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.     

Growth and competitive effects of Centaurea stoebe populations in response to simulated nitrogen deposition

Increased resource availability can promote invasion by exotic plants, raising concerns over the potential effects of global increases in the deposition of nitrogen (N). It is poorly understood why increased N favors exotics over natives. Fast growth may be a general trait of good invaders and these species may have exceptional abilities to increase growth rates in response to N deposition. Additionally, invaders commonly displace locals, and thus may have inherently greater competitive abilities. The mean growth response of Centaurea stoebe to two N levels was significantly greater than that of North American (NA) species. Growth responses to N did not vary among C. stoebe populations or NA species. Without supplemental N, NA species were better competitors than C. stoebe, and C. stoebe populations varied in competitive effects. The competitive effects of C. stoebe populations increased with N whereas the competitive effects of NA species decreased, eliminating the overall competitive advantage demonstrated by NA species in soil without N added. These results suggest that simulated N deposition may enhance C. stoebe invasion through increasing its growth and relative competitive advantage, and also indicate the possibility of local adaptation in competitive effects across the introduced range of an invader.     

Diffuse knapweed (Centaurea diffusa Lam.) seedling emergence and establishment in a Colorado grassland

Knapweeds (Centaurea spp.) are damaging invaders of grasslands and other North American rangelands. A field study was conducted to determine conditions that promote diffuse knapweed (C. diffusa) emergence and establishment in a native Colorado grassland (North America). Knapweed was planted in native grassland under treatments with different opening sizes, levels of competition, knapweed seed burial and season of seeding. There was no effect of opening size where competing natives were alive, but knapweed emergence in 5- and 15-cm openings was higher than 0-cm openings where natives were killed. Reducing competition reduced fall diffuse knapweed emergence, but did not affect spring emergence. Seed burial increased knapweed emergence, but the effect varied by season. Although diffuse knapweed emergence reached 35%, only four plants survived from 3,600 seeds. This native grassland did not prevent knapweed emergence or establishment, but both were so low that rapid knapweed invasion is unlikely.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Effects of nitrogen availability on competition between Bromus tectorum and Bouteloua gracilis

Exotic plant invasions are a serious concern for land managers and conservationists. There is evidence that increased nitrogen availability favors exotic species and decreased nitrogen availability favors non-weedy native species. This study was conducted to test the effect of nitrogen availability on competition between two grass species with contrasting life histories, cheatgrass (Bromus tectorum), a North American exotic, and blue grama (Bouteloua gracilis), a North American native. We investigated the effects of nitrogen availability and competition on aboveground biomass, belowground biomass, height, and % nitrogen tissue concentrations by growing the two species in the greenhouse under five levels of nitrogen and six levels of competition. Nitrogen availability affected competition between Bromus tectorum and Bouteloua gracilis. At the lowest level of N availability, neither species was affected by competition. As N availability increased, aboveground biomass gain of Bromus was more negatively affected by intraspecific competition relative to interspecific competition while the opposite occurred for Bouteloua. At the competition level at which each species gained the most aboveground biomass, Bromus had a linear response to increasing N availability while the response of Bouteloua was asymptotic. Our results do provide some support for the theory that fast growing exotic species have a rapid response to nutrient enrichment while native non-weedy species do not, and that low N levels can reduce competitive pressure from the exotic on the native.     

Extended leaf phenology may drive plant invasion through direct and apparent competition

Invasive plants can inflict great harm, yet drivers of successful invasion remain unclear. Many invaders of North American deciduous forests exhibit extended leaf phenology (ELP), or longer growing season relative to natives. ELP may grant invaders competitive advantages, but we argue that ELP more potently drives invasion in the presence of herbivores. ELP invaders can support herbivores by lessening starvation during winter; consequently, native plants may suffer when attacked later through apparent competition. As modeled here, even short ELP can promote competitive success of invaders, and apparent competition sharply enhances ELP invader dominance. In ‘partial enemy escape’ scenarios, a less palatable ELP invader nearly excludes a preferred native where an invader without ELP could not. Together, ELP and apparent competition enhance invasion even when biotic resistance should suppress it, i.e. when the invader competes weakly or provides preferred forage. Thus, ELP‐apparent competition interactions grant invaders considerable success while challenging core tenets of invasion ecology.     

