Mowing during rainfall enhances the control of Cirsium arvense

Pastoral farmers in New Zealand have described dramatic demises in populations of the weed Cirsium arvense (L.) Scop., a perennial herb indigenous to Eurasia, following its mowing during rainfall. To test the hypotheses (1) that the mowing of C. arvense during rainfall increases the control of this weed and (2) that the causal organism in this ‘mowing-in-the-rain’ effect is the vascular wilt fungus Verticillium dahliae, two series of field experiments were carried out in C. arvense-infested pastures in New Zealand, one in autumn 2008 (Experiment series 1, 9 farms), and another in spring/summer 2008–2009 (Experiment series 2, 12 farms). The effect of mowing in the rain as compared to mowing in the dry was to reduce the % ground cover of the thistle in the spring following treatment by 21 and 32% in Experiment series 1 and 2, respectively. Correlations of this ‘rain versus dry’ effect with the incidence of V. dahliae in the subterranean parts of C. arvense shoots sampled in each of the two field experiment series provided no statistical evidence that the effect increased with V. dahliae incidence. Thus these experiments provide no support for the hypothesis that V. dahliae is the biological mechanism for the ‘mowing-in-the-rain’ effect. Nevertheless, they do support mowing during rainfall as a simple and effective management tactic for C. arvense.     

Effects of pasture competition and specialist herbivory on the performance of Cirsium arvense

Combining specialist herbivory with interspecific plant competition can be an effective means of controlling pasture weeds. Cirsium arvense (Canada thistle, Californian thistle, creeping thistle) is one of the worst weeds of pastoral production systems in New Zealand (NZ). The oligophagous leaf-feeding beetle, Cassida rubiginosa, was recently released in NZ for control of C. arvense. To assess the impact of this biocontrol agent we conducted an outdoor potted-plant experiment with low and high densities of Cassida larvae combined with different levels of interspecific competition from typical NZ pasture species. Secondly, we carried out a field-release experiment to quantify the impact of high densities of Cassida under more natural conditions. Interspecific competition reduced all measured plant parameters of C. arvense except mean shoot height and base diameter. Herbivory by Cassida only reduced root biomass, and showed a weak additive response when combined with competition. All other measured parameters of C. arvense showed a substitutive response, with competition being the only factor having a significant impact on the weed. There were no significant synergistic interactions with competition and herbivory on C. arvense. Interestingly, the number of root buds per plant was significantly greater in the presence of herbivory by Cassida, suggesting that C. arvense may compensate for defoliation. Similar to the potted-plant experiment, Cassida had no significant effect on shoot growth and development in the field-release experiment. The results of this study indicate that competition from typical NZ pasture species is a more important factor than herbivory by Cassida, and unless Cassida reaches outbreak densities, it will likely have an insignificant impact on this weed.     

Occurrence of Ditylenchus weischeri and Not D. dipsaci in Field Pea Harvest Samples and Cirsium arvense in the Canadian Prairies

The stem nematode, a parasite of the herbaceous perennial weed, Cirsium arvense (L.) Scop. and identified as Ditylenchus dipsaci (Kühn) Filipjev, was reported in the Canadian prairies in 1979. Recently, D. weischeri Chizhov parasitizing Cirsium arvense was described in Russia, and it has been shown that this species is not an agricultural pest. In this study, we examined Ditylenchus species found in field pea (Pisum sativum L.) grain harvest samples in 2009 and 2010 and from C. arvense shoots in pea fields in the Saskatchewan, Alberta, and Manitoba provinces. Samples from 538 fields (mainly yellow pea) were provided by 151 growers throughout the main pea-growing area of the Canadian prairies. Of the samples collected, 2% were positive for Ditylenchus. The population density of the nematode ranged between 4 and 1,500 nematodes kg^-1 pea harvest sample and related to presence of C. arvense seeds. Positive samples occurred in 2009 but not in 2010 and were from throughout the pea-growing area of the Canadian prairies and not related to cropping history. C. arvense collected from yellow pea fields in Saskatchewan and Manitoba, but not Alberta, were infested with Ditylenchus. Morphological and molecular (ITS-PCR-RFLP) traits indicated that this species belongs to D. weischeri. The results indicated the stem nematode found in yellow pea grain is D. weischeri which resided with C. arvense seeds and debris to pea samples. Unlike D. dipsaci, D. weischeri is not a nematode pest of economic importance; therefore, its presence in the pea harvest samples was not a concern.     

