David A. Coomes

Biodiversity Conservation: Challenges Beyond 2010

Biodiversity Convention In October 2010, the Convention on Biological Diversity will meet to assess the current condition of global biodiversity and to propose and agree on priorities for its future conservation. In this context, Rands et al. (p. 1298; see the News Focus section; see the cover) review recent patterns of biodiversity conservation, highlighting successes, as well as current and future threats. They argue that biodiversity should be treated as a public good, with responsibility for its conservation integrated across sectors of society and government, rather than be confined to the business of environmental agencies, and review the conditions under which this goal might be achieved. The continued growth of human populations and of per capita consumption have resulted in unsustainable exploitation of Earth’s biological diversity, exacerbated by climate change, ocean acidification, and other anthropogenic environmental impacts. We argue that effective conservation of biodiversity is essential for human survival and the maintenance of ecosystem processes. Despite some conservation successes (especially at local scales) and increasing public and government interest in living sustainably, biodiversity continues to decline. Moving beyond 2010, successful conservation approaches need to be reinforced and adequately financed. In addition, however, more radical changes are required that recognize biodiversity as a global public good, that integrate biodiversity conservation into policies and decision frameworks for resource production and consumption, and that focus on wider institutional and societal changes to enable more effective implementation of policy.

Impacts of root competition in forests and woodlands: a theoretical framework and review of experiments.

Light is widely considered to be the most important factor limiting the performance of plants on the floors of forests and woodlands, but the roles of nutrient availability and water supply remain poorly defined. We seek to predict the types of forest in which root competition affects seedling performance, and the types of plants that respond most strongly to release from root competition. We then test our predictions by reviewing experiments in which tree seedlings and forest herbs are released from belowground competition, usually by cutting trenches to sever the roots of surrounding trees. First, we provide a worldwide review of changes in canopy form and fine-root mass along gradients of soil fertility and seasonal drought, keeping in mind the stages of forest development. Our review shows that penetration of light is least in forests on moist soils providing large amounts of major nutrients. The changes are far more complex than those considered by allocation models. Dry woodlands typically allow 20 ...

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.

Rate of tree carbon accumulation increases continuously with tree size

Forests are major components of the global carbon cycle, providing substantial feedback to atmospheric greenhouse gas concentrations. Our ability to understand and predict changes in the forest carbon cycle—particularly net primary productivity and carbon storage—increasingly relies on models that represent biological processes across several scales of biological organization, from tree leaves to forest stands. Yet, despite advances in our understanding of productivity at the scales of leaves and stands, no consensus exists about the nature of productivity at the scale of the individual tree, in part because we lack a broad empirical assessment of whether rates of absolute tree mass growth (and thus carbon accumulation) decrease, remain constant, or increase as trees increase in size and age. Here we present a global analysis of 403 tropical and temperate tree species, showing that for most species mass growth rate increases continuously with tree size. Thus, large, old trees do not act simply as senescent carbon reservoirs but actively fix large amounts of carbon compared to smaller trees; at the extreme, a single big tree can add the same amount of carbon to the forest within a year as is contained in an entire mid-sized tree. The apparent paradoxes of individual tree growth increasing with tree size despite declining leaf-level and stand-level productivity can be explained, respectively, by increases in a tree’s total leaf area that outpace declines in productivity per unit of leaf area and, among other factors, age-related reductions in population density. Our results resolve conflicting assumptions about the nature of tree growth, inform efforts to undertand and model forest carbon dynamics, and have additional implications for theories of resource allocation and plant senescence.

Colonization, tolerance, competition and seed-size variation within functional groups

Abstract Ecologists interested in seed size have generally contrasted functional groups of plants but, recently, some have focussed on explaining the large range of seed size found within a functional group. A potentially important theoretical advance was the idea that seed number–seedling survival tradeoffs could explain the coexistence of species, in particular colonization–competition tradeoffs where smaller-seeded species are superior colonizers and larger-seeded species are superior competitors. However, recent models have placed limits on the potential of this approach, chiefly by showing that the asymmetry of competition must be strong. Also, although there is evidence from studies within functional groups that seed size does trade off against number of seeds and dispersal of those seeds, and that seed size is correlated with competitive ability among seedlings and tolerance of hazards during establishment, the available evidence suggests that SNSS tradeoffs do not make possible long-term coexistence without other forms of niche differentiation.

