Benjamin W. Sullivan

Carbon and water fluxes from ponderosa pine forests disturbed by wildfire and thinning

Disturbances alter ecosystem carbon dynamics, often by reducing carbon uptake and stocks. We compared the impact of two types of disturbances that represent the most likely future conditions of currently dense ponderosa pine forests of the southwestern United States: (1) high-intensity fire and (2) thinning, designed to reduce fire intensity. High-severity fire had a larger impact on ecosystem carbon uptake and storage than thinning. Total ecosystem carbon was 42% lower at the intensely burned site, 10 years after burning, than at the undisturbed site. Eddy covariance measurements over two years showed that the burned site was a net annual source of carbon to the atmosphere whereas the undisturbed site was a sink. Net primary production (NPP), evapotranspiration (ET), and water use efficiency were lower at the burned site than at the undisturbed site. In contrast, thinning decreased total ecosystem carbon by 18%, and changed the site from a carbon sink to a source in the first posttreatment year. Thinning also decreased ET, reduced the limitation of drought on carbon uptake during summer, and did not change water use efficiency. Both disturbances reduced ecosystem carbon uptake by decreasing gross primary production (55% by burning, 30% by thinning) more than total ecosystem respiration (TER; 33-47% by burning, 18% by thinning), and increased the contribution of soil carbon dioxide efflux to TER. The relationship between TER and temperature was not affected by either disturbance. Efforts to accurately estimate regional carbon budgets should consider impacts on carbon dynamics of both large disturbances, such as high-intensity fire, and the partial disturbance of thinning that is often used to prevent intense burning. Our results show that thinned forests of ponderosa pine in the southwestern United States are a desirable alternative to intensively burned forests to maintain carbon stocks and primary production.

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

Significance Biological nitrogen fixation (BNF) is the largest natural source of new nitrogen (N) to terrestrial ecosystems. Tropical forest ecosystems are a putative global hotspot of BNF, but direct, spatially explicit measurements in the biome are virtually nonexistent. Nonetheless, robust estimates of tropical forest BNF are critical for understanding how these important ecosystems may respond to global change and assessing human perturbations to the N cycle. Here, we introduce a spatial sampling method to assess BNF and present evidence that tropical forest BNF is much lower than previously assumed. Our results imply that humans have roughly doubled N inputs to the tropical forest biome relative to N inputs through BNF. 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.

Interactions among nitrogen fixation and soil phosphorus acquisition strategies in lowland tropical rain forests.

Paradoxically, symbiotic dinitrogen (N2 ) fixers are abundant in nitrogen (N)-rich, phosphorus (P)-poor lowland tropical rain forests. One hypothesis to explain this pattern states that N2 fixers have an advantage in acquiring soil P by producing more N-rich enzymes (phosphatases) that mineralise organic P than non-N2 fixers. We assessed soil and root phosphatase activity between fixers and non-fixers in two lowland tropical rain forest sites, but also addressed the hypothesis that arbuscular mycorrhizal (AM) colonisation (another P acquisition strategy) is greater on fixers than non-fixers. Root phosphatase activity and AM colonisation were higher for fixers than non-fixers, and strong correlations between AM colonisation and N2 fixation at both sites suggest that the N-P interactions mediated by fixers may generally apply across tropical forests. We suggest that phosphatase enzymes and AM fungi enhance the capacity of N2 fixers to acquire soil P, thus contributing to their high abundance in tropical forests.

A comparison of plot‐based satellite and Earth system model estimates of tropical forest net primary production

