Differences in Phyllotreta cruciferae and Phyllotreta striolata (Coleoptera: Chrysomelidae) responses to neonicotinoid seed treatments

Insecticidal seed treatments are used commonly throughout the Northern Great Plains of North America to systemically protect seedlings of canola (Brassica napus L. and Brassica rapa L.) from attack by the flea beetles Phyllotreta cruciferae (Goeze) and Phyllotreta striolata (F.) (Coleoptera: Chrysomelidae). Here, we investigated differential responses by the two flea beetle species to the neonicotinoid seed treatments thiamethoxam (Helix and Helix XTra) and clothianidin (Prosper 400) in greenhouse experiments. P. cruciferae experienced higher mortality and fed less when exposed to these compounds than did P. striolata. Beetles of the overwintered and the summer generations responded differently when feeding on seedlings that developed with insecticidal seed treatments, with mortality higher for P. cruciferae in May than in August. When the two flea beetle species were held together at equal densities and allowed to feed on seedlings affected by the seed treatments, mortality of P. cruciferae significantly exceeded that of P. striolata. Differences in efficacies of these compounds for these beetles have ramifications for management strategies in regions where these insects occur sympatrically. Competitive release of P. striolata was previously reported to occur when P. cruciferae was excluded from brassicaceous crops; consequently, the consistent use of these seed treatments over millions of hectares of canola cropland may be a factor that contributes to a shift in prevalence of flea beetle pest species from P. cruciferae toward P. striolata.     

Phyllotreta cruciferae and Phyllotreta striolata responses to insecticidal seed treatments with different modes of action

The strategy used most commonly in western North America to protect seedlings of canola (Brassica rapa L. and Brassica napus L.) from attack by adults of the flea beetles Phyllotreta cruciferae (Goeze) and Phyllotreta striolata (Fabricius) (Coleoptera: Chrysomelidae) involves planting seed coated with insecticide for systemic activity. Previous research determined that the two beetle species responded differently to the most commonly used neonicotinoid seed dressings. However, other insecticides that exploit different modes of action have commercial potential for managing infestations of these pests, but no information exists on their efficacies for these flea beetle species. Studies were conducted to compare effects of the neonicotinoid compounds, thiamethoxam and imidacloprid, to spinosyn and fipronil as systemic seed treatments for reducing feeding damage to canola seedlings and increasing mortality of P. cruciferae and P. striolata. Phyllotreta cruciferae experienced greater mortality and caused less feeding damage than P. striolata to seedlings treated with the neonicotinoid compounds. Mortality increased and feeding damage decreased significantly when beetles fed upon seedlings treated with fipronil, indicating its potential usefulness for control of these pests. However, spinosyn seed treatment was relatively ineffective against either beetle species. Higher rates of P. striolata mortality with fipronil than thiamethoxam suggest that fipronil may provide improved flea beetle control over hundreds of thousands of hectares in western North America where flea beetle populations are dominated by P. striolata, and control with thiamethoxam has been suboptimal.     

TWO HERBIVORES AND CONSTRAINTS ON SELECTION FOR RESISTANCE IN BRASSICA RAPA

Although most plants experience herbivory by several insect species, there has been little empirical work directed toward understanding plant responses to these simultaneous selection pressures. In an experiment in which herbivory by flea beetles (Phyllotreta cruciferae) and diamondback moths (Plutella xylostella) was manipulated in a factorial design, I found that selection for resistance to these herbivores is not independent in Brassica rapa. Specifically, the effect of flea beetle damage on B. rapa fitness depends on the amount of diamondback moth damage a plant experiences: damage by these herbivores has a nonadditive effect on plant fitness. When diamondbacks are abundant, plants that sustain high levels of damage by flea beetles are favored by natural selection, but when diamondbacks are rare, a low level of damage by flea beetles is favored. However, resistance to the later-feeding diamondback moth is not affected by the presence or absence of damage by early-feeding flea beetles. Thus, there are no plant-mediated ecological interactions between these herbivores that affect the outcome of selection for resistance. Because these herbivores do not independently affect plant fitness, neither is likely to develop a pairwise coevolutionary relationship with its host. Instead, coevolution is diffuse.     

