1. Carbon storage in vegetation and soil underpins climate regulation through carbon sequestration. Because plant species differ in their ability to capture, store and release carbon, the collective functional characteristics of plant communities (functional diversity) should be a major driver of carbon accumulation in terrestrial ecosystems.
2. Three major components of plant functional diversity could be put forward as drivers of carbon storage in ecosystems: the most abundant functional trait values, the variety of functional trait values and the abundance of particular species that could have additional effects not incorporated in the first two components.
3. We tested for associations between these components and carbon storage across 16 sites in the Chaco forest of Argentina under the same climate and on highly similar parental material. The sites differed in their plant functional diversity caused by different long-term land-use regimes.
4. We measured six plant functional traits in 27 species and weighted them by the species abundance at each site to calculate the community-weighted mean (CWM) and the functional divergence (FDvar) of each single trait and of multiple traits (FDiv). We also measured plant and soil carbon storage. Using a stepwise multiple regression analysis, we assessed which of the functional diversity components best explained carbon storage.
5. Both CWM and FDvar of plant height and wood-specific gravity, but no leaf traits, were retained as predictors of carbon storage in multiple models. Relationships of FDvar of stem traits and FDiv with carbon storage were all negative. The abundance of five species improved the predictive power of some of the carbon storage models.
6. Synthesis. All three major components of plant functional diversity contributed to explain carbon storage. What matters the most to carbon storage in these ecosystems is the relative abundance of plants with tall, and to a lesser extent dense, stems with a narrow range of variation around these values. No consistent link was found between carbon storage and the leaf traits usually associated with plant resource use strategy. The negative association of trait divergence with carbon storage provided no evidence in support to niche complementarity promoting carbon storage in these forest ecosystems.

Based on empirical evidence from the literature we propose that, in nature, phenotypic plasticity in plants is usually expressed at a subindividual level. While reaction norms (i.e. the type and the degree of plant responses to environmental variation) are a property of genotypes, they are expressed at the level of modular subunits in most plants. We thus contend that phenotypic plasticity is not a whole-plant response, but a property of individual meristems, leaves, branches and roots, triggered by local environmental conditions. Communication and behavioural integration of interconnected modules can change the local responses in different ways: it may enhance or diminish local plastic effects, thereby increasing or decreasing the differences between integrated modules exposed to different conditions. Modular integration can also induce qualitatively different responses, which are not expressed if all modules experience the same conditions. We propose that the response of a plant to its environment is the sum of all modular responses to their local conditions plus all interaction effects that are due to integration. The local response rules to environmental variation, and the modular interaction rules may be seen as evolving traits targeted by natural selection. Following this notion, whole-plant reaction norms are an integrative by-product of modular plasticity, which has far-reaching methodological, ecological and evolutionary implications.

Leaf N and P stoichiometry covaries with many aspects of plant biology, yet the drivers of this trait at biogeographic scales remain uncertain. Recently we reported the patterns of leaf C and N based on systematic census of 213 species over 199 research sites in the grassland biomes of China. With the expanded analysis of leaf P, here we report patterns of leaf P and N:P ratios, and analyze the relative contribution of climatic variables and phylogeny in structuring patterns of leaf N:P stoichiometry. Average values of leaf P and N:P ratio were 1.9 mg g(-1) and 15.3 (mass ratio), respectively, consistent with the previous observation of a higher N:P ratio in China's flora than the global averages (ca. 13.8), resulting from a lower leaf P. Climatic variables had very little direct correlation with leaf P and N:P ratios, with growing season precipitation and temperature together explaining less than 2% of the variation, while inter-site differences and within-site phylogenetic variation explained 55 and 26% of the total variation in leaf P and N:P ratios. Across all sites and species, leaf N and P were highly positively correlated at all levels. However, the within-site, within-species covariations of leaf N and P were weaker than those across sites and across species. Leaf N and P relationships are driven by both variation between sites at the landscape scale (explaining 58% of the variance) and within sites at the local scale (explaining 24%), while the climatic factors exerted limited influence (explaining less than 3%). In addition, leaf N:P ratios in two dominant genera Kobresia and Stipa had different responses to precipitation. This study suggests that geographic variation and between-species variation, rather than climatic variation, are the major determinants of grassland foliar stoichiometry at the biome level.

