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.     

Reintroduction of Nassella pulchra to California coastal grasslands: Effects of topsoil removal,plant neighbour removal and grazing

Question: What is the most appropriate combination of treatments to reintroduce Nassella pulchra, a perennial bunchgrass, into degraded mediterranean coastal grasslands? Location: Central coast of California, USA. Methods: N. pulchra was sown from seeds and transplanted into a degraded grassland in a multi‐factorial experiment testing the effects of (1) two grazing intensities (lightly grazed by native mammal species or ungrazed); (2) topsoil removal and (3) reduction of plant neighbours. The experiment was carried out on two types of surrounding vegetation (exotic annual grasses and exotic forbs). Results: Topsoil removal greatly enhanced establishment from seeds and transplant survival, mainly because it reduced the exotic vegetation and thus reduced competition. While removing neighbours was essential when topsoil was left intact, it had a negative effect on N. pulchra when surrounding species included exotic forbs (Brassica spec, and Asteraceae) at low density (after topsoil removal). Moderate grazing by native mammals (deer, rabbits and gophers) did not affect N. pulchra. Conclusion: Our results suggest that seeding after topsoil has been removed is a promising method to reintroduce N. pulchra to highly degraded sites where there is little to no native seed bank.     

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.     

Mechanisms of influence of invasive grass litter on germination and growth of coexisting species in California

In grasslands, litter has been recognized as an important factor promoting grass persistence and the suppression of forbs. The invasive European annual grass Bromus diandrus (ripgut brome) is widespread throughout California, where it produces a persistent and thick litter layer. The native grass, Stipa pulchra, is also common in some grassland settings and can also produce persistent litter, yet it is typically associated with more forbs. Very little is known about the mechanisms through which these two common grass species influence seedling establishment of both exotic invasive and native herbs. Here, we evaluated the effect of B. diandrus and S. pulchra litter on seedling establishment of two invasive (the grass B. diandrus and the forb Centaurea melitensis) and two native (the grass S. pulchra, and the forb Clarkia purpurea) herbaceous plants in a greenhouse setting. Our results showed that B. diandrus litter cover hindered seedling establishment of the four species tested, but that the degree and mechanism of inhibition was dependent on which species was tested, life form (e.g. monocot/dicot) and seed size. Seedling emergence of the two forb species was more vulnerable to litter cover than either grass species and both forbs had smaller seed size. After germination, only seedling biomass of B. diandrus itself was reduced by litter (both B. diandrus and S. pulchra). We found no significant effects of leachate of either grass species on seedling emergence of any species, while a high concentration of B. diandrus leachates inhibited root growth of all species including B. diandrus seedlings. Stipa pulchra litter leachates did not affect S. pulchra or C. melitensis seedlings although it did suppress B. diandrus and C. purpurea seedling growth. Our findings provide direct experimental evidence for the mechanism of effect of litter on these coexisting invasive and native species. Such evidence helps advance our understanding of role of B. diandrus and S. pulchra litter in California grassland.     

The Role of Light and Soil Moisture in Plant Community Resistance to Invasion by Yellow Starthistle (Centaurea solstitialis)

To resist establishment by an invasive plant, a community may require one or more species functionally similar to the invader in their resource acquisition pattern. In this study, communities consisting of native winter annual forbs, non‐native annual grasses, native perennials, or a combination of the two native communities were established with and without Centaurea solstitialis to determine the effect of soil moisture and light availability on plant community invasion resistance. The annual plant communities were unable to resist invasion by C. solstitialis. In the native winter annual forb community, senescence in late spring increased light penetration (>75%) to the soil surface, allowing seeded C. solstitialis to quickly establish and dominate the plots. In addition, native annual forbs utilized only shallow soil moisture, whereas C. solstitialis used shallow and deep soil moisture. In communities containing native perennials, only Elymus glaucus established well and eventually dominated the plots. During the first 2 years of establishment, water use pattern of perennial communities was similar to native annual forbs and resistance to invasion was associated with reduced light availability during the critical stages of C. solstitialis establishment. In later years, however, water use pattern of perennial grass communities was similar or greater than C. solstitialis‐dominated plots. These results show that Central Valley grasslands that include E. glaucus resist C. solstitialis invasion by a combination of light suppression and soil water competition. Spatiotemporal resource utilization patterns, and not just functional similarity, should be considered when developing restoration strategies to resist invasion by many non‐native species.     