Anatomical Injury Induced by the Eriophyid Mite Aceria anthocoptes on the Leaves of Cirsium arvense

Anatomical injury of the leaves of the invasive species, Cirsium arvense (L.) Scop., caused by the eriophyid mite Aceria anthocoptes (Nal.), which is the only eriophyid mite that has been recorded on C. arvense worldwide, is described. The injury induced by the mite feeding on the leaves of C. arvense results in visible russeting and bronzing of the leaves. Other conspicuous deformations are folding and distortion of the leaf blade and curling of leaf edge, as well as gradual drying of leaves. The anatomical injury of the mature leaves of field-collected plants was limited to the epidermis of the lower leaf surface. However, on young leaves of experimentally infested plants, rust mite injuries extend to epidermal cells on both leaf surfaces and to those of deeper mesophyll layers. On these leaves, lesions on the lower leaf surface even affected the phloem of the vascular bundles. Leaf damage induced by A. anthocoptes is discussed with regard to the mite’s potential as a biological control agent of C. arvense.     

Mowing strategies for controlling Cirsium arvense in a permanent pasture in New Zealand compared using a matrix model

Defoliation has frequently been proposed as a means of controlling Cirsium arvense (L.) Scop. (Californian thistle, Canada thistle, creeping thistle, perennial thistle), an economically damaging pastoral weed in temperate regions of the world, but its optimization has remained obscure. We developed a matrix model for the population dynamics of C. arvense in sheep‐grazed pasture in New Zealand that accounts for the effects of aerial shoot defoliation on a population's photosynthetic opportunity and consequential overwintered root biomass, enabling mowing regimes varying in the seasonal timing and frequency of defoliation to be compared. The model showed that the long‐term population dynamics of the weed is influenced by both the timing and frequency of mowing; a single‐yearly mowing, regardless of time of year, resulted in stasis or population growth, while in contrast, 14 of 21 possible twice‐yearly monthly mowing regimes, mainly those with mowing in late spring, summer, and early autumn, resulted in population decline. Population decline was greatest (with population density halving each year) with twice‐yearly mowing either in late spring and late summer, early summer and late summer, or early summer and early autumn. Our results indicate that mowing can be effective in reducing populations of C. arvense in pasture in the long term if conducted twice each year when the initial mowing is conducted in mid spring followed by a subsequent mowing from mid summer to early autumn. These mowing regimes reduce the photosynthetic opportunity of the C. arvense population and hence its ability to form the overwintering creeping roots upon which population growth depends.     

Spatial Segregation of Congeneric Invaders in Central Pennsylvania, USA

Carduus acanthoides and Carduus nutans (plumeless and musk thistles) are among the most noxious weeds in the United States of America, presenting a serious challenge in cropping and pasture systems. Unfortunately, a lack of detailed spatial distribution information hampers both our ability to understand the factors affecting their invasive success, and the effectiveness of monitoring and management efforts. To examine patterns of distribution and co-occurrence at a local level, we sampled a 5000 km² area of central Pennsylvania that cut a transect across known areas of C. acanthoides and C. nutans infestation. A number of potential environmental explanatory variables were recorded and analyzed to examine whether they correlated with observed species distribution patterns. Patterns of forest density and spatial aggregation of the thistles were the primary covariates that significantly impacted both species’ distributions. The survey established that the frequency of sightings for each species diminished as the ranges converged, with only brief overlap: the two species are strongly negatively correlated in space. Understanding environmental correlates of infestation and the pattern of spatial dissociation of these two invasive species is an important step towards an improved understanding of the mechanisms underlying their invasive potential, and hence towards effective weed control.     

