Plant community composition steers grassland vegetation via soil legacy effects

Soil legacy effects are commonly highlighted as drivers of plant community dynamics and species co‐existence. However, experimental evidence for soil legacy effects of conditioning plant communities on responding plant communities under natural conditions is lacking. We conditioned 192 grassland plots using six different plant communities with different ratios of grasses and forbs and for different durations. Soil microbial legacies were evident for soil fungi, but not for soil bacteria, while soil abiotic parameters did not significantly change in response to conditioning. The soil legacies affected the composition of the succeeding vegetation. Plant communities with different ratios of grasses and forbs left soil legacies that negatively affected succeeding plants of the same functional type. We conclude that fungal‐mediated soil legacy effects play a significant role in vegetation assembly of natural plant communities.     

Maintenance of leaf N controls the photosynthetic CO2 response of grassland species exposed to 9 years of free‐air CO2 enrichment

Determining underlying physiological patterns governing plant productivity and diversity in grasslands are critical to evaluate species responses to future environmental conditions of elevated CO₂ and nitrogen (N) deposition. In a 9‐year experiment, N was added to monocultures of seven C₃ grassland species exposed to elevated atmospheric CO₂ (560 μmol CO₂ mol^?1) to evaluate how N addition affects CO₂ responsiveness in species of contrasting functional groups. Functional groups differed in their responses to elevated CO₂ and N treatments. Forb species exhibited strong down‐regulation of leaf N_mass concentrations (?26%) and photosynthetic capacity (?28%) in response to elevated CO₂, especially at high N supply, whereas C₃ grasses did not. Hence, achieved photosynthetic performance was markedly enhanced for C₃ grasses (+68%) in elevated CO₂, but not significantly for forbs. Differences in access to soil resources between forbs and grasses may distinguish their responses to elevated CO₂ and N addition. Forbs had lesser root biomass, a lower distribution of biomass to roots, and lower specific root length than grasses. Maintenance of leaf N, possibly through increased root foraging in this nutrient‐poor grassland, was necessary to sustain stimulation of photosynthesis under long‐term elevated CO₂. Dilution of leaf N and associated photosynthetic down‐regulation in forbs under elevated [CO₂], relative to the C₃ grasses, illustrates the potential for shifts in species composition and diversity in grassland ecosystems that have significant forb and grass components.     

The type of competition modulates the ecophysiological response of grassland species to elevated CO2 and drought

The effects of elevated CO₂ and drought on ecophysiological parameters in grassland species have been examined, but few studies have investigated the effect of competition on those parameters under climate change conditions. The objective of this study was to determine the effect of elevated CO₂ and drought on the response of plant water relations, gas exchange, chlorophyll a fluorescence and aboveground biomass in four grassland species, as well as to assess whether the type of competition modulates that response. Elevated CO₂ in well‐watered conditions increased aboveground biomass by augmenting CO₂ assimilation. Drought reduced biomass by reducing CO₂ assimilation rate via stomatal limitation and, when drought was more severe, also non‐stomatal limitation. When plants were grown under the combined conditions of elevated CO₂ and drought, drought limitation observed under ambient CO₂ was reduced, permitting higher CO₂ assimilation and consequently reducing the observed decrease in aboveground biomass. The response to climate change was species‐specific and dependent on the type of competition. Thus, the response to elevated CO₂ in well‐watered grasses was higher in monoculture than in mixture, while it was higher in mixture compared to monoculture for forbs. On the other hand, forbs were more affected than grasses by drought in monoculture, while in mixture the negative effect of drought was higher in grasses than in forbs, due to a lower capacity to acquire water and mineral nutrients. These differences in species‐level growth responses to CO₂ and drought may lead to changes in the composition and biodiversity of the grassland plant community in future climate conditions.     

Restoration of grazing management and its effect on vegetation in an upland grassland