<Emphasis Type="Italic">Acroptilon repens</Emphasis>, an Asian invader,has stronger competitive effects on species from America than species from its native range

The ability to competitively suppress native species is key to successful invasion. Since invasions involve an increase in abundance or dominance of a species in its non-native range, competitive effects might be expected to be stronger in the non-native range of an invader; however, there have been few comparisons of the competitive effects of invasive plants on species from invaded ranges versus species from native ranges. We compared the competitive and allelopathic effects of Acroptilon repens on native North American species to effects on related species from the native range of Acroptilon in Uzbekistan. We also compared the competitive interactions among these North American and Eurasian species, in the absence of Acroptilon, examining the hypothesis that particular regional species pools may show differences in competitive ability. The results showed that Acroptilon had stronger competitive effects against native North American species than against species native to Uzbekistan. There was no difference in the competitive effects among Eurasians and North Americans. The effects of leachates collected from Acroptilon roots were weak but more negative on species from North America than on species from Uzbekistan. Our results suggest that inherently stronger competitive and allelopathic effects of Acroptilon on North American plants than on plants from its native range may contribute to its invasive success.     

Nitrogen acquisition by annual and perennial grass seedlings: testing the roles of performance and plasticity to explain plant invasion

Differences in resource acquisition between native and exotic plants is one hypothesis to explain invasive plant success. Mechanisms include greater resource acquisition rates and greater plasticity in resource acquisition by invasive exotic species compared to non-invasive natives. We assess the support for these mechanisms by comparing nitrate acquisition and growth of invasive annual and perennial grass seedlings in western North America. Two invasive exotic grasses (Bromus tectorum and Taeniatherum caput-medusae) and three perennial native and exotic grasses (Pseudoroegneria spicata, Elymus elymoides, and Agropyron cristatum) were grown at various temperatures typical of autumn and springtime when resource are abundant and dominance is determined by rapid growth and acquisition of resources. Bromus tectorum and perennial grasses had similar rates of nitrate acquisition at low temperature, but acquisition by B. tectorum significantly exceeded perennial grasses at higher temperature. Consequently, B. tectorum had the highest acquisition plasticity, showcasing its ability to take advantage of transient warm periods in autumn and spring. Nitrate acquisition by perennial grasses was limited either by root production or rate of acquisition per unit root mass, suggesting a trade-off between nutrient acquisition and allocation of growth to structural tissues. Our results indicate the importance of plasticity in resource acquisition when temperatures are warm such as following autumn emergence by B. tectorum. Highly flexible and opportunistic nitrate acquisition appears to be a mechanism whereby invasive annual grasses exploit soil nitrogen that perennials cannot use.     

Established native perennial grasses out-compete an invasive annual grass regardless of soil water and nutrient availability

Competition and resource availability influence invasions into native perennial grasslands by non-native annual grasses such as Bromus tectorum. In two greenhouse experiments we examined the influence of competition, water availability, and elevated nitrogen (N) and phosphorus (P) availability on growth and reproduction of the invasive annual grass B. tectorum and two native perennial grasses (Elymus elymoides, Pascopyrum smithii). Bromus tectorum aboveground biomass and seed production were significantly reduced when grown with one or more established native perennial grasses. Conversely, average seed weight and germination were significantly lower in the B. tectorum monoculture than in competition native perennial grasses. Intraspecific competition reduced per-plant production of both established native grasses, whereas interspecific competition from B. tectorum increased production. Established native perennial grasses were highly competitive against B. tectorum, regardless of water, N, or P availability. Bromus tectorum reproductive potential (viable seed production) was not significantly influenced by any experimental manipulation, except for competition with P. smithii. In all cases, B. tectorum per-plant production of viable seeds exceeded parental replacement. Our results show that established plants of Elymus elymoides and Pascopyrum smithii compete successfully against B. tectorum over a wide range of soil resource availability.     

An unfortunate alliance: Native shrubs increase the abundance,performance, and apparent impacts of Bromus tectorum across a regional aridity gradient

Interspecific facilitation contributes to the assembly of desert plant communities. However, we know little of how desert communities invaded by exotic species respond to facilitation along regional-scale aridity gradients. These measures are essential for predicting how desert plant communities might respond to concomitant plant invasion and environmental change. Here, we evaluated the potential for Bromus tectorum (a dominant invasive plant species) and the broader herbaceous plant community to form positive associations with native shrubs along a substantial aridity gradient across the Great Basin, Mojave, and San Joaquin Deserts in North America. Along this gradient, we sampled metrics of abundance and performance for B. tectorum, all native herbaceous species combined, all exotic herbaceous species combined, and the total herbaceous community using 180 pairs of shrub and open microsites. Across the gradient, B. tectorum formed strong positive associations with native shrubs, achieving 1.6–2.2 times greater abundance, biomass, and reproductive output under native shrubs than away from shrubs, regardless of relative aridity. In contrast, the broader herbaceous community was not positively associated with native shrubs. Interestingly, increasing B. tectorum abundance corresponded to decreasing native abundance, native species richness, exotic species richness, and total species richness under but not away from shrubs. Taken together, these findings suggest that native shrubs have considerable potential to directly (by increasing abundance and performance) and indirectly (by increasing competitive effects on neighbors) facilitate B. tectorum invasion across a large portion of the non-native range.     

