The Mark of Zorro: Effects of the Exotic Annual Grass Vulpia myuros on California Native Perennial Grasses

Native perennial grasses were once common in California prairies that are now dominated by annual grasses introduced from Europe. Competition from exotics may be a principal impediment to reestablishment of native perennial grasses. Introduced annual grasses, such as Vulpia myuros (zorro fescue), are often included with native perennial species in revegetation seed mixtures used in California. To examine the potential suppressive effect of this graminoid, we evaluated the growth and performance of a mixture of California native perennial grasses and resident weeds when grown with varying densities of V. myuros. The annual fescue exhibited a strongly plastic growth response to plant density, producing similar amounts of above‐ground biomass at all seeding densities. Perennial grass seedling survival and above‐ ground biomass decreased and individuals became thinner (i.e., reduced weight‐to‐height ratio) with increasing V. myuros seeding density. V. myuros also significantly suppressed above‐ground biomass and densities of weeds and had a more negative effect on weed densities than on native perennial grass densities. Biomass of native grasses and weeds was not differentially affected by increasing densities of V. myuros. Overall, because V. myuros significantly reduced the survival and performance of the mixture of native perennial grasses and this effect increased with increasing V. myuros density, we conclude that including this exotic annual in native seed mixtures is counterproductive to restoration efforts.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

Direct and indirect effects of exotic annual grasses on species composition of a South Australian grassland

Abstract Invasion by Mediterranean annual grasses, such as Avena L. spp. and Bronms L. spp, is one of the major threats to temperate perennial grassland. This study investigated the effects of annual grasses and their litter on the species composition of a grassland near Burra, South Australia. The placement of annual grass litter on soil samples in the glasshouse decreased the establishment or growth of several exotic annual dicots. In the field the addition of annual grass litter slightly decreased the frequency of Danthonia Lam. & DC. tussocks. Furthermore, litter strongly reduced the species richness from 13 species in plots with no litter to nine species in plots with the highest litter level, mainly by decreasing the frequency of common exotic dicots. Native dicot frequency similarly appeared to be decreased by litter addition. In addition to the negative effects of their litter, annual grasses also directly competed with perennial grasses. The magnitude of the competitive effect varied systematically along a slope, suggesting that other factors such as soil properties may control competitive inter actions. The biomass of annual grasses also tended to increase with the addition of their own litter. This combination of positive and negative feedback mechanisms suggests that brief periods favourable for annual grasses, either through management changes or environmental conditions, can lead to persistent changes in the species composition of the system.     

Native cover crops suppress exotic annuals and favor native perennials in a greenhouse competition experiment

In a greenhouse experiment, we examined the effectiveness of four native cover crops for controlling four exotic, invasive species and increasing success of four western North American grassland species. Planting the annual cover crops, annual ragweed (Ambrosia artemisiifolia) and common sunflower (Helianthus annuus), reduced the biomass of the exotic species cheatgrass (Bromus tectorum), Japanese brome (Bromus japonicus), Canada thistle (Cirsium arvense), and whitetop (Cardaria draba). The annual cover crops also reduced the desired species biomass in competition with the perennial exotics, but either increased or did not affect the desired species biomass in competition with the annual exotics. Planting the perennial cover crops, Canada goldenrod (Solidago canadensis) and littleleaf pussytoes (Antennaria microphylla), rarely inhibited exotic species, but did increase the desired species biomass. Field experiments are needed to test the cover crops under more ecologically relevant conditions, but our results suggested that the annual cover crops may be effective for controlling invasive annuals and for facilitating native perennials.     

Effects of resource availability and propagule supply on native species recruitment in sagebrush ecosystems invaded by Bromus tectorum

