Monoamine oxidase binding not expected to significantly affect [18F]flortaucipir PET interpretation.

PURPOSE: [18F]-labeled positron emission tomography (PET) radioligands permit in vivo assessment of Alzheimer's disease biomarkers, including aggregated neurofibrillary tau (NFT) with [18F]flortaucipir. Due to structural similarities of flortaucipir with some monoamine oxidase A (MAO-A) inhibitors, this study aimed to evaluate flortaucipir binding to MAO-A and MAO-B and any potential impact on PET interpretation. METHODS: [18F]Flortaucipir autoradiography was performed on frozen human brain tissue slices, and PET imaging was conducted in rats. Dissociation constants were determined by saturation binding, association and dissociation rates were measured by kinetic binding experiments, and IC50 values were determined by competition binding. RESULTS: Under stringent wash conditions, specific [18F]flortaucipir binding was observed on tau NFT-rich Alzheimer's disease tissue and not control tissue. In vivo PET experiments in rats revealed no evidence of [18F]flortaucipir binding to MAO-A; pre-treatment with MAO inhibitor pargyline did not impact uptake or wash-out of [18F]flortaucipir. [18F]Flortaucipir bound with low nanomolar affinity to human MAO-A in a microsomal preparation in vitro but with a fast dissociation rate relative to MAO-A ligand fluoroethyl-harmol, consistent with no observed in vivo binding in rats of [18F]flortaucipir to MAO-A. Direct binding of flortaucipir to human MAO-B was not detected in a microsomal preparation. A high concentration of flortaucipir (IC50 of 1.3 µM) was found to block binding of the MAO-B ligand safinamide to MAO-B on microsomes suggesting that, at micromolar concentrations, flortaucipir weakly binds to MAO-B in vitro. CONCLUSION: These data suggest neither MAO-A nor MAO-B binding will contribute significantly to the PET signal in cortical target areas relevant to the interpretation of [18F]flortaucipir.

Intraspecific trait variability shapes leaf trait response to altered fire regimes.

BACKGROUND AND AIMS: Understanding impacts of altered disturbance regimes on community structure and function is a key goal for community ecology. Functional traits link species composition to ecosystem functioning. Changes in the distribution of functional traits at community scales in response to disturbance can be driven not only by shifts in species composition, but also by shifts in intraspecific trait values. Understanding the relative importance of these two processes has important implications for predicting community responses to altered disturbance regimes. METHODS: We experimentally manipulated fire return intervals in replicated blocks of a fire-adapted, longleaf pine (Pinus palustris) ecosystem in North Carolina, USA and measured specific leaf area (SLA), leaf dry matter content (LDMC) and compositional responses along a lowland to upland gradient over a 4 year period. Plots were burned between zero and four times. Using a trait-based approach, we simulate hypothetical scenarios which allow species presence, abundance or trait values to vary over time and compare these with observed traits to understand the relative contributions of each of these three processes to observed trait patterns at the study site. We addressed the following questions. (1) How do changes in the fire regime affect community composition, structure and community-level trait responses? (2) Are these effects consistent across a gradient of fire intensity? (3) What are the relative contributions of species turnover, changes in abundance and changes in intraspecific trait values to observed changes in community-weighted mean (CWM) traits in response to altered fire regime? KEY RESULTS: We found strong evidence that altered fire return interval impacted understorey plant communities. The number of fires a plot experienced significantly affected the magnitude of its compositional change and shifted the ecotone boundary separating shrub-dominated lowland areas from grass-dominated upland areas, with suppression sites (0 burns) experiencing an upland shift and annual burn sites a lowland shift. We found significant effects of burn regimes on the CWM of SLA, and that observed shifts in both SLA and LDMC were driven primarily by intraspecific changes in trait values. CONCLUSIONS: In a fire-adapted ecosystem, increased fire frequency altered community composition and structure of the ecosystem through changes in the position of the shrub line. We also found that plant traits responded directionally to increased fire frequency, with SLA decreasing in response to fire frequency across the environmental gradient. For both SLA and LDMC, nearly all of the observed changes in CWM traits were driven by intraspecific variation.

