Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Understory response to varying fire frequencies after 20 years of prescribed burning in an upland oak forest

Ecosystems in the eastern United States that were shaped by fire over thousands of years of anthropogenic burning recently have been subjected to fire suppression resulting in significant changes in vegetation composition and structure and encroachment by invasive species. Renewed interest in use of fire to manage such ecosystems will require knowledge of effects of fire regime on vegetation. We studied the effects of one aspect of the fire regime, fire frequency, on biomass, cover and diversity of understory vegetation in upland oak forests prescribe-burned for 20 years at different frequencies ranging from zero to five fires per decade. Overstory canopy closure ranged from 88 to 96% and was not affected by fire frequency indicating high tolerance of large trees for even the most frequent burning. Understory species richness and cover was dominated by woody reproduction followed in descending order by forbs, C3 graminoids, C4 grasses, and legumes. Woody plant understory cover did not change with fire frequency and increased 30% from one to three years after a burn. Both forbs and C3 graminoids showed a linear increase in species richness and cover as fire frequency increased. In contrast, C4 grasses and legumes did not show a response to fire frequency. The reduction of litter by fire may have encouraged regeneration of herbaceous plants and helped explain the positive response of forbs and C3 graminoids to increasing fire frequency. Our results showed that herbaceous biomass, cover, and diversity can be managed with long-term prescribed fire under the closed canopy of upland oak forests.     

Fire and seedling population dynamics in western Oregon prairies

Abstract. Through an experiment in three prairie vegetation types in western Oregon, USA the effect of prescribed fire on the timing and rates of seedling emergence and mortality was examined. Seeds of common exotic and native prairie species were sown into burned and unburned plots in late September, 1995. Emerged seedlings were censussed the following winter, early spring and late spring. Results indicated that spring population levels could not be forecast by fall seedling flushes, as winter survival was important in seedling establishment. The bulk of emergence for all grass and annual forb species occurred in the fall, followed by low to severe winter mortality. Perennial forbs were more variable in emergence times but, once emerged, perennial forb seedlings were likely to become established. Burning caused a statistically significant increase in seedling accumulation through emergence and survival in 11 of 23 cases. Burning improved seedling winter survival for most grass and short‐lived forb species and increased emergence of perennial forb species. These patterns were most conspicuous on the two sites dominated by exotic species, where burning significantly improved the accumulation of seedlings from most native species tested. Thus, prescribed burning might be a useful restoration tool in these communities. In contrast, two of the three species increased by burning in the native bunchgrass site were exotic pest plants, suggesting that fire should be prescribed with caution.     

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.     

Temporal and spatial patterns in emergence and early survival of perennial plants in the Sonoran Desert

Seedling emergence and survival of 15 perennial species were studied for six years in a 557-m² permanent plot at Tumamoc Hill, Arizona, USA, an ungrazed site in the northern Sonoran Desert. The minimum rain required for germination and emergence ranged from 17.5 to 35.6 mm. Few species emerged in every year of the study. First-year survival averaged across all 15 species was 3.7%; only 0.1% of seedlings lived as long as four years. The odds of survival in the first year improved with increased rain. About three times as many seedlings died from predation as desiccation. In 2-m² subplots, mortality of three woody species in the first 30 days after emergence appeared to be independent of seedling density. Short-, moderate-, and long-lived species displayed distinct survival strategies. Long-lived species compensated for generally poor seedling survival by frequent germination and emergence. Moderate-lived species exhibited highly episodic germination and emergence, a potentially risky behavior that might have been offset to some extent by relatively good long-term survival. Short-lived species had the highest seedling survival. Because these species can bloom in their first year, good early survival meant that some individuals were able to reproduce before they died.     

Influence of Grazing on Soil Seed Banks Determines the Restoration Potential of Aboveground Vegetation in a Semi‐arid Savanna of Ethiopia

Species composition, number of emerging seedlings, species diversity and functional group of the soil seed banks, and the influence of grazing on the similarity between the soil seed banks and aboveground vegetation, were studied in 2008 and 2009 in a semi‐arid savanna of Ethiopia. We tested whether the availability of persistent seeds in the soil could drive the transition from a degraded system under heavy grazing to healthy vegetation with ample perennial grasses. A total of 77 species emerged from the soil seed bank samples: 21 annual grasses, 12 perennial grasses, 4 herbaceous legumes, 39 forbs, and 1 woody species. Perennial grass species dominated the lightly grazed sites, whereas the heavily grazed sites were dominated by annual forbs. Heavy grazing reduced the number of seeds that can germinate in the seed bank. Species richness in the seed bank was, however, not affected by grazing. With increasing soil depth, the seed density and its species richness declined. There was a higher similarity in species composition between the soil seed bank and aboveground vegetation at the lightly grazed sites compared with the heavily grazed sites. The mean similarity between the seed banks and aboveground vegetation was relatively low, indicating the effect of heavy grazing. Moreover, seeds of perennial grasses were less abundant in the soil seed banks under heavy grazing. We concluded that restoration of grass and woody species from the soil seed banks in the heavily grazed areas could not be successful in semi‐arid savannas of Ethiopia.     