Intraspecific and interspecific pair-wise seedling competition between exotic annual grasses and native perennials: plant–soil relationships

Few studies have examined plant–soil relationships in competitive arenas between exotic and native plants in the western United States. A pair-wise competitive design was used to evaluate plant–soil relationships between seedlings of the exotic annual grasses Bromus tectorum and Taeniatherium caput-medusae and the native perennial grasses Elymus elymoides and Pseudoroegneria spicata. Two soils were tested: an arid soil (argid) occupied by E. elymoides and presently invaded by B. tectorum and a high elevation, high organic matter, soil (aquept) where none of the tested species would typically occur. Plant growth proceeded for 85 days at which time above-ground biomass and tissue nutrient concentrations were quantified. Soil also was collected from the rooting zone beneath each species and analyzed for various nutrient pools. The exotic species had significantly greater above-ground biomass than the natives and grew far better in the aquept soil than the argid soil. Growth of B. tectorum, and to some degree, T. caput-medusae was suppressed in intraspecific competition and enhanced, especially in the aquept soil, when competing with the natives. Although not significant, biomass of natives strongly trended downward when competing with the exotic grasses. Overall, concentrations of tissue nutrients were minimally affected by competition, but natives tended to be more negatively affected by competition with exotics. Except for phosphorus (P), all species had significantly greater nutrient concentrations when growing in the aquept soil compared to the argid soil. In both soils, exotics had significant greater tissue concentrations of manganese (Mn), magnesium (Mg), and iron (Fe), while natives had significantly greater nitrogen (N). Species affects on soil nutrient pools occurred mostly in the aquept soil with exotic species significantly decreasing pools of available N, potentially available N, and soil-solution pools of calcium (Ca²⁺), potassium (K⁺), and magnesium (Mg²⁺) relative to natives. Overall, the data suggest that, in the seedling state, B. tectorum is a superior competitor. Moreover, when the natives compete intra- or interspecifically, particularly in the aquept soil, availability of N and other nutrients in their rooting zone is consistently greater than when they compete interspecifically with the exotic grasses. These data suggest the exotics are able to co-opt nutrients in the rooting zone of the natives and perhaps gain a competitive advantage.     

Direct Effects of Soil Amendments on Field Emergence and Growth of the Invasive Annual Grass Bromus tectorum L. and the Native Perennial Grass Hilaria jamesii (Torr.) Benth

Bromus tectorum L. is a non-native, annual grass that has invaded western North America. In SE Utah, B. tectorum generally occurs in grasslands dominated by the native perennial grass, Hilaria jamesii (Torr.) Benth. and rarely where the natives Stipa hymenoides Roem. and Schult. and S. comata Trin. & Rupr. are dominant. This patchy invasion is likely due to differences in soil chemistry. Previous laboratory experiments investigated using soil amendments that would allow B. tectorum to germinate but would reduce B. tectorum emergence without affecting H. jamesii. For this study we selected the most successful treatments (CaCl₂, MgCl₂, NaCl and zeolite) from a previous laboratory study and applied them in the field in two different years at B. tectorum-dominated field sites. All amendments except the lowest level of CaCl₂ and zeolite negatively affected B. tectorum emergence and/or biomass. No amendments negatively affected the biomass of H. jamesii but NaCl reduced emergence. Amendment effectiveness depended on year of application and the length of time since application. The medium concentration of zeolite had the strongest negative effect on B. tectorum with little effect on H. jamesii. We conducted a laboratory experiment to determine why zeolite was effective and found it released large amounts of Na⁺, adsorbed Ca²⁺, and increased Zn²⁺, Fe²⁺, Mn²⁺, Cu²⁺, exchangeable Mg²⁺, exchangeable K, and NH₄⁺ in the soil. Our results suggest several possible amendments to control B. tectorum. However, variability in effectiveness due to abiotic factors such as precipitation and soil type must be accounted for when establishing management plans.     

