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.     

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.     

Effects of perennial semi-arid bunchgrass spatial patterns on performance of the invasive annual cheatgrass (Bromus tectorum L.)

Plant spatial patterns critically influence community dynamics, including plant interactions, resource distribution, and community invasibility. Research suggests that resistance of western US plant communities to further invasion by the exotic annual grass Bromus tectorum may be linked to the positions of, and spacing between, perennial plants. In particular, gaps between aggregated clusters of perennial plants may facilitate B. tectorum invasion by providing safe sites for seed germination and establishment. We tested the effects of random, regular, and aggregated bunchgrass patterns, manipulated at both community (plot) and neighborhood scales, on B. tectorum biomass and spikelet production after experimental seed addition. We found strong evidence of treatment effects on both biomass and spikelets, which varied between treatments by approximately 2.5-fold. Mean biomass and spikelet counts were lowest in plots in which bunchgrasses were aggregated at both community and neighborhood scales, likely due to the increased competition. Although not statistically distinguishable from most other treatments, B. tectorum biomass and spikelet counts were highest in plots with bunchgrass patterns that were random at the community scale and aggregated at the neighborhood scale. These plots were characterized by relatively large gaps between bunchgrass clusters, suggesting that B. tectorum may exploit gaps between aggregated perennial plants. Our results support the emerging hypothesis that community resistance to B. tectorum invasion could be increased through manipulation of perennial vegetation to reduce basal gap size and connectivity.     

Experimental removal of intraspecific competitors — effects on water relations and productivity of a desert bunchgrass,Hilaria rigida

Summary Intraspecific competition in the C₄ bunchgrass Hilaria rigida was examined on a Sonoran Desert site in southeastern California. Potential competition within monospecific stands was experimentally altered by removal of the aboveground portions of all plants within a 1.5 m radius of a monitored plant. Compared with unaltered plots, altered plots had less negative soil water potentials during periods of soil drying. Leaf blades on monitored plants of altered plots remained green longer and had greater stomatal conductances than those on monitored plants on unaltered plots. Production of new culms was twice as great on altered plots. Greater root biomass and root length were observed in altered plots, and root extension into soil areas formerly occupied by roots of neighboring plants occurred within one year after treatment. The results indicate that removal of the aboveground biomass of neighboring plants reduces the competition for limited available soil water in this desert environment.     

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.     

Evaluation of Herbicides for Restoring Native Grasses in Buffelgrass-Dominated Grasslands

Buffelgrass ( Pennisetum ciliare ) is an exotic grass that threatens arid and semiarid ecosystems. The objective of this study was to determine effectiveness of several herbicides at reducing competition from buffelgrass to enhance establishment of planted native grasses. In Duval County, Texas, plots were delineated in two experiments in a buffelgrass-dominated pasture and mowed on 2 September 2002. On 18 September 2002 and 7 October 2002, a 41% glyphosate (N-(phosphonomethyl) glycine) herbicide was applied to all plots. A mixture of three native grasses—green sprangletop ( Leptochloa dubia ), plains bristlegrass ( Setaria leucopila ), and four-flower trichloris ( Chloris pluriflora )—was planted on 8 October 2002. On 9 October 2002, 1.12 and 2.24 kg/ha of a 80% tebuthiuron ( N -[5-(1,1-dimethyethyl)-1,3,4-thiadiazol-2-yl]- N , N '-dimethylurea) herbicide was applied preemergence to the first experiment, and all other herbicides were applied postemergence on 27 July 2003 to the second experiment. Percent canopy cover of vegetation was estimated with a 20 × 50–cm sampling frame during April, June, and October 2003 and August 2004. Postemergent herbicides had no significant effect on canopy cover of buffelgrass or planted species ( p ≥ 0.05). Canopy cover of native grasses did not exceed 8% on any treatment or sampling date, and buffelgrass cover returned to pre-treatment conditions in less than 1 year; however, the 2.24 kg/ha rate of tebuthiuron suppressed ( p < 0.05) canopy cover of buffelgrass compared with controls and increased ( p < 0.05) native grasses almost 2 years past application. Tebuthiuron may have potential value in reducing buffelgrass canopy cover and increasing cover of native grasses, particularly Chloris spp.     

