Use of Risk Analysis for Screening Weed Biocontrol Agents: Altica carduorum Guer. (Coleoptera: Chrysomelidae) from China as a Biocontrol Agent of Cirsium arvense (L.) Scop. in North America

Candidate weed biocontrol agents must be screened to exclude those that could threaten desirable plants . Traditionally , this has been done by rejecting species that develop on economically important plants in laboratory no - choice tests . However , because congeneric plants often support development in these tests , even when they are not utilized in nature , the tests do not meet legislated requirements for rare plant species or the increasing public concern for native plants . Plant suitability for larval development is a poor predictor of host range because insects use a sequence of steps in which the early steps , such as host finding and acceptance for oviposition , tend to be stronger than the later ones , such as suitability for development . This study is a trial of a new approach to screening insects as weed biocontrol agents that uses risk analysis to quantify the suitability of a plant as a host on the basis of inset performance at various stages in its life cycle . The insects used for the study was a NW Chinese biotype of the leaf beetle , Altica carduorum, which in terms of climate adaptation and damage it inflicts on the weedy thistle Cirsium arvense, is a promising biocontrol agent for Canada . However , its ability to develop on all North American Cirsium spp . in laboratory no - choice tests currently excludes its release . We show by risk and factor analyses , with five sequential host - selection parameters , that the suitability of these thistles to A. carduorum is so low that the beetle would not jeopardize the continued existence of rare native thistles , so its release should not be a problem .     

Modelling and Mapping Geographic Ranges to Evaluate Weed Biocontrol Agents: A Case Study Using Altica carduorum (Coleoptera: Chrysomelidae) and Cirsium arvense (Asteraceae)

Leaf - feeding beetles , Altica carduorum, from a population in NW China have been identified as candidate agents for biocontrol of Canada thistle , Cirsium arvense. This paper assesses the potential of beetles from this population to establish on the Canadian prairies . A phenological model is applied to determine where sufficient heat accumulates to allow completion of at least one generation per year . The model was applied both with and without a submodel of adult thermoregulation . The model was driven using meteorological data from a grid covering the agricultural region of the Canadian prairies , i . e . ca . 670 000 km 2 , at a resolution of (50 50) km . In each grid square , the percentage of the years 1960 - 89 for which the model indicated that at least one generation could have been completed was calculated; these proportions were categorized (0 - 80%; 80 - 99% and 100%) and mapped . Maps of C. arvense density over the simulation region were also produced , and compared to the maps of predicted beetle distribution . The model suggests that A. carduorum could establish over much of the prairies . Without thermoregulation , the predicted range (i . e . the area in which one generation was completed in every year) is restricted to the warmer parts of the prairies , where C. arvense is generally sparse . Inclusion of thermoregulation almost doubled the predicted range to cover most of the range of C. arvense on the Canadian prairies , missing only the cooler peripheral areas . These results suggest that this strain of A. carduorum has potential to overlap the range of C. arvense on the Canadian prairies , and so has potential to control this weed .     

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.     

A methodology for risk analysis of plurivorous fungi in biological weed control: Sclerotinia sclerotiorum as a model

In New Zealand, research is being performed on the use of the ubiquitous plurivorous ascomycete Sclerotinia sclerotiorum, as a biocontrol agent for Cirsium arvense in pasture. As a consequence of the wide host range of this fungus, the proposed biocontrol may pose a risk to non-target arable plants. Crop disease risk is primarily due to the formation of sclerotia. These perennating bodies may directly infect susceptible crops sown at the biocontrol site in years after the biocontrol event. Additionally they may cause infection in neighbouring crops through the formation of apothecia from which airborne ascospores are dispersed. A methodology for a risk analysis is proposed, embodying both temporal and spatial processes, where the relative disease risk from biocontrol is defined as the ratio of added to naturally occurring inoculum.Exponential decay is proposed as a suitable model for the survival of sclerotia in soils beneath pastures, and data from an ongoing study are used in a preliminary parameterisation. The model is applied to published data on initial sclerotium population size and unpublished data on natural levels of soilborne sclerotia, to reveal how a withholding period, defined as the interval of time that must elapse for the ratio of added to natural inoculum to equal unity, may be calculated.For quantifying escape and aerial dispersal of ascospores from a treated pasture, three mechanistic models are proposed. Firstly, a simple one-layer model of escape, giving a provisional estimation of 81% of released ascospores escaping the pasture canopy, is discussed. Secondly, ascospore dispersal within and close to the biocontrol site (<30 m) is described by a model revealing abrupt decline in the atmospheric concentration of ascospores within short distances from the biocontrol site. Thirdly, for predicting ascospore concentrations at greater distances (>100 m), a Gaussian plume model is proposed. In a first approximation, these models are used in an example calculation to show how a hypothetical safety zone might be determined. A map of the province of Canterbury showing the locations of different agricultural activities and infestations of dense populations of C. arvense, revealed that potential biocontrol sites mostly do not border arable land and that a safety zone would therefore be feasible.The study presented here provides conceptual and mathematical frameworks for risk analysis of the use of plurivorous fungi in biological weed control, using S. sclerotiorum as an example. However, before a judgement may be made about the phytosanitory risks associated with the use of S. sclerotiorum to control C. arvense in pastures in Canterbury, the models developed in this paper must be parameterized using data collected under a wide range of conditions in the field.     

