Interannual climate variability mediates changes in carbon and nitrogen pools caused by annual grass invasion in a semiarid shrubland

Exotic plant invasions alter ecosystem properties and threaten ecosystem functions globally. Interannual climate variability (ICV) influences both plant community composition (PCC) and soil properties, and interactions between ICV and PCC may influence nitrogen (N) and carbon (C) pools. We asked how ICV and non-native annual grass invasion covary to influence soil and plant N and C in a semiarid shrubland undergoing widespread ecosystem transformation due to invasions and altered fire regimes. We sampled four progressive stages of annual grass invasion at 20 sites across a large (25,000 km²) landscape for plant community composition, plant tissue N and C, and soil total N and C in 2013 and 2016, which followed 2 years of dry and wet conditions, respectively. Multivariate analyses and ANOVAs showed that in invasion stages where native shrub and perennial grass and forb communities were replaced by annual grass-dominated communities, the ecosystem lost more soil N and C in wet years. Path analysis showed that high water availability led to higher herbaceous cover in all invasion stages. In stages with native shrubs and perennial grasses, higher perennial grass cover was associated with increased soil C and N, while in annual-dominated stages, higher annual grass cover was associated with losses of soil C and N. Also, soil total C and C:N ratios were more homogeneous in annual-dominated invasion stages as indicated by within-site standard deviations. Loss of native shrubs and perennial grasses and forbs coupled with annual grass invasion may lead to long-term declines in soil N and C and hamper restoration efforts. Restoration strategies that use innovative techniques and novel species to address increasing temperatures and ICV and emphasize maintaining plant community structure—shrubs, grasses, and forbs—will allow sagebrush ecosystems to maintain C sequestration, soil fertility, and soil heterogeneity.     

内蒙古锡林河流域草原群落植物功能群组成沿水热梯度变化的样带研究

 用样带法调查了草原群落植物功能群组成沿水热梯度的变化。结果表明,随着海拔高度的降低,降水量的减少,热量和干燥度的增加,群落植物生活型功能群组成中,多年生杂类草的相对多度逐渐减少,而多年生丛生禾草的相对多度逐渐增加,多年生丛生禾草对多年生杂类草具有强烈的生态替代作用。在植物生态类群功能群组成中,中旱生植物的作用逐渐减弱,而旱生植物的作用显著增强。草地的利用方式对群落组成具有重要的影响,永久样地和割草场中不耐牧种的相对生物量显著地高于放牧场,而放牧场中耐牧种的相对生物量则显著地高于永久样地和割草场。受草地利用     

Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap

Preventing invasion by exotic species is one of the key goals of restoration, and community assembly theory provides testable predictions about native community attributes that will best resist invasion. For instance, resource availability and biotic interactions may represent “filters” that limit the success of potential invaders. Communities are predicted to resist invasion when they contain native species that are functionally similar to potential invaders; where phenology may be a key functional trait. Nutrient reduction is another common strategy for reducing invasion following native species restoration, because soil nitrogen (N) enrichment often facilitates invasion. Here, we focus on restoring the herbaceous community associated with coastal sage scrub vegetation in Southern California; these communities are often highly invaded, especially by exotic annual grasses that are notoriously challenging for restoration. We created experimental plant communities composed of the same 20 native species, but manipulated functional group abundance (according to growth form, phenology, and N‐fixation capacity) and soil N availability. We fertilized to increase N, and added carbon to reduce N via microbial N immobilization. We found that N reduction decreased exotic cover, and the most successful seed mix for reducing exotic abundance varied depending on the invader functional type. For instance, exotic annual grasses were least abundant when the native community was dominated by early active forbs, which matched the phenology of the exotic annual grasses. Our findings show that nutrient availability and the timing of biotic interactions are key filters that can be manipulated in restoration to prevent invasion and maximize native species recovery.     

The Role of Light and Soil Moisture in Plant Community Resistance to Invasion by Yellow Starthistle (Centaurea solstitialis)

