Effect of burning and mowing on grass and forb diversity in a long‐term grassland experiment

Abstract. Disturbance may be an important determinant of plant community composition and diversity owing to its effects on competitive interactions, resources, dominance and vigour. The effect of type, timing and frequency of disturbance on grass and forb species richness was examined using data from a long‐term (> 50 yr) grassland burning and mowing experiment in KwaZulu‐Natal, South Africa. Grass species richness declined considerably (> 50%) in the absence of disturbance, whereas forb species richness was unaffected. Annual burning in sites not mown in summer tended to increase grass species richness relative to triennial burning (22% increase) with the reverse being true in sites mown twice in summer (37% decline). Forb species richness declined by 25% in sites mown twice in summer relative to sites mown in early summer only. Disturbance was necessary to achieve maximum grass species richness presumably by removing litter and increasing the availability of light. The interaction of time of mowing in summer (early versus late) and time of burning during the dormant period (spring versus winter) had the most dramatic effect on species richness. Time of burning had no effect on richness in sites mown in early summer, but winter burning resulted in a dramatic decline (27–42%) in richness in sites mown in late summer. This effect may be related to possible greater soil desiccation with this combination of disturbances.     

Aboveground Biomass Removal by Burning and Raking Increases Diversity in a Reconstructed Prairie

Prescribed spring burning often contributes to a predominance of C₄ grasses and low forb abundance and is impractical at many sites, especially near development. We tested raking after mowing as an alternative to prescribed burning in a reconstructed Minnesota prairie. We also tested mowing without raking as a possible means of maintaining prairie communities. Frequency, flowering stem abundance, and cover were measured for all plant species and native functional groups (C₄ grasses, C₃ graminoids, forbs, legumes, and annual or biennial forbs). Mowing alone did not differ from the control in its effect on any functional groups of plants. Round‐headed bush clover (Lespedeza capitata), a legume, and Black‐eyed Susan (Rudbeckia hirta), a biennial, increased in frequency with treatments that removed biomass (i.e., fire or raking), but they did not have significantly more flowering stems. Thus, new plants established well from seed, whereas the vitality of mature plants did not change. Raking had similar effects to burning on most functional groups, although flowering stems of C₄ grasses were significantly more abundant after fire than after raking. Burning reduced some C₃ forbs and grasses and favored the dominance of C₄ grasses. Therefore, raking after mowing in the spring provides an alternative to prescribed burning that has many of the same positive aspects as fire but does not promote aggressive C₄ grasses to the same extent.     

Plant Community Biomass Shifts in Response to Mowing and Fencing in Invaded Oak Meadows with Non‐Native Grasses and Abundant Ungulates

Many herbaceous meadows are dominated by competitive non‐native grasses and subject to ungulate herbivory, ecological processes that shift the proportional biomass of plant groups in the community. Predicting the outcome of restoration is complicated because herbivory and competition can interact. We examined the relationship between herbivory by native black‐tailed deer and domestic sheep and dominance of non‐native grasses in Garry oak meadows, one of North America's most endangered habitat types. A 3‐year factorial experiment tested the effects of mowing and fencing on plant community biomass, categorized into eight groups by geographic origin (native/non‐native), growth form (annual/perennial), and plant type (forb/grass). To test if the rarity of native plant groups was related to herbivory, we estimated ungulate foraging preferences for each plant group. Mowing and fencing treatments interacted for annual and perennial non‐native grasses. Dominance was shifted from non‐native to native grasses only when both mowing and fencing were applied. Fencing increased the total biomass, whereas mowing had no overall effect; however, fencing alone did not affect any individual plant group. Mowing shifted dominance from grasses to forbs, although both native and non‐native forbs benefited from the increased light availability. We also noted that herbivore fecal pellet densities were greatest in the spring, which coincided with the peak season of their preferred plant group, native perennial forbs. Overall, applying both mowing and fencing was the most effective restoration treatment to increase native plant groups and biomass.     

