The lesser of two weevils: physiological responses of spotted knapweed (Centaurea stoebe) to above- and belowground herbivory by Larinus minutus and Cyphocleonus achates

The physiological responses of plants to variable levels of root and shoot herbivory in the field may yield valuable insights regarding potential compensation or tolerance responses for herbivory. In an infestation of Centaurea stoebe (spotted knapweed) located in the Colorado foothills, we measured physiology, biomass, and flower production of individual plants undergoing a natural range of herbivory by the above- and belowground biological control insects, Larinus minutus and Cyphocleonus achates. Over the growing season, net carbon assimilation rate, transpiration, stomatal conductance, and intercellular leaf [CO₂] (C _i) all decreased, while water use efficiency increased. The decrease in these physiological traits was due to an increase in the intensity of L. minutus damage over time; effects of C. achates root damage to plant physiology, including transpiration were only marginally significant. The effects of these two species on plant physiology were not interactive, and Larinus minutus was found to exert larger negative effects on this invasive plant in terms of plant physiology and potential reproductive output than C. achates. While previous studies have shown C. achates to have significant negative effects on population densities of spotted knapweed, the addition of Larinus minutus to the suite of insects used in biological control of spotted knapweed should facilitate continued or enhanced reduction in densities of this noxious weed.     

Testing biological control agent compatibility: Cyphocleonus achates and Larinus minutus on diffuse knapweed

While weed biological control success is typically achieved with one agent, multiple agents are invariably introduced. Biological control agents that share a host–plant may interact either directly or indirectly through changes in host–plant quality. Negative interactions could reduce the impacts of the agents on the density of their host–plant while positive interactions (facilitation) could improve biological control success.In the Okanagan Valley of British Columbia, Canada, initial declines in the invasive rangeland weed, diffuse knapweed (Centaurea diffusa) were attributed to the introduction of the weevil Larinus minutus. A second weevil, Cyphocleonus achates has recently become common on diffuse knapweed. We sought to determine if the recent increase of C. achates could threaten the success of L. minutus. We considered whether L. minutus colonisation or performance remained the same when C. achates was present, and whether the two agents acted independently to reduce plant performance.Neither changes in colonisation rates nor competitive interactions were apparent between C. achates and L. minutus. Both insects reduced plant performance and, for all metrics, the reduction in plant performance by one species was independent of the second. The two agents appear to be compatible and both should contribute to the control of diffuse knapweed. To assess how biological control agents interact requires understanding both their competitive interactions and their joint effects on the shared host.     

Interactions and effects of multiple biological control insects on diffuse and spotted knapweed in the Front Range of Colorado

Abundances and interactions among biological control insects and their effects on target invasive plants were monitored within the flower heads and roots of diffuse knapweed, Centaurea diffusa, and in spotted knapweed, Centaurea stoebe, along the Colorado Front Range. Flower weevils, (Larinus species) and root-feeders (Cyphocleonus achates and Sphenoptera jugoslavica) were released on knapweed that already supported biological control gall flies (Urophora species). At a single monitoring site, seed production by C. diffusa declined from 4400 seeds m⁻² in 1997 to zero seeds m⁻² on the monitoring sites in 2006, while the flowering stem density of C. diffusa declined from a peak of almost 30 stems m⁻² in 2000 to zero stems m⁻² in 2006. The average abundance of Urophora and Larinus in flower heads fluctuated independently during the 2001–2006 interval, while the relative abundance of C. achates and S. jugoslavica in roots exhibited a weak inverse relationship that appeared driven by climate effects. The relative abundance of insects on a population of C. stoebe was monitored for five years as Larinus species and C. achates became established on spotted knapweed that already supported Urophora species. Spotted knapweed seed production on our monitoring site declined from 4600 seeds m⁻² in 2003 to zero seeds m⁻² in 2006. Unlike C. diffusa, substantial numbers of rosettes of C. stoebe remained present. Larinus consumed almost all Urophora encountered in C. diffusa, and consumed about 40% of the Urophora in co-infested flower heads of C. stoebe (ca. 10–15% of the total Urophora population). No negative correlations between the relative densities of flower head and root-feeding insects were observed. The effects of these insects on target plants have produced results consistent with the ‘cumulative stress hypothesis’ for biological control of Centaurea species.     

