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Nonpoint-source nutrients contribute to eutrophication of surface waters. While effects of particular management actions are difficult to identify at the watershed scale, assessing nutrient fluxes over time can illuminate the net impact of trends in land use and management. We investigated nutrient fluxes in upper Kamm Creek in northwest Washington State to determine if historical changes in land management, responding to economic and regulatory shifts, had impacted nutrient export. We compared current (October 2015 to September 2018) nitrogen and phosphorus fluxes in Kamm Creek to data from a previous study (1993 to 1998). We found significantly higher current fluxes of nitrate, and significantly lower current fluxes of orthophosphate (P) and total phosphorus (TP), compared to the historical sampling period. The increased annual nitrate flux resulted from slightly higher average discharge and significantly higher nitrate concentrations throughout the year. In contrast, current P and TP concentrations were significantly lower throughout the year than previously. The Nooksack River, which receives water from the study stream, showed similar patterns for P and TP, but not nitrate. Kamm Creek had reduced phosphorus fluxes and increased nitrate fluxes between the 1993 to1998 and 2015 to 2018 time periods. The watershed experienced several concurrent land-use changes, including reductions in hay and corn acreage, increases in berry crop acreage, legislation to reduce manure application, and altered use of nitrate-enriched groundwater. We cannot currently distinguish among these. Further understanding relationships between specific management changes and nutrient fluxes will help to maintain local agricultural productivity and improve water quality.

Watershed nitrogen (N) budgets provide insights into drivers and solutions for groundwater and surface water N contamination. We constructed a comprehensive N budget for the transboundary Nooksack River Watershed (British Columbia, Canada and Washington, US) using locally-derived data, national statistics and standard parameters. Feed imports for dairy (mainly in the US) and poultry (mainly in Canada) accounted for 30 and 29% of the total N input to the watershed, respectively. Synthetic fertilizer was the next largest source contributing 21% of inputs. Food imports for humans and pets together accounted for 9% of total inputs, lower than atmospheric deposition (10%). N imported by returning salmon representing marine derived nutrients accounted for

Plant diversity can increase biomass production in plot-scale studies, but applying these results to ecosystem carbon (C) storage at larger spatial and temporal scales remains problematic. Other ecosystem controls interact with diversity and plant production, and may influence soil pools differently from plant pools. We integrated diversity with the state-factor framework, which identifies key controls, or 'state factors', over ecosystem properties and services such as C storage. We used this framework to assess the effects of diversity, plant traits and state factors (climate, topography, time) on live tree, standing dead, organic horizon and total C in Québec forests. Four patterns emerged: (1) while state factors were usually the most important model predictors, models with both state and biotic factors (mean plant traits and diversity) better predicted C pools; (2) mean plant traits were better predictors than diversity; (3) diversity increased live tree C but reduced organic horizon C; (4) different C pools responded to different traits and diversity metrics. These results suggest that, where ecosystem properties result from multiple processes, no simple relationship may exist with any one organismal factor. Integrating biodiversity into ecosystem ecology and assessing both traits and diversity improves our mechanistic understanding of biotic effects on ecosystems.

International audience; Global species extinction rates are orders of magnitude above the background rate documented in the fossil record. However, recent data syntheses have found mixed evidence for patterns of net species loss at local spatial scales. For example, two recent data meta-analyses have found that species richness is decreasing in some locations and is increasing in others. When these trends are combined, these papers argued there has been no net change in species richness, and suggested this pattern is globally representative of biodiversity change at local scales. Here we reanalyze results of these data syntheses and outline why this conclusion is unfounded. First, we show the datasets collated for these syntheses are spatially biased and not representative of the spatial distribution of species richness or the distribution of many primary drivers of biodiversity change. This casts doubt that their results are representative of global patterns. Second, we argue that detecting the trend in local species richness is very difficult with short time series and can lead to biased estimates of change. Reanalyses of the data detected a signal of study duration on biodiversity change, indicating net biodiversity loss is most apparent in studies of longer duration. Third, estimates of species richness change can be biased if species gains during post-disturbance recovery are included without also including species losses that occurred during the disturbance. Net species gains or losses should be assessed with respect to common baselines or reference communities. Ultimately, we need a globally coordinated effort to monitor biodiversity so that we can estimate and attribute human impacts as causes of biodiversity change. A combination of technologies will be needed to produce regularly updated global datasets of local biodiversity change to guide future policy. At this time the conclusion that there is no net change in local species richness is not the consensus state of knowledge.

