Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

New techniques and findings in the study of a candidate allelochemical implicated in invasion success

Allelopathy has been hypothesized to promote the success of invasive plants. Support for the role of allelopathy in invasions has emerged from research on the candidate allelochemical (?)‐catechin, which is secreted by spotted knapweed. Here we describe new methods to quantify catechin in liquid and soil. With a new technique, we assayed catechin production by individual plants in liquid media and found levels up to two orders of magnitude less than previously reported. An acetone/water solution provided consistent recovery of catechin from soil, with percent recovery depending upon soil type. We evaluated soils from two spotted knapweed sites in Montana, USA, but found no measurable catechin. Idaho fescue, a native species reportedly sensitive to catechin, only exhibited slightly reduced growth at concentrations 10 times higher than previously reported to cause 100% mortality. Our results emphasize that more research is required to clarify the role of catechin in the invasion of spotted knapweed.     

模拟气候变暖对不同地区多斑矢车菊形态属性的影响

以北美及欧洲7个不同地区种源恶性杂草多斑矢车菊(Centaurea maculosa)为材料,采用红外辐射加热器模拟增温方法研究增温2℃(地上10 cm)对其生长及形态属性的影响。结果表明:与对照相比,增温使乌克兰、阿肯色、加拿大组多斑矢车菊株高显著增加,马里兰组株高显著减小。增温使多斑矢车菊叶片数量明显增加1~5片(乌克兰、马里兰组除外),叶片形态趋于变小;同时使加拿大组总叶面积显著增加,佛蒙特组总叶面积显著减小。增温使蒙大拿组多斑矢车菊地上生物量显著增加,罗马尼亚组却显著降低。增温使乌克兰、罗马尼亚、佛蒙特组多斑矢车菊地下生物量显著降低;除阿肯色组根冠比增加外,增温使其他地区多斑矢车菊根冠比下降。研究发现,7种多斑矢车菊对增温的响应(从促进到抑制)顺序依次为阿肯色组、加拿大组、蒙大拿组、乌克兰组、佛蒙特组、马里兰组、罗马尼亚组;不同地区多斑矢车菊对增温效应的适应能力不同,在生长及形态发生上存在差异,其潜在入侵潜力也可能不相同。     

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

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

Adaptive plasticity and niche expansion in an invasive thistle

Phenotypic differentiation in size and fecundity between native and invasive populations of a species has been suggested as a causal driver of invasion in plants. Local adaptation to novel environmental conditions through a micro‐evolutionary response to natural selection may lead to phenotypic differentiation and fitness advantages in the invaded range. Local adaptation may occur along a stress tolerance trade‐off, favoring individuals that, in benign conditions, shift resource allocation from stress tolerance to increased vigor and fecundity and, therefore, invasiveness. Alternately, the typically disturbed invaded range may select for a plastic, generalist strategy, making phenotypic plasticity the main driver of invasion success. To distinguish between these hypotheses, we performed a field common garden and tested for genetically based phenotypic differentiation, resource allocation shifts in response to water limitation, and local adaptation to the environmental gradient which describes the source locations for native and invasive populations of diffuse knapweed (Centaurea diffusa). Plants were grown in an experimental field in France (naturalized range) under water addition and limitation conditions. After accounting for phenotypic variation arising from environmental differences among collection locations, we found evidence of genetic variation between the invasive and native populations for most morphological and life‐history traits under study. Invasive C. diffusa populations produced larger, later maturing, and therefore potentially fitter individuals than native populations. Evidence for local adaptation along a resource allocation trade‐off for water limitation tolerance is equivocal. However, native populations do show evidence of local adaptation to an environmental gradient, a relationship which is typically not observed in the invaded range. Broader analysis of the climatic niche inhabited by the species in both ranges suggests that the physiological tolerances of C. diffusa may have expanded in the invaded range. This observation could be due to selection for plastic, “general‐purpose” genotypes with broad environmental tolerances.     

Interspecific interactions between the gall-fly Urophora affinis Frfld. (Diptera: Tephritidae) and the weevil Larinus minutus Gyll. (Coleoptera: Curculionidae), two biological control agents released against spotted knapweed, Centaurea stobe L. ssp. micranthos

Interspecific competition has been suggested as an explanation for the failure of some insects as biological control agents for weeds. Enclosure and exclusion cages were used, in southern British Columbia, Canada to evaluate the importance of interspecific competition between a seedhead weevil, Larinus minutus, and a gall-inducing fly, Urophora affinis, two biocontrol agents released against spotted knapweed in North America. At the seedhead scale, U. affinis, which is an inferior biological control agent based on knapweed seed mortality, was the superior competitor. Larinus minutus attack rates were significantly lower in the presence of U. affinis compared to release treatments where L. minutus was attacking alone. Reduced L. minutus attack rates were apparent in seed heads expected to contain both species, assuming insect distributions were random, but instead only contained U. affinis. L. minutus did not significantly affect U. affinis density. Although overall attack rates on knapweed seedheads were higher when both species were together at a site, the consequence of the antagonistic interaction is that overall seed destruction was not as high as it could have been if the weevil were attacking on its own. These results support minimizing the number of biocontrol agents released that use similar resources on the target weed, to avoid negative interactions between control agents and potential reductions in biocontrol efficacy resulting from competitive exclusion.     

