Sean C. Thomas

The worldwide leaf economics spectrum

Bringing together leaf trait data spanning 2,548 species and 175 sites we describe, for the first time at global scale, a universal spectrum of leaf economics consisting of key chemical, structural and physiological properties. The spectrum runs from quick to slow return on investments of nutrients and dry mass in leaves, and operates largely independently of growth form, plant functional type or biome. Categories along the spectrum would, in general, describe leaf economic variation at the global scale better than plant functional types, because functional types overlap substantially in their leaf traits. Overall, modulation of leaf traits and trait relationships by climate is surprisingly modest, although some striking and significant patterns can be seen. Reliable quantification of the leaf economics spectrum and its interaction with climate will prove valuable for modelling nutrient fluxes and vegetation boundaries under changing land-use and climate.

Increasing carbon storage in intact African tropical forests

The response of terrestrial vegetation to a globally changing environment is central to predictions of future levels of atmospheric carbon dioxide. The role of tropical forests is critical because they are carbon-dense and highly productive. Inventory plots across Amazonia show that old-growth forests have increased in carbon storage over recent decades, but the response of one-third of the world’s tropical forests in Africa is largely unknown owing to an absence of spatially extensive observation networks. Here we report data from a ten-country network of long-term monitoring plots in African tropical forests. We find that across 79 plots (163 ha) above-ground carbon storage in live trees increased by 0.63 Mg C ha-1 yr-1 between 1968 and 2007 (95% confidence interval (CI), 0.22–0.94; mean interval, 1987–96). Extrapolation to unmeasured forest components (live roots, small trees, necromass) and scaling to the continent implies a total increase in carbon storage in African tropical forest trees of 0.34 Pg C yr-1 (CI, 0.15–0.43). These reported changes in carbon storage are similar to those reported for Amazonian forests per unit area, providing evidence that increasing carbon storage in old-growth forests is a pan-tropical phenomenon. Indeed, combining all standardized inventory data from this study and from tropical America and Asia together yields a comparable figure of 0.49 Mg C ha-1 yr-1 (n = 156; 562 ha; CI, 0.29–0.66; mean interval, 1987–97). This indicates a carbon sink of 1.3 Pg C yr-1 (CI, 0.8–1.6) across all tropical forests during recent decades. Taxon-specific analyses of African inventory and other data suggest that widespread changes in resource availability, such as increasing atmospheric carbon dioxide concentrations, may be the cause of the increase in carbon stocks, as some theory and models predict.

Competition and Allometry in Three Species of Annual Plants

Comparisons between competing and noncompeting populations of three annual plant species demonstrate that plant allometry is altered by competition. When plants are grown in isolation, relationships between stem diameter, height, and plant mass generally show simple allometry (i.e., the relationships are linear on log-log scale). When plants are competing, however, these relationships are curvilinear or discontinuous. We attempt to clarify the relationship between interindividual allometry (the usual source of allometric data) and allometric growth. When plants are not competing, the allometric relationships among individuals of different sizes at one point in time and the allometry of individual plants as they grow appear to be similar, but these two classes of allometric relationships are very different for competing plants. We present a simple model that explains both static and dynamic patterns of plant allometry in terms of (a) the allometric responses of individual plants to competition, and (b) the size dependence of growth after the onset of competition. Our results illustrate the importance of reciprocal interactions between competition and allometry, and emphasize the difficulty in making inferences from one size measure to another. We conclude that the commonly held assumption that plant size is a single entity, which can be reflected by any of several measures, may not be justified.

The Importance of Demographic Niches to Tree Diversity

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

Assessing Evidence for a Pervasive Alteration in Tropical Tree Communities

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.

A Reassessment of Carbon Content in Tropical Trees

Accurate knowledge of carbon (C) content in live wood is essential for quantifying tropical forest C stocks, yet generic assumptions (such as biomass consisting of 50% carbon on a weight/weight basis) remain widely used despite being supported by little chemical analysis. Empirical data from stem cores of 59 Panamanian rainforest tree species demonstrate that wood C content is highly variable among co-occurring species, with an average (47.4±2.51% S.D.) significantly lower than widely assumed values. Prior published values have neglected to account for volatile C content of tropical woods. By comparing freeze- and oven-dried wood samples, we show that volatile C is non-negligible, and excluding the volatile fraction underestimates wood C content by 2.48±1.28% (S.D.) on average. Wood C content varied substantially among species (from 41.9–51.6%), but was neither strongly phylogenetically conserved, nor correlated to ecological (i.e. wood density, maximum tree height) or demographic traits (i.e. relative growth rate, mortality rate). Overall, assuming generic C fractions in tropical wood overestimates forest C stocks by ∼3.3–5.3%, a non-trivial margin of error leading to overestimates of 4.1–6.8 Mg C ha−1 in a 50-ha forest dynamics plot on Barro Colorado Island, Panama. In addition to addressing other sources of error in tropical forest C accounting, such as uncertainties in allometric models and belowground biomass, compilation and use of species-specific C fractions for tropical tree species would substantially improve both local and global estimates of terrestrial C stocks and fluxes.