Interannual variability in aboveground net primary production (ANPP) was assessed with long-term (mean = 12 years) data from 11 Long Term Ecological Research sites across North America. The greatest interannual variability in ANPP occurred in grasslands and old fields, with forests the least variable. At a continental scale, ANPP was strongly correlated with annual precipitation. However, interannual variability in ANPP was not related to variability in precipitation. Instead, maximum variability in ANPP occurred in biomes where high potential growth rates of herbaceous vegetation were combined with moderate variability in precipitation. In the most dynamic biomes, ANPP responded more strongly to wet than to dry years. Recognition of the fourfold range in ANPP dynamics across biomes and of the factors that constrain this variability is critical for detecting the biotic impacts of global change phenomena.

In recent years, steppe degradation in North China has become a serious environmental problem. Most research on steppe degradation is conducted at the level of communities or at the scale of small regions. To better understand the spatio-temporal variation and driving factors of grassland degradation, monitoring and analysis at broad regional scales are needed. This paper systematically describes the state and characteristics of steppe degradation at the Xilinhot plateau, makes an in-depth empirical analysis of the natural and man-made causes leading to degradation, and analyzes what driving factors have influenced degradation in this typical steppe region over the last 20 years. Ten biophysical and socio-economic variables, including altitude, slope, precipitation, temperature, soil conditions, distance to river, distance to highway, population density, sheep unit density, and fencing policy, were evaluated on their impact on observed patterns of degradation. The results indicate that all of these factors had a significant influence on the process of steppe degradation. During the first 10 years, from 1991 to 2000, steppe degradation increased, but after 2000, the degradation trend has, to some extent, reversed. The analysis indicates that the measures taken by the government, such as fencing vulnerable areas, played an important role in this change. The results advance the understanding of grassland degradation and contribute to constructing an empirical and theoretical base for grassland management and planning.

Abstract

The semi-private property rights arrangement called the Household Production Responsibility System (HPRS) was started in the early 1980s in Xilingol pasture of Inner Mongolia (China), and stimulated the development of stockbreeding. The grassland has been degrading severely with increasing numbers of livestock. Based on a historical review of property rights regimes in Inner Mongolia and empirical surveys in Xilingol pasture during 2001–2003, this paper assesses the implementation of HPRS and its impacts on incomes of households as well as the environmental impact on the grassland. It was found that HPRS does not mitigate the “Tragedy of the Commons”, instead it has exacerbated the situation. It was also found that co-management of grassland and livestock among a few households presents a sustainable use of grassland to develop livestock breeding. We conclude with the recommendation that small-scale collective property rights systems should be encouraged in Xilingol pasture of Inner Mongolia.

A number of studies show contrasting results in how plant species with specific life-history strategies respond to fragmentation, but a general analysis on whether traits affect plant species occurrences in relation to habitat area and isolation has not been performed. We used published data from forests and grasslands in north-central Europe to analyse if there are general patterns of sensitivity to isolation and dependency of area for species using three traits: life-span, clonality, and seed weight. We show that a larger share of all forest species was affected by habitat isolation and area as compared to grassland species. Persistence-related traits, life-span and clonality, were associated to habitat area and the dispersal and recruitment related trait, seed weight, to isolation in both forest and grassland patches. Occurrence of clonal plant species decreased with habitat area, opposite to non-clonal plant species, and long-lived plant species decreased with grassland area. The directions of these responses partly challenge some earlier views, suggesting that further decrease in habitat area will lead to a change in plant species community composition, towards relatively fewer clonal and long-lived plants with large seeds in small forest patches and fewer clonal plants with small seeds in small grassland patches. It is likely that this altered community has been reached in many fragmented European landscapes consisting of small and isolated natural and semi-natural patches, where many non-clonal and short-lived species have already disappeared. Our study based on a large-scale dataset reveals general and useful insights concerning area and isolation effects on plant species composition that can improve the outcome of conservation and restoration efforts of plant communities in rural landscapes.

