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.     

Post hoc assessment of an operational biocontrol program: efficacy of the flea beetle Aphthona lacertosa Rosenhauer (Chrysomelidae: Coleoptera), an introduced biocontrol agent for leafy spurge

Mixed populations of Aphthona lacertosa and Aphthona czwalinae were released at more than 50 locations in Alberta in 1997. Two and 3 years post-release, beetle populations were primarily A. lacertosa, with A. czwalinae forming less than 0.5% of the sampled populations. Beetle densities were moderate (10–70 beetles per m²) or high (>70 beetles per m²) at 14% and more than 60% of the sampled sites in 1999 and 2000, respectively. Larger beetles had greater instantaneous egg loads (r²=0.424,P=0.003). In 2000, the largest beetles were found at moderate density sites and there was a significant negative relationship between beetle size and the time taken to accumulate a degree day threshold of 1230 (for females: r²=0.678,P=0.001). Sites with the most rapid accumulation of degree days have the greatest potential for beetle population growth based on potential fecundity. Changes in leafy spurge percent cover, stem density, and canopy height from 1997 to 2000 were assessed across sites with low (<10 beetles per m²), moderate, and high beetle densities in 2000. Sites with high beetle densities had significantly greater reductions of leafy spurge within 5 m of the release point than sites with low beetle densities (P<0.017). Damage caused by the beetles at high-density sites was often visible as a halo-shaped patch of dead leafy spurge stems. The significant overall reduction of leafy spurge within release patches makes A. lacertosa a promising biocontrol agent for leafy spurge in Alberta.     

Prerelease efficacy assessment, in quarantine, of a tephritid gall fly being considered as a biological control agent for Cape-ivy (Delairea odorata)

Due to the long-standing emphasis on only releasing host-specific agents, classical biological control of weeds has an enviable track record of few direct impacts to nontargets. However, even an agent whose host-range is restricted solely to the target weed can have indirect impacts. Such indirect impacts are most likely if, after release, the populations of the agent build up to high numbers without causing accompanying declines in the populations of the target weed. Therefore, it is advisable, prior to release, to demonstrate that the candidate agent is not only host-specific, but that it has clear potential to depress populations of the target weed. Prerelease efficacy assessments (PREA) of potential weed biocontrol agents are not yet common, and are most easily done in the region where both the target and the potential agent are native. We present an example of a PREA performed under strict containment conditions of an approved quarantine facility. A gall-forming fly, Parafreutreta regalis, from South Africa is being considered for release in California to control Cape-ivy, Delairea odorata. We conducted two trials exposing test Cape-ivy plants to two different densities of this fly, and, after approximately two months, comparing the growth of the galled vines to similar vines that had not been exposed to flies. Under both the high density (10 pairs of flies/plant) and low density (2 pairs/plant) treatments, the galled vines exhibited visible stunting, and the ungalled stems were longer, and had more nodes and larger leaves. These trials confirmed that relatively subtle, sublethal impacts on the target can be quantified, even under strict containment conditions, and this should encourage others to assess, prior to release, the potential impact of prospective agents on their proposed target.     

Impact of defoliation by the biocontrol agentZygogramma bicolorataon the weed Partheniumhysterophorus in Australia

The leaf-feeding beetle Zygogrammabicolorata Pallister was introduced from Mexico intoAustralia in 1980 as a biocontrol agent for the weedParthenium hysterophorus L. (Asteraceae). Z. bicolorata became abundant in 1990, and since 1992there has been regular outbreaks resulting in thedefoliation of the weed in central Queensland. In thisstudy we evaluated the impact of defoliation by Z. bicolorata on P. hysterophorus from 1996 to1998. Z. bicolorata caused 91–100% defoliationresulting in reductions in weed density by 32–93%,plant height by 18–65%, plant biomass by 55–89%,flower production by 75–100%, soil seed-bank by13–86% and seedling emergence in the following seasonby 73–90%. At sites with continued outbreaks ofZ. bicolorata, it is expected that the existing soilseed-bank will be minimised, resulting in reduceddensity of parthenium in 6 to 7 years.     