Effects of Phenology at Burn Time on Post‐Fire Recovery in an Invasive C4 Grass

The spread of non‐indigenous, C₄ grasses threatens global conservation of savannas and subtropical grasslands. Identifying control methods to selectively target these invasives has proven difficult. Here, we tested the hypothesis that the effectiveness of prescribed burns for control is determined, in part, by the phenology of the target species at burn time. We conducted two experiments in a subhumid, C₄ grassland in central Texas. The focal invasive was the C₄, perennial bunchgrass Bothriochloa ischaemum (L.) Keng (KR bluestem). Burns were conducted in early and late fall when plants were in different phenological states. In addition, we attempted to manipulate phenological state through temporary rainout shelters to expedite maturation. The two experiments differed in the timing of the rainout shelter application (experiment 1: May to July, experiment 2: August and September), but otherwise had the same complete factorial design (burn time × shelter). Across experiments, when at least 50% of all tillers were pre‐reproductive at burn time, either due to shelter treatment or time of year, spring tiller densities were significantly lower than when plants were burned in a more advanced reproductive state. Trends in fall biomass generally followed trends in spring tiller densities, with one exception where plants in no‐shelter plots burned in October had lower biomass than expected based on tiller densities. Treatment responses for the native C₄ grass B. laguroides were consistent with those of B. ischaemum. These findings suggest that strategic burns can be used to reduce the subsequent recovery of invasive C₄ grasses while not disadvantaging native grasses.     

Restoring a Mediterranean‐climate shrub community with perennial species reduces future invasion

Successful restoration of an invaded landscape to a diverse, invasion‐resistant native plant community requires determining the optimal native species mix to add to the landscape. We manipulated native seed mix (annuals, perennials, or a combination of the two), while controlling the growth of non‐native species to test the hypothesis that altering native species composition can influence native establishment and subsequent non‐native invasion. Initial survival of native annuals and perennials was higher when seeded in separate mixes than when combined, and competition between the native perennials and annuals led to lower perennial cover in year 2 of mixed‐seeded plots. The plots with the highest perennial cover had the highest resistance to invasion by Brassica nigra. To clarify interactions among different functional groups of natives and B. nigra, we measured competitive interactions in pots. We grew one native annual, one native perennial, and B. nigra alone or with different competitors and measured biomass after 12 weeks. Brassica nigra was the strongest competitor, limiting the growth of all native species, and was not impacted by competition with native annuals or perennial seedlings. Results from the potted plant experiment demonstrated the strong negative influence of B. nigra on native seedlings. Older native perennials were the strongest competitors against invasive species in the field, yet perennial seedling survival was limited by competition with native annuals and B. nigra. Management action that maximizes perennial growth in early years may lead to a relatively more successful restoration and the establishment of an invasion‐resistant community.     

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

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

Nutrient enrichment enhances hidden differences in phenotype to drive a cryptic plant invasion

Many mechanisms of invasive species success have been elucidated, but those driving cryptic invasions of non‐native genotypes remain least understood. In one of the most successful cryptic plant invasions in North America, we investigate the mechanisms underlying the displacement of native Phragmites australis by its Eurasian counterpart. Since invasive Phragmites’ populations have been especially prolific along eutrophic shorelines, we conducted a two‐year field experiment involving native and invasive genotypes that manipulated nutrient level and competitor identity (inter‐ and intra‐genotypic competition) to assess their relative importance in driving the loss of native Phragmites. Inter‐genotypic competition suppressed aboveground biomass of both native and invasive plants regardless of nutrient treatment (～ 27%), while nutrient addition disproportionately enhanced the aboveground biomass (by 67%) and lateral expansion (by > 3 × farther) of invasive Phragmites. Excavation of experimental plots indicated that nutrient addition generates these differences in aboveground growth by differentially affecting rhizome production in invasive vs native plants; invasive rhizome biomass and rhizome length increased by 595% and 32% with nutrient addition, respectively, while natives increased by only 278% and 15%. Regardless of nutrient level, native rhizomes produced twice as many roots compared to invasives, which field surveys revealed are heavily infected with mycorrhizal symbionts. These results suggest that native Phragmites competes well under nutrient‐limited conditions because its rhizomes are laden with nutrient‐harvesting roots and mycorrhizae. Invasive Phragmites’ vigorous aboveground response to nutrients and scarcity of lateral roots, in contrast, may reflect its historic distribution in eutrophic Eurasian wetlands and correspond to its prevalence in New England marshes characterized by elevated nutrient availability and relaxed nutrient competition. These findings reveal that discrete differences in phenotype can interact with anthropogenic modification of environmental conditions to help explain the success of cryptic invaders.     

