Restoration of a Native Shrubland Impacted by Exotic Grasses, Frequent Fire, and Nitrogen Deposition in Southern California

Natural ecosystems globally are often subject to multiple human disturbances that are difficult to restore. A restoration experiment was done in an urban fragment of native coastal sage scrub vegetation in Riverside, California that has been subject to frequent fire, high anthropogenic nitrogen deposition, and invasion by Mediterranean annual weeds. Hand cultivation and grass‐specific herbicide were both successful in controlling exotic annual grasses and promoting establishment of seeded coastal sage scrub vegetation. There was no native seedbank left at this site after some 30 years of conversion to annual grassland, and the only native plants that germinated were the seeded shrubs, with the exception of one native summer annual. The city green‐waste mulch used in this study (C:N of 39:1) caused short‐term N immobilization but did not result in decreased grass density or increased native shrub establishment. Seeding native shrubs was successful in a wet year in this Mediterranean‐type climate but was unsuccessful in a dry year. An accidental spring fire did not burn first‐year shrubs, although adjacent plots dominated by annual grass did burn. The shrubs continued to exclude exotic grasses into the second growing season, suggesting that successful shrub establishment may reduce the frequency of the fire return interval.     

Restoration of Native Perennials in a California Annual Grassland after Prescribed Spring Burning and Solarization

Grasslands dominated by exotic annual grasses have replaced native perennial vegetation types in vast areas of California. Prescribed spring fires can cause a temporary replacement of exotic annual grasses by native and non‐native forbs, but generally do not lead to recovery of native perennials, especially where these have been entirely displaced for many years. Successful reintroduction of perennial species after fire depends on establishment in the postfire environment. We studied the effects of vegetation changes after an April fire on competition for soil moisture, a key factor in exotic annual grass dominance. As an alternative to fire, solarization effectively kills seeds of most plant species but with a high labor investment per area. We compared the burn to solarization in a study of establishment and growth of seeds and transplants of the native perennial grass Purple needlegrass (Nassella pulchra) and coastal sage species California sagebrush (Artemisia californica). After the fire, initial seed bank and seedling densities and regular percent cover and soil moisture (0–20 cm) data were collected in burned and unburned areas. Burned areas had 96% fewer viable seeds of the dominant annual grass, Ripgut brome (Bromus diandrus), leading to replacement by forbs from the seed bank, especially non‐native Black mustard (Brassica nigra). In the early growing season, B. diandrus dominating unburned areas consistently depleted soil moisture to a greater extent between rains than forbs in burned areas. However, B. diandrus senesced early, leaving more moisture available in unburned areas after late‐season rains. Nassella pulchra and A. californica established better on plots treated with fire and/or solarization than on untreated plots. We conclude that both spring burns and solarization can produce conditions where native perennials can establish in annual grasslands. However, the relative contribution of these treatments to restoration appears to depend on the native species being reintroduced, and the long‐term success of these initial restoration experiments remains to be determined.     

California native and exotic perennial grasses differ in their response to soil nitrogen,exotic annual grass density,and order of emergence

Early emergence of plant seedlings can offer strong competitive advantages over later-germinating neighbors through the preemption of limiting resources. This phenomenon may have contributed to the persistent dominance of European annual grasses over native perennial grasses in California grasslands, since the former species typically germinate earlier in the growing season than the latter and grow rapidly after establishing. Recently, European perennial grasses have been spreading into both non-native annual and native perennial coastal grass stands in California. These exotic perennials appear to be less affected by the priority effects arising from earlier germination by European annual grasses. In addition, these species interactions in California grasslands may be mediated by increasing anthropogenic or natural soil nitrogen inputs. We conducted a greenhouse experiment to test the effects of order of emergence and annual grass seedling density on native and exotic perennial grass seedling performance across different levels of nitrogen availability. We manipulated the order of emergence and density of an exotic annual grass (Bromus diandrus) grown with either Nassella pulchra (native perennial grass), Festuca rubra (native perennial grass), or Holcus lanatus (exotic perennial grass), with and without added nitrogen. Earlier B. diandrus emergence and higher B. diandrus density resulted in greater reduction in the aboveground productivity of the perennial grasses. However, B. diandrus suppressed both native perennials to a greater extent than it did H. lanatus. Nitrogen addition had no effect on the productivity of native perennials, but greatly increased the growth of the exotic perennial H. lanatus, grown with B. diandrus. These results suggest that the order of emergence of exotic annual versus native perennial grass seedlings could play an important role in the continued dominance of exotic annual grasses in California. The expansion of the exotic perennial grass H. lanatus in coastal California may be linked to its higher tolerance of earlier-emerging annual grasses and its ability to access soil resources amidst high densities of annual grasses.     