Importance of temperature for the performance and biocontrol efficiency of the parasitoid Perilitus brevicollis (Hymenoptera: Braconidae) on Salix

With the prospect of warmer temperatures as a consequence of ongoing climate change, it is important to investigate how such increases will affect parasitoids and their top-down suppression of herbivory in agroecosystems. Here we studied how the performance and biocontrol efficiency of the willow “bodyguard” Perilitus brevicollis Haliday (Hymenoptera: Braconidae) were affected at different constant temperatures (10, 15, 20, 25°C) when parasitizing a pest insect, the blue willow beetle (Phratora vulgatissima L., Coleoptera: Chrysomelidae). Parasitism did not reduce herbivory at all at 10°C, indicating poor biocontrol efficiency at low temperatures. At higher temperatures, however, parasitism reduced herbivory substantially, implying that biocontrol may be promoted by a warmer climate. Parasitoid performance (survival and development rate) generally increased with increasing temperature up to 20°C. The only exception was body size, which followed the temperature–size rule and decreased with increasing temperature. Our results indicate that a warmer climate may enhance the biocontrol of the blue willow beetle in environments that currently are cooler than the parasitoid’s optimal temperature for development.     

Effect of Herbivore Density, Timing of Attack and Plant Community on Performance of Creeping Thistle Cirsium arvense (L.) Scop. (Asteraceae)

Creeping thistle, Cirsium arvense (L.) Scop. (Asteraceae), is one of the most serious weeds in ecological compensation areas (within-field or border refugia) in Europe. Since conventional weed control measures are restricted in compensation areas, augmenting indigenous agents for biological control of the weed may be a feasible alternative. In this paper, we studied the effect of density of the shield beetle, Cassida rubiginosa Muller (Coleoptera, Chrysomelidae), and two vegetation types typical for ecological compensation areas, on the performance of creeping thistle plants in an open field experiment. Early-season larval feeding had no measurable effect on creeping thistle growth, while late-season feeding significantly reduced shoot growth. These findings were attributed to higher feeding rates of the herbivores at higher ambient temperatures late in the season. Defoliation had a strong effect on the above-ground performance of C. arvense plants, but not on the below-ground performance. In contrast, the plant community affected all below-ground performance parameters measured, but only some of the above-ground performance parameters of creeping thistle. A combination of high levels of plant competition and herbivory by C. rubiginosa larvae led to 50% mortality in C. arvense plants during the growing season. We conclude that augmentation of indigenous herbivores of C. arvense in combination with breaking up the root system by tillage and the establishment of a highly competitive plant community of beneficial herbs may be a feasible way to control this problematic weed in ecological compensation areas.     

Evidence for rapid evolutionary change in an invasive plant in response to biological control

We present evidence that populations of an invasive plant species that have become re‐associated with a specialist herbivore in the exotic range through biological control have rapidly evolved increased antiherbivore defences compared to populations not exposed to biocontrol. We grew half‐sib families of the invasive plant Lythrum salicaria sourced from 17 populations near Ottawa, Canada, that differed in their history of exposure to a biocontrol agent, the specialist beetle Neogalerucella calmariensis. In a glasshouse experiment, we manipulated larval and adult herbivory to examine whether a population's history of biocontrol influenced plant defence and growth. Plants sourced from populations with a history of biocontrol suffered lower defoliation than naïve, previously unexposed populations, strongly suggesting they had evolved higher resistance. Plants from biocontrol‐exposed populations were also larger and produced more branches in response to herbivory, regrew faster even in the absence of herbivory and were better at compensating for the impacts of herbivory on growth (i.e. they exhibited increased tolerance). Furthermore, resistance and tolerance were positively correlated among genotypes with a history of biocontrol but not among naïve genotypes. Our findings suggest that biocontrol can rapidly select for increased defences in an invasive plant and may favour a mixed defence strategy of resistance and tolerance without an obvious cost to plant vigour. Although rarely studied, such evolutionary responses in the target species have important implications for the long‐term efficacy of biocontrol programmes.     