Effectiveness of repeated autumn and spring fires for understorey restoration in weed‐invaded temperate eucalypt woodlands

Question. Can strategic burning, targeting differing ecological characteristics of native and exotic species, facilitate restoration of native understorey in weed‐invaded temperate grassy eucalypt woodlands? Location. Gippsland Plains, eastern Victoria, Australia. Methods. In a replicated, 5‐year experimental trial, the effects of repeated spring or autumn burning were evaluated for native and exotic plants in a representative, degraded Eucalyptus tereticornis grassy woodland. Treatments aimed to reduce seed banks and modify establishment conditions of exotic annual grasses, and to exhaust vegetative reserves of exotic perennial grasses. Treatments were applied to three grassland patch types, dominated by the native grass Austrodanthonia caespitosa, ubiquitous exotic annuals, or the common exotic perennial grass Paspalum dilatatum. Results. The dominant native grass Austrodanthonia caespitosa and native forbs were resilient to repeated fires, and target exotic annuals and perennials were suppressed differentially by autumn and spring fires. Exotic annuals were also suppressed by drought, reducing the overall treatment effects but indicating important opportunities for restoration. The initially sparse exotic geophyte Romulea rosea increased in cover with fire and the impact of this species on native forbs requires further investigation. There was minimal increase in diversity of subsidiary natives with fire, probably owing to lack of propagules. Conclusions. While fire is often considered to increase ecosystem invasibility, our study showed that strategic use of fire, informed by the relative responses of available native and exotic taxa, is potentially an effective step towards restoration of weed‐invaded temperate eucalypt woodlands.     

Factors Affecting Revegetation of Oil Field Access Roads in Semiarid Grassland

We assessed vegetation recovery on access roads removed after well abandonment in an active oil‐producing region of northern Great Plains grasslands. We compared extant vegetation on 58 roads, restored 3–22 years previously, to records of species seeded on each and to adjacent, undisturbed prairie, to evaluate main differences between the restored and adjacent community and to explore patterns in the restored plant community over time. The restored plant community was dominated by low richness of seeded non‐native and native grasses and forbs, whereas adjacent prairie had numerous, abundant native graminoids and shrubs and higher richness of native forbs. Cover of seeded species on roads was double that of colonizing species. Disparity in cover of dominant native grasses between the adjacent community and relatively narrow restored roadway suggests that conditions for germination and survival in roadbeds are poor. This is at least partly due to persistence of seeded species. Differences in restored plant composition over time were best explained by changes in species seeded, from non‐natives to natives, and secondarily by successional shifts from ruderal to perennial non‐seeded species. Of the 30 species seeded at least once on these roads, only 10 were commonly used. The long‐term influence of seeding choices in grassland road restorations implies that improvements in these practices will be critical to reversing ecological impacts of roads.     

Plant population and community responses to removal of dominant species in the shortgrass steppe

Question: What are the plant population‐ and community‐level effects of removal of dominant plant species in the shortgrass steppe? Location: The Shortgrass Steppe Long‐Term Ecological Research site in northern Colorado, USA. Methods: We annually measured plant cover and density by species for 10 years after a one‐time aboveground removal of the dominant perennial grass, Bouteloua gracilis. Removal and control plots (3 m × 3 m) were within grazed and ungrazed locations to assess the influence of grazing on recovery dynamics. Our analyses examined plant species, functional type, and community responses to removal, paying special attention to the dynamics of subdominant and rare species. Results: Basal cover of B. gracilis increased by an average of 1% per year, but there was significantly less plant cover in treatment compared to control plots for 5 years following removal. In contrast to the lower cover in treatment plots, the plant density (number of plants m^?2) of certain subdominant perennial grasses, herbaceous perennial and annual forbs, a dwarf shrub, and cactus increased after removal of the dominant species, with no major change in species richness (number of species per 1 m × 1 m) or diversity. Subdominant species were more similar between years than rare species, but dominant removal resulted in significantly lower similarity of the subdominant species in the short term and increased the similarity of rare species in the long term. Conclusions: Removal of B. gracilis, the dominant perennial grass in the shortgrass steppe, increased the absolute density of subdominant plants, but caused little compensation of plant cover by other plants in the community and changes in species diversity.     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

