Attraction of Agrilus planipennis (Coleoptera: Buprestidae) to a volatile pheromone: effects of release rate, host volatile, and trap placement

Attraction of emerald ash borer, Agrilus planipennis Fairmaire, to a volatile pheromone was demonstrated in three field experiments using baited green sticky traps. A dose-response curve was generated for male A. planipennis to increasing release rates of (3Z)-dodecen-12-olide ((3Z)-lactone) in combination with the green leaf volatile, (3Z)-hexenol. Only the lowest release rate (<2.50 μg/d) of (3Z)-lactone significantly increased captures of male A. planipennis, as compared with traps baited with (3Z)-hexenol alone. Effect of trap height, (3Z)-lactone, and (3Z)-hexenol and their interactions on the trap capture of A. planipennis was determined in a factorial experiment. Number of males per trap was significantly and positively affected by (3Z)-lactone, (3Z)-hexenol, and trap height whereas number of females per trap was affected by trap height only; none of the interactions were significant. As predicted, the greatest mean catch of males was in traps baited with (3Z)-lactone and (3Z)-hexenol placed high in the canopy. Electroantennogram tests on the bark volatile, 7-epi-sesquithujene, demonstrated the ability of male and female A. planipennis antennae to detect and respond to this compound, particularly the (+)-7-epi-sesquithujene isomer. Results from an olfactometer bioassay and field testing did not demonstrate attraction of either males or females to (+)-7-epi-sesquithujene. These data increase our understanding of the pheromone ecology of the invasive emerald ash borer, provide further confirmation of the behavioral activity of the female-produced lactone pheromone, and should increase the ability to detect A. planipennis infestations where they are present.     

白蜡窄吉丁虫在中国的研究现状与分布调查

2002年在美国密执安州发现了一重大外来入侵害虫,白蜡窄吉丁Agrilus planipennis Fairmaire, 由于该虫新近在美国发现,危害白蜡树,故英文名称为Emerald Ash Borer (EAB)。初步分析显示,该虫至少在5年前就已传入美国,只是现在刚爆发被发现。目前该虫在美国密执安州和加拿大安大略省可致死各种大小的白蜡树。该虫在我国曾被定名为花曲柳窄吉丁Agrilus marcopoli Obenberger,为鞘翅目吉丁科。据资料报道分布于黑龙江、吉林、辽宁、山东、内蒙古和台湾等地;在国外分布于朝鲜,日本,蒙古,俄罗斯远东地区。危害木樨科属树木。两年一代或一年一代,因地而已。在东北三省主要危害水曲柳和花曲柳等树木,但不是主要害虫。天津市1993年在引种白蜡树上发现了白蜡窄吉丁,1998年严重爆发造成大面积白蜡树死亡。在亚洲的其他分布区,该虫还危害另外一些阔叶树,如多种蜡树、榆树等。鉴于该虫的广泛分布和寄主树种多及毁灭性危害的特点,美国一些专家认为其潜在危害不亚于1996年在美国发现的光肩星天牛。因此,美国已全面开展对该虫的研究,包括资料收集,风险性评估,研究各种控制措施等。 本文是中美对此虫合作研究的一部分,旨在对该虫在我国的分布、 危害、 研究基础作一调查与回顾, 为下一步深入研究提供信息与参考资料。白蜡窄吉丁入侵北美再一次显示随着贸易的全球化发展, 外来入侵种问题也越来越严重。国际合作共同开展研究是控制入侵种的重要举措之一。     

Susceptibility of Agrilus planipennis (Coleoptera: Buprestidae) to Beauveria bassiana and Metarhizium anisopliae