Competitive Control of an Exotic Mangrove Species: Restoration of Native Mangrove Forests by Altering Light Availability

Recently, many studies have focused on the possibility of restoring mangrove ecosystems by introducing fast‐growing mangroves. However, methods for managing an exotic fast‐growing species to restore mangrove ecosystems and at the same time preventing invasion by introduced species remains unclear. Sonneratia apetala Buch‐Ham is one example of an exotic mangrove with both high ecological value and potential risk for invasion after introduction. To investigate the possibility of reducing the potential for invasion by altering light availability, we simulated different irradiances of S. apetala understory in the greenhouse. For each irradiance treatment, three levels of competition between S. apetala and native mangroves Aegiceras corniculatum (L.) were used: no competition, intraspecific competition and interspecific competition. Compared with A. corniculatum, S. apetala showed a significantly higher growth rate for both height and biomass accumulation under full irradiation. Compared to the full irradiation treatment, the shading treatment significantly reduced the height, total biomass and biomass allocation to leaves of S. apetala by 61.31, 71.0, and 76.2%, respectively, whereas the growth of A. corniculatum was not affected. The results suggested that lowering light availability could inhibit the growth of S. apetala and increase the competitiveness of A. corniculatum. Planting introduced fast‐growing mangroves at a density of approximately 2,000 plants/hm² is an effective strategy for preventing potential invasion and restoring wetland habitats. By taking advantage of the differences in shade tolerance between fast‐growing exotic mangroves and native mangroves, introduction of fast‐growing mangroves in coastal areas could have huge potential for reforesting mangrove ecosystems.     

Mycorrhizae transfer carbon from a native grass to an invasive weed: evidence from stable isotopes and physiology

Invasive exotic weeds pose one of the earth's most pressing environmental problems. Although many invaders completely eliminate native plant species from some communities, ecologists know little about the mechanisms by which these exotics competitively exclude other species. Mycorrhizal fungi radically alter competitive interactions between plants within natural communities, and a recent study has shown that arbuscular mycorrhizal (AM) fungi provide a substantial competitive advantage to spotted knapweed, Centaurea maculosa, a noxious perennial plant that has spread throughout much of the native prairie in the northwestern U.S. Here we present evidence that this advantage is potentially due to mycorrhizally mediated transfer of carbon from a native bunchgrass, Festuca idahoensis, to Centaurea. Centaurea maculosa, Festuca idahoensis (Idaho fescue, C₃), and Bouteloua gracilis (blue gramma, C₄) were grown in the greenhouse either alone or with Centaurea in an incomplete factorial design with and without AM fungi. Centaurea biomass was 87–168% greater in all treatments when mycorrhizae were present in the soil (P < 0.0001). However, Centaurea biomass was significantly higher in the treatment with both mycorrhizae and Festuca present together than in any other treatment combination (P < 0.0001). This high biomass was attained even though Centaurea photosynthetic rates were 14% lower when grown with Festuca and mycorrhizae together than when grown with Festuca without mycorrhizae. Neither biomass nor photosynthetic rates of Centaurea were affected by competition with the C₄ grass Bouteloua either with or without mycorrhizae. The stable isotope signature of Centaurea leaves grown with Festuca and mycorrhizae was more similar to that of Festuca, than when Centaurea was grown alone with mycorrhizae (P = 0.06), or with Festuca but without mycorrhizae (P = 0.09). This suggests that carbon was transferred from Festuca to the invasive weed. We estimated that carbon transferred from Festuca by mycorrhizae contributed up to 15% of the aboveground carbon in Centaurea plants. Our results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.     

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.     

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.     

Promoting Native Vegetation and Diversity in Exotic Annual Grass Infestations

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

Burning and Grazing Management in a California Grassland: Growth, Mortality, and Recruitment of Nassella pulchra

Abstract Annual grasslands in California are often managed with seasonal grazing and prescribed burning on the assumption that such practices have long‐term benefits for native species. Mature native perennial bunchgrasses, particularly Nassella pulchra (purple needlegrass), are often the focal species, although very little is known about responses at different life history stages. Thus, important questions remain about long‐term population dynamics of both mature plants and seedling recruitment. In plots receiving repeated grazing and burning events over 7 years, mortality of mature plants was threefold higher on mounds than on intermounds and likely reflected increased competition intensity associated with increased resource availability in deeper soil. Burning and grazing treatments had strong positive effects on basal area of mature N. pulchra. However, plants in grazed plots that were not burned contained considerable standing dead biomass. Topographic location strongly influenced growth as intermound plants grew relatively more than mound plants, but the effects on growth of burning and grazing did not vary with topographic location. In mapped plots N. pulchra recruitment was very low, and overall density dropped an average of 31%. However, a significant time‐by‐burning effect indicated that survival was significantly higher in burned plots. After 7 years of repeated treatments, effects of burning and grazing management on mature N. pulchra were positive but not for all phenological stages. Understanding long‐term influence of management on bunchgrass populations may not be easy to determine because short‐term results may not reflect long‐term responses and some life cycle dynamics may be observed only over very long periods.     

