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.     

Reduced soil compaction enhances establishment of non-native plant species

Many studies have shown that soil disturbance facilitates establishment of invasive, non-native plant species, and a number of mechanisms have been isolated that contribute to the process. To our knowledge no studies have isolated the role of altered soil compaction, a likely correlate of many types of soil disturbance, in facilitating invasion. To address this, we measured the response of seeded non-native and native plant species to four levels of soil compaction in mesocosms placed in an abandoned agricultural field in the Methow Valley, Washington, USA. Soil compaction levels reflected the range of resistance to penetration (0.1–3.0 kg cm⁻²) measured on disturbed soils throughout the study system prior to the experiment. Percent cover of non-native species, namely Bromus tectorum and Centaurea diffusa, decreased by 34% from the least to the most compacted treatments, whereas percent cover of native species, mostly Pseudoroegneria spicata and Lupinus spp., did not respond to compaction treatments. Experimental results were supported by a survey of soil penetration resistance and percent cover by species in 18 abandoned agricultural fields. Percent cover of B. tectorum was negatively related to soil compaction levels, whereas none of the native species showed any response to soil compaction. These results highlight a potentially important, though overlooked, aspect of soil disturbance that may contribute to subsequent non-native plant establishment.     

Carbon Addition as a Countermeasure Against Biological Invasion by Plants

Increased nitrogen availability is known to favor invasion by non-native plants into natural grasslands. This suggests that decreasing nitrogen availability might serve as a countermeasure against invasion. One way to at least temporarily decrease nitrogen availability to plants is to increase microbial nitrogen uptake by adding carbon to the soil, and sawdust is a carbon source whose low cost could make it a practical conservation tool. To test whether adding sawdust to soil can counter the tendency of nitrogen enrichment to promote invasions by non-native plants, we hand-tilled 1.5kg sawdust/m² into the upper soil of the bare, nitrogen-rich patches left by dead shrubs of the nitrogen-fixing shrub Lupinus arboreus in two nearby areas with contrasting levels of invasion in a coastal grassland in northern California. After two years, in both areas, patches with sawdust had 40% less biomass of non-native plants than patches without sawdust, whereas biomass of native plants was not affected by sawdust addition. The more negative effect of sawdust on non-native species was specifically due to an effect on non-native grasses; adding sawdust increased the frequency of both native and non-native forbs. Results suggest that adding carbon as sawdust to soil can help counter invasion of grassland by non-native plants when invasion is being promoted by increased nitrogen availability, especially when the major invasive species are grasses.     

Controls over invasion of Bromus tectorum: The importance of climate,soil, disturbance and seed availability

Question: Predicting the future abundance and distribution of invasive plants requires knowing how they respond to environmental conditions. In arid and semi‐arid ecosystems where water is a limiting resource, environmental conditions and disturbance patterns influence invasions by altering acquisition and utilization of water over space and time. We ask: 1. How do variations in climatic and soil properties influence temporal soil water dynamics? 2. How does this variation affect the establishment of Bromus tectorum (cheatgrass), a cool‐season annual grass that has successfully colonized much of the U.S. Great Basin? Location: Short‐grass Steppe in northeastern Colorado, USA; Arid Lands Ecology reserve in southeastern Washington, USA; and the Patagonian steppe of the Chubut province in Argentina. Methods: We utilized a soil water model to simulate seasonal soil water dynamics in multiple combinations of climatic and soil properties. In addition, we utilized a gap dynamics model to simulate the impact of disturbance regime and seed availability on competition between B. tectorum and native plants. Results: Our results suggest that climate is very important, but that soil properties do not significantly influence the probability of observing conditions suitable for B. tectorum establishment. Results of the plant competition model indicate that frequent disturbance causes more Bromus tectorum in invaded areas and higher seed availability causes faster invasion. Conclusions: These results imply a general framework for understanding Bromus tectorum invasion in which climatic conditions dictate which areas are susceptible to invasion, disturbance regime dictates the severity of invasion and seed availability dictates the speed of invasion.     