Fire Effects on Cover and Dietary Resources of Sage-Grouse Habitat

Abstract: We evaluated 6 years of vegetation response following prescribed fire in Wyoming big sagebrush (Artemisia tridentata spp. wyomingensis) steppe on vegetation cover, productivity, and nutritional quality of forbs preferred by greater sage-grouse (Centrocercus urophasianus), and abundance of common arthropod orders. Habitat cover (shrubs and tall herbaceous cover [>18 cm ht]) was about 50% lower after burning compared to unburned controls because of the loss of sagebrush. Perennial grasses and an invasive annual forb, pale alyssum (Alyssum alyssoides), increased in cover or yield after fire. There were no increases in yield or nutritional quality of forb species important in diets of sage-grouse. Abundance of ants (Hymenoptera), a significant component in the diet of young sage-grouse, decreased after fire. These results suggest that prescribed fire will not improve habitat characteristics for sage-grouse in Wyoming big sagebrush steppe where the community consists of shrubs, native grasses, and native forbs.     

Effects of Frequent Mowing on Survival and Persistence of Forbs Seeded into a Species-Poor Grassland

Many early attempts at tallgrass prairie reconstruction failed to achieve the high species diversity of remnant prairies, and instead consist primarily of C₄ grasses. We hypothesized that frequent mowing of established prairie grasses could create sufficient gaps in the aboveground and belowground environment to allow for the establishment of native forbs from seed. We studied forb seedling establishment in a 25‐year‐old prairie planting in northern Iowa that was dominated by native warm‐season grasses. In winter 1999, 23 species of native forbs were broadcast into the recently burned sod at a rate of 350 viable seeds/m². Treatment plots were mowed weekly for either one or two growing seasons, and control plots were unmowed. Mowed plots had greater light availability than controls, especially when warm‐season grasses began to flower. Overwinter seedling mortality was 3% in mowed treatments compared to 29% in the controls. Forbs in mowed plots had significantly greater root and shoot mass than those in control plots in the first and second growing seasons but were not significantly more abundant. By the fourth growing season, however, forbs were twice as abundant in the mowed treatments. No lasting negative impacts of frequent mowing on the grass population were observed. Mowing a second year influenced species composition but did not change total seedling establishment. Experimental evidence is consistent with the idea that mowing reduced competition for light from large established grasses, allowing forb seedlings the opportunity to reach sufficient size to establish, survive, and flower in the second and subsequent years.     

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.     

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.     

Mowing Reduces Exotic Annual Grasses but Increases Exotic Forbs in a Semiarid Grassland

Cheatgrass (Bromus tectorum) and other exotic winter‐active plants can be persistent invaders in native grasslands, growing earlier in the spring than native plants and pre‐empting soil resources. Effective management strategies are needed to reduce their abundance while encouraging the reestablishment of desirable native plants. In this 4‐year study, we investigated whether mowing and seeding with native perennial grasses could limit growth of exotic winter‐actives, and benefit growth of native plants in an invaded grassland in Colorado, United States. We established a split‐plot experiment in October 2008 with 3 mowing treatments: control, spring‐mowed, and spring/summer‐mowed (late spring, mid‐summer, and late summer), and 3 within‐plot seeding treatments: control, added B. tectorum seeds, and added native grass seeds. Cover of plant species and aboveground biomass were measured for 3 years. In March and June of 2010, 2011, and March of 2012, B. tectorum and other winter‐annual grasses were half as abundant in both mowing treatments as in control plots; however, cover of non‐native winter‐active forbs increased 2‐fold in spring‐mowed plots and almost 3‐fold in spring/summer‐mowed plots relative to controls. These patterns remained consistent 1 year after termination of treatments. Native cool‐season grasses were most abundant in spring‐mowed plots, and least abundant in control plots. There was higher cover of native warm‐season grasses in spring/summer‐mowed plots than in control plots in July 2011 and 2012. The timing of management can have strong effects on plant community dynamics in grasslands, and this experiment indicates that adaptive management can target the temporal niche of undesirable invasive species.     