The susceptibility of Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) to selected strains of the entomopathogenic fungi Beauveria bassiana (Balsamo) Vuillemin and Metarhizium anisopliae (Metschnikoff) Sorokin was evaluated through bioassays with direct immersion or foliar exposure under laboratory conditions. Results showed that A. planipennis adults were susceptible to B. bassiana and M. anisoplae. Significant time-mortality response was found for each isolates. Isolate B. bassiana GHA killed A. planipennis adults at a faster rate compared with other isolates tested, with the lowest average time-to-death values. The LC50 values estimated under direct immersion method ranged from 1.7 x 10(5) to 1.9 x 10(7), 3.5 x 10(4) to 5.3 x 10(5), and 4.1 x 10(3) to 2.9 x 10(5) conidia/ml for B. basissiana and from 3.2 x 10(6) to 1.1 x 10(7), 4.5 x 10(3) to 4.5 x 10(5), and 1.4 x 10(2) to 1.2 x 10(5) conidia/ml for M. anisopliae at 4, 5, and 6 d after treatment, respectively. By days 5 and 6, B. bassiana GHA outperformed all other isolates tested except ARSEF 7234, followed by ARSEF 7152, 6393, and 7180. Significant concentration-mortality response was also observed for two B. bassiana GHA formulations, BotaniGard ES and Mycotrol O, and M. anisopliae F52 when insects were treated through foliar exposure. The LC50 values ranged from 114.5 to 309.6, 18.4 to 797.3, and 345.3 to 362.0 conidia/cm2 for BotaniGard, Mycotrol, and M. anisopliae F52, respectively. Based on the results of these bioassays, the efficacy of both B. bassiana GHA formulations and M. anisopliae F52 were similar against adult A. planipennis. The potential use of entomopathogenic fungi for management of A. planipennis in North America is discussed.     

The Role of Olfactory Cues in Short-Range Mate Finding by the Emerald Ash Borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae)

We investigated the relative importance of olfaction versus vision in the mate-finding behavior of Agrilus planipennis. When coupled in male–female, male–male and female–female pairs, attempts to mate occurred only in the male–female pairs, suggesting that beetles can identify the opposite sex before attempting to mate. In a set of sensory deprivation experiments with male–female pairs, we evaluated whether males could find females when deprived of their sense of olfaction, vision or both. Males whose antennae were blocked with model paint took significantly longer to find females and spent less time in copula compared to untreated males. Males whose eyes were similarly blocked did not differ in their mate finding capacity compared to untreated males. In a third experiment that compared both olfaction and vision, olfactorily impaired beetles never mated whereas the mate finding potential of visually impaired beetles did not differ from that of untreated beetles. Our results indicate that males can identify females before coming into physical contact with them, and that at short range (≤5 cm), volatile cues detected by olfaction are involved in mate finding by A. planipennis.     

Host range of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) in North America: results of multiple-choice field experiments

Emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae), an invasive phloem-feeding pest, was identified as the cause of widespread ash (Fraxinus) mortality in southeast Michigan and Windsor, Ontario, Canada, in 2002. A. planipennis reportedly colonizes other genera in its native range in Asia, including Ulmus L., Juglans L., and Pterocarya Kunth. Attacks on nonash species have not been observed in North America to date, but there is concern that other genera could be colonized. From 2003 to 2005, we assessed adult A. planipennis landing rates, oviposition, and larval development on North American ash species and congeners of its reported hosts in Asia in multiple-choice field studies conducted at several southeast Michigan sites. Nonash species evaluated included American elm (U. americana L.), hackberry (Celtis occidentalis L.), black walnut (J. nigra L.), shagbark hickory [Carya ovata (Mill.) K.Koch], and Japanese tree lilac (Syringa reticulata Bl.). In studies with freshly cut logs, adult beetles occasionally landed on nonash logs but generally laid fewer eggs than on ash logs. Larvae fed and developed normally on ash logs, which were often heavily infested. No larvae were able to survive, grow, or develop on any nonash logs, although failed first-instar galleries occurred on some walnut logs. High densities of larvae developed on live green ash and white ash nursery trees, but there was no evidence of larval survival or development on Japanese tree lilac and black walnut trees in the same plantation. We felled, debarked, and intensively examined >28 m2 of phloem area on nine American elm trees growing in contact with or adjacent to heavily infested ash trees. We found no sign of A. planipennis feeding on any elm.     