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.     

Mortality of exotic and native seeds in invaded and uninvaded habitats

We examined seed survival in exotic- and native-dominated grasslands by placing seeds of a once-pervasive native grass species, Nassella pulchra, and two of the most common, widespread exotic grass species, Avena fatua and Bromus hordeaceus, in mesh bags in the field for 3 months. Compared to germination of unexposed seeds not placed in the field, exotic species experienced an approximately 40% reduction in viability, whereas the mortality experienced by the native species was <20%. Despite these differences, germination rates of exposed seeds were similar between native and exotic species because native N. pulchra seeds had lower initial viability prior to entering the seed bank. Seed mortality did not differ based on whether seeds were placed in habitats dominated by exotic or native grasses. Rather, our results suggest that re-establishment of native N. pulchra must focus on maximizing seed viability and survival, and that A. fatua and B. hordeaceus overcome relatively higher losses of viable seeds in the seed bank, potentially by producing large numbers of highly viable seeds.     

Burning and Grazing Management in a California Grassland: Effect on Bunchgrass Seed Viability

Prescribed fire is an important management tool for reducing the dominance of non‐native species in annual grasslands; both annual and perennial native species show strong vegetative responses in the subsequent growing season. However, although the post‐fire contribution of native species to the seed bank is assumed to be larger than in pretreatment years, the effects on seed quality, particularly viability and longevity, are not well understood. In this study, I germinated Nassella pulchra (purple needlegrass) seed that had been stored for 10 years after collection from target plants receiving treatment combinations of summer burning and grazing by sheep. Seeds from burned plants were larger and had higher germinability than seed from unburned plants. Seeds from plants that were both burned and grazed had the highest germination. The strong relationship between long‐term viability and seed size suggests greater maternal provisioning and increased seed quality subsequent to burning and grazing. I conclude that managing for seed quality may be a useful approach for conservation of native species in California's critically endangered grassland habitats.     

Postwildfire seeding to restore native vegetation and limit exotic annuals: an evaluation in juniper‐dominated sagebrush steppe

Reestablishment of perennial vegetation is often needed after wildfires to limit exotic species and restore ecosystem services. However, there is a growing body of evidence that questions if seeding after wildfires increases perennial vegetation and reduces exotic plants. The concern that seeding may not meet restoration goals is even more prevalent when native perennial vegetation is seeded after fire. We evaluated vegetation cover and density responses to broadcast seeding native perennial grasses and mountain big sagebrush (Artemisia tridentata Nutt. spp. vaseyana [Rydb.] Beetle) after wildfires in the western United States in six juniper (Juniperus occidentalis ssp. occidentalis Hook)‐dominated mountain big sagebrush communities for 3 years postfire. Seeding native perennial species compared to not seeding increased perennial grass and sagebrush cover and density. Perennial grass cover was 4.3 times greater in seeded compared to nonseeded areas. Sagebrush cover averaged 24 and less than 0.1% in seeded and nonseeded areas at the conclusion of the study, respectively. Seeding perennial species reduced exotic annual grass and annual forb cover and density. Exotic annual grass cover was 8.6 times greater in nonseeded compared to seeded areas 3 years postfire. Exotic annual grass cover increased over time in nonseeded areas but decreased in seeded areas by the third‐year postfire. Seeded areas were perennial‐dominated and nonseeded areas were annual‐dominated at the end of the study. Establishing perennial vegetation may be critical after wildfires in juniper‐dominated sagebrush steppe to prevent the development of annual‐dominated communities. Postwildfire seeding increased perennial vegetation and reduced exotic plants and justifies its use.     

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.     