Carbon Dioxide Exchange Between an Old-growth Forest and the Atmosphere

Eddy-covariance and biometeorological methods show significant net annual carbon uptake in an old-growth Douglas-fir forest in southwestern Washington, USA. These results contrast with previous assumptions that old-growth forest ecosystems are in carbon equilibrium. The basis for differences between conventional biomass-based carbon sequestration estimates and the biometeorologic estimates are discussed. Annual net ecosystem exchange was comparable to younger ecosystems at the same latitude, as quantified in the AmeriFlux program. Net ecosystem carbon uptake was significantly correlated with photosynthetically active radiation and air temperature, as well as soil moisture and precipitation. Optimum ecosystem photosynthesis occurred at relatively cool temperatures (5°–10°C). Understory and soil carbon exchange always represented a source of carbon to the atmosphere, with a strong seasonal cycle in source strength. Understory and soil carbon exchange showed a Q10 temperature dependence and represented a substantial portion of the ecosystem carbon budget. The period of main carbon uptake and the period of soil and ecosystem respiration are out of phase, however, and driven by different climatic boundary conditions. The period of strongest ecosystem carbon uptake coincides with the lowest observed values of soil and ecosystem respiration. Despite the substantial contribution of soil, the overall strength of the photosynthetic sink resulted in the net annual uptake. The net uptake estimates here included two correction methods, one for advection and the other for low levels of turbulence.

Three-dimensional Structure of an Old-growth Pseudotsuga-Tsuga Canopy and Its Implications for Radiation Balance, Microclimate, and Gas Exchange

We describe the three-dimensional structure of an old-growth Douglas-fir/western hemlock forest in the central Cascades of southern Washington, USA. We concentrate on the vertical distribution of foliage, crowns, external surface area, wood biomass, and several components of canopy volume. In addition, we estimate the spatial variation of some aspects of structure, including the topography of the outer surface, and of microclimate, including the within-canopy transmittance of photosynthetically active radiation (PAR). The crowns of large stems, especially of Douglas-fir, dominate the structure and many aspects of spatial variation. The mean vertical profile of canopy surfaces, estimated by five methods, generally showed a single maximum in the lower to middle third of the canopy, although the height of that maximum varied by method. The stand leaf area index was around 9 m2 m−2, but also varied according to method (from 6.3 to 12.3). Because of the deep narrow crowns and numerous gaps, the outer canopy surface is extremely complex, with a surface area more than 12 times that of the ground below. The large volume included below the outer canopy surface is very porous, with spaces of several qualitatively distinct environments. Our measurements are consistent with emerging concepts about the structure of old-growth forests, where a high degree of complexity is generated by diverse structural features. These structural characteristics have implications for various ecosystem functions. The height and large volume of the stand indicate a large storage component for microclimatic variables. The high biomass influences the dynamics of those variables, retarding rates of change. The complexity of the canopy outer surface influences radiation balance, particularly in reducing short-wave reflectance. The bottom-heaviness of the foliage profile indicates much radiation absorption and gas exchange activity in the lower canopy. The high porosity contributes to flat gradients of most microclimate variables. Most stand respiration occurs within the canopy and is distributed over a broad vertical range.

Scale‐dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

Relative size at onset of maturity in rain forest trees : a comparative analysis of 37 Malaysian species

Relative size at onset of maturity (RSOM), defined as the size at reproductive onset divided by asymptotic maximal size, has been found to be remarkably constant among species in many animal taxa that display indeterminate growth. The present study extends this general approach to plants, examining 37 species of Malaysian rain forest trees. Most of these species exhibit pronounced reproductive size thresholds; however, in contrast to previous studies, values of RSOM vary substantially among closely related species. The relationship among species between height at reproductive onset and asymptotic maximal height deviates significantly from linearity: larger tree species display larger reproductive size thresholds on both an absolute and a relative basis. Larger tree species also generally display more distinct size thresholds, as quantified by the exponent parameter of a modified logistic regression function describing probability of reproduction as a function of size, than do smaller tree species. Statistically significant values of RSOM (on a height basis) vary among species from 0.20 to 0.75, with a mean of 0.42. Differences in RSOM among genera were not significant; however, there was some evidence that species associated with riparian areas have higher average values for RSOM than non-riparian species. Compared with other organisms that have been examined, tropical tree species exhibit a remarkable range of values for RSOM. Such variation has important implications to theories explaining canopy stratification, life-history variation, and niche differentiation in tropical forests.