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Cheatgrass is favored by warming but not CO2 enrichment in a semi‐arid grassland

Elevated CO₂ and warming may alter terrestrial ecosystems by promoting invasive plants with strong community and ecosystem impacts. Invasive plant responses to elevated CO₂ and warming are difficult to predict, however, because of the many mechanisms involved, including modification of phenology, physiology, and cycling of nitrogen and water. Understanding the relative and interactive importance of these processes requires multifactor experiments under realistic field conditions. Here, we test how free‐air CO₂ enrichment (to 600 ppmv) and infrared warming (+1.5 °C day/3 °C night) influence a functionally and phenologically distinct invasive plant in semi‐arid mixed‐grass prairie. Bromus tectorum (cheatgrass), a fast‐growing Eurasian winter annual grass, increases fire frequency and reduces biological diversity across millions of hectares in western North America. Across 2 years, we found that warming more than tripled B. tectorum biomass and seed production, due to a combination of increased recruitment and increased growth. These results were observed with and without competition from native species, under wet and dry conditions (corresponding with tenfold differences in B. tectorum biomass), and despite the fact that warming reduced soil water. In contrast, elevated CO₂ had little effect on B. tectorum invasion or soil water, while reducing soil and plant nitrogen (N). We conclude that (1) warming may expand B. tectorum's phenological niche, allowing it to more successfully colonize the extensive, invasion‐resistant northern mixed‐grass prairie, and (2) in ecosystems where elevated CO₂ decreases N availability, CO₂ may have limited effects on B. tectorum and other nitrophilic invasive species.     

Soil amendment effects on the exotic annual grass Bromus tectorum L. and facilitation of its growth by the native perennial grass Hilaria jamesii (Torr.) Benth

Greenhouse experiments were undertaken to identify soil factors that curtail growth of the exotic annual grass Bromus tectorum L. (cheatgrass) without significantly inhibiting growth of native perennial grasses (here represented by Hilaria jamesii [Torr.] Benth). We grew B. tectorum and H. jamesii alone (monoculture pots) and together (combination pots) in soil treatments that manipulated levels of soil phosphorus, potassium, and sodium. Hilaria jamesii showed no decline when its aboveground biomass in any of the applied treatments was compared to the control in either the monoculture or combination pots. Monoculture pots of B. tectorum showed a decline in aboveground biomass with the addition of Na₂HPO₄ and K₂HPO_4. Interestingly, in pots where H. jamesii was present, the negative effect of these treatments was ameliorated. Whereas the presence of B. tectorum generally decreased the aboveground biomass of H. jamesii (comparing aboveground biomass in monoculture versus combination pots), the presence of H. jamesii resulted in an enhancement of B. tectorum aboveground biomass by up to 900%. We hypothesize that B. tectorum was able to obtain resources from H. jamesii, an action that benefited B. tectorum while generally harming H. jamesii. Possible ways resources may be gained by B. tectorum from native perennial grasses include (1) B. tectorum is protected from salt stress by native plants or associated soil biota; (2) when B. tectorum is grown with H. jamesii, the native soil biota is altered in a way that favors B. tectorum growth, including B. tectorum tapping into the mycorrhizal network of native plants and obtaining resources from them; (3) B. tectorum can take advantage of root exudates from native plants, including water and nutrients released by natives via hydraulic redistribution; and (4) B. tectorum is able to utilize some combination of the above mechanisms. In summary, land managers may find adding soil treatments can temporarily suppress B. tectorum and enhance the establishment of native plants. However, the extirpation of B. tectorum is unlikely, as many native grasses are likely to facilitate its growth.     

Soil conditioning and plant–soil feedbacks affect competitive relationships between native and invasive grasses

Understanding how competition from invasive species and soil conditions individually and interactively affect native performance will increase knowledge of invasion dynamics and can be used to improve the success of restoration plans. This study, conducted in Reno NV, USA, uses a two-phase plant?Csoil feedback experiment coupled with a target-neighbor competition design to examine the individual and interactive effects of both soil conditions and invasive neighbors on native performance. Study species include invasive species (Bromus tectorum and Agropyron cristatum) and native species (Elymus elymoides and Pseudoroegneria spicata). Results indicate that both plant performance and competitive interactions were influenced by species-specific soil conditioning. Specifically, invasive B. tectorum generated a larger competitive effect on natives than invasive A. cristatum; however, only A. cristatum conditioned soil in a manner that increased competitive effects of conspecifics on natives. Native P. spicata was relatively unaffected by soil conditioning and conversely, E. elymoides was strongly affected by soil conditioning. Few previous studies have examined soil conditioning and the interaction of soil conditioning and neighbor effects that both are potentially important mechanisms in structuring plant communities and influencing plant invasion.     