Direct and Indirect Effects of a Shoot-Base Boring Weevil and Plant Competition on the Performance of Creeping Thistle, Cirsium arvense

Creeping thistle or Canada thistle, Cirsium arvense (L.) Scop., is considered one of the world's worst weeds and the third most important weed in Europe. Biological control of this indigenous weed in Europe by use of native agents may provide a low-cost alternative to use of chemical or mechanical control measures and contribute to a more sustainable weed management. We investigated the potential of a shoot-base boring weevil, Apion onopordi Kirby (Coleoptera: Apionidae), for biological weed control, in the presence or absence of plant competition by three grass species. Infestation of thistle shoots by A. onopordi at natural infestation levels reduced above- and belowground plant performance after 2 years. Plant competition at natural levels had an overall greater effect than that of herbivory, significantly reducing both above- and belowground thistle performance in both years, thereby slowing the propagation of the weed. Weevil infestation and grass competition had a synergistic effect on C. arvense growth; the combined effects of the two factors was greater than the sum of both single-factor effects. The experiment revealed that A. onopordi promotes systemic infections of the rust fungus Puccinia punctiformis (Str.) Röhl in the year following weevil infestation. Systemically infected thistle shoots died before the end of the growing season. Although the direct effect of A. onopordi may not be sufficient to control creeping thistle, the synergistic interaction with plant competition and the indirect effect via promotion of systemic rust infections makes A. onopordi a promising agent for the biological control of this weed.     

Host finding and recognition by Altica carduorum, a defoliator of Cirsium arvense

Altica carduorum Guer. (Coleoptera: Chrysomelidae) populations aggregate on vigorously growing stands of its host, the thistle Cirsium arvense (L.) Scop. (Asteraceae). The beetle needs contact to recognize a Cirsium species and biting to distinguish its host C. arvense. Feeding or mechanical damage on C. arvense, but not other Cirsium species, aggregates the beetles. Also feces from feeding on C. arvense, but not other Cirsium species, aggregates the opposite sex. Adults also aggregate on larval feces. We suggest that initial host location in the spring depends on random encounter by beetles overwintering in a stand of C. arvense. The beetles commonly make short flights after feeding and either land on another C. arvense plant, which establishes a secondary aggregation centre, or return to the original one. In this manner beetles released on a stand of C. arvense in China spread a radius of 600 m in a year, but those released 35 m away from their host failed to find it. Altica carduorum will develop on any Cirsium species to which it is confined, but is monophagous in the field because host finding is dependent on aggregation to wound and feces substances that are specific to C. arvense.     

Field assessment of a frond-feeding weevil, a successful biological control agent of red waterfern, Azolla filiculoides, in southern Africa

Azolla filiculoides (red waterfern) is a small, floating fern native to South America, that has invaded aquatic habitats, predominantly water resevoirs in southern Africa. A frond-feeding weevil, Stenopelmus rufinasus Gyllenhal (Coleoptera: Curculionidae), was imported from Florida, USA, and released as a biological control agent against this weed in South Africa at the end of 1997. To date, 24,700 weevils have been released, which has resulted in local extinction of red waterfern at 81% of the 112 release sites. The weevil has not failed to control a single site. Several sites were, however, lost due to flooding or drainage of dams. The surface area of weed controlled totalled 203.5 ha. On average, A. filiculoides was controlled in infested sites in 6.9 (±4.3) months. The weed recolonized at 22 of the sites (through either spore germination or dispersal by waterfowl), but the weevils subsequently spread to all of these sites and successfully caused local extinction of the weed at 18 of the sites. Five years after the release of the weevil, the weed no longer poses a threat to aquatic systems in southern Africa. In comparison to other biological control programs of aquatic weeds, the program against A. filiculoides in southern Africa ranks among the most successful cases anywhere in the world.     