Reproduction and Development of the Biocontrol AgentHydrellia pakistanae(Diptera: Ephydridae) on Monoecious Hydrilla

Previous investigations of the laboratory biology and host range ofHydrellia pakistanae,a biological control ofHydrilla verticillata(hydrilla), used the dioecious hydrilla biotype common to Florida, Texas, and California. A monoecious biotype that is now spreading throughout the mid-Atlantic states, California, and Washington was not investigated during these original studies. We therefore compared the dioecious and monoecious hydrilla biotypes as hosts forH. pakistanae.FemaleH. pakistanaeaccepted the two biotypes equally as ovipositional substrates. Overall developmental success differed little: 42% of eggs oviposited on monoecious hydrilla produced adults compared to 39% of eggs oviposited on dioecious hydrilla. Fly development required about 33 days on both biotypes (at 22 ± 2°C), but larvae that completed development mined 1.6 times as many leaves on monoecious hydrilla as on dioecious plants. These data suggest thatH. pakistanaewould be a useful biocontrol agent of monoecious hydrilla, should this plant invade areas where it can grow as a perennial.     

Effect of temperature on the survival of Aconophora compressa Walker (Hemiptera: Membracidae): implications for weed biocontrol

Abstract  In weed biocontrol, similarity of abiotic factors between the native and introduced range of a biocontrol agent is critical to its establishment and effectiveness. This is particularly the case for weeds that have a wide geographical distribution in the native range. For such weeds, the choice of a specialist insect that has narrow tolerance limits to important abiotic factors can diminish its ability to be an effective biocontrol agent. The membracid Aconophora compressa was introduced in Australia from Mexico for biocontrol of Lantana camara , a plant with a wide climatic tolerance. In this study we investigated the effect of constant and alternating temperatures on A. compressa survival. Longevity of adults and nymphs declined with increasing temperatures, and at 39°C individuals survived for less than a day. At lower temperatures, nymphs survived longer than adults. Survival at alternating temperatures was longer than at constant temperatures, but the general trend of lower survival at higher temperatures remained. Spatially and temporally, the climatic tolerance of A. compressa appears to be a subset of that of lantana, thereby limiting its potential impact.     

The host range of four new biotypes of Dactylopius tomentosus (Hemiptera: Dactylopiidae) from southern USA and their potential as biological control agents of Cylindropuntia spp. (Cactaceae) in Australia: Part II

Eight Cylindropuntia species have naturalised in Australia and pose serious economic, environmental and social impacts. The host range of four additional biotypes of D. tomentosus from southern USA was investigated. Feeding and development were restricted to the genus Cylindropuntia. However, they showed differences in specificity within this genus and some biotypes discriminated between the provenances of Cylindropuntia rosea and Cylindropuntia tunicata. Efficacy trials were conducted to determine whether populations of each biotype could be sustained on the naturalised Cylindropuntia species and if these populations could retard the growth or kill these plants. The ‘acanthocarpa’ biotype offers potential control of C. rosea (Lorne Station), while the ‘cylindropuntia sp.’ biotype shows great potential to control C. rosea (Grawin). The ‘cylindropuntia sp.’ biotype also had a high impact on Cylindropuntia kleiniae and Cylindropuntia imbricata, and a moderate impact on Cylindropuntia leptocaulis and Cylindropuntia prolifera. The ‘acanthocarpa?×?echinocarpa’ biotype had its greatest impact on C. tunicata (Grawin), killing this plant in 18 weeks. A fourth biotype, ‘leptocaulis’, was damaging to some species, but was less effective than the other biotypes. Cylindropuntia spinosior is the only naturalised species in Australia where no effective biocontrol agent has been found.     

