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1. Grasses using the C4 photosynthetic pathway dominate today's savanna ecosystems and account for ~20% of terrestrial carbon fixation. However, this dominant status was reached only recently, during a period of C4 grassland expansion in the Late Miocene and Early Pliocene (4–8 Myr ago). Declining atmospheric CO2 has long been considered the key driver of this event, but new geological evidence casts doubt on the idea, forcing a reconsideration of the environmental cues for C4 plant success. 2. Here, I evaluate the current hypotheses and debate in this field, beginning with a discussion of the role of CO2 in the evolutionary origins, rather than expansion, of C4 grasses. Atmospheric CO2 starvation is a plausible selection agent for the C4 pathway, but a time gap of around 10 Myr remains between major decreases in CO2 during the Oligocene, and the earliest current evidence of C4 plants. 3. An emerging ecological perspective explains the Miocene expansion of C4 grasslands via changes in climatic seasonality and the occurrence of fire. However, the climatic drivers of this event are debated and may vary among geographical regions. 4. Uncertainty in these areas could be reduced significantly by new directions in ecological research, especially the discovery that grass species richness along rainfall gradients shows contrasting patterns in different C4 clades. By re-evaluating a published data set, I show that increasing seasonality of rainfall is linked to changes in the relative abundance of the major C4 grass clades Paniceae and Andropogoneae. I propose that the explicit inclusion of these ecological patterns would significantly strengthen climate change hypotheses of Miocene C4 grassland expansion. Critically, they allow a new series of testable predictions to be made about the fossil record. 5. Synthesis. This paper offers a novel framework for integrating modern ecological patterns into theories about the geological history of C4 plants. [ABSTRACT FROM AUTHOR]

Asexual reproduction plays a fundamental role in the structure, dynamics and persistence of perennial grasslands. Thus, assessing how asexual reproductive traits of plant communities respond to drought may be key for understanding grassland resistance to drought and recovery following drought.Here, we quantified three asexual reproductive traits (i.e. above‐ground tiller abundance, below‐ground bud abundance and the ratio of tillers to buds) during a 4‐year severe drought and a 2‐year drought recovery period in four grasslands that spanned an aridity gradient in northern China. We also assessed the relationship between these traits and the resistance and recovery of above‐ground net primary productivity (ANPP).We found that drought had limited and largely inconsistent effects on asexual reproduction among drought and recovery years and grasslands overall. Drought increased tiller abundance in the first treatment year and reduced bud banks by the fourth treatment year across grasslands. However, neither of the three asexual reproductive traits were correlated with drought resistance of ANPP. Drought legacies differed among the four grasslands with positive, negative and no legacies evident for the three asexual reproductive traits, and no clear relationship with aridity. Bud banks and tiller to bud ratio decreased and increased, respectively, in the first recovery year, but not in the second recovery year. In contrast to drought resistance, community bud abundance was strongly related to recovery, such that communities with higher bud abundance had greater ANPP recovery following drought.Synthesis. These results suggest that asexual reproductive traits may be important drivers of ecosystem recovery after drought, but that variable responses of these asexual reproduction traits during drought complicates predictions of overall grassland responses. [ABSTRACT FROM AUTHOR]