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.     

The relationships among root and leaf traits of 76 grassland species and relative abundance along fertility and disturbance gradients

For 76 annual, biennial, and perennial species common in the grasslands of central Minnesota, USA, we determined the patterns of correlations among seven organ-level traits (specific leaf area, leaf thickness, leaf tissue density, leaf angle, specific root length, average fine root diameter, and fine root tissue density) and their relationships with two traits relating to growth form (whether species existed for part of the growing season in basal, non-caulescent form and whether species were rhizomatous or not). The first correlation of traits showed that grasses had thin, dense leaves and thin roots while forbs had thick, low-density leaves and thick roots without any significant differences in growth form or life history. The second correlation of traits showed a gradient of species from those with high-density roots and high-density erect leaves to species with low-density roots and low-density leaves that were held parallel to the ground. High tissue density species were more likely to exist as a basal rosette for part of the season, were less likely to be rhizomatous, and less likely to be annuals. We examined the relationships between the two axes that represent the correlations of traits and previously collected data on the relative abundance of species across gradients of nitrogen addition and disturbance. Grasses were generally more abundant than forbs and the relative abundance of grasses and forbs did not change with increasing nitrogen addition or soil disturbance. High tissue density species became less common as fertility and disturbance increased.     

Determinants of C3 forb growth and production in a C4 dominated grassland

Forbs are the most abundant species within the vascular flora of tallgrass prairie and they make the greatest contribution to biodiversity of all growth forms. However, little is known about the factors that determine their productivity and growth rates. The objective of this study was to assess the controls of forb growth (absolute and relative) and production in tallgrass prairie from a long-term burning experiment at the Konza Prairie in NE Kansas. Over the 14-year study, forbs comprised 9% of the total biomass production on sites with a high fire frequency vs. 29% on the low fire frequency site, with gramminoids accounting for the remainder. Although interannual variations in peak biomass of the grasses was strongly correlated with environmental variables related to water availability, there were no similar relationships for forbs, suggesting that production of forbs and grasses responded to interannual variations in climate in different ways. Multivariate analysis of climatic controls on growth rates of grasses and forbs yielded similar results. Although forbs had low biomass and absolute growth per unit ground area in frequently burned prairie, their relative growth rates were highest in such sites. Thus, it appears that reduced growth rates of individual forbs per se do not limit forb success in annually burned prairie. Instead, direct negative effects of fire on forbs (increased mortality) may be responsible. Determinants of forb growth and productivity in unburned prairie remain unresolved.     

Emergence and survival of herbaceous seedlings in temperate grassy woodlands: Recruitment limitations and regeneration niche

Abstract Field experiments examined herbaceous seedling emergence and survival in temperate grassy woodlands on the New England Tablelands of New South Wales. Effects of intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds, grazing and seed theft by ants on seedling emergence and survival were studied in two field experiments. Thirteen species with a range of traits were used in the experiments and their cumulative emergence was compared with laboratory germination studies. Field emergence correlated to laboratory germination but all species had lower emergence in the field. Little natural emergence of native species was observed in the field in unsown treatments. Short‐lived forbs had the highest emergence, followed by perennial grasses; rhizomatous graminoids and perennial forbs had the lowest emergence. Soil surface and cover treatments did not markedly enhance emergence suggesting that intertussock spaces were not prerequisites for forb emergence. No consistent pattern of enhanced emergence was found for any treatment combination across all species. Seedling survival varied among species, with perennial grasses and short‐lived forbs having the highest seedling mortality. Low mortality rates in the graminoids and rhizomatous forbs appeared partially to compensate for lower seedling emergence. All perennial grasses and some short‐lived forbs showed increased risk of mortality with grazing. Differences in emergence and survival of species were related to ground cover heterogeneity, soil surfaces and, to some extent, herbivory. The complexity of these patterns when superimposed on temporal variability suggests that no generalizations can be made about the regeneration niche of herbaceous species groups. Strong recruitment limitation and partitioning of resources in the regeneration niche may reduce competition among native species and explain the high species richness of the herbaceous layer in the temperate grassy communities of eastern Australia.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