Effects of constant and fluctuating temperatures on development of the coconut hispine beetle,Brontispa longissima (Coleoptera: Chrysomelidae) and two species of parasitoid

Abstract

The larval parasitoid Asecodes hispinarum Bou?ek has been used to control the coconut hispine beetle, Brontispa longissima (Gestro) in Vietnam. Although A. hispinarum has succeeded in suppressing B. longissima in southern Vietnam, biocontrol has not succeeded in central Vietnam. One factor underlying this failure might be the high temperature during the hot season. In the present work, we examined whether A. hispinarum parasitises the beetles within the range of temperatures that occur during the hot season in central Vietnam and we evaluated the potential use of the pupal parasitoid Tetrastichus brontispae Ferrière. Development of the beetle and two parasitoids was investigated at two constant temperatures (28 and 30°C) and at fluctuating temperatures from 25 to 35°C (corresponding to temperatures during the hot season in central Vietnam). B. longissima reached adulthood in all the temperature treatments. More than 70% of A. hispinarum emerged from hosts at 28°C, but none emerged at 30°C or under fluctuating temperature conditions, indicating that A. hispinarum cannot be used as a biological control agent in central Vietnam. However, T. brontispae could reproduce the next generation at all temperatures. These results suggest that T. brontispae is a potential agent for continuous biological control of B. longissima in central Vietnam.     

Strong effects of hydrologic environment and weak effects of elevated CO<Subscript>2</Subscript> on the invasive weed <Emphasis Type="Italic">Alternanthera philoxeroides</Emphasis> and the biocontrol beetle <Emphasis Type="Italic">Agasicles hygrophila</Emphasis>

Global change, such as elevated CO₂, may alter interactions between invasive plants and biocontrol agents, impacting biocontrol efficacy. Here, we conducted four experiments in Texas, USA to test how elevated CO₂ influences an invasive plant (Alternanthera philoxeroides) and its interactions with an introduced biocontrol beetle (Agasicles hygrophila) in terrestrial (well-watered) and flooded environments. We grew plants for 9 months in ambient or elevated CO₂ (800 ppm) chambers in continuously flooded or well-watered conditions. In no-choice trials, flooding increased leaf toughness and decreased beetle consumption but beetles only oviposited on ambient CO₂ leaves. In choice trials, beetles preferred to feed and oviposit on terrestrial plants but were also less likely to damage elevated CO₂ leaves. Caged beetle populations were larger in terrestrial conditions than aquatic conditions for a second set of plants grown in the chambers. With a third set of plants grown in the ambient or elevated CO₂ chambers, damage for plants placed in the field (aquatic setting) was higher for plants grown in terrestrial conditions vs. flooded conditions at ambient CO₂. Our results suggest that elevated CO₂ will have minor effects on the efficacy of this biocontrol agent by decreasing oviposition and number of leaves damaged, and hydrologic environment may affect invasive plant performance by altering herbivore oviposition and feeding preferences. A broader understanding of the effects of global change on biocontrol will help prevent and manage future spread of invasive plants.     

Isolation and characterization of microsatellite loci from Lochmaea suturalis, the heather beetle (Coleoptera: Chrysomelidae), a pest in Europe and a biocontrol agent in New Zealand

The heather beetle Lochmaea suturalis which is native to northwest Europe has been released as a biocontrol agent for heather in New Zealand. We have isolated and optimized eight microsatellite loci from New Zealand beetles. These loci provide markers with high polymorphism ranging from four to 20 alleles per locus. Observed heterozygosity averaged 0.631 per locus. These results suggest the markers are useful for population studies that will contribute to assessment of L. suturalis as a biocontrol agent.     

Biocontrol potential of tortoise beetle (Aspidomorpha miliaris) (Coleoptera: Chrysomelidae) on Ipomoea carnea in Assam,India

Ipomoea carnea Jacq. is a noxious aquatic weed with rapid vegetative growth and high invading powers. Infestation and population distribution of Aspidomorpha miliaris F. in Assam, India, showed that the beetle occurred on I. carnea at high densities (2.8–163.3 individuals/plant) causing damage up to 84%. Laboratory studies also support it as potential biocontrol agent for I. carnea.     

Host Specificity of Altica carduorum Guer. Chrysomelidae: Coleoptera , a Defoliator of Cirsium arvense L. Scop. Asteraceae from North-western China

Traditionally, insects are not approved for release in North America as agents for the biological control of weeds if the larvae complete development on desirable plants in no-choice tests. This criterion excludes the use of the leaf-feeding beetle Altica carduroum Guer. since it can complete development on all Cirsium spp., including rare North American species, although in nature it is only known from the target weed Cirsium arvense. Host suitability is the result of a sequence of selection factors, and this study shows that most discrimination between thistle species is the result of host finding and egg production, and not larval developmental ability. In no-choice tests, the larvae completed development on 18 Cirsium species in five sections of the genus that were tested and on Silybum marianum, but not on other genera in the tribe Cardueae or more distantly related Asteraceae. The adult food consumption on C. arvense was two to eight times greater than on other Cirsium species, many of which, such as C. pitcheri and C. drummondii, received no feeding by over 50 of the beetles in the first 24 h. The best North American thistle for egg production was C. scariosum, but this was only 28 of that on C. arvense. The preference for C. arvense in multiple-choice tests increased with the size 2 of the arena. In the largest, an area of 6 m , 80 of the beetles and eggs were found on C. arvense and none were found on plants outside the subtribe Carduineae. Thus, as theory predicts, host discrimination between thistle species takes place at earlier stages than suitability for larval development, and this should be considered when determining the safety of candidate agents.     