Effects of seeding date and canola species on seedling damage by flea beetles in three ecoregions

Flea beetles ( Phyllotreta striolata (Fabricius), Phyllotreta cruciferae (Goeze) and Psylliodes punctulata Melsheimer) (Coleoptera: Chrysomelidae) are chronic threats to canola seedling establishment throughout the Northern Great Plains of North America. We conducted field studies in AB, Canada, from 2001 to 2004 at Lethbridge, Lacombe and Beaverlodge in the southern, central and northern regions of the province, respectively, to assess the impacts of seeding date (fall, April, May), and canola species ( Brassica rapa L., Brassica napus L.) on flea beetle damage to canola. Flea beetle damage was generally low at Lacombe but reached economic threshold levels at other sites. Although our results were not consistent in all years, seeding date had opposite effects on flea beetle damage to canola seedlings at the southern and northern sites. In the south, both canola species planted in April escaped flea beetle damage unlike the May-seeded plots, which required insecticide spraying in 2003. The numbers of flea beetles on sticky cards sampled during the seedling stage of canola were also lower in early-planted plots than in those planted in May. In the north, flea beetle damage was lower in the May-planted plots compared with those planted earlier. To maximize yields, canola growers need to plant as early as possible in all agro-ecoregions; our study showed that crops in the central and northern regions will be at greater risk of flea beetle damage than in southern regions.     

Temporal and host-related variation in frequencies of genes that enable Phyllotreta nemorum to utilize a novel host plant, Barbarea vulgaris

The flea beetle, Phyllotreta nemorum L. (Coleoptera: Chrysomelidae), is an intermediate specialist feeding on a small number of plants within the family Brassicaceae. The most commonly used host plant is Sinapis arvensis L., whereas the species is found more rarely on Cardaria draba (L.) Desv., Barbarea vulgaris R.Br., and cultivated radish (Raphanus sativus L.). The interaction between flea beetles and Barbarea vulgaris ssp. arcuata (Opiz.) Simkovics seems to offer a good opportunity for experimental studies of coevolution. The plant is polymorphic, as it contains one type (the P‐type) that is susceptible to all flea beetle genotypes, and another type (the G‐type) that is resistant to some genotypes. At the same time, the flea beetle is also polymorphic, as some genotypes can utilize the G‐type whereas others cannot. The ability to utilize the G‐type of B. vulgaris ssp. arcuata is controlled by major dominant genes (R‐genes). The present investigation measured the frequencies of flea beetles with R‐genes in populations living on different host plants in 2 years (1999 and 2003). Frequencies of beetles with R‐genes were high in populations living on the G‐type of B. vulgaris ssp. arcuata in both years. Frequencies of beetles with R‐genes were lower in populations living on other host plants, and declining frequencies were observed in five out of six populations living on S. arvensis. Selection in favour of R‐genes in populations living on B. vulgaris is the most likely mechanism to account for the observed differences in the relative abundance of R‐genes in flea beetle populations utilizing different host plants. A geographic mosaic with differential levels of interactions between flea beetles and their host plants was demonstrated.     

Feeding by flea beetles (Coleoptera: Chrysomelidae; Phyllotreta spp.) is decreased on canola (Brassica napus) seedlings with increased trichome density