Rapid deforestation of a coastal landscape driven by sea-level rise and extreme events.

Climate change is driving ecological shifts in coastal regions of the world, where low topographic relief makes ecosystems particularly vulnerable to sea-level rise, salinization, storm surge, and other effects of global climate change. The consequences of rising water tables and salinity can penetrate well inland, and lead to particularly dramatic changes in freshwater forested wetlands dominated by tree species with low salt tolerance. The resulting loss of coastal forests could have significant implications to the coastal carbon cycle. We quantified the rates of vegetation change including land loss, forest loss, and shrubland expansion in North Carolina's largest coastal wildlife refuge over 35 yr. Despite its protected status, and in the absence of any active forest management, 32% (31,600 hectares) of the refuge area has changed landcover classification during the study period. A total of 1,151 hectares of land was lost to the sea and ~19,300 hectares of coastal forest habitat was converted to shrubland or marsh habitat. As much as 11% of all forested cover in the refuge transitioned to a unique land cover type-"ghost forest"-characterized by standing dead trees and fallen tree trunks. The formation of this ghost forest transition state peaked prominently between 2011 and 2012, following Hurricane Irene and a 5-yr drought, with 4,500 ± 990 hectares of ghost forest forming during that year alone. This is the first attempt to map and quantify coastal ghost forests using remote sensing. Forest losses were greatest in the eastern portion of the refuge closest to the Croatan and Pamlico Sounds, but also occurred much further inland in low-elevation areas and alongside major canals. These unprecedented rates of deforestation and land cover change due to climate change may become the status quo for coastal regions worldwide, with implications for wetland function, wildlife habitat, and global carbon cycling.

Temperature accelerates the rate fields become forests.

Secondary succession, the postdisturbance transition of herbaceous to woody-dominated ecosystems, occurs faster at lower latitudes with important ramifications for ecosystem processes. This pattern could be driven by the direct effect of temperature on tree growth; however, an alternative mechanism is tree-herb competition, which may be more intense in more fertile northern soils. We manipulated soil fertility and herbaceous species composition in identical experiments at six sites spanning the Eastern United States (30-43° N) and monitored the growth and survival of four early successional trees. Tree seedling mass 2 years after sowing was strongly associated with site differences in mean growing season temperature, regardless of species or soil treatment. The effect of temperature was twofold: seedlings grew faster in response to warmer site temperatures, but also due to the reduction of competitive interference from the herbaceous community, which was inhibited in warmer sites. Our results suggest that increasing temperatures will promote a faster transition of fields to forests in temperate ecosystems.

Warming reverses top-down effects of predators on belowground ecosystem function in Arctic tundra.

Predators can disproportionately impact the structure and function of ecosystems relative to their biomass. These effects may be exacerbated under warming in ecosystems like the Arctic, where the number and diversity of predators are low and small shifts in community interactions can alter carbon cycle feedbacks. Here, we show that warming alters the effects of wolf spiders, a dominant tundra predator, on belowground litter decomposition. Specifically, while high densities of wolf spiders result in faster litter decomposition under ambient temperatures, they result, instead, in slower decomposition under warming. Higher spider densities are also associated with elevated levels of available soil nitrogen, potentially benefiting plant production. Changes in decomposition rates under increased wolf spider densities are accompanied by trends toward fewer fungivorous Collembola under ambient temperatures and more Collembola under warming, suggesting that Collembola mediate the indirect effects of wolf spiders on decomposition. The unexpected reversal of wolf spider effects on Collembola and decomposition suggest that in some cases, warming does not simply alter the strength of top-down effects but, instead, induces a different trophic cascade altogether. Our results indicate that climate change-induced effects on predators can cascade through other trophic levels, alter critical ecosystem functions, and potentially lead to climate feedbacks with important global implications. Moreover, given the expected increase in wolf spider densities with climate change, our findings suggest that the observed cascading effects of this common predator on detrital processes could potentially buffer concurrent changes in decomposition rates.

Succession, regression and loss: does evidence of saltwater exposure explain recent changes in the tree communities of North Carolina's Coastal Plain?