Many studies have examined how intensity of grazing and patterns of precipitation individually and interactively influence the spatial and temporal dynamics of grassland vegetation, such as dominance, succession, coexistence, and spatial heterogeneity. However existing models have rarely considered the diet preferences of grazers and how they interact with variation in precipitation amount and timing. We examined how plant community structure responds to the individual and combined effects of grazing intensity, selective grazing, and patterns of precipitation, based on a six-year grazing experiment with seven levels of field-manipulated grazing intensity in a typical steppe of Inner Mongolia. The palatable species, mainly forbs, were most severely damaged at intermediate levels of grazing intensity; given that these species are the major contributors to plant community diversity, a U-shaped diversity-grazing intensity relationship resulted. In contrast, spatial heterogeneity of aboveground biomass and species composition peaked at intermediate levels of grazing intensity. Cold season precipitation positively correlated with the abundance of the dominant C3 grasses and correlated negatively with the subdominant forbs and C4 plants. Thus, when cold season precipitation increased, plant community species diversity decreased. Grazing intensity and precipitation did not interact in their effects on species richness. These findings contrast with the predictions from current disturbance–diversity models and indicate that diet selection of grazing animals is an important factor shaping the diversity-grazing intensity relationship in semi-arid grasslands. Future grassland biodiversity conservation and management practices should take diet preference of grazing animals into account.

Biodiversity confers economic value by enhancing carbon storage in grasslands, an economic argument for biodiversity conservation., Carbon storage by ecosystems is valuable for climate protection. Biodiversity conservation may help increase carbon storage, but the value of this influence has been difficult to assess. We use plant, soil, and ecosystem carbon storage data from two grassland biodiversity experiments to show that greater species richness increases economic value: Increasing species richness from 1 to 10 had twice the economic value of increasing species richness from 1 to 2. The marginal value of each additional species declined as species accumulated, reflecting the nonlinear relationship between species richness and plant biomass production. Our demonstration of the economic value of biodiversity for enhancing carbon storage provides a foundation for assessing the value of biodiversity for decisions about land management. Combining carbon storage with other ecosystem services affected by biodiversity may well enhance the economic arguments for conservation even further.

Degradation of semiarid ecosystems from overgrazing threatens a variety of ecosystem services. Rainfall and nitrogen commonly co-limit production in semiarid grassland ecosystems; however, few studies have reported how interactive effects of precipitation and nitrogen addition influence the recovery of grasslands degraded by overgrazing. We conducted a 6-year experiment manipulating precipitation (natural precipitation and simulated wet year precipitation) and nitrogen (0, 25 and 50 kg N ha−1) addition at two sites with different histories of livestock grazing (moderately and heavily grazed) in Inner Mongolian steppe. Our results suggest that recovery of plant community composition and recovery of production can be decoupled. Perennial grasses provide long-term stability of high-quality forage production in this system. Supplemental water combined with exclosures led, in the heavily grazed site, to the strongest recovery of perennial grasses, although widespread irrigation of rangeland is not a feasible management strategy in many semiarid and arid regions. N fertilization combined with exclosures, but without water addition, increased dominance of unpalatable annual species, which in turn retarded growth of perennial species and increased inter-annual variation in primary production at both sites. Alleviation of grazing pressure alone allowed recovery of desired perennial species via successional processes in the heavily grazed site. Our experiments suggest that recovery of primary production and desirable community composition are not necessarily correlated. The use of N fertilization for the management of overgrazed grassland needs careful and systematic evaluation, as it has potential to impede, rather than aid, recovery.

Evidence is mounting that extinctions are altering key processes important to the productivity and sustainability of Earth’s ecosystems 1–4 . Further species loss will accelerate change in ecosystem processes 5–8 , but it is unclear how these effects compare to the direct effects of other forms of environmental change that are both driving diversity loss and altering ecosystem function. Here we use a suite of meta-analyses of published data to show that the effects of species loss on productivity and decomposition—two processes important in all ecosystems—are of comparable magnitude to the effects of many other global environmental changes. In experiments, intermediate levels of species loss (21–40%) reduced plant production by 5–10%, comparable to previously documented effects of ultraviolet radiation and climate warming. Higher levels of extinction (41–60%) had effects rivalling those of ozone, acidification, elevated CO2 and nutrient pollution. At intermediate levels, species loss generally had equal or greater effects on decomposition than did elevated CO2 and nitrogen addition. The identity of species lost also had a large effect on changes in productivity and decomposition, generating a wide range of plausible outcomes for extinction. Despite the need for more studies on interactive effects of diversity loss and environmental changes, our analyses clearly show that the ecosystem consequences of local species loss are as quantitatively significant as the direct effects of several global change stressors that have mobilized major international concern and remediation efforts 9 .

