Biotic constraints on the invasion of diffuse knapweed (Centaurea diffusa) in North American grasslands

Knapweeds (Centaurea spp.) are among the most invasive of non-indigenous plant species that have colonized western North America over the last century. We conducted a 4-year experiment in a reconstructed grassland to test hypotheses related to the ability of grasslands to resist the invasion of diffuse knapweed (C. diffusa). We experimentally invaded C. diffusa and three native species into areas where we manipulated soil nitrogen (N) and phosphorus (P) availability and removed extant grasses to reduce competition. We evaluated the growth response of these species to these resources and competitive manipulations. Of the native species that were experimentally added, only one species, Ratibida pinnata (prairie coneflower), established in any numbers. Establishment values in intact vegetation were low for both species, but establishment by C. diffusa (0.02%) clearly outperformed that of R. pinnata (0.001%). Under reduced grass competition, establishment was enhanced, but the values for C. diffusa (0.68%) were not statistically different from those of R. pinnata (0.57%). Neither species performed better under higher soil nutrients in the presence of competing grasses. In plots with both species, biomass of the two planted species was positively correlated, but the biomass of both species was negatively correlated with non-added weedy species. Subsequent harvests of C. diffusa indicated that establishment was enhanced in treatments with higher soil nutrients but that the biomass of these plants could only be enhanced when plant competition was also reduced. These results indicate that C. diffusa can establish in intact grasslands at rates higher than natives, but opportunism rather than competitive ability best describes the invasiveness of C. diffusa. Thus, the mechanisms contributing to the establishment of this knapweed species are different from factors identified as contributing to the dominance of this invader.     

The effects of size of opening in vegetation and litter cover on seedling establishment of goldenrods (Solidago spp.)

We investigated the effects of size of opening in the vegetation and litter cover on seedling establishment of two species of goldenrods (Solidago spp.) in an abandoned field in southwestern Michigan, U.S.A. Seeds of S. canadensis and S. juncea were sown into clipped plots, ranging from 0 cm (control, unclipped) to 100 cm in diameter, with and without litter. Seedling emergence, survival and growth were followed for one year. Soil moisture was not significantly different among the opening sizes, but, within a size, tended to be lower when litter was removed. Light intensity at the soil surface was positively related to opening size early in the growing season, but later in the growing season reached a maximum in intermediate-sized openings and then leveled off. Litter strongly inhibited seedling emergence in both species. Emergence of S. canadensis seedlings was lower in 0 and 10 cm openings than in the larger openings, while emergence of S. juncea seedlings was lower in the largest openings (100 cm) than in all the smaller openings. In contrast, seedling growth and probability of survival increased with diameter of opening for both species. Some seedlings of S. juncea did survive in complete vegetation cover (controls, 0 cm openings) while seedlings of S. canadensis survived only in openings of at least 30 cm diameter. Thus, S. juncea had a smaller minimum opening size for seedling establishment than S. canadensis, although both species grew and survived best in the largest openings made in the experiment.     

How do biological control and hybridization affect enemy escape?

Two mechanisms often linked with plant invasions are escape from enemies and hybridization. Classical biological control aims to reverse enemy escape and impose top-down population control. However, hybridization has the potential to alter interactions with enemies and thus affect biological control. We examined how introductions of biological control agents affect enemy escape by comparing specialist enemy loads between the native and introduced ranges of two noxious weeds (spotted and diffuse knapweed; Centaurea stoebe L. and C. diffusa Lam.) that have been the targets of an extensive biological control program. Hybrids between spotted and diffuse knapweed are often found within diffuse knapweed sites in North America, so we also compared enemy loads on plants that appeared morphologically like diffuse knapweed and hybrids. Finally, we tested the preference for diffuse knapweed, hybrids, and spotted knapweed of one of the agents thought to be instrumental in control of diffuse knapweed (Larinus minutus; Curculionidae). In North America spotted knapweed has largely escaped its root herbivores, while seedhead herbivore loads are comparable in the introduced and native ranges. Diffuse knapweed exhibited seedhead herbivore loads five times higher in the introduced compared to native range. While this pattern of seedhead herbivory is expected with successful biological control, increased loads of specialist insect herbivores in the introduced range have rarely been reported in the literature. This finding may partially explain the better population control of diffuse vs. spotted knapweed. Within North American diffuse knapweed sites, typical diffuse knapweed and hybrid plants carried similar herbivore loads. However, in paired feedings trials, the specialist L. minutus demonstrated a preference for newly created artificial hybrids over North American diffuse knapweed and for European diploid spotted knapweed over North American tetraploid spotted knapweed. Overall though, hybridization does not appear to disrupt biological control in this system.     