Net primary production (NPP) by plants represents the largest annual flux of carbon dioxide (CO2) from the atmosphere to the terrestrial biosphere, playing a critical role in the global carbon (C) cycle and the Earths climate. Rates of NPP in tropical forests are thought to be among the highest on Earth, but debates about the magnitude, patterns, and controls of NPP in the tropics highlight uncertainty in our understanding of how tropical forests may respond to environmental change. Here, we compared tropical NPP estimates generated using three common approaches: (1) field-based methods scaled from plot-level measurements of plant biomass, (2) radiation-based methods that model NPP from satellite-derived radiation absorption by plants, (3) and biogeochemical model-based methods. For undisturbed tropical forests as a whole, the three methods produced similar NPP estimates (i.e., ~ 10 Pg C yr−1). However, the three different approaches produced vastly different patterns of NPP both in space and through time, suggesting that our understanding of tropical NPP is poor and that our ability to predict the response of NPP in the tropics to environmental change is limited. To address this shortcoming, we suggest the development of an expanded, high-density, permanent network of sites where NPP is continuously evaluated using multiple approaches. Well-designed NPP megatransects that include a high-density plot network would significantly increase the accuracy and certainty in the observed rates and patterns of tropical NPP and improve the reliability of Earth system models used to predict NPP–carbon cycle–climate interactions into the future.

Assessing nutrient limitation in complex forested ecosystems: alternatives to large-scale fertilization experiments

Quantifying nutrient limitation of primary productivity is a fundamental task of terrestrial ecosystem ecology, but in a high carbon dioxide environment it is even more critical that we understand potential nutrient constraints on plant growth. Ecologists often manipulate nutrients with fertilizer to assess nutrient limitation, yet for a variety of reasons, nutrient fertilization experiments are either impractical or incapable of resolving ecosystem responses to some global changes. The challenges of conducting large, in situ fertilization experiments are magnified in forests, especially the high-diversity forests common throughout the lowland tropics. A number of methods, including fertilization experiments, could be seen as tools in a toolbox that ecologists may use to attempt to assess nutrient limitation, but there has been no compilation or synthetic discussion of those methods in the literature. Here, we group these methods into one of three categories (indicators of soil nutrient supply, organismal indicators of nutrient limitation, and lab-based experiments and nutrient depletions), and discuss some of the strengths and limitations of each. Next, using a case study, we compare nutrient limitation assessed using these methods to results obtained using large-scale fertilizations across the Hawaiian Archipelago. We then explore the application of these methods in high-diversity tropical forests. In the end, we suggest that, although no single method is likely to predict nutrient limitation in all ecosystems and at all scales, by simultaneously utilizing a number of the methods we describe, investigators may begin to understand nutrient limitation in complex and diverse ecosystems such as tropical forests. In combination, these methods represent our best hope for understanding nutrient constraints on the global carbon cycle, especially in tropical forest ecosystems.

Does dissolved organic carbon regulate biological methane oxidation in semiarid soils

In humid ecosystems, the rate of methane (CH4 ) oxidation by soil-dwelling methane-oxidizing bacteria (MOB) is controlled by soil texture and soil water holding capacity, both of which limit the diffusion of atmospheric CH4 into the soil. However, it remains unclear whether these same mechanisms control CH4 oxidation in more arid soils. This study was designed to measure the proximate controls of potential CH4 oxidation in semiarid soils during different seasons. Using a unique and well-constrained 3-million-year-old semiarid substrate age gradient, we were able to hold state factors constant while exploring the relationship between seasonal potential CH4 oxidation rates and soil texture, soil water holding capacity, and dissolved organic carbon (DOC). We measured unexpectedly higher rates of potential CH4 oxidation in the wet season than the dry season. Although other studies have attributed low CH4 oxidation rates in dry soils to desiccation of MOB, we present several lines of evidence that this may be inaccurate. We found that soil DOC concentration explained CH4 oxidation rates better than soil physical factors that regulate the diffusion of CH4 from the atmosphere into the soil. We show evidence that MOB facultatively incorporated isotopically labeled glucose into their cells, and MOB utilized glucose in a pattern among our study sites that was similar to wet-season CH4 oxidation rates. This evidence suggests that DOC, which is utilized by MOB in other environments with varying effects on CH4 oxidation rates, may be an important regulator of CH4 oxidation rates in semiarid soils. Our collective understanding of the facultative use of DOC by MOB is still in its infancy, but our results suggest it may be an important factor controlling CH4 oxidation in soils from dry ecosystems.

