Future Ecosystem Functioning Research Articles

How Light Competition between Plants Affects their Response to Climate Change

How plants respond to climate change is of major concern, as plants will strongly impact future ecosystem functioning, food production and climate. Here we investigate how vegetation structure and functioning may be influenced by predicted increases in annual temperatures and atmospheric CO2 concentration, and modeled the extent to which local plant-plant interactions may modify these effects.A canopy model was developed, which calculates photosynthesis as a function of light, nitrogen, temperature, CO2 and water availability, and considers different degrees of light competition between neighboring plants through canopy mixing; soybean (Glycine max) was used as a reference system.The model predicts increased net photosynthesis and reduced stomatal conductance and transpiration under atmospheric CO2 increase. When CO2 elevation is combined with warming, photosynthesis is increased more, but transpiration is reduced less. Intriguingly, when competition is considered the optimal response shifts to producing larger leaf areas, but with lower stomatal conductance and associated vegetation transpiration than when competition is not considered. Furthermore, only when competition is considered are the predicted effects of elevated CO2 on LAI well within the range of observed effects obtained by Free Air CO2 Enrichment (FACE) experiments. Together, our results illustrate how competition between plants may modify vegetation responses to climate change.

Native and exotic plant cover vary inversely along a climate gradient 11 years following stand-replacing wildfire in a dry coniferous forest, Oregon, USA.

Community re-assembly following future disturbances will often occur under warmer and more moisture-limited conditions than when current communities assembled. Because the establishment stage is regularly the most sensitive to climate and competition, the trajectory of recovery from disturbance in a changing environment is uncertain, but has important consequences for future ecosystem functioning. To better understand how ongoing warming and rising moisture limitation may affect recovery, we studied native and exotic plant composition 11 years following complete stand-replacing wildfire in a dry coniferous forest spanning a large gradient in climatic moisture deficit (CMD) from warm and dry low elevation sites to relatively cool and moist higher elevations sites. We then projected future precipitation, temperature and CMD at our study locations for four scenarios selected to encompass a broad range of possible future conditions for the region. Native perennials dominated relatively cool and moist sites 11 years after wildfire, but were very sparse at the warmest and driest (high CMD) sites, particularly when combined with high topographic sun exposure. In contrast, exotic species (primarily annual grasses) were dominant or co-dominant at the warmest and driest sites, especially with high topographic sun exposure. All future scenarios projected increasing temperature and CMD in coming decades (e.g., from 4.5% to 29.5% higher CMD by the 2080's compared to the 1971-2000 average), even in scenarios where growing season (May-September) precipitation increased. These results suggest increasing temperatures and moisture limitation could facilitate longer term (over a decade) transitions toward exotic-dominated communities after severe wildfire when a suitable exotic seed source is present.

Long-term exposure to elevated CO2 enhances plant community stability by suppressing dominant plant species in a mixed-grass prairie.

Climate controls vegetation distribution across the globe, and some vegetation types are more vulnerable to climate change, whereas others are more resistant. Because resistance and resilience can influence ecosystem stability and determine how communities and ecosystems respond to climate change, we need to evaluate the potential for resistance as we predict future ecosystem function. In a mixed-grass prairie in the northern Great Plains, we used a large field experiment to test the effects of elevated CO2, warming, and summer irrigation on plant community structure and productivity, linking changes in both to stability in plant community composition and biomass production. We show that the independent effects of CO2 and warming on community composition and productivity depend on interannual variation in precipitation and that the effects of elevated CO2 are not limited to water saving because they differ from those of irrigation. We also show that production in this mixed-grass prairie ecosystem is not only relatively resistant to interannual variation in precipitation, but also rendered more stable under elevated CO2 conditions. This increase in production stability is the result of altered community dominance patterns: Community evenness increases as dominant species decrease in biomass under elevated CO2. In many grasslands that serve as rangelands, the economic value of the ecosystem is largely dependent on plant community composition and the relative abundance of key forage species. Thus, our results have implications for how we manage native grasslands in the face of changing climate.