Field cage assessment of interference among insects attacking seed heads of spotted and diffuse knapweed

Field studies were conducted to determine the competitive interactions between introduced biological control agents that attack the seed heads of spotted knapweed (Centaurea stoebe ssp. micranthos) and diffuse knapweed (Centaurea diffusa). Two weevils, Bangasternus fausti and Larinus minutus (Coleoptera: Curculionidae), were each paired with the previously established fly, Urophora affinis (Diptera: Tephritidae). Each species was released either alone or in pair-wise combinations inside screen cages placed over existing knapweed plants at six field sites in Montana and one in Oregon. Larinus minutus produced almost three times as many progeny on diffuse knapweed as on spotted knapweed. Larinus minutus reproduction was not affected by competition with U. affinis, but U. affinis reproduction was reduced by the presence of L. minutus (by 71% on spotted and 77% on diffuse knapweed). Bangasternus fausti reproduction generally was not affected by competition with U. affinis, nor was U. affinis affected by B. fausti on either host plant. There were extremely few cases of successful production of both weevil and fly in the same capitulum, which was probably because weevil larvae consume the developing flies. Both weevils increased the total proportion of seed heads infested on diffuse knapweed, and B. fausti increased it on spotted knapweed. However, the release of either weevil did not significantly further reduce seed production on either plant. The results and experimental design are discussed in light of the subsequent establishment and impact of these agents.     

Honey bees and wild pollinators differ in their preference for and use of introduced floral resources

Introduced plants may be important foraging resources for honey bees and wild pollinators, but how often and why pollinators visit introduced plants across an entire plant community is not well understood. Understanding the importance of introduced plants for pollinators could help guide management of these plants and conservation of pollinator habitat. We assessed how floral abundance and pollinator preference influence pollinator visitation rate and diversity on 30 introduced versus 24 native plants in central New York. Honey bees visited introduced and native plants at similar rates regardless of floral abundance. In contrast, as floral abundance increased, wild pollinator visitation rate decreased more strongly for introduced plants than native plants. Introduced plants as a group and native plants as a group did not differ in bee diversity or preference, but honey bees and wild pollinators preferred different plant species. As a case study, we then focused on knapweed (Centaurea spp.), an introduced plant that was the most preferred plant by honey bees, and that beekeepers value as a late‐summer foraging resource. We compared the extent to which honey bees versus wild pollinators visited knapweed relative to coflowering plants, and we quantified knapweed pollen and nectar collection by honey bees across 22 New York apiaries. Honey bees visited knapweed more frequently than coflowering plants and at a similar rate as all wild pollinators combined. All apiaries contained knapweed pollen in nectar, 86% of apiaries contained knapweed pollen in bee bread, and knapweed was sometimes a main pollen or nectar source for honey bees in late summer. Our results suggest that because of diverging responses to floral abundance and preferences for different plants, honey bees and wild pollinators differ in their use of introduced plants. Depending on the plant and its abundance, removing an introduced plant may impact honey bees more than wild pollinators.     

Is spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis> L.) patch size related to the effect on soil and vegetation properties?

Spotted knapweed (Centaurea stoebe L. subsp. Micranthos (Gugler) Hayek) was first introduced in the 1890s from Europe into western North America, where it now occupies over three million hectares of rangeland and pasture in 14 states and two Canadian provinces, reducing forage production and causing economic damage. Despite many reported effects spotted knapweed can have on soils and native vegetation, it is not known whether patch size is correlated with these ecosystem-level effects. The objective of our study was to determine whether the effects of spotted knapweed on plant composition and soil properties was related to spotted knapweed patch size. We asked the following questions: (1) Are there differences in plant species richness and diversity between small and large knapweed patches? and (2) Do soil water and soil mineral nutrient properties change depending on knapweed patch size? Twenty-four knapweed patches, and paired natural grassland plots, were randomly selected within Lac du Bois Provincial Park, British Columbia, Canada. Knapweed patch size ranged from 6 to 366 m². Sampling and analysis revealed a significant effect of knapweed patch size on soil and vegetation properties. Soil P, soil temperature, and total dry plant biomass (g/0.25 m²) increased, while soil N, soil C, and soil moisture decreased with patch size. Since our results show that spotted knapweed patch size is related to degree of soil alteration, it is important to consider size of patch when modeling the impact of spotted knapweed in North America. Since large patches of spotted knapweed seem to have a proportionately greater effect on soil chemistry properties, large patches may move the system further away from a point where it is possible to restore the site to pre-invasion conditions.     