Question: How are plant species and functional group composition, and potential sward height affected by implementation of different grazing regimes on previously abandoned semi-natural grassland? Location: The Jizerské mountains, northern Czech Republic. Methods: We established a randomized block experiment with the following treatments: unmanaged control (U), intensive (IG) and extensive (EG) continuous grazing, first cut followed by intensive (ICG) and first cut followed by extensive (ECG) continuous grazing for the rest of the growing season. The percentage cover of all vascular plant species was recorded in 40 permanent plots. Results: Total plant species richness increased in all managed treatments, whereas species number was reduced in U at the end of the experiment. Tall forbs (Aegopodium podagraria, Galium album, Anthriscus sylvestris, Cirsium arvense) as well as tall grasses (Elytrigia repens and Alopecurus pratensis) were more abundant in U. Species associated with both grazing treatments (IG, EG) were Dactylis glomerata, Festuca rubra agg. and Phleum pratense. Agrostis capillaris, Taraxacum spp., Trifolium repens, Ranunculus repens and Cirsium vulgare were promoted by ECG and ICG. Abundance of tall grasses and tall forbs reflected the intensity of management in the order U>EG, ECG>IG, ICG. Prostrate forbs, on the other hand, increased their cover with increasing intensity: ICG>IG>ECG>EG. Conclusions: Plant species composition of semi-natural grasslands is affected by the defoliation regime. Continuous grazing on abandoned grassland alters the sward structure towards a permanent pasture with short, light-sensitive grasses and prostrate forbs. To maintain or enhance plant species richness in semi-natural grasslands, understanding the effects of different grazing regimes on plant species composition is necessary.     

Trampling enhances the dominance of graminoids over forbs in flooded grassland mesocosms

Questions: What are the interactive effects of flooding and cattle trampling upon the structural attributes and the floristic composition of a plant community? Do the effects on the plant community persist over an extended recovery period? Location: Flooding Pampa grasslands, Argentina (36°30′ S, 58°30′ W). Methods: We assessed the effects of 40‐d of flooding, trampling and the combination thereof on plant cover and biomass, vertical distribution of foliage and floristic composition in lowland grassland mesocosms. We considered a 120‐d recovery period to evaluate the persistence of flooding and trampling effects on the plant community. Results: Flooding, with or without trampling, increased cover and biomass of the graminoid species, especially marsh grasses, which developed a taller canopy, whereas most of the forb species were negatively affected. This was enhanced by trampling, as the aerial biomass of the dominant legume Lotus tenuis decreased by 90%, while three major forb species disappeared. Trampling under flooding conditions did not reduce the total above‐ground biomass production, as the growth enhancement of graminoids was enough to compensate for the breakdown of the forbs. Below‐ground biomass was lower when both perturbations occurred simultaneously. After 120‐d of recovery, graminoids continued to be dominant while the remaining forbs (including L. tenuis) recovered only partially. Below‐ground biomass recovered fully at the end of the growing season. Conclusions: The combination of flooding and trampling shifts the community co‐dominance of graminoids and forbs towards a persistent dominance of graminoid species. When both perturbations are combined, the above‐ground production of the grassland is unaffected and root biomass is rapidly recovered. However, the loss of the legume L. tenuis deserves attention because this is the unique nitrogen‐fixing species of the ecosystem, which improves the forage quality for livestock production.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Root exudation rate as functional trait involved in plant nutrient‐use strategy classification

Plants adopt a variety of life history strategies to succeed in the Earth's diverse environments. Using functional traits which are defined as “morphological, biochemical, physiological, or phonological” characteristics measurable at the individual level, plants are classified according to their species’ adaptative strategies, more than their taxonomy, from fast growing plant species to slower‐growing conservative species. These different strategies probably influence the input and output of carbon (C)‐resources, from the assimilation of carbon by photosynthesis to its release in the rhizosphere soil via root exudation. However, while root exudation was known to mediate plant‐microbe interactions in the rhizosphere, it was not used as functional trait until recently. Here, we assess whether root exudate levels are useful plant functional traits in the classification of plant nutrient‐use strategies and classical trait syndromes? For this purpose, we conducted an experiment with six grass species representing along a gradient of plant resource‐use strategies, from conservative species, characterized by low biomass nitrogen (N) concentrations and a long lifespans, to exploitative species, characterized by high rates of photosynthesis and rapid rates of N acquisition. Leaf and root traits were measured for each grass and root exudate rate for each planted soil sample. Classical trait syndromes in plant ecology were found for leaf and root traits, with negative relationships observed between specific leaf area and leaf dry matter content or between specific root length and root dry matter content. However, a new root trait syndrome was also found with root exudation levels correlating with plant resource‐use strategy patterns, specifically, between root exudation rate and root dry matter content. We therefore propose root exudation rate can be used as a key functional trait in plant ecology studies and plant strategy classification.     

