Stephen H. Roxburgh

THE INTERMEDIATE DISTURBANCE HYPOTHESIS: PATCH DYNAMICS AND MECHANISMS OF SPECIES COEXISTENCE

The intermediate disturbance hypothesis (IDH) has been used for several decades as an explanation for the coexistence of species in ecological communities. It is intuitively simple, but deceptively so. We show, via discussion and examples, that the IDH is not one mechanism of coexistence, but rather summarizes a set of similar phenomena that can arise from the action of several different coexistence mechanisms. These underlying mechanisms are defined by the various ways in which species differ in their response to disturbance-induced spatial and temporal variability in resources and environmental conditions. As an example, the original specification of the IDH required patchy disturbances for coexistence. However, because the underlying mechanisms of coexistence can also operate at the within-patch scale, patchy disturbances are not a necessary requirement for coexistence under intermediate-disturbance regimes. These conclusions are illustrated through the analysis of three models: a spatial within-patch model, a spatial between-patch model, and a purely temporal model. All three generate similar patterns of coexistence under intermediate disturbance, yet underlying that coexistence lie at least two quite-distinct mechanisms of species coexistence: the storage effect and relative nonlinearity. The results from our analyses suggest that, as a promoter of species coexistence, the IDH is both broader in scope and richer in detail than has previously been recognized.

How frequency and intensity shape diversity–disturbance relationships

Understanding the relationship between disturbance regimes and species diversity has been of central interest to ecologists for decades. For example, the intermediate disturbance hypothesis proposes that diversity will be highest at intermediate levels of disturbance. Although peaked (hump-shaped) diversity–disturbance relationships (DDRs) have been documented in nature, many other DDRs have been reported as well. Here, we begin to theoretically unify these diverse empirical findings by showing how a single simple model can generate several different DDRs, depending on the aspect of disturbance that is considered. Additionally, we elucidate the competition-mediated mechanism underlying our results. Our findings have the potential to reconcile apparently conflicting empirical results on the effects of disturbance on diversity.

A NEW METHOD FOR DETECTING SPECIES ASSOCIATIONS WITH SPATIALLY AUTOCORRELATED DATA

Many organisms display patchiness in their distribution patterns over a wide range of spatial scales. Patchy distribution patterns can be caused by processes such as growth, migration, reproduction, and mortality, which result in neighboring areas being more likely to contain a species than distant areas, a phenomenon known as positive spatial autocorrelation. When species are patchily distributed, the within-species spatial random- ness assumptions of the standard statistical tests for detecting species associations are seriously violated. Using these tests under such circumstances can lead to incorrect rejection of the null hypothesis. To address this problem we introduce a new test for detecting species associations—the random patterns test. This test takes into account spatial autocorrelation by including the characteristics of the spatial pattern of each species into the null model. A randomization procedure was used to generate the null distribution of the test statistic. The random patterns test is illustrated with data collected from an herbaceous understory community of a Eucalyptus forest near Canberra, Australia.

A critical overview of model estimates of net primary productivity for the Australian continent

Net primary production links the biosphere and the climate system through the global cycling of carbon, water and nutrients. Accurate quantification of net primary productivity (NPP) is therefore critical in understanding the response of the worlds ecosystems to global climate change, and how changes in ecosystems might themselves feed back to the climate system.

Modelling the potential for prescribed burning to mitigate carbon emissions from wildfires in fire-prone forests of Australia

Prescribed fire can potentially reduce carbon emissions from unplanned fires. This potential will differ among ecosystems owing to inherent differences in the efficacy of prescribed burning in reducing unplanned fire activity (or ‘leverage’, i.e. the reduction in area of unplanned fire per unit area of prescribed fire). In temperate eucalypt forests, prescribed burning leverage is relatively low and potential for mitigation of carbon emissions from unplanned fires via prescribed fire is potentially limited. Simulations of fire regimes accounting for non-linear patterns of fuel dynamics for three fuel types characteristic of eucalypt forests in south-eastern Australia supported this prediction. Estimated mean annual fuel consumption increased with diminishing leverage and increasing rate of prescribed burning, even though average fire intensity (prescribed and unplanned fires combined) decreased. The results indicated that use of prescribed burning in these temperate forests is unlikely to yield a net reduction in carbon emissions. Future increases in burning rates under climate change may increase emissions and reduce carbon sequestration. A more detailed understanding of the efficacy of prescribed burning and dynamics of combustible biomass pools is required to clarify the potential for mitigation of carbon emissions in temperate eucalypt forests and other ecosystems.

