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Alan R. Townsend

Publications and source records attributed to Alan R. Townsend.

9 recordsLinked to original sources

Spatially robust estimates of biological nitrogen (N) fixation imply substantial human alteration of the tropical N cycle

Biological nitrogen fixation (BNF) is the largest natural source of exogenous nitrogen (N) to unmanaged ecosystems and also the primary baseline against which anthropogenic changes to the N cycle are measured. Rates of BNF in tropical rainforest are thought to be among the highest on Earth, but they are notoriously difficult to quantify and are based on little empirical data. We adapted a sampling strategy from community ecology to generate spatial estimates of symbiotic and free-living BNF in secondary and primary forest sites that span a typical range of tropical forest legume abundance. Although total BNF was higher in secondary than primary forest, overall rates were roughly five times lower than previous estimates for the tropical forest biome. We found strong correlations between symbiotic BNF and legume abundance, but we also show that spatially free-living BNF often exceeds symbiotic inputs. Our results suggest that BNF in tropical forest has been overestimated, and our data are consistent with a recent top-down estimate of global BNF that implied but did not measure low tropical BNF rates. Finally, comparing tropical BNF within the historical area of tropical rainforest with current anthropogenic N inputs indicates that humans have already at least doubled reactive N inputs to the tropical forest biome, a far greater change than previously thought. Because N inputs are increasing faster in the tropics than anywhere on Earth, both the proportion and the effects of human N enrichment are likely to grow in the future.

Proceedings of the National Academy of Sciences of

Effects of canopy tree species on belowground biogeochemistry in a lowland wet tropical forest

Tropical rain forests are known for their high biological diversity, but the effects of plant diversity on important ecosystem processes in this biome remain unclear. Interspecies differences in both the demand for nutrients and in foliar and litter nutrient concentrations could drive variations in both the pool sizes and fluxes of important belowground resources, yet our understanding of the effects and importance of aboveground heterogeneity on belowground biogeochemistry is poor, especially in the species-rich forests of the wet tropics. To investigate the effects of individual tree species on belowground biogeochemical processes, we used both field and laboratory studies to examine how carbon (C), nitrogen (N), and phosphorus (P) cycles vary under nine different canopy tree species – including three legume and six non-legume species – that vary in foliar nutrient concentrations in a wet tropical forest in southwestern Costa Rica. We found significant differences in belowground C, N and P cycling under different canopy tree species: total C, N and P pools in standing litter varied by species, as did total soil and microbial C and N pools. Rates of soil extracellular acid phosphatase activity also varied significantly among species and functional groups, with higher rates of phosphatase activity under legumes. In addition, across all tree species, phosphatase activity was significantly positively correlated with litter N/P ratios, suggesting a tight coupling between relative N and P inputs and resource allocation to P acquisition. Overall, our results suggest the importance of aboveground plant community composition in promoting belowground biogeochemical heterogeneity at relatively small spatial scales.

Soil Biology and Biochemistry

Accumulation of atmospheric sulfur in some Costa Rican soils

Sulfur is one of the macronutrient elements whose sources to terrestrial ecosystems should shift from dominance by rock-weathering to atmospheric deposition as soils and underlying substrate undergo progressive weathering and leaching. However, the nature and timing of this transition is not well known. We investigated sources of sulfur to tropical rain forests growing on basalt-derived soils in the Osa Peninsula region of Costa Rica. Sulfur sources were examined using stable isotope ratios (δ 34 S) and compared to chemical indices of soil development. The most weathered soils, and the forests they supported, are dominated by atmospheric sulfur, while a less weathered soil type contains both rock-derived and atmospheric sulfur. Patterns of increasing δ 34 S with increasing soil sulfur concentration across the landscape suggest atmospheric sulfur is accumulating, and little rock-derived sulfur has been retained. Soil sulfur, minus adsorbed sulfate, is correlated with carbon and nitrogen, implying that sulfur accumulation occurs as plants and microbes incorporate sulfur into organic matter. Only the lower depth increments of the more weathered soils contained significant adsorbed sulfate. The evidence suggests a pattern of soil development in which sulfur-bearing minerals in rock, such as sulfides, weather early relative to other minerals, and the released sulfate is leached away. Sulfur added via atmospheric deposition is retained as organic matter accumulates in the soil profile. Adsorbed sulfate accumulates later, driven by changes in soil chemistry and mineralogy. These aspects of sulfur behavior during pedogenesis in this environment may hasten the transition to dominance by atmospheric sources.