Predators determine herbivore and plant biomass via so-called trophic cascades, and the strength of such effects is influenced by ecosystem productivity. To determine whether evolutionary trade-offs among plant traits influence patterns of trophic control, we manipulated predators and soil fertility and measured impacts of a major herbivore (the aphid Aphis nerii) on 16 milkweed species (Asclepias spp.) in a phylogenetic field experiment. Herbivore density was determined by variation in predation and trade-offs between herbivore resistance and plant growth strategy. Neither herbivore density nor predator effects on herbivores predicted the cascading effects of predators on plant biomass. Instead, cascade strength was strongly and positively associated with milkweed response to soil fertility. Accordingly, contemporary patterns of trophic control are driven by evolutionary convergent trade-offs faced by plants.

Theoretical and laboratory research suggests that phenotypic plasticity can evolve under selection. However, evidence for its evolutionary potential from the wild is lacking. We present evidence from a Dutch population of great tits (Parus major) for variation in individual plasticity in the timing of reproduction, and we show that this variation is heritable. Selection favoring highly plastic individuals has intensified over a 32-year period. This temporal trend is concurrent with climate change causing a mismatch between the breeding times of the birds and their caterpillar prey. Continued selection on plasticity can act to alleviate this mismatch.

Temperature data over the past five decades show faster warming of the global land surface during the night than during the day. This asymmetric warming is expected to affect carbon assimilation and consumption in plants, because photosynthesis in most plants occurs during daytime and is more sensitive to the maximum daily temperature, Tmax, whereas plant respiration occurs throughout the day and is therefore influenced by both Tmax and the minimum daily temperature, Tmin. Most studies of the response of terrestrial ecosystems to climate warming, however, ignore this asymmetric forcing effect on vegetation growth and carbon dioxide (CO2) fluxes. Here we analyse the interannual covariations of the satellite-derived normalized difference vegetation index (NDVI, an indicator of vegetation greenness) with Tmax and Tmin over the Northern Hemisphere. After removing the correlation between Tmax and Tmin, we find that the partial correlation between Tmax and NDVI is positive in most wet and cool ecosystems over boreal regions, but negative in dry temperate regions. In contrast, the partial correlation between Tmin and NDVI is negative in boreal regions, and exhibits a more complex behaviour in dry temperate regions. We detect similar patterns in terrestrial net CO2 exchange maps obtained from a global atmospheric inversion model. Additional analysis of the long-term atmospheric CO2 concentration record of the station Point Barrow in Alaska suggests that the peak-to-peak amplitude of CO2 increased by 23 +/- 11% for a +1 degrees C anomaly in Tmax from May to September over lands north of 51 degrees N, but decreased by 28 +/- 14% for a +1 degrees C anomaly in Tmin. These lines of evidence suggest that asymmetric diurnal warming, a process that is currently not taken into account in many global carbon cycle models, leads to a divergent response of Northern Hemisphere vegetation growth and carbon sequestration to rising temperatures.

We quantified the biomass allocation patterns to leaves, stems and roots in vegetative plants, and how this is influenced by the growth environment, plant size, evolutionary history and competition. Dose-response curves of allocation were constructed by means of a meta-analysis from a wide array of experimental data. They show that the fraction of whole-plant mass represented by leaves (LMF) increases most strongly with nutrients and decreases most strongly with light. Correction for size-induced allocation patterns diminishes the LMF-response to light, but makes the effect of temperature on LMF more apparent. There is a clear phylogenetic effect on allocation, as eudicots invest relatively more than monocots in leaves, as do gymnosperms compared with woody angiosperms. Plants grown at high densities show a clear increase in the stem fraction. However, in most comparisons across species groups or environmental factors, the variation in LMF is smaller than the variation in one of the other components of the growth analysis equation: the leaf area : leaf mass ratio (SLA). In competitive situations, the stem mass fraction increases to a smaller extent than the specific stem length (stem length : stem mass). Thus, we conclude that plants generally are less able to adjust allocation than to alter organ morphology.

