Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain

1. We examined the effect of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain in northern Australia. Macroinvertebrates were sampled from four grass communities: (1) para grass, (2) hymenachne (Hymenachne acutigluma), a native perennial; (3) rice (Oryza meridionalis), a native annual, and (4) areas where para grass had been sprayed with herbicide. 2. Macroinvertebrate richness, abundance and community similarity showed very few differences among the grass communities, particularly in the epiphytic habitat. Benthic invertebrates showed some differences among grasses, with lower richness and abundance and different community structure associated with hymenachne. Herbicide control of para grass had no apparent effect on benthic invertebrates but reduced the abundance of epiphytic invertebrates in the short term. 3. The results of this study indicate that para grass has very little impact on macroinvertebrate communities, despite the changes to macrophyte communities. This is probably because para grass has similar physical structure to the native grasses and because none of these grasses contribute directly to aquatic food webs. Control of para grass using herbicide has little impact on aquatic invertebrates. This suggests that predicting the impact of weed invasion in wetlands requires an understanding of both the functional properties of macrophytes and the habitat preferences of the macroinvertebrates.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

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

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

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.     

Effects of exotic grass invasion on spatial heterogeneity in the ground-layer of grassy woodlands

We quantified the effects of exotic annual grass invasion on the ground-layer structure of grassy eucalypt woodlands, with the aim of determining if weed invasion decreased gap size and plant basal area leading to reduced spatial heterogeneity. We measured plant density, distance between plants and basal plant area in woodland sites which ranged from zero to 100% exotic plant cover in the ground-layer. The ground-layer in uninvaded woodlands was heterogeneous, with a large variation in basal plant area and distance between plants. Exotic annual grass density was positively correlated with total plant density, whereas native plant density was negatively correlated. Total plant basal area decreased as total plant density increased, with a lower total plant area in exotic dominated transects compared to native dominated. Variation in basal plant area decreased with increasing plant density. Exotic annual grasses were more closely spaced together (smaller gap size) and had a smaller basal area than the native grasses and rushes. There was also less variation in basal area and gap size with individual exotic annual grasses compared to the native grasses. Inter-plant distance was greater for both the native and exotic grasses when they had native grasses neighbouring them instead of exotic grasses. These findings show that woodlands invaded by exotic annual grasses have relatively less spatial heterogeneity in the ground-layer. These results have implications for other aspects of perennial grassy ecosystems invaded by annual grasses, including plant recruitment and restoration strategies.     

Grouping plant species by shared native range, and not by native status, predicts response to an exotic herbivore

Differences among exotic species can be as large as differences between native and exotic species. Typically, however, only the distinction between native and exotic is made when predicting responses in a community. In this paper, I examine the response of plant species to experimental disturbance and exclusion of invasive European rabbits (Oryctolagus cuniculus) in a grassland community with exotic plants originating from five continents. I explore group responses based on native status, shared native range with rabbits, having a congener from the native range of rabbits, life-history (e.g., annual), and life-form (e.g., grass). Individual species responses to rabbits were idiosyncratic, but group responses were predicted by continent of origin, not native status. Native status did predict response to disturbance with almost uniform responses within groups. Exotic species, regardless of origin, were positively affected by disturbance. Native species, in contrast, were negatively affected by disturbance. These results suggest that grouping plant species by native status is valid for questions of disturbance, but when analyzing outcomes of interactions, factors other than native status, such as shared evolutionary history, should be considered.     

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.     

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.     

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.     

Biotic resistance to plant invasions? Native herbivores prefer non‐native plants

In contrast to expectations of the enemy release hypothesis, but consistent with the notion of biotic resistance, we found that native generalist crayfishes preferred exotic over native freshwater plants by a 3 : 1 ratio when plants were paired by taxonomic relatedness. Native crayfishes also preferred exotic over native plants when tested across 57 native and 15 exotic plants found growing sympatrically at 11 sites throughout the southeastern USA. Exotic grass carp that share little evolutionary history with most of these plants exhibited no preference for native vs. exotic species. Analyses of three terrestrial data sets showed similar patterns, with native herbivores generally preferring exotic plants, while exotic herbivores rarely exhibited a preference. Thus, exotic plants may escape their coevolved herbivores only to be preferentially consumed by the native generalist herbivores in their new ranges, suggesting that native herbivores may provide biotic resistance to plant invasions.     

Biogeochemical and Microbial Legacies of Non‐Native Grasses Can Affect Restoration Success

The restoration of disturbed ecosystems is challenging and often unsuccessful, particularly when non‐native plants are abundant. Ecosystem restoration may be hindered by the effects of non‐native plants on soil biogeochemical characteristics and microbial communities that persist even after plants are removed. To examine the importance of soil legacy effects, we used experimental restorations of Florida shrubland habitat that had been degraded by the introduction of non‐native grasses coupled with either mechanical disturbance or pasture conversion. We removed non‐native grasses and inoculated soils with native microbial communities at each degraded site, then examined how habitat structure, soil nitrogen, soil microbial abundances, and native seed germination responded over two years compared to undisturbed native sites. Grass removal treatments effectively restored some aspects of native habitat structure, including decreased exotic grass cover, increased bare ground, and reduced litter cover. Soil fungal abundance was also somewhat restored by grass removals, but soil algal abundance was unaffected. In addition, grass removal and microbial inoculation improved seed germination rates in degraded sites, but these remained quite low compared to native sites. High soil nitrogen persisted throughout the experiment regardless of treatment. Many treatment effects were site‐specific, however, with legacies in the more degraded vegetation type tending to be more difficult to overcome. These results support the need for context‐dependent restoration approaches and suggest that the degree of soil legacy effects may be a good indicator of restoration potential.     