Competitive impacts and responses of an invasive weed: dependencies on nitrogen and phosphorus availability

Changes in competitive interactions under conditions of enhanced resource availability could explain the invasion success of some problematic plant species. For one invader of North American grasslands, Centaurea diffusa (diffuse knapweed), we test three hypotheses: (1) under ambient (high resource) conditions, C. diffusa is better able to tolerate competition from the resident community (competitive response), (2) under ambient conditions, C. diffusa strong impacts the competitive environment (competitive effect), and (3) reductions in nitrogen and/or phosphorus availability diminish these advantages. In support of our first hypothesis, C. diffusa was the most tolerant to neighbor competition of the four focal species under current resource conditions. In opposition to our second hypothesis, however, neighborhoods that contained C. diffusa and those where C. diffusa had been selectively removed did not differ in their impact on the performance of target transplant individuals or on resource conditions. Reduction in resource availability influenced competitive tolerance but not competitive impact, in partial support of our last hypothesis. Reduction in soil nitrogen (via sucrose carbon addition) enhanced the degree of neighbor competition experienced by all species but did not change their relative rankings; C. diffusa remained the best competitor under low nitrogen conditions. Reduction of soil phosphorus (via gypsum addition) weakened the ability of C. diffusa to tolerate neighbor competition proportionately more than the other focal species. Consequently, under low phosphorus conditions, C. diffusa lost its competitive advantage and tolerated neighbor competition similarly to the other focal species. We conclude that C. diffusa invasion may be double-edged: C. diffusa is less limited by nitrogen than the other focal species and is better able to utilize phosphorus to its competitive advantage.     

Competition between two grass species with and without grazing over a productivity gradient

Soil nutrient-level and herbivory are predicted to have opposing effects on the allocation pattern of the competitive dominant plant species. Lower stem and higher leaf allocation are favoured when plants are grazed, whereas a higher stem allocation is favoured at high nutrient levels. Grazing by hares and geese can prevent invasion of the tall Elymus athericus, into short vegetation of Festuca rubra, at unproductive stages of salt-marsh succession but not at more productive stages. We hypothesise that the negative effect of herbivory on Elymus decreases due to increasing soil nitrogen levels and shifts the competitive balance towards this species. We tested how simulated grazing and nitrogen availability affected the competitive balance between adult plants of both grass species in a greenhouse experiment. Elymus had a higher above-ground biomass production, invested relatively more in stem and root tissue and had a larger shoot length than Festuca. The above-ground relative yield of Elymus in mixtures of both species increased with increasing nitrogen levels. This indicates that Elymus was the superior competitor at high soil fertility. Although clipping removed relatively more biomass from Elymus than from Festuca and exceeded the observed biomass removal in field conditions, it did not change the competitive balance between both species. Decreasing effects of herbivory due to increasing nitrogen levels are not a likely explanation for the invasion of Elymus in productive marshes. The results suggest that once Elymus has established it can easily invade vegetation dominated by Festuca irrespective of grazing by herbivores such as hares and geese. Herbivory by small herbivores may mainly retard the invasion of this plant by influencing establishment itself.     

Responses to fertility and disturbance in a low-diversity grassland

We examined variation in species composition in a low-diversity, anthropogenic grassland in response to 11 years of nitrogen (N) manipulation and disturbance. The species-poor grassland (2–3 species/0.5 m²) represents a wide spread vegetation type (>10 million ha in North America) dominated by the introduced perennial grasses Bromus inermis and Agropyron cristatum. Four levels of N and three of soil disturbance were applied in all combinations to plots (5 × 15 m, N = 120) in a completely randomized design each year. Seeds or transplants of 47 species were added to ensure that dispersal was not a barrier to changes in species composition. After 11 years of treatment, all but the most disturbed plots continued to be dominated by B. inermis. The cover of the second-most abundant species, A. cristatum, decreased with disturbance but did not vary significantly with N. Despite the lack of changes in the identity of the dominant species, our environmental manipulations strongly influenced ecosystem characteristics. Added N increased soil available N, and decreased the cover of bare ground and light availability. Soil disturbance decreased aboveground biomass, and increased the cover of bare ground and light availability. Sawdust application, designed to decrease N availability, significantly reduced community biomass, and increased light availability and the cover of bare ground, but did not alter nutrient availability or species composition. The results highlight the difficulty of restoring diversity in species-poor, anthropogenic communities dominated by introduced species, and thus the importance of conserving remnants of diverse natural grasslands.     