Forest–grassland mosaics, with abrupt boundaries between the two vegetation types, occur across the globe. Fire and herbivory are widely considered primary drivers that maintain these mosaics by limiting tree establishment in grasslands, while edaphic factors and frosts are generally considered to be secondary factors that reinforce these effects. However, the relative importance of these drivers likely varies across systems. In particular, although frost is known to occur in many montane tropical mosaics, experimental evidence for its role as a driving factor is limited.We used replicated in situ transplant and warming experiments to examine the role of microclimate (frost and freezing temperatures) and soil in influencing germination and seedling survival of both native forest trees and alien invasive Acacia trees in grasslands of a tropical montane forest–grassland mosaic in the Western Ghats of southern India.Seed germination of both native and alien tree species was higher in grasslands regardless of soil type, indicating that germination was not the limiting stage to tree establishment. However, irrespective of soil type, native seedlings in grasslands incurred high mortality following winter frosts and freezing temperatures relative to native seedlings in adjoining forests where freezing temperatures did not occur. Seedling survival through the tropical winter was thus a primary limitation to native tree establishment in grasslands. In contrast, alien Acacia seedlings in grasslands incurred much lower levels of winter mortality. Experimental night‐time warming in grasslands significantly enhanced over‐winter survival of all tree seedlings, but increases were much greater for alien Acacia than for native tree seedlings.Synthesis. Our results provide evidence for a primary role for frost and freezing temperatures in limiting tree establishment in grasslands of this tropical montane forest–grassland mosaic. Future increases in temperature are likely to release trees from this limitation and favour tree expansion into grasslands, with rates of expansion of non‐native Acacia likely to be much greater than that of native trees. We suggest that studies of frost limitation to plant establishment are needed across a range of tropical ecosystems to re‐evaluate the general importance of frost as a driver of vegetation transitions in the tropics. [ABSTRACT FROM AUTHOR]

Plant traits can provide unique insights into plant performance at the community scale. Functional composition, defined by both functional diversity and community‐weighted trait means (CWMs), can affect the stability of above‐ground net primary production (ANPP) in response to climate extremes. Further complexity arises, however, when functional composition itself responds to environmental change. The duration of climate extremes, such as drought, is expected to increase with rising global temperatures; thus, understanding the impacts of long‐term drought on functional composition and the corresponding effect that has on ecosystem function could improve predictions of ecosystem sensitivity to climate change.We experimentally reduced growing season precipitation by 66% across six temperate grasslands for 4 years and measured changes in three indices of functional diversity (functional dispersion, richness and evenness), community‐weighted trait means and phylogenetic diversity (PD). Specific leaf area (SLA), leaf nitrogen content (LNC) and (at most sites) leaf turgor loss point (πTLP) were measured for species cumulatively representing ~90% plant cover at each site.Long‐term drought led to increased community functional dispersion in three sites, with negligible effects on the remaining sites. Species re‐ordering following the mortality/senescence of dominant species was the main driver of increased functional dispersion. The response of functional diversity was not consistently matched by changes in phylogenetic diversity. Community‐level drought strategies (assessed as CWMs) largely shifted from drought tolerance to drought avoidance and/or escape strategies, as evidenced by higher community‐weighted πTLP, SLA and LNC. Lastly, ecosystem drought sensitivity (i.e. relative reduction in ANPP in drought plots) was positively correlated with community‐weighted SLA and negatively correlated with functional diversity.Synthesis. Increased functional diversity following long‐term drought may stabilize ecosystem functioning in response to future drought. However, shifts in community‐scale drought strategies may increase ecosystem drought sensitivity, depending on the nature and timing of drought. Thus, our results highlight the importance of considering both functional diversity and abundance‐weighted traits means of plant communities as their collective effect may either stabilize or enhance ecosystem sensitivity to drought. [ABSTRACT FROM AUTHOR]

Assembly of grassland communities has long been scrutinized through the lens of functional diversity. Studies generally point to an overwhelming influence of climate on observed patterns of functional diversity, despite experimental evidence demonstrating the importance of biotic interactions. We postulate that this is because most observational studies neglect both scale dependencies of assembly processes and phenotypic variation between individuals. Here, we test for changes in the importance of abiotic filtering and biotic interactions along a stress gradient by explicitly accounting for different scales. In addition to quantifying intraspecific trait variability (ITV), we also vary the two components of spatial scale, including grain (i.e. community size) and extent (i.e. the geographical area that defines the species pool)., We sampled 20 grassland communities in ten sites distributed along a 975-m elevation gradient. At each site, we measured seven functional traits for a total of 2020 individuals at different spatial grains. We related community functional diversity metrics to the main environmental gradient of our study area, growing season length (GSL), and assessed the dependence of these relationships on spatial grain, spatial extent and ITV., At large spatial grain and extent, the imprint of environmental filtering on functional diversity became more important with increasing stress (i.e. functional diversity decreased with shorter GSL). At small spatial grain and extent, we found a convex relationship between functional diversity and GSL congruent with the hypothesis that competition is dominant at low-stress levels while facilitative interactions are dominant at high-stress levels (i.e. high functional diversity at both extremes of the stress gradient). Importantly, the effect of intraspecific variability on assembly rules was noticeable only at small spatial grain and extent., Synthesis. Our study reveals how the combination of abiotic stress and biotic interactions shapes the functional diversity of alpine grasslands at different spatial scales, and highlights the importance of phenotype variation between individuals for community assembly processes at fine spatial scale. Our results suggest that studies analysing trait-based assembly rules but ignoring ITV and focusing on a single spatial scale are likely to miss essential features of community diversity patterns. [ABSTRACT FROM AUTHOR]

