Drew W. Purves

Identification of 100 fundamental ecological questions

Summary 1. Fundamental ecological research is both intrinsically interesting and provides the basic knowledge required to answer applied questions of importance to the management of the natural world. The 100th anniversary of the British Ecological Society in 2013 is an opportune moment to reflect on the current status of ecology as a science and look forward to high-light priorities for future work.

Predictive Models of Forest Dynamics

Dynamic global vegetation models (DGVMs) have shown that forest dynamics could dramatically alter the response of the global climate system to increased atmospheric carbon dioxide over the next century. But there is little agreement between different DGVMs, making forest dynamics one of the greatest sources of uncertainty in predicting future climate. DGVM predictions could be strengthened by integrating the ecological realities of biodiversity and height-structured competition for light, facilitated by recent advances in the mathematics of forest modeling, ecological understanding of diverse forest communities, and the availability of forest inventory data.

Assessing uncertainties in a second-generation dynamic vegetation model caused by ecological scale limitations

*Second-generation Dynamic Global Vegetation Models (DGVMs) have recently been developed that explicitly represent the ecological dynamics of disturbance, vertical competition for light, and succession. Here, we introduce a modified second-generation DGVM and examine how the representation of demographic processes operating at two-dimensional spatial scales not represented by these models can influence predicted community structure, and responses of ecosystems to climate change. *The key demographic processes we investigated were seed advection, seed mixing, sapling survival, competitive exclusion and plant mortality. We varied these parameters in the context of a simulated Amazon rainforest ecosystem containing seven plant functional types (PFTs) that varied along a trade-off surface between growth and the risk of starvation induced mortality. *Varying the five unconstrained parameters generated community structures ranging from monocultures to equal co-dominance of the seven PFTs. When exposed to a climate change scenario, the competing impacts of CO(2) fertilization and increasing plant mortality caused ecosystem biomass to diverge substantially between simulations, with mid-21st century biomass predictions ranging from 1.5 to 27.0 kg C m(-2). *Filtering the results using contemporary observation ranges of biomass, leaf area index (LAI), gross primary productivity (GPP) and net primary productivity (NPP) did not substantially constrain the potential outcomes. We conclude that demographic processes represent a large source of uncertainty in DGVM predictions.

SCALING FROM TREES TO FORESTS: TRACTABLE MACROSCOPIC EQUATIONS FOR FOREST DYNAMICS

Individual-based forest simulators, such as TASS and SORTIE, are spatial stochastic processes that predict properties of populations and communities by simulating the fate of every plant throughout its life cycle. Although they are used for forest management and are able to predict dynamics of real forests, they are also analytically intractable, which limits their usefulness to basic scientists. We have developed a new spatial individual-based forest model that includes a perfect plasticity formulation for crown shape. Its structure allows us to derive an accurate approximation for the individual-based model that predicts mean densities and size structures using the same parameter values and functional forms, and also it is analytically tractable. The approximation is represented by a system of von Foerster partial differential equations coupled with an integral equation that we call the perfect plasticity approximation (PPA). We have derived a series of analytical results including equilibrium abundances for trees of different crown shapes, stability conditions, transient behaviors, such as the constant yield law and self-thinning exponents, and two species coexistence conditions.

Predicting and understanding forest dynamics using a simple tractable model

The perfect-plasticity approximation (PPA) is an analytically tractable model of forest dynamics, defined in terms of parameters for individual trees, including allometry, growth, and mortality. We estimated these parameters for the eight most common species on each of four soil types in the US Lake states (Michigan, Wisconsin, and Minnesota) by using short-term (≤15-year) inventory data from individual trees. We implemented 100-year PPA simulations given these parameters and compared these predictions to chronosequences of stand development. Predictions for the timing and magnitude of basal area dynamics and ecological succession on each soil were accurate, and predictions for the diameter distribution of 100-year-old stands were correct in form and slope. For a given species, the PPA provides analytical metrics for early-successional performance (H20, height of a 20-year-old open-grown tree) and late-successional performance (Ẑ*, equilibrium canopy height in monoculture). These metrics predicted which species were early or late successional on each soil type. Decomposing Ẑ* showed that (i) succession is driven both by superior understory performance and superior canopy performance of late-successional species, and (ii) performance differences primarily reflect differences in mortality rather than growth. The predicted late-successional dominants matched chronosequences on xeromesic (Quercus rubra) and mesic (codominance by Acer rubrum and Acer saccharum) soil. On hydromesic and hydric soils, the literature reports that the current dominant species in old stands (Thuja occidentalis) is now failing to regenerate. Consistent with this, the PPA predicted that, on these soils, stands are now succeeding to dominance by other late-successional species (e.g., Fraxinus nigra, A. rubrum).