Resistance and resilience of a grassland ecosystem to climate extremes

Climate change forecasts of more frequent climate extremes suggest that such events will become increasingly important drivers of future ecosystem dynamics and function. Because the rarity and unpredictability of naturally occurring climate extremes limits assessment of their ecological impacts, we experimentally imposed extreme drought and a mid‐summer heat wave over two years in a central U.S. grassland. While the ecosystem was resistant to heat waves, it was not resistant to extreme drought, which reduced aboveground net primary productivity (ANPP) below the lowest level measured in this grassland in almost 30 years. This extreme reduction in ecosystem function was a consequence of reduced productivity of both C4grasses and C3forbs. However, the dominant forb was negatively impacted by the drought more than the dominant grass, and this led to a reordering of species abundances within the plant community. Although this change in community composition persisted post‐drought, ANPP recovered completely the year after drought due to rapid demographic responses by the dominant grass, compensating for loss of the dominant forb. Overall, these results show that an extreme reduction in ecosystem function attributable to climate extremes (e.g., low resistance) does not preclude rapid ecosystem recovery. Given that dominance by a few species is characteristic of most ecosystems, knowledge of the traits of these species and their responses to climate extremes will be key for predicting future ecosystem dynamics and function.

Convergence of terrestrial plant production across global climate gradients.

Variation in terrestrial net primary production (NPP) with climate is thought to originate from a direct influence of temperature and precipitation on plant metabolism. However, variation in NPP may also result from an indirect influence of climate by means of plant age, stand biomass, growing season length and local adaptation. To identify the relative importance of direct and indirect climate effects, we extend metabolic scaling theory to link hypothesized climate influences with NPP, and assess hypothesized relationships using a global compilation of ecosystem woody plant biomass and production data. Notably, age and biomass explained most of the variation in production whereas temperature and precipitation explained almost none, suggesting that climate indirectly (not directly) influences production. Furthermore, our theory shows that variation in NPP is characterized by a common scaling relationship, suggesting that global change models can incorporate the mechanisms governing this relationship to improve predictions of future ecosystem function.

How light competition between plants affects their response to climate change

How plants respond to climate change is of major concern, as plants will strongly impact future ecosystem functioning, food production and climate. Here, we investigated how vegetation structure and functioning may be influenced by predicted increases in annual temperatures and atmospheric CO2 concentration, and modeled the extent to which local plant-plant interactions may modify these effects. A canopy model was developed, which calculates photosynthesis as a function of light, nitrogen, temperature, CO2 and water availability, and considers different degrees of light competition between neighboring plants through canopy mixing; soybean (Glycine max) was used as a reference system. The model predicts increased net photosynthesis and reduced stomatal conductance and transpiration under atmospheric CO2 increase. When CO2 elevation is combined with warming, photosynthesis is increased more, but transpiration is reduced less. Intriguingly, when competition is considered, the optimal response shifts to producing larger leaf areas, but with lower stomatal conductance and associated vegetation transpiration than when competition is not considered. Furthermore, only when competition is considered are the predicted effects of elevated CO2 on leaf area index (LAI) well within the range of observed effects obtained by Free air CO2 enrichment (FACE) experiments. Together, our results illustrate how competition between plants may modify vegetation responses to climate change.