Impact of plant functional group, plant species, and sampling time on the composition of nirK-type denitrifier communities in soil

We studied the influence of eight nonleguminous grassland plant species belonging to two functional groups (grasses and forbs) on the composition of soil denitrifier communities in experimental microcosms over two consecutive years. Denitrifier community composition was analyzed by terminal restriction fragment length polymorphism (T-RFLP) of PCR-amplified nirK gene fragments coding for the copper-containing nitrite reductase. The impact of experimental factors (plant functional group, plant species, sampling time, and interactions between them) on the structure of soil denitrifier communities (i.e., T-RFLP patterns) was analyzed by canonical correspondence analysis. While the functional group of a plant did not affect nirK-type denitrifier communities, plant species identity did influence their composition. This effect changed with sampling time, indicating community changes due to seasonal conditions and a development of the plants in the microcosms. Differences in total soil nitrogen and carbon, soil pH, and root biomass were observed at the end of the experiment. However, statistical analysis revealed that the plants affected the nirK-type denitrifier community composition directly, e.g., through root exudates. Assignment of abundant T-RFs to cloned nirK sequences from the soil and subsequent phylogenetic analysis indicated a dominance of yet-unknown nirK genotypes and of genes related to nirK from denitrifiers of the order Rhizobiales. In conclusion, individual species of nonleguminous plants directly influenced the composition of denitrifier communities in soil, but environmental conditions had additional significant effects.     

Urban domestic gardens (III): Composition and diversity of lawn floras

Question: How do lawn floras compare with those of semi‐natural grasslands? Are the compositions of lawn floras determined by local, within‐garden factors (e.g. lawn management and size) or by regional factors (e.g. climate and location)? Are lawn floras nested (like semi‐natural grasslands) or not (like cultivated parts of gardens)? Are there gradients of species composition within lawns? Location: Sheffield, UK. Methods: We examined the composition of the floras of entire lawns and of two 1‐m² lawn quadrats in 52 gardens. Results: A total of 159 species of vascular plants was recorded, 60 of them only once. Most lawn species were forbs, but most lawn cover consisted of grasses. Lawn species were predominantly natives. Bigger lawns had more species, but richness was not closely linked to other environmental or management variables. Composition of lawn floras varied with altitude, with woodland and wetland plants more common in the higher west of the city, and weeds of waste ground in the east. The species‐area curve derived from 1‐m² lawn quadrats was very similar to that of semi‐natural grasslands. Lawn quadrats were significantly nested, with rarer species mostly confined to more species‐rich quadrats. Trampling‐tolerant Poa annua was more abundant in the part of the lawn nearer the house. Conclusions: In most respects, lawns behaved much more like semi‐natural grasslands than like cultivated flower beds and borders. Species composition of lawns is strongly influenced by local climate. Most lawns show an internal gradient of composition, linked to a gradient of intensity of use.     

Seed traits,seed‐reserve utilization and offspring performance across pre‐industrial to future CO2 concentrations in a Mediterranean community

Atmospheric CO₂ enrichment can affect plants directly via impacts on their performance, and indirectly, by environment‐specific traits passed down from the mother plant to the offspring. Such maternal effects can significantly alter plant species composition, especially in annual ecosystems where the entire community is recruited from seeds each year. This study assessed impacts of future, high CO₂ (440 and 600 ppm) and pre‐industrial, low CO₂ (280 ppm) on seed traits and offspring performance in three plant functional groups (grasses, legumes, forbs) comprising 17 annual species of a semi‐arid Mediterranean community. In grasses, seed size and seed‐reserve utilization as expressed by root elongation tended to be higher at high than at low maternal CO₂, but total seed protein concentration and protein pool decreased with increasing maternal CO₂. The response of seed size to high CO₂ increased with increasing leaf‐mass fraction in grasses, and decreased with decreasing concentration of leaf non‐structural carbohydrates in legumes. Offspring development was studied at ambient CO₂, and showed reduced emergence success of high‐CO₂ progeny compared with low‐CO₂ progeny in forbs. Total biomass was lower in high‐CO₂ than in low‐CO₂ offspring across all functional groups. The biomass response to high maternal CO₂ in legume offspring correlated inversely with seed size, resulting in up to 25% lower biomass in large‐seeded species. Under the scenario of maternal effects combined with projected changes in biomass and seed production under direct exposure to high CO₂, legumes might gain and forbs and grasses might lose from future CO₂ enrichment. Most changes in seed traits and offspring performance were greater between pre‐industrial and near‐future CO₂ than between near‐ and remote‐future CO₂ concentrations. Hence, maternal effects of increasing CO₂ may contribute to current changes in plant productivity and species composition, and they need to be considered when predicting impacts of global change on plant communities.     