Radial Dispersion of Neighbors and the Small-Scale Competitive Impact of Two Annual Grasses on a Native Perennial Grass

In California's Mediterranean type grasslands, native perennial grasses such as Nassella pulchra are surrounded by introduced annual species and these annuals are thought to have displaced natives through much of their range. Amongst other invaders, two grasses Lolium multiflorum and Bromus hordeaceus, commonly dominate portions of the grassland with potential for N. pulchra restoration. We hypothesized that competitor species differences and small‐scale gaps (150 cm²) could be important determinants of N. pulchra survival and performance on these sites. Lolium multiflorum and B. hordeaceus were planted in 20 cm diameter circular plots at a constant rate of 1 seed per cm² surrounding newly transplanted N. pulchra plants. Nassella pulchra showed no significant effect of the species of competitor or from the distribution of the competitors. Both interspersion of patches of bare ground and separation of competitors into patches did not increase N. pulchra pre‐dawn water potential, basal area change, number of seeds produced, or average weight of seeds. The presence of L. multiflorum was associated with a decrease in N. pulchra survival compared with plots with only B. hordeaceus. Plants with increases in basal area of less than 0.75 cm² during the growing season had 74% mortality compared with no mortality in plants with more growth. However, initial N. pulchra plant size was not a good predictor of mortality. Limiting competition from annuals may increase survival of N. pulchra plantings, but 60% of the plants survived for at least 1 year, despite being transplanted into soil containing substantial annual grass seed.     

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.     

Preferences for 15N-ammonium, 15N-nitrate,and 15N-glycine differ among dominant exotic and subordinate native grasses from a California oak woodland

Following an invasion of exotic annual grasses into California oak woodlands, grass species dominance shifted from native perennials to exotic annuals. In combination with other ecosystem and species characteristics, species-specific N preferences may influence species coexistence and dominance. If species N preferences follow dominance patterns in California oak woodlands, then the more dominant exotic grasses should prefer the most abundant inorganic soil N form (NH₄⁺), while the subordinate native grasses should prefer the less available inorganic (NO₃^?) or organic (glycine) soil N forms. To investigate this prediction, we applied ¹⁵N-labeled NH₄⁺, NO₃^?, and glycine to soil and measured % ¹⁵N recovery by two dominant annual grasses (Bromus diandrus and Bromus hordeaceus) and two subordinate perennial grasses (Elymus glaucus and Nassella pulchra). As expected, shoots of B. diandrus recovered more ¹⁵N-NH₄⁺ (74%) than either ¹⁵N-NO₃^? (51%) or ¹⁵N-glycine (39%). B. diandrus also captured at least 3.2 times more ¹⁵N-NH₄⁺ than subordinate grasses. Dominant B. hordeaceus, however, demonstrated no N form preferences. As hypothesized, shoots of subordinate E. glaucus and N. pulchra recovered 2.1–2.3 times more ¹⁵N-NO₃^? than ¹⁵N-NH₄⁺ and increased %N by 4.8–5.7% in response to the application of ¹⁵N-NO₃^?. Both subordinate grasses did not prefer ¹⁵N-glycine over ¹⁵N-NH₄⁺, suggesting that the importance of this N form in structuring species coexistence in California oak woodlands is minimal. These results support our theory that species N preferences follow dominance patterns in California oak woodlands. To further understand the role of these species-specific N preferences in structuring dominance, the importance of N form versus such characteristics as rooting distribution and species phenologies needs to be explored in the presence of interspecific competition.     