A knapweed biological control perspective

Diffuse and spotted knapweed (Centaurea diffusa Lam. and C. stoebe micranthos (Gugler) Hayek) are Eurasian plants that devastate dry and mesic North American grasslands. They have a mutualistic association with arbuscular mycorrhizal fungal (AMF) phylotypes with hyphal links to nearby plants and a nutrient flux to the strongest sink, usually knapweed. They displace many AMF beneficial to grass and affect knapweed nutrient allocation, biology, knapweed insects and probably root necrosis and emergence of ant buried seed. AMF determined nutrient root or shoot allocation determines nutrient shoot and root allocation and the benefit to root or seed-head insect species and whether C. diffusa is an annual–biannual or a semelparous perennial needing 5 or more years to flower. Both knapweeds do well without its AMF phylotypes without competition in fertile soil. In grass in Eurasia, they have a community of seven seed-head species segregated by head development stage. Prolonged seed dormancy buffered knapweed decline that resulted in release of a surfeit seed-head species. The presence of an eliasome on the seed and vigorous seedling clumps suggests burial by myrmecochorous ants with AMF supplied carbon supporting their growth. The root species community is segregated by habitat, climate, root part, and size. With larval induced compensatory growth and AMF nutrient sharing, the growth of plants with and without a larva was the same. On feeding completion, a nutrient out flux from the attacked plants reduced growth; but without killing. This needs a dual species or a repeated single species attack. Root species packing increases knapweed utilization; but the four approved species are insufficient for maximum utilization. Two additions are suggested. The aim of the paper is to provide enough understanding of the AMF and its plant and insect interactions to facilitate knapweed biological control and avoid past mistakes.     

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.     

Impact of clear-cutting and prescribed burning on microbial diversity and community structure in a Jack pine (Pinus banksiana Lamb.) clear-cut using Biolog Gram-negative microplates

Three isolates ofFusarium avenaceum are pathogenic on spotted knapweed(Centaurea maculosa), a major weed plant of pasturelands and rangelands of the Pacific Northwestern USA. One isolate (no. 1) obtained from the European centre of origin of knapweed and isolate no. 365 native to Montana, did not significantly affect knapweed seed germination. However,F. avenaceum no. 1003, another Montana native isolate, caused a 100% decrease in seed germination and hence, no seedling emergence. When formulated, isolate no. 1003, could be recovered from treated soils after 7 days and caused a significant reduction in seedling emergence or seedling dry weight. This organism had no effect on the germination ofTriticum aestivum orMedicago sativa, but did affect the germination of other plant species.F. avenaceum appears to be a candidate for the biocontrol of spotted knapweed, however, a native isolate is potentially more effective than an isolate obtained from the centre of origin ofC. maculosa.     

Mid-Spring Burning Reduces Spotted Knapweed and Increases Native Grasses during a Michigan Experimental Grassland Establishment

Infestations of the exotic perennial Spotted knapweed (Centaurea maculosa Lam.) hinder the restoration and management of native ecosystems on droughty, infertile sites throughout the Midwestern United States. We studied the effects of annual burning on knapweed persistence on degraded, knapweed‐infested gravel mine spoils in western Michigan. Our experiment included 48, 4‐m² plots seeded to native warm‐season grasses in 1999 using a factorial arrangement of initial herbicide and fertility treatments. Beginning in 2003, we incorporated fire as an additional factor and burned half of the plots in late April or May for 3 years (2003–2005). Burning increased the dominance of warm‐season grasses and decreased both biomass and dominance of knapweed in most years. Burning reduced adult knapweed densities in all 3 years of the study, reduced seedling densities in the first 2 years, and reduced juvenile densities in the last 2 years. Knapweed density and biomass also declined on the unburned plots through time, suggesting that warm‐season grasses may effectively compete with knapweed even in the absence of fire. By the end of the study, mean adult knapweed densities on both burned (0.4‐m²) and unburned (1.3‐m²) plots were reduced to levels where the seeded grasses should persist with normal management, including the use of prescribed fire. These results support the use of carefully timed burning to help establish and maintain fire‐adapted native plant communities on knapweed‐infested sites in the Midwest by substantially reducing knapweed density, biomass, and seedling recruitment and by further shifting the competitive balance toward native warm‐season grasses.     