Contrasting sensitivities of two dominant C4 grasses to heat waves and drought

Heat waves and droughts are predicted to increase in frequency and intensity with climate change. However, we lack a mechanistic understanding of the independent and interactive effects of severe heat and water stress for most ecosystems. In a mesic tallgrass prairie ecosystem, we used a factorial experimental approach to assess ecophysiological and productivity responses of two dominant C4 grasses, Andropogon gerardii and Sorghastrum nutans, to a season-long drought and a mid-summer heat wave at four intensities. We hypothesized that drought would have greater impacts than heat waves, that combined effects would be greater than either factor alone, and that the dominant grasses would differ in their responses to heat and water heat stress. We detected significant reductions in photosynthesis, leaf water potential, and productivity with drought but few direct responses to the heat waves. Surprisingly, there was no additive effect of heat and water stress on any plant response. However, S. nutans was more sensitive than A. gerardii to drought. In this grassland, water stress will likely dominate photosynthetic and productivity responses caused by discrete drought and heat wave events, rather than direct or additive effects of heat stress, with differential sensitivity in these grasses altering future ecosystem structure and function.

The Tree of Life in ecosystems: evolution of plant effects on carbon and nutrient cycling

Summary The time is now ripe for ecologists and evolutionary biologists together to tackle a huge and important new challenge: mapping plant species effects on ecosystem functions onto the Tree of Life. This new research agenda is now tractable because of major recent advances in (i) screening plant species world‐wide for the traits that support these functions, as well as in our understanding of how these traits support these functions; (ii) genetic screening and bioinformatics to build huge molecular plant phylogenies; and (iii) the comparative methods tools to analyse traits and phylogenies together. Understanding the evolutionary dynamics of traits related to biogeochemical cycling requires concerted research activity at a range of different phylogenetic scales, from analyses that consider data from all land plants to micro‐evolution and other sources of intraspecific variation. This effort also requires study of the different plant organs involved in carbon and nutrient uptake, processing and transport, including the coordination between them. This overall effort should reveal (i) which lineages are associated with fast and which with slow turnover of carbon and nutrients (and water); (ii) how they are associated with fast versus slow turnover in terms of the traits involved; (iii) when in evolutionary history key transitions between fast and slow occurred; and (iv) what were and are the consequences of such variation in plant carbon and nutrient traits and turnover rates for past, present and future ecosystem functioning and services. Here, we introduce seven papers that together help set up this research agenda, each at a different phylogenetic level and resolution, and for different plant organs. Recent studies have begun to reveal evolutionary patterns in traits that matter to carbon and nutrient cycling for other groups of organisms, such as decomposing fungi and macrodetritivores and herbivorous animals. Ultimately such studies will all help towards the goal of mapping effects of organisms on biogeochemical cycling onto the entire Tree of Life.

Climate change and grazing interact to alter flowering patterns in the Mongolian steppe

Socio-economic changes threaten nomadic pastoralism across the world, changing traditional grazing patterns. Such land-use changes will co-occur with climate change, and while both are potentially important determinants of future ecosystem functioning, interactions between them remain poorly understood. We investigated the effects of grazing by large herbivores and climate manipulation using open-top chambers (OTCs) on flower number and flowering species richness in mountain steppe of northern Mongolia. In this region, sedentary pastoralism is replacing nomadic pastoralism, and temperature is predicted to increase. Grazing and OTCs interacted to affect forb flowering richness, which was reduced following grazing removal, and reduced by OTCs in grazed plots only. This interaction was directly linked to the soil moisture and temperature environments created by the experimental treatments: most species flowered when both soil moisture and temperature levels were high (i.e. in grazed plots without OTCs), while fewer species flowered when either temperature, or moisture, or both, were low. Removal of grazing increased the average number of graminoid flowers produced at peak flowering in Year 1, but otherwise grazing removal and OTCs did not affect community-level flower composition. Of four abundant graminoid species examined individually, three showed increased flower number with grazing removal, while one showed the reverse. Four abundant forb species showed no significant response to either treatment. Our results highlight how climate change effects on mountain steppe could be contingent on land-use, and that studies designed to understand ecosystem response to climate change should incorporate co-occurring drivers of change, such as altered grazing regimes.