Influence of fire and mechanical sagebrush reduction treatments on restoration seedings in Utah,United States

Overabundance of woody plants in semiarid ecosystems can degrade understory herbaceous vegetation and often requires shrub reduction and seeding to recover ecosystem services. We used meta‐analysis techniques to assess the effects of fire and mechanical shrub reduction over two post‐treatment timeframes (1–4 and 5–10 years) on changes in cover and frequency of 15 seeded species at 63 restoration sites with high potential for recovery. Compared to mechanical treatments, fire resulted in greater increases in seeded species. Native shrubs did not increase, and forbs generally declined over time; however, large increases in perennial grasses were observed, suggesting that seeding efforts contributed to enhanced understory herbaceous conditions. We found greater increases in a few non‐native species than native species across all treatments, suggesting the possibility that interference among seeded species may have influenced results of this regional assessment. Differences among treatments and species were likely driven by seedbed conditions, which should be carefully considered in restoration planning. Site characteristics also dictated seeded species responses: while forbs showed greater increases in cover over the long term at higher elevation sites considered to be more resilient to disturbance, surprisingly, shrubs and grasses had greater increases in cover and frequency at lower elevation sites where resilience is typically much lower. Further research is needed to understand the causes of forb mortality over time, and to decipher how greater increases of non‐native relative to native seeded species will influence species diversity and successional trajectories of restoration sites.     

Herbaceous species responses to long-term effects of prescribed fire, grazing and selective tree cutting in the savanna-woodlands of West Africa

Grazing, fire and selective tree cutting are major disturbances that shape species diversity in savanna ecosystems, yet their effects are highly variable. We carried out a factorial experiment with two levels to examine the effects of grazing, fire and selective tree cutting on herbaceous species richness, abundance and diversity on two sites in the Sudanian savanna-woodlands of Burkina Faso for 10 years (1994–2003). The results showed significant inter-annual variation in species richness, abundance and diversity at both sites (p<0.001), while main or combined effects of fire, grazing and selective cutting were very limited and varied between life forms and sites. Grazing tended to favour the diversity of perennial grasses; fire tended to influence the richness of annual grasses and abundance and diversity of perennial grasses while selective tree cutting had no effect on any of the vegetation attributes assessed. The combined effect of grazing, fire and selective cutting tended to increase the diversity of forbs. In many cases, the responses of herbaceous species to treatments were clearer on the site with deeper soils than the one with shallow soils. Depending on the site and treatments, the inter-annual variation in vegetation attributes was partly related to amount and/or frequency of rainfall and partly to inter-annual variation in grazing or fire intensity. It can be concluded that both disturbances and climatic condition influence the structure and diversity of herbaceous flora in the Sudanian savanna-woodland ecosystem. The responses were site-specific, which accentuates the importance of landscape-scale approaches to understand the impacts of disturbances on composition, structure and diversity of savanna ecosystems.     

冷蒿草原土壤可萌发种子库特征及其对放牧的响应

（１）采用萌发试验法对内蒙古冷蒿（Ａｒｔｅｍｉｓｉａ ｆｒｉｇｉｄａ）草原土壤种子特征及其在不同放牧压力下的变化进行了研究。（２）冷蒿草原可萌发种子库由４种多年生禾草,１１种种多年生杂草及３种１年生植物所组成。（３）每样中的可萌发种子是变化于４５到３０５之间,平均１０２／样。多年生禾草、多年生杂类草及１年生植物的土壤中种子量分别占总可萌发种子库的４６．５７％、５１．９６％和１．５７％。多年和禾 草     

Seedling emergence patterns and dormancy/germination physiology of <Emphasis Type="Italic">Primula modesta</Emphasis> in a subalpine region

On the basis of the germination/dormancy responses of seeds to temperature and light, and local seed rain, we attempted to interpret the seedling emergence patterns of Primula modesta Bisset et Moore (Primulaceae) in two different types of habitats in a subalpine zone of Mt. Asama: an oligotrophic flat moor and a grassland with relatively dense herbaceous vegetation cover. The seasonal pattern of seedling emergence was well explained by the dormancy/germination physiology revealed in laboratory germination tests. The seeds were demonstrated to have a strict light requirement even after experiencing moist chilling, which might facilitate the incorporation of the seeds into the soil seed bank. Despite sufficient seed production, the seedlings emerging were far less at the grassy site than the moor site, but the number of seedlings was significantly dependent on the seed rain within previous season and on the litter cover of the microsite in both sites. Therefore, the spatiotemporal patterns of seedling emergence in the habitats could be well explained by the spatiotemporal patterns of seed rain and safe-sites for germination.     