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.     

Climate warming affects biological invasions by shifting interactions of plants and herbivores

Plants and herbivorous insects can each be dramatically affected by temperature. Climate warming may impact plant invasion success directly but also indirectly through changes in their natural enemies. To date, however, there are no tests of how climate warming shifts the interactions among invasive plants and their natural enemies to affect invasion success. Field surveys covering the full latitudinal range of invasive Alternanthera philoxeroides in China showed that a beetle introduced for biocontrol was rare or absent at higher latitudes. In contrast, plant cover and mass increased with latitude. In a 2‐year field experiment near the northern limit of beetle distribution, we found the beetle sustained populations across years under elevated temperature, dramatically decreasing A. philoxeroides growth, but it failed to overwinter in ambient temperature. Together, these results suggest that warming will allow the natural enemy to expand its range, potentially benefiting biocontrol in regions that are currently too cold for the natural enemy. However, the invader may also expand its range further north in response to warming. In such cases where plants tolerate cold better than their natural enemies, the geographical gap between plant and herbivorous insect ranges may not disappear but will shift to higher latitudes, leading to a new zone of enemy release. Therefore, warming will not only affect plant invasions directly but also drive either enemy release or increase that will result in contrasting effects on invasive plants. The findings are also critical for future management of invasive species under climate change.     

Morphological variation in different populations of <Emphasis Type="Italic">Aceria anthocoptes</Emphasis> (Acari: Eriophyoidea) associated with the Canada thistle, <Emphasis Type="Italic">Cirsium arvense</Emphasis>, in Serbia

The russet mite, Aceria anthocoptes (Nal.), is the only eriophyid that has been recorded on Canada thistle, Cirsium arvense (L.) Scop. It has been noted in several European countries and recently in the USA. With its apparent host specificity and because of the damage it causes to its host plant, A. anthocoptes is being studied as a potential candidate for classical biological control. The aim of the present study was to examine quantitative morphological traits in four populations of A. anthocoptes living on two infraspecific host plant taxa (C. arvense var. arvense and C. arvense var. vestitum) in two geographically separate areas of Serbia in order to test the hypothesis of absence of the possible host plant impact on mite morphology. MANOVA analysis revealed significant differences between populations from different localities in Serbia. Populations of A. anthocoptes inhabiting two thistle varieties in the vicinity of Belgrade differed significantly from mites inhabiting the same two host varieties in the vicinity of the town of Ivanjica. Canonical discriminant analysis showed that the trait which best discriminates the populations of A. anthocoptes is the number of dorsal annuli. It was not possible to ascribe morphological differences to the impact of the host plant.     

不同环境变量下银胶菊叶甲的生命表(英文)

在室内研究了不同环境变量(如食物、温度、光周期和不同光波长)下银胶菊杂草Parthenium hysterophorus L.的食叶昆虫和生物防治因子银胶菊叶甲Zygogramma bicolorata Pallister 的生命表。不同发育阶段的该种叶甲取食银胶菊不同部位时,取食花的甲虫死亡率指标Kappa值最低,其次是取食叶片和茎;而取食花时甲虫世代存活率最高。温度显著影响主要发育阶段该种叶甲的死亡率和存活率。27℃下饲养的未成熟期甲虫的Kappa值最低,其次是30, 25, 20 和35℃。世代存活和存活率表现相同的趋势。不同光周期显著影响死亡率,在14L∶10D (长日照)下世代存活最好,其次是 12L∶12D (昼夜相等), 10L∶14D (短日照), 24L∶0D (连续光照) 和0L∶24D (连续黑暗)。甲虫对不同波长光的反应上,在白光（广谱）下Kappa值最低,世代存活率最高,其次是黄光（λ≈570 nm)、蓝光 (λ≈475 nm) 和红光(λ≈650 nm)。卵的死亡率最高。不同发育阶段的甲虫在27℃长日照白光下用银胶菊花饲养最佳。死亡率趋势具有严格和显著的阶段特异性,表现出内在的存活效应,与研究的因素无关。