Establishment and dispersal of the biological control weevil Rhinoncomimus latipes on mile-a-minute weed, Persicaria perfoliata

Mile-a-minute weed, Persicaria perfoliata (L.) H. Gross (Polygonaceae), is an annual vine from Asia that has invaded the eastern US where it can form dense monocultures and outcompete other vegetation in a variety of habitats. The host-specific Asian weevil Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae) was first released in the US in 2004 as part of a classical biological control program. The weevil was intensively monitored in three release arrays over 4 years, and field cages at each site were used to determine the number of generations produced. The weevil established at all three sites and produced three to four generations before entering a reproductive diapause in late summer. Weevils dispersed at an average rate of 1.5–2.9 m wk⁻¹ through the 50 m diameter arrays, which had fairly contiguous mile-a-minute cover. Weevils dispersing in the broader, more variable landscape located both large monocultures and small isolated patches of mile-a-minute 600–760 m from the release within 14 months. Weevil density ranged from fewer than 10 to nearly 200 weevils m⁻² mile-a-minute weed. Mile-a-minute cover decreased at the site with the highest weevil density. The production of P. perfoliata seed clusters decreased with increasing weevil populations at two sites, and seedling production declined over time at two sites by 75% and 87%. The ability of the weevil to establish, produce multiple generations per season, disperse to new patches, and likelihood of having an impact on plants in the field suggests that R. latipes has the potential to be a successful biological control agent.     

Direct and ecological costs of resistance and tolerance in the stinging nettle

Plant resistance and tolerance to herbivores, parasites, pathogens, and abiotic factors may involve two types of costs. First, resistance and tolerance may be costly in terms of plant fitness. Second, resistance and tolerance to multiple enemies may involve ecological trade-offs. Our study species, the stinging nettle ( Urtica dioica L.) has significant variation among seed families in resistance and tolerance as well as costs of resistance and tolerance to the holoparasitic plant Cuscuta europaea L. Here we report on variation among seed families (i.e. genetic) in tolerance to nutrient limitation and in resistance to both mammalian herbivores (i.e. number of stinging trichomes) and an invertebrate herbivore (i.e. inverse of the performance of a generalist snail, Arianta arbustorum). Our results indicate direct fitness costs of snail resistance in terms of host reproduction whereas we did not detect fitness costs of mammalian resistance or tolerance to nutrient limitation. We further tested for ecological trade-offs among tolerance or resistance to the parasitic plant, herbivore resistance, and tolerance to nutrient limitation in the stinging nettle. Tolerance of nettles to nutrient limitation and resistance to mammalian herbivores tended to correlate negatively. However, there were no significant correlations among resistance and tolerance to the different natural enemies (i.e. parasitic plants, snails, and mammals). The results of this greenhouse study thus suggest that resistance and tolerance of nettles to diverse enemies are free to evolve independently of each other but not completely without direct costs in terms of plant fitness.     

Monitored releases of Rhinoncomimus latipes (Coleoptera: Curculionidae), a biological control agent of mile-a-minute weed (Persicaria perfoliata), 2004–2008

Mile-a-minute weed, Persicaria perfoliata (L.) H. Gross, is an invasive annual vine of Asian origin that has developed extensive monocultures, especially in disturbed open areas in the Mid-Atlantic region of the United States. A host-specific Asian weevil, Rhinoncomimus latipes Korotyaev, was approved for release in North America in 2004, and weevils have been reared at the New Jersey Department of Agriculture Beneficial Insect Laboratory since then. By the end of 2007 more than 53,000 weevils had been reared and released, mostly in New Jersey, but also in Delaware, Maryland, Pennsylvania, and West Virginia. The beetles established at 63 out of 65 sites (96.9%) where they were released between 2004 and 2007, with successful releases consisting of as few as 200 weevils. Weevils were recorded at 30 additional non-release sites in New Jersey, where they had dispersed at an average rate of 4.3 km/year. Standardized monitoring of fixed quadrats was conducted in paired release and control sites at eight locations. Significant differences in mile-a-minute weed populations in the presence and absence of weevils were found at three locations, with reduction in spring densities to 25% or less of what they had been at the start within 2–3 years at release sites, while weed densities at control sites were largely unchanged. Mile-a-minute weed populations at a fourth site were similarly reduced at the release site, but without control data for comparison due to rapid colonization of the paired control site. At the other four locations, all on islands, mile-a-minute weed populations were reduced at both release and control sites without large weevil populations developing, apparently due to environmental conditions such as late frost and extreme drought.     