Shrub succession and invasibility in a New Zealand montane grassland

Abstract Two successive shrub invasions of a short tussock grassland induced by grazing and burning were examined in montane South Island, New Zealand. The first invasion was by a native shrub, matagouri (Discaria toumatou Raoul). The second invasion was by an exotic shrub, Scotch broom (Cytisus scoparius (L.) Link), which invaded the matagouri shrubland that had developed over the grassland. The invasions were investigated using analysis of spatial patterns of both shrubs and tussocks, and age, growth rates and size structure of the shrubs. Competition between the two shrub species was examined using spatial patterns and comparing allometric relationships. After initial invasion by matagouri of the grasslands, stand density increased by consolidation about its initial colonization points. Current matagouri distribution is often negatively associated with tussocks. Scotch broom occurs most frequently in a dense sward of introduced grasses and occasionally in tussocks in interstices among matagouri shrubs. Despite the palatability of Scotch broom to sheep that graze the site, there was no evidence that the spiny matagouri facilitates invasion by protecting Scotch broom seedlings; rather there was negative association between the shrub species. The two species probably compete for above-ground space. However, diameter and height growth rates of Scotch broom far exceed those of matagouri so Scotch broom is likely to increase in biomass rapidly at the site. The autogenic organization and disturbance history of the resident plant communities have rendered each vulnerable to successive invasions.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.     

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.     

Abiotic constraints on the competitive ability of exotic and native grasses in a Pacific Northwest prairie

In prairie ecosystems, abiotic constraints on competition can structure plant communities; however, the extent to which competition between native and exotic plant species is constrained by environmental factors is still debated. The objective of our study was to use paired field and greenhouse experiments to evaluate the competitive dynamics between two native (Danthonia californica and Deschampsia cespitosa) and two exotic (Schedonorus arundinaceus and Lolium multiflorum) grass species under varying nutrient and moisture conditions in an upland prairie in the Willamette Valley, Oregon. We hypothesized the two invasive, exotic grasses would be more competitive under high-nutrient, moderate-moisture conditions, resulting in the displacement of native grasses from these environments. In the field, the experimental reduction of competition resulted in shorter, wider plants, but only the annual grass, Lolium multiflorum, produced more aboveground biomass when competition was reduced. In the greenhouse, the two exotic grasses produced more total biomass than the two native grasses. Competitive hierarchies were influenced by nutrient and/or moisture treatments for the two exotic grasses, but not for the two native grasses. L. multiflorum dominated competitive interactions with all other grasses across treatments. In general, S. arundinaceus dominated when in competition with native grasses, and D. cespitosa produced the most biomass in monoculture or under interspecific competition with the other native grass, D. californica. D. californica, D. cespitosa, and S. arundinaceus all produced more biomass in high-moisture, high-nutrient environments, and D. cespitosa, L. multiflorum, and S. arundinaceus allocated more biomass belowground in the low nutrient treatment. Taken together, these experiments suggest the competitive superiority of the exotic grasses, especially L. multiflorum, but, contrary to our hypothesis, the native grasses were not preferentially excluded from nutrient-rich, moderately wet environments. Laurel Pfeifer-Meister and Esther M. Cole contributed equally to this work.     

Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to mediterranean coastal sage scrub of California

This study examined the growth responses of exotic annuals and native shrubs to elevated N levels to test the hypothesis that increased N availability favors nitrophilous annuals over the slower-growing shrubs. The vegetation structure of the coastal sage scrub ecosystems in southern California is shifting from shrubland to annual grasslands. Over the last 30 years large tracts of wildlands, particularly those adjacent to urban centers, have lost significant native shrub cover, which has been replaced by exotic annuals native to the Mediterranean Basin. During this same time, air pollution has led to increased terrestrial eutrophication by atmospheric deposition. Changes in vegetation are often the result of changes in resource availability. The results of our experiments showed the three native shrubs tested to be more nitrophilous than the three annuals tested, which contrasts with most models of perennial species' adaptation to stressful environments. Under greenhouse conditions the annual grasses exhibited yield depression at the highest N treatments of 80 g g^–1 in soil. The three shrub species evaluated continued to increase shoot biomass at 80 g g^–1 N in soil. The grasses also exhibited increased tissue N concentrations with increased soil N in contrast with the shrubs where there was little difference in tissue N concentrations with increasing availability. Although the differential yield responses to elevated N do not explain the success of the annual vegetation in replacing shrubs, the inability of the shrubs to regulate growth under elevated N levels may explain the poor survival of mature individuals.     

The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

The Influence of Spatial Patterns of Soil Moisture on the Grass and Shrub Responses to a Summer Rainstorm in a Chihuahuan Desert Ecotone

The cycling of surface water, energy, nutrients, and carbon is different between semiarid grassland and shrubland ecosystems. Although differences are evident when grasslands are compared to shrublands, the processes that contribute to this transition are more challenging to document. We evaluate how surface redistribution of precipitation and plant responses to the resulting infiltration patterns could contribute to the changes that occur during the transition from grassland to shrubland. We measured soil water potential under grasses (Bouteloua eriopoda), shrubs (Larrea tridentata) and bare soil and changes in plant water relations and gas exchange following a 15 mm summer storm in the grassland–shrubland ecotone at the Sevilleta National Wildlife Refuge in central New Mexico USA. Following the storm, soil water potential (Ψ_s) increased to 30 cm depth beneath both grass and shrub canopies, with the greatest change observed in the top 15 cm of the soil. The increase in Ψ_s was greater beneath grass canopies than beneath shrub canopies. Ψ_s under bare soil increased only to 5 cm depth. The substantial redistribution of rainfall and different rooting depths of the vegetation resulted in high Ψ_s throughout most of the rooting volume of the grasses whereas soil moisture was unchanged throughout a large portion of the shrub rooting volume. Consistent with this pattern, predawn water potential (Ψ_PD) of grasses increased more than 5 MPa to greater than −1 MPa whereas Ψ_PD of shrubs increased to −2.5 MPa, a change of less than 2 MPa. Transpiration increased roughly linearly with Ψ_PD in both grasses and shrubs. In grasses, assimilation was strongly correlated with Ψ_PD whereas there was no relationship in shrubs where assimilation showed no significant response to the pulse of soil moisture following the storm. These data show that preferential redistribution of water to grass canopies enhances transpiration and assimilation by grasses following large summer storms. This process may inhibit shrubland expansion at the ecotone during periods without extreme drought.     

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.     

Invasive grass density negatively impacts chaparral seedling establishment

Type conversion from native shrubland to invasive annual grassland is on the rise due to global change factors such as prolonged drought and increasing fire frequency. Efforts to restore chaparral ecosystems are limited by current understanding of competitive interactions between shrub seedlings and invasive grasses as well as soil moisture requirements of chaparral seedlings. We set up a restoration experiment where we out-planted Adenostoma fasciculatum seedlings, manipulated invasive grass density, monitored soil moisture at two depths, and tracked seedling survival and biomass. We found that higher invasive grass cover was associated with higher rates of seedling mortality but found no difference in biomass per surviving plant. Soil moisture was higher at 15 cm under the 100% weeded treatment than the 50% weeded and control treatments during January. Lower invasive cover resulted in higher richness of annual native plant species, as plots with 100% invasive removal had higher richness than 50% removal and unplanted control plots. Future restoration efforts in the chaparral will likely be more successful in increasing initial seedling establishment if invasive grass removal is included.     