Herbivory and novel weapons: no evidence for enhanced competitive ability or allelopathy induction of <Emphasis Type="Italic">Centaurea diffusa</Emphasis> by biological controls

Biological control of weeds by arthropod herbivores is thought to work by reducing the competitive ability of the weed relative to the surrounding vegetation. However, the assumption that herbivory reduces plant competitive ability has not been tested in most biological control systems, and counter to expectation, recent research on the impact of biological control agents on invasive Centaurea species suggests that this genus may respond to herbivory by increased competitive ability through enhanced plant re-growth and/or by inducing increased production of phytotoxic allelochemicals. We examined the impact of two biological control agents of the invasive plant diffuse knapweed (C. diffusa) to see if feeding by either of these insects would enhance the plant’s competitive ability or allelochemical output. Sub-lethal herbivory by either of the biological control agents significantly reduced knapweed performance when the plant was grown in competition with either of two native species. Competition with knapweed significantly reduced the performance of both native species (Artemisia frigida and Bouteloua gracilis), and herbivory by one of the biocontrol agents resulted in a small but significant increase in both native species’ performance. Diffuse knapweed’s putative allelochemical 8-hydroxyquinoline was not detected in experimental or field collected soils from knapweed-infested sites. In contrast to other studies on the impacts of biological control on other Centaurea species, these data support the premise that biological control agents may reduce invading plant competitive ability. We find no evidence for diffuse knapweed allelopathy mediated by 8-hydroxyquinoline or enhanced allelopathy in response to herbivory by biological control agents.     

Effects of Nitrogen and Salinity on Growth and Competition Between a Native Grass and an Invasive Congener

Numerous studies show that an increase in the availability of limiting resources can increase invasion by non-native plants into natural communities. One possible explanation is that the ability of natives to compete with non-natives tends to decrease when resource availability is increased. We tested this hypothesis in a competition experiment using two closely matched plant species and two environmental factors related to limiting resources in a coastal grassland system on Bodega Head in northern California. We grew the native grass Bromus carinatus and the non-native grass B. diandrus together and apart at different levels of soil nitrogen crossed with different levels of soil salinity. Both species are abundant in the grassland and previous work suggested that the abundance of B. carinatus is lower and the abundance of B. diandrus is higher on soil that has been enriched with nitrogen. Salinity has been shown to be negatively associated with invasion by B. diandrus into another California grassland, and to vary significantly over short distances in the grassland at Bodega Head, where it could affect water availability, which strongly limits plant growth during the dry season. Contrary to our prediction that low resource availabilities would increase the relative competitive ability of the native, the ability of B. carinatus to compete with B. diandrus was not greater when nitrogen availability was lower or when soil salinity was higher. Instead, high salinity increased the relative competitive ability of the non-native, and low nitrogen had little effect on competition. This suggests that preventing resource enrichment will not suffice to control invasion by non-native plant species in this grassland.     

The invasive grass,Melinis minutiflora,inhibits tree regeneration in a Neotropical savanna

Abstract Exotic grasses are becoming increasingly abundant in Neotropical savannas, with Melinis minutiflora Beauv. being particularly invasive. To better understand the consequences for the native flora, we performed a field study to test the effect of this species on the establishment, survival and growth of seedlings of seven tree species native to the savannas and forests of the Cerrado region of Brazil. Seeds of the tree species were sown in 40 study plots, of which 20 were sites dominated by M. minutiflora, and 20 were dominated by native grasses. The exotic grass had no discernable effect on initial seedling emergence, as defined by the number of seedlings present at the end of the first growing season. Subsequent seedling survival in plots dominated by M. minutiflora was less than half that of plots dominated by native species. Consequently, at the end of the third growing season, invaded plots had only 44% as many seedlings as plots with native grasses. Above‐ground grass biomass of invaded plots was more than twice that of uninvaded plots, while seedling survival was negatively correlated with grass biomass, suggesting that competition for light may explain the low seedling survival where M. minutiflora is dominant. Soils of invaded plots had higher mean Ca, Mg and Zn, but these variables did not account for the higher grass biomass or the lower seedling survival in invaded plots. The results indicate that this exotic grass is having substantial effects on the dynamics of the tree community, with likely consequences for ecosystem structure and function.     

