Collaborative agent-based modeling for managing shrub encroachment in an Afroalpine grassland.

We co-designed an agent-based model of an Afroalpine grassland in Ethiopia that is experiencing unwanted shrub encroachment. The goal was to enable managers of a community conservation area to better understand the drivers of shrub encroachment and to test possible management actions for controlling shrubs. Due to limited site-specific data, we parameterized this model using insights from published literature, remote sensing, and expert opinion from scientists and local managers. We therefore sought to explore potential future scenarios rather than make highly accurate predictions, focusing on facilitating discussions and learning among the diverse co-management team. We evaluated three social-ecological scenarios with our model, examining: (1) the impact of changing precipitation regimes on vegetation, (2) whether changing the frequency of guassa grass harvests would improve the long-term sustainability of the grassland, and (3) whether the combination of grass harvest and shrub removal would affect shrub encroachment. We found that the model was highly sensitive to the amount of grass harvested each year for local use. Our results indicate that the guassa grass was more resilient than shrubs during persistent dry climatic conditions, whereas a reduction in only the early spring rains (known as the "belg") resulted in considerable loss of grass biomass. While our modeling results lacked the quantitative specificity desired by managers, participants in the collaborative modeling process learned new approaches to planning and management of the conservation area and expanded their knowledge of the ecological complexity of the system. Several participants used the model as a boundary object, interpreting it in ways that reinforced their cultural values and goals for the conservation area. Our work highlights the lack of detailed scientific knowledge of Afroalpine ecosystems, and urges managers to reconnect with traditional ecological management of the conservation area in their pursuit of shrub encroachment solutions. The decline or absence of the belg rains is becoming increasingly common in the Ethiopian highlands, and our results underscore the need for more widespread understanding of how this changing climatic regime impacts local environmental management. This work lays a foundation for social-ecological research to improve both understanding and management of these highly threatened ecosystems.

Warming has a minor effect on surface soil organic carbon in alpine meadow ecosystems on the Qinghai-Tibetan Plateau.

The alpine meadow ecosystem on the Qinghai-Tibetan Plateau (QTP) is very sensitive to warming and plays a key role in regulating global carbon (C) cycling. However, how warming affects the soil organic carbon (SOC) pool and related C inputs and outputs in alpine meadow ecosystems on the QTP remains unclear. Here, we combined two field experiments and a meta-analysis on field experiments to synthesize the responses of the SOC pool and related C cycling processes to warming in alpine meadow ecosystems on the QTP. We found that the SOC content of surface soil (0-10 cm) showed a minor response to warming, but plant respiration was accelerated by warming. In addition, the warming effect on SOC was not correlated with experimental and environmental variables, such as the method, magnitude and duration of warming, initial SOC content, mean annual temperature, and mean annual precipitation. We conclude that the surface SOC content is resistant to climate warming in alpine meadow ecosystems on the QTP.

Warming and land use change concurrently erode ecosystem services in Tibet.

Alpine meadows on the Tibetan Plateau comprise the largest alpine ecosystem in the world and provide critical ecosystem services, including forage production and carbon sequestration, on which people depend from local to global scales. However, the provision of these services may be threatened by climate warming combined with land use policies that are altering if and how pastoralists can continue to graze livestock, the dominant livelihood practice in this region for millennia. We synthesized findings from a climate warming and yak grazing experiment with landscape-level observations in central Tibet to gain insight into the trajectories of change that Tibet's alpine meadows will undergo in response to expected changes in climate and land use. We show that within 5 years, experimental warming drove an alpine community with intact, sedge-dominated turfs into a degraded state. With removal of livestock, consistent with policy intended to reverse degradation, a longer-term shift to a more shrub-dominated community will likely occur. Neither degraded nor shrub meadows produce forage or sequester carbon to the same degree as intact meadows, indicating that climate warming and drying will reduce the ability of Tibet's alpine meadows to provide key ecosystem services, and that livestock reduction policies intended to counteract trajectories of land degradation instead endanger contemporary livelihoods on the Tibetan Plateau.

