Mark T. Bulling

Predictable waves of sequential forest degradation and biodiversity loss spreading from an African city

Tropical forest degradation emits carbon at a rate of ~0.5 Pg·y−1, reduces biodiversity, and facilitates forest clearance. Understanding degradation drivers and patterns is therefore crucial to managing forests to mitigate climate change and reduce biodiversity loss. Putative patterns of degradation affecting forest stocks, carbon, and biodiversity have variously been described previously, but these have not been quantitatively assessed together or tested systematically. Economic theory predicts a systematic allocation of land to its highest use value in response to distance from centers of demand. We tested this theory to see if forest exploitation would expand through time and space as concentric waves, with each wave targeting lower value products. We used forest data along a transect from 10 to 220 km from Dar es Salaam (DES), Tanzania, collected at two points in time (1991 and 2005). Our predictions were confirmed: high-value logging expanded 9 km·y−1, and an inner wave of lower value charcoal production 2 km·y−1. This resource utilization is shown to reduce the public goods of carbon storage and species richness, which significantly increased with each kilometer from DES [carbon, 0.2 Mg·ha−1; 0.1 species per sample area (0.4 ha)]. Our study suggests that tropical forest degradation can be modeled and predicted, with its attendant loss of some public goods. In sub-Saharan Africa, an area experiencing the highest rate of urban migration worldwide, coupled with a high dependence on forest-based resources, predicting the spatiotemporal patterns of degradation can inform policies designed to extract resources without unsustainably reducing carbon storage and biodiversity.

Bacterial biodiversity-ecosystem functioning relations are modified by environmental complexity

Background With the recognition that environmental change resulting from anthropogenic activities is causing a global decline in biodiversity, much attention has been devoted to understanding how changes in biodiversity may alter levels of ecosystem functioning. Although environmental complexity has long been recognised as a major driving force in evolutionary processes, it has only recently been incorporated into biodiversity-ecosystem functioning investigations. Environmental complexity is expected to strengthen the positive effect of species richness on ecosystem functioning, mainly because it leads to stronger complementarity effects, such as resource partitioning and facilitative interactions among species when the number of available resource increases. Methodology/Principal Findings Here we implemented an experiment to test the combined effect of species richness and environmental complexity, more specifically, resource richness on ecosystem functioning over time. We show, using all possible combinations of species within a bacterial community consisting of six species, and all possible combinations of three substrates, that diversity-functioning (metabolic activity) relationships change over time from linear to saturated. This was probably caused by a combination of limited complementarity effects and negative interactions among competing species as the experiment progressed. Even though species richness and resource richness both enhanced ecosystem functioning, they did so independently from each other. Instead there were complex interactions between particular species and substrate combinations. Conclusions/Significance Our study shows clearly that both species richness and environmental complexity increase ecosystem functioning. The finding that there was no direct interaction between these two factors, but that instead rather complex interactions between combinations of certain species and resources underlie positive biodiversity ecosystem functioning relationships, suggests that detailed knowledge of how individual species interact with complex natural environments will be required in order to make reliable predictions about how altered levels of biodiversity will most likely affect ecosystem functioning.

Habitat structure mediates biodiversity effects on ecosystem properties

Much of what we know about the role of biodiversity in mediating ecosystem processes and function stems from manipulative experiments, which have largely been performed in isolated, homogeneous environments that do not incorporate habitat structure or allow natural community dynamics to develop. Here, we use a range of habitat configurations in a model marine benthic system to investigate the effects of species composition, resource heterogeneity and patch connectivity on ecosystem properties at both the patch (bioturbation intensity) and multi-patch (nutrient concentration) scale. We show that allowing fauna to move and preferentially select patches alters local species composition and density distributions, which has negative effects on ecosystem processes (bioturbation intensity) at the patch scale, but overall positive effects on ecosystem functioning (nutrient concentration) at the multi-patch scale. Our findings provide important evidence that community dynamics alter in response to localized resource heterogeneity and that these small-scale variations in habitat structure influence species contributions to ecosystem properties at larger scales. We conclude that habitat complexity forms an important buffer against disturbance and that contemporary estimates of the level of biodiversity required for maintaining future multi-functional systems may need to be revised.

