Susan K. Wiser

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.

Community structure and forest invasion by an exotic herb over 23 years

We studied the invasion of a New Zealand mountain beech (Nothofagus solandri var. cliffortioides) forest by the exotic perennial herb, Hieracium lepidulum. We used data from 250 randomly located permanent plots (400 m2) established in 1970 that sampled 9000 ha of forest. Frequency of H. lepidulum was 11%, 43%, and 57% in 1970, 1985, and 1993, respectively. For each year of measurement, invasion patterns were related to (a) distance to the forest margin as a measure of dispersal limitation, (b) community structure, (c) stem biomass dynamics indicating disturbance history, and (d) environmental characteristics. In 1970, invaded plots had more species and lower potential solar radiation, and they were closer to the forest margin; however, invaded plots were only weakly predicted by these site variables. H. lepidulum also invaded relatively species-rich subplots (0.75 m2) showing that community structure was also significant at a microsite scale. Using the same sets of variables, the ability to predict which ...

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

Mapping tree density at a global scale

166 billion to

Functional trait space and the latitudinal diversity gradient

490 billion per year. Although representing only a small percentage of the total value of biodiversity, this value is two to six times as much as it would cost to effectively implement conservation globally. These results highlight the necessity to reassess biodiversity valuation and the potential benefits of integrating and promoting biological conservation in forest resource management and forestry practices worldwide. Global effect of tree species diversity on forest productivity. Ground-sourced data from 777,126 global forest biodiversity permanent sample plots (dark blue dots, left), which cover a substantial portion of the global forest extent (white), reveal a consistent positive and concave-down biodiversity-productivity relationship across forests worldwide (red line with pink bands representing 95% confidence interval, right). The biodiversity-productivity relationship (BPR) is foundational to our understanding of the global extinction crisis and its impacts on ecosystem functioning. Understanding BPR is critical for the accurate valuation and effective conservation of biodiversity. Using ground-sourced data from 777,126 permanent plots, spanning 44 countries and most terrestrial biomes, we reveal a globally consistent positive concave-down BPR, showing that continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The value of biodiversity in maintaining commercial forest productivity alone—US

PREDICTION OF RARE-PLANT OCCURRENCE: A SOUTHERN APPALACHIAN EXAMPLE

166 billion to 490 billion per year according to our estimation—is more than twice what it would cost to implement effective global conservation. This highlights the need for a worldwide reassessment of biodiversity values, forest management strategies, and conservation priorities.

High-elevation rock outcrop vegetation of the Southern Appalachian Mountains

The global extent and distribution of forest trees is central to our understanding of the terrestrial biosphere. We provide the first spatially continuous map of forest tree density at a global scale. This map reveals that the global number of trees is approximately 3.04 trillion, an order of magnitude higher than the previous estimate. Of these trees, approximately 1.30 trillion exist in tropical and subtropical forests, with 0.74 trillion in boreal regions and 0.66 trillion in temperate regions. Biome-level trends in tree density demonstrate the importance of climate and topography in controlling local tree densities at finer scales, as well as the overwhelming effect of humans across most of the world. Based on our projected tree densities, we estimate that over 15 billion trees are cut down each year, and the global number of trees has fallen by approximately 46% since the start of human civilization.

IMMEDIATE DAMAGE BY AN EARTHQUAKE TO A TEMPERATE MONTANE FOREST

Significance We present a conceptual framework for testing theories for the latitudinal gradient of species richness in terms of variation in functional diversity at the alpha, beta, and gamma scales. We compared ecological community theory with large-scale observational data of tree species richness and functional diversity. We found that the patterns of functional trait diversity are not consistent with any one theory of species diversity. These conflicting results indicate that none of the broad classes of biodiversity theory considered here is alone able to explain the latitudinal gradient of species diversity in terms of functional trait space. The processes causing the latitudinal gradient in species richness remain elusive. Ecological theories for the origin of biodiversity gradients, such as competitive exclusion, neutral dynamics, and environmental filtering, make predictions for how functional diversity should vary at the alpha (within local assemblages), beta (among assemblages), and gamma (regional pool) scales. We test these predictions by quantifying hypervolumes constructed from functional traits representing major axes of plant strategy variation (specific leaf area, plant height, and seed mass) in tree assemblages spanning the temperate and tropical New World. Alpha-scale trait volume decreases with absolute latitude and is often lower than sampling expectation, consistent with environmental filtering theory. Beta-scale overlap decays with geographic distance fastest in the temperate zone, again consistent with environmental filtering theory. In contrast, gamma-scale trait space shows a hump-shaped relationship with absolute latitude, consistent with no theory. Furthermore, the overall temperate trait hypervolume was larger than the overall tropical hypervolume, indicating that the temperate zone permits a wider range of trait combinations or that niche packing is stronger in the tropical zone. Although there are limitations in the data, our analyses suggest that multiple processes have shaped trait diversity in trees, reflecting no consistent support for any one theory.

Habitat area and climate stability determine geographical variation in plant species range sizes

Ecologically sound efforts to manage or reintroduce populations of rare species require detailed knowledge of species habitat requirements. However, the fact that such species are rare implies that the data needed for habitat characterization are sparse and that species might well be absent from favorable sites due to chance aspects of dispersal or mortality. We use four rare plant species endemic to southern Appalachian high-elevation rock outcrops, to illustrate how nonparametric and parametric logistic regression can yield predictive models of the probability that a species will occur, given certain site conditions. Models were constructed for each species at two scales: 100-m2 plots and 1-m2 subplots. At the 100-m2 plot scale, absences beyond the current geographic range were excluded. At the 1-m2 subplot scale, absences from subplots were only included if the species occurred elsewhere on the 100-m2 plot. Six significant models resulted; no significant model could be constructed for Solidago spithama...

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

Species composition patterns and vegetation-envi- ronment relationships were quantified for high-elevation rock outcrops of the Southern Appalachian Mountains, an infre- quent and insular habitat in a forested landscape. Outcrops occur over a wide geographic range encompassing extensive variation in both geology and climate. Geographic-scale fac- tors interact with site-scale factors to produce variation in vegetation among outcrops. Similarly, site-scale factors inter- act with micro-scale factors to produce variation in vegetation within outcrops. To provide a quantitatively-based classifica- tion of outcrop vegetation we used a TWINSPAN analysis of 154 100-m2 plots. We recognized nine communities that pri- marily correspond to different combinations of elevation, bedrock type, geography, and moisture. Within outcrops of a single bedrock type, vegetation composition of 100-m 2 plots was consistently correlated with elevation and solar radiation, but relationships to soil nutrients varied with bedrock type. Both site-scale (100 m 2 ) factors (e.g. elevation, slope, aspect, and bedrock type) and plot-scale (1-m 2 ) microsite factors (e.g. soil depth, vegetation height, soil nutrients) were strongly correlated with species composition at the 1-m 2 level. Envi- ronment can be used to predict composition more effectively for 100-m 2 plots on a single bedrock type than either across bedrock types or at a 1-m 2 scale. Composition-environment relationships resemble those described for outcrop systems from other regions with pronounced topographic relief more than they do those described for the nearby but flatter and lower-elevation outcrops of the Southeastern Piedmont. There is strong spatial autocorrelation in this community, perhaps owing to dispersal limitation. Consequently, a comprehensive conservation strategy must include reservation of both a range of geologic types and a range of geographic locations.