1. Grasslands cover ~30% of the Earth's terrestrial surface and provide many ecosystem services. Many grasslands are heavily managed to maximize these services for human benefit, but the outcome of management is anticipated to be increasingly influenced by various aspects of climate change and elevated atmospheric CO2. The relative importance of global change vs. land management on grasslands is largely unknown. 2. A meta-analysis is used here to examine drivers at both scales primarily targeting services provided by grasslands relating to plant productivity (above- and below-ground biomass) and soil processes (nutrients and soil respiration) in 38 manipulative experiments published in the last decade. We specifically target effects of (i) single and combined land management practices (LMs), (ii) single and combined factors relating to broad-scale climate change and elevated CO2, and (iii) combined management practices and changes to climate and CO2. Collectively, this examines the general efficacy of global change models in predicting changes to grassland functioning. 3. We found that combinations of management practices had approximately double the explanatory power for variation in grassland services compared with individual or interactive effects of factors associated with climate change and CO2. These interacting management practices such as nutrient additions and defoliation predominantly influenced functions associated with productivity or biomass both below and above ground. The effects of interacting factors of climate and CO2 influenced a wider range of ecosystem functions, but the magnitude of these effects was relatively smaller. 4. Interactions between management practices or between climate change/CO2 factors always had higher explanatory power than any factor in isolation indicating that multivariate synergistic models of environmental change can better describe impacts on ecosystem function in plant communities (e.g. relative to univariate climate-based models). Given that the magnitude and direction (positive or negative) of the interactions varied widely, this also implies that the outcomes of these multivariate interactions can vary spatially, temporally or by immediate context (e.g. management prescriptions). 5. Synthesis. Although our work confirms how climate change and CO2 can affect many ecosystembased functional attributes, it suggests that combinations of LMs remain the dominant set of factors in determining the performance of grassland plant communities. Land management may thus be critical for influencing projected responses to future climate change and elevated CO2 in models of grassland function at least for factors relating to primary production. [ABSTRACT FROM AUTHOR]

Under climate change, alterations to precipitation and temperature regimes are expected to impact ecosystem structure and function. These impacts may in part be determined by feedbacks between plants and associated microbes, including both endophytic fungal symbionts and viral pathogens., To test potential key components of these feedbacks in a grassland system, we conducted a field experiment that factorially manipulated temperature regime, precipitation regime and endophyte infection of a dominant species, Schedonorus arundinaceus. After one growing season, we evaluated the four most frequently occurring grass species for infection by three species of barley and cereal yellow dwarf viruses (B/ CYDVs). For S. arundinaceus, we also tracked changes in viral prevalence over an additional two growing seasons, for a total of three growing seasons., Plant species identity played a strong role in determining virus prevalence; Poa pratensis and Paspalum dilatatum were more frequently infected than either S. arundinaceus (endophyte infected or endophyte free) or Eleusine indica. Both elevated temperature and elevated precipitation impacted virus prevalence, but effects varied among years and interacted with fungal endophyte symbiosis for S. arundinaceus., In the final year of data collection, elevated precipitation increased virus prevalence in endophyte-infected, but not endophyte-free, S. arundinaceus. In the same year, but regardless of endophyte infection, elevated temperature by itself increased virus prevalence for S. arundinaceus. This effect of temperature on virus prevalence may have been driven by vector abundance because the viruses are obligately aphid-transmitted, and aphids were found more frequently in heated plots., Synthesis. Overall, our investigation experimentally demonstrates strong combined impacts of biotic and abiotic factors on disease dynamics in a grassland system. Impacts of climate change on virus prevalence in grasslands may depend on the responses of aphid vectors and the presence of endophytic fungal symbionts. [ABSTRACT FROM AUTHOR]