BACKGROUND AND AIMS: Coastal plant communities globally are highly vulnerable to future sea-level rise and storm damage, but the extent to which these habitats are affected by the various environmental perturbations associated with chronic salinization remains unclear. In this study, we examine the relationship between North Carolina wetland tree community composition and basal area change and indicators of salinization such as soil salt ion content and elevation. METHODS: We surveyed 34 forest plots in forested, freshwater wetlands across the Albemarle-Pamlico Peninsula. A subset of our study sites had been sampled during the previous decade as part of the Carolina Vegetation Survey, enabling us to investigate the environmental effects on current community structure, and community change over time. KEY RESULTS: Multi-variate (ordination) analysis and linear regression models of tree community composition revealed that elevation and soil salt content were correlated with changes in total site tree basal area. Shifts in tree community composition were, however, only weakly correlated with indicators of salinization, specifically elevation, soil sulphate and sodium, but not chloride. While the majority of plots experienced gains in basal area over the past decade, consistent with secondary succession, sites with high soil salt content or low elevation experienced basal area (biomass) loss during the same period. CONCLUSIONS: The key factors associated with chronic saltwater intrusion (soil ion content) likely explain recent changes in tree biomass, and potential shifts in community composition in low-elevation sites along the North Carolina coast. Not only is it probable that other coastal forest ecosystems worldwide will experience similar stressors and shifts in community biomass and structure as sea levels rise, but the ability of these habitats to deliver key ecosystem services like carbon sequestration and flood defence will be compromised as a result.

Functional trait similarity predicts survival in rare plant reintroductions.

Rare species reintroductions are an increasingly common conservation strategy, but often result in poor survival of reintroduced individuals. Reintroduction sites are chosen primarily based on historical occupancy and/or abiotic properties of the site, with much less consideration given to properties of the larger biotic community. However, ecological niche theory suggests that the ability to coexist with other species is determined in part by the degree of functional similarity between species. The degree to which functional similarity affects the survival of reintroduced plants is poorly understood, but has important implications for the allocation of limited conservation resources. We collected a suite of abiotic, biotic, and functional trait variables centered on outplanted individuals from four reintroduced rare plant species and used logistic regression and model selection to assess their influence on individual survival. We show that higher functional similarity between reintroduced individuals and the local community, measured by differences between their multivariate functional traits and the community-weighted mean traits of their immediate neighbors, increases survival and is a stronger predictor of survival than local variation in abiotic factors, suggesting that the functional composition of the biotic community should be incorporated into site selection to improve reintroduction success.

Impacts of female body size on cannibalism and juvenile abundance in a dominant arctic spider.

Body size influences an individual's physiology and the nature of its intra- and interspecific interactions. Changes in this key functional trait can therefore have important implications for populations as well. For example, among invertebrates, there is typically a positive correlation between female body size and reproductive output. Increasing body size can consequently trigger changes in population density, population structure (e.g. adult to juvenile ratio) and the strength of intraspecific competition. Body size changes have been documented in several species in the Arctic, a region that is warming rapidly. In particular, wolf spiders, one of the most abundant arctic invertebrate predators, are becoming larger and therefore more fecund. Whether these changes are affecting their populations and role within food webs is currently unclear. We investigated the population structure and feeding ecology of the dominant wolf spider species Pardosa lapponica at two tundra sites where adult spiders naturally differ in mean body size. Additionally, we performed a mesocosm experiment to investigate how variation in wolf spider density, which is likely to change as a function of body size, influences feeding ecology and its sensitivity to warming. We found that juvenile abundance is negatively associated with female size and that wolf spiders occupied higher trophic positions where adult females were larger. Because female body size is positively related to fecundity in P. lapponica, the unexpected finding of fewer juveniles with larger females suggests an increase in density-dependent cannibalism as a result of increased intraspecific competition for resources. Higher rates of density-dependent cannibalism are further supported by the results from our mesocosm experiment, in which individuals occupied higher trophic positions in plots with higher wolf spider densities. We observed no changes in wolf spider feeding ecology in association with short-term experimental warming. Our results suggest that body size variation in wolf spiders is associated with variation in intraspecific competition, feeding ecology and population structure. Given the widespread distribution of wolf spiders in arctic ecosystems, body size shifts in these predators as a result of climate change could have implications for lower trophic levels and for ecosystem functioning.