Global meta-analysis of wood decomposition rates: a role for trait variation among tree species?

The carbon flux from woody debris, a crucial uncertainty within global carbon-climate models, is simultaneously affected by climate, site environment and species-based variation in wood quality. In the first global analysis attempting to explicitly tease out the wood quality contribution to decomposition, we found support for our hypothesis that, under a common climate, interspecific differences in wood traits affect woody debris decomposition patterns. A meta-analysis of 36 studies from all forested continents revealed that nitrogen, phosphorus, and C : N ratio correlate with decomposition rates of angiosperms. In addition, gymnosperm wood consistently decomposes slower than angiosperm wood within common sites, a pattern that correlates with clear divergence in wood traits between the two groups. New empirical studies are needed to test whether this difference is due to a direct effect of wood trait variation on decomposer activity or an indirect effect of wood traits on decomposition microsite environment. The wood trait-decomposition results point to an important role for changes in the wood traits of dominant tree species as a driver of carbon cycling, with likely feedback to atmospheric CO(2) particularly where angiosperm species replace gymnosperms regionally. Truly worldwide upscaling of our results will require further site-based multi-species wood trait and decomposition data, particularly from low-latitude ecosystems.

The effects of sampling bias and model complexity on the predictive performance of MaxEnt species distribution models.

Species distribution models (SDMs) trained on presence-only data are frequently used in ecological research and conservation planning. However, users of SDM software are faced with a variety of options, and it is not always obvious how selecting one option over another will affect model performance. Working with MaxEnt software and with tree fern presence data from New Zealand, we assessed whether (a) choosing to correct for geographical sampling bias and (b) using complex environmental response curves have strong effects on goodness of fit. SDMs were trained on tree fern data, obtained from an online biodiversity data portal, with two sources that differed in size and geographical sampling bias: a small, widely-distributed set of herbarium specimens and a large, spatially clustered set of ecological survey records. We attempted to correct for geographical sampling bias by incorporating sampling bias grids in the SDMs, created from all georeferenced vascular plants in the datasets, and explored model complexity issues by fitting a wide variety of environmental response curves (known as “feature types” in MaxEnt). In each case, goodness of fit was assessed by comparing predicted range maps with tree fern presences and absences using an independent national dataset to validate the SDMs. We found that correcting for geographical sampling bias led to major improvements in goodness of fit, but did not entirely resolve the problem: predictions made with clustered ecological data were inferior to those made with the herbarium dataset, even after sampling bias correction. We also found that the choice of feature type had negligible effects on predictive performance, indicating that simple feature types may be sufficient once sampling bias is accounted for. Our study emphasizes the importance of reducing geographical sampling bias, where possible, in datasets used to train SDMs, and the effectiveness and essentialness of sampling bias correction within MaxEnt.

Angiosperm wood structure: Global patterns in vessel anatomy and their relation to wood density and potential conductivity

Woody stems comprise a large biological carbon fraction and determine water transport between roots and leaves; their structure and function can influence both carbon and hydrological cycles. While angiosperm wood anatomy and density determine hydraulic conductivity and mechanical strength, little is known about interrelations across many species. We compiled a global data set comprising two anatomical traits for 3005 woody angiosperms: mean vessel lumen area (Ā) and number per unit area (N). From these, we calculated vessel lumen fraction (F = ĀN) and size to number ratio (S = Ā/N), a new vessel composition index. We examined the extent to which F and S influenced potential sapwood specific stem conductivity (K(S)) and wood density (D; dry mass/fresh volume). F and S varied essentially independently across angiosperms. Variation in K(S) was driven primarily by S, and variation in D was virtually unrelated to F and S. Tissue density outside vessel lumens (D(N)) must predominantly influence D. High S should confer faster K(S) but incur greater freeze-thaw embolism risk. F should also affect K(S), and both F and D(N) should influence mechanical strength, capacitance, and construction costs. Improved theory and quantification are needed to better understand ecological costs and benefits of these three distinct dimensions.