Journal of Geophysical Research: Biogeosciences

Stoichiometric patterns in foliar nutrient resorption across multiple scales

*Nutrient resorption is a fundamental process through which plants withdraw nutrients from leaves before abscission. Nutrient resorption patterns have the potential to reflect gradients in plant nutrient limitation and to affect a suite of terrestrial ecosystem functions. *Here, we used a stoichiometric approach to assess patterns in foliar resorption at a variety of scales, specifically exploring how N : P resorption ratios relate to presumed variation in N and/or P limitation and possible relationships between N : P resorption ratios and soil nutrient availability. *N : P resorption ratios varied significantly at the global scale, increasing with latitude and decreasing with mean annual temperature and precipitation. In general, tropical sites (absolute latitudes < 23°26′) had N : P resorption ratios of < 1, and plants growing on highly weathered tropical soils maintained the lowest N : P resorption ratios. Resorption ratios also varied with forest age along an Amazonian forest regeneration chronosequence and among species in a diverse Costa Rican rain forest. *These results suggest that variations in N : P resorption stoichiometry offer insight into nutrient cycling and limitation at a variety of spatial scales, complementing other metrics of plant nutrient biogeochemistry. The extent to which the stoichiometric flexibility of resorption will help regulate terrestrial responses to global change merits further investigation.

New Phytologist

Relationships among net primary productivity, nutrients and climate in tropical rain forest: A pan-tropical analysis

Tropical rain forests play a dominant role in global biosphere-atmosphere CO 2 exchange. Although climate and nutrient availability regulate net primary production (NPP) and decomposition in all terrestrial ecosystems, the nature and extent of such controls in tropical forests remain poorly resolved. We conducted a meta-analysis of carbon-nutrient-climate relationships in 113 sites across the tropical forest biome. Our analyses showed that mean annual temperature was the strongest predictor of aboveground NPP (ANPP) across all tropical forests, but this relationship was driven by distinct temperature differences between upland and lowland forests. Within lowland forests (< 1000 m), a regression tree analysis revealed that foliar and soil-based measurements of phosphorus (P) were the only variables that explained a significant proportion of the variation in ANPP, although the relationships were weak. However, foliar P, foliar nitrogen (N), litter decomposition rate (k), soil N and soil respiration were all directly related with total surface (0–10 cm) soil P concentrations. Our analysis provides some evidence that P availability regulates NPP and other ecosystem processes in lowland tropical forests, but more importantly, underscores the need for a series of large-scale nutrient manipulations – especially in lowland forests – to elucidate the most important nutrient interactions and controls.

Ecology Letters

Functional ecology of free-living nitrogen fixation: A contemporary perspective

Nitrogen (N) availability is thought to frequently limit terrestrial ecosystem processes, and explicit consideration of N biogeochemistry, including biological N 2 fixation, is central to understanding ecosystem responses to environmental change. Yet, the importance of free-living N 2 fixation—a process that occurs on a wide variety of substrates, is nearly ubiquitous in terrestrial ecosystems, and may often represent the dominant pathway for acquiring newly available N—is often underappreciated. Here, we draw from studies that investigate free-living N 2 fixation from functional, physiological, genetic, and ecological perspectives. We show that recent research and analytical advances have generated a wealth of new information that provides novel insight into the ecology of N 2 fixation as well as raises new questions and priorities for future work. These priorities include a need to better integrate free-living N 2 fixation into conceptual and analytical evaluations of the N cycle's role in a variety of global change scenarios.