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.     

Resolving the native provenance of invasive fireweed (Senecio madagascariensis Poir.) in the Hawaiian Islands as inferred from phylogenetic analysis

Accurate identification of weedy species is critical to the success of biological control programs seeking host-specific control agents. Phylogenetic relationships based on internal transcribed spacer region (ITS1, ITS2) DNA sequence data were used to elucidate the most likely origin and taxonomic placement of Senecio madagascariensis Poir. (fireweed; Asteraceae) in the Hawaiian archipelago. Putative S. madagascariensis populations from Madagascar, South Africa, Swaziland, and Hawaii were included in the analysis. Different phylogenetic models (maximum parsimony and maximum likelihood) were congruent in suggesting that Hawaiian fireweed is most closely related to populations from the KwaZulu-Natal region in South Africa. Phylogenetic divergence and morphological data (achene characteristics) suggest that the S. madagascariensis complex is in need of revised alpha-level taxonomy. Taxonomic identity of invasive fireweed in Hawaii is important for finding effective biological control agents as native range populations constitute different biotypic variants across a wide geographical area. Based on our phylogenetic results, research directed at biological control of Hawaiian infestations should focus on areas in the KwaZulu-Natal region in South Africa where host-specific natural enemies are most likely to be found. Our results show that phylogeographical analysis is a potential powerful and efficient tool to address questions relevant to invasion biology of plants.     

Temporal pattern of africanization in a feral honeybee population from Texas inferred from mitochondrial DNA

The invasion of Africanized honeybees (Apis mellifera L.) in the Americas provides a window of opportunity to study the dynamics of secondary contact of subspecies of bees that evolved in allopatry in ecologically distinctive habitats of the Old World. We report here the results of an 11-year mitochondrial DNA survey of a feral honeybee population from southern United States (Texas). The mitochondrial haplotype (mitotype) frequencies changed radically during the 11-year study period. Prior to immigration of Africanized honeybees, the resident population was essentially of eastern and western European maternal ancestry. Three years after detection of the first Africanized swarm there was a mitotype turnover in the population from predominantly eastern European to predominantly A. m. scutellata (ancestor of Africanized honeybees). This remarkable change in the mitotype composition coincided with arrival of the parasitic mite Varroa destructor, which was likely responsible for severe losses experienced by colonies of European ancestry. From 1997 onward the population stabilized with most colonies of A. m. scutellata maternal origin.     

Soil space and nutrients differentially promote the growth and competitive advantages of two invasive plants

Aims Invasive plants commonly occupy disturbed soils, thereby providing a stage for understanding the role of disturbance-enhanced resources in plant invasions. Here, we addressed how soil space and soil nutrients affect the growth and competitive effect of invasive plants and whether this effect varies with different invaders.Methods We conducted an experiment in which two invasive plants (Bromus tectorum and Centaurea maculosa) and one native species (Poa pratensis) were grown alone or together in four habitats consisting of two levels of soil space and nutrients. At the end of the experiment, we determined the total biomass, biomass allocation and relative interaction intensity of B. tectorum, C. maculosa and P. pratensis .Important findings Across two invaders, B. tectorum and C. maculosa, increased soil nutrients had greater positive effects on their growth than increased soil space, the effects of soil space on root weight ratio were greater than those of soil nutrients, and their competitive effect decreased with soil space but increased with soil nutrients. These findings suggest that changing soil space and nutrients differentially influence the growth and competitive advantages of two invaders. Bromus tectorum benefited more from increased soil resources than C. maculosa. Soil space and nutrients affected the biomass allocation of C. maculosa but not B. tectorum. The competitive effect of B. tectorum was unaffected by soil space and soil nutrients, but the opposite was the case for C. maculosa. Thus, the effects of soil space and nutrients on growth and competitive ability depend on invasive species identity.     

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.     