To resist establishment by an invasive plant, a community may require one or more species functionally similar to the invader in their resource acquisition pattern. In this study, communities consisting of native winter annual forbs, non‐native annual grasses, native perennials, or a combination of the two native communities were established with and without Centaurea solstitialis to determine the effect of soil moisture and light availability on plant community invasion resistance. The annual plant communities were unable to resist invasion by C. solstitialis. In the native winter annual forb community, senescence in late spring increased light penetration (>75%) to the soil surface, allowing seeded C. solstitialis to quickly establish and dominate the plots. In addition, native annual forbs utilized only shallow soil moisture, whereas C. solstitialis used shallow and deep soil moisture. In communities containing native perennials, only Elymus glaucus established well and eventually dominated the plots. During the first 2 years of establishment, water use pattern of perennial communities was similar to native annual forbs and resistance to invasion was associated with reduced light availability during the critical stages of C. solstitialis establishment. In later years, however, water use pattern of perennial grass communities was similar or greater than C. solstitialis‐dominated plots. These results show that Central Valley grasslands that include E. glaucus resist C. solstitialis invasion by a combination of light suppression and soil water competition. Spatiotemporal resource utilization patterns, and not just functional similarity, should be considered when developing restoration strategies to resist invasion by many non‐native species.     

Grass invasion causes rapid increases in ecosystem carbon and nitrogen storage in a semiarid shrubland

Accurately predicting terrestrial carbon (C) and nitrogen (N) storage requires understanding how plant invasions alter cycling and storage. A common, highly successful type of plant invasion occurs when the invasive species is of a distinctly different functional type than the native dominant plant, such as shrub encroachment throughout the western United States and annual grass invasions in Mediterranean shrublands, as studied here. Such invasions can dramatically transform landscapes and have large potential to alter C and N cycling by influencing storage in multiple pools. We used a manipulation of non‐native annual grass litter within a shrub‐dominated habitat in southern California (coastal sage scrub, CSS) to study how grass invasion alters ecosystem C and N storage. We added, removed, or left unchanged grass litter in areas of high and low invasion, then followed soil and vegetation changes. Grass litter greatly increased C and N storage in soil, aboveground native and non‐native biomass. Aboveground litter storage increased due to the greater inputs and slower decomposition of grass litter relative to shrub litter; shading by grass litter further reduced decomposition of both non‐native and native litter, which may be due to reduced photodegradation. Soil C and N pools in areas of high litter increased ～20% relative to low litter areas in the two years following manipulation and were generally sinks for C and N, while areas with low litter were sources. We synthesize our results into a C cycle of invaded and uninvaded areas of CSS and link changes in storage to increases in the soil fungi : bacteria ratio, increased plant inputs, and decreased litter loss. Overall, we show that grasses, especially through their litter, control important abiotic and biotic mechanisms governing C and N storage, with widespread implications for C sequestration and N storage in semiarid systems undergoing grass or shrub invasions.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Repeated mowing to restore remnant native grasslands invaded by nonnative annual grasses: upsides and downsides above and below ground

California grasslands have been severely impacted by the invasion of nonnative annual grasses, which often limit restoration of this important ecosystem. In this study, we explored the use of mowing as a restoration tool for native perennial grasslands at the Santa Rosa Plateau Ecological Reserve in southern California. We sought to evaluate if, over time, mowing would reduce nonnative annual grass cover and benefit native species, especially the native bunchgrass Stipa pulchra. We hypothesized that repeated mowing, carefully timed to target nonnative annual grasses prior to seed maturation, would reduce nonnative seed inputs into the soil and eventually lead to diminished abundance of these species. We monitored vegetation in mowed and unmowed plots for 4 years, and conducted a seed bank study after 5 years to better understand the cumulative effects of mowing on native and nonnative seed inputs. Consistent with our hypotheses, we found that mowing successfully reduced nonnative annual grass cover and benefitted some native species, including S. pulchra. However, we also found that nonnative forb species showed progressive increases in mowed plots over time. We observed similar patterns of species composition in the soil seed bank. Together, these results suggest that mowing can be used to control nonnative annual grasses and increase the abundance of native bunchgrasses, but that this method may also have the unintended consequence of increasing nonnative forb species.     

Fungal community assembly in soils and roots under plant invasion and nitrogen deposition

Fungal community composition in the Anthropocene is driven by rapid changes in environmental conditions caused by human activities. This study examines the relative importance of two global change drivers – atmospheric nitrogen (N) deposition and annual grass invasion – on structuring fungal communities in a California chaparral ecosystem, with emphasis on arbuscular mycorrhizal fungi. We used molecular markers, functional groupings, generalized linear statistics and joint distribution modeling, to examine how environmental variables structure taxonomic and functional composition of fungal communities. Invasive grasses had a lower richness and relative abundance of symbiotic fungi (both AMF and other fungi) compared to native shrubs. We found a higher richness and abundance of rhizophilic (e.g. Glomeraceae) and edaphophilic (e.g. Gigasporaceae) AMF with increasing soil NO₃. Our findings suggest that invasive persistence may decrease the presence of multiple soil symbionts that native species depend on for pathogen protection and increased access to soil resources.     