Laboratory and field studies were undertaken to determine the effects of increased numbers of trichomes on seedling stems, petioles, and first true leaves of Brassica napus L., canola, on the feeding and behavior of the crucifer flea beetle Phyllotreta cruciferae (Goeze) (Coleoptera: Chrysomelidae). Seedlings of 'Westar' canola with genes inserted from Arabidopsis thaliana L. for increased trichome production, called Hairyl, were tested against Westar seedlings in no-choice and choice laboratory tests, and against parental plants and other cultivars grown from seed with and without insecticide in field trials at Saskatoon and Lethbridge, Canada. Analyses ofprefeeding and feeding behavior in no-choice tests of first true leaves found that flea beetles interacted with their host while off Hairyl leaves more so than beetles presented with leaves of Westar. Beetles required twice as much time to reach satiation when feeding on leaves with increased pubescence than on Westar leaves. In laboratory choice tests, flea beetles fed more on cotyledons and second true leaves of Westar than on comparable tissues of the transgenic line. In field trials, variations in feeding patterns were seen over time on cotyledons of the line with elevated trichomes. However, all four young true leaves of Hairyl seedlings were fed upon less than were the parental lines. Feeding on Hairyl plants frequently occurred at levels equal to or less than on cultivars grown from insecticide-treated seed. This study highlights the first host plant resistance trait developed in canola, dense pubescence, with a strong potential to deter feeding by crucifer flea beetles.     

Explaining differences in flea beetle Phyllotreta cruciferae goeze densities in simple and mixed broccoli cropping systems as a function of individual behavior

Diversification of habitat has proved to be an efficient way to reduce insect pest levels in agroecosystems. Some general theories used to explain this fact, such as the natural enemies and the resource concentration hypotheses, do not always clearly apply because, in many cases, pest individuals and population response seem controlled by more specific insect-plant interactions. In non replicated plots, we found substantially lower flea beetle densities in mixed broccoli-Vicia cropping systems compared to broccoli monoculture. These results were consistent with those from controlled experiments reported in the literature. To investigate if beetle behavior was related to such population reduction, the movement behavior of marked individuals of Phyllotreta cruciferae Goeze released in plots composed solely of broccoli plants and of broccoli mixed with Vicia faba or Vicia sativa plants, was followed and analyzed. The mean tenure time of beetles was longer in simple than in mixed cultures. Also, more beetles tended to fly out and leave mixed cultures compared to monoculture. This resulted in faster reduction of artificially introduced flea beetle populations in the mixed systems.Flea beetles landing on cover crop plants spent considerable time entangled in Vicia sativa branches or attempting to reach the upper part of the tall Vicia faba plants from which they could fly away. It is possible that the beetles characteristic movement on these two species of cover crops increased their risk of predation and the time and energy expended before they reached suitable host plants. Nevertheless, it seems that the detected flea beetle emigration rates were more than sufficient to account for the population trends observed.     

马缨丹提取物对黄曲条跳甲成虫的生物活性

测定了马缨丹(Lantana camaraL.)地上部位(茎叶花)的不同溶剂提取物及萃取物对黄曲条跳甲Phyllotreta striolata(Fabricius)成虫的取食忌避、拒食、胃毒和触杀作用。结果表明:处理后48 h,以马缨丹乙醇回流提取物效果最好,其对黄曲条跳甲成虫的取食忌避率达94.94%;非选择性拒食率为85.48%;除正丁醇萃取物对黄曲条跳甲成虫的胃毒作用达到56.67%外,其它处理对黄曲条跳甲成虫的胃毒作用不明显;石油醚萃取物、石油醚冷浸物、乙醇回流物对黄曲条跳甲成虫的触杀作用效果最好,分别为56.67,53.33和53.33%。据实验结果可知马缨丹不同溶剂提取物及萃取物对黄曲条跳甲成虫的生物活性主要表现在对成虫较强的忌避和拒食作用。     

饲养黄曲条菜跳甲实验种群的一种新方法

介绍一种简便可行的饲养黄曲条菜跳甲Phyllotretastriolata(F.)的新方法———整苗大瓶罩养法。用5种植物根部饲养越冬代,黄曲条菜跳甲均可成功地完成其生活周期,未成熟期最高存活率为63.3%,成虫平均寿命最长为219.2d,平均产卵量最大为192.4粒雌。     

Heterogeneity of plant phenotypes caused by herbivore-specific induced responses influences the spatial distribution of herbivores