Ecological niche modelling of an invasive alien plant and its potential biological control agents

Invasive alien plants are of concern in South Africa. Pompom weed (Campuloclinium macrocephalum) is currently invading the Grassland and Savannah biomes of South Africa and is likely to continue spreading in the southern African sub- region. Two possible biological control agents (Liothrips tractabilis and Cochylis campuloclinium) have been identified for control of pompom weed. We used ecological niche modelling to predict which areas in southern Africa are likely to be suitable for pompom weed and the two potential biological control agents. The overlap between areas predicted to be highly suitable for pompom weed and areas suitable for the biological control agents was assessed. Methods of reducing sampling bias in a data set used for calibrating models were also compared. Finally, the performance of models calibrated using only native range data, only invaded range data and both were also compared. Models indicate that pompom weed is likely to spread across a greater region of southern Africa than it currently occupies, with the Savannah and Grassland biomes being at greatest risk of invasion. Poor overlap was found between the areas predicted to be highly suitable for pompom weed and those areas predicted to be suitable for the biological control agents. However, models of the potential distribution of the biological control agents are interpreted with caution due to the very small sample size of the data set used to calibrate the models. Models calibrated using both native range and invaded range data were found to perform best whilst models calibrated using only native range data performed the worst. There was little difference found between models that were calibrated using spatially reduced (selecting only one record per 30 min grid cell) and randomly reduced (randomly selecting 50% of available records) biased data sets.     

Enemy release does not increase performance of <Emphasis Type="Italic">Cirsium</Emphasis> <Emphasis Type="Italic">arvense</Emphasis> in New Zealand

Cirsium arvense (L.) Scop. (Californian, Canada, or creeping thistle) is an exotic perennial herb indigenous to Eurasia that successfully established in New Zealand (NZ) approximately 130 years ago. Presently, C. arvense is considered one of the worst invasive weeds in NZ arable and pastoral productions systems. A mechanism commonly invoked to explain the apparent increased vigour of introduced weeds is release from natural enemies. The enemy-release hypothesis (ERH) predicts that plants in an introduced range should experience reduced herbivory, particularly from specialists, and that release from this natural enemy pressure facilitates increased plant performance in the introduced range. In 2007, surveys were carried out in 13 populations in NZ (7 in the North Island and 6 in the South Island) and in 12 populations in central Europe to quantify and compare growth characteristics of C. arvense in its native versus introduced range. Altitude and mean annual precipitation for each population were used as covariates in an attempt to explain differences or similarities in plant traits among ranges. All plant traits varied significantly among populations within a range. Shoot dry weight was greater in the South Island compared to Europe, which is in line with the prediction of increased plant performance in the introduced range; however, this was explained by environmental conditions. Contrary to expectations, the North Island was not different from Europe for all plant traits measured, and after adjustment for covariates showed decreased shoot density and dry weight compared to the native range. Therefore, environmental factors appear to be more favourable for growth of C. arvense in both the North and South Islands. In accordance with the ERH, there was significantly greater endophagous herbivory in the capitula and stems of shoots in Europe compared to both NZ ranges. In NZ, capitulum attack from Rhinocyllus conicus was found only in the North Island, and no stem-mining attack was found anywhere in NZ. Thus, although C. arvense experiences significantly reduced natural enemy pressure in both the North and South Islands of NZ there is no evidence that it benefits from this enemy release.     