Establishment and Management of Native Functional Groups in Restoration

The limiting similarity hypothesis predicts that communities should be more resistant to invasion by non‐natives when they include natives with a diversity of traits from more than one functional group. In restoration, planting natives with a diversity of traits may result in competition between natives of different functional groups and may influence the efficacy of different seeding and maintenance methods, potentially impacting native establishment. We compare initial establishment and first‐year performance of natives and the effectiveness of maintenance techniques in uniform versus mixed functional group plantings. We seeded ruderal herbaceous natives, longer‐lived shrubby natives, or a mixture of the two functional groups using drill‐ and hand‐seeding methods. Non‐natives were left undisturbed, removed by hand‐weeding and mowing, or treated with herbicide to test maintenance methods in a factorial design. Native functional groups had highest establishment, growth, and reproduction when planted alone, and hand‐seeding resulted in more natives as well as more of the most common invasive, Brassica nigra. Wick herbicide removed more non‐natives and resulted in greater reproduction of natives, while hand‐weeding and mowing increased native density. Our results point to the importance of considering competition among native functional groups as well as between natives and invasives in restoration. Interactions among functional groups, seeding methods, and maintenance techniques indicate restoration will be easier to implement when natives with different traits are planted separately.     

Native Consumers and Seed Limitation Constrain the Restoration of a Native Perennial Grass in Exotic Habitats

Native consumers and seed limitation may be particularly important in the restoration of native plants where they have been displaced by exotic plants. We used experimental exclosures and seed additions to examine the role of native mammalian consumers and seeding density (500 or 1,000 seeds/m²) in affecting the establishment of a native perennial grass, Purple needlegrass ( Nassella pulchra ), in the grasslands of California. To focus solely on consumers and propagule density, experimental areas were tilled and weeded. Consumers were important determinants of restoration success: averaged across propagule density, consumers reduced N. pulchra seedling recruitment by nearly 30%, reduced seedling height by 44%, reduced plant establishment by 52%, and reduced reproductive tiller length by 43%. Small rodents affected seedling establishment, especially where seeding density was high but did not affect seedling height. Plots accessible by squirrels and rabbits exhibited significantly decreased seedling height and plant establishment, whereas there was no additional impact of allowing large consumers (i.e., deer) access. Despite strong, spatially variable effects of consumers, doubling seed density led to nearly doubled N. pulchra establishment on average. Consumer effects were persistent, shaping N. pulchra abundance in the subsequent growing season and remaining evident over 18 months after the experiment was initiated. Our work suggests that, despite strong consumer effects, seed addition may be a viable strategy for restoration of N. pulchra in invaded areas where it has been displaced by exotic plants, especially when combined with restoration strategies that reduce competition with exotic plants.     

Nitrogen deposition enhances Bromus tectorum invasion: biogeographic differences in growth and competitive ability between China and North America

Increased resource supply commonly facilitates invasion by exotic plants, raising concerns over atmospheric nitrogen (N) deposition; fast‐growing annual invaders may have exceptional abilities to outperform native perennials in response to N pulses. However, it remains unclear whether this advantage is due to growth differences or to shifts in competitive outcomes, and whether annual invaders are favored by N deposition in their introduced range over native range. We conducted an experiment to compare the growth and competitive ability of Bromus tectorum and its native perennial grasses either at three different N regimes or between China and North America. The soil used in this experiment was from mountain grasslands as a neutral growth medium. The total biomass of three natives from China and North America did not increase along the N deposition gradient. Nitrogen addition enhanced the growth of North American B. tectorum instead of Chinese B. tectorum. Nitrogen addition increased the competitive ability of B. tectorum, but had no effect on that of natives. North American B. tectorum was bigger and had greater competitive ability and root weight ratio than Chinese B. tectorum. In contrast, North American natives were less competitive than Chinese natives. There was a significantly positive correlation between the growth of B. tectorum grown alone and its competitive ability. These findings suggest that N deposition may enhance the B. tectorum invasion through disproportionally increasing the growth and maintaining inherent competitive advantages of North American B. tectorum, further increasing threats to introduced ranges. There were differences in the growth and competitive ability of B. tectorum and natives between China and North America, which explains why B. tectorum is a minor component at home and becomes a successful invader abroad.     