The roles of exotic grasses and forbs when restoring native species to highly invaded southern California annual grassland

Many semi-arid shrublands in the western US have experienced invasion by a suite of exotic grasses and forbs that have altered community structure and function. The effect of the exotic grasses in this area has been studied, but little is known about how exotic forbs influence the plant community. A 3-year experiment in southern California coastal sage scrub (CSS) now dominated by exotic grasses was done to investigate the influence of both exotic grasses (mainly Bromus spp.) and exotic forbs (mainly Erodium spp.) on a restoration seeding (9 species, including grasses, forbs, and shrubs). Experimental plots were weeded to remove one, both, or neither group of exotic species and seeded at a high rate with a mix of native species. Abundance of all species varied with precipitation levels, but seeded species established best when both groups of exotic species were removed. The removal of exotic grasses resulted in an increase in exotic and native forb cover, while removal of exotic forbs led to an increase in exotic grass cover and, at least in one year, a decrease in native forb cover. In former CSS now converted to exotic annual grassland, a competitive hierarchy between exotic grasses and forbs may prevent native forbs from more fully occupying the habitat when either group of exotics is removed. This apparent competitive hierarchy may interact with yearly variation in precipitation levels to limit restoration seedings of CSS/exotic grassland communities. Therefore, management of CSS and exotic grassland in southern California and similar areas must consider control of both exotic grasses and forbs when restoration is attempted.     

Impact of Plant Functional Group, Plant Species, and Sampling Time on the Composition of nirK-Type Denitrifier Communities in Soil

We studied the influence of eight nonleguminous grassland plant species belonging to two functional groups (grasses and forbs) on the composition of soil denitrifier communities in experimental microcosms over two consecutive years. Denitrifier community composition was analyzed by terminal restriction fragment length polymorphism (T-RFLP) of PCR-amplified nirK gene fragments coding for the copper-containing nitrite reductase. The impact of experimental factors (plant functional group, plant species, sampling time, and interactions between them) on the structure of soil denitrifier communities (i.e., T-RFLP patterns) was analyzed by canonical correspondence analysis. While the functional group of a plant did not affect nirK-type denitrifier communities, plant species identity did influence their composition. This effect changed with sampling time, indicating community changes due to seasonal conditions and a development of the plants in the microcosms. Differences in total soil nitrogen and carbon, soil pH, and root biomass were observed at the end of the experiment. However, statistical analysis revealed that the plants affected the nirK-type denitrifier community composition directly, e.g., through root exudates. Assignment of abundant T-RFs to cloned nirK sequences from the soil and subsequent phylogenetic analysis indicated a dominance of yet-unknown nirK genotypes and of genes related to nirK from denitrifiers of the order Rhizobiales. In conclusion, individual species of nonleguminous plants directly influenced the composition of denitrifier communities in soil, but environmental conditions had additional significant effects.     

Methane yield of biomass from extensive grassland is affected by compositional changes induced by order of arrival

Low‐input grassland biomass from marginal and other slightly more fertile sites can be used for energy production without competing with food or fodder production. The effect of grassland diversity on methane yield has received some attention, but we do not know how community assembly may affect methane yield from grassland biomass. However, methane yields determine the potential economic value of a bioenergy substrate. Hence, a better understanding of how plant community assembly affects methane yield would be important. We measured biomass production and methane yield in the second year of a grassland field experiment which manipulated the order of arrival of different plant functional groups (forbs, grasses or legumes sown first and all sown simultaneously) and sown diversity (9 vs. 21 species). The order of arrival of the plant functional groups significantly determined the relative dominance of each group which in turn mainly explained the variance in aboveground biomass production. Differences in area‐specific methane yields were driven by differences in biomass production and which plant functional groups dominated a plot. When grasses were sown first, legumes and grasses codominated a plot and the highest area‐specific methane yield was obtained. Overall, the results indicate that altering the order of arrival affected the community functional and species composition (and hence methane yields) much more than sown diversity. Our study shows that a combined use of positive biodiversity effects and guided plant community assembly may be able to optimize methane yields under field conditions. This may allow a guided, sustainable, and lucrative use of grassland biomass for biogas production in the future.     

Is leaf-level photosynthesis related to plant success in a highly productive grassland?

