Chemistry of Cirsium and Carduus: a role in ecological risk assessment for biological control of weeds?

Prediction of host plant range and ecological impact of exotic phytophagous insects, such as insects for classical biological control of weeds, represents a major challenge. Recently, the flowerhead weevil (Rhinocyllus conicus Fröl.), introduced from Europe into North America to control exotic thistles (Carduus spp.), has become invasive. It feeds heavily on some, but not all species of native North American thistles (Cirsium spp.). We hypothesized that such non-target use among native plants could be better predicted by knowledge of characteristic chemical profiles of secondary compounds to supplement the results of host specificity testing. To evaluate this hypothesis, we reviewed the literature on the chemistry of Cirsium and Carduus thistles. We asked what compounds are known to be present, what is known about their biological activity, and whether such information on chemical profiles would have better predicted realized host range and ecological effects of R. conicus in North America. We found an extensive, but incomplete literature on the chemistry of true thistles. Two main patterns emerged. First, consistent chemical similarities and interesting differences occur among species of thistles. Second, variation occurs in biologically active groups of characteristic compounds, specifically flavonoids, sterols, alkaloids and phenolic acids, that are known to influence host plant acceptance, selection, and feeding by phytophagous insects. Surprisingly, sesquiterpene lactones, which are characterisitic in closely related Asteraceae, have not been extensively reported for Cirsium or Carduus. The minimal evidence on sesquiterpene lactones may reflect extraction methods vs. true absence. In summary, our review suggests further research on thistle chemistry in insect feeding is warranted. Also, since the exotic Canada thistle (Cirsium arvense) is an invasive thistle of current concern in North America, such research on mechanisms underlying host range expansion by exotic insects would be useful.     

Tests of the antibiotic properties of the invasive vine Vincetoxicum rossicum against bacteria, fungi and insects

We tested extracts of Vincetoxicum rossicum for inhibition of bacterial and fungal growth and for anti-insect activities that may account for the invasive characteristics of this introduced species in North America. Bioassay-guided fractionation was used to identify (−)-antofine as the principle inhibitor of bacteria and fungi in root extracts. This compound had especially pronounced antifungal activity, inhibiting the growth of diverse taxa that include yeast-like and filamentous fungi and, notably, broad-host-range plant pathogens. A second compound(s), that is as yet uncharacterized but distinct from (−)-antofine, was detected as having antifeedant activity against a larval hymenopteran, rose sawfly (Allantus cinctus), and toxicity to two larval lepidopterans, the masked birch caterpillar (Drepana arcuata) and the European corn borer (Ostrinia nubilalis). That V. rossicum contains potent inhibitors of plant pathogenic fungi, diverse bacteria, and herbivorous insects likely contributes to its success as an invasive species.     

Application of a frequency distribution method for determining instars of Eucryptorrhynchus brandti (Coleoptera: curculionidae) from several morphological variables

Eucryptorrhynchus brandti (Harold) (Coleoptera: Curculionidae) is a major pest of Ailanthus altissima (Mill.) Swingle (Simaroubaceae), commonly known as tree-of-heaven, and A. altissima var. Qiantouchun in China. It is considered a potential biological control agent for tree-of-heaven in North America. The aim of this study was to use a frequency distribution method to determine the instars of field-collected larvae of E. brandti. We collected larval samples of various sizes from both the field and the laboratory and measured five morphological variables, including antenna spacing, mandible width, head-capsule width, ocellus spacing, and pronotum width. Based on the results of a frequency distribution method and Dyar’s rule, the larvae of E. brandti were divided into seven instars. Of the five variables measured, the width of the head capsule provided the best measurement for determining instar stage. The regression equation between the head-capsule widths and the instar number was y?=?0.324e^0.096x (R²?=?0.970).     

入侵害虫西花蓟马的生物学、危害及防治技术

西花蓟马Frankliniellaoccidentalis最早发生于北美地区,现已广泛分布于世界各地,是园艺作物最重要的害虫之一。早在1996年,西花蓟马就被我国农业部列为进境植物检疫潜在的危险性害虫,2003年夏季西花蓟马在北京部分保护地首次发现。该文从分布及危害、寄主植物、生物学特性和防治方法4个方面对该害虫进行了概述。     