Experimental evidence rejects pairwise modelling approach to coexistence in plant communities

Competition is often invoked as the cause of plant species loss with increasing system productivity. Experimental results for multispecies assemblages are virtually absent and mathematical models are thus used to explore the relationship between competition and coexistence. Modelling approaches to coexistence and diversity in competitive communities commonly employ Lotka–Volterra-type (LV) models with additive pairwise competitive effects. Using pairwise plant competition experiments, we calibrate the LV system and use it to predict plant biomass and coexistence in six three-species and one seven-species experimental mixture. Our results show that five out of the six three-species sets and the seven-species set deviate significantly from LV model predictions. Fitting an additional non-additive competition coefficient resulted in predictions that more closely matched the experimental results, with stable coexistence suggested in all but one case. These results are discussed with particular reference to the possible underlying mechanisms of coexistence in our experimental community. Modelling the effect of competition intensity on stability indicates that if non-additive effects occur, they will be relevant over a wide range of community sizes. Our findings caution against relying on coexistence predictions based on LV models.

The Effect of Carbon Credits on Savanna Land Management and Priorities for Biodiversity Conservation

Carbon finance offers the potential to change land management and conservation planning priorities. We develop a novel approach to planning for improved land management to conserve biodiversity while utilizing potential revenue from carbon biosequestration. We apply our approach in northern Australias tropical savanna, a region of global significance for biodiversity and carbon storage, both of which are threatened by current fire and grazing regimes. Our approach aims to identify priority locations for protecting species and vegetation communities by retaining existing vegetation and managing fire and grazing regimes at a minimum cost. We explore the impact of accounting for potential carbon revenue (using a carbon price of US

An ecoclimatic framework for evaluating the resilience of vegetation to water deficit

14 per tonne of carbon dioxide equivalent) on priority areas for conservation and the impact of explicitly protecting carbon stocks in addition to biodiversity. Our results show that improved management can potentially raise approximately US

Diversity–disturbance relationships: frequency and intensity interact

5 per hectare per year in carbon revenue and prevent the release of 1–2 billion tonnes of carbon dioxide equivalent over approximately 90 years. This revenue could be used to reduce the costs of improved land management by three quarters or double the number of biodiversity targets achieved and meet carbon storage targets for the same cost. These results are based on generalised cost and carbon data; more comprehensive applications will rely on fine scale, site-specific data and a supportive policy environment. Our research illustrates that the duel objective of conserving biodiversity and reducing the release of greenhouse gases offers important opportunities for cost-effective land management investments.

Vegetation state change and consequent carbon dynamics in savanna woodlands of Australia in response to grazing, drought and fire: a scenario approach using 113 years of synthetic annual fire and grassland growth

The surge in global efforts to understand the causes and consequences of drought on forest ecosystems has tended to focus on specific impacts such as mortality. We propose an ecoclimatic framework that takes a broader view of the ecological relevance of water deficits, linking elements of exposure and resilience to cumulative impacts on a range of ecosystem processes. This ecoclimatic framework is underpinned by two hypotheses: (i) exposure to water deficit can be represented probabilistically and used to estimate exposure thresholds across different vegetation types or ecosystems; and (ii) the cumulative impact of a series of water deficit events is defined by attributes governing the resistance and recovery of the affected processes. We present case studies comprising Pinus edulis and Eucalyptus globulus, tree species with contrasting ecological strategies, which demonstrate how links between exposure and resilience can be examined within our proposed framework. These examples reveal how climatic thresholds can be defined along a continuum of vegetation functional responses to water deficit regimes. The strength of this framework lies in identifying climatic thresholds on vegetation function in the absence of more complete mechanistic understanding, thereby guiding the formulation, application and benchmarking of more detailed modelling.