Field assessment of the risk posed by Diorhabda elongata, a biocontrol agent for control of saltcedar (Tamarix spp.), to a nontarget plant, Frankenia salina

The biological control program for saltcedar (Tamarix spp.) has led to open releases of a specialist beetle (Chrysomelidae: Diorhabda elongata) in several research locations, but the controversy over potential impacts to native, nontarget plants of the genus Frankenia remains unresolved. To assess the potential for nontarget impacts under field conditions, we installed cultivated Frankenia spp. (primarily two forms of Frankenia salina but also including Frankenia jamesii) at locations in Nevada and Wyoming where D. elongata densities and saltcedar defoliation were expected to be very high, so insects would be near starvation with high probability of attacking nontargets if these were suitable hosts. Subsequent insect abundance was high, and only minor impact (<4% foliar damage) was observed on both forms of F. salina under these ‘worst case’ conditions; there was no impact to F. jamesii. No oviposition nor larval development were observed on any plants, there was no dieback of damaged F. salina stems, and plants continued growing once insect populations subsided. These results under ‘natural’ field conditions contrast with caged host-range tests in which feeding, development and minor oviposition occurred on the nontarget plant. Other ecological factors, such as distance from target plants to natural Frankenia spp. populations, inhospitable conditions for agent survival in such sites, and intrinsic insect behavior that makes colonization and/or genetic adaptation highly unlikely, lead us to conclude that nontarget impacts following program implementation will be insignificant or absent. Host range testing of new agents, while necessary to ensure safety, must put greater attention on assessing the ecological context where agents will be establishing, and on balancing speculated risks against potential benefits of biological control.     

Effects of biocontrol agents on Fusarium verticillioides count and fumonisin content in the maize agroecosystem: Impact on rhizospheric bacterial and fungal groups

The present study tested the ability of Bacillus amyloliquefaciens and Microbacterium oleovorans to reduce Fusarium verticillioides populations and fumonisin accumulation in the maize agroecosystem. The impact of releasing these biocontrol agents on rhizospheric bacterial and fungal groups was also evaluated through isolation and identification of culturable microorganisms. When applied as seed coatings at a concentration of 10⁷ CFU ml⁻¹ both agents were effective in reducing F. verticillioides counts and fumonisin B₁ and B₂ content from maize grains. Rhizospheric counts of the pathogen were also decreased by use of B. amyloliquefaciens at 10⁷ CFU ml⁻¹. Richness and diversity indexes calculated for bacteria and fungi inhabiting the rhizosphere of maize remained unchanged following the addition of both biocontrol agents to seeds. Our research is being continued to further characterize the bacterial and fungal isolates with additional field assays.     

Predicted Impact of an Exotic Generalist Predator on Monarch Butterfly (Lepidoptera: Nymphalidae) Populations: A Quantitative Risk Assessment

The multicolored Asian lady beetle, Harmonia axyridis (Pallas), was recently identified as a potential hazard to monarch butterflies, Danaus plexippus (L.). A quantitative risk assessment for the potential impact of H. axyridis on D. plexippus in Minnesota corn and soybean fields was developed using risk analysis software. This assessment considered a potential worst-case scenario for the impact of H. axyridis on D. plexippus. Habitat-specific recruitment of D. plexippus eggs was determined empirically. Subsequently, simulated abundance of D. plexippus in each habitat was reduced by two classes of stage-specific mortality: predation by H. axyridis and causes other than H. axyridis. Predation was modeled as a function of D. plexippus exposure to H. axyridis, and predation rate of D. plexippus by H. axyridis. Exposure and subsequent risk varied considerably by habitat, with a low risk of H. axyridis adversely affecting D. plexippus populations developing in corn fields, but a moderate to high risk in soybean fields. Predicted rates of D. plexippus mortality attributable to H. axyridis were greater in soybean compared to corn fields, possibly due to the numerical response of H. axyridis to soybean aphid, Aphis glycines Matsumura. This study demonstrates that H. axyridis has the potential to have a strong adverse impact on D. plexippus populations. However, the likelihood of occurrence for this worst-case scenario remains uncertain. To evaluate the landscape-level risk of H. axyridis impacting D. plexippus, further data on recruitment of D. plexippus in other habitats and exposure estimates specific to additional habitats are needed.     

Integration of biological control agents with other weed management technologies: Successes from the leafy spurge (Euphorbia esula) IPM program