Large Shift in Symbiont Assemblage in the Invasive Red Turpentine Beetle

Changes in symbiont assemblages can affect the success and impact of invasive species, and may provide knowledge regarding the invasion histories of their vectors. Bark beetle symbioses are ideal systems to study changes in symbiont assemblages resulting from invasions. The red turpentine beetle (Dendroctonus valens) is a bark beetle species that recently invaded China from its native range in North America. It is associated with ophiostomatalean fungi in both locations, although the fungi have previously been well-surveyed only in China. We surveyed the ophiostomatalean fungi associated with D. valens in eastern and western North America, and identified the fungal species using multi-gene phylogenies. From the 307 collected isolates (147 in eastern North America and 160 in western North America), we identified 20 species: 11 in eastern North America and 13 in western North America. Four species were shared between eastern North America and western North America, one species (Ophiostoma floccosum) was shared between western North America and China, and three species (Grosmannia koreana, Leptographium procerum, and Ophiostoma abietinum) were shared between eastern North America and China. Ophiostoma floccosum and O. abietinum have worldwide distributions, and were rarely isolated from D. valens. However, G. koreana and L. procerum are primarily limited to Asia and North America respectively. Leptographium procerum, which is thought to be native to North America, represented >45% of the symbionts of D. valens in eastern North America and China, suggesting D. valens may have been introduced to China from eastern North America. These results are surprising, as previous population genetics studies on D. valens based on the cytochrome oxidase I gene have suggested that the insect was introduced into China from western North America.     

How do biological control and hybridization affect enemy escape?

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

薇甘菊的生物防治及其天敌在中国的新发现

薇甘菊(Mikania mixeantha H.B.K.)为原产中南美洲的菊科假泽兰属攀缘草本植物,现在广泛分布于东南亚地区并对我国南方部分地区的农林业生产造成严重危害。由于施用化学除草剂会对环境造成污染,所以生物防治更为可靠。大量工作表明,一些真菌和昆虫有望被用来进行薇甘菊的生物控制。本文中报道的薇甘菊天敌小蓑蛾(Acanthopsyche sp.)为国内首次发现,为薇甘菊的生物防治提供了新的线索和启示。     

The structure of the phytophagous insect fauna on the introduced weed Solidago altissima in Switzerland

Solidago altissima L. was introduced into Europe as an ornamental plant from North America more than 100 years ago and the phytophagous insect fauna of it was recently examined in Switzerland where it has become an important weed in disturbed habitats. Rhizomes and aerial parts were examined and all insects collected in summer were tested in a no-choice feeding test. 18 out of 55 phytophagous insects were found feeding on S. altissima in Switzerland and for the remaining 37 the relation with the plant was not determined. The insects that have expanded their host range to feed on S. altissima since its introduction to Switzerland are almost solely opportunistic, unspecialized ectophages not closely attuned to the growth cycle of S. altissima. Only 4% of the insects were specialists and 9% endophagous. In contrast in North America S. altissima supports 25% specialists and 17% endophages. The native Solidago virgaurea L. in Switzerland supports many more specialists (28%) and endophages (23%) than S. altissima here. Possible reasons why almost none of these have switched to S. altissima are discussed. A comparison is made between the number of insects recruited by S. altissima in Switzerland with the number recruited by the exotic plants Heracleum mantegazzianum Som. & Lev. (Apiaceae) in Switzerland and Xanthium occidentale Bertol. (Asteraceae) in Australia. Reasons for different levels of insect recruitment are discussed. The high number of phytophagous insect species found on S. altissima in North America shows that there should be no shortage of possible control agents and any insects imported into Switzerland for the biological control of S. altissima would find largely unexploited food sources awaiting them.     

Comparison between a native and exotic adelgid as hosts for Laricobius rubidus (Coleoptera: Derodontidae)

Laricobius rubidus LeConte, native to eastern North America whose primary host is Pineus strobi Hartig, has been observed in association with Adelges tsugae, an exotic and lethal pest of hemlock trees in eastern North America. Pineus strobi and A. tsugae were used to evaluate host preference and suitability of this predatory beetle to determine in part its suitability as a potential biological control agent of A. tsugae. In a paired-choice oviposition test, L. rubidus preferred to oviposit in P. strobi ovisacs over those of A. tsugae. However, in the no-choice oviposition test, there was no significant difference in the mean number of eggs laid by L. rubidus females in P. strobi or A. tsugae ovisacs. There were no significant differences in larval developmental time or survivorship for L. rubidus reared on a diet of either P. strobi or A. tsugae. Laricobius rubidus completed development to the adult stage on A. tsugae indicating that it is a suitable host and therefore has the potential to contribute to biological control of A. tsugae in the eastern United States. The introduction of a congener, Laricobius nigrinus to eastern North America may result in competition with L. rubidus on A. tsugae, but the inability of L. nigrinus to complete development on P. strobi suggests that the two congeners will not compete on the primary host of L. rubidus.     