Fire, native species, and soil resource interactions influence the spatio-temporal invasion pattern of Bromus tectorum

Bromus tectorum (cheatgrass) is an invasive annual that occupies perennial grass and shrub communities throughout the western United States. Bronus tectorum exhibits an intriguing spatio‐temporal pattern of invasion in low elevation ponderosa pine Pinus ponderosa/bunchgrass communities in western Montana where it forms dense rings beneath solitary pines following fire. This pattern provides a unique opportunity to investigate several indirect effects of native vegetation that influence the invasion pattern of B. tectorum, and specifically how native species, disturbance, and soil resources interact to influence the spatio‐temporal pattern of invasion. We established four replicate field sites, each containing burned‐tree, burned‐grass, unburned‐tree, and unburned‐grass sampling locations, and initiated a series of field sampling and greenhouse experiments utilizing these locations. The objective of our first greenhouse experiment was to identify whether belowground factors contributed to the pattern of B. tectorum biomass observed in these field locations. This experiment generated a B. tectorum biomass response that was nearly identical to the invasion pattern observed in the field, suggesting further investigation of belowground factors was necessary. We measured resin‐sorbed NH₄⁺ and NO₃⁻ during one generation of B. tectorum, and measured a suite of P fractions through a sequential extraction procedure from these soils. These data revealed that a resource island of high N and P exists beneath pine trees. Through a second greenhouse experiment, we determined that N limited B. tectorum biomass in tree soil, whereas P limited biomass in bunchgrass soil. Finally, through a germination experiment we determined that pine litter strongly inhibited B. tectorum germination. These data suggest B. tectorum is regulated by P in bunchgrass soil, and by N and inhibition by pine litter beneath trees, effects that are likely alleviated by fire. These data demonstrate the combined role of direct and indirect interactions between native and invasive species in regulating biological invasions.     

Are alien plants more competitive than their native conspecifics? A test using Hypericum perforatum L.

The evolution of increased competitive ability hypothesis predicts that introduced plants that are long liberated from their natural enemies may lose costly herbivore defense, enabling them to reallocate resources previously spent on defense to traits that increase competitive superiority. We tested this prediction by comparing the competitive ability of native St John's wort ( Hypericum perforatum) from Europe with introduced St John's wort from central North America where plants have long grown free of specialist herbivores, and introduced plants from western North America where plants have been subjected to over 57 years of biological control. Plants were grown in a greenhouse with and without competition with Italian ryegrass ( Lolium multiflorum). St John's wort from the introduced range were not better interspecific competitors than plants from the native range. The magnitude of the effect of ryegrass on St John's wort was similar for introduced and native genotypes. Furthermore, introduced plants were not uniformly larger than natives; rather, within each region of origin there was a high variability in size between populations. Competition with ryegrass reduced the growth of St John's wort by >90%. In contrast, St John's wort reduced ryegrass growth <10%. These results do not support the contention that plants from the introduced range evolve greater competitive ability in the absence of natural enemies.     

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.     

Flexible N uptake and assimilation mechanisms may assist biological invasion by Alliaria petiolata

Although nitrogen has historically limited terrestrial plant productivity in the northern hemisphere, accelerated industrial activity is changing the availability of N, with consequences for ecosystem properties including altered susceptibility to biological invasion. Alliaria petiolata (Bieb.) Cavara & Grande is an increasingly problematic invader in forests of eastern North America. Population growth rate of this species is especially high in N-rich habitats, and it produces a variety of N-based compounds that have been shown to interfere with the growth and reproduction of native plants. To investigate how increases and shifts in forms of N will impact A. petiolata, seedlings were transplanted to the greenhouse from the field and grown in sand culture. We applied three concentrations of N (0.25, 1 and 2 mM) using five different ratios of NH₄ ⁺ and NO₃ ⁻ (100/0, 75/25, 50/50, 25/75, 0/100) in a crossed design to yield fifteen different treatments. Plants were measured throughout the growing season and a final harvest yielded measures of biomass and tissue quality. Plant growth increased significantly in response to increased concentration of total N. These increases were similar for all combinations of N. This flexibility in uptake ability may facilitate the invasion of this species, not only by increasing the range of habitats A. petiolata can occupy but also by enhancing N uptake that can lead to the production of secondary compounds disrupting other species’ belowground mutualisms. We suggest that this species’ ability to respond rapidly to changes in N availability, regardless of its form, may modify competitive interactions with natives and intensify its negative impacts.