• Energy Research

Abstract Novel ecosystems can differ from the native systems they replaced. We used phenology measures to compare ecosystem functioning between novel exotic‐dominated and native‐dominated grasslands in the central U.S. Phenology, or timing of biological events, is affected by climate and land use changes. We assessed how phenology shifts are being altered by exotic species dominance by comparing remotely sensed Normalized Difference Vegetation Index within growing seasons at exotic‐ and native‐dominated sites along a latitudinal gradient. Exotic species were dominated by the C3 species functional group in the north and the C4 species functional group in the south. Date of senescence was an average of 36 days later in exotic than native‐dominated grasslands, and this effect was consistent across latitudes. Exotic‐dominated grasslands greened‐up an average of 10.7 days earlier than native‐dominated grasslands, but this effect was highly dependent on latitude and the plant functional group that dominated at that latitude. Green‐up differed between native and exotic sites the most in central and northern regions that had dominant C3 grasses. We estimated the effects of an increase in global temperatures on green‐up and senescence with a space‐for‐time substitution, and by comparing growing degree day differences between historical average temperatures and +2.5°C. Green‐up was significantly earlier and senescence was significantly later with a 2.5°C increase in temperature. The native–exotic difference was significantly greater than the difference due to increased temperature for senescence, but not for green‐up. Synthesis and applications. Native to exotic plant conversions in central U.S. grasslands have led to highly altered phenology, especially in terms of senescence, and this effect should be considered along with global warming in models moving forward. This conversion will have to be considered in developing estimates of how global change will affect phenology in locations where exotics are present, especially in cases where their abundance is increasing concurrent with climate change. Global change models and policy should consider exotic species invasion as an additional widespread factor behind changes in phenology.

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.

Summary During community assembly, early arriving exotic species might suppress other species to a greater extent than do native species. Because most exotics were intentionally introduced, we hypothesize there was human selection on regeneration traits during introduction. This could have occurred at the across‐ or within‐species level (e.g. during cultivar development). We tested these predictions by seeding a single species that was either native, exotic ‘wild‐type’ (from their native range), or exotic ‘cultivated’ using 28 grassland species in a glasshouse experiment. Priority effects were assessed by measuring species’ effect on establishment of species from a seed mix added 21 d later. Exotic species had higher germination and earlier emergence dates than native species, and differences were found in both ‘wild’ and ‘cultivated’ exotics. Exotic species reduced biomass and species diversity of later arriving species much more than native species, regardless of seed source. Results indicate that in situations in which priority effects are likely to be strong, effects will be greater when an exotic species arrives first than when a native species arrives first; and this difference is not merely a result of exotic species cultivation, but might be a general native–exotic difference that deserves further study.

It remains unclear whether biodiversity buffers ecosystems against climate extremes, which are becoming increasingly frequent worldwide. Early results suggested that the ecosystem productivity of diverse grassland plant communities was more resistant, changing less during drought, and more resilient, recovering more quickly after drought, than that of depauperate communities. However, subsequent experimental tests produced mixed results. Here we use data from 46 experiments that manipulated grassland plant diversity to test whether biodiversity provides resistance during and resilience after climate events. We show that biodiversity increased ecosystem resistance for a broad range of climate events, including wet or dry, moderate or extreme, and brief or prolonged events. Across all studies and climate events, the productivity of low-diversity communities with one or two species changed by approximately 50% during climate events, whereas that of high-diversity communities with 16-32 species was more resistant, changing by only approximately 25%. By a year after each climate event, ecosystem productivity had often fully recovered, or overshot, normal levels of productivity in both high- and low-diversity communities, leading to no detectable dependence of ecosystem resilience on biodiversity. Our results suggest that biodiversity mainly stabilizes ecosystem productivity, and productivity-dependent ecosystem services, by increasing resistance to climate events. Anthropogenic environmental changes that drive biodiversity loss thus seem likely to decrease ecosystem stability, and restoration of biodiversity to increase it, mainly by changing the resistance of ecosystem productivity to climate events.

AbstractTheory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.