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.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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Long‐term experiment manipulating community assembly results in favorable restoration outcomes for invaded prairies

Invasive species are a common problem in restoration projects. Manipulating soil fertility and species arrival order has the potential to lower their abundance and achieve higher abundances of seeded native species. In a 7‐year experiment in Missouri, United States, we tested how nutrient addition and the timing of arrival of the invasive legume Lespedeza cuneata and seeded native prairie grass and forb species influenced overall community composition. Treatments that involved early arrival of seeded forb and grass species and late arrival of L. cuneata were most successful at creating community structure that fulfilled our restoration goals, displaying high abundance of seeded native forb species, low abundances of L. cuneata, and non‐native species. There were few treatment interactions, with the exception that timing seeded native forbs and timing of L. cuneata arrival interactively influenced the abundance of seeded native forbs. This suggests that the individual treatments are supporting the restoration goals, such as creating a community with low abundance of L. cuneate or high abundance of native seeded species, without restricting each other. This study demonstrates the importance of priority effects in disturbed habitats prone to invasion, the lasting effects of initial seeding on long‐term community composition, and the potential for fertilization to positively benefit restoration of degraded grasslands.     

Disentangling direct and indirect effects of a legume shrub on its understorey community

Direct and indirect interactions among plants contribute to shape community composition through above‐ and belowground processes. However, we have not disentangled yet the direct and indirect soil and canopy effects of dominants on understorey species. We addressed this issue in a semi‐arid system from southeast Spain dominated by the legume shrub Retama sphaerocarpa. During a year with an exceptionally dry spring, we removed the shrub canopy to quantify aboveground effects and compared removed‐canopy plots to open plots between shrubs to quantify soil effects, both with and without watering. We added a grass removal treatment in order to separate direct from indirect shrub effects and quantified biomass, abundance, richness and composition of the forb functional group. With watering, changes in forb biomass were primarily driven by indirect shrub effects, with contrasting negative soil and positive aboveground indirect effects; changes in forb abundance and composition were more influenced by direct shrub soil effects with contrasting species composition between open and Retama patches. As community composition was different between open and Retama patches the indirect effects of Retama on forb species did not concern forbs from the open community but forbs from Retama patches. Indirect effects are, thus, important at the functional group level rather than at the species level. Without watering, there were no significant interactions. Changes in species richness between treatments were weak and seldom significant. We conclude that shrub effects on understorey forbs are primarily due to their influence on soil properties, directly affecting forb species composition but indirectly affecting the biomass of the forbs of the Retama patches, and only with sufficient water.     

Original Articles: Nitrogen Availability and Old-Field Succession in a Shortgrass Steppe

The relationship between soil nitrogen (N) availability and plant community structure was investigated in old-fields in the shortgrass steppe of Colorado. Nitrogen availability was manipulated by N or sucrose additions for 4 years at three old-fields (early-seral, mid-seral, and late-seral) and at an uncultivated control site. The addition of N generally resulted in increased abundance of annual forbs and grasses relative to perennials at all of the previously cultivated sites. Conversely, experimental reduction of N availability generally increased the relative abundance of perennials. Despite a lack of detectable differences in N mineralization between sites and treatments, ion-exchange resin bags confirmed that sucrose additions reduced plant-available N and that N additions increased plant-available N. This was evidenced further by similar observations for plant tissue N content. The degree to which N additions increased N availability at the various sites supported the idea that late-seral plant communities are less effective at N capture relative to earlier-seral communities. The mid-seral old-field had the lowest rates of litter decomposition and a relatively large accumulation of litter on the soil surface. This mid-seral old-field was dominated by an exotic annual grass (Bromus tectorum), which appears to be a major hindrance to redevelopment of the plant-soil system. By experimentally reducing N availability at this stage, we were able, in 4 years, to change the plant community into one that more closely resembled the late-seral community. We also observed that the natural recruitment of weedy annual species on the uncultivated site during an unusually wet year was suppressed by reducing N availability. Our results suggest that available N is an important factor controlling the rate and course of plant and soil community redevelopment on abandoned croplands in the shortgrass steppe, and that manipulation of N availability might be useful in restoration of rangeland vegetation. Received 19 May 1998; accepted 27 August 1999.     