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.

Over the past several decades, a rapidly expanding field of research known as biodiversity and ecosystem functioning has begun to quantify how the world's biological diversity can, as an independent variable, control ecological processes that are both essential for, and fundamental to, the functioning of ecosystems. Research in this area has often been justified on grounds that (1) loss of biological diversity ranks among the most pronounced changes to the global environment and that (2) reductions in diversity, and corresponding changes in species composition, could alter important services that ecosystems provide to humanity (e.g., food production, pest/disease control, water purification). Here we review over two decades of experiments that have examined how species richness of primary producers influences the suite of ecological processes that are controlled by plants and algae in terrestrial, marine, and freshwater ecosystems. Using formal meta-analyses, we assess the balance of evidence for eight fundamental questions and corresponding hypotheses about the functional role of producer diversity in ecosystems. These include questions about how primary producer diversity influences the efficiency of resource use and biomass production in ecosystems, how primary producer diversity influences the transfer and recycling of biomass to other trophic groups in a food web, and the number of species and spatial /temporal scales at which diversity effects are most apparent. After summarizing the balance of evidence and stating our own confidence in the conclusions, we outline several new questions that must now be addressed if this field is going to evolve into a predictive science that can help conserve and manage ecological processes in ecosystems.

Scheduled to begin in 2011, the removal of two dams on the Elwha River, Washington, will be one of the largest dam removal and river restoration projects undertaken in the United States. One challenge associated with this project is to understand how exposure and downstream deposition of sediments presently detained behind the dams will influence patterns of revegetation and invasive species colonization following dam removal. We conducted two greenhouse experiments assessing the potential effects of reservoir sediments on germination success and growth of (1) propagules dispersed naturally via seed rain, and (2) seeds of selected native and invasive species. Observed summer seed rain density was relatively low (

Summary 1. Recent debates about the role of biotic resistance in controlling invasion success have focused on effects of species richness. However, functional composition could be a stronger control: species already in the community with similar functional traits to those of the invaders should have the greatest competitive effect on invaders. Still, experiments assessing effects of functional similarity have found contradictory results. 2. We used experimental communities in a serpentine grassland in California, USA, to assess the extent to which functional composition and functional diversity influenced success of two different types of invading plants: early season annuals (E) and late-season annuals (L) that have been previously shown to differ in patterns of resource acquisition. 3. We seeded known quantities of seed of six different species (three in each functional group) into experimental plots containing established communities differing in functional composition and functional diversity. The experimental communities contained different combinations of E, L, perennial bunchgrass (P) and nitrogen-fixer (N) functional groups, with functional diversity ranging from 0 to 4 groups. Each invading species was seeded into a separate quadrat within each plot to minimize competitive effects of invaders on each other. We measured both seedling and adult success of invaders for two full growing seasons to further understand mechanisms underlying biotic resistance. 4. More functionally diverse communities were less invaded overall, as measured by the average success of individual invaders. However, assessment of invaders by functional groups was more informative: Es in the extant community suppressed E invaders the most, and Ls in the extant community suppressed L invaders the most. 5. We observed a variety of interactions among extant functional groups in reducing invader success, including synergism, complementarity and ‘basement’ effects, where two or more groups negatively affected invaders, but combinations of groups were no more suppressive than single groups. The extant community influenced invaders more strongly through suppression of adult plant growth than through effects on seedling establishment. 6. Synthesis. Contrary to predictions from neutral theory, these results indicate that niche overlap was an important component of biotic resistance in these experimental plant communities and summed up to significant effects of species richness.

Predicting ecosystem responses to global change is a major challenge in ecology. A critical step in that challenge is to understand how changing environmental conditions influence processes across levels of ecological organization. While direct scaling from individual to ecosystem dynamics can lead to robust and mechanistic predictions, new approaches are needed to appropriately translate questions through the community level. Species invasion, loss, and turnover all necessitate this scaling through community processes, but predicting how such changes may influence ecosystem function is notoriously difficult. We suggest that community-level dynamics can be incorporated into scaling predictions using a trait-based response-effect framework that differentiates the community response to environmental change (predicted by response traits) and the effect of that change on ecosystem processes (predicted by effect traits). We develop a response-and-effect functional framework, concentrating on how the relationships among species' response, effect, and abundance can lead to general predictions concerning the magnitude and direction of the influence of environmental change on function. We then detail several key research directions needed to better scale the effects of environmental change through the community level. These include (1) effect and response trait characterization, (2) linkages between response-and-effect traits, (3) the importance of species interactions on trait expression, and (4) incorporation of feedbacks across multiple temporal scales. Increasing rates of extinction and invasion that are modifying communities worldwide make such a research agenda imperative.

