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Diversity and dispersal interactively affect predictability of ecosystem function.

Theory and small-scale experiments predict that biodiversity losses can decrease the magnitude and stability of ecosystem services such as production and nutrient cycling. Most of this research, however, has been isolated from the immigration and emigration (dispersal) processes that create and maintain diversity in nature. As common anthropogenic drivers of biodiversity change--such as habitat fragmentation, species introductions and climate change--are mediated by these understudied processes, it is unclear how environmental degradation will affect ecosystem services. Here we tested the interactive effects of mobile grazer diversity and dispersal on the magnitude and stability of ecosystem properties in experimental seagrass communities that were either isolated or connected by dispersal corridors. We show that, contrary to theoretical predictions, increasing the number of mobile grazer species in these metacommunities increased the spatial and temporal variability of primary and secondary production. Moreover, allowing grazers to move among and select patches reduced diversity effects on production. Finally, effects of diversity on stability differed qualitatively between patch and metacommunity scales. Our results indicate that declining biodiversity and habitat fragmentation synergistically influence the predictability of ecosystem functioning.

Animal Migration↗

Plant litter decomposition in a semi-arid ecosystem controlled by photodegradation.

The carbon balance in terrestrial ecosystems is determined by the difference between inputs from primary production and the return of carbon to the atmosphere through decomposition of organic matter. Our understanding of the factors that control carbon turnover in water-limited ecosystems is limited, however, as studies of litter decomposition have shown contradictory results and only a modest correlation with precipitation. Here we evaluate the influence of solar radiation, soil biotic activity and soil resource availability on litter decomposition in the semi-arid Patagonian steppe using the results of manipulative experiments carried out under ambient conditions of rainfall and temperature. We show that intercepted solar radiation was the only factor that had a significant effect on the decomposition of organic matter, with attenuation of ultraviolet-B and total radiation causing a 33 and 60 per cent reduction in decomposition, respectively. We conclude that photodegradation is a dominant control on above-ground litter decomposition in this semi-arid ecosystem. Losses through photochemical mineralization may represent a short-circuit in the carbon cycle, with a substantial fraction of carbon fixed in plant biomass being lost directly to the atmosphere without cycling through soil organic matter pools. Furthermore, future changes in radiation interception due to decreased cloudiness, increased stratospheric ozone depletion, or reduced vegetative cover may have a more significant effect on the carbon balance in these water-limited ecosystems than changes in temperature or precipitation.

Argentina↗

Hydrogen and bioenergetics in the Yellowstone geothermal ecosystem.

The geochemical energy budgets for high-temperature microbial ecosystems such as occur at Yellowstone National Park have been unclear. To address the relative contributions of different geochemistries to the energy demands of these ecosystems, we draw together three lines of inference. We studied the phylogenetic compositions of high-temperature (>70 degrees C) communities in Yellowstone hot springs with distinct chemistries, conducted parallel chemical analyses, and carried out thermodynamic modeling. Results of extensive molecular analyses, taken with previous results, show that most microbial biomass in these systems, as reflected by rRNA gene abundance, is comprised of organisms of the kinds that derive energy for primary productivity from the oxidation of molecular hydrogen, H2. The apparent dominance by H2-metabolizing organisms indicates that H2 is the main source of energy for primary production in the Yellowstone high-temperature ecosystem. Hydrogen concentrations in the hot springs were measured and found to range up to >300 nM, consistent with this hypothesis. Thermodynamic modeling with environmental concentrations of potential energy sources also is consistent with the proposed microaerophilic, hydrogen-based energy economy for this geothermal ecosystem, even in the presence of high concentrations of sulfide.

Archaea↗

Artificial ecosystem selection.

Artificial selection has been practiced for centuries to shape the properties of individual organisms, providing Darwin with a powerful argument for his theory of natural selection. We show that the properties of whole ecosystems can also be shaped by artificial selection procedures. Ecosystems initiated in the laboratory vary phenotypically and a proportion of the variation is heritable, despite the fact that the ecosystems initially are composed of thousands of species and millions of individuals. Artificial ecosystem selection can be used for practical purposes, illustrates an important role for complex interactions in evolution, and challenges a widespread belief that selection is most effective at lower levels of the biological hierarchy.

Biological Evolution↗

Phenotypic diversity and ecosystem functioning in changing environments: a theoretical framework.