Combined effects of precipitation and nitrogen deposition on native and invasive winter annual production in California deserts

Primary production in deserts is limited by soil moisture and N availability, and thus is likely to be influenced by both anthropogenic N deposition and precipitation regimes altered as a consequence of climate change. Invasive annual grasses are particularly responsive to increases in N and water availabilities, which may result in competition with native forb communities. Additionally, conditions favoring increased invasive grass production in arid and semi-arid regions can increase fire risk, negatively impacting woody vegetation that is not adapted to fire. We conducted a seeded garden experiment and a 5-year field fertilization experiment to investigate how winter annual production is altered by increasing N supply under a range of water availabilities. The greatest production of invasive grasses and native forbs in the garden experiment occurred under the highest soil N (inorganic N after fertilization = 2.99 g m^?2) and highest watering regime, indicating these species are limited by both water and N. A classification and regression tree (CART) analysis on the multi-year field fertilization study showed that winter annual biomass was primarily limited by November–December precipitation. Biomass exceeded the threshold capable of carrying fire when inorganic soil N availability was at least 3.2 g m^?2 in piñon-juniper woodland. Due to water limitation in creosote bush scrub, biomass exceeded the fire threshold only under very wet conditions regardless of soil N status. The CART analyses also revealed that percent cover of invasive grasses and native forbs is primarily dependent on the timing and amount of precipitation and secondarily dependent on soil N and site-specific characteristics. In total, our results indicate that areas of high N deposition will be susceptible to grass invasion, particularly in wet years, potentially reducing native species cover and increasing the risk of fire.     

Early-successional plants regulate grassland productivity and species composition: a removal experiment

Grime's (1998) "mass-ratio" hypothesis holds that ecosystem processes depend in the short term on functional properties of dominant plants and in the longer term on how resident species influence the recruitment of dominants. The latter of these effects may be especially important among early-successional species in disturbed ecosystems, but experimental tests are few. We removed two groups of early-successional species, an annual forb Gutierrezia dracunculoides (DC.) S. F. Blake and annual species (mostly grasses) that complete growth early in the growing season [early-season (ES) species], from a heavily-grazed grassland in central Texas, USA dominated by a C₄ perennial grass. We sought to determine effects of annuals on grassland functioning [productivity, water balance, soil and plant nitrogen (N)] and composition. Removals did not impact N retention in the soil/plant system during the two years of this study, but removing ES annuals increased the amount of water between 30 and 120 cm in the soil profile early in each growing season. Production and N accumulation by vegetation declined following the removal of ES annuals in approximate proportion to the contribution of annuals to aboveground biomass and N, consistent with the mass-ratio hypothesis. By the second year, production and N uptake by initially sub-dominant species increased to fully compensate for the loss of annuals. These results are consistent with the view that ecosystem functions are more strongly linked to species attributes than to diversity per se. Longer-term effects of annuals on grassland composition were evident in a dramatic increase in biomass of perennial forbs after annuals were removed. Because perennial forbs differ from the dominant grass in this grassland in traits that influence ecosystem functioning, ES annuals may affect grassland functioning more by regulating the composition of vegetation than by directly affecting process rates.     

Mechanisms of resistance of Mediterranean annual communities to invasion by Conyza bonariensis: effects of native functional composition

Recent studies have shown that a high species or functional group richness may not always lead to a greater resistance of plant communities to invasion, whereas species and/or functional group composition can more reliably predict invasion resistance. The aim of this study was to understand the mechanisms through which functional group composition can influence the resistance of Mediterranean annual communities to invasion by the exotic Conyza bonariensis . To analyse the effects of functional composition on the performance of individuals introduced as seedlings we first examined the relationships between the demographic and vegetative parameters of C. bonariensis and the biomass achieved by each functional group (grasses, legumes and Asteraceae rosettes) in synthetic communities. As a further step to approach the mechanisms involved in community resistance to invasion, we included in the analyses measurements of functional variables taken within the synthetic communities.
In agreement with earlier results and theory suggesting that high nutrient availability can favour invasions, an abundant legume biomass in communities increased the final biomass and net fecundity of C. bonariensis , due to positive effects on soil nitrate concentration. Survival and establishment of C. bonariensis were mainly favoured by a high biomass of Asteraceae. Additional results from measurements of herbivory suggested that C. bonariensis survival wasn't related to abiotic conditions but may be owed to a protection against herbivores in plots with abundant Asteraceae . Establishment was on the other hand likely to be hindered by the effects of abundant grass and legume foliage on light quality, and therefore easier within an Asteraceae canopy.
We conclude that invasion of Mediterranean old fields by species with biologies similar to C. bonariensis could be limited by favouring communities dominated by annual grasses.     