Plant species in grasslands are often separated into groups (C(4) and C(3) grasses, and forbs) with presumed links to ecosystem functioning. Each of these in turn can be separated into native and introduced (i.e., exotic) species. Although numerous studies have compared plant traits between the traditional groups of grasses and forbs, fewer have compared native versus introduced species. Introduced grass species, which were often introduced to prevent erosion or to improve grazing opportunities, have become common or even dominant species in grasslands. By virtue of their abundances, introduced species may alter ecosystems if they differ from natives in growth and allocation patterns. Introduced grasses were probably selected nonrandomly from the source population for forage (aboveground) productivity. Based on this expectation, aboveground production is predicted to be greater and root mass fraction to be smaller in introduced than native species. We compared root and shoot distribution and tissue quality between introduced and native C(4) grass species in the Blackland Prairie region of Central Texas, USA, and then compared differences to the more well-studied divergence between C(4) grasses and forbs. Comparisons were made in experimental monocultures planted with equal-sized transplants on a common soil type and at the same density. Aboveground productivity and C:N ratios were higher, on average, in native grasses than in native forbs, as expected. Native and introduced grasses had comparable amounts of shallow root biomass and tissue C:N ratios. However, aboveground productivity and total N were lower and deep root biomass and root mass fraction were greater in native than introduced grasses. These differences in average biomass distribution and N could be important to ecosystems in cases where native and introduced grasses have been exchanged. Our results indicate that native-introduced status may be important when interpreting species effects on grassland processes like productivity and plant N accumulation.

Climate change drivers affect plant community productivity via three pathways: (i) direct effects of drivers on plants; (ii) the response of species abundances to drivers (community response); and (iii) the feedback effect of community change on productivity (community effect). The contribution of each pathway to driver-productivity relationships depends on functional traits of dominant species. We used data from three experiments in Texas, USA, to assess the role of community dynamics in the aboveground net primary productivity (ANPP) response of C4 grasslands to two climate drivers applied singly: atmospheric CO2 enrichment and augmented summer precipitation. The ANPP-driver response differed among experiments because community responses and effects differed. ANPP increased by 80-120g m(-2) per 100 μl l(-1) rise in CO2 in separate experiments with pasture and tallgrass prairie assemblages. Augmenting ambient precipitation by 128mm during one summer month each year increased ANPP more in native than in exotic communities in a third experiment. The community effect accounted for 21-38% of the ANPP CO2 response in the prairie experiment but little of the response in the pasture experiment. The community response to CO2 was linked to species traits associated with greater soil water from reduced transpiration (e.g. greater height). Community effects on the ANPP CO2 response and the greater ANPP response of native than exotic communities to augmented precipitation depended on species differences in transpiration efficiency. These results indicate that feedbacks from community change influenced ANPP-driver responses. However, the species traits that regulated community effects on ANPP differed from the traits that determined how communities responded to drivers.

To predict the ecological consequences of biodiversity loss, researchers have spent much time and effort quantifying how biological variation affects the magnitude and stability of ecological processes that underlie the functioning of ecosystems. Here we add to this work by looking at how biodiversity jointly impacts two aspects of ecosystem functioning at once: (1) the production of biomass at any single point in time (biomass/area or biomass/volume), and (2) the stability of biomass production through time (the CV of changes in total community biomass through time). While it is often assumed that biodiversity simultaneously enhances both of these aspects of ecosystem functioning, the joint distribution of data describing how species richness regulates productivity and stability has yet to be quantified. Furthermore, analyses have yet to examine how diversity effects on production covary with diversity effects on stability. To overcome these two gaps, we reanalyzed the data from 34 experiments that have manipulated the richness of terrestrial plants or aquatic algae and measured how this aspect of biodiversity affects community biomass at multiple time points. Our reanalysis confirms that biodiversity does indeed simultaneously enhance both the production and stability of biomass in experimental systems, and this is broadly true for terrestrial and aquatic primary producers. However, the strength of diversity effects on biomass production is independent of diversity effects on temporal stability. The independence of effect sizes leads to two important conclusions. First, while it may be generally true that biodiversity enhances both productivity and stability, it is also true that the highest levels of productivity in a diverse community are not associated with the highest levels of stability. Thus, on average, diversity does not maximize the various aspects of ecosystem functioning we might wish to achieve in conservation and management. Second, knowing how biodiversity affects productivity gives no information about how diversity affects stability (or vice versa). Therefore, to predict the ecological changes that occur in ecosystems after extinction, we will need to develop separate mechanistic models for each independent aspect of ecosystem functioning.