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Feature: Improving our knowledge of drought‐induced forest mortality through experiments, observations, and modeling

Combined effects of warming and elevated CO2 on the impact of drought in grassland species

Aims Drought is a major growth limiting factor in the majority of terrestrial ecosystems and is expected to become more frequent in the future. Therefore, resolving the drought response of plants under changing climate conditions is crucial to our understanding of future ecosystem functioning. This study responds to the need for experimental research on the combined effects of warming, elevated CO2 and drought, and aims to determine whether the response to drought is altered under future climate conditions.

Resource limitation, tolerance, and the future of ecological plant classification.

Throughout the evolutionary history of plants, drought, shade, and scarcity of nutrients have structured ecosystems and communities globally. Humans have begun to drastically alter the prevalence of these environmental factors with untold consequences for plant communities and ecosystems worldwide. Given limitations in using organ-level traits to predict ecological performance of species, recent advances using tolerances of low resource availability as plant functional traits are revealing the often hidden roles these factors have in structuring communities and are becoming central to classifying plants ecologically. For example, measuring the physiological drought tolerance of plants has increased the predictability of differences among species in their ability to survive drought as well as the distribution of species within and among ecosystems. Quantifying the shade tolerance of species has improved our understanding of local and regional species diversity and how species have sorted within and among regions. As the stresses on ecosystems continue to shift, coordinated studies of whole-plant growth centered on tolerance of low resource availability will be central in predicting future ecosystem functioning and biodiversity. This will require efforts that quantify tolerances for large numbers of species and develop bioinformatic and other techniques for comparing large number of species.

Impairment of C4 photosynthesis by drought is exacerbated by limiting nitrogen and ameliorated by elevated [CO2] in maize

Predictions of future ecosystem function and food supply from staple C(4) crops, such as maize, depend on elucidation of the mechanisms by which environmental change and growing conditions interact to determine future plant performance. To test the interactive effects of elevated [CO(2)], drought, and nitrogen (N) supply on net photosynthetic CO(2) uptake (A) in the world's most important C(4) crop, maize (Zea mays) was grown at ambient [CO(2)] (∼385 ppm) and elevated [CO(2)] (550 ppm) with either high N supply (168 kg N ha(-1) fertilizer) or limiting N (no fertilizer) at a site in the US Corn Belt. A mid-season drought was not sufficiently severe to reduce yields, but caused significant physiological stress, with reductions in stomatal conductance (up to 57%), A (up to 44%), and the in vivo capacity of phosphoenolpyruvate carboxylase (up to 58%). There was no stimulation of A by elevated [CO(2)] when water availability was high, irrespective of N availability. Elevated [CO(2)] delayed and relieved both stomatal and non-stomatal limitations to A during the drought. Limiting N supply exacerbated stomatal and non-stomatal limitation to A during drought. However, the effects of limiting N and elevated [CO(2)] were additive, so amelioration of stress by elevated [CO(2)] did not differ in magnitude between high N and limiting N supply. These findings provide new understanding of the limitations to C(4) photosynthesis that will occur under future field conditions of the primary region of maize production in the world.

Distinct seasonal assemblages of arbuscular mycorrhizal fungi revealed by massively parallel pyrosequencing

• Understanding the dynamics of rhizosphere microbial communities is essential for predicting future ecosystem function, yet most research focuses on either spatial or temporal processes, ignoring combined spatio-temporal effects. • Using pyrosequencing, we examined the spatio-temporal dynamics of a functionally important community of rhizosphere microbes, the arbuscular mycorrhizal (AM) fungi. We sampled AM fungi from plant roots growing in a temperate grassland in a spatially explicit manner throughout a year. • Ordination analysis of the AM fungal assemblages revealed significant temporal changes in composition and structure. Alpha and beta diversity tended to be negatively correlated with the climate variables temperature and sunshine hours. Higher alpha diversity during colder periods probably reflects more even competitive interactions among AM fungal species under limited carbon availability, a conclusion supported by analysis of beta diversity which highlights how resource limitation may change localized spatial dynamics. • Results reveal distinct AM fungal assemblages in winter and summer at this grassland site. A seasonally changing supply of host-plant carbon, reflecting changes in temperature and sunshine hours, may be the driving force in regulating the temporal dynamics of AM fungal communities. Climate change effects on seasonal temperatures may therefore substantially alter future AM fungal community dynamics and ecosystem functioning.