Plant phenology and life span influence soil pool dynamics: Bromus tectorum invasion of perennial C3–C4 grass communities

In water-limited ecosystems, small rainfall events can have dramatic impacts on microbial activity and soil nutrient pools. Plant community phenology and life span also affect soil resources by determining the timing and quantity of plant nutrient uptake, storage, and release. Using the replacement of C₃–C₄ perennial grasses by the invasive annual grass Bromus tectorum as a case study, we investigated the influence of phenology and life span on pulse responses and sizes of soil carbon (C) and nitrogen (N) pools. We hypothesized that available and microbial C and N would respond to small rainfall events and that B. tectorum invasion would increase soil C and N pools by reducing inter-annual plant C and N storage and alter seasonal pool dynamics by changing the timing of plant uptake and litter inputs. We tested our hypotheses by simulating small rainfall events in B. tectorum and perennial grass communities three times during the growing season. Microbial pools responded strongly to soil moisture and simulated rainfall events, but labile C and N pools were affected weakly or not at all. All pools were larger beneath B. tectorum than perennial grasses. Soil C and N pools increased after senescence in both communities. Our results suggest that transforming a perennial into a B. tectorum dominated community increases the overall size of soil C and N pools by decreasing plant C and N storage and changes seasonal pool dynamics by altering dominant plant phenology. Our results indicate strong roles for water, life span and phenology in controlling soil C and N pools and begin to elucidate the biogeochemical effects of altering plant community phenology and life span.     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Dominant Functional Group Effects on the Invasion Resistance at Different Resource Levels

Background

Functional group composition may affect invasion in two ways the effect of abundance, i.e. dominance of functional group; and the effect of traits, i.e. identity of functional groups. However, few studies have focused on the role of abundance of functional group on invasion resistance. Moreover, how resource availability influences the role of the dominant functional group in invasion resistance is even less understood.

Methodology/Principal Findings

In this experiment, we established experimental pots using four different functional groups (annual grass, perennial grass, deciduous shrub or arbor and evergreen shrub or arbor), and the dominant functional group was manipulated. These experimental pots were respectively constructed at different soil nitrogen levels (control and fertilized). After one year of growth, we added seeds of 20 different species (five species per functional group) to the experimental pots. Fertilization significantly increased the overall invasion success, while dominant functional group had little effect on overall invasion success. When invaders were grouped into functional groups, invaders generally had lower success in pots dominated by the same functional group in the control pots. However, individual invaders of the same functional group exhibited different invasion patterns. Fertilization generally increased success of invaders in pots dominated by the same than by another functional group. However, fertilization led to great differences for individual invaders.

Conclusions/Significance

The results showed that the dominant functional group, independent of functional group identity, had a significant effect on the composition of invaders. We suggest that the limiting similarity hypothesis may be applicable at the functional group level, and limiting similarity may have a limited role for individual invaders as shown by the inconsistent effects of dominant functional group and fertilization.     

Grazing-induced changes in plant composition affect litter quality and nutrient cycling in flooding Pampa grasslands

Changes in plant community composition induced by vertebrate grazers have been found to either accelerate or slow C and nutrient cycling in soil. This variation may reflect the differential effects of grazing-promoted (G+) plant species on overall litter quality and decomposition processes. Further, site conditions associated with prior grazing history are expected to influence litter decay and nutrient turnover. We studied how grazing-induced changes in plant life forms and species identity modified the quality of litter inputs to soil, decomposition rate and nutrient release in a flooding Pampa grassland, Argentina. Litter from G+ forbs and grasses (two species each) and grazing-reduced (G−) grasses (two species) was incubated in long-term grazed and ungrazed sites. G+ species, overall, showed higher rates of decomposition and N and P release from litter. However, this pattern was primarily driven by the low-growing, high litter-quality forbs included among G+ species. Forbs decomposed and released nutrients faster than either G+ or G− grasses. While no consistent differences between G+ and G− grasses were observed, patterns of grass litter decay and nutrient release corresponded with interspecific differences in phenology and photosynthetic pathway. Litter decomposition, N release and soil N availability were higher in the grazed site, irrespective of species litter type. Our results contradict the notion that grazing, by reducing more palatable species and promoting less palatable ones, should decrease nutrient cycling from litter. Plant tissue quality and palatability may not unequivocally link patterns of grazing resistance and litter decomposability within a community, especially where grazing causes major shifts in life form composition. Thus, plant functional groups defined by species’ “responses” to grazing may only partially overlap with functional groups based on species “effects” on C and nutrient cycling.