Successful biological control of mist flower (Ageratina riparia) in New Zealand: Agent establishment, impact and benefits to the native flora

The white smut fungus (Entyloma ageratinae) and the gall fly (Procecidochares alani) were released in New Zealand in 1998 and 2001 respectively to suppress mist flower (Ageratina riparia). The fungus established and spread rapidly, crossing 80 km of sea to Great Barrier Island within 2 years. The mean number of P. alani galls increased exponentially to 1.96/stem at release sites, but dispersal was slow. The impact of the biocontrol agents was monitored once annually from 1998/99 to 2003/04, at up to 51 sites in the North Island. The mean percentage of live leaves infected with fungus rapidly reached nearly 60%. Maximum plant height declined significantly. In heavy infestations, mean percentage cover of mist flower declined from 81 to 1.5%. Galls were only recorded towards the end of the impact study, and at low mean numbers. As mist flower declined, the species richness and mean percentage cover of native plants increased. In contrast, the species richness and mean percentage cover of exotic plants (excluding mist flower) did not change significantly. Many plant species colonizing the plots were important native mid- or late-successional shrubs or trees. With few exceptions, the exotic plant species common in the plots were not weeds that appeared to threaten native forest habitats. There was only a weak “replacement weed effect” from the potentially serious invader African club moss (Selaginella kraussiana). These data, together with reports of reduced threats to rare endemic plants from mist flower, suggest this rapid, well-monitored weed biocontrol program was very successful.     

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.     

AreAltica carduorumandAltica cirsicola(Coleoptera: Chrysomelidae) Different Species? Implications for the Release ofA. cirsicolafor the Biocontrol of Canada Thistle in Canada

Altica carduorumandAltica cirsicolaare two species of leaf-feeding beetle (Coleoptera: Chrysomelidae) which appear to be morphologically indistinguishable and reportedly hybridize in the laboratory. A European population ofA. carduorumwas previously screened for host-plant specificity and released in North America for the control of Canada thistle,Cirsium arvense.A population ofA. cirsicolafrom China is currently being considered as a biocontrol agent forC. arvenseand, as a different beetle species, must be screened using host-specificity tests similar to those used forA. carduorum.IfA. carduorumandA. cirsicolaare, in fact, one species, the screening requirements forA. cirsicolacould be significantly reduced. Hence, we investigated the taxonomic relationship betweenA. carduorumandA. cirsicolausing morphometric analyses, hybridization experiments, and DNA fingerprinting using random amplified polymorphic DNA (RAPD) analysis. Discriminant function analyses indicate thatA. carduorumandA. cirsicolacannot be reliably distinguished by their morphologies, and interspecific matings produce fertile F₁offspring. However, because interspecific matings produce significantly fewer offspring than intraspecific matings, and because of clear differences in their DNA profiles, we conclude thatA. carduorumandA. cirsicolaare separate species. This study serves to highlight the value of genetic analyses in taxonomic studies and their role in biological control programs.     

The potential for indirect effects between a weed, one of its biocontrol agents and native herbivores: A food web approach

The safety of biological control is a contentious issue. We suggest that constructing and analyzing food webs may be a valuable addition to standard biological control research techniques, as they offer a means of assessing the post-release safety of control agents. Using preliminary data to demonstrate the value of food webs in biocontrol programs, we quantified the extent to which a key agent has infiltrated natural communities in Australia and, potentially, impacted on non-target species. Using these data, we also demonstrate how food webs can be used to generate testable hypotheses regarding indirect interactions between introduced agents and non-target species. We developed food webs in communities invaded to varying degrees by an exotic weed, bitou bush, Chrysanthemoides monilifera ssp. rotundata, and a key biocontrol agent for this weed in Australia, the tephritid fly, Mesoclanis polana. Three food webs were constructed during springtime showing the interactions between plants, seed-feeding insects and their parasitoids. One food web was constructed in a plot of native Australian vegetation that was free of bitou bush (‘bitou-free’), another in a plot of Australian vegetation surrounded by an invasion of bitou bush (‘bitou-threatened’) and a third from a plot infested with a monoculture of bitou bush (‘bitou-infested’). The bitou-free web contained 36 species, the bitou-threatened plot 9 species and the bitou-infested web contained 6 species. One native Australian herbivore attacked the seeds of bitou bush. M. polana, a seed-feeding fly, was heavily attacked by native parasitoids, these being more abundant than the parasitoids feeding on the native seed feeders. A surprising result is that none of the three species of native parasitoids reared from M. polana were reared from any of the native herbivores. The food webs revealed how a highly host-specific biocontrol agent, such as M. polana has the potential to change community structure by increasing the abundance of native parasitoids. The webs also suggest that indirect interactions between M. polana and native non-target species are possible, these been mediated by shared parasitoids. The experiments necessary to determine the presence of these interactions are outlined.     