Annual Review of Ecology, Evolution, and Systemati

Experimental drought in a tropical rain forest increases soil carbon dioxide losses to the atmosphere

Climate models predict precipitation changes for much of the humid tropics, yet few studies have investigated the potential consequences of drought on soil carbon (C) cycling in this important biome. In wet tropical forests, drought could stimulate soil respiration via overall reductions in soil anoxia, but previous research suggests that litter decomposition is positively correlated with high rainfall fluxes that move large quantities of dissolved organic matter (DOM) from the litter layer to the soil surface. Thus, reduced rainfall could also limit C delivery to the soil surface, reducing respiration rates. We conducted a throughfall manipulation experiment to investigate how 25% and 50% reductions in rainfall altered both C movement into soils and the effects of those DOM fluxes on soil respiration rates. In response to the experimental drought, soil respiration rates increased in both the −25% and −50% treatments. Throughfall fluxes were reduced by 26% and 55% in the −25% and −50% treatments, respectively. However, total DOM fluxes leached from the litter did not vary between treatments, because the concentrations of leached DOM reaching the soil surface increased in response to the simulated drought. Annual DOM concentrations averaged 7.7 ± 0.8, 11.2 ± 0.9, and 15.8 ± 1.2 mg C/L in the control, −25%, and −50% plots, respectively, and DOM concentrations were positively correlated with soil respiration rates. A laboratory incubation experiment confirmed the potential importance of DOM concentration on soil respiration rates, suggesting that this mechanism could contribute to the increase in CO 2 fluxes observed in the reduced rainfall plots. Across all plots, the data suggested that soil CO 2 fluxes were partially regulated by the magnitude and concentration of soluble C delivered to the soil, but also by soil moisture and soil oxygen availability. Together, our data suggest that declines in precipitation in tropical rain forests could drive higher CO 2 fluxes to the atmosphere both via increased soil O 2 availability and through responses to elevated DOM concentrations.

Golfo Dulce Forest Reserve

Ecological science and sustainability for the 21st century

Ecological science has contributed greatly to our understanding of the natural world and the impact of humans on that world. Now, we need to refocus the discipline towards research that ensures a future in which natural systems and the humans they include coexist on a more sustainable planet. Acknowledging that managed ecosystems and intensive exploitation of resources define our future, ecologists must play a greatly expanded role in communicating their research and influencing policy and decisions that affect the environment. To accomplish this, they will have to forge partnerships at scales and in forms they have not traditionally used. These alliances must act within three visionary areas: enhancing the extent to which decisions are ecologically informed; advancing innovative ecological research directed at the sustainability of the planet; and stimulating cultural changes within the science itself, thereby building a forward-looking and international ecology. We recommend: (1) a research initiative to enhance research project development, facilitate large-scale experiments and data collection, and link science to solutions; (2) procedures that will improve interactions among researchers, managers, and decision makers; and (3) efforts to build public understanding of the links between ecosystem services and humans.

Frontiers in Ecology and the Environment

Ecology for a crowded planet

Within the next 50 to 100 years, the support and maintenance of an extended human family of 8 to 11 billion people will be difficult at best. The authors of this Policy Forum describe changes that are required if we hope to meet the needs and aspirations of humans while improving the health of our planet's ecosystems. Problems as diverse as disease transmission and global climate change have benefited substantially from advances in ecology. Such advances have set the stage for emergence of a proactive ecological science in which social and political realities are acknowledged and attention is turned decisively toward the future. The ecological sciences must chart an understanding of how ecosystem services can persist given their extensive human use. Innovative research on the sciences of ecosystem services, ecological restoration, and ecological design must be massively accelerated and must be accompanied by more effective communication of ecological knowledge to society.

Science