Biodiversity plays a vital role for ecosystem functioning in a changing environment. Yet theoretical approaches that incorporate diversity into classical ecosystem theory do not provide a general dynamic theory based on mechanistic principles. In this paper, we suggest that approaches developed for quantitative genetics can be extended to ecosystem functioning by modeling the means and variances of phenotypes within a group of species. We present a framework that suggests that phenotypic variance within functional groups is linearly related to their ability to respond to environmental changes. As a result, the long-term productivity for a group of species with high phenotypic variance may be higher than for the best single species, even though high phenotypic variance decreases productivity in the short term, because suboptimal species are present. In addition, we find that in the case of accelerating environmental change, species succession in a changing environment may become discontinuous. Our work suggests that this phenomenon is related to diversity as well as to the environmental disturbance regime, both of which are affected by anthropogenic activities. By introducing new techniques for modeling the aggregate behavior of groups of species, the present approach may provide a new avenue for ecosystem analysis.

Biomass↗

ET743: Chemical analysis of the sea squirt Ecteinascidia turbinata ecosystem.

The sea squirt Ecteinascidia turbinata produces the powerful drug ET743. In this study Inductively Coupled Plasma-Atomic Emission Spectroscopy (ICP-AES) and Matrix Assisted Laser Desorption Ionization-Mass Spectrometry (MALDI-MS) are systematically used to measure elemental and molecular species in a Florida Keys mangrove ecosystem that contains the sea squirt. ICP-AES is used to measure the concentration of 27 elements down to the parts per billion level in 16 organisms and 3 sediment samples that reside in the mangrove ecosystem including turtle grass, blue crabs, fire sponge, and lettuce slugs. MALDI-MS is used to search for ET743 in these same organisms and sediment samples. A mass spectral feature corresponding to ET743 is identified in the extract of the sea squirt, red mangrove root (Rhizophera mangle), the schoolmaster snapper (Lutjanus griseus), and a sediment sample taken from the ecosystem. We use MALDI-MS to study the impact that various environmental conditions, such as UV light, I(2), cation binding (Fe(+3), Zn(+2), Pb(+2), Cu(+2)), metal oxide nanoparticles (FeO, CuO, TiO(2), ZnO, Al(2)O(3)), a common mineral (CaCO(3)), and extremes in acidity (0.1 M HCl, 0.1 M NaOH) have on the ET743 structure. The data provide potential structures (degradation products, metal-ligand complexes, etc.) that might be present in organism or sedimentary extracts that are similar to ET743. We are studying the marine geochemistry of this ecosystem so a broth can be developed and tested for producing this marine natural product.

Animals↗

Soil water balance and ecosystem response to climate change.

Some essential features of the terrestrial hydrologic cycle and ecosystem response are singled out by confronting empirical observations of the soil water balance of different ecosystems with the results of a stochastic model of soil moisture dynamics. The simplified framework analytically describes how hydroclimatic variability (especially the frequency and amount of rainfall events) concurs with soil and plant characteristics in producing the soil moisture dynamics that in turn impact vegetation conditions. The results of the model extend and help interpret the classical curve of Budyko, which relates evapotranspiration losses to a dryness index, describing the partitioning of precipitation into evapotranspiration, runoff, and deep infiltration. They also provide a general classification of soil water balance of the world ecosystems based on two governing dimensionless groups summarizing the climate, soil, and vegetation conditions. The subsequent analysis of the links among soil moisture dynamics, plant water stress, and carbon assimilation offers an interpretation of recent manipulative field experiments on ecosystem response to shifts in the rainfall regime, showing that plant carbon assimilation crucially depends not only on the total rainfall during the growing season but also on the intermittency and magnitude of the rainfall events.

Carbon↗

Biological stoichiometry: a chemical bridge between ecosystem ecology and evolutionary biology.

The mission of the American Society of Naturalists is "to advance and diffuse knowledge of organic evolution and other broad biological principles so as to enhance the conceptual unification of the biological sciences." In this article, I argue that the area of biology least integrated with knowledge of organic evolution is the field of ecosystem ecology, as evidenced by a semiquantitative literature survey of use of terms in the scientific literature. I present an overview of recent theoretical developments and empirical findings in the emerging field of biological stoichiometry (the study of the balance of energy and multiple chemical elements in living systems). These developments hold some promise as a means to conceptually integrate ecosystem ecology, with its emphasis on flows and pools of energy and chemical elements, with evolutionary biology, with its emphasis on genetic fitness and the biochemical products of the genome. For example, recent evidence indicates that organismal C : P and N : P ratios have a major impact on biologically mediated flows of energy and phosphorus; in turn, variations among taxa in these ratios are connected to evolved differences in organismal growth rate because of the connection between growth rate and the need for increased allocation to P-rich ribosomal RNA. In this way, evolutionary change in growth-related traits, by altering organismal P requirements, has direct biogeochemical implications, while ecosystem conditions can constrain evolutionary acceleration of growth rates by imposing a direct P limitation on production of the needed biochemical machinery of growth. Thus, stoichiometric theory provides a broad biological principle that can interconvert the currencies and concerns of ecosystem ecology and evolutionary biology, facilitating integration of diverse fields of study and contributing to conceptual unification of the biological sciences.