Pinyon pine (Pinus edulis) mortality and response to water addition across a three million year substrate age gradient in northern Arizona, USA

Background and aimsPinyon pine (Pinus edulis Engelm.) is an important tree species in the western United States that has experienced large-scale mortality during recent severe drought. The influence of soil conditions on pinyon pine response to water availability is poorly understood. We investigated patterns of tree mortality and response of tree water relations and growth to experimental water addition at four sites across a three million year soil-substrate age gradient.MethodsWe measured recent pinyon mortality at four sites, and tree predawn water potential, leaf carbon isotope signature, and branch, leaf, and stem radial growth on 12 watered and unwatered trees at each site. Watered trees recieved fifty percent more than growing season precipitation for 6 years.ResultsSubstrate age generally had a greater effect on tree water stress and growth than water additions. Pinyon mortality was higher on intermediate-aged substrates (50–55%) than on young (15%) and old (17%) substrates, and mortality was positively correlated with pinyon abundance prior to drought.ConclusionsThese results suggest high soil resource availability and consequent high stand densities at intermediate-age substrates predisposes trees to drought-induced mortality in semi-arid regions. The response of tree water relations to water addition was consistent with the inverse texture hypothesis; watering reduced tree water stress most in young, coarsely textured soil, likely because water rapidly penetrated deep in the soil profile where it was protected from evapotranspiration.

Pulse Emissions of Carbon Dioxide during Snowmelt at a High-Elevation Site in Northern Arizona, U.S.A.

Abstract The paradigm that winter is a dormant period of soil biogeochemical activity in high elevation or high latitude ecosystems has been amply refuted by recent research. Carbon dioxide (CO2) released from cold or snow-covered soil is a substantial component of total annual ecosystem carbon fluxes. Recent investigations have shown that the late-winter/early-spring transition is a period of high biogeochemical activity. However, little is known about the temporal dynamics of CO2 from the snowpack itself during periods of snowmelt. We present a case study of three snowmelt events at a high-elevation site in northern Arizona during which we measured changes in CO2 concentrations and fluxes above the snow and within the soil profile, and characterized the soil physical environment and site meteorological variables. We show that large pulses of CO2 were emitted to the atmosphere during snowmelt, and we present evidence that these pulses came from CO2 stored in snowpack. Earlier and more frequent snowmelts worldwide caused by climate change have the potential to alter the timing of release of CO2 from land to atmosphere.

What is the relationship between soil methane oxidation and other C compounds

What is the relationship between soil methane oxidation and other C compounds? BEN JAMIN W . SULL IVAN a , PAUL C . S ELMANTS 2 and STEPHEN C. HART School of Forestry, Northern Arizona University, 200 E. Pine Knoll Drive, Flagstaff, AZ 86011, USA, Department of Natural Resources and Environmental Management, University of Hawai’I at Manoa, Honolulu, HI 96822, USA, School of Natural Sciences & Sierra Nevada Research Institute, University of California, 5200 North Lake Road, Merced, CA 95343, USA

Nutrient limitation of soil microbial activity during the earliest stages of ecosystem development

A dominant paradigm in ecology is that plants are limited by nitrogen (N) during primary succession. Whether generalizable patterns of nutrient limitation are also applicable to metabolically and phylogenetically diverse soil microbial communities, however, is not well understood. We investigated if measures of N and phosphorus (P) pools inform our understanding of the nutrient(s) most limiting to soil microbial community activities during primary succession. We evaluated soil biogeochemical properties and microbial processes using two complementary methodological approaches—a nutrient addition microcosm experiment and extracellular enzyme assays—to assess microbial nutrient limitation across three actively retreating glacial chronosequences. Microbial respiratory responses in the microcosm experiment provided evidence for N, P and N/P co-limitation at Easton Glacier, Washington, USA, Puca Glacier, Peru, and Mendenhall Glacier, Alaska, USA, respectively, and patterns of nutrient limitation generally reflected site-level differences in soil nutrient availability. The activities of three key extracellular enzymes known to vary with soil N and P availability developed in broadly similar ways among sites, increasing with succession and consistently correlating with changes in soil total N pools. Together, our findings demonstrate that during the earliest stages of soil development, microbial nutrient limitation and activity generally reflect soil nutrient supply, a result that is broadly consistent with biogeochemical theory.