Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0–10 cm² basal area), medium (10–30 cm²), and large ( > 30 cm²) clones were initially mapped in five 0.91-m² quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.     

How season of grazing and herbivore selectivity influence monsoon tall-grass communities of northern Australia

Abstract. The hypothesis that season of defoliation and herbivore selectivity may be as important as level of use in determining plant community response to grazing was tested in a monsoon grassland in northern Australia. Plots, dominated by the tussock grasses Themeda triandra and Chrysopogon fallax, were grazed by cattle at low, medium and high rates of utilization in either the early wet, late wet or dry seasons. Effects of grazing on species composition were greatest in the early wet season when high rates of utilization significantly reduced the proportion and occurrence of Themeda and increased the proportion of forbs. Grazing in the dry season had no significant effect on composition. At medium and high levels of utilization in the early wet season, the pasture responded negatively to defoliation, only partially compensating for plant tissue lost to herbivory. The negative response to defoliation carried over to the next wet season when these same medium and high-grazing treatments produced only 80 % and 60 % growth, respectively, of that in treatments grazed at low levels of utilization or those grazed during the dry season. The frequency of Themeda was still lower, and that of annual grasses and non-leguminous forbs higher, in plots that had been grazed at a high rate of utilization for just eight weeks in the early wet season two years previously. Species richness and diversity were also significantly affected by this grazing disturbance. If species composition is to be maintained in these grasslands then stocking rates must be set at low levels to cope with the combined effect of undercompensation in response to defoliation in the wet season and strong dietary preferences for grazing sensitive species.     

Old-field successional dynamics on the Inland Pampa,Argentina

Abstract. The first 10 yr of old-field successional dynamics on the Argentine Inland Pampa were studied on a series of adjacent plots established consecutively between 1978 and 1989. We examined differences in species abundance patterns among plots in order to detect the spatial and temporal variability of succession. Perennial grasses steadily increased in cover and replaced the dominant annual species after 5 yr. Pioneer dicots persisted in older seral stages with 20 — 23 species/plot. Overall, exotic species (mostly the grasses Lolium multiflorum and Cynodon dactylon) contributed much to the plant cover in these communities. Native grasses comprised 45 % of total cover at years 7 — 10 of succession, but occurred with less than 7 species/plot. Substantial variation was found in the successional pathway, which reflected the particular sequence from annual forbs to short-lived and perennial grasses in the various plots. The course of succession was apparently influenced by a 2-yr period of unusually high rainfall. Deyeuxia viridiflavescens, a native perennial grass virtually absent before the wet period, spread over the study area and dominated seral communities for 3 yr, irrespective of plot age. Climatic conditions thus affected the successional turnover of life forms by increasing the rate of colonization by perennial grasses. We further point out the constraints imposed on secondary succession by the life histories of ‘available’ species.     

Foraging ecology of roan antelope: key resources during critical periods

The use of landscape zones and grass species by roan antelope, a species threatened with local extirpation within South Africa's Kruger National Park, were investigated. Plant‐based observations of grazing were made within a 300 ha enclosure in the roan range, where 40 roan antelope were confined at high density in the absence of other grazers. The study spanned the dry seasons of two years, one with average rainfall and one with low rainfall. We recorded changes in the extent of grazing of different grass species, height differences between grazed and ungrazed tillers and intensity of cropping per tuft. In the average year, the grazing pressure in the bottomland grassland was twice that in the upland savanna, with two tall grass species bearing the brunt of the grazing through the dry season. Two highly palatable upland grasses were also extensively grazed by the mid dry season. In the dry year, the extent of grazing in the upland exceeded that in the bottomland, and several upland grass species little used the previous year became heavily grazed. Roan antelope appeared to be separated ecologically from more common grazers by their selective use of tall grasses growing in the drainage line grassland during the critical dry season months. However, their grazing expansion into the upland savanna during the dry year potentially brought them into competition with these grazers. Nevertheless, their population performance did not suffer despite the high‐density conditions. Heightened predation pressure following an influx of these grazers, rather than resource limitation, appeared to be primarily responsible for the drastic decline of this species in the park.     

Effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, South Africa

Moisture, nutrients, fire and herbivory are the principal factors governing tree–grass cover ratios of savannas. We investigated tree (Acacia mellifera) recruitment after fire and under conditions of maximum-recorded rainfall, nitrogen addition and grazing in a completely-crossed field experiment. We employed a similar garden experiment with the exception of the fire treatment. Tree germination in the field was extremely low, probably due to below-average natural rainfall in plots that only received natural rain, and insufficient watering frequency in irrigated plots. Due to low germination in the field experiment, no treatment significantly affected tree recruitment. In the garden experiment, frequent watering, nutrient control (i.e. no nitrogen addition) and grazing enhanced tree recruitment with significant interactions between rain, nitrogen and grazing. We infer that above-average rainfall years with frequent rainfall events are required for mass tree recruitment. Grass defoliation makes space and resources available for tree seedlings. Nitrogen enrichment increases the competitive ability of fast-growing grasses more than that of the N₂-fixing tree component. In contrast to conventional wisdom that grazing alone causes encroachment, we suggest that there are complex interactions between the above-mentioned factors and ‘triggering’ events such as unusually high rainfall.     

Testing the limits of resistance: a 19‐year study of Mediterranean grassland response to grazing regimes

A synthesis of a long‐term (19 years) study assessing the effects of cattle grazing on the structure and composition of a Mediterranean grassland in north‐eastern Israel is presented, with new insights into the response of the vegetation to grazing management and rainfall. We hypothesized that the plant community studied would be resistant to high grazing intensities and rainfall variability considering the combined long history of land‐use and unpredictable climatic conditions where this community evolved. Treatments included manipulations of stocking densities (moderate, heavy, and very heavy) and of grazing regimes (continuous vs. seasonal), in a factorial design. The effect of interannual rainfall variation on the expression of grazing impacts on the plant community was minor. The main effects of grazing on relative cover of plant functional groups were related to early vs. late seasonal grazing. Species diversity and equitability were remarkably stable across all grazing treatments. A reduction in tall grass cover at higher stocking densities was correlated with increased cover of less palatable groups such as annual and perennial thistles, as well as shorter and prostrate groups such as short annual grasses. This long‐term study shows that interannual fluctuations in plant functional group composition could be partly accounted for by grazing pressure and timing, but not by the measured rainfall variables. Grazing affected the dominance of tall annual grasses. However, the persistence of tall grasses and more palatable species over time, despite large differences in grazing pressure and timing, supports the idea that Mediterranean grasslands are highly resistant to prolonged grazing. Indeed, even under the most extreme grazing conditions applied, there were no signs of deterioration or collapse of the ecosystem. This high resistance to grazing intensity and interannual fluctuation in climatic conditions should favor the persistence of the plant community under forecasted increasing unpredictability due to climate change.     

Growth and nutrient content of perennial grasslands following burning in semi-arid,sub-tropical Australia

Fire often increases the productivity of perennial tussock grasslands inmesic environments but can reduce growth for one or more growing seasons inaridand semi-arid environments. We examined effects of single-burns on growth andnutrient content of grasslands in sub-tropical, northwestern Australia. Thesegrasslands were dominated by Themeda triandra, a speciesoften managed by regular burning in wetter temperate and tropical zones. Burnswere in the late dry season and were replicated using small plots (5 ×5-m) within fenced areas at two sites.Total projective cover and aboveground biomass were significantly less in burntplots relative to controls for 2.5 years after burning despite four growingseasons, including the first summer, of above-average rainfall. Recovery ofburnt plots was hindered by an extended dry period in the second year,demonstrating that rainfall in subsequent seasons can be as important asrainfall in the first season in determining post-burn productivity ofgrasslandsin semi-arid environments. Greater decreases in grass cover in burnt plotsduring the extended dry period may have been due to less standing dead andlitter than controls, and therefore less insulation from extreme summertemperatures, although relationships between cover changes and cover at thestart of the period were weak. With the exception of increased pH near grasstussocks, burning had little effect on chemical characteristics of surfacesoilsin the first week. Concentrations of N, and particularly P, in abovegroundplantmaterial were greater in burnt plots four months after burning, followingsummerrains, but were either less than or similar to those in controls withincreasingly dry conditions. Significantly lower concentrations of P in greenfoliage from burnt plots during dry seasons, when uptake from soil pools wouldbe minimal, indicated that burning decreased P retranslocation from plantstores. However, we found no evidence that single-burns increased nutrientlimitations to growth because plant contents of N and P were comparable inburntand control plots during periods of adequate water supply. Our data supportprevious generalizations that prescribed burning of perennial tussockgrasslandsin semi-arid environments is mostly unnecessary because putative benefits ofincreased productivity and forage quality, characteristic of more mesicenvironments, were not realized.     