Marine biodiversity–ecosystem functions under uncertain environmental futures

Anthropogenic activity is currently leading to dramatic transformations of ecosystems and losses of biodiversity. The recognition that these ecosystems provide services that are essential for human well-being has led to a major interest in the forms of the biodiversity–ecosystem functioning relationship. However, there is a lack of studies examining the impact of climate change on these relationships and it remains unclear how multiple climatic drivers may affect levels of ecosystem functioning. Here, we examine the roles of two important climate change variables, temperature and concentration of atmospheric carbon dioxide, on the relationship between invertebrate species richness and nutrient release in a model benthic estuarine system. We found a positive relationship between invertebrate species richness and the levels of release of NH4-N into the water column, but no effect of species richness on the release of PO4-P. Higher temperatures and greater concentrations of atmospheric carbon dioxide had a negative impact on nutrient release. Importantly, we found significant interactions between the climate variables, indicating that reliably predicting the effects of future climate change will not be straightforward as multiple drivers are unlikely to have purely additive effects, resulting in increased levels of uncertainty.

Species effects on ecosystem processes are modified by faunal responses to habitat composition

Heterogeneity is a well-recognized feature of natural environments, and the spatial distribution and movement of individual species is primarily driven by resource requirements. In laboratory experiments designed to explore how different species drive ecosystem processes, such as nutrient release, habitat heterogeneity is often seen as something which must be rigorously controlled for. Most small experimental systems are therefore spatially homogeneous, and the link between environmental heterogeneity and its effects on the redistribution of individuals and species, and on ecosystem processes, has not been fully explored. In this paper, we used a mesocosm system to investigate the relationship between habitat composition, species movement and sediment nutrient release for each of four functionally contrasting species of marine benthic invertebrate macrofauna. For each species, various habitat configurations were generated by selectively enriching patches of sediment with macroalgae, a natural source of spatial variability in intertidal mudflats. We found that the direction and extent of faunal movement between patches differs with species identity, density and habitat composition. Combinations of these factors lead to concomitant changes in nutrient release, such that habitat composition effects are modified by species identity (in the case of NH4–N) and by species density (in the case of PO4–P). It is clear that failure to accommodate natural patterns of spatial heterogeneity in such studies may result in an incomplete understanding of system behaviour. This will be particularly important for future experiments designed to explore the effects of species richness on ecosystem processes, where the complex interactions reported here for single species may be compounded when species are brought together in multi-species combinations.

Influence of macrofaunal assemblages and environmental heterogeneity on microphytobenthic production in experimental systems

Despite the complexity of natural systems, heterogeneity caused by the fragmentation of habitats has seldom been considered when investigating ecosystem processes. Empirical approaches that have included the influence of heterogeneity tend to be biased towards terrestrial habitats; yet marine systems offer opportunities by virtue of their relative ease of manipulation, rapid response times and the well-understood effects of macrofauna on sediment processes. Here, the influence of heterogeneity on microphytobenthic production in synthetic estuarine assemblages is examined. Heterogeneity was created by enriching patches of sediment with detrital algae (Enteromorpha intestinalis) to provide a source of allochthonous organic matter. A gradient of species density for four numerically dominant intertidal macrofauna (Hediste diversicolor, Hydrobia ulvae, Corophium volutator, Macoma balthica) was constructed, and microphytobenthic biomass at the sediment surface was measured. Statistical analysis using generalized least squares regression indicated that heterogeneity within our system was a significant driving factor that interacted with macrofaunal density and species identity. Microphytobenthic biomass was highest in enriched patches, suggesting that nutrients were obtained locally from the sediment–water interface and not from the water column. Our findings demonstrate that organic enrichment can cause the development of heterogeneity which influences infaunal bioturbation and consequent nutrient generation, a driver of microphytobenthic production.

