Restoration of Native Perennials in a California Annual Grassland after Prescribed Spring Burning and Solarization

Grasslands dominated by exotic annual grasses have replaced native perennial vegetation types in vast areas of California. Prescribed spring fires can cause a temporary replacement of exotic annual grasses by native and non‐native forbs, but generally do not lead to recovery of native perennials, especially where these have been entirely displaced for many years. Successful reintroduction of perennial species after fire depends on establishment in the postfire environment. We studied the effects of vegetation changes after an April fire on competition for soil moisture, a key factor in exotic annual grass dominance. As an alternative to fire, solarization effectively kills seeds of most plant species but with a high labor investment per area. We compared the burn to solarization in a study of establishment and growth of seeds and transplants of the native perennial grass Purple needlegrass (Nassella pulchra) and coastal sage species California sagebrush (Artemisia californica). After the fire, initial seed bank and seedling densities and regular percent cover and soil moisture (0–20 cm) data were collected in burned and unburned areas. Burned areas had 96% fewer viable seeds of the dominant annual grass, Ripgut brome (Bromus diandrus), leading to replacement by forbs from the seed bank, especially non‐native Black mustard (Brassica nigra). In the early growing season, B. diandrus dominating unburned areas consistently depleted soil moisture to a greater extent between rains than forbs in burned areas. However, B. diandrus senesced early, leaving more moisture available in unburned areas after late‐season rains. Nassella pulchra and A. californica established better on plots treated with fire and/or solarization than on untreated plots. We conclude that both spring burns and solarization can produce conditions where native perennials can establish in annual grasslands. However, the relative contribution of these treatments to restoration appears to depend on the native species being reintroduced, and the long‐term success of these initial restoration experiments remains to be determined.     

The effect of soil nitrogen on competition between native and exotic perennial grasses from northern coastal California

The invasion of European perennial grasses represents a new threat to the native coastal prairie of northern California. Many coastal prairie sites also experience anthropogenic nitrogen (N) deposition or increased N availability as a result of invasion by N-fixing shrubs. We tested the hypothesis that greater seedling competitive ability and greater responsiveness to high N availability of exotic perennial grasses facilitates their invasion in coastal prairie. We evaluated pairwise competitive responses and effects, and the occurrence of asymmetrical competition, among three common native perennial grasses (Agrostis oregonensis, Festuca rubra, and Nassella pulchra) and three exotic perennial grasses (Holcus lanatus, Phalaris aquatica, and Festuca arundinacea), at two levels of soil N. We also compared the root and shoot biomass and response to fertilization of singly-grown plants, so we could evaluate how performance in competition related to innate plant traits. Competitive effects and responses were negatively correlated and in general varied continuously across native and exotic species. Two exceptions were the exotic species Holcus, which had large effects on neighbors and small responses to them, and competed asymmetrically with all other species in the experiment, and the native grass Nassella, which had strong responses to but little effect on neighbors, and was out-competed by all but one other species in the experiment. High allocation to roots and high early relative growth rate appear to explain Holcus’s competitive dominance, but its shoot biomass when grown alone was not significantly greater than those of the species it out-competed. Competitive dynamics were unaffected by fertilization. Therefore, we conclude that seedling competitive ability alone does not explain the increasing dominance of exotic perennial grasses in California coastal prairie. Furthermore, since native and exotic species responded individualistically, grouping species as ‘natives’ and ‘exotics’ obscured underlying variation within the two categories. Finally, elevated soil N does not appear to influence competition among the native and exotic perennial grasses studied, so reducing soil N pools may not be a critical step for the restoration of California coastal prairie.     

California perennial grasses are physiologically distinct from both Mediterranean annual and perennial grasses

In the Central Valley of California, native perennial grass species have been largely replaced by Eurasian annual species, while in many parts of the Mediterranean Basin native perennial grasses continue to dominate, even on disturbed or degraded sites. We assessed whether differences in summer rainfall patterns have lead to the development of different plant-water strategies between grasses from these two regions. We compared six measures of plant-water physiology for three guilds of grasses: California perennial grasses, Mediterranean perennial grasses, and Mediterranean annual grasses. Discriminant analysis distinguished between the three guilds; Mediterranean perennial grasses were characterized by a more conservative water-relations physiology than Mediterranean annual grasses, whereas California perennial grasses were in some ways intermediate between the two Mediterranean grass guilds. For individual traits, California perennial grasses were either intermediate or more like Mediterranean annuals than Mediterranean perennials. Our results suggest California perennials are more drought tolerant than Mediterranean annuals but less drought tolerant than Mediterranean perennials, despite the fact that California??s Central Valley has a more intense summer drought than the Mediterranean Basin. These patterns may help explain why Mediterranean annuals, but not Mediterranean perennials, have been more successful invaders of interior California grasslands.     

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.     

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.     