Plant species' origin predicts dominance and response to nutrient enrichment and herbivores in global grasslands.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

Herbivores and nutrients control grassland plant diversity via light limitation.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Eutrophication weakens stabilizing effects of diversity in natural grasslands.

Studies of experimental grassland communities have demonstrated that plant diversity can stabilize productivity through species asynchrony, in which decreases in the biomass of some species are compensated for by increases in others. However, it remains unknown whether these findings are relevant to natural ecosystems, especially those for which species diversity is threatened by anthropogenic global change. Here we analyse diversity-stability relationships from 41 grasslands on five continents and examine how these relationships are affected by chronic fertilization, one of the strongest drivers of species loss globally. Unmanipulated communities with more species had greater species asynchrony, resulting in more stable biomass production, generalizing a result from biodiversity experiments to real-world grasslands. However, fertilization weakened the positive effect of diversity on stability. Contrary to expectations, this was not due to species loss after eutrophication but rather to an increase in the temporal variation of productivity in combination with a decrease in species asynchrony in diverse communities. Our results demonstrate separate and synergistic effects of diversity and eutrophication on stability, emphasizing the need to understand how drivers of global change interactively affect the reliable provisioning of ecosystem services in real-world systems.

Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

Climate change and water use partitioning by different plant functional groups in a grassland on the Tibetan Plateau.

The Tibetan Plateau (TP) is predicted to experience increases in air temperature, increases in snowfall, and decreases in monsoon rains; however, there is currently a paucity of data that examine the ecological responses to such climate changes. In this study, we examined the effects of increased air temperature and snowfall on: 1) water use partitioning by different plant functional groups, and 2) ecosystem CO2 fluxes throughout the growing season. At the individual plant scale, we used stable hydrogen isotopes (δD) to partition water use between shallow- and deep-rooted species. Prior to the arrival of summer precipitation (typically mid-July), snowmelt was the main water source in the soils. During this time, shallow and deep-rooted species partitioned water use by accessing water from shallow and deep soils, respectively. However, once the monsoon rains arrived, all plants used rainwater from the upper soils as the main water source. Snow addition did not result in increased snowmelt use throughout the growing season; instead, snowmelt water was pushed down into deeper soils when the rains arrived. At the larger plot scale, CO2 flux measurements demonstrated that rain was the main driver for net ecosystem productivity (NEP). NEP rates were low during June and July and reached a maximum during the monsoon season in August. Warming decreased NEP through a reduction in gross primary productivity (GPP), and snow additions did not mitigate the negative effects of warming by increasing NEP or GPP. Both the isotope and CO2 flux results suggest that rain drives productivity in the Nam Tso region on the TP. This also suggests that the effects of warming-induced drought on the TP may not be mitigated by increased snowfall. Further decreases in summer monsoon rains may affect ecosystem productivity, with large implications for livestock-based livelihoods.

Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet.

Global climate change is predicted to have large impacts on the phenology and reproduction of alpine plants, which will have important implications for plant demography and community interactions, trophic dynamics, ecosystem energy balance, and human livelihoods. In this article we report results of a 3-year, fully factorial experimental study exploring how warming, snow addition, and their combination affect reproductive phenology, effort, and success of four alpine plant species belonging to three different life forms in a semiarid, alpine meadow ecosystem on the central Tibetan Plateau. Our results indicate that warming and snow addition change reproductive phenology and success, but responses are not uniform across species. Moreover, traits associated with resource acquisition, such as rooting depth and life history (early vs. late flowering), mediate plant phenology, and reproductive responses to changing climatic conditions. Specifically, we found that warming delayed the reproductive phenology and decreased number of inflorescences of Kobresia pygmaea C. B. Clarke, a shallow-rooted, early-flowering plant, which may be mainly constrained by upper-soil moisture availability. Because K. pygmaea is the dominant species in the alpine meadow ecosystem, these results may have important implications for ecosystem dynamics and for pastoralists and wildlife in the region.

Global assessment of experimental climate warming on tundra vegetation: heterogeneity over space and time.

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Productivity is a poor predictor of plant species richness.

For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters(-2)) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.

Abundance of introduced species at home predicts abundance away in herbaceous communities.