Revised methods for the mass-rearing of the spotted knapweed biological control agent,Cyphocleonus achates (Coleoptera: Curculionidae), in field corrals

Cyphocleonus achates (Fahraeus), a root-feeding weevil introduced from Eurasia, is an effective biological control agent against spotted knapweed, Centaurea stoebe L. ssp. micranthos. Because C. achates is univoltine and does not fly, distribution of the weevil has been slow. To hasten the weevil's distribution, a rearing effort using field corrals was initiated at a facility in Corvallis, Montana. Procedures for mass-rearing the weevil in field corrals are described, with an emphasis on improvements over earlier methods. The described field-corral approach is effective and appropriate for producing C. achates for distribution in the western United States.     

Linking field based studies with greenhouse experiments: the impact of <Emphasis Type="Italic">Centaurea stoebe</Emphasis> (=<Emphasis Type="Italic">C. maculosa</Emphasis>) in British Columbia grasslands

The grassland community of Lac du Bois Provincial Park in the interior of British Columbia has become increasingly invaded by Centaurea stoebe (=Centaurea maculosa; spotted knapweed). Allelopathy, through the production of the phytotoxin (±)-catechin by C. stoebe roots is believed to be partly responsible for knapweed’s invasive success. We used field sampling and greenhouse experiments to examine two questions: (1) Is increasing abundance of C. stoebe negatively associated with the abundance of specific native grassland species? (2) Do species that exhibit a negative correlation with C. stoebe abundance in the field demonstrate increased levels of susceptibility to application of (±)-catechin during germination? Thirty-eight plots were sampled in the grassland, encompassing areas of low—high knapweed abundance. Seeds from eight native species, exhibiting positive, neutral or negative correlation with knapweed abundance, were treated with three concentrations (0, 0.5, 2.0 mg/mL) of (±)-catechin. Root growth and percent germination were measured over a 6-week period. The results indicate that C. stoebe abundance is negatively correlated with native plant species abundance and may alter plant community composition. Moreover root radical growth was significantly negatively affected by treatment with (±)-catechin in all four native plant species that exhibited a negative correlation with knapweed abundance in the field. Past studies have failed to conclusively link greenhouse results with plant community patterns. Here, we provide a correlative link between plant community composition and tolerance to a phytotoxin.     

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.     

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.     

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.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

Effects of plant competition,seed predation,and nutrient limitation on seedling survivorship of spotted knapweed (<Emphasis Type="Italic">Centaurea stoebe</Emphasis>)

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m² (ranging from 0 to 16,364 seeds/m²), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568–2,272 seeds m⁻² year⁻¹), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of ‘safe sites’ within undisturbed vegetation can limit densities of C. stoebe.     

Influence of plant phenostage and ploidy level on oviposition and feeding of two specialist herbivores of spotted knapweed,Centaurea stoebe

A caged field experiment was used to determine how Centaurea stoebe L. phenostage (rosette, single-stem, multiple-stem) and ploidy level (diploid = 2× and tetraploid = 4×) influence oviposition and feeding of two biological control agents, Agapeta zoegana (Lep.: Cochylidae) and Cyphocleonus achates (Col.: Curculionidae). Ploidy level did not influence oviposition patterns of A. zoegana but rosette and one-stem plants had significantly more eggs than multiple-stem (4×) plants. Differences in oviposition levels did not translate into differences in larval densities, but 2× plants (particularly large one-stem plants) had significantly more larvae than 4× plants. There was a significant positive correlation between numbers of larvae and root diameter. Ploidy level and phenostage both had a significant effect on C. achates feeding damage, with adults feeding more frequently on multiple-stem plants. No C. achates larvae were observed when the roots were dissected. Furthermore, the generalist herbivore Arion lusitanicus, naturally present in the garden plots, was predominantly associated with young rosette plants, a stage at which survival rate is acknowledged to be the most important determinant of knapweed density. These results indicate that the combined damage caused by A. zoegana and C. achates, superimposed on damage caused by generalist herbivores in the local community, could provide effective control for C. stoebe.     

Sex‐biased seed predation in gynodioecious Dianthus superbus var. longicalycinus (Capryophyllaceae) and differential influence of two seed predator species on the floral traits