Invasion biologists often suggest that phenotypic plasticity plays an important role in successful plant invasions. Assuming that plasticity enhances ecological niche breadth and therefore confers a fitness advantage, recent studies have posed two main hypotheses: (1) invasive species are more plastic than non-invasive or native ones; (2) populations in the introduced range of an invasive species have evolved greater plasticity than populations in the native range. These two hypotheses largely reflect the disparate interests of ecologists and evolutionary biologists. Because these sciences are typically interested in different temporal and spatial scales, we describe what is required to assess phenotypic plasticity at different levels. We explore the inevitable tradeoffs of experiments conducted at the genotype vs. species level, outline components of experimental design required to identify plasticity at different levels, and review some examples from the recent literature. Moreover, we suggest that a successful invader may benefit from plasticity as either (1) a Jack-of-all-trades, better able to maintain fitness in unfavourable environments; (2) a Master-of-some, better able to increase fitness in favourable environments; or (3) a Jack-and-master that combines some level of both abilities. This new framework can be applied when testing both ecological or evolutionary oriented hypotheses, and therefore promises to bridge the gap between the two perspectives.

Abstract

Grazing by large herbivores, in interplay with environmental productivity, is a key driver of the composition of the vegetation with important consequences on the ecosystem and, consequently, for land management. We tested the predictions of the resource availability – resource–acquisition theory by assessing the extent to which community averages of plant traits, known to be related to plant growth, competitive ability and response to grazing were correlated with resource gradients within local (200 km²) geographical ranges. Second, we assessed the applicability of the same set of plant traits to make inferences on ecological effects of grazing by sheep in alpine ecosystems in Norway, using a data set consisting of 16 sites in central Norway. We estimated grazing intensity by free-ranging sheep based on GPS telemetry, soil properties, plant species composition and species traits i.e. specific leaf area (SLA), leaf dry matter content (LDMC), leaf size and plant height. Soil fertility and the interaction between soil fertility and grazing, but not grazing intensity alone, were significantly related to plant species and traits composition. Generally, average SLA showed lower correspondence with soil fertility and grazing than the other traits. Leaf size and plant height were lowest at sites with high grazing intensity and in sites with low fertility, and increased with soil fertility in little and moderately grazed sites, but declined at high fertility sites when grazing was intense. LDMC showed the opposite trend. Grazing intensity was more related to the variability in plant composition and average plant traits when environmental productivity was high. Our results therefore are indicative of a convergence of responses to grazing and nutrient limitation.

Zusammenfassung

Die Beweidung durch große Herbivore stellt im Zusammenspiel mit der Produktivität der Umwelt einen Schlüsselfaktor für die Zusammensetzung der Vegetation dar und hat damit große Auswirkungen auf das Ökosystem und dementsprechend für die Landpflege. Wir untersuchten die Vorhersagen der Ressourcenverfügbarkeits-Ressourcenerwerbs-Theorie indem wir abschätzten, in welchem Ausmaß die durchschnittlichen Pflanzeneigenschaften einer Gesellschaft, die erwiesenermaßen mit dem Pflanzenwachstum, der Konkurrenzfähigkeit und der Reaktion auf Beweidung zusammenhängen, mit den Ressourcengradienten innerhalb von lokalen geografischen Bereichen (200 km²) korreliert sind. Wir bewerteten zweitens die Anwendbarkeit der gleichen Auswahl von Pflanzeneigenschaften für die Abschätzung von ökologischen Effekten der Schafsbeweidung auf alpine Ökosysteme in Norwegen, indem wir einen Datensatz verwendeten, der aus 16 Probeflächen in Zentralnorwegen bestand. Wir erfassten die Beweidungsintensität von freilaufenden Schafen mithilfe der GPS-Telemetrie, die Bodenfaktoren, die Zusammensetzung der Pflanzenarten und die Arteigenschaften, d. h. die spezifische Blattfläche (SLA), die Blatttrockenmasse (LDMC), die Blattgröße und Pflanzenhöhe. Die Bodenfruchtbarkeit und die Interaktion zwischen der Bodenfruchtbarkeit und der Beweidung, jedoch nicht die Beweidungsintensität an sich, waren mit den Pflanzenarten und der Zusammensetzung der Eigenschaften signifikant korreliert. Im Allgemeinen zeigte die durchschnittliche SLA einen geringeren Zusammenhang mit der Bodenfruchtbarkeit und der Beweidung als die anderen Eigenschaften. Die Blattfläche und die Pflanzenhöhe waren auf Flächen mit starker Beweidung und auf Flächen mit geringer Fruchtbarkeit, am geringsten, nahmen aber mit der Bodenfruchtbarkeit in gering und mittelstark beweideten Flächen zu und nahmen in fruchtbaren Flächen bei starker Beweidung ab. LDMC zeigte den gegenläufigen Trend. Die Beweidungsintensität war stärker mit der Variabilität in der Pflanzenzusammensetzung und den durchschnittlichen Pflanzeneigenschaften verbunden, wenn die Produktivität der Umwelt hoch war. Unsere Ergebnisse weisen daher auf eine Konvergenz der Reaktionen auf Beweidung und Nährstofflimitierung hin.