In search of microbial indicator taxa: shifts in stream bacterial communities along an urbanization gradient.

A majority of environmental studies describe microbiomes at coarse scales of taxonomic resolution (bacterial community, phylum), ignoring key ecological knowledge gained from finer-scales and microbial indicator taxa. Here, we characterized the distribution of 940 bacterial taxa from 41 streams along an urbanization gradient (0%-83% developed watershed area) in the Raleigh-Durham area of North Carolina (USA). Using statistical approaches derived from macro-organismal ecology, we found that more bacterial taxa were classified as intolerant than as tolerant to increasing watershed urbanization (143 vs 48 OTUs), and we identified a threshold of 12.1% developed watershed area beyond which the majority of intolerant taxa were lost from streams. Two bacterial families strongly decreased with urbanization: Acidobacteriaceae (Acidobacteria) and Xanthobacteraceae (Alphaproteobacteria). Tolerant taxa were broadly distributed throughout the bacterial phylogeny, with members of the Comamonadaceae family (Betaproteobacteria) presenting the highest number of tolerant taxa. Shifts in microbial community structure were strongly correlated with a stream biotic index, based on macroinvertebrate composition, suggesting that microbial assemblages could be used to establish biotic criteria for monitoring aquatic ecosystems. In addition, our study shows that classic methods in community ecology can be applied to microbiome datasets to identify reliable microbial indicator taxa and determine the environmental constraints on individual taxa distributions along environmental gradients.

Species' traits do not converge on optimum values in preferred habitats.

Plant trait expression is shaped by filters, which can alter trait means and variances, theoretically driving species toward an "optimum" trait value for a set of environmental conditions. Recent research has highlighted the ubiquity of intraspecific variation in functional traits, which can cause plants to diverge from a hypothesized "optimum". We examined whether species occurring in "core" habitats (where they occur frequently, abundantly, and consistently) express traits that are nearer to "optimum", as captured by the community-weighted mean (CWM). We also asked whether trait variance showed signs of environmental filtering. We used cluster analysis to group plots based on environmental factors along a wet-to-dry ecotone. We used indicator species analysis to identify species with strong associations within each cluster. Trait means and variances were compared, and evidence of variance filtering was tested using a null-model approach. Trait means and trait variances respond to local-scale environmental filtering and species in core habitats were not necessarily nearer to the CWM than in other habitats. Intraspecific trait variability shows a strong signal of filtering, as variability was reduced for nearly all species and all traits compared to estimates of variability generated in the absence of environmental filtering. Our results provide strong evidence that species traits are not necessarily near "optimum" trait values in core habitats, and that trait distributions within species are strongly shaped by the environment. Future analyses should account for this divergence when calculating metrics of functional diversity, and extrapolating to ecosystem function.

Invasive species' leaf traits and dissimilarity from natives shape their impact on nitrogen cycling: a meta-analysis.

Many exotic species have little apparent impact on ecosystem processes, whereas others have dramatic consequences for human and ecosystem health. There is growing evidence that invasions foster eutrophication. We need to identify species that are harmful and systems that are vulnerable to anticipate these consequences. Species' traits may provide the necessary insights. We conducted a global meta-analysis to determine whether plant leaf and litter functional traits, and particularly leaf and litter nitrogen (N) content and carbon: nitrogen (C : N) ratio, explain variation in invasive species' impacts on soil N cycling. Dissimilarity in leaf and litter traits among invaded and noninvaded plant communities control the magnitude and direction of invasion impacts on N cycling. Invasions that caused the greatest increases in soil inorganic N and mineralization rates had a much greater litter N content and lower litter C : N in the invaded than the reference community. Trait dissimilarities were better predictors than the trait values of invasive species alone. Quantifying baseline community tissue traits, in addition to those of the invasive species, is critical to understanding the impacts of invasion on soil N cycling.