Influence of Centaurea biebersteinii patch size on Urophora quadrifasciata (Dipt. Tephritidae) in Michigan, USA

Abstract:  Host plant patch size plays an important role in the distribution and densities of many insect species. Understanding how the host plant patch size influences the population of a biological control agent is necessary to monitor the success of a biological control programme. Urophora quadrifasciata (Dipt. Tephritidae) was released in the early 1970s to control the spotted knapweed, Centaurea biebersteinii (Asteraceae) in North America. The studies reported here investigated the influences of spotted knapweed patch size and stem density on U. quadrifasciata larval populations overwintering within capitula in the Upper Peninsula of Michigan, USA. An inverse relationship was found between the percentage of capitula with U. quadrifasciata larvae and spotted knapweed patch size in 2002 from Houghton county, MI. In 2003, an inverse relationship was also found in the south region of the study area in Houghton and Keweenaw counties, Michigan. The percentage of capitula with U. quadrifasciata larvae and spotted knapweed patch size were positively related for the central region. No relationship was found for the North region in 2003. Dispersal of U. quadrifasciata is directed north-east along the Keweenaw Peninsula, Michigan. With this directed spread, areas with well-established populations of U. quadrifasciata would provide source individuals for new colonization. Spotted knapweed patch size may play an important role in the overwintering populations of U. quadrifasciata as a result of dispersal.     

Local pre‐adaptation to disturbance and inbreeding–environment interactions affect colonisation abilities of diploid and tetraploid Centaurea stoebe

• Primary colonisation in invasive ranges most commonly occurs in disturbed habitats, where anthropogenic disturbance may cause physical damage to plants. The tolerance to such damage may differ between cytotypes and among populations as a result of differing population histories (adaptive differentiation between ruderal verus natural habitats). Moreover, founder populations often experience inbreeding depression, the effects of which may increase through physical damage due to inbreeding–environment interactions. We aimed to understand how such colonisation processes differ between diploid and tetraploid Centaurea stoebe populations, with a view to understanding why only tetraploids are invasive.
• We conducted a clipping experiment (frequency: zero, once or twice in the growing season) on inbred versus outbred offspring originating from 37 C. stoebe populations of varying cytotype, range and habitat type (natural versus ruderal). Aboveground biomass was harvested at the end of the vegetation period, while re‐sprouting success was recorded in the following spring.
• Clipping reduced re‐sprouting success and biomass, which was significantly more pronounced in natural than in ruderal populations. Inbreeding depression was not detected under benign conditions, but became increasingly apparent in biomass when plants were clipped. The effects of clipping and inbreeding did not differ between cytotypes.
• Adaptive differentiation in disturbance tolerance was higher among populations than between cytotypes, which highlights the potential of pre‐adaptation in ruderal populations during early colonisation on anthropogenically disturbed sites. While the consequences of inbreeding increased through clipping‐mediated stress, they were comparable between cytotypes, and consequently do not contribute to understanding the cytotype shift in the invasive range.

     

Measuring the effects of invasive species on the demography of a rare endemic plant

The impacts of invasive species are among the greatest threats to the persistence of native species and communities. Yet most work on rare plants has focused on issues such as habitat fragmentation and genetic diversity, while few studies have quantified the impacts of invasive plants on native ones or investigated the underlying mechanisms of those impacts. I used removal experiments to assess the effects of invasive grasses on the seedling and adult demography of an endangered California endemic, Oenothera deltoidesssp. howellii. Invasive plant removal significantly increased O. deltoides seedling recruitment, but had no effect on adult plants. Differences in seedling recruitment were primarily driven by greatly increased seedling emergence rates in removal plots, although there was also some evidence of higher seedling survivorship with invasives removal. Differences in habitat type strongly influenced both the effects of removal treatments and O. deltoides demography, with areas that support natural recruitment showing weaker treatment effects and higher overall recruitment, but lower adult survivorship, compared to those under restoration through planting. These results indicate that inhibition of germination due to reduced soil disturbance, rather than resource competition, appears to be the strongest impact of invasive plants on this rare endemic. Although previous work has documented the importance of changed disturbance regimes in generating invasion impacts, invader effects on rare plants are generally presumed to result from resource competition. Studies like this one highlight the need for a greater emphasis on understanding the mechanisms by which invasive plants impact native ones, and the importance of such information in designing conservation and management strategies.     

Comparison of the effect of elevated CO2 on an invasive species (Centaurea solstitialis) in monoculture and community settings

The ongoing increase in atmospheric CO₂ concentration ([CO₂]) is likely to change the species composition of plant communities. To investigate whether growth of a highly invasive plant species, Centaurea solstitialis (yellow starthistle), was affected by elevated [CO₂], and whether the success of this species would increase under CO₂ enrichment, I grew the species in serpentine soil microcosms, both as a monoculture and as a component of a grassland community. Centaurea grown in monoculture responded strongly to [CO₂] enrichment of 350 mol mol^–1, increasing aboveground biomass production by 70%, inflorescence production by 74%, and midday photosynthesis by an average of 132%. When grown in competition with common serpentine grassland species, Centaurea responded to CO₂ enrichment with similar but nonsignificant increases (+69% aboveground biomass, +71% inflorescence production), while total aboveground biomass of the polyculture increased by 28%. Centaurea's positive CO₂ response in monoculture and parallel (but non-significant) response in polyculture provoke questions about possible consequences of increasing CO₂ for more typical California grasslands, where the invader already causes major problems.     