An invasive weed can occupy a variety of environments and ecological niches and generally no single control method can be used across all areas the weed is found. Biological control agents integrated with other methods can increase and/or improve site-specific weed control, but such combinatorial approaches have not been widely utilized. The successful leafy spurge (Euphorbia esula L.) control program provides examples for future integrated weed programs that utilize biological control agents with traditional methods. Weed control methods can be used separately, such as when the leafy spurge gall midge (Spurgia esulae Gagné) reduced seed production in wooded areas while herbicides prevented further spread outside the tree line. Traditional methods also can be used directly with biological control agents. Incorporation of Aphthona spp. with herbicides has resulted in more rapid and complete leafy spurge control than either method used alone. Also, the insect population often increased rapidly following herbicide treatment, especially in areas where Aphthona spp. were established for several years but had been ineffective. Incorporation of Aphthona spp. with sheep or goat grazing has resulted in a larger decline in leafy spurge production than insects alone and in weed density than grazing alone. Controlled burns can aid establishment of biological control agents in marginally suitable environments, but timing of the fire must be coordinated to the insect’s life-cycle to ensure survival. Integration of biological control agents with revegetation programs required the agent to be the last method introduced because the cultivation and herbicide treatments necessary to establish desirable grasses and forbs were destructive to the insect. In a practical application, herbicides were combined with Aphthona spp. to help the insect establish and control leafy spurge in the habitat of the western prairie fringed orchid (Platanthera praeclara Sheviak and Bowles), an endangered species. Several experimental designs can be used to evaluate biological control agents with cultural, mechanical, and chemical control methods or with additional biological agents.     

Induction of PR proteins and resistance by the biocontrol agent Clonostachys rosea in wheat plants infected with Fusarium culmorum

Clonostachys rosea (CR) is a common worldwide saprophyte with destructive effect against several plant pathogenic fungi showing antagonistic features against a wide variety of pathogens. We recently isolated a strain of C. rosea, named CR47, from wheat crown infected with Fusarium culmorum (FC); this strain proved to be effective against Fusarium seed borne diseases of cereals under field condition. In this paper the function of C. rosea applied as seed treatment on wheat seedling growth was investigated. In addition, we investigated the expression pattern of peroxidases and chitinases as well as PR4 proteins following both CR treatments of seeds and FC infection and also in the three-component system pathogen–antagonist–wheat. Several chitinase isoforms were induced by CR-treatment both in coleoptiles and roots, whereas some peroxidase isoforms were induced only in the presence of both antagonist and pathogen. In the latter case, it seems that CR-treatment by itself promotes plant growth and reduces the peroxidase expression, while enhances some chitinase isoforms probably involved in cell wall disruption. Moreover, both the antagonist and the pathogen studied induced PR4 protein expression, which probably exerts its role on the invading microorganisms by a translation-inhibitory process that could be ascribed to their ribonuclease activity.     

Biocontrol of a perennial legume,Sesbania punicea,using a florivorous weevil,Trichapion lativentre: weed population dynamics with a scarcity of seeds

Summary The establishment in South Africa of a florivorous, apionid weevil, Trichapion lativentre, on Sesbania punicea, a leguminous weed of South American origin, has reduced seed production of the plants by >98%. Surveys of the age structure and density of plants in infestations of S. punicea throughout South Africa have shown that the rate of recruitment of seedlings has drastically declined within a few years in many areas, due to the weevils. However, there has unexpectedly not been a corresponding decline in the density of mature plants in extant infestations of S. punicea. In spite of this, T. lativentre has curtailed the rate of spread of the weed into uninvaded habitats and has impeded reinvasion into areas cleared of infestations by mechanical means or by another complimentary biocontrol agent.     

Host range of Ceutorhynchus assimilis (Coleoptera: Curculionidae), a candidate for biological control of Lepidium draba (Brassicaceae) in the USA

Ceutorhynchus assimilis has been selected as a potential biological control agent of Lepidium draba, which is a Eurasian invasive weed in North America. Preliminary studies indicated specificity of this weevil collected in southern France on L. draba. This result was in discord with the pest status of C. assimilis found in the literature. Host-specificity tests based both on field and laboratory experiments showed heterogeneity in the host spectrum of the weevils reared from different host-plants as determined by larval development. However, no distinguishable morphological differences could be visually detected between the populations feeding on different host-plants. All sampled populations of weevils were polyphagous as adults. Weevils reared from L. draba were specific to this plant for their complete larval development. Conversely, populations living on other wild and cultivated Brassicaceae species were not able to use L. draba as a host plant. Such differentiation is further highlighted by other biological aspects such as plant infestation rates, sex-ratio, duration of larval development, and differences in the timing of their life cycles. These results demonstrate that C. assimilis, an insect species formerly considered as a pest of Brassicaceae, is characterized by its host-range variability, with one population being potentially useful in the biological control of L. draba. Moreover, this example points to the need to test multiple populations of biological control agents in assessing risk.     

