model" to describe the features of a community that are dependent only upon the size of the host population and features of the life histories of the constituent parasite species. This model predicts that parasite species will exhibit a nested pattern of local and global relative abundance. The observed data correspond fairly well to this pattern. The absences of individual parasite species from communities where they might be expected to be present emphasizes the role of stochastic colonization and extinction events in delineating the constituent members of the community on any island.Statistical analysis of the distribution of parasite species per host illustrates that this pattern is random in habitats where parasite species diversity is low, but decreasingly variable in habitats where more diverse parasite communities occur. Increased parasite diversity also leads to an increase in the proportion of hosts that contain mixed species infections. Comparisons of worm burdens from single and mixed species infections within individual hosts suggest that interactions between parasite species only rarely leads to reduced worm burdens.

Effects of competitive asymmetry on a local density model of plant interference.

Although competition between plants is usually asymmetric (i.e. larger plants have a disproportionate effect on smaller plants) almost all models of plant competition at the local level have assumed symmetric competition. We add a simple version of competitive asymmetry to the local density neighborhood models of plant interference and population dynamics developed by Pacala & Silander (1985, Am. Nat. 125, 385-411; 1987, Oikos 48, 217-224) by assuming that plants within a neighborhood can be put in a linear dominance hierarchy based upon their initial size. The size of a focal plant is a function of the number of dominant and the number of subordinate neighbors within its neighborhood, with subordinate neighbors having less of an effect than dominant ones. Asymmetry prevents precipitous changes in focal plant size with changes in local density, making the relationship between focal plant size and local density hyperbolic, even if the symmetric model is not hyperbolic. Thus, asymmetry makes the model conform to the law of constant final yield, irrespective of the form of the relationship between plant size and local crowding. Asymmetry also prevents population dynamic oscillations in the model in cases in which it would occur in the absence of asymmetry. The results show that asymmetry has major effects on a model of local interference in plants, and point to the importance of including it in such models.

Stochastic simulation of clonal growth in the tall goldenrod, Solidago altissima.

As clonal plants grow they move through space. The movement patterns that result can be complex and difficult to interpret without the aid of models. We developed a stochastic simulation model of clonal growth in the tall goldenrod, Solidago altissima. Our model was calibrated with field data on the clonal expansion of both seedlings and established clones, and model assumptions were verified by statistical analyses.When simulations were based on empirical distributions with long rhizome lengths, there was greater dispersal, less leaf overlap, and less spatial aggregation than when simulations were based on distributions with comparatively short rhizome lengths. For the field data that we utilized, variation in rhizome lengths had a greater effect than variation for either branching angles or "rhizome initiation points" (see text). We also found that observed patterns of clonal growth in S. altissima did not cause the formation of "fairy rings". However, simulations with an artificial distribution of branching angles demonstrate that "fairy rings" can result solely from a plant's clonal morphology.Stochastic simulation models that incorporated variation in rhizome lengths, branching angles, and rhizome initiation points produced greater dispersal and less leaf overlap than deterministic models. Thus, variation for clonal growth parameters may increase the efficiency of substrate exploration by increasing the area covered and by decreasing the potential for intraclonal competition. We also demonstrated that ramet displacements were slightly, but consistently lower in stochastic simulation models than in random-walk models. This difference was due to the incorporation of details on rhizome bud initiation into stochastic simulation models, but not random-walk models. We discuss the advantages and disadvantages of deterministic, stochastic simulation, and random-walk models of clonal growth.

Host-parasitoid associations in patchy environments.

Studies of insect host-parasitoid interactions have contributed much to the consensus that spatial patchiness is important in the regulation of natural populations. A variety of theoretical models predict that host and parasitoid populations, although unstable in the absence of environmental heterogeneity, may persist at roughly steady overall densities in a patchy environment owing to variation in levels of parasitism from patch to patch. Observed patterns of parasitism, however, have a variety of forms (with variation in attack rates among patches depending directly or indirectly on host density, or showing variation uncorrelated with host density). There is some confusion about the dynamical consequences of these different forms. Here we first show how the dynamical effects of all these forms of environmental heterogeneity can be assessed by a common criterion. This 'CV2 greater than 1 rule' states that the overall population densities will remain roughly steady from generation to generation if the coefficient of variation squared (CV2) of the density of searching parasitoids in the vicinity of each host exceeds approximately unity. By partitioning CV2 into components, we show that both direct and inverse patterns of dependence on host density, and density-independent patterns, all contribute to population regulation in the same way. Second, we show how a maximum-likelihood method can be applied to the kind of field data that are usually available (that is, percentage parasitism versus local host density) to estimate the components of CV2. This analysis indicates that heterogeneity is large enough to stabilize dynamics in 9 of 34 published studies, and that density-independent heterogeneity is the main factor in most cases.

The relation between the number of parasites/host and host age: population dynamic causes and maximum likelihood estimation.

We examined dynamical factors that shape the distribution of the number of parasites/host in constant or temporally varying environments, and with or without host-age dependent variation in host susceptibility and parasite mortality. We predict properties of the parasite distribution in the absence of density-dependent factors such as density-dependent mortality of recruitment and parasite-induced host mortality. These properties provide a criterion for the detection of density dependence in temporally variable systems with host-age dependent interactions. We have then introduced methods to estimate and statistically evaluate the effects of host age or size on the distribution of parasites/host. The methods are based on a maximum likelihood protocol for linear and non-linear regression when data are negatively binomially distributed. We have illustrated the use of the theoretical results and statistical methods by re-analysing the data of Halvorsen & Andersen (1984) on cestode infections in Norwegian arctic charr and by analysing new data on nematode infections in Caribbean Anolis lizards.