Impact of biodiversity-climate futures on primary production and metabolism in a model benthic estuarine system

BackgroundUnderstanding the effects of anthropogenically-driven changes in global temperature, atmospheric carbon dioxide and biodiversity on the functionality of marine ecosystems is crucial for predicting and managing the associated impacts. Coastal ecosystems are important sources of carbon (primary production) to shelf waters and play a vital role in global nutrient cycling. These systems are especially vulnerable to the effects of human activities and will be the first areas impacted by rising sea levels. Within these coastal ecosystems, microalgal assemblages (microphytobenthos: MPB) are vital for autochthonous carbon fixation. The level of in situ production by MPB mediates the net carbon cycling of transitional ecosystems between net heterotrophic or autotrophic metabolism. In this study, we examine the interactive effects of elevated atmospheric CO2 concentrations (370, 600, and 1000 ppmv), temperature (6°C, 12°C, and 18°C) and invertebrate biodiversity on MPB biomass in experimental systems. We assembled communities of three common grazing invertebrates (Hydrobia ulvae, Corophium volutator and Hediste diversicolor) in monoculture and in all possible multispecies combinations. This experimental design specifically addresses interactions between the selected climate change variables and any ecological consequences caused by changes in species composition or richness.ResultsThe effects of elevated CO2 concentration, temperature and invertebrate diversity were not additive, rather they interacted to determine MPB biomass, and overall this effect was negative. Diversity effects were underpinned by strong species composition effects, illustrating the importance of individual species identity.ConclusionsOverall, our findings suggest that in natural systems, the complex interactions between changing environmental conditions and any associated changes in invertebrate assemblage structure are likely to reduce MPB biomass. Furthermore, these effects would be sufficient to affect the net metabolic balance of the coastal ecosystem, with important implications for system ecology and sustainable exploitation.

Dirichlet neighbours: revisiting Dirichlet tessellation for neighbourhood analysis

Abstract The analysis of neighbourhood for point features is reviewed in terms of requirements and available techniques; the application of Dirichlet/Voronoi/Thiessen tessellation techniques are discussed. Robin Sibsons Natural Neighbours (Sibson, 1980, Natural neighbourhood interpolation. In V. D. Brunner, Graphical methods for multivariate data . New York: Brunner/Mazel) technique for Dirichlet Tessellation is discussed and benefits demonstrated in the consequently available neighbourhood descriptors of distance, bearing and nearest neighbours. The application of Dirichlet Tessellation derived neighbourhood analysis is shown in terms of the study of badger territories and other potential applications in the sciences and social sciences are outlined.

Geographical variation in the response to nitrogen deposition in Arabidopsis lyrata petraea

The adaptive responses to atmospheric nitrogen deposition for different European accessions of Arabidopsis lyrata petraea were analysed using populations along a strong atmospheric N-deposition gradient. Plants were exposed to three N-deposition rates, reflecting the rates at the different locations, in a full factorial design. Differences between accessions in the response to N were found for important phenological and physiological response variables. For example, plants from low-deposition areas had higher nitrogen-use efficiencies (NUE) and C : N ratios than plants from areas high in N deposition when grown at low N-deposition rates. The NUE decreased in all accessions at higher experimental deposition rates. However, plants from high-deposition areas showed a limited capacity to increase their NUE at lower experimental deposition rates. Plants from low-deposition areas had faster growth rates, higher leaf turnover rates and shorter times to flowering, and showed a greater increase in growth rate in response to N deposition than those from high-deposition areas. Indications for adaptation to N deposition were found, and results suggest that adaptation of plants from areas high in N deposition to increased N deposition has resulted in the loss of plasticity.

Role of functionally dominant species in varying environmental regimes: evidence for the performance-enhancing effect of biodiversity.

BackgroundTheory suggests that biodiversity can act as a buffer against disturbances and environmental variability via two major mechanisms: Firstly, a stabilising effect by decreasing the temporal variance in ecosystem functioning due to compensatory processes; and secondly, a performance enhancing effect by raising the level of community response through the selection of better performing species. Empirical evidence for the stabilizing effect of biodiversity is readily available, whereas experimental confirmation of the performance-enhancing effect of biodiversity is sparse.ResultsHere, we test the effect of different environmental regimes (constant versus fluctuating temperature) on bacterial biodiversity-ecosystem functioning relations. We show that positive effects of species richness on ecosystem functioning are enhanced by stronger temperature fluctuations due to the increased performance of individual species.ConclusionsOur results provide evidence for the performance enhancing effect and suggest that selection towards functionally dominant species is likely to benefit the maintenance of ecosystem functioning under more variable conditions.