Abstract.  1. Herbivory can induce resistance in a plant and the induced phenotype may be disfavoured by subsequent herbivores. Yet, as the distance between plants in a population increases, limited mobility may make a herbivore more likely to feed and oviposit on host plants in its immediate surroundings.
2. The present study tested whether a herbivore's preference and distribution across plants with different induced phenotypes was influenced by the spatial distribution of plants. A fragmented population of Solanum dulcamara plants was created. This consisted of discrete, spatially separated patches with different histories of damage, either herbivory from adult flea beetles ( Psylliodes affinis ), tortoise beetles ( Plagiometriona clavata ), or mechanical damage. Each patch was separated by 7 m and consisted of 12 plants that were spaced 30 cm apart. Then a fixed number of adult tortoise beetles were introduced to each patch, and movement and oviposition within and between spatially separate homogeneous patches (receiving one type of damage) were compared with movement and oviposition within heterogeneous patches (containing all three types of damage) over the growing season.
3. Flea beetle and tortoise beetle herbivory consistently induced different phytochemical responses in S. dulcamara (polyphenol oxidase and peroxidase), and adult tortoise beetles avoided oviposition on the flea beetle induced plants within heterogeneous patches. However, between homogeneous patches, plant phenotype did not influence oviposition. Colonisation by naturally occurring flea beetle adults followed a similar pattern.
4. These results suggest that the heterogeneity of plant phenotypes can influence herbivore choice and distribution at small but not large spatial scales.     

黄曲条跳甲人工饲养技术改进

通过室内人工饲养建立实验种群,是开展黄曲条跳甲Phyllotreta striolata(Fabricius)各种研究的基础,解决其幼虫阶段的人工饲养技术是建立其实验种群的关键。介绍经改进后的黄曲条跳甲室内人工饲养技术,该方法简单易行,可以方便地获取试虫的不同虫态;采用"小菜苗法"和"萝卜薄片堆叠法"室内饲养黄曲条跳甲幼虫,最高存活率可达90%以上。     

Specificity of a Y-linked gene in the flea beetle Phyllotreta nemorum for defences in Barbarea vulgaris

A Y-linked gene (R-gene) in the flea beetle Phyllotreta nemorum L. (Coleoptera: Chrysomelidae: Alticinae) confer the ability of larvae to survive on types of the plant Barbarea vulgaris R.Br. (Brassicaceae) which are immune to attack by susceptible conspecifics. Two near-isogenic flea beetle lines were developed. The YE-line contained the Y-linked R-gene, and male larvae from this line survived on B. vulgaris. The ST-line did not contain the gene and did not survive on the plant. The YE-line had been developed through 8–9 generations of backcrosses (YE-males with ST-females) and the two lines were considered to be isogenic except for genes located on the Y-chromosome.A single copy of the Y-linked gene is sufficient to transfer a susceptible genotype (ST) into a resistant genotype (YE) which is able to utilize a plant that is immune to attack by specimens without R-genes. The Y-linked gene had no effects on survival on other plant species tested. The gene did not have any effect on developmental times and weights of adult beetles reared on other plants than B. vulgaris. Developmental times of larvae with the Y-linked gene were longer on B. vulgaris than on normal host plants, R. sativus and S. arvensis, but the adults obtained the same size on these plant species. No trade-offs of the Y-linked gene were discovered. The results suggest that the occurrence of the Y-linked gene is a derived trait which has enabled the flea beetle to expand its host plant range. The evolution of a host shift to B. vulgaris seems not to be favoured by the presence of this single gene.     