Effects of Established Perennial Grasses on Introduction of Native Forbs in California

Prairie restoration is not complete without the establishment of both grasses and forbs. However, if desirable forbs and grasses are seeded simultaneously, control of broadleaf weeds is problematic. If possible, a two‐step process of introducing forbs after establishing grasses would allow use of broadleaf‐specific herbicides at the critical early stages of grass growth. We conducted experiments to investigate methods for introducing forbs into previously restored native perennial grasslands on rural roadsides in the Sacramento Valley of California. In one experiment, we studied the effects of background vegetation (established perennial grasses or tilled ground) on seven native forb species planted from seed. In a second experiment, we evaluated the effects of background vegetation (existing perennial grasses or tilled ground) and container size (36 ml or 105 ml) with excavation technique (excavation by core removal [core] or by creating an impression [dibble]) on the growth of transplants of the native perennial forbs Asclepias fascicularis and Sisyrinchium bellum. The presence of established perennial grasses reduced the growth of seeded forbs, but did not affect transplants, indicating the vulnerability of seedling forbs to interference. When compared to control plots that had been tilled in the autumn, weed canopy cover was significantly lower in the presence of perennial grasses if seeded with forbs, but not in the presence of perennial grasses alone. Both transplanted species grew better in the large container/core treatment than the small container/dibble treatment; however, existing grasses eliminated these positive effects. Asclepias fascicularis performed better when grown in large containers than in small containers, but its growth was not affected by excavation method; S. bellum performed better when planted with the core method than the dibble method of excavation, but container size made no difference. We attribute differences in the responses of the species to interactions between phenological differences and expansive clay soils that naturally de‐compact upon drying.     

Contrasting effects of grazing on the early stages of woody encroachment in a Neotropical savanna

Woody encroachment in savannas represents an ecological process of current global interest given its negative impact on ecosystem functioning, particularly on forage production. Traditional savanna models propose competition and niche differentiation as the main mechanisms allowing tree-grass coexistence. Demographic models, instead, propose abiotic and biotic factors as bottlenecks controlling vital rates and transitions from seeds to adult trees. The role played by domestic grazing on woody encroachment is yet controversial. Here, using a multistage tree life approach, we combine both models and evaluate the role of grazing and herbaceous vegetation on woody recruitment in a Neotropical savanna dominated by Vachellia caven, a successful and widely spread encroacher tree species. We performed three experiments to evaluate seed predation, seedling emergence and survival of V. caven by manipulating cattle grazing (grazed and ungrazed areas) and herbaceous vegetation presence (vegetated and unvegetated). Finally, we combined the results of the three experiments to estimate the probability of plant recruitment across these experimental factors. Grazing decreased seed predation by half, did not modify seedling emergence and decreased seedling survival. Herbaceous vegetation did not affect seed predation nor seedling emergence rate, but increased seedling survival. Overall, the net effect of grazing on V. caven recruitment was neutral since the increase in seed availability due to the reduction in seed predation rate was compensated by the negative effect of grazing on seedling survival. Our analysis revealed that cattle grazing and herbaceous vegetation had contrasting effects on the seed and seedling life stages. We propose that in order to restrain the early stages of encroachment, cattle grazing pressure could be managed following the seasonality of demographic tree transitions. Through rotational grazing amongst paddocks, stocking rates could be relaxed during the primary dispersal stage to maximize granivory, and then increased to enhance the chance of seedling consumption and trampling.     

Post-pastoral changes in composition and guilds in a semi-arid conservation area, Central Otago, New Zealand

Changes in the vegetation of Flat Top Hill, a highly modified conservation area in semi;arid Central Otago, New Zealand, are described four years after the cessation of sheep and rabbit grazing. Unusually moist weather conditions coincide with the four-year period of change in response to the cessation of grazing. Between 1993 and 1997, the average richness and diversity (H') of species increased, and the average proportion of native species decreased significantly. The vegetation was significantly richer in exotic annual and perennial grass species, exotic perennial forbs, exotic woody species and native tussock grasses in 1997 than in 1993. Eight response guilds of species are identified. Most "remnant" native shrubs and forbs were stable, in that they remained restricted to local refugia and showed little change in local frequency. However, taller native grass species increased, some locally, and others over wide environmental ranges. Rare native annual forbs and several native perennial species from "induced" xeric communities decreased, and this may be a consequence of competition from exotic perennial grasses in the absence of grazing. The invasive exotic herb Sedum acre decreased in abundance between 1993 and 1997, but several other prominent exotic species increased substantially in range and local frequency over a wide range of sites. Exotic woody species, and dense, sward-forming grasses are identified as potential threats to native vegetation recovery.     