Response of a Native Perennial Grass Stand to Disturbance in California's Coast Range Grassland

To assess the potential for enhancing an existing stand of native perennial grasses on a California Coast Range Grassland site, we experimentally manipulated the seasonal timing and presence of grazing for 3 years (1994 through 1996) and of autumn burning for 2 years (1994 and 1995) and measured species cover for 6 years (1993 through 1998). We subjected the species matrix to classification (TWINSPAN) and ordination (CCA) and tested the ordination site scores as well as diversity indices with linear mixed effects models. Four distinct plant community groups emerged from the classification. Two of these were dominated by annual grasses and two by perennial grasses. No treatment effects were observed on diversity. For composition, temporal and spatial random effects were important mixed effects model parameters, as was the fixed effect covariate, pre‐treatment CCA site score, indicating the importance of random environmental variation and initial starting conditions. Incorporation of these random effects and initial condition terms made for more powerful tests of the fixed effects, grazing season, and burning. We found no significant burning effects. Grazing removal imparted a shift in plant community from more annual‐dominated toward more perennial‐dominated vegetation. Individual perennial grass species responded differently according to genus and species. Nassella spp. increased gradually over time regardless of grazing treatment. Nassella pulchra (purple needlegrass) increase was greatest under spring grazing and N. lepida (foothill needlegrass) was greatest with grazing removal. Danthonia californica (California oatgrass) had little response over time under seasonal grazing treatments, but increased with grazing removal. Under relatively mesic weather conditions it appears that grazing removal from Coast Range Grasslands with existing native perennial grass populations can increase their cover. However if N. pulchra is the sole existing population, spring season‐restricted grazing should be equally effective at enhancing cover of the native grass species.     

Continuous grazing disrupts desert grass-soil seed bank composition under variable rainfall

Long-term field studies help unveil mechanisms of grass soil seed bank (SSB) persistence and resilience. We wonder whether grazing is a disruptive force that changes grass SSB composition and size, while heavy rainfall is a positive bottom-up force that restores the grass SSB in grazed areas of the Monte desert, Argentina. We used core sampling to study the grass SSB in cattle-free (17 year) and grazed (10 year) grasslands. The SSB size in cattle-free sites usually responded in a continuous positive fashion to rainfall, and the proportion of palatable grasses in the SSB was similar among years. Most grass species formed a transient SSB, but the presence of a stand of perennial plants would prevent their local extinction. In contrast, the SSB size in the grazed habitats showed a threshold response to rainfall (i.e. it did not increase under low-moderate rainfall), the seeds of palatable grasses were always scarce, and the proportion of seeds of less palatable grasses increased under extreme rainy conditions. Seed production from grasses that grew tillers from axillary buds during a drought was zero, suggesting that the bank of plants could have a role in the replenishment of the grass SSB only in mesic years. Within the time span assessed here, continuous heavy grazing together with scarce as well as heavy rainfall were disruptive forces that reduced the number and proportion of seeds from palatable grasses in the SSB, which are also the species preferred by seed-eating animals.

     

Defoliation and competition effects in a productivity gradient for a semiarid Mediterranean annual grassland community

Grazing and competition are two main factors shaping range plant communities; however, few studies have investigated their interaction. The current study aimed to investigate the effects of defoliation, competition and their interaction on production of annual grasses in semiarid Mediterranean areas. Competition treatments (absence/presence of neighbors) were combined with three defoliation intensities (0%, 30% and 60%) in a complete factorial design. Competition significantly reduced grass biomass. However, the role of competition was eliminated under heavy defoliation or under dry growth conditions. Defoliation showed variable results on final biomass (FB) and cumulative biomass (CB). While heavy defoliation (60% clipping intensity) reduced grass FB down to 80% during the two growing seasons, light defoliation (30%) significantly increased CB. Results showed that competition may limit the direct effect of defoliation on dominant grass species. Further, the relationship between site productivity and competition effect was best explained by a negative linear model. This hypothesized model may suggest that facilitation and competition alternatively affect grassland communities along a productivity gradient. The results suggest that light grazing may sustain or even enhance grassland productivity. The results also indicated the suitability of annual grass species to re-vegetate degraded rangeland in semi-arid climate. Further, optimum grazing practices to conserve biodiversity of Avena grassland may involve moderate stocking rate.     

Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin,USA

As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether “nurse plants” will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220–350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25–33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.     