Prefeeding Behavior of the Crucifer Flea Beetle, Phyllotreta cruciferae, on Host and Nonhost Crucifers

The prefeeding behaviours of adult crucifer flea beetles, Phyllotreta cruciferae (Coleoptera: Chrysomelidae: Alticinae), were determined on seedlings of the host plant, Brassica napus, and compared to behaviors on seedlings of the nonhost crucifers, Crambe abyssinica, Sinapis alba, and Camelina sativa. Three stages of prefeeding behaviour, i.e., acclimation, stimulation, and initial feeding, were distinguished through observation of filmed beetles. Both antennal and tarsal chemoreceptors are important in determination of host plant quality by the crucifer flea beetle. The results of this study suggest that the sequence of prefeeding behaviors plays a crucial role in the onset of feeding. Differences in time spent on plant tissue and the frequency and duration of prefeeding behaviors provide insight into possible mechanisms of resistance to flea beetles in the non-Brassica crucifers. The nonpreferred hosts C. abyssinica and S. alba contain deterrent phytochemicals that partially inhibit feeding. These deterrent compounds appear to be volatile in nature in S. alba but nonvolatile in C. abyssinica. CFB resistance in the nonhost C. sativa may result from either the presence of repellent or the absence of stimulatory volatile phytochemicals.     

Enhanced β-ionone emission in Arabidopsis over-expressing AtCCD1 reduces feeding damage in vivo by the crucifer flea beetle

Plant carotenoid derived β-ionone has been shown to have diverse biological effects on some insect herbivores and herbivore parasitoids. In this study, Arabidopsis transgenic plants over-expressing a carotenoid cleavage dioxygenase1 gene (AtCCD1) were generated to test whether β-ionone emissions could be enhanced and used to control feeding by the crucifer flea beetle (Phyllotreta cruciferae Goeze). The transgenic plants exhibited a morphological phenotype indistinguishable from the wild type (WT) control over their complete life cycle. Gas chromatography and mass spectrometry analyses of headspace volatiles collected from 6-wk-old intact flowering plants revealed substantially enhanced β-ionone emission from transgenic plants compared with WT, but no β-ionone enhancement occurred at a young vegetative stage (4-wk-old seedlings). Bioassays in an enclosed environment showed that AtCCD1 over-expression resulted in ?50% less leaf area damage by flea beetles on transgenic plants compared with WT plants. The mean number of damaged transgenic leaves per plant also was significantly lower in transgenic plants (P<0.05). Our results indicate that AtCCD1 over-expression and induced β-ionone emission might find application in the control of pests for Brassica crops grown in greenhouse operations. Potentially, β-ionone also could be used on crops grown in open-air ecosystems if this allomone is released in sufficient quantities to discourage herbivore foragers.     

The evolution of plant response to herbivory: simultaneously considering resistance and tolerance in Brassica rapa

Although the evolution of plant response to herbivory can involve either resistance (a decrease in susceptibility to herbivore damage) or tolerance (a decrease in the per unit effect of herbivory on plant fitness), until recently few studies have explicitly incorporated both of these characters. Moreover, theory suggests these characters do not evolve independently, and also that the pattern of natural selection acting on resistance and tolerance depends on their costs and benefits. In a genotypic selection analysis on an experimental population of Brassica rapa (Brassicaceae) I found a complex set of correlational selection gradients acting on resistance and tolerance of damage by flea beetles (Phyllotreta cruciferae: Chrysomelidae) and weevils (Ceutorhynchus assimilis: Curculionidae), as well as directional and stabilizing selection on resistance to attack by weevils. Evolution of response to flea beetle attack is constrained by a strong allocation cost of tolerance, and this allocation cost may be caused by a complex correlation among weevil resistance, weevil tolerance, flea beetle resistance, and flea beetle tolerance. Thus, one important conclusion of this study is that ecological costs may involve complex correlations among multiple characters, and for this reason these costs may not be detectable by simple pairwise correlations between characters. The evolution of response to weevil attack is probably constrained by a series of correlations between weevil resistance, weevil tolerance, and fitness in the absence of weevil damage, and possibly by a cost of tolerance of weevil damage. However, the nature of these constraints is complicated by apparent overcompensation for weevil damage. Because damage by both flea beetles and weevils had non-linear effects on plant fitness, standard measures of tolerance were not appropriate. Thus, a second important contribution of this study is the use of the area under the curve defined by the regression of fitness on damage and damage-squared as a measure of tolerance. This revised version was published online in July 2006 with corrections to the Cover Date.     