Defoliation of Centaurea solstitialis Stimulates Compensatory Growth and Intensifies Negative Effects on Neighbors

Efforts to arrest the spread of invasive weeds with herbivory may be hindered by weak effects of the herbivores or strong compensatory responses of the invaders. We conducted a greenhouse experiment to study the effects of defoliation and soil fungi on competition between the invasive weed Centaurea solstitialis and C. solstitialis and Avena barbata, a naturalized Eurasian annual grass, and Nassella pulchra, a native California bunchgrass. Surprisingly, considering the explosive invasion of grasslands by C. solstitialis, Avena and Nassella were strong competitors and reduced the invader’s biomass by 80.2% and 80.1% over all defoliation and soil fungicide treatments, respectively. However, our experiments were conducted in artificial environments where competition was probably accentuated. When fungicide was applied to the soil, the biomass of C. solstitialis was reduced in all treatment combinations, but reduction in the biomass of the invader had no corollary impact on the grasses. There was no overall effect of defoliation on the final biomass of C. solstitialis as the invader compensated fully for severe clipping. In fact, the directional trend of the clipping effect was +6.4% over all treatments after eight weeks. A significant neighbor × soil fungicide × clipping effect suggested that the compensatory response was the strongest without soil fungicide and when C. solstitialis was alone (+ 19%). Our key finding was that the compensatory response of C. solstitialis in all treatments was associated with an increase in the weed’s negative effects on Nassella and Avena – there was a significant decrease in the total biomass of both grasses and the reproductive biomass of Avena in pots with clipped C. solstitialis. Our results were obtained in controlled conditions that may have been conducive to compensatory growth, but they suggest the existence of mechanisms that may allow C. solstitialis, like other Centaurea species, to resist herbivory.     

Impacts of invasive plants on resident animals across ecosystems,taxa, and feeding types: a global assessment

As drivers of global change, biological invasions have fundamental ecological consequences. However, it remains unclear how invasive plant effects on resident animals vary across ecosystems, animal classes, and functional groups. We performed a comprehensive meta‐analysis covering 198 field and laboratory studies reporting a total of 3624 observations of invasive plant effects on animals. Invasive plants had reducing (56%) or neutral (44%) effects on animal abundance, diversity, fitness, and ecosystem function across different ecosystems, animal classes, and feeding types while we could not find any increasing effect. Most importantly, we found that invasive plants reduced overall animal abundance, diversity and fitness. However, this significant overall effect was contingent on ecosystems, taxa, and feeding types of animals. Decreasing effects of invasive plants were most evident in riparian ecosystems, possibly because frequent disturbance facilitates more intense plant invasions compared to other ecosystem types. In accordance with their immediate reliance on plants for food, invasive plant effects were strongest on herbivores. Regarding taxonomic groups, birds and insects were most strongly affected. In insects, this may be explained by their high frequency of herbivory, while birds demonstrate that invasive plant effects can also cascade up to secondary consumers. Since data on impacts of invasive plants are rather limited for many animal groups in most ecosystems, we argue for overcoming gaps in knowledge and for a more differentiated discussion on effects of invasive plant on native fauna.     

Control of the invasive weed Ambrosia artemisiifolia with Ophraella communa and Epiblema strenuana

Ambrosia artemisiifolia L. is native to North America, and was unintentionally introduced into China in the 1930s, where it has become invasive. The two insect species Epiblema strenuana and Ophraella communa have been considered as biological control agents of A. artemisiifolia in China. The purpose of this study was to examine the control effect of O. communa + E. strenuana on A. artemisiifolia in the field. The mortality of A. artemisiifolia plants increased with time and increasing initial release densities of O. communa and/or E. strenuana in 2008 and 2009. The treatments of 0.53 O. communa + 0.53 E. strenuana per plant and 12 O. communa + 16 E. strenuana per plant at early (60–70 cm tall) and later (90–100 cm tall) growth stages could subsequently kill all plants prior to seed production in both 2008 and 2009. Thus, the two initial densities of the two insect species may be recommended when they are jointly used for biological control of A. artemisiifolia at the two growth stages. However, all or some plants could survive and bear seeds in any other treatment and in the non-treated control plots. This implies that biological control of A. artemisiifolia with the two biological control agents will be recommended in the areas invaded by A. artemisiifolia in China.