A recent debate among ecologists has focused on mechanisms by which species diversity might affect net primary productivity. Communities with more species could use a greater variety of resource capture characteristics, leading to greater use of limiting resources (complementarity) and therefore greater productivity (overyielding). Recent experiments, however, have shown a variety of relationships between diversity and productivity. In an experiment on serpentine grassland communities spanning 8 years, we found that overyielding increased several years after plot establishment. Overyielding varied greatly depending on the functional characteristics of the species involved and the biotic and abiotic environment (particularly water availability). While functional differences among species led to strong complementarity and facilitation, these effects were not sufficient to cause significant transgressive overyielding or consistent increases in productivity with increased plant diversity. These results suggest that greater absolute production with greater diversity may be restricted to particular species combinations or environmental conditions.

Climatic change may influence decomposition dynamics in arctic and boreal ecosystems, affecting both atmospheric CO2 levels, and the flux of dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) to aquatic systems. In this study, we investigated landscape-scale controls on potential production of these compounds using a one-year laboratory incubation at two temperatures (10° and 30 °C). We measured the release of CO2, DOC and DON from tundra soils collected from a variety of vegetation types and climatic regimes: tussock tundra at four sites along a latitudinal gradient from the interior to the north slope of Alaska, and soils from additional vegetation types at two of those sites (upland spruce at Fairbanks, and wet sedge and shrub tundra at Toolik Lake in northern Alaska). Vegetation type strongly influenced carbon fluxes. The highest CO2 and DOC release at the high incubation temperature occurred in the soils of shrub tundra communities. Tussock tundra soils exhibited the next highest DOC fluxes followed by spruce and wet sedge tundra soils, respectively. Of the fluxes, CO2 showed the greatest sensitivity to incubation temperatures and vegetation type, followed by DOC. DON fluxes were less variable. Total CO2 and total DOC release were positively correlated, with DOC fluxes approximately 10% of total CO2 fluxes. The ratio of CO2 production to DOC release varied significantly across vegetation types with Tussock soils producing an average of four times as much CO2 per unit DOC released compared to Spruce soils from the Fairbanks site. Sites in this study released 80–370 mg CO2-C g soil C−1 and 5–46 mg DOC g soil C−1 at high temperatures. The magnitude of these fluxes indicates that arctic carbon pools contain a large proportion of labile carbon that could be easily decomposed given optimal conditions. The size of this labile pool ranged between 9 and 41% of soil carbon on a g soil C basis, with most variation related to vegetation type rather than climate.

The LI-COR 6200 portable photosynthesis system (LI-6200) is commonly used in combination with large chambers to measure ecosystem level CO2 flux in ecosystems with small-statured canopies (agriculture, tundra, grasslands, forest understory, etc.). Two problems with the methodology lead to artifactually low estimates of rates of net ecosystem assimilation of CO2 (or overestimates of ecosystem respiration). The first is that accuracy of the equations used by the LI-6200 to calculate photosynthesis depends on a constant vapor pressure in the chamber. This assumption is commonly violated with large ecosystem chambers when evapotranspiration rates are high. We provide equations that correct this problem and permit recalculation of the LI-COR fluxes. The second problem is that of boundary layer formation under still conditions, such as at night. As high concentrations of CO2 close to the ground surface become mixed by chamber fans, exceptionally high values of net ecosystem respiration result. Substantial mixing time is necessary for rates to stabilize. As ecologists attempt to understand how global change might affect whole-ecosystem carbon balance, both of these technical problems must be addressed to get accurate results.

The ecological consequences of biodiversity loss have aroused considerable interest and controversy during the past decade. Major advances have been made in describing the relationship between species diversity and ecosystem processes, in identifying functionally important species, and in revealing underlying mechanisms. There is, however, uncertainty as to how results obtained in recent experiments scale up to landscape and regional levels and generalize across ecosystem types and processes. Larger numbers of species are probably needed to reduce temporal variability in ecosystem processes in changing environments. A major future challenge is to determine how biodiversity dynamics, ecosystem processes, and abiotic factors interact.