不同载畜率对荒漠草原群落结构和功能群生产力的影响

采用随机区组试验研究了内蒙古高原荒漠草原亚带短花针茅(S tip a brev if lora)草原群落不同载畜率对草原群落结构和功能群生产力的影响。结果表明:随着载畜率的增加,群落的植物种数逐渐减少,在每个载畜率水平下,冷蒿(A rtem isia f rig id a)占有绝对优势地位,优势度为35.66%~41.95%;群落的地上生物量随载畜率的增加而降低,各功能群地上生物量组成中,灌木类处于主体地位,分别占据了群落68%(CK)、77.40%(LG)、73.25%(M G)、76.91%(HG)的生物量;在植物生活型功能组成中,灌木类和杂类草、多年生丛生禾草和杂类草功能群,在生物量上具有生态互补效应(n iche com p lem en tary effect)。     

Longleaf Pine (Pinus palustris P. Mill.) Restoration Using Herbicides: Overstory and Understory Vegetation Responses on a Coastal Plain Flatwoods Site in Florida,U.S.A.

A study was conducted on a Coastal Plain flatwoods site in Florida to determine the effects of common forestry herbicides on Longleaf pine seedling survival and growth and on the understory vegetation. Following removal of the overstory slash pine, five low‐rate herbicide treatments were applied over the top of planted Longleaf pine seedlings to provide short‐term understory vegetation control and accelerate seedling growth. The objective was to increase Longleaf pine growth by reducing the shrub competition while increasing the herbaceous ground cover. Despite causing reduction in seedling survival over the control treatment, imazapyr (0.21 ae kg/ha) resulted in the highest seedling growth (height and volume). The significant reduction of shrub cover, density, and height by imazapyr was believed to be responsible for the improved seedling growth in this treatment. Both hexazinone (0.56 ai kg/ha) and sulfometuron methyl (0.26 ai kg/ha) + hexazinone (0.56 ai kg/ha) treatments also reduced cover of Runner oak, a major shrub species, but the response was evident only 8 months after treatment. Although sulfometuron methyl (0.26 ai kg/ha) and sulfometuron methyl + hexazinone treatments did not result in any significant change in overall grass, forb, and shrub cover, both treatments resulted in greater Longleaf pine growth compared to the control. None of the herbicides significantly affected the major understory grasses and forbs. Overall, imazapyr provided the best desired results with significant increase in seedling growth and better control of shrub species with no significant effects on grass and other herbaceous species cover.     

Grazing effects on plant functional group diversity in Mediterranean shrublands

Grazing is one of the prevalent human activities that even today are taking place inside protected areas with direct or indirect effects on ecosystems. In this study we analyzed the effects of grazing on plant species diversity, plant functional group (PFG) diversity and community composition of shrublands. We analyzed plant diversity data from 582 sampling plots located in 66 protected areas of the Greek Natura 2000 network, containing in total 1102 plant species and subspecies. We also classified a priori all plant species in seven PFGs: annual forbs, annual grasses/sedges, legumes, perennial forbs, perennial grasses/sedges, small shrubs and tall shrubs. For each site, grazing intensity was estimated in four classes (no grazing, low, medium and high grazing intensity). We found that, at the spatial and temporal scale of this study, as grazing intensity increased, so did total species richness. However, each PFG displayed a different response to grazing. Short-lived species (annual grasses or forbs and legumes) benefited from grazing and their species richness and proportion in the community increased with grazing. Perennial grasses and forbs species richness increased with grazing intensity, but their dominance decreased, since their proportion in the community declined. Short shrub species richness remained unaffected by grazing, while tall shrub diversity decreased. Finally, in sites without grazing the spatial pattern of species richness of the different PFGs was not congruent with each other, while in grazed sites they were significantly positively correlated (with the exception of tall shrubs). This finding may imply that grazing is a selective pressure organizing the community structure, and imposing a certain contribution of each PFG. So, in Mediterranean shrublands in protected areas with a long historical record of grazing, it seems that grazing promotes species diversity and its continuation on a portion of the landscape may be a necessary part of an effective management plan.