Gynodioecy, the co‐occurrence of hermaphrodite and female individuals within a species, is maintained by differential reproductive success between sexes. Recently, researchers recognized that not only pollinators but also herbivores are important agents in the evolution and maintenance of gynodioecy, when herbivory is hermaphrodite biased. In this study, we investigated whether there is hermaphrodite‐biased herbivory in a gynodioecious plant, Dianthus superbus var. longicalycinus, and if so, what floral traits influenced hermaphrodite‐biased herbivory. We measured flower morphology (flower diameter, calyx tube length, corolla height and petal width) and phenology of flowers of female individuals, hermaphrodites and gynomonoecious individuals in a natural population. We also investigated seed predation and predator species. At the study site, Sibinia weevils (Curculionidae; Coleoptera) and Coleophora moths (Coleophoridae; Lepidoptera) were common pre‐dispersal seed predators in this species. The weevil appeared early in the flowering season, and weevil predation correlated with flower phenology. Because female individuals did not flower early in the season, weevil predation was less frequent in female individuals. Moth predation correlated with calyx length. The calyx length of flowers of female individuals was smaller than those of hermaphrodites, but a direct comparison of moth predation rates failed to find a significant difference among sex morphs. We found that the two seed predators had different effects on floral traits in D. superbus var. longicalycinus. We suggest that weevil predation contributes to the maintenance of gynodioecy because female individuals successfully escaped weevil predation by flowering late. It remains unclear why flower phenology is different among sex morphs.     

The case against (-)-catechin involvement in allelopathy of Centaurea stoebe (spotted knapweed)

Proving allelopathic chemical interference is a daunting endeavor, in that production and movement of a phytotoxin from a donor plant to a receiving plant must be demonstrated in the substrate in which the plants grow, which is usually a complex soil matrix. The soil levels or soil flux levels of the compound generated by the donor must be proven to be sufficient to adversely affect the receiving plant. Reports of (-)-catechin to be the novel weapon used by Centaurea stoebe (spotted knapweed) to invade new territories are not supported by the paper featured in this Addendum, nor by papers produced by two other laboratories. These papers find that (-)-catechin levels in soil in which C. stoebe grows are orders of magnitude below levels that cause only minor growth effects on reported sensitive species. Furthermore, the claim that (-)-catechin acts as a phytotoxin through causing oxidative damage is refuted by the fact that the molecule is a strong antioxidant and is quickly degraded by extracellular root enzymes.Key words: allelochemical, allelopathy, catechin, chemical interference, phytotoxin, spotted knapweed     

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.     

Biogeographic differences in the effects of <Emphasis Type="Italic">Centaurea stoebe</Emphasis> on the soil nitrogen cycle: novel weapons and soil microbes

The success of some invasive plants may be due in part to native organisms lacking adaptation to species-specific biochemical traits of invaders—the Novel Weapons Hypothesis. We tested this hypothesis in the context of soil microbial communities by comparing the effects of Centaurea stoebe and the root exudate (±)-catechin, on ammonification and nitrification in both the non-native and native ranges of this species. In a non-native range (Montana), soil nitrate (NO₃ ⁻) concentrations were lower in invaded than uninvaded grasslands. This did not appear to be due only to higher uptake rates as both C. stoebe plants and catechin significantly reduced resin extractable NO₃ ⁻, the maximum rate of nitrification, and gross nitrification in Montana soils. Thus, reduced NO₃ ⁻ in invaded communities may be due in part to the inhibition of nitrifying bacteria by secondary metabolites produced by C. stoebe. The effects of C. stoebe on N-related processes were different in Romanian grasslands, where C. stoebe is native. In Romanian soil, C. stoebe had no effect on resin extractable NH₄ ⁺ or NO₃ ⁻ (compared to other plant species), the maximum rate of nitrification, nor gross nitrification. A relatively high concentration of catechin reduced the maximum rate of nitrification in situ, but substantially less than in Montana. In vivo, gross ammonification was lowest when treated with catechin. Our results suggest biogeographic differences in the way a plant species alters nitrogen cycling through the direct effects of root exudates and adds to a growing body of literature demonstrating the important belowground effects of invasive plants.     

Growth and development of the knapweed root weevil,Cyphocleonus achates,on a meridic larval diet

Cyphocleonus achates, the knapweed weevil, is an effective biological control agent of the invasive weed, Centaurea maculosa Lam. A meridic diet was developed and tested for the rearing of the larval stage of this insect. Using this diet, C. achates was reared for over three generations, with the adults being offered knapweed plants for feeding and oviposition in greenhouse conditions. Slight or no differences were seen between insects reared on a standard meridic diet formulation and one containing knapweed tissues. The following life history parameters were monitored over the three generations: percent egg hatch (ranging from 42.9 to 59.1%), time to egg hatch (20.0–23.2 days), time to adult emergence (52.0–54.1 days), adult weights 3 days post-eclosion (101.9–117.0 mg), percent adult emergence (48.3–58.6%), and percent mortality/deformity in the different stages (with mortality occurring primarily in the early larval stages). Additionally, a study involving low temperature and short day conditions suggested that C. achates could be maintained for longer periods of time in larval diet cells when placed in growth-retarding conditions, although percent adult emergence was lower. External morphology was also studied in order to distinguish between the sexes to ensure that each adult cage had a similar ratio of females to males. Abdominal features were found to be the most dependable characteristics for use when determining the sex of adult C. achates.     