To investigate how plant diversity affects ecosystem-level processes such as primary production and nutrient cycling, I established an experimental plant diversity gradient in serpentine grassland using four functional groups of plants: early season annual forbs (E), late season annual forbs (L), perennial bunchgrasses (P), and nitrogen fixers (N). These groups differ in growth form, phenology, and other traits relevant to nutrient cycling (e.g., rooting depth, litter C:N ratio). Two or three species of each type were planted in single-group treatments, and in two-, three-, and four-way combinations, giving a range of richness from zero to nine species. I tested the hypothesis that, because of complementary resource use, increasing functional group diversity will lead to higher net primary production. At the scale of this experiment (α-diversity and yearly production), more diverse treatments were not necessarily the most productive. Live plant biomass varied more within than among levels of diversity. In most two-, three-, and four-way mixtures of functional groups, overall productivity did not differ significantly from the average of the yields of component one-group treatments. This pattern apparently resulted from competition: early season annuals and late season annuals reduced the biomass of perennial bunchgrasses (the most productive group in monoculture) below levels expected from monoculture yields. Relative Yield Totals (RYT) indicated complementary resource use in the EL and LP two-way and ELPN four-way mixtures. In the EL mixture, complementarity appeared to result from temporal rather than spatial partitioning of resources. Because of shifts in root:shoot ratio in mixtures, however, only the LP treatment had consistently significant RYT >1 when assessing total (roots plus shoots) productivity and nitrogen yield. These results show that (1) composition (the identity of the species present) can be at least as important as richness (the number of species present) in effects on ecosystem processes; (2) competition during critical parts of the growing season may prevent absolute increases in net primary production with increasing diversity, despite complementary resource use at other times of the year; and (3) shifts in belowground allocation in species mixtures can have significant effects on estimates of productivity and resource use as species diversity changes.

We evaluated the effects of plant functional group richness on seasonal patterns of soil nitrogen and phosphorus cycling, using serpentine grassland in south San Jose, California. We established experimental plots with four functional types of plants: early-season annual forbs (E), late-season annual forbs (L), nitrogen-fixers (N), and perennial bunchgrasses (P). These groups differ in several traits relevant to nutrient cycling, including phenology, rooting depth, root:shoot ratio, size, and leaf C:N content. Two or three species of each group were planted in single functional group (SFG) treatments, and in two-, three-, and four-way combinations of functional groups. We analyzed available nutrient pool sizes, microbial biomass nitrogen and phosphorus, microbial nitrogen immobilization, nitrification rates, and leaching losses. We used an index of “relative resource use” that incorporates the effects of plants on pool sizes of several depletable soil resources: inorganic nitrogen in all seasons, availabl...

F. Stuart Chapin (e-mail: fschapin@socrates. berkeley.edu) is a professor and David U. Hooper is a postdoctoral fellow in the Department of Integrative Biology, University of California, Berkeley, CA 94720. Osvaldo E. Sala is a professor at the Faculty of Agronomy, University of Buenos Aires, Av. San Martin 4453, Buenos Aires 1417, Argentina. Ingrid C. Burke is an associate professor in the Department of Forest Sciences, Colorado State University, Ft. Collins, CO 80523. J. Phillip Grime is a professor in the Unit of Comparative Plant Ecology, The University of Sheffield, Sheffield S10 2TN, UK. William K. Lauenroth is a professor in the Rangeland Ecosystem Science Department, Natural Resource Ecology Laboratory, Colorado State University, Ft. Collins, CO 80523. Amanda Lombard is a research scientist at the FitzPatrick Institute, University of Cape Town, Cape Town, 7700 South Africa. Harold A. Mooney is a professor in the Department of Biological Sciences, Stanford University, Stanford, CA 94305. Arvin R. Mosier is a research scientist in the USDA Agricultural Research Service, P. O. Box E, Ft. Collins, CO 80522. Shahid Naeem is an assistant professor and David Tilman is a professor in the Department of Ecology, Evolution, and Behavior, University of Minnesota, St. Paul, MN 55108. Stephen W. Pacala is a professor in the Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ 08544. Jacques Roy is director of research at the Centre d'Ecologie Fonctionnelle et Evolutive, CNRS, F-34293 Montpellier Cedex 5, France. William L. Steffen is executive officer of GCTE, CSIRO, Division of Wildlife and Ecology, Lyneham ACT 2602, Australia. Changes in biodiversity can have significant

The relative effects of plant richness (the number of plant functional groups) and composition (the identity of the plant functional groups) on primary productivity and soil nitrogen pools were tested experimentally. Differences in plant composition explained more of the variation in production and nitrogen dynamics than did the number of functional groups present. Thus, it is possible to identify and differentiate among potential mechanisms underlying patterns of ecosystem response to variation in plant diversity, with implications for resource management.