Functional traits of the understory plant community of a pyrogenic longleaf pine forest across environmental gradients.

Understanding and predicting the response of plant communities to environmental changes and disturbances such as fire requires an understanding of the functional traits present in the system, including within and across species variability, and their dynamics over time. These data are difficult to obtain as few studies provide comprehensive information for more than a few traits or species, rarely cover more than a single growing season, and usually present only summary statistics of trait values. As part of a larger study seeking to understand the dynamics of plant communities in response to different prescribed fire regimes, we measured the functional traits of the understory plant communities located in over 140 permanent plots spanning strong gradients in soil moisture in a pyrogenic longleaf pine forest in North Carolina, USA, over a four-year period from 2011 and 2014. We present over 120,000 individual trait measurements from over 130 plant species representing 91 genera from 47 families. We include data on the following 18 traits: specific leaf area, leaf dry matter content, leaf area, leaf length, leaf width, leaf perimeter, plant height, leaf nitrogen, leaf carbon, leaf carbon to nitrogen ratio, water use efficiency, time to ignition, maximum flame height, maximum burn temperature, mass-specific burn time, mass-specific smolder time, branching architecture, and the ratio of leaf matter consumed by fire. We also include information on locations, soil moisture, relative elevation, soil bulk density, and fire histories for each site.

Increased grassland arthropod production with mammalian herbivory and eutrophication: a test of mediation pathways.

Increases in nutrient availability and alterations to mammalian herbivore communities are a hallmark of the Anthropocene, with consequences for the primary producer communities in many ecosystems. While progress has advanced understanding of plant community responses to these perturbations, the consequences for energy flow to higher trophic levels in the form of secondary production are less well understood. We quantified arthropod biomass after manipulating soil nutrient availability and wild mammalian herbivory, using identical methods across 13 temperate grasslands. Of experimental increases in nitrogen, phosphorus, and potassium, only treatments including nitrogen resulted in significantly increased arthropod biomass. Wild mammalian herbivore removal had a marginal, negative effect on arthropod biomass, with no interaction with nutrient availability. Path analysis including all sites implicated nutrient content of the primary producers as a driver of increased arthropod mean size, which we confirmed using 10 sites for which we had foliar nutrient data. Plant biomass and physical structure mediated the increase in arthropod abundance, while the nitrogen treatments accounted for additional variation not explained by our measured plant variables. The mean size of arthropod individuals was 2.5 times more influential on the plot-level total arthropod biomass than was the number of individuals. The eutrophication of grasslands through human activity, especially nitrogen deposition, thus may contribute to higher production of arthropod consumers through increases in nutrient availability across trophic levels.

Trait space of rare plants in a fire-dependent ecosystem.

The causes of species rarity are of critical concern because of the high extinction risk associated with rarity. Studies examining individual rare species have limited generality, whereas trait-based approaches offer a means to identify functional causes of rarity that can be applied to communities with disparate species pools. Differences in functional traits between rare and common species may be indicative of the functional causes of species rarity and may therefore be useful in crafting species conservation strategies. However, there is a conspicuous lack of studies comparing the functional traits of rare species and co-occurring common species. We measured 18 important functional traits for 19 rare and 134 common understory plant species from North Carolina's Sandhills region and compared their trait distributions to determine whether there are significant functional differences that may explain species rarity. Flowering, fire, and tissue-chemistry traits differed significantly between rare and common, co-occurring species. Differences in specific traits suggest that fire suppression has driven rarity in this system and that changes to the timing and severity of prescribed fire may improve conservation success. Our method provides a useful tool to prioritize conservation efforts in other systems based on the likelihood that rare species are functionally capable of persisting.

Joint effects of nutrient addition and enemy exclusion on exotic plant success.