Resource availability and propagule supply are major factors influencing establishment and persistence of both native and invasive species. Increased soil nitrogen (N) availability and high propagule inputs contribute to the ability of annual invasive grasses to dominate disturbed ecosystems. Nitrogen reduction through carbon (C) additions can potentially immobilize soil N and reduce the competitiveness of annual invasive grasses. Native perennial species are more tolerant of resource limiting conditions and may benefit if N reduction decreases the competitive advantage of annual invaders and if sufficient propagules are available for their establishment. Bromus tectorum, an exotic annual grass in the sagebrush steppe of western North America, is rapidly displacing native plant species and causing widespread changes in ecosystem processes. We tested whether nitrogen reduction would negatively affect B. tectorum while creating an opportunity for establishment of native perennial species. A C source, sucrose, was added to the soil, and then plots were seeded with different densities of both B. tectorum (0, 150, 300, 600, and 1,200 viable seeds m⁻²) and native species (0, 150, 300, and 600 viable seeds m⁻²). Adding sucrose had short-term (1 year) negative effects on available nitrogen and B. tectorum density, biomass and seed numbers, but did not increase establishment of native species. Increasing propagule availability increased both B. tectorum and native species establishment. Effects of B. tectorum on native species were density dependent and native establishment increased as B. tectorum propagule availability decreased. Survival of native seedlings was low indicating that recruitment is governed by the seedling stage.     

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.     

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.     

Control of Exotic Annual Grasses to Restore Native Forbs in Abandoned Agricultural Land

Exotic annual grasses are a major challenge to successful restoration in temperate and Mediterranean climates. Experiments to restore abandoned agricultural fields from exotic grassland to coastal sage scrub habitat were conducted over two years in southern California, U.S.A. Grass control methods were tested in 5 m² plots using soil and vegetation treatments seeded with a mix of natives. The treatments compared grass‐specific herbicide, mowing, and black plastic winter solarization with disking and a control. In year two, herbicide and mowing treatments were repeated on the first‐year plots, plus new control and solarization plots were added. Treatments were evaluated using percent cover, richness and biomass of native and exotic plants. Disking alone reduced exotic grasses, but solarization was the most effective control in both years even without soil sterilization, and produced the highest cover of natives. Native richness was greatest in solarization and herbicide plots. Herbicide application reduced exotics and increased natives more than disking or mowing, but produced higher exotic forb biomass than solarization in the second year. Mowing reduced grass biomass and cover in both years, but did not improve native establishment more than disking. Solarization was the most effective restoration method, but grass‐specific herbicide may be a valuable addition or alternative. Solarization using black plastic could improve restoration in regions with cool, wet summers or winter growing seasons by managing exotic seedbanks prior to seeding. While solarization may be impractical at very large scales, it will be useful for rapid establishment of annual assemblages on small scales.     

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.     

Can Carbon Addition Increase Competitiveness of Native Grasses? A Case Study from California

There is growing interest in the addition of carbon (C) as sucrose or sawdust to the soil as a tool to reduce plant‐available nitrogen (N) and alter competitive interactions among species. The hypothesis that C addition changes N availability and thereby changes competitive dynamics between natives and exotics was tested in a California grassland that had experienced N enrichment. Sawdust (1.2 kg/m) was added to plots containing various combinations of three native perennial bunchgrasses, exotic perennial grasses, and exotic annual grasses. Sawdust addition resulted in higher microbial biomass N, lower rates of net N mineralization and net nitrification, and higher concentrations of extractable soil ammonium in the soil. In the first year sawdust addition decreased the degree to which exotic annuals competitively suppressed the seedlings of Nassella pulchra and, to a lesser extent, Festuca rubra, both native grasses. However there was no evidence of reduced growth of exotic grasses in sawdust‐amended plots. Sawdust addition did not influence interactions between the natives and exotic perennial grasses. In the second year, however, sawdust addition did not affect the interactions between the natives and either group of exotic grasses. In fact, the native perennial grasses that survived the first year of competition with annual grasses significantly reduced the aboveground productivity of annual grasses even without sawdust addition. These results suggest that the addition of sawdust as a tool in the restoration of native species in our system provided no significant benefit to natives over a 2‐year period.     

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.     

Optimal prescribed burn frequency to manage foundation California perennial grass species and enhance native flora

Grasslands can be diverse assemblages of grasses and forbs but not much is known how perennial grass species management affects native plant diversity except in a few instances. We studied the use of late-spring prescribed burns over a span of 11 years where the perennial grass Poa secunda was the foundation species, with four additional years of measurements after the final burn. We evaluated burn effects on P. secunda, the rare native annual forb Amsinckia grandiflora and local native and exotic species. Annual burning maintained P. secunda number, resulted in significant expansion, the lowest thatch and exotic grass cover, the highest percentage of bare ground, but also the lowest native forb and highest exotic forb cover. Burning approximately every 3 years maintained a lower number of P. secunda plants, allowed for expansion, and resulted in the highest native forb cover with a low exotic grass cover. Burning approximately every 5 years and the control (burned once from a wildfire) resulted in a decline in P. secunda number, the highest exotic grass and thatch cover and the lowest percentage of bare ground. P. secunda numbers were maintained up to 4 years after the final burn. While local native forbs benefited from burning approximately every 3 years, planted A. grandiflora performed best in the control treatment. A. grandiflora did not occur naturally at the site; therefore, no seed bank was present to provide across-year protection from the effects of the burns. Thus, perennial grass species management must also consider other native species life history and phenology to enhance native flora diversity.     