A retrospective analysis of known and potential risks associated with exotic toadflax-feeding insects

To date, eight exotic toadflax-feeding insect species have been accidentally or intentionally introduced to North America. Reports on their establishment and impact have been recorded for more than 60 years. Environmental risks linked to biological control of toadflax were identified in terms of host resources and undesirable impacts on the target species through the critical review of this record. Data gaps revealed during this retrospective analysis are addressed through suggestions for future research and associated experimental methodologies. Known and potential impacts of toadflax-feeding insects on both invasive toadflax and non-target species are examined. Recent programmatic demands for demonstrated agent efficacy and stringent host selectivity during the prerelease screening process clearly illustrate that classical biological control of invasive toadflax in North America is progressing beyond the so-called lottery approach.     

Insect herbivores associated with the introduced weed Bidens frondosa L. (Asteraceae) in Korea,and their potential role as augmentative biological control agents

The annual herb Bidens frondosa L., native to North America, is an invasive weed. Currently no information is available on the insect herbivores associated with this weed in Korea. A survey was carried out at two‐weekly intervals from May to October 2008 at two sites, and the incidence and abundance of various insect herbivores studied. A total nine species of insects was recorded and among them the defoliating caterpillar Hadjina chinensis (Wallengren) (Lepidoptera) was the only species known to have host plants restricted to genus Bidens. Further host specificity studies are required to evaluate the potential of this insect as a candidate for augmentative biological control agent for B. frondosa in Korea. All other insect species are either polyphagous or known crop pests.     

Predictable risk to native plants in weed biological control

Data on field host use of 112 insects, 3 fungi, 1 mite, and 1 nematode established for biological control of weeds in Hawaii, the continental United States, and the Caribbean indicate that the risk to native flora can be judged reliably before introduction. Virtually all risk is borne by native plant species that are closely related to target weeds. Fifteen species of insects introduced for biological control use 41 native plant species; 36 of which are congeneric with target weeds, while 4 others belong to two closely allied genera. Only 1 of 117 established biological organisms uses a native plant unrelated to the target weed. Thus the elements of protection for the native flora are the selection of weed targets that have few or no native congeners and the introduction of biological control organisms with suitably narrow diets.     

Avoiding conflicts between insect and weed biological control: selection of non-target species to assess host specificity of cabbage seedpod weevil parasitoids

Abstract:  Classical biological control of insect pests and weeds may lead to potential conflicts, where insect pests are closely related to weed biological control agents. Such a conflict may occur in the classical biological control of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) in North America, which belongs to the same subfamily, Ceutorhynchinae, as a number of agents introduced or proposed for introduction against non-indigenous invasive weed species. We propose a step-by-step procedure to select non-target species and thereby to develop a non-target species test list for screening candidate entomophagous biological control agents of a herbivore pest insect in a way that would simultaneously evaluate non-target potential on weed biological control agents and other non-target species. Using these recommendations, we developed a non-target test list for host specificity evaluations in the area of origin (Europe) and the area of introduction (North America) for cabbage seedpod weevil parasitoids. Scientifically based predictions on expected host–parasitoid interactions and ecological information about the ecological host range in the area of origin can help avoid conflicts, while still allowing the introduction of safe and effective agents against both insect pests and weeds.     

我国南方不同产量的八角树内生真菌群落研究

八角Illicium verum原产于我国华南和越南北部,是我国传统的香料和药用植物。内生真菌可提高植物生长与产量。然而,八角的产果量与其内生真菌间的关系尚未有人报道。为揭示内生真菌与八角产量的关系,本文分析了广西壮族自治区不同八角产量植株的内生真菌定殖率和群落组成。从60棵植株的叶片中分离到412株内生真菌,并根据形态学和分子生物学方法将其鉴定为22个真菌分类单元。内生真菌群落的优势类群分别为Colletotrichum gloeosporioides、Dermateaceae sp.和Phyllosticta sp. 1。内生真菌定殖率的变化范围为6%-100%,平均值为41%。内生真菌的定殖率与八角产量的回归分析表明,在较低和中等产量的八角植株中,内生真菌定殖率与产量呈正相关,而在高产的八角植株中,内生真菌定殖率与产量呈负相关。     

The colonization of native phytophagous insects in North America by exotic parasitoids