Biotic Interactions in the Tropics: Ecological drift in niche-structured communities: neutral pattern does not imply neutral process

Introduction The neutral vs. structure debate We can define a neutral community as one in which all species, and so all individuals, are equivalent, in the sense that they are interchangeable at all times and under all conditions. In contrast, we can define a structured community as one in which species are not equivalent, and species-specific differences affect the population dynamics, and therefore the behaviour, of the community. This distinction is an important one, because in a neutral community the biodiversity, as measured by species richness and abundance patterns, has nothing to do with the biogeochemical functioning of the community (e.g. carbon fixation and nutrient-cycling). In fact, in a truly neutral community one could eliminate all but one species without affecting the biogeochemical functioning of the community at all. In contrast, much of the species-specific variation in biological traits observed in reality (see below) has direct relevance for the functioning of the community. For example, the short-term carbon uptake of a forest depends on the growth rates of the individual trees, and the long-term carbon storage depends on adult life-span and wood density, and there is wide species-specific variation in these traits. In niche-structured communities, the biodiversity and functioning are intimately linked, and some combination of at least some species is required to maintain the functioning of the community.

Influences of forest structure, climate and species composition on tree mortality across the eastern US.

Few studies have quantified regional variation in tree mortality, or explored whether species compositional changes or within-species variation are responsible for regional patterns, despite the fact that mortality has direct effects on the dynamics of woody biomass, species composition, stand structure, wood production and forest response to climate change. Using Bayesian analysis of over 430,000 tree records from a large eastern US forest database we characterised tree mortality as a function of climate, soils, species and size (stem diameter). We found (1) mortality is U-shaped vs. stem diameter for all 21 species examined; (2) mortality is hump-shaped vs. plot basal area for most species; (3) geographical variation in mortality is substantial, and correlated with several environmental factors; and (4) individual species vary substantially from the combined average in the nature and magnitude of their mortality responses to environmental variation. Regional variation in mortality is therefore the product of variation in species composition combined with highly varied mortality-environment correlations within species. The results imply that variation in mortality is a crucial part of variation in the forest carbon cycle, such that including this variation in models of the global carbon cycle could significantly narrow uncertainty in climate change predictions.

Crown Plasticity and Competition for Canopy Space: A New Spatially Implicit Model Parameterized for 250 North American Tree Species

Background Canopy structure, which can be defined as the sum of the sizes, shapes and relative placements of the tree crowns in a forest stand, is central to all aspects of forest ecology. But there is no accepted method for deriving canopy structure from the sizes, species and biomechanical properties of the individual trees in a stand. Any such method must capture the fact that trees are highly plastic in their growth, forming tessellating crown shapes that fill all or most of the canopy space. Methodology/Principal Findings We introduce a new, simple and rapidly-implemented model–the Ideal Tree Distribution, ITD–with tree form (height allometry and crown shape), growth plasticity, and space-filling, at its core. The ITD predicts the canopy status (in or out of canopy), crown depth, and total and exposed crown area of the trees in a stand, given their species, sizes and potential crown shapes. We use maximum likelihood methods, in conjunction with data from over 100,000 trees taken from forests across the coterminous US, to estimate ITD model parameters for 250 North American tree species. With only two free parameters per species–one aggregate parameter to describe crown shape, and one parameter to set the so-called depth bias–the model captures between-species patterns in average canopy status, crown radius, and crown depth, and within-species means of these metrics vs stem diameter. The model also predicts much of the variation in these metrics for a tree of a given species and size, resulting solely from deterministic responses to variation in stand structure. Conclusions/Significance This new model, with parameters for US tree species, opens up new possibilities for understanding and modeling forest dynamics at local and regional scales, and may provide a new way to interpret remote sensing data of forest canopies, including LIDAR and aerial photography.

Animal Versus Wind Dispersal and the Robustness of Tree Species to Deforestation

Studies suggest that populations of different species do not decline equally after habitat loss. However, empirical tests have been confined to fine spatiotemporal scales and have rarely included plants. Using data from 89,365 forest survey plots covering peninsular Spain, we explored, for each of 34 common tree species, the relationship between probability of occurrence and the local cover of remaining forest. Twenty-four species showed a significant negative response to forest loss, so that decreased forest cover had a negative effect on tree diversity, but the responses of individual species were highly variable. Animal-dispersed species were less vulnerable to forest loss, with six showing positive responses to decreased forest cover. The results imply that plant-animal interactions help prevent the collapse of forest communities that suffer habitat destruction.

Facultative nitrogen fixation by canopy legumes in a lowland tropical forest

Symbiotic dinitrogen (N2) fixation is often invoked to explain the N richness of tropical forests as ostensibly N2-fixing trees can be a major component of the community. Such arguments assume N2 fixers are fixing N when present. However, in laboratory experiments, legumes consistently reduce N2 fixation in response to increased soil N availability. These contrasting views of N2 fixation as either obligate or facultative have drastically different implications for the N cycle of tropical forests. We tested these models by directly measuring N2-fixing root nodules and nitrogenase activity of individual canopy-dominant legume trees (Inga sp.) across several lowland forest types. Fixation was substantial in disturbed forests and some gaps but near zero in the high N soils of mature forest. Our findings suggest that canopy legumes closely regulate N2 fixation, leading to large variations in N inputs across the landscape, and low symbiotic fixation in mature forests despite abundant legumes.