Soil Biological and Chemical Properties in Restored Perennial Grassland in California

Restoration of California native perennial grassland is often initiated with cultivation to reduce the density and cover of non‐native annual grasses before seeding with native perennials. Tillage is known to adversely impact agriculturally cultivated land; thus changes in soil biological functions, as indicated by carbon (C) turnover and C retention, may also be negatively affected by these restoration techniques. We investigated a restored perennial grassland in the fourth year after planting Nassella pulchra, Elymus glaucus, and Hordeum brachyantherum ssp. californicum for total soil C and nitrogen (N), microbial biomass C, microbial respiration, CO₂ concentrations in the soil atmosphere, surface efflux of CO₂, and root distribution (0‐ to 15‐, 15‐ to 30‐, 30‐ to 60‐, and 60‐ to 80‐cm depths). A comparison was made between untreated annual grassland and plots without plant cover still maintained by tillage and herbicide. In the uppermost layer (0‐ to 15‐cm depth), total C, microbial biomass C, and respiration were lower in the tilled, bare soil than in the grassland soils, as was CO₂ efflux from the soil surface. Root length near perennial bunchgrasses was lower at the surface and greater at lower depths than in the annual grass–dominated areas; a similar but less pronounced trend was observed for root biomass. Few differences in soil biological or chemical properties occurred below 15‐cm depth, except that at lower depths, the CO₂ concentration in the soil atmosphere was lower in the plots without vegetation, possibly from reduced production of CO₂ due to the lack of root respiration. Similar microbiological properties in soil layers below 15‐cm depth suggest that deeper microbiota rely on more recalcitrant C sources and are less affected by plant removal than in the surface layer, even after 6 years. Without primary production, restoration procedures with extended periods of tillage and herbicide applications led to net losses of C during the plant‐free periods. However, at 4 years after planting native grasses, soil microbial biomass and activity were nearly the same as the former conditions represented by annual grassland, suggesting high resilience to the temporary disturbance caused by tillage.     

Disturbance,drought and dynamics of desert dune grassland,South Africa

A seven-year study of marked plants and plots in Stipagrostis ciliata (Desf.) de Winter dune grassland, in the arid (<100 mm yr^–1) Bushmanland area of the Northern Cape province of South Africa, was designed to test the hypothesis that establishment of ephemeral plants, and recruitment of perennial grasses was dependent upon disturbances that reduced the density of living perennial grass tussocks. In 1989, eight 4 m² plots were cleared of perennial vegetation by uprooting and removing all plants so as to resemble small-scale disturbances made by burrowing mammals or territorial antelope. The vegetation on the cleared plots and surroundings was monitored until 1996. Initial results supported our hypothesis. In wet years, when ephemeral plants were abundant, their average fresh mass was 2–3 times greater per unit area on the cleared plots than in control plots in adjacent, undisturbed grassland. Many Stipagrostis seedlings established in the cleared plots over the two years following clearing but were rare in adjacent areas among established conspecifics. However, a drought in 1992 (11 mm of rain over 12 months) lead to widespread mortality of the perennial grass, killing 56% (range 22–79%) of established tufts. High densities of Stipagrostis seedlings appeared following the drought-breaking rains in January 1993, both in the disturbed plots and in the surrounding `undisturbed' dune grassland. Ephemeral plants established in large numbers throughout the area during the high rainfall year of 1996 and were generally more numerous in the old disturbances than in control plots. Seven years after clearing the biomass of grass on the cleared plots was approximately 34% of the mass removed from the plots in 1989 whereas in the undisturbed grassland biomass was 66% of 1989 levels. Drought had little long-term effect on community composition, and Stipagrostis ciliata constituted 94–98% of plant community before and after drought. Cleared plots were recolonised by S. ciliata, but the contribution of other grass species increased by 6–9%. Synchronous recruitment following occasional drought-induced mortality can generate even-aged populations of the dominant desert dune 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.     