The impact of herbivore exclusion on forb diversity: Comparing species and functional responses during a drought

Rainfall and herbivory shape savannah herbaceous communities, but these disturbances are being altered globally. To assess potential consequences of such alterations, we evaluated herbivore effects on species and functional diversity during an episodic drought in a sodic savannah using data collected from long-term herbivore exclosures in the Kruger National Park, South Africa. Herbaceous life forms are rarely acknowledged as distinct functional entities. Moreover, the functional ecology of forbs remains elusive. Here, we present disturbances–responses by forbs separately from grasses. We hypothesised that combinations of intense utilisation and drought would be associated with low diversity and high dominance at species and functional levels for both life forms. Contrary to our hypothesis, low forb and grass diversity was associated with long-term herbivore exclusion, which exceeded expected undesirable effects of intense utilisation and drought. Grasses responded less sensitively, suggesting that forbs respond dynamically to changes in herbivore assemblage when these alterations are combined with drought. Consistent with patterns in savannah systems, forbs contributed significantly to species and functional trait diversity. High forb diversity is suggested to enhance resilience of this nutrient-rich ecosystem against declines in its functioning when subjected to drought and alterations in herbivory.     

Trait–environment relations for dominant grasses in South African mesic grassland support a general leaf economic model

Question: Following the framework of Suding et al. (2003), we examined whether morphological traits (organismal response), tolerance and competitive effect (specific process response) were associated with grass dominance (abundance response) on burning, mowing, fertilization and soil depth gradients in KwaZulu‐Natal (KZN), South Africa. Location: University of KwaZulu‐Natal, Pietermaritzburg, South Africa. Methods: Using several pot experiments involving 29 grass species in total, we determined the vegetative traits, competitive effect and response, and tolerance to shading for grasses common in closed, tufted mesic grassland in KZN. Results: The primary axis of grass–trait variation was most strongly related to a negative correlation (trade‐off) between growth rate and specific leaf area (SLA), with broad‐leaved, rapidly‐growing grasses (high SLA) occupying one extreme and narrow‐leaved, slow‐growing grasses (low SLA) the other extreme of the first principal component. The low SLA, slow‐growth strategy was found to be a relatively general strategy among grasses dominant in undisturbed, high litter grassland, as well as those adapted to moisture‐stressed habitats. In contrast, grasses dominant in highly productive habitats with some form of disturbance, e.g. mowing, had a broad‐leaved, rapid‐growth strategy. Intermediate combinations of the SLA–growth rate trade‐off were common among grasses dominant under other combinations of disturbance and soil resource availability. Conclusions: Distinct patterns of organismal (SLA, growth rate) and specific process (competitive effect and response, as well as tolerance of shading) responses appeared to be associated with grasses dominant on gradients of burning, mowing, fertilization and soil depth. These organismal and specific process responses were similar to those for North American and European grasses dominant under the same environmental influences, suggesting that some general trait–environment patterns exist at an inter‐continental scale. This general trait–environment relationship appears to be driven by functional adaptive selection along the SLA–environment continuum and its unavoidable trade‐off with growth rate.     

Manipulating disturbance regimes and seeding to restore mesic Mediterranean grasslands

Question: Can managing disturbance regimes alone or in combination with seeding native species serve to shift the balance from exotic towards native species? Location: Central coast of California, USA. Methods: We measured vegetation composition for 10 yr in a manipulative experiment replicated at three sites. Treatments included no disturbance, grazing and clipping at three frequencies with and without litter removal. We seeded eight native species into clipped plots and compared cover in comparable plots with no seeding. Results: Regardless of frequency, clipping generally shifted community dominance from exotic annual grasses to exotic annual forbs, rather than consistently favoring native species. At one site, perennial grass cover decreased in no‐disturbance plots, but only after 4 yr. Litter removal had minimal impact on litter depth and plant community composition. Grazing had a highly variable effect on the abundance of different plant guilds across sites and years. Seeding increased abundance of only two of eight native species. Conclusions: Managing disturbance regimes alone is insufficient to restore native species guilds in highly‐invaded grasslands and seeding native species has highly variable success.     