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.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

Variation in the impact of exotic grasses on native plant composition in relation to fire across an elevation gradient in Hawaii

The impact that an exotic species can have on the composition of the community it enters is a function of its abundance, its particular species traits and characteristics of the recipient community. In this study we examined species composition in 14 sites burned in fires fuelled by non‐indigenous C4 grasses in Hawaii Volcanoes National Park, Hawaii. We considered fire intensity, time since fire, climatic zone of site, unburned grass cover, unburned native cover and identity of the most abundant exotic grass in the adjacent unburned site as potential predictor variables of the impact of fire upon native species. We found that climatic zone was the single best variable for explaining variation in native cover among burned sites and between burned and unburned pairs. Fire in the eastern coastal lowlands had a very small effect on native plant cover and often stimulated native species regeneration, whereas fire in the seasonal submontane zone consistently caused a decline in native species cover and almost no species were fire tolerant. The dominant shrub, Styphelia tameiameia, in particular was fire intolerant. The number of years since fire, fire intensity and native cover in reference sites were not significantly correlated with native species cover in burned sites. The particular species of grass that carried the fire did however, have a significant effect on native species recovery. Where the African grass Melinis minutiflora was a dominant or codominant species, fire impacts were more severe than where it was absent regardless of climate zone. Overall, the impacts of exotic grass‐fuelled fires on native species composition and cover in seasonally dry Hawaiian ecosystems was context specific. This specificity is best explained by differences between the climatic zones in which fire occurred. Elevation was the main physical variable that differed among the climatic zones and it alone could explain a large percentage of the variation in native cover among sites. Rainfall, by contrast, did not vary systematically with elevation. Elevation is associated with differences in composition of the native species assemblages. In the coastal lowlands, the native grass Heteropogon contortus, was largely responsible for positive changes in native cover after fire although other native species also increased. Like the exotic grasses, this species is a perennial C4 grass. It is lacking in the submontane zone and there are no comparable native species there and almost all native species in the submontane zone were reduced by fire. The lack of fire tolerant species in the submontane zone thus clearly contributes to the devastating impact of fire upon native cover there.     

Before Cattle: A Comparative Floristic Study of Eucalyptus Savanna Grazed by Macropods and Cattle in North Queensland,Australia1

Floristic differences between cattle-grazed and macropod-grazed sites (protected from domestic stock by enclosure within a Holocene lava flow) were examined in Eucalyptus savanna in north Queensland, Australia. While macropod-grazed sites have had a lower incidence of fire than the cattle-grazed sites, it seems probable that grazing impacts per se are the major differences between these treatments. One dominant (Capillipedium parviflorum) and another common (Sehima nervosa) native perennial grass in the macropod-grazed treatment were not recorded within the cattle-grazed treatment. There was generally a lower species richness and Shannon-Weaver diversity of perennial forbs in the cattle-grazed treatment compared to the macropod-grazed treatment. There was higher richness, diversity, and abundance of annual grasses in the cattle-grazed treatment compared to the macropod-grazed treatment. The relative richness of native and exotic species was not significantly affected by cattle grazing.     

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.     

Promoting Native Vegetation and Diversity in Exotic Annual Grass Infestations

Exotic plant invasions are especially problematic because reestablishment of native perennial vegetation is rarely successful. It may be more appropriate to treat exotic plant infestations that still have some remaining native vegetation. We evaluated this restoration strategy by measuring the effects of spring burning, fall burning, fall applied imazapic, spring burning with fall applied imazapic, and fall burning with fall applied imazapic on the exotic annual grass, medusahead (Taeniatherum caput‐medusae (L.) Nevski), and native vegetation at six sites in Oregon for 2 years post‐treatment. Medusahead infestations included in this study had some residual native perennial bunchgrasses and forbs. Burning followed by imazapic application provided the best control of medusahead and resulted in the greatest increases in native perennial vegetation. However, imazapic application decreased native annual forb cover the first year post‐treatment and density the first and second year post‐treatment. The spring burn followed by imazapic application produced an almost 2‐fold increase in plant species diversity compared to the control. The fall burn followed by imazapic application also increased diversity compared to the control. Results of this study indicate that native plants can be promoted in medusahead invasions; however, responses vary by plant functional group and treatment. Our results compared to previous research suggest that restoration of plant communities invaded by exotic annual grass may be more successful if efforts focus on areas with some residual native perennial vegetation. Thus, invasive plant infestations with some native vegetation remaining should receive priority for restoration efforts over near monocultures of invasive plant species.     

Fire and seedling population dynamics in western Oregon prairies

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