Classical biological control could have a major environmental cost if introduced natural enemies colonize and disrupt native systems. Although quantifying these impacts is difficult for systems already colonized by natural enemies, the a priori condition for such impacts can be evaluated based on the extent to which exotics have acquired native hosts. We use native host records for exotic parasitoids introduced into North America for biological control to document the number of exotic species that have been recorded from at least one native insect species. We also evaluate the ability of six biological and ecological variables to predict whether or not a parasitoid will move onto natives. Sixteen percent of 313 parasitoid species introduced against holometabolous pests are known from natives. Further, the likelihood that a parasitoid had colonized native hosts was largely unpredictable with respect to the independent variables. We conclude that given the quality of the data available either now or in the foreseeable future, coupled with inherent stochasticity in host shifts by parasitoids, there are no rules of thumb to assist biological control workers in evaluating if an introduced parasitoid will colonize native insect communities. Received: 2 July 1997 / Accepted: 3 August 1997     

Contemporary evolution of host plant range expansion in an introduced herbivorous beetle Ophraella communa

Host range expansion of herbivorous insects is a key event in ecological speciation and insect pest management. However, the mechanistic processes are relatively unknown because it is difficult to observe the ongoing host range expansion in natural population. In this study, we focused on the ongoing host range expansion in introduced populations of the ragweed leaf beetle, Ophraella communa, to estimate the evolutionary process of host plant range expansion of a herbivorous insect. In the native range of North America, O. communa does not utilize Ambrosia trifida, as a host plant, but this plant is extensively utilized in the beetle's introduced range. Larval performance and adult preference experiments demonstrated that native O. communa beetles show better survival on host plant individuals from introduced plant populations than those from native plant populations and they also oviposit on the introduced plant, but not on the native plant. Introduced O. communa beetles showed significantly higher performance on and preference for both introduced and native A. trifida plants, when compared with native O. communa. These results indicate the contemporary evolution of host plant range expansion of introduced O. communa and suggest that the evolutionary change of both the host plant and the herbivorous insect involved in the host range expansion.     

丛枝菌根真菌与植食性昆虫的相互作用

丛枝菌根(arbuscular mycorrhizal AM)真菌与昆虫均是陆地生态系统中的重要组分,同植物关系密切,对植物的影响和作用是巨大的。生态系统中则以AM真菌-植物-昆虫互作体系参预食物网与生态过程。早在20世纪80年代,人们已开始研究AM真菌对昆虫的影响。进入21世纪人们越来越重视AM真菌与昆虫的相互作用。总结了AM真菌对昆虫取食偏好、生长、繁殖和对植物危害等方面的影响、以及昆虫对AM真菌侵染、扩展和产孢的影响;分析了植物营养状况、昆虫性别、昆虫龄期和AM真菌种类等对AM真菌与昆虫相互作用的影响特点;探讨了AM真菌与昆虫相互作用的机制;展望了利用AM真菌抑制植食性害虫、及促进天敌昆虫和部分传粉昆虫作用的可能性,旨在丰富菌根学研究内容、促进AM真菌与昆虫互作领域的深入研究、为探索生物防控农林业害虫的新途径提供依据。     

Climate warming increases biological control agent impact on a non‐target species

Climate change may shift interactions of invasive plants, herbivorous insects and native plants, potentially affecting biological control efficacy and non‐target effects on native species. Here, we show how climate warming affects impacts of a multivoltine introduced biocontrol beetle on the non‐target native plant Alternanthera sessilis in China. In field surveys across a latitudinal gradient covering their full distributions, we found beetle damage on A. sessilis increased with rising temperature and plant life history changed from perennial to annual. Experiments showed that elevated temperature changed plant life history and increased insect overwintering, damage and impacts on seedling recruitment. These results suggest that warming can shift phenologies, increase non‐target effect magnitude and increase non‐target effect occurrence by beetle range expansion to additional areas where A. sessilis occurs. This study highlights the importance of understanding how climate change affects species interactions for future biological control of invasive species and conservation of native species.     

Founder effects and phenotypic variation in Adelges cooleyi, an insect pest introduced to the eastern United States

Introduced organisms experience founder effects including genetic bottlenecks that result in significant reductions in genetic variation. Genetic bottlenecks may constrain the evolution of phenotypic traits that facilitate success in novel habitats. We examined the effect of introduction into novel environments on genetic diversity of an insect pest, Adelges cooleyi, which was introduced into the eastern United States during the mid nineteenth century. We compared variation in mitochondrial and nuclear genomes in native and introduced samples to determine the effect of introduction on genetic variation experienced by this insect. We also measured an ecologically important phenotype, variation in host preference, in both native and introduced samples to compare variation in that trait with molecular genetic variation. To further investigate the relationship between genetic and phenotypic variation, we examined the degree to which mtDNA haplotypes provide information about host preference. Adelges cooleyi in eastern North America has significantly reduced genetic and phenotypic variation, but this low variation does not appear to have prevented persistence in a novel environment. Introduced insects appear to have retained host preference phenotypes similar to those of insects found where introductions likely originated.     

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.