We addressed the question: “Are short-term, leaf-level measurements of photosynthesis correlated with long-term patterns of plant success?” in a productive grassland where interspecific competitive interactions are important. To answer this question, seasonal patterns of leaf-level photosynthesis were measured in 27 tallgrass prairie species growing in sites that differed in species composition and productivity due to differences in fire history. Our specific goals were to assess the relationship between gas exchange under field conditions and success (defined as aerial plant cover) for a wide range of species, as well as for these species grouped as dominant and sub-dominant grasses, forbs, and woody plants. Because fire increases productivity and dominance by grasses in this system, we hypothesized that any relationship between photosynthesis and success would be strongest in annually burned sites. We also predicted that regardless of fire history, the dominant species (primarily C₄ grasses) would have higher photosynthetic rates than the less successful species (primarily C₃ grasses, forbs and woody plants). Because forbs and woody species are less abundant in annually burned sites, we expected that these species would have lower photosynthetic rates in annually burned than in infrequently burned sites. As expected, the dominant C₄?grasses had the highest cover on all sites, relative to?other growth forms, and they had the highest maximum and seasonally averaged photosynthetic rates (17.6 ± 0.42 μmol m^?2 s^?1). Woody species had the lowest average cover as well as the lowest average photosynthetic rates, with subdominant grasses and forbs intermediate in both cover and photosynthesis. Also as predicted, the highest overall photosynthetic rates were found on the most productive annually burned site. Perhaps most importantly, a positive relationship was found between leaf-level photosynthesis and cover for a core group of species when data were combined across all sites. These data support the hypothesis that higher instantaneous rates of leaf-level photosynthesis are indicative of long-term plant success in this grassland. However, in contrast to our predictions, the subdominant grasses, forbs and woody species on the annually burned site had higher photosynthetic rates than in the less frequently burned sites, even though their average cover was lower on annually burned sites, and hence they were less successful. The direct negative effect of fire on plant cover and species-specific differences in the availability of resources may explain why photosynthesis was high but cover was low in some growth forms in annually burned sites.     

Independent effects of arbuscular mycorrhiza and earthworms on plant diversity and newcomer plant establishment

Questions: How do arbuscular mycorrhiza and earthworms affect the structure and diversity of a ruderal plant community? Is the establishment success of newcomer plants enhanced by these soil organisms and their interactions? Methods: We grew a native ruderal plant community composed of different functional groups (grasses, legumes and forbs) in the presence and absence of arbuscular mycorrhizal fungi (AMF) and endogeic earthworms in mesocosms. We introduced seeds of five, mainly exotic, plant species from the same functional groups after a disturbance simulating mowing. The effects of the soil organisms on the native ruderal plant community and seedling establishment of the newcomer plants were assessed. Results: After disturbance, the total above‐ground regrowth of the native plant community was not affected by the soil organisms. However, AMF increased plant diversity and shoot biomass of forbs, but decreased shoot biomass of grasses of the native plant community. Earthworms led to a reduction in total root biomass. Establishment of the introduced newcomer plants increased in the presence of AMF and earthworms. Especially, seedling establishment of the introduced non‐native legume Lupinus polyphyllus and the native forb Plantago lanceolata was promoted in the presence of AMF and earthworms, respectively. The endogeic earthworms gained more weight in the presence of AMF and led to increased extraradical AMF hyphal length in soil. However, earthworms did not seem to modify the effect of AMF on the plant community. Conclusion: The present study shows the importance of mutualistic soil organisms in mediating the establishment success of newcomer plants in a native plant community. Mutualistic soil organisms lead to changes in the structure and diversity of the native plant community and might promote newcomer plants, including exotic species.     

Estimates of root system topology of five plant species grown at steady-state nutrition

Results from a controlled growth-analysis experiment were used to illustrate some methods for measuring and describing root system topology. The experiment was performed in a nutrient solution system with an exponential nutrient supply and steady-state growth, to achieve well-defined levels of whole-plant nutrient status. Five naturally coexisting grassland species were included: The slow-growing forbs Polygala vulgaris L. and Crepis praemorsa (L.) F. L. Walth., and the grass Danthonia decumbens (L.) DC. were compared with the more common, fast-growing grasses Agrostis capillaris L. and Dactylis glomerata L. The most marked difference in morphological indices was a much higher specific root length in the grasses than in the forbs, which implies thinner roots. In contrast to the conclusions of previous studies, an index of the topology for the grasses was very similar to that for the forbs. The specific root lenght and link length apparently vary more between species and nutrient levels than topology does, and may therefore be more ecologically important. The only clear plastic response to growth-limiting nitrogen supply was a markedly increased link length in P. vulgaris. There were also indications that nitrogen limitation led to more herringbone-like root systems in P. vulgaris and C. praemorsa. In general, there was a clear tendency for the estimates of topology to change with plant size, which may make many topological indices, especially those based on regression slopes, very difficult to interpret. Until interactions with plant size, other morphological parameters and among-plant competition can be properly understood, the relevance of root topology for plant performance remains unclear. This revised version was published online in June 2006 with corrections to the Cover Date.     