Pre-release monitoring of Alliaria petiolata (garlic mustard) invasions and the impacts of extant natural enemies in southern Michigan forests

Monitoring of populations of a target weed species prior to releasing natural enemies has the potential to improve the rigor and safety of biological control and to determine the invader’s impacts on native communities while creating a reference point for evaluating the efficacy of subsequent biocontrol agent releases. Eight populations of garlic mustard, Alliaria petiolata (M. Bieb) Cavara and Grande (Brassicaceae), an invasive weed in southern Michigan, were monitored in anticipation of releases of classical biological control agents. The A. petiolata populations were shown to be expanding with 44.4% of initially uninvaded sampling quadrats becoming invaded after four years. While 88.2% of quadrats with A. petiolata showed evidence of foliar damage from pathogens and browsing by mammals, insects and other invertebrates, levels of damage were low and had little impact on rosette or seedling survival. Contrary to expectations, damage was positively correlated with A. petiolata fecundity (P = 0.0465). Given the continued expansion of A. petiolata and the lack of significant herbivore damage by acquired natural enemies, a biological control program should be considered against this invasive plant. If biological control agents are released, the results of this study will provide a benchmark for evaluating their performance.     

What Magnitude Are Observed Non-Target Impacts from Weed Biocontrol?

A systematic review focused by plant on non-target impacts from agents deliberately introduced for the biological control of weeds found significant non-target impacts to be rare. The magnitude of direct impact of 43 biocontrol agents on 140 non-target plants was retrospectively categorized using a risk management framework for ecological impacts of invasive species (minimal, minor, moderate, major, massive). The vast majority of agents introduced for classical biological control of weeds (>99% of 512 agents released) have had no known significant adverse effects on non-target plants thus far; major effects suppressing non-target plant populations could be expected to be detectable. Most direct non-target impacts on plants (91.6%) were categorized as minimal or minor in magnitude with no known adverse long-term impact on non-target plant populations, but a few cacti and thistles are affected at moderate (n = 3), major (n = 7) to massive (n = 1) scale. The largest direct impacts are from two agents (Cactoblastis cactorum on native cacti and Rhinocyllus conicus on native thistles), but these introductions would not be permitted today as more balanced attitudes exist to plant biodiversity, driven by both society and the scientific community. Our analysis shows (as far as is known), weed biological control agents have a biosafety track record of >99% of cases avoiding significant non-target impacts on plant populations. Some impacts could have been overlooked, but this seems unlikely to change the basic distribution of very limited adverse effects. Fewer non-target impacts can be expected in future because of improved science and incorporation of wider values. Failure to use biological control represents a significant opportunity cost from the certainty of ongoing adverse impacts from invasive weeds. It is recommended that a simple five-step scale be used to better communicate the risk of consequences from both action (classical biological control) and no action (ongoing impacts from invasive weeds).     

Conidial germination and germ tube elongation of Phomopsis amaranthicola and Microsphaeropsis amaranthi on leaf surfaces of seven Amaranthus species: Implications for biological control