Shrubs as ecosystem engineers in a coastal dune: influences on plant populations,communities and ecosystems

Question: How do two shrubs with contrasting life‐history characteristics influence abundance of dominant plant taxa, species richness and aboveground biomass of grasses and forbs, litter accumulation, nitrogen pools and mineralization rates? How are these shrubs – and thus their effects on populations, communities and ecosystems – distributed spatially across the landscape? Location: Coastal hind‐dune system, Bodega Head, northern California. Methods: In each of 4 years, we compared vegetation, leaf litter and soil nitrogen under canopies of two native shrubs –Ericameria ericoides and the nitrogen‐fixing Lupinus chamissonis– with those in adjacent open dunes. Results: At the population level, density and cover of the native forb Claytonia perfoliata and the exotic grass Bromus diandrus were higher under shrubs than in shrub‐free areas, whereas they were lower under shrubs for the exotic grass Vulpia bromoides. In contrast, cover of three native moss species was highest under Ericameria and equally low under Lupinus and shrub‐free areas. At community level, species richness and aboveground biomass of herbaceous dicots was lower beneath shrubs, whereas no pattern emerged for grasses. At ecosystem level, areas beneath shrubs accumulated more leaf litter and had larger pools of soil ammonium and nitrate. Rates of nitrate mineralization were higher under Lupinus, followed by Ericameria and then open dune. At landscape level, the two shrubs – and their distinctive vegetation and soils – frequently had uniform spatial distributions, and the distance separating neighbouring shrubs increased as their combined sizes increased. Conclusions: Collectively, these data suggest that both shrubs serve as ecosystem engineers in this coastal dune, having influences at multiple levels of biological organization. Our data also suggest that intraspecific competition influenced the spatial distributions of these shrubs and thus altered the distribution of their effects throughout the landscape.     

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.     

Mid-Spring Burning Reduces Spotted Knapweed and Increases Native Grasses during a Michigan Experimental Grassland Establishment

Infestations of the exotic perennial Spotted knapweed (Centaurea maculosa Lam.) hinder the restoration and management of native ecosystems on droughty, infertile sites throughout the Midwestern United States. We studied the effects of annual burning on knapweed persistence on degraded, knapweed‐infested gravel mine spoils in western Michigan. Our experiment included 48, 4‐m² plots seeded to native warm‐season grasses in 1999 using a factorial arrangement of initial herbicide and fertility treatments. Beginning in 2003, we incorporated fire as an additional factor and burned half of the plots in late April or May for 3 years (2003–2005). Burning increased the dominance of warm‐season grasses and decreased both biomass and dominance of knapweed in most years. Burning reduced adult knapweed densities in all 3 years of the study, reduced seedling densities in the first 2 years, and reduced juvenile densities in the last 2 years. Knapweed density and biomass also declined on the unburned plots through time, suggesting that warm‐season grasses may effectively compete with knapweed even in the absence of fire. By the end of the study, mean adult knapweed densities on both burned (0.4‐m²) and unburned (1.3‐m²) plots were reduced to levels where the seeded grasses should persist with normal management, including the use of prescribed fire. These results support the use of carefully timed burning to help establish and maintain fire‐adapted native plant communities on knapweed‐infested sites in the Midwest by substantially reducing knapweed density, biomass, and seedling recruitment and by further shifting the competitive balance toward native warm‐season grasses.     

Nitrogen limitation, 15N tracer retention, and growth response in intact and Bromus tectorum-invaded Artemisia tridentata ssp. wyomingensis communities

Annual grass invasion into shrub-dominated ecosystems is associated with changes in nutrient cycling that may alter nitrogen (N) limitation and retention. Carbon (C) applications that reduce plant-available N have been suggested to give native perennial vegetation a competitive advantage over exotic annual grasses, but plant community and N retention responses to C addition remain poorly understood in these ecosystems. The main objectives of this study were to (1) evaluate the degree of N limitation of plant biomass in intact versus B. tectorum-invaded sagebrush communities, (2) determine if plant N limitation patterns are reflected in the strength of tracer ¹⁵N retention over two growing seasons, and (3) assess if the strength of plant N limitation predicts the efficacy of carbon additions intended to reduce soil N availability and plant growth. Labile C additions reduced biomass of exotic annual species; however, growth of native A. tridentata shrubs also declined. Exotic annual and native perennial plant communities had divergent responses to added N, with B. tectorum displaying greater ability to use added N to rapidly increase aboveground biomass, and native perennials increasing their tissue N concentration but showing little growth response. Few differences in N pools between the annual and native communities were detected. In contrast to expectations, however, more ¹⁵N was retained over two growing seasons in the invaded annual grass than in the native shrub community. Our data suggest that N cycling in converted exotic annual grasslands of the northern Intermountain West, USA, may retain N more strongly than previously thought.     