Worldwide, ecosystems are increasingly dominated by exotic plant species, a shift hypothesized to result from numerous ecological factors. Two of these, increased resource availability and enemy release, may act in concert to increase exotic success in plant communities (Resource-Enemy Release Hypothesis, R-ERH). To test this, we manipulated the availability of soil nutrients and access of vertebrate herbivores, insect herbivores, and fungal pathogens to intact grassland communities containing both native and exotic species. Our results supported both conditions necessary for R-ERH. First, exotics were less damaged than natives, experiencing less foliar damage (insect herbivory and fungal disease) than native species, particularly in communities where soil nutrients were added. Second, fertilization increased foliar damage on native species, but not exotic species. As well as fulfilling both conditions for R-ERH, these results demonstrate the importance of considering the effects of resource availability when testing for enemy release. When both conditions are fulfilled, R-ERH predicts that increasing resource availability will increase exotic abundance only in the presence of enemies. Our results fully supported this prediction for vertebrate herbivores: fertilization increased exotic cover only in communities exposed to vertebrate herbivores. Additionally, the prediction was partially supported for insect herbivores and fungal pathogens, excluding these enemies reduced exotic cover as predicted, but inconsistent with R-ERH, this effect occurred only in unfertilized communities. These results highlight the need to consider the influence of multiple enemy guilds on community processes like exotic plant invasions. Moreover, this study experimentally demonstrates that resource availability and natural enemies can jointly influence exotic success in plant communities.

Restored forested wetland surprisingly resistant to experimental salinization.

Salinization of coastal freshwater wetlands is an increasingly common and widespread phenomenon resulting from climate change. The ecosystem consequences of added salinity are poorly constrained and highly variable across prior observational and experimental studies. We added 1.8 metric tons of marine salts to replicated 200 m2 plots within a restored forested wetland in Eastern North Carolina over the course of four years. Based on prior small-scale experiments at this site, we predicted that salinization would lead to slower tree growth and suppressed soil carbon cycling. Results from this large-scale field experiment were subtle and inconsistent over space and time. By the fourth year of the experiment, we observed the predicted suppression of soil respiration and a reduction of water extractable carbon from soils receiving salt treatments. However, we found no cumulative effects of four years of salinization on total soil carbon stocks, tree growth, or root biomass. We observed substantial variation in soil solution chemistry (notably, pH and base saturation) across replicated treatment blocks; the effective salt levels, ionic composition, and pH varied following treatment depending upon pre-existing differences in edaphic factors. Our multi-year monitoring also revealed an underlying trend of wetland acidification across the entire site, a suspected effect of ecosystem recovery following wetland restoration on former agricultural land. The overwhelming resistance to our salt treatments could be attributed to the vigor of a relatively young, healthy wetland ecosystem. The heterogeneous responses to salt that we observed over space and time merits further investigation into the environmental factors that control carbon cycling in wetlands. This work highlights the importance of multi-year, large-scale field experiments for investigating ecosystem responses to global environmental change.

Transitions from functionalization to fragmentation reactions of laboratory secondary organic aerosol (SOA) generated from the OH oxidation of alkane precursors.

Functionalization (oxygen addition) and fragmentation (carbon loss) reactions governing secondary organic aerosol (SOA) formation from the OH oxidation of alkane precursors were studied in a flow reactor in the absence of NO(x). SOA precursors were n-decane (n-C10), n-pentadecane (n-C15), n-heptadecane (n-C17), tricyclo[5.2.1.0(2,6)]decane (JP-10), and vapors of diesel fuel and Southern Louisiana crude oil. Aerosol mass spectra were measured with a high-resolution time-of-flight aerosol mass spectrometer, from which normalized SOA yields, hydrogen-to-carbon (H/C) and oxygen-to-carbon (O/C) ratios, and C(x)H(y)+, C(x)H(y)O+, and C(x)H(y)O(2)+ ion abundances were extracted as a function of OH exposure. Normalized SOA yield curves exhibited an increase followed by a decrease as a function of OH exposure, with maximum yields at O/C ratios ranging from 0.29 to 0.74. The decrease in SOA yield correlates with an increase in oxygen content and decrease in carbon content, consistent with transitions from functionalization to fragmentation. For a subset of alkane precursors (n-C10, n-C15, and JP-10), maximum SOA yields were estimated to be 0.39, 0.69, and 1.1. In addition, maximum SOA yields correspond with a maximum in the C(x)H(y)O+ relative abundance. Measured correlations between OH exposure, O/C ratio, and H/C ratio may enable identification of alkane precursor contributions to ambient SOA.