Phenology and dispersal of Larinus minutus Gyllenhal and Larinus obtusus Gyllenhal (Coleoptera: Curculionidae), two biological control agents of Centaurea stoebe ssp. micranthos (spotted knapweed) in Michigan

Centaurea stoebe L. ssp. micranthos (Gugler) (spotted knapweed) is an invasive plant that has been the target of classical biological control in North America for more than four decades. The seedhead-feeding weevils Larinus minutus Gyllenhal and Larinus obtusus Gyllenhal (Coleoptera: Curculionidae) are two of the most-widely released C. stoebe control agents, and have more recently been introduced into the eastern US. While there have been many studies focusing on their ability to impact C. stoebe in the western US and Canada, there have been few studies from eastern North America, and basic knowledge of important aspects of their biology is lacking. Here we report on the phenology and dispersal of L. minutus and L. obtusus in Michigan. We regularly sampled two established Larinus spp. populations in southern Michigan in 2012 and 2013, and found that while adult abundance fluctuates during the growing season, they remained at easily detectable levels from mid-June through the end of August. We also used previously established populations of L. minutus and L. obtusus released in 1996 (n = 1), 2007 (n = 2), and 2010 (n = 5) to determine how dispersal of Larinus spp. into the surrounding landscape changes with time since release. Populations of Larinus spp. weevils showed little dispersal for 2 years post-release. However, after initial establishment dispersal rates increased rapidly, resulting in average dispersal rates that increased exponentially with time since release. These findings can inform future biological control release and sampling programs for Larinus spp. in eastern North America.     

Inter-annual variation in above- and belowground herbivory on a native,annual legume

The relative importance of subterranean versus aboveground insect damage to plants is not well understood. In particular, the simultaneous effects of above- and belowground herbivory, and the importance of highly variable abiotic factors such as rainfall, have received little attention in diverse natural ecosystems. We investigated the influence of both above- and belowground herbivory on Lupinus nanus (Fabaceae), an annual plant native to coastal California. A number of insect species damage L. nanus aboveground, and a weevil larva consumes nodules belowground. To manipulate herbivory in the field, we employed a combination of insecticides and simulated herbivory during two different years. In 1997, simulated belowground damage reduced L. nanus survival, and insecticide application to roots increased seed production and seed mass. By contrast, in 1998, only aboveground folivory significantly reduced L. nanus reproduction, and, in combination, above- and belowground insecticides did not affect flower or seed number relative to controls. A growth chamber experiment conducted in the absence of herbivory revealed that the aboveground insecticide marginally reduced flower production and the belowground insecticide marginally increased flower production compared to controls; these non-target effects made our field experiments for aboveground herbivory conservative. Finally, ambient levels of herbivory differed among years (1997, 1998, and 2000), which varied greatly in rainfall due to the effects of El Nino. The results suggest that the impacts of herbivores are temporally variable and that abiotic factors, particularly those related to large-scale changes in weather patterns, may be more important than insect herbivory to L. nanus in some years. This revised version was published online in August 2006 with corrections to the Cover Date.     

Do invasive plants structure microbial communities to accelerate decomposition in intermountain grasslands?

Invasive plants are often associated with greater productivity and soil nutrient availabilities, but whether invasive plants with dissimilar traits change decomposer communities and decomposition rates in consistent ways is little known. We compared decomposition rates and the fungal and bacterial communities associated with the litter of three problematic invaders in intermountain grasslands; cheatgrass (Bromus tectorum), spotted knapweed (Centaurea stoebe) and leafy spurge (Euphorbia esula), as well as the native bluebunch wheatgrass (Pseudoroegneria spicata). Shoot and root litter from each plant was placed in cheatgrass, spotted knapweed, and leafy spurge invasions as well as remnant native communities in a fully reciprocal design for 6 months to see whether decomposer communities were species‐specific, and whether litter decomposed fastest when placed in a community composed of its own species (referred to hereafter as home‐field advantage–HFA). Overall, litter from the two invasive forbs, spotted knapweed and leafy spurge, decomposed faster than the native and invasive grasses, regardless of the plant community of incubation. Thus, we found no evidence of HFA. T‐RFLP profiles indicated that both fungal and bacterial communities differed between roots and shoots and among plant species, and that fungal communities also differed among plant community types. Synthesis. These results show that litter from three common invaders to intermountain grasslands decomposes at different rates and cultures microbial communities that are species‐specific, widespread, and persistent through the dramatic shifts in plant communities associated with invasions.