几种十字花科蔬菜害虫生态位的研究

以Levins生态位宽度指数和Pianka生态位重叠指数估计发生在同一营养阶层的 4种主要十字花科植物害虫营养生态位、时间生态位和空间生态位的生态位宽度和重叠度。结果表明小菜蛾、黄曲条跳甲、菜粉蝶和小猿叶甲 4种昆虫的生态位并不完全重叠。     

印楝素乳油对黄曲条跳甲种群控制作用评价

在生命表方法分析的基础上。建立了害虫累积为害量指标,用以描述和比较某一时间段内害虫动态持续为害力大小．应用该指标分析印楝素乳油对黄曲条跳甲(Phyllotreta striolata(F.))种群控制作用,结果表明,对成虫显著的忌避作用是其对种群控制的主要机理．印楝素0．50ml·L^-1对成虫的忌避率为83．75％,种群趋势指数I值降为0．3354,干扰作用控制指数为0．0863．采用田间试验获得的结果,避免了室内测定的空间限制,因而是评价异源植物次生化合物对害虫忌避作用的主要依据．该指标在以害虫控制为目标的动态研究中具有较好的应用前景．     

Impact of planting dates and insecticide strategies for managing crucifer flea beetles (Coleoptera: Chrysomelidae) in spring-planted canola

Integration of cultural practices, such as planting date with insecticide-based strategies, was investigated to determine best management strategy for flea beetles (Phyllotreta spp.) (Coleoptera: Chrysomelidae) in canola (Brassica napus L.). We studied the effect of two spring planting dates of B. napus and different insecticide-based management strategies on the feeding injury caused by fleabeetles in North Dakota during 2002-2003. Adult beetle peak emergence usually coincided with the emergence of the early planted canola, and this resulted in greater feeding injury in the early planted canola than later planted canola. Use of late-planted canola may have limited potential for cultural control of flea beetle, because late-planted canola is at risk for yield loss due to heat stress during flowering. Flea beetle injury ratings declined when 1) the high rate of insecticide seed treatment plus a foliar insecticide applied 21 d after planting was used, 2) the high rate of insecticide seed treatment only was used, or 3) two foliar insecticide sprays were applied. These insecticide strategies provided better protection than the low rates of insecticide seed treatments or a single foliar spray, especially in areas with moderate-to-high flea beetle populations. The foliar spray on top of the seed treatment controlled later-emerging flea beetles as the seed treatment residual was diminishing and the crop became vulnerable to feeding injury. The best insecticide strategy for management of flea beetle was the high rate of insecticide seed treatment plus a foliar insecticide applied at 21 d after planting, regardless of planting date.     

Species spectrum of flea beetles (Phyllotreta spp., Coleoptera, Chrysomelidae) attracted to allyl isothiocyanate-baited traps

In field tests in Hungary, Slovenia and Bulgaria, in allyl isothiocyanate-baited traps significantly more beetles of Phyllotreta cruciferae, Ph. vittula, Ph. undulata, Ph. nigripes, Ph. nodicornis, Ph. balcanica, Ph. atra, Ph. procera, Ph. ochripes, Ph. diademata and Psylliodes chrysocephalus (Coleoptera, Chrysomelidae, Halticinae) were captured than in unbaited control traps. With the exception of Ph. cruciferae, this is the first report on significant field attraction by allyl isothiocyanate for these species. The species spectrum captured included six important agricultural pests. At all sites a great portion of the catch (ranging from ca 30 to 98%) was Ph. cruciferae, irrespective of the plant culture. The second most abundant species present at most sites was Ph. vittula. The present results are very promising from the point of view of applicability of allyl isothiocyanate in Europe as a bait in cabbage flea beetle traps for detection and monitoring.