Summary Stream conservation and restoration strategies commonly focus on preserving extant riparian forest and restoring riparian habitat. Though these practices are beneficial to stream habitat, they may not suffice to restore stream condition in catchments heavily influenced by intensive land use. To evaluate this hypothesis, we measured invertebrate populations in 12 streams affected by different land-use types (cultivated, developed, forested and grassland). Within each stream, we compared reaches with and without riparian buffers of remnant mature forest. Within each reach, we calculated the relative abundance of individuals from the insect orders Ephemeroptera, Plecoptera and Trichoptera (%EPT) and assessed specific conductance, dissolved oxygen (DO), temperature and substratum composition. Forested catchments had the highest %EPT (average 23.8) followed by grassland (16.1), cultivated (1.96) and developed (0.31). Reaches with forested buffers present were not associated with higher %EPT in forested, cultivated or developed catchments. In grassland catchments, however, %EPT was about eightfold higher in reaches with forested buffers than those without. High values of %EPT were associated with large stream substratum and low specific conductance and temperature, which were most common in forested catchments. Developed and cultivated catchments did not follow this pattern, %EPT being uniformly low, even where abiotic conditions were similar to forested catchments. These results confirm that intensive urban and cultivated land uses at the catchment scale degrade stream biological communities. They also indicate that patchy forested riparian corridors, as found in many restoration projects, are probably insufficient to mitigate severe, large-scale biological degradation.

Extensive research shows that more species-rich assemblages are generally more productive and efficient in resource use than comparable assemblages with fewer species. But the question of how diversity simultaneously affects the wide variety of ecological functions that ecosystems perform remains relatively understudied, and it presents several analytical and empirical challenges that remain unresolved. In particular, researchers have developed several disparate metrics to quantify multifunctionality, each characterizing different aspects of the concept, and each with pros and cons. We compare four approaches to characterizing multifunctionality and its dependence on biodiversity, quantifying 1) magnitudes of multiple individual functions separately, 2) the extent to which different species promote different functions, 3) the average level of a suite of functions, and 4) the number of functions that simultaneously exceed a critical threshold. We illustrate each approach using data from the pan-European BIODEPTH experiment and the R multifunc package developed for this purpose, evaluate the strengths and weaknesses of each approach, and implement several methodological improvements. We conclude that a extension of the fourth approach that systematically explores all possible threshold values provides the most comprehensive description of multifunctionality to date. We outline this method and recommend its use in future research., Comment: This article has been submitted to Methods in Ecology & Evolution for review

The most unique feature of Earth is the existence of life, and the most extraordinary feature of life is its diversity. Approximately 9 million types of plants, animals, protists and fungi inhabit the Earth. So, too, do 7 billion people. Two decades ago, at the first Earth Summit, the vast majority of the world's nations declared that human actions were dismantling the Earth's ecosystems, eliminating genes, species and biological traits at an alarming rate. This observation led to the question of how such loss of biological diversity will alter the functioning of ecosystems and their ability to provide society with the goods and services needed to prosper.

Long-term livestock over-grazing causes nitrogen outputs to exceed inputs in Inner Mongolia, suggesting that low levels of nitrogen fertilization could help restore grasslands degraded by overgrazing. However, the effectiveness of such an approach depends on the response of production and species composition to the interactive drivers of nitrogen and water availability. We conducted a five-year experiment manipulating precipitation (NP: natural precipitation and SWP: simulated wet year precipitation) and nitrogen (0, 25 and 50 kg N ha(-1) yr(-1)) addition in Inner Mongolia. We hypothesized that nitrogen fertilization would increase forage production when water availability was relatively high. However, the extent to which nitrogen would co-limit production under average or below average rainfall in these grasslands was unknown.Aboveground net primary production (ANPP) increased in response to nitrogen when precipitation was similar to or higher than the long-term average, but not when precipitation was below average. This shift in limitation was also reflected by water and nitrogen use efficiency. Belowground live biomass significantly increased with increasing water availability, but was not affected by nitrogen addition. Under natural precipitation (NP treatment), the inter-annual variation of ANPP was 3-fold greater than with stable water availability (CV(ANPP) = 61±6% and 17±3% for NP and SWP treatment, respectively) and nitrogen addition increased CV(ANPP) even more (89±14%). This occurred in part because fertilizer nitrogen left in the soil in dry years remained available for uptake during wet years and because of high production by unpalatable annual species in wet years in the NP treatment. In summary, plant growth by residual fertilizer nitrogen could lead to sufficient yields to offset lack of additional production in dry years. However, the utility of fertilization for restoration may be constrained by shifts in species composition and the lack of response by belowground biomass, which reduces replacement of soil carbon and nitrogen.