Plant–community responses to shrub cover in a páramo grassland released from grazing and burning

Shrub encroachment can follow grazing or burning release in páramo grasslands. While encroachment decreases herbaceous species richness in some grassland systems, the effects of this process on the herbaceous community in páramo grasslands are currently unknown. We collected data on shrub cover, herbaceous‐species cover and species composition in a páramo grassland 12 years after release from burning and cattle grazing near Zuleta, Ecuador. Topographic and soil measures were also included as predictor variables of differences in community composition. Contrary to studies in other systems, shrub cover did not have a significant effect on herbaceous‐species richness, whereas shrub‐species richness significantly increased with shrub cover. However, shrub cover was associated with significant shifts in herbaceous–community composition. Most notably, there was an increase in some shade‐tolerant forbs and tall‐statured wetland grasses with increasing shrub cover, and a corresponding decrease in some short‐statured grasses and early successional forbs. These results could indicate that the ameliorative effects of shrubs (e.g. frost and wind protection) in harsh alpine environments may partially compensate for the expected competitive effect of shrubs due to shading.     

Nitrogen addition decreases seed germination in a temperate steppe

Seed germination and seedling establishment play an important role in driving the responses of plant community structure and function to global change. Nitrogen (N) deposition is one of the driving factors of global change, which often leads to a loss in species richness in grassland ecosystems. However, how seed germination responds to N addition remains unclear. A pot incubation test was conducted in a semi‐arid grassland in the Mongolian Plateau, Northern China, to investigate the effect of N addition (0, 5, 10, 20, 40, and 80 g N/m²) on seed germination from May to October 2016. Twenty species germinated under all treatments; however, the responses of the 20 species to N addition were different. The densities of Stipa krylovii, Leymus chinensis, and Artemisia frigida, which are the dominant species in this temperate steppe, decreased significantly as the amount of N addition. Moreover, N addition significantly suppressed seedling densities of the community, perennial forbs, perennial grasses, and annuals and biennials. Furthermore, species richness of the community, perennial forbs, and annuals and biennials decreased sharply with increasing N addition level, but perennial grass species richness did not change. The Shannon–Wiener diversity index also decreased as the amount of N addition increased. Our results suggest that N enrichment plays an important role in the seed germination stage and decreases supplements of seedlings to adult plants. These findings may help explain the causes of species loss by atmospheric N deposition in grassland ecosystems.     

Restoration of Native Warm Season Grassland Species in a Tall Fescue Pasture Using Prescribed Fire and Herbicides

Pastures dominated by tall fescue (Schedonorus phoenix (Scop.) Holub) cover much of the eastern United States, and there are increasing efforts to restore native grassland plant species to some of these areas. Prescribed fire and herbicide are frequently used to limit the growth of tall fescue and other non‐natives, while encouraging native grasses and forbs. A fungal endophyte, commonly present in tall fescue, can confer competitive advantages to the host plant, and may play a role in determining the ability of tall fescue plants to persist in pastures following restoration practices. We compared vegetation composition among four actively restored subunits of a tall fescue pasture (each receiving different combinations of prescribed fire and/or herbicide) and a control. We also measured the rate of endophyte infection in tall fescue present within each restoration treatment and control to determine if restoration resulted in lower tall fescue cover but higher endophyte infection rates (i.e. selected for endophyte‐infected individuals). Tall fescue cover was low in all restoration treatments and the control (1.1–17.9%). The control (unmanaged) had higher species richness than restoration treatments and plant community composition was indicative of succession to forest. Restoration practices resulted in higher cover of native warm season grasses, but in some cases also promoted a different undesirable species. We found no evidence of higher fungal endophyte presence in tall fescue following restoration, as all subunits had low endophyte infection rates (2.2–9.3%). Restoration of tall fescue systems using prescribed fire and herbicide may be used to promote native grassland species.