【Objective】The distribution characteristics of soil organic carbon (SOC) storage were studied for the purpose to determine the reasonable grazing management way and provide a basis for the regional carbon estimation in grassland ecosystem. 【Method】The SOC storage of 0-100 cm soil depth was measured by using field method of stratified sampling in the typical steppe.【Result】The SOC storage was between 9.72-14.84 kg•m-2 in the three study plots. No significant difference was found in SOC storage in three study plots of spatial distance. The significant difference of SOC storage was found between four different grazing managements treatments in study area, and the order was moderate grazing (MG)＞light grazing (LG)＞heavy grazing (HG)＞control area (CK). The SOC storage decreased with the increasing of soil depth, and a significant difference was detected between SOC among different soil layers. A positive significant relationship was found between SOC storage and soil layer (P＜0.01), and the relationship could be described by using logarithmic and linear equations.【Conclusion】The SOC storage was relatively stable in the typical steppe, and presented a vertical descending characteristics. The effect of different grazing managements on SOC storage in the same vegetation type was significantly higher than the effect of spatial distance. Moderate grazing is benefit for carbon fixation of grassland ecosystem.

【Objective】The distribution characteristics of soil organic carbon (SOC) storage were studied for the purpose to determine the reasonable grazing management way and provide a basis for the regional carbon estimation in grassland ecosystem. 【Method】The SOC storage of 0-100 cm soil depth was measured by using field method of stratified sampling in the typical steppe.【Result】The SOC storage was between 9.72-14.84 kg•m-2 in the three study plots. No significant difference was found in SOC storage in three study plots of spatial distance. The significant difference of SOC storage was found between four different grazing managements treatments in study area, and the order was moderate grazing (MG)＞light grazing (LG)＞heavy grazing (HG)＞control area (CK). The SOC storage decreased with the increasing of soil depth, and a significant difference was detected between SOC among different soil layers. A positive significant relationship was found between SOC storage and soil layer (P＜0.01), and the relationship could be described by using logarithmic and linear equations.【Conclusion】The SOC storage was relatively stable in the typical steppe, and presented a vertical descending characteristics. The effect of different grazing managements on SOC storage in the same vegetation type was significantly higher than the effect of spatial distance. Moderate grazing is benefit for carbon fixation of grassland ecosystem.

The use of livestock to manage vegetation composition has become a common element of conservation planning in many regions of the world. Similar to many arid and semi-arid grasslands throughout the world, California grasslands have a history of invasion by non-native grasses and forbs. Attempts to restore native plant populations using managed grazing are common, despite the lack of an overarching quantitative basis for assessing livestock effects on different plant groups. Given the wide range of soils, climate and topography over which grasslands are found, it is important to understand the context-dependency of grazing effects across a region. We performed a meta-analysis of livestock grazing within California grasslands to investigate the response of different plant groups to grazing relative to precipitation, grassland type, soil and grazing regime. We found that exotic forbs showed a dramatic, uniform increase in cover with grazing, but no increase in richness. By contrast, native forbs increased in richness yet their cover response was weak and variable depending on grazing regime and precipitation. Exotic grass cover was unaffected while richness was enhanced by grazing. Native grass cover generally increased with grazing, although the high variation among studies was not predicted by the explanatory variables we evaluated. These results lend support to the use of grazing to enhance native forb richness and native grass cover in some settings although this must be weighed against increases in the cover of exotic forbs. (C) 2012 Elsevier Ltd.