Influence of separate rearing on adult longevity and fecundity of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae)

白蜡窄吉丁Agrilus planipennis Fairmaire是近年来发生和危害严重的国际性检疫害虫.为了了解不同饲养条件和种内个体间相互感知对成虫生物学的影响,本研究采用群体饲养、单对饲养、无隔离单雌饲养、视觉隔离单雌饲养和嗅觉隔离单雌饲养等5种处理,在室内测定了白蜡窄吉丁成虫的寿命、取食量、产卵量和产卵历期等参数的变化.结果显示:隔离饲养降低了白蜡窄吉丁成虫的寿命和繁殖力,且不同处理下白蜡窄吉丁成虫的寿命和取食量均存在显著差异(P＜0.05),但产卵量的差异不明显(P＞0.05).其中无隔离单雌饲养时成虫的平均寿命最长,为32.40 d,群体饲养的寿命最短,为20.77 d,且单雌饲养的寿命均比单对饲养和群体饲养的长.单雌饲养的3种情况下,视觉隔离条件下的日均取食量最大,为156.16 mm2,嗅觉隔离的最小,为107.35 mm2;无隔离饲养条件下雌虫的产卵量最大,产卵历期也最长,分别为98.33 d和21.33 d.这些研究结果表明,白蜡窄吉丁种内个体间相互感知的阻断对成虫的取食、发育和繁殖活动存在明显影响,且嗅觉在其种群内的交流活动中作用最重要,其次是视觉.此外,个体之间的相互干扰或竞争对白蜡窄吉丁成虫的寿命和繁殖力也可能存在一定的影响.这为生产上采用隔离措施如营造混交林带防治该虫提供了理论基础.     

隔离饲养对白蜡窄吉丁成虫寿命和繁殖力的影响

白蜡窄吉丁Agrilus planipennis Fairmaire是近年来发生和危害严重的国际性检疫害虫。为了了解不同饲养条件和种内个体间相互感知对成虫生物学的影响, 本研究采用群体饲养、 单对饲养、 无隔离单雌饲养、 视觉隔离单雌饲养和嗅觉隔离单雌饲养等5种处理, 在室内测定了白蜡窄吉丁成虫的寿命、 取食量、 产卵量和产卵历期等参数的变化。结果显示： 隔离饲养降低了白蜡窄吉丁成虫的寿命和繁殖力, 且不同处理下白蜡窄吉丁成虫的寿命和取食量均存在显著差异（P<0.05）, 但产卵量的差异不明显（P>0.05）。其中无隔离单雌饲养时成虫的平均寿命最长, 为32.40 d, 群体饲养的寿命最短, 为20.77 d, 且单雌饲养的寿命均比单对饲养和群体饲养的长。单雌饲养的3种情况下, 视觉隔离条件下的日均取食量最大, 为156.16 mm², 嗅觉隔离的最小, 为107.35 mm²; 无隔离饲养条件下雌虫的产卵量最大, 产卵历期也最长, 分别为98.33 d和21.33 d。这些研究结果表明, 白蜡窄吉丁种内个体间相互感知的阻断对成虫的取食、 发育和繁殖活动存在明显影响, 且嗅觉在其种群内的交流活动中作用最重要, 其次是视觉。此外, 个体之间的相互干扰或竞争对白蜡窄吉丁成虫的寿命和繁殖力也可能存在一定的影响。这为生产上采用隔离措施如营造混交林带防治该虫提供了理论基础。     

Development of a host-based semiochemical lure for trapping emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae)

Bark volatiles from green ash Fraxinus pennsylvanica were tested for electrophysiological activity by Agrilus planipennis using gas chromatographic-electroantennographic detection (GC-EAD) and for behavioral activity using baited purple traps in Michigan. GC-EAD analysis of the headspace volatiles of bark tissue samples from 0- and 24-h-old fully girdled (stressed) ash trees showed that the latter had elevated sesquiterpene levels. Six of the elevated compounds consistently elicited antennal responses by both male and female A. planipennis. Five of the antennally active compounds were identified as alpha-cubebene, alpha-copaene, 7-epi-sesquithujene, trans-beta-caryophyllene, and alpha-humulene (alpha-caryophyllene). The sixth EAD-active compound remains unidentified. We monitored capture of adult A. planipennis on traps baited with several combinations of ash tree volatiles. Treatments included two natural oil distillates (Manuka and Phoebe oil) that were found to contain, respectively, high concentrations of four and five of the six antennally active ash bark volatiles. A four-component leaf lure developed by the USDA Forest Service and Canadian Forest Service was also tested. In three separate field studies, Manuka oil-baited traps caught significantly more adult beetles than unbaited traps. Lures designed to release 5, 50, and 500 mg of Manuka oil per day all caught more insects than unbaited traps. In a field test comparing and combining Phoebe oil with Manuka oil, Phoebe oil-baited traps caught significantly more beetles than either Manuka oil-baited traps or unbaited traps. We hypothesize that the improved attractancy of Phoebe oil to A. planipennis over Manuka oil is caused by the presence of the antennally active sesquiterpene, 7-epi-sesquithujene.     