Drivers of secondary succession rates across temperate latitudes of the Eastern USA: climate, soils, and species pools.

Climate change is widely expected to induce large shifts in the geographic distribution of plant communities, but early successional ecosystems may be less sensitive to broad-scale climatic trends because they are driven by interactions between species that are only indirectly related to temperature and rainfall. Building on a biogeographic analysis of secondary succession rates across the Eastern Deciduous Forest (EDF) of North America, we describe an experimental study designed to quantify the relative extent to which climate, soil properties, and geographic species pools drive variation in woody colonization rates of old fields across the EDF. Using a network of five sites of varying soil fertility spanning a latitudinal gradient from central New York to northern Florida, we added seeds of nine woody pioneer species to recently tilled old fields and monitored first-year growth and survivorship. Results suggest seedlings of southern woody pioneer species are better able to quickly establish in fields after abandonment, regardless of climate regime. Sites of lower soil fertility also exhibited faster rates of seedling growth, likely due to the slower development of the successional herbaceous community. We suggest that climate plays a relatively minor role in community dynamics at the onset of secondary succession, and that site edaphic conditions are a stronger determinant of the rate at which ecosystems develop to a woody-dominated state. More experimental research is necessary to determine the nature of the herbaceous-woody competitive interface and its sensitivity to environmental conditions.

Effects of biodiversity on the functioning of trophic groups and ecosystems.

Over the past decade, accelerating rates of species extinction have prompted an increasing number of studies to reduce species diversity experimentally and examine how this alters the efficiency by which communities capture resources and convert those into biomass. So far, the generality of patterns and processes observed in individual studies have been the subjects of considerable debate. Here we present a formal meta-analysis of studies that have experimentally manipulated species diversity to examine how it affects the functioning of numerous trophic groups in multiple types of ecosystem. We show that the average effect of decreasing species richness is to decrease the abundance or biomass of the focal trophic group, leading to less complete depletion of resources used by that group. At the same time, analyses reveal that the standing stock of, and resource depletion by, the most species-rich polyculture tends to be no different from that of the single most productive species used in an experiment. Of the known mechanisms that might explain these trends, results are most consistent with what is called the 'sampling effect', which occurs when diverse communities are more likely to contain and become dominated by the most productive species. Whether this mechanism is widespread in natural communities is currently controversial. Patterns we report are remarkably consistent for four different trophic groups (producers, herbivores, detritivores and predators) and two major ecosystem types (aquatic and terrestrial). Collectively, our analyses suggest that the average species loss does indeed affect the functioning of a wide variety of organisms and ecosystems, but the magnitude of these effects is ultimately determined by the identity of species that are going extinct.

Impacts of plant diversity on biomass production increase through time because of species complementarity.

Accelerating rates of species extinction have prompted a growing number of researchers to manipulate the richness of various groups of organisms and examine how this aspect of diversity impacts ecological processes that control the functioning of ecosystems. We summarize the results of 44 experiments that have manipulated the richness of plants to examine how plant diversity affects the production of biomass. We show that mixtures of species produce an average of 1.7 times more biomass than species monocultures and are more productive than the average monoculture in 79% of all experiments. However, in only 12% of all experiments do diverse polycultures achieve greater biomass than their single most productive species. Previously, a positive net effect of diversity that is no greater than the most productive species has been interpreted as evidence for selection effects, which occur when diversity maximizes the chance that highly productive species will be included in and ultimately dominate the biomass of polycultures. Contrary to this, we show that although productive species do indeed contribute to diversity effects, these contributions are equaled or exceeded by species complementarity, where biomass is augmented by biological processes that involve multiple species. Importantly, both the net effect of diversity and the probability of polycultures being more productive than their most productive species increases through time, because the magnitude of complementarity increases as experiments are run longer. Our results suggest that experiments to date have, if anything, underestimated the impacts of species extinction on the productivity of ecosystems.