Variations in mountain vegetation use by reindeer (Rangifer tarandus) affects dry heath but not grass heath

Question: Are differences in landscape use of semi‐domesticated reindeer reflected in the vegetation of summer grazing grounds? Location: Alpine heaths, central east Sweden. Methods: Dry heath and grass heath vegetation plots with inferred grazing intensities (high, intermediate and low) were selected a priori from an interpolated pellet count map compiled in 2002. In each plot, faecal pellets were counted, environmental variables measured and vegetation sampled by listing presence and absence. Species composition was compared with a detrended correspondence analysis, and a canonical correspondence analysis was used to infer relations between species composition and environmental variables. Plots were also clustered to provide groupings for an indicator species analysis. Results: Significant differences in faecal pellet count were present between the highest and lowest grazing intensities for both vegetation types, showing that the pattern in the interpolated pellet maps was robust. Differences in species composition between grazing intensities were found for the dry heath only. Here, there was an apparent grazing gradient, with lichens and mosses in the low‐use plots and grasses and herbs in the high‐use plots. No such gradient was found for the grass heath. Conclusions: Within the dry heath vegetation type, grazing levels had a subtle effect on the vegetation, while no effects were seen in the grass heath, probably as a result of the dominance of more grazing‐tolerant graminoids. Even in the dry heath, species richness did not differ between grazing levels, but the relative abundances of species differed.     

Ecosystem engineering varies spatially: a test of the vegetation modification paradigm for prairie dogs

Colonial, burrowing herbivores can be engineers of grassland and shrubland ecosystems worldwide. Spatial variation in landscapes suggests caution when extrapolating single‐place studies of single species, but lack of data and the need to generalize often leads to ‘model system’ thinking and application of results beyond appropriate statistical inference. Generalizations about the engineering effects of prairie dogs (Cynomys sp.) developed largely from intensive study at a single complex of black‐tailed prairie dogs C. ludovicianus in northern mixed prairie, but have been extrapolated to other ecoregions and prairie dog species in North America, and other colonial, burrowing herbivores. We tested the paradigm that prairie dogs decrease vegetation volume and the cover of grasses and tall shrubs, and increase bare ground and forb cover. We sampled vegetation on and off 279 colonies at 13 complexes of 3 prairie dog species widely distributed across 5 ecoregions in North America. The paradigm was generally supported at 7 black‐tailed prairie dog complexes in northern mixed prairie, where vegetation volume, grass cover, and tall shrub cover were lower, and bare ground and forb cover were higher, on colonies than at paired off‐colony sites. Outside the northern mixed prairie, all 3 prairie dog species consistently reduced vegetation volume, but their effects on cover of plant functional groups varied with prairie dog species and the grazing tolerance of dominant perennial grasses. White‐tailed prairie dogs C. leucurus in sagebrush steppe did not reduce shrub cover, whereas black‐tailed prairie dogs suppressed shrub cover at all complexes with tall shrubs in the surrounding habitat matrix. Black‐tailed prairie dogs in shortgrass steppe and Gunnison's prairie dogs C. gunnisoni in Colorado Plateau grassland both had relatively minor effects on grass cover, which may reflect the dominance of grazing‐tolerant shortgrasses at both complexes. Variation in modification of vegetation structure may be understood in terms of the responses of different dominant perennial grasses to intense defoliation and differences in foraging behavior among prairie dog species. Spatial variation in the engineering role of prairie dogs suggests spatial variation in their keystone role, and spatial variation in the roles of other ecosystem engineers. Thus, ecosystem engineering can have a spatial component not evident from single‐place studies.     

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

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

California perennial grasses are physiologically distinct from both Mediterranean annual and perennial grasses

In the Central Valley of California, native perennial grass species have been largely replaced by Eurasian annual species, while in many parts of the Mediterranean Basin native perennial grasses continue to dominate, even on disturbed or degraded sites. We assessed whether differences in summer rainfall patterns have lead to the development of different plant-water strategies between grasses from these two regions. We compared six measures of plant-water physiology for three guilds of grasses: California perennial grasses, Mediterranean perennial grasses, and Mediterranean annual grasses. Discriminant analysis distinguished between the three guilds; Mediterranean perennial grasses were characterized by a more conservative water-relations physiology than Mediterranean annual grasses, whereas California perennial grasses were in some ways intermediate between the two Mediterranean grass guilds. For individual traits, California perennial grasses were either intermediate or more like Mediterranean annuals than Mediterranean perennials. Our results suggest California perennials are more drought tolerant than Mediterranean annuals but less drought tolerant than Mediterranean perennials, despite the fact that California??s Central Valley has a more intense summer drought than the Mediterranean Basin. These patterns may help explain why Mediterranean annuals, but not Mediterranean perennials, have been more successful invaders of interior California grasslands.     

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.     

Do rhizome severing and shoot defoliation affect clonal growth of Leymus chinensis at ramet population level?