Disturbance: a double-edged sword for restoration in a changing climate

Ecological restoration often relies on disturbance as a tool for establishing target plant communities, but disturbance can be a double-edged sword, at times initiating invasion and unintended outcomes. Here we test how fire disturbance, designed to enhance restoration seeding success, combines with climate and initial vegetation conditions to shift perennial versus annual grass dominance and overall community diversity in Pacific Northwest grasslands. We seeded both native and introduced perennial grasses and native forbs in paired, replicated burned-unburned plots in three sites along a latitudinal climate gradient from southern Oregon to central-western Washington. Past restoration and climate manipulations at each site had increased the variation of starting conditions between plots. Burning promoted the expansion of extant forbs and perennial grasses across all sites. Burning also enhanced the seeding success of native perennial grass and native forbs at the northern and central site, and the success of introduced perennial grasses across all three sites. Annual grass dominance was driven more by latitude than burning, with annuals maintaining their dominance in the south and perennials in the north. At the same time, unrestored grasslands surrounding all sites remained dominated by perennial grasses, suggesting that initial plot clearing may have allowed for annual grass invasion in the southern site. When paired with disturbance, further warming may increase the risk of annual grass dominance, a potentially persistent state.     

The response of native species to removal of invasive exotic grasses in a seasonally dry Hawaiian woodland

Abstract. Non-native perennial grasses form 30% of the live understory biomass in seasonally dry, submontane forests in Hawaii Volcanoes National Park, yet their effects on native species are unknown. We removed these grasses from plots of 20 m × 20 m in 1991 and maintained removal and control areas over the next three years. Two fast growing shrub species, Dodonaea viscosa and Osteomeles anthylidifolia, increased in size significantly more in removal areas than in controls. Individuals of the most abundant shrub species, Styphelia tameiameia showed no net growth response to grass removal. They did, however, change their architecture: many branches along the mid and upper sections of the main trunk died and a proliferation of new leaves and shoots occurred in the lower 40 cm of trunk. Basal diameter increase was very small in Metrosideros polymorpha, the dominant tree species in these sites. All species except Styphelia had significantly increased leaf tissue nitrogen in removal plots by 18 months after removal when compared to shrubs in control areas suggesting that removal plot shrubs had greater access to soil nitrogen. Available soil-N pools, which were generally higher in the removal plots, support this interpretation. Light levels near the soil surface were also higher where grasses were removed than where they were present which may have contributed to increased shrub growth. By contrast, soil moisture was consistently lower where grasses were removed than where they were still present. Shrub tissue carbon isotope values were consistent with the interpretation that shrubs in removal plots had less rather than more water available to them. Hence, the increased growth observed in removal plot shrubs could not be due to release from moisture competition. Lastly, our results showed that seedlings of all woody species except Metrosideros were significantly more abundant in removal plots at both one and three years after removal and initially high sapling mortality was balanced by high recruitment into the sapling class. We believe that over time this will result in increased densities of native shrubs if grasses are kept out. With the presence of grasses, shrub growth in these woodlands is reduced and biomass is shifting towards grasses.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Semiarid grassland responses to short-term variation in water availability

Standing crop and species composition in semiarid grassland are linked to long-term patterns of water availability, but grasslands are characterized by large single-season variability in rainfall. We tested whether a single season of altered water availability influenced the proportions of grasses and shrubs in a semiarid grassland near the northern edge of the North American Great Plains. We studied stands of the clonal shrub snowberry (Symphoricarpos occidentalis) and adjacent grassland dominated by the native grasses Stipa spartea and Bouteloua gracilis. Rain was excluded and water supplied in amounts corresponding to years of low, medium, and high rainfall, producing a 2 − 4-fold range in monthly precipitation among water supply treatments. There were ten replicate plots of each water treatment in both snowberry stands and grassland. Grass standing crop increased significantly with water availability in grassland but not inside snowberry stands. Total standing crop and shrub stem density increased significantly with water supply, averaged across both communities. In contrast, water had no effect on shrub standing crop or light penetration. In summary, our finding that water has significant effects on a subset of components of grassland vegetation is consistent with long-term, correlational studies, but we also found that a single season of altered water supply had no effect on other important aspects of the ecosystem. This revised version was published online in June 2006 with corrections to the Cover Date.