The LI-COR 6200 portable photosynthesis system (LI-6200) is commonly used in combination with large chambers to measure ecosystem level CO

Human alteration of the global environment has triggered the sixth major extinction event in the history of life and caused widespread changes in the global distribution of organisms. These changes in biodiversity alter ecosystem processes and change the resilience of ecosystems to environmental change. This has profound consequences for services that humans derive from ecosystems. The large ecological and societal consequences of changing biodiversity should be minimized to preserve options for future solutions to global environmental problems. Fil: Chapin III, F. Stuart. University Of Alaska; Estados Unidos Fil: Zavaleta, Erika S.. University of Stanford; Estados Unidos Fil: Eviner, Valerie T.. University of California at Berkeley; Estados Unidos Fil: Naylor, Rosamond L.. University of Stanford; Estados Unidos Fil: Vitousek, Peter M.. University of Stanford; Estados Unidos Fil: Reynolds, Heather L.. Kalamazoo College; Estados Unidos Fil: Hooper, David U.. Western Washington University; Estados Unidos Fil: Lavorel, Sandra. Centre National de la Recherche Scientifique; Francia Fil: Sala, Osvaldo Esteban. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura; Argentina Fil: Hobbie, Sarah E.. University of Minnesota; Estados Unidos Fil: Mack, Michelle C.. University Of Alaska; Estados Unidos Fil: Díaz, Sandra Myrna. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Instituto Multidisciplinario de Biología Vegetal. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Instituto Multidisciplinario de Biología Vegetal; Argentina

We investigated the relationship between plant nitrogen limitation and water availability in dryland ecosystems. We tested the hypothesis that at lower levels of annual precipitation, aboveground net primary productivity (ANPP) is limited primarily by water whereas at higher levels of precipitation, it is limited primarily by nitrogen. Using a literature survey of fertilization experiments in arid, semi-arid, and subhumid ecosystems, we investigated the response of ANPP to nitrogen addition as a function of variation in precipitation across geographic gradients, as well as across year-to-year variation in precipitation within sites. We used four different indices to assess the degree of N limitation: (1) Absolute Increase of plant production in response to fertilization (the slope of ANPP vs. amount of added N at different levels of annual precipitation); (2) Relative Response (the percent increase in fertilized over control ANPP at different levels of N addition); (3) Fertilizer Use Efficiency (FUE, the absolute gain in productivity per amount of fertilizer N), and (4) Maximum Response (the greatest absolute increase in ANPP at saturating levels of N addition). Relative Response to fertilization did not significantly increase with increasing precipitation either across the geographic gradient or across year-to-year variation within sites. Nor did the Maximum Response to fertilization increase with increasing precipitation across the geographic gradient. On the other hand, there was a significant increase in the Absolute Increase and FUE indices with both geographical and temporal variation in precipitation. Together, these results indicate that there is not necessarily a shift of primary limitation from water to N across the geographic water availability gradient. Instead, our results support the hypothesis of co-limitation. The apparently contradictory results from the four indices of N limitation can best be explained by an integration of plant ecophysiological, community, and ecosystem mechanisms whereby plants are co-limited by multiple resources, species shifts occur in response to changing resource levels, and nitrogen and water availability are tightly linked through biogeochemical feedbacks.

Nature 486, 59–67 (2012); doi:10.1038/nature11148 In Table 1 and Supplementary Table 2 of this Review, under the ‘Category of service’ called ‘Regulating’, the first two ‘Measures of service provision’ related to ‘Biocontrol’ should read ‘Abundance of herbivorous pests’ instead of ‘Control of herbivorous pests’.