Leymus chinensis (Trin.) Tzvel. is a perennial species of Gramineae, usually subject to defoliation from grazing and mowing. We examined whether shoot defoliation and rhizome severing affected rhizome and ramet growth, and vegetative bud outgrowth of L. chinensis ramet populations. We also tested the hypothesis that clonal growth of the ramets subject to defoliation would benefit from clonal integration between interconnected ramets besides from possible compensatory growth. To 48 experimental plots, we applied six treatments resulting from interactions between two rhizome connection states (unsevered/severed) and three defoliation regimes (non-defoliated, mildly-defoliated and heavily-defoliated). Defoliation affected rhizome growth and bud outgrowth, but had little effect on shoot growth. Mild and heavy defoliation exerted similar effects on rhizome growth. Only heavy defoliation significantly reduced bud outgrowth while mild defoliation did not. The fact that shoot growth did not change after defoliation and that the bud numbers remained unchanged after mild defoliation suggest that the compensatory response enable the species to tolerate grazing to some extent. Neither rhizome severing nor the interaction of rhizome severing and defoliation had effect on any tested variables. Lack of the effect of rhizome severing falsified the first half of our hypothesis, that is, clonal integration was unimportant in our experiment. The probable reasons were suspected to be the short duration of the experiment and/or the buffer effect of carbohydrate reserves in rhizomes for shoot growth and bud production in time of defoliation.     

Abundance of trees and grasses in a woodland savanna in relation to environmental factors

The relationships of plant species associations and underlying environmental factors in a woodland savanna in South Africa were investigated. 40 plots were included with 25 tree and 17 grass species dominating the arboreal and ground cover. Correspondence Analysis described the relationships between soil moisture retention, soil nutrients and the abundance of trees and grasses. Dry matter indices represented the accumulated effects of rainfall, fire and grazing of the herbaceous layer. Variations in the abundance of plants corresponded to well-defined gradients of soil nutrients. The distribution of grass and tree species along the ordination axes indicated that soils with high water retention capacity and high nutrient contents provided a suitable substrate for many of the tree species sampled. However, grass species abundance was high in plots with porous soils and poor nutrient availability.     

Water relations in competitive interactions of Mediterranean grasses and shrubs

In Mediterranean ecosystems, competition between opportunistic grasses and slower-growing woody species may affect the speed and path of ecosystem recovery and the success of restoration plantings after natural or human-induced disturbance. In this experiment, competitive interactions between Mediterranean annual and perennial grass species (Avena fatua and Brachypodium retusum, respectively) and an important Mediterranean shrub (Rosmarinus offlcinalis) were examined under semi-controlled conditions simulating wet and dry Mediterranean rainfall regimes. The identity of the grass competitor and the level of water availability in the plots interacted to produce differing rates of R. offlcinalis growth but similar levels of mortality. In particular, competition with the perennial grass resulted in very low rates of R. offlcinalis growth at both irrigation levels. Measurements of soil water content showed that both grasses reduced soil moisture to low levels, though this effect was temporary in the case of the winter annual grass. Resistance to hydraulic flow in roots was highest in the perennial grass, smaller but of similar magnitude in the shrub, and much lower in the annual grass. Transpirational response to decreasing leaf water potential was a quick, sharp drop in conductance in R. offlcinalis, in contrast to a moderated decline from much lower initial transpiration rates in B. retusum. The annual grass largely maintained both leaf water potential and transpiration through leaf-tip senescence and death. Quantification of the rate of hydric recuperation of leaves after irrigation of drought-stressed plants showed that the perennial grass recovered at a rate four times that of R. offlcinalis, suggesting a strategy for making quick use of rare summer rains that may contribute to its competitive success. The appropriateness of planting or suppressing grasses in restoration of disturbed sites in Mediterranean Spain is discussed.     

Not novel, just better: competition between native and non-native plants in California grasslands that share species traits

Invasive plants have often been shown to possess novel traits such as the ability to fix nitrogen, access unused resource pools, or the ability to exude allelopathic chemicals. We describe a case of a successful invasion where the native and non-native species are very similar in most life-history characteristics including their growth forms, lifespan, and degree of summertime activity. Data from permanent transects suggest that exotic perennial grass invaders can establish into intact native-dominated grasslands, achieving cover values from 6 to 71% over several years. We also established a 4-year competition experiment to test the effect of each group—the native and non-native perennial grasses—on the other. Competitive interactions were found to consistently favor the non-native grasses: native perennial grass productivity was significantly lower in plots with exotic perennial grasses as compared to plots without exotic perennial grasses. By contrast, productivity of the exotic perennial grasses was not reduced by the presence of the native perennial grasses. These results suggest that competitive ability, rather than a unique trait, has contributed to the success of the exotic perennial grasses in our system. Management tools to control exotic perennial grass invasions are likely to negatively influence native perennial grass populations, as strategies that succeed against the invasive species may kill or reduce the native species as well.