A native nitrogen-fixing shrub facilitates weed invasion

Invasions by exotic weedy plants frequently occur in highly disturbed or otherwise anthropogenically altered habitats. Here we present evidence that, within California coastal prairie, invasion also can be facilitated by a native nitrogen-fixing shrub, bush lupine (Lupinus arboreus). Bush lupines fix nitrogen and grow rapidly, fertilizing the sandy soil with nitrogen-rich litter. The dense lupine canopy blocks light, restricting vegetative growth under bushes. Heavy insect herbivory kills lupines, opening exposed nitrogen-rich sites within the plant community. Eventual re-establishment of lupine occurs because of an abundant and long-lived seed bank. Lupine germination, rapid growth, shading and fertilization of sites, and then death after only a few years, results in a mosaic of nutrient-rich sites that are available to invading species. To determine the role of bush lupine death and nitrogen enrichment in community composition, we examined nutrient dynamics and plant community characteristics within a site only recently colonized by lupine, comparing patches where lupines had recently died or were experimentally killed with adjacent areas lacking lupine. In experimentally killed patches, instantaneous pool sizes of exchangeable ammonium and nitrate nitrogen were higher than in adjacent sites free of lupine. Seedlings of the introduced grass Bromus diandrus accumulated 48% greater root biomass and 93% more shoot biomass when grown in a greenhouse in soil collected under experimentally killed lupines compared to B. diandrus seedlings grown in soil collected at least 1 m away from lupines. At the end of the spring growing season, total above-ground live plant biomass was more than twice as great in dead lupine patches as in the adjacent lupine-free grassland, but dead lupine patches contained 47% fewer plant species and 57% fewer native species. Sites where lupines have repeatedly died and reestablished during recent decades support an interstitial grassland community high in productivity but low in diversity, composed of mostly weedy introduced annual plants. In contrast, at a site only recently colonized by bush lupines, the interstitial grassland consists of a less productive but more diverse set of native and introduced species. We suggest that repeated bouts of lupine germination, establishment, and death can convert a rich native plant community into a less diverse collection of introduced weeds.     

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C₃ grass, and Boutelouagracilis (HBK.) Lag., a native C₄ grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.     

Contrasting effects of resource availability and plant mortality on plant community invasion by Bromus tectorum L.

The positive effect of disturbance on plant community invasibility is one of the more consistent results in invasion ecology. It is generally attributed to a coincident increase in available resources (due to the disturbance) that allows non-resident plant species to establish (Davis MA, Grime JP Thompson K, J Ecol 88:528–534, 2000). However, most research addressing this issue has been in artificial or highly modified plant communities. Our goal in this study was to investigate the interactive effects of resource availability and plant mortality disturbance on the invasion of natural plant communities. We conducted a series of experiments that examined the response of Bromus tectorum L., a highly invasive annual grass, to experimentally created gradients of resource availability [nitrogen (N) and water] and resident plant species mortality. We found that B. tectorum biomass was co-limited by N and water. Biomass at the end of the growing season was a saturating function (i.e., increased to a maximum) of water, which determined maximum biomass, and N, which determined the rate at which maximum biomass was attained. Despite that fact that plant mortality increased N availability, it had a negative impact on invasion success. Plant mortality also decreased foliar cover, standing dead biomass, and soil cover by litter. In harsh environments, removing foliar and soil cover may increase germination and seedling stress by increasing soil temperatures and water loss. Across all treatments, B. tectorum success decreased with decreasing foliar cover and standing dead biomass. This, in combination with the strong limitation of B. tectorum biomass by water in this experiment, suggests that our plant mortality disturbance removed soil cover that may have otherwise aided B. tectorum invasion into this semi-arid plant community by reducing water stress.