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 Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

Variation in the establishment of a non-native annual grass influences competitive interactions with Mojave Desert perennials

Competition between native and non-native species can change the composition and structure of plant communities, but in deserts, the highly variable timing of resource availability also influences non-native plant establishment, thus modulating their impacts on native species. In a field experiment, we varied densities of the non-native annual grass Bromus madritensis ssp. rubens around individuals of three native Mojave Desert perennials—Larrea tridentata, Achnatherum hymenoides, and Pleuraphis rigida—in either winter or spring. For comparison, additional plots were prepared for the same perennial species and seasons, but with a mixture of native annual species as neighbors. Growth of perennials declined when Bromus was established in winter because Bromus stands had 2–3 months of growth and high water use before perennial growth began. However, water potentials for the perennials were not significantly reduced, suggesting that direct competition for water may not be the major mechanism driving reduced perennial growth. The impact of Bromus on Larrea was lower than for the two perennial grasses, likely because Larrea maintains low growth rates throughout the year, even after Bromus has completed its life cycle. This result contrasts with the perennial grasses, whose phenology completely overlaps with (Achnatherum) or closely follows (Pleuraphis) that of Bromus. In comparison, Bromus plants established in spring were smaller than those established in winter and thus did not effectively reduce growth of the perennials. Growth of perennials with mixed annuals as neighbors also did not differ from those with Bromus neighbors of equivalent biomass, but stands of these native annuals did not achieve the high biomass of Bromus stands that were necessary to reduce perennial growth. Seed dormancy and narrow requirements for seedling survivorship of native annuals produce densities and biomass lower than those achieved by Bromus; thus, impacts of native Mojave Desert annuals on perennials are expected to be lower than those of Bromus.     

Managing contingency in semiarid grassland restoration through repeated planting

The success of grassland restoration in semiarid regions is contingent on good establishment years. Here it is asked whether success rate can be increased by planting repeatedly among years, or by consistently using high planting densities. I planted seeds of a dominant native perennial grass, Elymus lanceolatus, at five densities (range: 30–3000 seeds/m²) in each of 3 years. Elymus seedling emergence and cover increased significantly with planting density. Cover after three growing seasons was maximized (i.e. not significantly different from the greatest cover) at planting densities of 300–600 seeds/m². On the other hand, low germination in a dry warm year resulted in density having no effect on cover 3 years later. Two favorable planting years showed trajectories to native dominance after three growing seasons. In contrast, one unfavorable planting year showed a trajectory of non‐native dominance. Emergence was increased modestly by both herbivore and interspecific neighbor removal. Overall, the most economical process for ensuring success (assuming that seed costs are high and planting and site‐preparation costs are low) may be to plant at moderate density (300–600 seeds/m²) in repeated years until a favorable year is encountered.     

Exotic Grass Competition in Suppressing Native Shrubland Re-establishment

Disturbance of coastal sage scrub in southern California has led to extensive displacement of native shrubs by exotic annual grasses. The initial conversion from shrubland to exotic grassland is typically associated with disturbance caused by intense grazing, high fire frequency, or mechanical vegetation removal. While native shrubs have been shown to recolonize annual grasslands under some conditions, other annual grasslands are persistent and show no evidence of shrub recolonization. This study examined the mechanisms by which annual grasses may exclude native shrubs and persist after release from disturbance. Grass density was manipulated in experimental plots to achieve a series of prescribed densities. Artemisia californica, a dominant native shrub, was seeded or planted into the plots and responses to the grass density treatments were measured over two growing seasons. A. californica germination, first season growth, and survival were all negatively related to the density of neighboring annual grasses. The most probable mechanism underlying the reduction of first season growth and survival was depletion of soil water by the grasses. The effects of the grasses on A. californica were no longer significant in the second season. The results of this study indicate that Mediterranean annual grasses reduce recruitment and can persist by inhibiting post-disturbance establishment of A. californica from seed. Although succession alone may not return disturbed annual grasslands to their former shrubland composition, the results suggest that restoration can be achieved by using container plantings or grass removal followed by seeding.