Biological Evolution↗

Habitat connectivity and ecosystem productivity: implications from a simple model.

The import of resources (food, nutrients) sustains biological production and food webs in resource-limited habitats. Resource export from donor habitats subsidizes production in recipient habitats, but the ecosystem-scale consequences of resource translocation are generally unknown. Here, I use a nutrient-phytoplankton-zooplankton model to show how dispersive connectivity between a shallow autotrophic habitat and a deep heterotrophic pelagic habitat can amplify overall system production in metazoan food webs. This result derives from the finite capacity of suspension feeders to capture and assimilate food particles: excess primary production in closed autotrophic habitats cannot be assimilated by consumers; however, if excess phytoplankton production is exported to food-limited heterotrophic habitats, it can be assimilated by zooplankton to support additional secondary production. Transport of regenerated nutrients from heterotrophic to autotrophic habitats sustains higher system primary production. These simulation results imply that the ecosystem-scale efficiency of nutrient transformation into metazoan biomass can be constrained by the rate of resource exchange across habitats and that it is optimized when the transport rate matches the growth rate of primary producers. Slower transport (i.e., reduced connectivity) leads to nutrient limitation of primary production in autotrophic habitats and food limitation of secondary production in heterotrophic habitats. Habitat fragmentation can therefore impose energetic constraints on the carrying capacity of aquatic ecosystems. The outcomes of ecosystem restoration through habitat creation will be determined by both functions provided by newly created aquatic habitats and the rates of hydraulic connectivity between them.

Animals↗

The FASSET framework for assessment of environmental impact of ionising radiation in European ecosystems--an overview.

The FASSET project was launched in November 2000 under the EC 5th Framework Programme to develop a framework for the assessment of environmental impact of ionising radiation in European ecosystems. It involved 15 organisations in seven European countries and delivered its final report in spring 2004. The project set out to organise radioecological and radiobiological data into a logical structure that would facilitate the assessment of likely effects on non-human biota resulting from known or postulated depositions of radionuclides in the environment. The project included an overview of 20 pathway-based environmental assessment systems targeted at radioactive substances, or at hazardous substances in general. The resulting framework includes the following fundamental elements: source characterisation; description of seven major European ecosystems; selection of a number of reference organisms on the basis of prior ecosystem and exposure analysis; environmental transfer analysis; dosimetric considerations; effects analysis; and general guidance on interpretation including consideration of uncertainties. The project has used existing information supplemented with development in some areas, e.g. Monte Carlo calculations to derive dose conversion coefficients, model development, and the building of an effects database (FRED, the FASSET Radiation Effects Database). On the basis of experience from FASSET and other recent programmes, it can be concluded that (i) there is substantial agreement in terms of conceptual approaches between different frameworks currently in use or proposed, (ii) differences in technical approaches can be largely attributed to differences in ecosystems of concern or in national regulatory requirements, (iii) sufficient knowledge is available to scientifically justify assessments following the Framework structure, but (iv) significant data gaps exist for environmental transfer of key nuclides as well as for effects data for key wildlife groups at environmentally relevant dose rates. This paper briefly describes the overall content of the FASSET Framework, as well as highlighting a few important future challenges.

Ecosystem↗

Strategy for modeling putative multilevel ecosystems on Europa.

A general strategy for modeling ecosystems on other worlds is described. Two alternative biospheres beneath the ice surface of Europa are modeled, based on analogous ecosystems on Earth in potentially comparable habitats, with reallocation of biomass quantities consistent with different sources of energy and chemical constituents. The first ecosystem models a benthic biosphere supported by chemoautotrophic producers. The second models two concentrations of biota at the top and bottom of the subsurface water column supported by energy harvested from transmembrane ionic gradients. Calculations indicate the plausibility of both ecosystems, including small macroorganisms at the highest trophic levels, with ionotrophy supporting a larger biomass than chemoautotrophy.

Biomass↗

Effects of plant traits on ecosystem and regional processes: a conceptual framework for predicting the consequences of global change.