本地和外来草本物种对水分条件时间异质性的可塑响应

极端气候导致的干旱和水淹事件频发，影响了外来植物和本地植物的生长。为了解外来种和本地种植物对干旱和水淹事件发生顺序的响应，探讨草本植物适应水分时间异质性的策略，该文以美国蒙大拿州西部4种本地植物和4种外来植物为研究对象，将所有植物分别进行持续湿润(对照，CK)、水淹-干旱(I-D)和干旱-水淹(D-I)处理，并观测一系列形态和生物量特征的变化。结果表明：(1)与CK相比，D-I和I-D处理均显著降低了外来种的总生物量(P<0.05)。(2)D-I显著降低了本地种早期总生物量、后期地下生物量和根冠比，但显著提高了其后期的相对生长(P<0.05)。(3)D-I处理显著降低了所有植物的地下-地上生物量关系的异速指数，外来种异速指数显著高于本地种(P<0.05)。综上认为，极端事件(水淹和干旱)的发生顺序能改变外来植物和本地植物的生物量分配，早期干旱比后期干旱更容易减少植物生物量的积累，但能促进本地种后期的生长；本地种在环境胁迫下不被降低的总生物量表现说明维持表型稳定的能力较强；D-I处理下本地种和外来种地上和地下生物量关系的分配方式不同。     

Effects of perennial neighbors and nitrogen pulses on growth and nitrogen uptake by Bromus tectorum

An experiment was conducted to determine if growth and biomass responsesof the annual grass Bromus tectorum are affected by themagnitude and timing of nitrogen (N) pulses and if these responses areinfluenced by different perennial neighbor species. Nitrogen(NH₄:NO₃) was applied in three pulse treatments of varyinginterpulse length (3-d, 9-d, or 21-d between N additions). The total amount of Nadded was the same among treatments; hence, both the frequency and magnitude ofN pulses varied (i.e., the longer the interpulse period,the greater the amount of N added for a single pulse).Bromus showed little response to the different N-pulsetreatments. The only characteristic that varied among pulse treatments wasspecific leaf area (SLA), which was significantly greater whenBromus was grown under the 21-d N pulse than when grownunder the 3-d or 9-d N pulses. Bromus height, leaf andtiller numbers, leaf area and aboveground biomass were not affected by theN-pulse treatments nor were tissue-N contents and concentrations. However,Bromus production and tissue-N were significantly differentwhen Bromus was grown with different perennial neighborspecies. Tiller production, aboveground biomass, and seed numbers ofBromus were lowest when the perennial neighbor was thetussock grass Agropyron desertorum, intermediate when theneighbor was the evergreen shrub Artemisia tridentata, andgreatest when the neighbor was the deciduous shrub Chrysothamnusnauseosus. N contents of Bromus leaves were alsolowest when the neighbor was Agropyron. In contrast, root Nuptake capacities were greatest for Agropyron-Bromus rootmixes and lowest for Chrysothamnus-Bromus root mixes. Theseresults suggest that perennial neighbors affect growth, seed production, and Nuptake of Bromus to a greater extent than the timing andmagnitude of N pulses.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Invasion resistance and persistence: established plants win,even with disturbance and high propagule pressure

Disturbances and propagule pressure are key mechanisms in plant community resistance to invasion, as well as persistence of invasions. Few studies, however, have experimentally tested the interaction of these two mechanisms. We initiated a study in a southwestern ponderosa pine (Pinus ponderosa Laws.)/bunch grass system to determine the susceptibility of remnant native plant communities to cheatgrass (Bromus tectorum L.) invasion, and persistence of cheatgrass in invaded areas. We used a 2 × 2 factorial design consisting of two levels of aboveground biomass removal and two levels of reciprocal seeding. We seeded cheatgrass seeds in native plots and a native seed mixture in cheatgrass plots. Two biomass removal disturbances and sowing seeds over 3 years did not reverse cheatgrass dominance in invaded plots or native grass dominance in non-invaded native plots. Our results suggest that two factors dictated the persistence of the resident communities. First, bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey) was the dominant native herbaceous species on the study site. This species is typically a poor competitor with cheatgrass as a seedling, but is a strong competitor when mature. Second, differences in pretreatment levels of plant-available soil nitrogen and phosphorus may have favored the dominant species in each community. Annual species typically require higher levels of plant-available soil nutrients than perennial plants. This trend was observed in the annual cheatgrass community and perennial native community. Our study shows that established plants and soil properties can buffer the influences of disturbance and elevated propagule pressure on cheatgrass invasion.