Microsphaeropsis amaranthi and Phomopsis amaranthicola are potential biological control agents for several Amaranthus species. In an effort to understand the initial infection processes with these pathogens, a study was conducted of the conidial germination and germ tube length (μm) on the weed leaf surfaces at 21 °C and 28 °C. Weeds included Amaranthus rudis, A. palmeri, A. powellii, A. retroflexus, A. spinosus, A. hybridus, and A. albus. For P. amaranthicola, conidial germination and germ tube length varied among the seven weed species at both temperatures, while for M. amaranthi the differences in germ tube lengths were significant among weed species only at 21 °C. While the conidia of M. amaranthi and P. amaranthicola germinated on the leaf surfaces of all seven weed species, temperature appeared to impact the number and length of germ tubes on the leaf surfaces. The percentage of germinated conidia and the length of germ tubes at both temperatures were often greater for M. amaranthi than for P. amaranthicola. In order for the fungal pathogen to successfully infect and kill a weedy host, conidia must germinate and form a germ tube, two processes that vary with host species and temperature for M. amaranthi and P. amaranthicola. The extent to which successive infection processes, e.g., penetration, invasion and colonization, contribute to host specificity warrants study.     

Influence of insects and fungal pathogens on individual and population parameters of <Emphasis Type="Italic">Cirsium arvense</Emphasis> in its native and introduced ranges

Introduced weeds are hypothesized to be invasive in their exotic ranges due to release from natural enemies. Cirsium arvense (Californian, Canada, or creeping thistle) is a weed of Eurasian origin that was inadvertently introduced to New Zealand (NZ), where it is presently one of the worst invasive weeds. We tested the ‘enemy release hypothesis’ (ERH) by establishing natural enemy exclusion plots in both the native (Europe) and introduced (NZ) ranges of C. arvense. We followed the development and fate of individually labelled shoots and recorded recruitment of new shoots into the population over two years. Natural enemy exclusion had minimal impact on shoot height and relative growth rate in either range. However, natural enemies did have a significant effect on shoot population growth and development in the native range, supporting the ERH. In year one, exclusion of insect herbivores increased mean population growth by 2.1–3.6 shoots m⁻², and in year two exclusion of pathogens increased mean population growth by 2.7–4.1 shoots m⁻². Exclusion of insect herbivores in the native range also increased the probability of shoots developing from the budding to the reproductive growth stage by 4.0× in the first year, and 13.4× in the second year; but exclusion of pathogens had no effect on shoot development in either year. In accordance with the ERH, exclusion of insect herbivores and pathogens did not benefit shoot development or population growth in the introduced range. In either range, we found no evidence for an additive benefit of dual exclusion of insects and pathogens, and in no case was there an interaction between insect and pathogen exclusion. This study further demonstrates the value of conducting manipulative experiments in the native and introduced ranges of an invasive plant to elucidate invasion mechanisms.     

Can Invasive Species Enhance Competitive Ability and Restoration Potential in Native Grass Populations?

Native plant individuals often persist within communities dominated by exotics but the influence of this exposure on native populations is poorly understood. Selection for traits contributing to competitive ability may lead to native plant populations that are more tolerant of the presence of exotic invaders. In this way, long‐term coexistence with an exotic may confer competitive advantages to remnant (experienced) native populations and be potentially beneficial to restoration. In past studies we have documented genetic differentiation within native grass populations exposed to the exotic invader Russian knapweed (Acroptilon repens). Here, we examine populations of a cool‐season grass, needle‐and‐thread (Hesperostipa comata [Trin. & Rupr.]) and a warm season, alkali sacaton (Sporobolus airoides [Torr.]) collected from Russian knapweed‐invaded sites and adjacent noninvaded sites to assess their relative competitive ability against a novel exotic neighbor, Canada thistle (Cirsium arvense). Experienced S. airoides (from within A. repens invasions) appear to better tolerate (accumulate biomass, leaf nitrogen content, and to initiate new tillers) the presence of a novel competitor (C. arvense). Experienced and inexperienced H. comata genets differ in their response to the presence of C. arvense. Relative neighbor effects of native grasses on C. arvense were generally greater from experienced grasses. The ability to compete with novel neighbors may be driven by general competitive traits rather than species‐specific coevolutionary trajectories. Irrespective of competitive mechanisms, the conservation of native species populations within weed invasions may provide an important restoration tool by retaining unique components of native gene pools selected by competitive interactions with exotics.