Human activities are causing widespread changes in the species composition of natural and managed ecosystems, but the consequences of these changes are poorly understood. This paper presents a conceptual framework for predicting the ecosystem and regional consequences of changes in plant species composition. Changes in species composition have greatest ecological effects when they modify the ecological factors that directly control (and respond to) ecosystem processes. These interactive controls include: functional types of organisms present in the ecosystem; soil resources used by organisms to grow and reproduce; modulators such as microclimate that influence the activity of organisms; disturbance regime; and human activities. Plant traits related to size and growth rate are particularly important because they determine the productive capacity of vegetation and the rates of decomposition and nitrogen mineralization. Because the same plant traits affect most key processes in the cycling of carbon and nutrients, changes in plant traits tend to affect most biogeochemical cycling processes in parallel. Plant traits also have landscape and regional effects through their effects on water and energy exchange and disturbance regime.

Biomass↗

Carbon dioxide and water vapor exchange by young and old ponderosa pine ecosystems during a dry summer.

We investigated key factors controlling mass and energy exchange by a young (6-year-old) ponderosa pine (Pinus ponderosa Laws.) plantation on the west side of the Sierra Nevada Mountains and an old-growth ponderosa pine forest (mix of 45- and 250-year-old trees) on the east side of the Cascade Mountains, from June through September 1997. At both sites, we operated eddy covariance systems above the canopy to measure net ecosystem exchange of carbon dioxide and water vapor, and made concurrent meteorological and ecophysiological measurements. Our objective was to understand and compare the controls on ecosystem processes in these two forests. Precipitation is much higher in the young plantation than in the old-growth forest (1660 versus 550 mm year-1), although both forests experienced decreasing soil water availability and increasing vapor pressure deficits (D) as the summer of 1997 progressed. As a result, drought stress increased at both sites during this period, and changes in D strongly influenced ecosystem conductance and net carbon uptake. Ecosystem conductance for a given D was higher in the young pine plantation than in the old-growth forest, but decreased dramatically following several days of high D in late summer, possibly because of xylem cavitation. Net CO2 exchange generally decreased with conductance at both sites, although values were roughly twice as high at the young site. Simulations with the 3-PG model, which included the effect of tree age on fluxes, suggest that, during the fall through spring period, milder temperatures and ample water availability at the young site provide better conditions for photosynthesis than at the old pine site. Thus, over the long-term, the young site can carry more leaf area, and the climatic conditions between fall and spring offset the more severe limitations imposed by summer drought.

California↗

Rocky Mountain ecosystems: diversity, complexity and interactions.

The interior west of North America provides many opportunities to study ecosystem responses to climate change, biological diversity and management of disturbance regimes. These ecosystem responses are not unique to the Rocky Mountains, but they epitomize similar scientific problems throughout North America. Better management of these ecosystems depends on a thorough understanding of the underlying biology and ecological interactions of the species that occupy the diverse habitats of this region. This review highlights progress in research to understand aspects of this complex ecosystem.

Altitude↗

Component carbon fluxes and their contribution to ecosystem carbon exchange in a pine forest: an assessment based on eddy covariance measurements and an integrated model.

We used a combination of eddy flux, canopy, soil and environmental measurements with an integrated biophysical model to analyze the seasonality of component carbon (C) fluxes and their contribution to ecosystem C exchange in a 50-year-old Scots pine forest (Pinus sylvestris L.) in eastern Finland (62 degrees 47' N, 30 degrees 58' E) over three climatically contrasting years (2000-2002). Eddy flux measurements showed that the growing Scots pine forest was a sink for CO2, with annual net C uptakes of 131, 210 and 258 g C m-2> year-1 in 2000, 2001 and 2002, respectively. The integrated process model reproduced the annual course of daily C flux above the forest canopy as measured by the eddy covariance method once the site-specific component parameters were estimated. The model explained 72, 66 and 68% of the variation in daily net C flux in 2000, 2001 and 2002, respectively. Modeled annual C loss by respiration was 565, 629 and 640 g C m-2 year-1, accounting for 77, 77 and 65% of annual gross C uptake, respectively. Carbon fluxes from the forest floor were the dominant contributors to forest ecosystem respiration, with the fractions of annual respiration from the forest floor, foliage and wood being 46-62, 27-44 and 9-10%, respectively. The wide range in daily net C uptake during the growing season was largely attributable to day-to-day fluctuations in incident quantum irradiance. During just a few days in early spring and late autumn, ecosystem net C exchange varied between source and sink as a result of large daily changes in temperature. The forest showed a greater reduction in gross C uptake by photosynthesis than in C loss by respiration during the dry summer of 2000, indicating that interannual variability in ecosystem net C uptake at this site was modified mostly by summer rainfall and vapor pressure deficit.