Non-target effects of an introduced biological control agent on deer mouse ecology

Release of exotic insects as biological control agents is a common approach to controlling exotic plants. Though controversy has ensued regarding the deleterious direct effects of biological control agents to non-target species, few have examined the indirect effects of a ”well-behaved” biological control agent on native fauna. We studied a grassland in west-central Montana infested with spotted knapweed (Centaurea maculosa) to examine the effects of knapweed invasion and two gall flybiological control agents (Urophora affinis and U. quadrifasciata) on the native deer mouse (Peromyscus maniculatus). Stomach-content analysis revealed that Urophora were the primary food item in Peromyscus diets for most of the year and made up 84–86% of the winter diet. Stomach contents indicated that wild-caught mice consumed on average up to 247 Urophora larvae mouse^–1 day^–1, while feeding trials revealed that deer mice could depredate nearly 5 times as many larvae under laboratory conditions. In feeding trials, deer mice selected knapweed seedheads with greater numbers of galls while avoiding uninfested seedheads. When Urophora larvae were present in knapweed seedheads, deer mice selected microhabitats with moderately high (31–45% cover) and high knapweed infestation (≥46% cover). After Urophora emerged and larvae were unavailable to Peromyscus, mice reversed habitat selection to favor sites dominated by native-prairie with low knapweed infestation (0–15%). Establishment of the biological control agent, Urophora spp., has altered deer mouse diets and habitat selection by effecting changes in foraging strategies. Deer mice and other predators may reduce Urophora populations below a threshold necessary to effectively control spotted knapweed. Received: 04 May 1999 / Accepted: 14 August 1999     

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.     

Plant community changes after the reduction of an invasive rangeland weed, diffuse knapweed, Centaurea diffusa

The expected outcome of weed control in natural systems is that the decline of a dominant weed will result in an increase in diversity of the plant community but this has seldom been tested. Here we evaluate the response of the plant community following the decline of diffuse knapweed (Centaurea diffusa) in six different pastures at White Lake, BC, Canada over five years. This period followed the establishment, spread and high levels of attack by the introduced European weevil, Larinus minutus, as part of a biological control program. Knapweed declined immediately before and during the study period, but, contrary to expectations, the species richness and diversity of the rangeland plant community did not increase. The absolute cover of native and introduced forbs and grasses increased following knapweed decline, but only the introduced grasses showed a consistent increase in cover relative to the other life-forms. However, unlike in other studies, the native plants dominated the study site. We conclude that the changes in plant communities following successful biological control are variable among programs and that the impact of replacement species must be evaluated in assessing the success of ecological restoration programs that use biological control to manage an undesirable weed.     

Efficacy and non-target effects of herbicides in foothills grassland restoration are short-lived

Questions

Selective herbicides are frequently used in ecological restoration to control invasive non-native forbs and recover plant communities. However, the long-term efficacy of this practice, its non-target effects on native plants, and its role in facilitating secondary invasions are not well understood. Similarly, little is known about the extent to which herbicide drift may affect native plant communities.

Location

Foothills grasslands of Montana, USA.

Methods

We conducted a 6-year experiment to investigate changes in the abundance of a target invasive plant, knapweed (Centaurea stoebe subsp. micranthos) and plant community structure in response to the herbicides Tordon® (picloram) and Milestone® (aminopyralid), applied at a recommended rate and a diluted rate that simulated drift.

Results

Knapweed cover and the richness of native and non-native forb species declined in the first 3 years in response to treatment at recommended rates, but not drift rates. Secondary invasion by non-native monocots was significant but weak. The cover of native forbs and the cover and richness of native monocots did not differ among treatments but changed significantly with the year. Surprisingly, 6 years after treatments, there were no differences among treatments in the cover of the target invasive plant or community structure.

Conclusions

Our results demonstrate that the efficacy and non-target effects of herbicides in grassland restoration can be short-lived and idiosyncratic because of year effects. Restoration of knapweed invasions might require other active interventions, such as seeding or repeated spraying. Our study supports previous calls for long-term monitoring of herbicides application in ecological restoration.