1. Life table response experiment (LTRE) analyses have been widely used to examine the sources of differences in the long-term deterministic growth rate ( r = log λ) of stage-structured populations that live in spatially distinct habitats or under distinct experimental conditions. However, existing methods for LTRE analysis ignore the fact that persistent temporal variation in matrix elements results in a long-term stochastic growth rate ( a = log λs) that is different from the deterministic growth rate ( r) and thus do not take into account environmental stochasticity. 2. Here, we develop a stochastic extension of LTRE methods that can be used to compare stochastic growth rates among populations that differ in the observed variability of their matrix elements over time. We illustrate our method with actual data and explore a range of questions that may be addressed with these new tools. Specifically, we investigate how variability in weather conditions affected the population dynamics of the short-lived perennial plant species Anthyllis vulneraria and examine how differences in stochastic growth rates ( a) are determined by contributions of mean matrix elements and variability in matrix elements. 3. We find that, consistent with the life history of the species, differences in mean fertility and growth made the largest contribution to differences in a, whereas in terms of variability fertility made the largest contribution in most populations. However, we also find that in all populations, the magnitude of the total contribution of mean matrix elements outweighed that of variability. Finally, increasing soil depth significantly lowered contributions of variability in matrix elements, but it was not related to contributions of differences in mean matrix elements. 4. Synthesis. Stochastic life table response experiment analysis described here provides the first systematic way of incorporating observed differences in temporal variability into the comparison of natural populations. A key finding from this study is that populations occurring on relatively deeper soils were better buffered against climatic variation than populations occurring on shallow soils. We expect this new approach to analyse temporal variability to prove especially useful in the analysis of natural populations experiencing environmental change. [ABSTRACT FROM AUTHOR]

1 The species composition of fragmented semi-natural grasslands may change over time due to stochastic local extinction and colonization events, successional change and/or as a response to changing management or abiotic conditions. The resistance of vegetation to change may be mediated through the effects of topography (slope and aspect) on soils and microclimate. 2 To assess long-term vegetation change in British chalk grasslands, 92 plots first surveyed by F. H. Perring in 1952–53, and distributed across four climatic regions, were re-surveyed during 2001–03. Changes in vegetation since the original survey were assessed by comparing local colonization and extinction rates at the plot scale, and changes in species frequency at the subplot scale. Vegetation change was quantified using indirect ordination (Detrended Correspondence Analysis; DCA) and Ellenberg indicator values. 3 Across all four regions, there was a significant decrease in species number and a marked decline in stress-tolerant species typical of species-rich calcareous grasslands, both in terms of decreased plot occupancy and decreased frequency within occupied plots. More competitive species typical of mesotrophic grasslands had colonized plots they had not previously occupied, but had not increased significantly in frequency within occupied plots. 4 A significant increase in Ellenberg fertility values, which was highly correlated with the first DCA axis, was found across all regions. The magnitude of change of fertility and moisture values was found to decrease with angle of slope and with a topographic solar radiation index derived from slope and aspect. 5 The observed shift from calcareous grassland towards more mesotrophic grassland communities is consistent with the predicted effects of both habitat fragmentation and nutrient enrichment. It is hypothesized that chalk grassland swards on steeply sloping ground are more resistant to invasion by competitive grass species than those on flatter sites due to phosphorus limitation in shallow minerogenic rendzina soils, and that those with a southerly aspect are more resistant due to increased magnitude and frequency of drought events. [ABSTRACT FROM AUTHOR]

The article describes the microclimates of forest and savanna sites in the Olokemeji Forest Reserve in Nigeria. This description is based on rainfall and temperature data from Olokemeji and for other climatic elements, on data from Ibadan. The mean length of dry periods is less than 4 days during the wet season and rises to over 50 days during the dry season. The results show that Olokemeji has a climate typical of it area of the country and of the forest/derived savanna boundary area in west Africa.