Resistance to Centaurea solstitialis invasion from annual and perennial grasses in California and Argentina

A common explanation for Centaurea solstitialis invasion in California is that it occupies an ??empty niche?? created by the replacement of native perennial grasses by exotic annual grasses and concomitant increases in soil water availability. This hypothesis, however, cannot explain C. solstitialis invasion into perennial-dominated grasslands of central Argentina. We assessed invasibility of annual versus perennial grass communities in these regions through parallel field experiments where we created grass plots and, after one year of establishment, measured effects on water and light, and added C. solstitialis seeds in two successive trials. Additionally, we removed vegetation around naturally occurring C. solstitialis in both regions, and examined the performance of Californian and Argentinean C. solstitialis individuals when growing under common conditions simulating climate in California and Argentina. In California, both grass types offered high resistance to C. solstitialis invasion, water was generally greater under perennials than annuals, and light was similarly low beneath both types. In Argentina, invasibility was generally greater in annual than perennial plots, water was similar between groups, and light was much greater beneath annuals. Removal experiments showed that competition from annual grasses in California and perennial grasses in Argentina greatly reduce C. solstitialis performance. Additionally, Californian and Argentinean individuals did not exhibit genetic differentiation in studied traits. Our results suggest that dominant plant functional groups in both California and Argentina offer substantial resistance to C. solstitialis invasion. The success of this species might be tightly linked to a remarkable ability to take advantage of disturbance in both regions.     

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.     

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.     

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.     

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.     

Responses to invasion and invader removal differ between native and exotic plant groups in a coastal dune

The spread of exotic, invasive species is a global phenomenon that is recognized as a major source of environmental change. Although many studies have addressed the effects of exotic plants on the communities they invade, few have quantified the effects of invader removal on plant communities, or considered the degree to which different plant groups vary in response to invasion and invader removal. We evaluated the effects of an exotic succulent, iceplant (Carpobrotus edulis), on a coastal dune plant community in northern California, as well as the community responses to its removal. To assess possible mechanisms by which iceplant affects other plants, we also evaluated its above- and belowground influences on the germination and growth of a dominant exotic annual grass, Bromus diandrus. We found that iceplant invasion was associated with reduced native plant cover as well as increased cover and density of some exotic plants—especially exotic annual grasses. However, iceplant removal did not necessarily lead to a reversal of these effects: removal increased the cover and density of both native and exotic species. We also found that B. diandrus grown in iceplant patches, or in soil where iceplant had been removed, had poorer germination and growth than B. diandrus grown in soil not influenced by iceplant. This suggests that the influence of iceplant on this dune plant community occurs, at least in part, due to belowground effects, and that these effects remain after iceplant has been removed. Our study demonstrates the importance of considering how exotic invasive plants affect not only native species, but also co-occurring exotic taxa. It also shows that combining observational studies with removal experiments can lead to important insights into the influence of invaders and the mechanisms of their effects.     

Reducing Competitive Suppression of a Rare Annual Forb by Restoring Native California Perennial Grasslands

Populations of the rare annual forb Amsinckia grandiflora may be declining because of competitive suppression by exotic annual grasses, and may perform better in a matrix of native perennial bunchgrasses. We conducted a field competition experiment in which Amsinckia seedlings were transplanted into forty 0.64‐m² experimental plots of exotic annual grassland or restored perennial grassland. The perennial grassland plots were restored using mature 3 cm‐diameter plants of the native perennial bunchgrass Poa secunda planted in three densities. The exotic annual grassland plots were established in four densities through manual removal of existing plants. Both grass types reduced soil water potential with increasing biomass, but this reduction was not significantly different between grass types. Both grass types significantly reduced the production of Amsinckia inflorescences. At low and intermediate densities (dry biomass per unit area of 20–80 g/m²), the exotic annual grasses reduced Amsinckia inflorescence number to a greater extent than did Poa, although at high densities (>90 g/m²) both grass types reduced the number of Amsinckia inflorescences to the same extent. The response of Amsinckia inflorescence number to Poa biomass was linear, whereas the same response to the annual grass biomass is logarithmic, and appeared to be related to graminoid cover. This may be because of the different growth forms exhibited by the two grass types. Results of this research suggest that restored native perennial grasslands at intermediate densities have a high habitat value for the potential establishment of the native annual A. grandiflora.     

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.     

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.     

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.     

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.     

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.     

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.     

Stress responses of native and exotic grasses in a Neotropical savanna predict impacts of global change on invasion spread

Communities subject to stress, including those with low invasibility, may be dominated by exotic generalist species. African grasses are aggressive invasive species in Neotropical savannas, where their response to abiotic stress remains unknown. We assessed the role of waterlogging and canopy closure on the presence, abundance and reproductive tillering of African and native grasses in a Neotropical savanna in southeastern Brazil. We obtained abundance and reproductive tillering data of exotic (Melinis minutiflora, Melinis repens and Urochloa decumbens) and common native grasses in 20 sites. We also determined the groundwater depth, soil surface water potential and canopy cover at these sites. The grass species generally had a low frequency and performed poorly where soil remained waterlogged throughout the year, except for two native species. Most native species were exclusive to either well‐drained savannas or better drained wet grasslands. However, two species (Loudetiopsis chrysothrix and Trachypogon spicatus) occurred in both vegetation types. Two exotic species (M. minutiflora and M. repens) were less common but demonstrated reasonable performance in wet grasslands, possibly due to their root system plasticity. Furthermore, U. decumbens had a lower occurrence, density and reproductive tillering at these sites, but was successful at sites where the groundwater level was slightly deeper. Although the favourable water regime in the savannas increases their invasibility in general, resistance to invasion by African grasses may be greater at microsites with high canopy closure, where these species showed lower performance and did not affect the abundance of co‐occurring native grasses. In summary, the Brazilian savanna becomes more susceptible to the spread of African grasses when disturbances decrease canopy closure or lower rainfall associated with climate change reduces the average groundwater depth and consequently releases invasive species from soil waterlogging in grasslands.