Carbon↗

Using ergodic theory to assess the performance of ecosystem models.

Ecosystem simulation models are designed to assess the flux of energy, water, carbon and nitrogen according to a given vegetation type. The reliability of the modeled results is determined by model validations. Model validations are typically done using classical statistical methods like regression analysis of predicted versus observed values, paired t statistics and error assessment procedures to characterize the quality of current and future model predictions. All these validation efforts concentrate on static aspects of the model but fail to describe the model dynamics. In this paper, we introduce methods from ergodic theory to analyze the dynamic behavior of ecosystem models. We describe (1) how the attractor representation of model behavior can be reconstructed from a time series of model outputs, and (2) what we can learn from the attractor to assess the model dynamics. As an application example, we provide simulation results for two important pine forest ecosystems in Austria, i.e., 23 Cembran pine and 16 Scots pine stands. These stands were simulated with three model parameterizations: one representing a generic, evergreen needle-leaf forest, and two species- specific parameter sets, one for Cembran pine and one for Scots pine. First, we applied standard validation methods to get static measures of model accuracy and precision. Next, we used ergodic theory to assess model dynamics. A comparison of both analyses reveals important issues related to model dynamics, such as the finding that the occurrence of instabilities in model behavior may not be detected by standard validation methods. Using ergodic theory, we were able to reconstruct the attractors of model behavior. In the attractor describing model dynamics for Cembran pine, simulated with the generic, evergreen needle-leaf forest parameter set, we detected instability in model behavior. We identified this instability as a riddled basin configuration, which is a strong indicator for the occurrence of a chaotic model behavior that may result in random predictions. Our results suggest that ergodic theory is a useful tool for assessing inconsistencies in the dynamics of ecosystem model simulations that have not been detected by standard statistical validation methods.

Climate↗

Impacts of marine-derived nutrients on stream ecosystem functioning.

Energy and nutrient subsidies transported across ecosystem boundaries are increasingly appreciated as key drivers of consumer-resource dynamics. As purveyors of pulsed marine-derived nutrients (MDN), spawning salmon are one such cross-ecosystem subsidy to freshwaters connected to the north Pacific. We examined how salmon carcasses influenced detrital processing in an oligotrophic stream. Experimental manipulations of MDN inputs revealed that salmon carcasses indirectly reduced detrital processing in streams through temporarily decoupling the detrital resource-consumer relationship, in which detrital consumers shifted their diet to the high-nutrient resource, i.e. salmon carcasses. The average decomposition rate of alder leaves with salmon carcass addition was significantly lower than that without the carcass, which was associated with lower abundance and biomass of detritivorous Trichoptera on the carcass-treated leaves. There were generally larger in size Trichopteran detritivores on the carcasses than on leaves. These results imply that cross-boundary MDN subsidies indirectly retard the ecosystem processing of leaf litter within the short term, but may enhance those food-limited detritivorous consumers. Because unproductive freshwaters in the Pacific northwest are highly dependent upon the organic matter inputs from surrounding forests, this novel finding has implications for determining conservation and management strategies of salmon-related aquatic ecosystems, in terms of salmon habitat protection and fisheries exploitation.

Analysis of Variance↗

Protecting terrestrial ecosystems and the climate through a global carbon market.

Protecting terrestrial ecosystems through international environmental laws requires the development of economic mechanisms that value the Earth's natural systems. The major international treaties to address ecosystem protection lack meaningful binding obligations and the requisite financial instruments to affect large-scale conservation. The Kyoto Protocol's emissions-trading framework creates economic incentives for nations to reduce greenhouse-gas (GHG) emissions cost effectively. Incorporating GHG impacts from land-use activities into this system would create a market for an important ecosystem service provided by forests and agricultural lands: sequestration of atmospheric carbon. This would spur conservation efforts while reducing the 20% of anthropogenic CO(2) emissions produced by land-use change, particularly tropical deforestation. The Kyoto negotiations surrounding land-use activities have been hampered by a lack of robust carbon inventory data. Moreover, the Protocol's provisions agreed to in Kyoto made it difficult to incorporate carbon-sequestering land-use activities into the emissions-trading framework without undermining the atmospheric GHG reductions contemplated in the treaty. Subsequent negotiations since 1997 failed to produce a crediting system that provides meaningful incentives for enhanced carbon sequestration. Notably, credit for reducing rates of tropical deforestation was explicitly excluded from the Protocol. Ultimately, an effective GHG emissions-trading framework will require full carbon accounting for all emissions and sequestration from terrestrial ecosystems. Improved inventory systems and capacity building for developing nations will, therefore, be necessary.

Air Pollution↗