Plants can adapt to grazing environments by developing defensive traits, such as spines and toxins, or having a small phenotype, such as short and prostrate growth forms. This study examined facilitative and competitive interactions between species with different types of grazing adaptation. We predicted that large species with defensive traits sometimes protect grazing-adapted species without defensive traits from herbivores, but competitively suppress them overall. We conducted an experiment using fences and removals of an unpalatable plant in the long-term deer grazing habitat of Nara Park in Nara, Japan. We evaluated the seasonal variations in the facilitative and competitive effects of a defensive perennial, Urtica thunbergiana, on the growth, survival, reproduction, and final fitness of a small palatable annual species, Persicaria longiseta, during a growing season. The populations of the two species in the park have adapted to the grazed habitat by increasing the density of stinging hairs (Urtica) and developing inherently short shoots and small leaves (Persicaria). We found that Urtica individuals had facilitative effects on the growth of Persicaria individuals under grazing during a few periods of the growing season, but had neutral effects on survival and plant fitness throughout the season. In the fenced plots, Urtica had negative effects on the growth, survival, and reproduction of Persicaria. These results suggest that the relative importance of the facilitative and competitive effects of Urtica on Persicaria fluctuated due to seasonal variations in grazing pressure and vegetative productivity. Although well-defended plants often facilitate less-defended species, we conclude that the facilitative effects of Urtica on Persicaria are limited in a plant community with a long history of intensive grazing.

A major aim in ecology is identifying determinants of invasiveness. We performed a meta-analysis of 117 field or experimental-garden studies that measured pair-wise trait differences of a total of 125 invasive and 196 non-invasive plant species in the invasive range of the invasive species. We tested whether invasiveness is associated with performance-related traits (physiology, leaf-area allocation, shoot allocation, growth rate, size and fitness), and whether such associations depend on type of study and on biogeographical or biological factors. Overall, invasive species had significantly higher values than non-invasive species for all six trait categories. More trait differences were significant for invasive vs. native comparisons than for invasive vs. non-invasive alien comparisons. Moreover, for comparisons between invasive species and native species that themselves are invasive elsewhere, no trait differences were significant. Differences in physiology and growth rate were larger in tropical regions than in temperate regions. Trait differences did not depend on whether the invasive alien species originates from Europe, nor did they depend on the test environment. We conclude that invasive alien species had higher values for those traits related to performance than non-invasive species. This suggests that it might become possible to predict future plant invasions from species traits.

The non-equilibrium concept of rangeland dynamics predicts that the potential for grazing-induced degradation is low in rangelands with relatively variable precipitation. To date, evidence in support of the non-equilibrium concept has been inconsistent. Using a standardized protocol, including a newly developed global map of rainfall variability, we reviewed the incidence of degradation in relation to rainfall variability across 58 published studies. We distinguished between (1) zonal degradation (i.e., degradation independent of water and key resources), (2) degradation in the presence of key resources, and (3) degradation in the presence of water. For studies not affected by proximity to permanent water or key resources, we found strong support for the non-equilibrium concept for rangelands. Zonal degradation was absent at CV (coefficient of variation) values above 33%, which has been proposed as a critical threshold. Grazing degradation was almost entirely restricted to areas with relatively stable annual precipitation as expressed by a low CV, or to rangelands with key resources or water points nearby. To better understand rangeland dynamics, we recommend that future studies use globally comparable measures of degradation and rainfall variability. Our work underlines that rangelands with relatively stable rainfall patterns, and those with access to water or key resources, are potentially vulnerable to degradation. Grazing management in such areas should incorporate strategic rest periods. Such rest periods effectively mimic natural fluctuations in herbivore populations, which are a defining characteristic of non-degraded rangelands occurring under highly variable precipitation regimes.