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.     

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.     

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.     

The grass may not always be greener: projected reductions in climatic suitability for exotic grasses under future climates in Australia

Climate change presents a new challenge for the management of invasive exotic species that threaten both biodiversity and agricultural productivity. The invasion of exotic perennial grasses throughout the globe is particularly problematic given their impacts on a broad range of native plant communities and livelihoods. As the climate continues to change, pre-emptive long-term management strategies for exotic grasses will become increasingly important. Using species distribution modelling we investigated potential changes to the location of climatically suitable habitat for some exotic perennial grass species currently in Australia, under a range of future climate scenarios for the decade centred around 2050. We focus on eleven species shortlisted or declared as the Weeds of National Significance or Alert List species in Australia, which have also become successful invaders in other parts of the world. Our results indicate that the extent of climatically suitable habitat available for all of the exotic grasses modelled is projected to decrease under climate scenarios for 2050. This reduction is most severe for the three species of Needle Grass (genus Nassella) that currently have infestations in the south-east of the continent. Combined with information on other aspects of establishment risk (e.g. demographic rates, human-use, propagule pressure), predictions of reduced climatic suitability provide justification for re-assessing which weeds are prioritised for intensive management as the climate changes.     

Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.     

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.     

Soil ecosystem function under native and exotic plant assemblages as alternative states of successional grasslands

Old fields often become dominated by exotic plants establishing persistent community states. Ecosystem functioning may differ widely between such novel communities and the native-dominated counterparts. We evaluated soil ecosystem attributes in native and exotic (synthetic) grass assemblages established on a newly abandoned field, and in remnants of native grassland in the Inland Pampa, Argentina. We asked whether exotic species alter soil functioning through the quality of the litter they shed or by changing the decomposition environment. Litter decomposition of the exotic dominant Festuca arundinacea in exotic assemblages was faster than that of the native dominant Paspalum quadrifarium in native assemblages and remnant grasslands. Decomposition of a standard litter (Triticum aestivum) was also faster in exotic assemblages than in native assemblages and remnant grasslands. In a common garden, F. arundinacea showed higher decay rates than P. quadrifarium, which reflected the higher N content and lower C:N of the exotic grass litter. Soil respiration rates were higher in the exotic than in the native assemblages and remnant grasslands. Yet there were no significant differences in soil N availability or net N mineralization between exotic and native assemblages. Our results suggest that exotic grass dominance affected ecosystem function by producing a more decomposable leaf litter and by increasing soil decomposer activity. These changes might contribute to the extended dominance of fast-growing exotic grasses during old-field succession. Further, increased organic matter turnover under novel, exotic communities could reduce the carbon storage capacity of the system in the long term.     

Plant–soil feedbacks of exotic plant species across life forms: a meta-analysis

Invasive exotic plant species effects on soil biota and processes in their new range can promote or counteract invasions via changed plant–soil feedback interactions to themselves or to native plant species. Recent meta-analyses reveale that soil influenced by native and exotic plant species is affecting growth and performance of natives more strongly than exotics. However, the question is how uniform these responses are across contrasting life forms. Here, we test the hypothesis that life form matters for effects on soil and plant–soil feedback. In a meta-analysis we show that exotics enhanced C cycling, numbers of meso-invertebrates and nematodes, while having variable effects on other soil biota and processes. Plant effects on soil biota and processes were not dependent on life form, but patterns in feedback effects of natives and exotics were dependent on life form. Native grasses and forbs caused changes in soil that subsequently negatively affected their biomass, whereas native trees caused changes in soil that subsequently positively affected their biomass. Most exotics had neutral feedback effects, although exotic forbs had positive feedback effects. Effects of exotics on natives differed among plant life forms. Native trees were inhibited in soils conditioned by exotics, whereas native grasses were positively influenced in soil conditioned by exotics. We conclude that plant life form matters when comparing plant–soil feedback effects both within and between natives and exotics. We propose that impact analyses of exotic plant species on the performance of native plant species can be improved by comparing responses within plant life form.     

Invasion by Non-Native Annual Grasses: The Importance of Species Biomass, Composition, and Time Among California Native Grasses of the Central Valley

The Central Valley of California is noted for its dearth of remnant native grass populations and for low native grass seedling establishment within grasslands now dominated by non‐native annual species. In contrast, remnant populations are common along the coast, and studies have shown an ability for seedlings and adults to compete with non‐native annual grasses. The invasibility of well‐established populations of native grasses in the Central Valley remains unclear. The objectives of this study were to compare the invasibility of native grasses differing in density and species composition and, given the species in this study, to assess the ability of mixes with greater species richness to resist invasion relative to their abilities in monoculture. In the Sacramento Valley of California, six species of native grasses were planted at three densities in monospecific and mixed‐species plots. Percent cover of native perennial and non‐native annual grasses was measured in years 2 and 3, and biomass was sampled in year 5. Native grass biomass and, to a lesser extent, species composition were important in explaining variation in non‐native grass invasibility in the fifth year. Species‐rich treatments did not experience less invasion than would be expected by the proportional invasibility of each species in monoculture. However, invasibility of plots consisting of slower growing, shorter statured species decreased over time, suggesting a successional benefit to diverse communities. This study demonstrates that established stands of native grasses in the Sacramento Valley can resist invasion by non‐native annual grasses and that stand biomass is a particularly important factor in determining invasibility.