Can Carbon and Phosphorous Amendments Increase Native Forbs in a Restoration Process? A Case Study in the Northern Tall‐grass Prairie (U.S.A.)

In the northern Great Plains (United States), sites with less than 20% of native species are difficult to restore. We have experimented with a restoration method that shows some promise. It consists of systematically installing simulated small‐scale patches (8.0 m² in size) over 25% of an old field and then seeding these patches with native species. The working hypothesis is that these patches will generate a constant source of propagules which in time will lead to increases in native species diversity within the surrounding grass matrix. The objective of this paper was to determine whether soil amendments should be used to facilitate the establishment and persistence of native species (primarily forbs) within these patches. We seeded the patches with a mixture of native grass and forb species and applied four soil treatments: P fertilization, C additions, C + P, and a control (no amendments). Results for the first 5 years were as follows: (1) seeded forb richness was mostly unaffected by soil amendments; (2) seeded and nonseeded forb biomass and density were substantially reduced by C additions, whereas they were unaffected or increased under P additions; (3) both seeded and non‐native grass biomass substantially increased with C additions; and (4) there was an inverse relationship between native seeded forbs and non‐native grass biomass. Our conclusions are that: (1) P amendments are a potential tool for enhancing native seeded forb biomass in simulated small‐scale disturbance patches; and (2) C additions, although enhancing seeded grass biomass do not reduce the biomass of non‐native grasses.     

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.     

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.     

Plant response to disturbance in a Mediterranean grassland: How many functional groups?

Abstract. Data referring to changes in vegetation composition resulting from cattle exclosure and ploughing in a Portuguese pasture dominated by annuals were used to test hypotheses regarding the biology of species favoured or eliminated by disturbance in semi-natural herbaceous communities. These hypotheses were tested in two ways. First we compared the distribution of six a priori groups – grasses, small rosettes, large rosettes, small species with leafy stems, large species with leafy stems, legumes – across grazed, ploughed and undisturbed plots. In a second set of analyses we examined changes in the frequencies of individual biological attributes in response to grazing and ploughing. These analyses were carried out separately for grasses and dicot forbs. Overall, the species composition showed little response to either grazing or ploughing, though species dominance changed. This lack of responsiveness of species composition was attributed to the long history of intensive land use which has resulted in the loss of disturbance-intolerant species over entire landscapes. When considering a priori groups, small rosettes were indifferent to disturbance. grazing and ploughing showed that dominated. Large rosettes, large species with leafy stems and legumes were generally intolerant to both grazing and ploughing, though individual species may increase in response to disturbance. Small species with leafy stems were the only group favoured by grazing whereas ploughing favoured grasses. As to individual traits, grazing excluded large grass species with heavy seeds and promoted a flat rosette canopy structure and a small size, along with a moderate dormancy and protected inflorescences. In forbs, grazing favoured small species, as expected, while it excluded tall species, and, in contrast to earlier results, a rosette canopy. These attributes were consistent with responses of the a priori groups, though it would not have been possible to reconstruct groups directly from the attribute list. Ploughing had no effect on any of the forb traits. As to grass traits, flat- and short-statured species increased and heavy-seeded species decreased. Our analysis revealed two advantages of establishing plant functional classifications within life forms. Subgroups within forbs had contrasting types of behaviour. For the same trait patterns could differ within the grass group from within the forb group. Finally, this analysis emphasizes the need for plant functional classifications aiming at the identification of syndromes of co-occurring attributes rather than of lists of isolated traits of which actual combinations are not specified.