Microclimate moderates plant responses to macroclimate warming

Significance Around the globe, climate warming is increasing the dominance of warm-adapted species—a process described as “thermophilization.” However, thermophilization often lags behind warming of the climate itself, with some recent studies showing no response at all. Using a unique database of more than 1,400 resurveyed vegetation plots in forests across Europe and North America, we document significant thermophilization of understory vegetation. However, the response to macroclimate warming was attenuated in forests whose canopies have become denser. This microclimatic effect likely reflects cooler forest-floor temperatures via increased shading during the growing season in denser forests. Because standing stocks of trees have increased in many temperate forests in recent decades, microclimate may commonly buffer understory plant responses to macroclimate warming. Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., “thermophilization” of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that “climatic lags” may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12–67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass—e.g., for bioenergy—may open forest canopies and accelerate thermophilization of temperate forest biodiversity.

Positive biodiversity-productivity relationship predominant in global forests.

Global biodiversity and productivity The relationship between biodiversity and ecosystem productivity has been explored in detail in herbaceous vegetation, but patterns in forests are far less well understood. Liang et al. have amassed a global forest data set from >770,000 sample plots in 44 countries. A positive and consistent relationship can be discerned between tree diversity and ecosystem productivity at landscape, country, and ecoregion scales. On average, a 10% loss in biodiversity leads to a 3% loss in productivity. This means that the economic value of maintaining biodiversity for the sake of global forest productivity is more than fivefold greater than global conservation costs. Science, this issue p. 196 Global forest inventory records suggest that biodiversity loss would result in a decline in forest productivity worldwide. INTRODUCTION The biodiversity-productivity relationship (BPR; the effect of biodiversity on ecosystem productivity) is foundational to our understanding of the global extinction crisis and its impacts on the functioning of natural ecosystems. The BPR has been a prominent research topic within ecology in recent decades, but it is only recently that we have begun to develop a global perspective. RATIONALE Forests are the most important global repositories of terrestrial biodiversity, but deforestation, forest degradation, climate change, and other factors are threatening approximately one half of tree species worldwide. Although there have been substantial efforts to strengthen the preservation and sustainable use of forest biodiversity throughout the globe, the consequences of this diversity loss pose a major uncertainty for ongoing international forest management and conservation efforts. The forest BPR represents a critical missing link for accurate valuation of global biodiversity and successful integration of biological conservation and socioeconomic development. Until now, there have been limited tree-based diversity experiments, and the forest BPR has only been explored within regional-scale observational studies. Thus, the strength and spatial variability of this relationship remains unexplored at a global scale. RESULTS We explored the effect of tree species richness on tree volume productivity at the global scale using repeated forest inventories from 777,126 permanent sample plots in 44 countries containing more than 30 million trees from 8737 species spanning most of the global terrestrial biomes. Our findings reveal a consistent positive concave-down effect of biodiversity on forest productivity across the world, showing that a continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The BPR shows considerable geospatial variation across the world. The same percentage of biodiversity loss would lead to a greater relative (that is, percentage) productivity decline in the boreal forests of North America, Northeastern Europe, Central Siberia, East Asia, and scattered regions of South-central Africa and South-central Asia. In the Amazon, West and Southeastern Africa, Southern China, Myanmar, Nepal, and the Malay Archipelago, however, the same percentage of biodiversity loss would lead to greater absolute productivity decline. CONCLUSION Our findings highlight the negative effect of biodiversity loss on forest productivity and the potential benefits from the transition of monocultures to mixed-species stands in forestry practices. The BPR we discover across forest ecosystems worldwide corresponds well with recent theoretical advances, as well as with experimental and observational studies on forest and nonforest ecosystems. On the basis of this relationship, the ongoing species loss in forest ecosystems worldwide could substantially reduce forest productivity and thereby forest carbon absorption rate to compromise the global forest carbon sink. We further estimate that the economic value of biodiversity in maintaining commercial forest productivity alone is