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Plant species traits are the predominant control on litter decomposition rates within biomes worldwide.

Worldwide decomposition rates depend both on climate and the legacy of plant functional traits as litter quality. To quantify the degree to which functional differentiation among species affects their litter decomposition rates, we brought together leaf trait and litter mass loss data for 818 species from 66 decomposition experiments on six continents. We show that: (i) the magnitude of species-driven differences is much larger than previously thought and greater than climate-driven variation; (ii) the decomposability of a species' litter is consistently correlated with that species' ecological strategy within different ecosystems globally, representing a new connection between whole plant carbon strategy and biogeochemical cycling. This connection between plant strategies and decomposability is crucial for both understanding vegetation-soil feedbacks, and for improving forecasts of the global carbon cycle.

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes.

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.

Experimental warming, not grazing, decreases rangeland quality on the Tibetan Plateau.

We investigated experimental warming and simulated grazing (clipping) effects on rangeland quality, as indicated by vegetation production and nutritive quality, in winter-grazed meadows and summer-grazed shrublands on the Tibetan Plateau, a rangeland system experiencing climatic and pastoral land use changes. Warming decreased total aboveground net primary productivity (ANPP) by 40 g x m(-2) x yr(-1) at the meadow habitats and decreased palatable ANPP (total ANPP minus non-palatable forb ANPP) by 10 g x m(-2) x yr(-1) at both habitats. The decreased production of the medicinal forb Gentiana straminea and the increased production of the non-palatable forb Stellera chamaejasme with warming also reduced rangeland quality. At the shrubland habitats, warming resulted in less digestible shrubs, whose foliage contains 25% digestible dry matter (DDM), replacing more digestible graminoids, whose foliage contains 60% DDM. This shift from graminoids to shrubs not only results in lower-quality forage, but could also have important consequences for future domestic herd composition. Although warming extended the growing season in non-clipped plots, the reduced rangeland quality due to decreased vegetative production and nutritive quality will likely overwhelm the improved rangeland quality associated with an extended growing season. Grazing maintained or improved rangeland quality by increasing total ANPP by 20-40 g x m(-2) x yr(-1) with no effect on palatable ANPP. Grazing effects on forage nutritive quality, as measured by foliar nitrogen and carbon content and by shifts in plant group ANPP, resulted in improved forage quality. Grazing extended the growing season at both habitats, and it advanced the growing season at the meadows. Synergistic interactions between warming and grazing were present, such that grazing mediated the warming-induced declines in vegetation production and nutritive quality. Moreover, combined treatment effects were nonadditive, suggesting that we cannot predict the combined effect of global changes and human activities from single-factor studies. Our findings suggest that the rangelands on the Tibetan Plateau, and the pastoralists who depend on them, may be vulnerable to future climate changes. Grazing can mitigate the negative warming effects on rangeland quality. For example, grazing management may be an important tool to keep warming-induced shrub expansion in check. Moreover, flexible and opportunistic grazing management will be required in a warmer future.

Plant community responses to experimental warming across the tundra biome.

Recent observations of changes in some tundra ecosystems appear to be responses to a warming climate. Several experimental studies have shown that tundra plants and ecosystems can respond strongly to environmental change, including warming; however, most studies were limited to a single location and were of short duration and based on a variety of experimental designs. In addition, comparisons among studies are difficult because a variety of techniques have been used to achieve experimental warming and different measurements have been used to assess responses. We used metaanalysis on plant community measurements from standardized warming experiments at 11 locations across the tundra biome involved in the International Tundra Experiment. The passive warming treatment increased plant-level air temperature by 1-3 degrees C, which is in the range of predicted and observed warming for tundra regions. Responses were rapid and detected in whole plant communities after only two growing seasons. Overall, warming increased height and cover of deciduous shrubs and graminoids, decreased cover of mosses and lichens, and decreased species diversity and evenness. These results predict that warming will cause a decline in biodiversity across a wide variety of tundra, at least in the short term. They also provide rigorous experimental evidence that recently observed increases in shrub cover in many tundra regions are in response to climate warming. These changes have important implications for processes and interactions within tundra ecosystems and between tundra and the atmosphere.