Field assessment of host plant specificity and potential effectiveness of a prospective biological control agent, Aceria salsolae, of Russian thistle, Salsola tragus

The eriophyid mite, Aceria salsolae de Lillo and Sobhian, is being evaluated as a prospective classical biological control agent of invasive alien tumbleweeds, including Salsola tragus, S. collina, S. paulsenii and S. australis, in North America. Previous laboratory experiments to determine the host specificity of the mite indicated that it could sometimes persist and multiply on some nontarget plants, including Bassia hyssopifolia and B. scoparia. These are both European plants whose geographic range overlaps that of the mite, but the mite has never been observed on them in the field. A field experiment was conducted in Italy to determine if the mite would infest and damage these plants under natural outdoor conditions. The results indicate that this mite does not attain significant populations on these nontarget plants nor does it significantly damage them. Salsola tragus was heavily infested by A. salsolae, and plant size was negatively correlated to the level of infestation. Although S. kali plants were also infested, their size did not appear to be affected by the mites. The other nontarget plants were not as suitable for the mite in the field as in previous laboratory experiments. We conclude that there would be no significant risk to nontarget plants as a result of using A. salsolae as a biological agent to control Salsola species in North America.     

Osmotic stress adaptation, compatible solutes accumulation and biocontrol efficacy of two potential biocontrol agents on Fusarium head blight in wheat

Fusarium head blight (FHB) caused by Gibberella zeae (anamorph = Fusarium graminearum) is a devastating disease that causes extensive yield and quality losses to wheat in humid and semi-humid regions of the world. Biological control has been demonstrated to be effective under laboratory conditions but a few biocontrol products have been effective under field conditions. The improvement in the physiological quality of biocontrol agents may improve survival under field conditions, and therefore, enhance biocontrol activity. Bacillus subtilis RC 218 and Brevibacillus sp. RC 263 were isolated from wheat anthers and showed significant effect on control of FHB under greenhouse assays. This study showed the effect of water availability measured as water activity (a_W) using a growth medium modified with NaCl, glycerol and glucose on: (i) osmotic stress tolerance, (ii) viability in modified liquid medium, (iii) quantitative intracellular accumulation of betaine and ectoine and (iv) the biocontrol efficacy of the physiologically improved agents. Viability of B. subtilis RC 218 in NaCl modified media was similar to the control. Brevibacillus sp. RC 263 showed a limited adaptation to growth in osmotic stress. Betaine was detected in high levels in modified cells but ectoine accumulation was similar to the control cells. Biocontrol activity was studied in greenhouse assays on wheat inoculated at anthesis period with F. graminearum RC 276. Treatments with modified bacteria reduced disease severity from 60% for the control to below 20%. The physiological improvement of biocontrol agents could be an effective strategy to enhance stress tolerance and biocontrol activity under fluctuating environmental conditions.     

Morphological evolution and ecological diversification of the forest-dwelling poppies (Papaveraceae: Chelidonioideae) as deduced from a molecular phylogeny of the ITS region

Sequences of the ITS region of nrDNA were analyzed for the seven genera of Papaveraceae subf. Chelidonioideae s.str. Three major clades can be recognized. These are 1.Chelidonium/Hylomecon/Stylophorum, 2.Eomecon/Sanguinaria, and 3.Bocconia/Macleaya. The monophyly of genera in the first of these three clades is doubtful, and clades two and three are sister to each other. Use of the ITS phylogeny of the subfamily to trace its morphological and ecological evolution shows that morphological change is concentrated in theBocconia/Macleaya clade, and probably related to the evolution of wind-pollination from insect-pollination in these two genera after habitat shift.     

Secondary succession of weed communities in lowland habitats of Taiwan in relation to the introduction of wild-rice (Oryza perennis) populations

Wild-rice (Oryza perennis Moench) populations were introduced into abandoned lowland fields (and a ditch) at five sites in Taiwan. The weeds arising after clearance and tillage of the fields were observed with regard to various attributes of the community during three years. Among a total of 84 identified species, Leersia hexandra had the highest relative frequency. With secondary succession, the percentage biomass of perennial species, particularly of grasses, gradually increased and the dominance index (1-H′/H′ _max) also increased. Species present in the buried seed pool in soil samples showed a different pattern of successional changes. With species classified into grass, sedge, and dicot groups and their percentage of biomass data compared with corresponding data for the density of plants from buried seeds, a negative correlation was found which suggested differences in behavior among the plant groups. Experiments on the effects of weeding and inundation suggested that the niches of Oryza perennis and of Leersia hexandra largely overlap, and differed from that of Panicum repens. The wild-rice populations declined as Leersia dominated. This relationship was demonstrated by principal component analysis and standard partial regression analysis of the data.