Effects of multiple climate change factors on the tall fescue–fungal endophyte symbiosis: infection frequency and tissue chemistry

• Climate change (altered CO(2) , warming, and precipitation) may affect plant-microbial interactions, such as the Lolium arundinaceum-Neotyphodium coenophialum symbiosis, to alter future ecosystem structure and function. • To assess this possibility, tall fescue tillers were collected from an existing climate manipulation experiment in a constructed old-field community in Tennessee (USA). Endophyte infection frequency (EIF) was determined, and infected (E+) and uninfected (E-) tillers were analysed for tissue chemistry. • The EIF of tall fescue was higher under elevated CO(2) (91% infected) than with ambient CO(2) (81%) but was not affected by warming or precipitation treatments. Within E+ tillers, elevated CO(2) decreased alkaloid concentrations of both ergovaline and loline, by c. 30%; whereas warming increased loline concentrations 28% but had no effect on ergovaline. Independent of endophyte infection, elevated CO(2) reduced concentrations of nitrogen, cellulose, hemicellulose, and lignin. • These results suggest that elevated CO(2) , more than changes in temperature or precipitation, may promote this grass-fungal symbiosis, leading to higher EIF in tall fescue in old-field communities. However, as all three climate factors are likely to change in the future, predicting the symbiotic response and resulting ecological consequences may be difficult and dependent on the specific atmospheric and climatic conditions encountered.

Using niche‐based modelling to assess the impact of climate change on tree functional diversity in Europe

ABSTRACTRapid anthropogenic climate change is already affecting species distributions and ecosystem functioning worldwide. We applied niche‐based models to analyse the impact of climate change on tree species and functional diversity in Europe. Present‐day climate was used to predict the distributions of 122 tree species from different functional types (FT). We then explored projections of future distributions under one climate scenario for 2080, considering two alternative dispersal assumptions: no dispersal and unlimited dispersal. The species‐rich broadleaved deciduous group appeared to play a key role in the future of different European regions. Temperate areas were projected to lose both species richness and functional diversity due to the loss of broadleaved deciduous trees. These were projected to migrate to boreal forests, thereby increasing their species richness and functional diversity. Atlantic areas provided an intermediate case, with a predicted reduction in the numbers of species and occasional predicted gains in functional diversity. This resulted from a loss in species within the broadleaved deciduous FT, but overall maintenance of the group. Our results illustrate the fact that both species‐specific predictions and functional patterns should be examined separately in order to assess the impacts of climate change on biodiversity and gain insights into future ecosystem functioning.

Diversity components of impending primate extinctions.

Many extant species are at risk to go extinct. This impending loss of species is likely to cause changes in future ecosystem functions. Ecological components of diversity, such as dietary or habitat specializations, can be used to estimate the impact of extinctions on ecosystem functions. As an approach to estimate the impact of future extinctions, we tested interdependency between ecological and taxonomic change based on current predictions of extinction rates in primates. We analyzed the ecological characteristics of extant primate faunas having species in various categories of endangerment of extinction and forecasted the future primate faunas as if they were paleontological faunas. Predicting future faunas combines the wealth of ecological information on living primates with large, fossil record-like changes in diversity. Predicted extinction patterns of living primates in Africa, Asia, Madagascar, and South America show that changes in ecology differ among the regions in ways that are not reducible to taxonomic measures. The ecological effects of primate extinctions are initially least severe in South America and larger in Asia and Africa. Disproportionately larger ecological changes are projected for Madagascar. The use of taxonomy as a proxy for ecology can mislead when estimating competence of future primate ecosystems.