The effect of biological diversity on ecosystem biogeochemistry has not been widely studied in recent years, due in part to widespread recognition that our understanding of population/ecosystem interactions was insufficient to sustain the pioneering discussions of the late 1960s (cf. Woodwell and Smith 1969). Indeed, the fields of population biology and ecosystem studies largely pursued different approaches throughout much of the 1970s and 1980s. The gap between population and ecosystem ecology is now narrowing substantially, and the linked nature of human alterations to both biological diversity and ecosystem functioning is increasingly appreciated. These developments have led to renewed interest in the connections between diversity and many aspects of ecosystem function.

integration and collaboration as we meet the challenge of understanding the great complexity of biological systems. Ecological subdisciplines are rapidly combining and incorporating other biological, physical, mathematical, and sociological disciplines. The burgeoning base of theoretical and empirical work, made possible by new methods, technologies, and funding opportunities, is providing the opportunity to reach robust answers to major ecological questions. In December 1999 the National Science Foundation convened a white paper committee to evaluate what we know and do not know about important ecological processes, what hurdles currently hamper our progress, and what intellectual and conceptual interfaces need to be encouraged. The committee distilled the discussion into four frontiers in research on the ecological structure of the earth’s biological diversity and the ways in which ecological processes continuously shape that structure (i.e., ecological dynamics). This article summarizes the discussions of those frontiers and explains why they are crucial to our understanding of how ecological processes shape patterns and dynamics of global biocomplexity. The frontiers are 1. Dynamics of coalescence in complex communities 2. Evolutionary and historical determinants of ecological processes: The role of ecological memory 3. Emergent properties of complex systems: Biophysical constraints and evolutionary attractors 4. Ecological topology: Defining the spatiotemporal domains of causality for ecological structure and processes Each of the four research frontiers takes a different approach to the overall ecological dynamics of biocomplexity, and all require integration and collaboration among those approaches. These overlapping frontiers themselves are not necessarily new. Within each frontier, however, are emerging questions and approaches that will help us understand how ecological processes are interconnected over multiple spatial and temporal scales, from local community structure to global patterns.

Assistant professor in the Department of Biology at Western Washington University, Bellingham, Washington 98225-9160 10: Professor at the Laboratoire d'Ecologie de Sols Tropicaux, ORSTOM/Universite Paris VI, 32 Avenue Henri Varagnat, 93143 Bondy, France 11: Senior scientist at the Centre for Terrestrial Ecology, Netherlands Institute of Ecology, 6666 ZG Heteren, Netherlands Utrecht, Netherlands 12: Professor at the Department of Environmental Studies, University of Utrecht, Utrecht, Netherlands 13: Professor at the Institute of Soil Biology, Academy of Sciences of the Czech Republic, Na sadkach 7, 370 05 Ceske Budejovice, Czech Republic 14: Professor at the Department of Environmental Science, Policy,and Management, University of California, Berkeley, California 94720-3110 15: Professor at the Center for Microbial Ecology, Michigan State University, 540 Plant and Soil Science Building, East Lansing, Michigan 48824-1325 16: Professor at the Department of Animal Ecology, Justus Liebig University of Giessen, Heinrich-Buff-Ring 26-32 (IFZ), D-35392 Giessen, Germany 2: Professor at the Queen Mary and Westfield College, School of Biological Sciences, University of London, London E1 4NS, United Kingdom 3: Research professor and the director of the Centre for Agri-Environmental Research, Department of Agriculture, University of Reading, Earley Gate, Reading RG6 6AT, United Kingdom 4: Professor of Soil Biology and Biological Soil Quality and director of the Department of Environmental Sciences, Wageningen University, 6700 EC Wageningen, Netherlands 5: Professor at the Centre for Biodiversity and Bioresources, School of Biological Sciences, Macquarie University, Sydney, New South Wales 2109, Australia 6: Chair, SCOPE Committee on Soil and Sediment Biodiversity and Ecosystem Functioning, and professor and director, Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, Colorado 80523 7: Scientist at Landcare Research, Lincoln, New Zealand 8: Research professor in the Institute of Ecology at the University of Georgia, 102 Ecology Annex, Athens, Georgia 30602-2360 9: Professor at the Department of Soil Science and Agricultural Engineering, University of Zimbabwe, Mount Pleasant, Harare, Zimbabwe

IN the Travancore Court of the British Empire Exhibition at Wembley several interesting products of this enterprising State were displayed. Among the medicinal products there were two substances named “Kalpal” and “Kalmadom.” From a description drawn up by Dr. Kunjan Pillai, these are said to be deposits found in the clefts and hollows of high rocks “in the form of a brownish-white foam which on exposure becomes harder with time and has the consistence of lime-plaster.” The decomposed rock is washed with water to remove foreign matter, as lichens, leaves, etc., and the sediment is made up into balls and dried in the sun. The lumps reduced to powder are said to be a valuable medicine for kidney complaints.