基于特定时间调查数据组建的白蜡窄吉丁种群生命表

为了探明影响白蜡窄吉丁Agrilus planipennis Fairmaire自然种群变化的关键因子, 本研究于2010-2011年采用当年秋季和翌年春季两个特定时间调查取样的方法, 组建了天津、 北京和辽宁3个地区白蜡窄吉丁的自然种群生命表。结果显示, 这3个地区白蜡窄吉丁的自然种群数量在今后几年均将呈增长趋势, 但不同地区的增长速率存在较大差异。北京地区白蜡窄吉丁的种群数量增长速率最快（种群趋势指数I=12.9997）, 其次是天津地区（I=12.4388）, 这两个地区的寄主植物相同, 均为绒毛白蜡。增长最慢的是辽宁地区（I=3.6394）, 其寄主植物为水曲柳。另外, 通过3个地区白蜡窄吉丁生命表和排除控制指数可以看出, 天津地区白蜡窄吉丁最主要的致死因子是白蜡吉丁柄腹茧蜂Spathius agrili Yang（种群数量排除控制指数EIPC=1.5382）, 对种群数量变动的贡献最大; 北京地区的是啄木鸟（EIPC=1.4515）, 而辽宁地区白蜡窄吉丁最主要的致死因子是白蜡吉丁卵跳小蜂Oobius agrili Zhang et Huang的寄生作用（EIPC=1.4701）。这些研究结果表明, 寄主植物的差异是影响白蜡窄吉丁自然种群数量变动的主要因素, 同时寄生性天敌白蜡吉丁柄腹茧蜂对种群数量的影响作用也比较大。因此, 实际应用中可以通过培育抗性树种和人工繁育优势天敌等方法对该虫进行有效防治。     

Microbial control of Agrilus planipennis (Coleoptera: Buprestidae) with Beauveria bassiana strain GHA: field applications

The effects of Beauveria bassiana strain GHA, applied as BotaniGard ES, on newly colonised and well-established populations of emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae) were evaluated in the field using foliar and trunk sprays in Michigan in 2004–2005. Results from field trials at a newly colonised white ash site showed a 41% reduction in A. planipennis population in fungal-treated trees compared with that of untreated controls. In addition, fungal infection was also found in 20% of the larval population within 14 days of incubation under laboratory conditions. At a site with a well-established Agrilus planipennis population in green ash trees, larval density was reduced by 47% for trees treated with the fungus compared with that of the controls; 21% of larvae from the current generation were found infected after 14 days of laboratory incubation. Fungal-treated green ash trees also produced fewer adults emerging in the next generation, with a 63% reduction in adult density observed in treated trees compared to that of controls. As a result, fungal-treated trees sustained 42% less crown dieback than did controls. A. planipennis larval density was negatively correlated with trunk height above the ground, and positively correlated with log diameter. Results of laboratory leaf bioassays on A. planipennis adults showed that fungal conidia persisted well under field conditions, with mortality of 78–100% at 7 – days post – exposure for leaves collected between 2 and 264 h after application. Potential strategies for using B. bassiana strain GHA for managing A. planipennis are discussed.     

Dramatic expansion of the range of the invasive ash pest,buprestid beetle Agrilus planipennis Fairmaire, 1888 (Coleoptera,Buprestidae) in European Russia

The emerald ash borer (Agrilus planipennis Fairmaire, 1888) is a dangerous invasive pest of ashes. It was previously believed that in European Russia it occurs only in Moscow, Moscow Province, and in the eastern regions of Smolensk Province. An examination of ash trees in 12 cities of European Russia has revealed a much wider area of the emerald ash borer invasion. The pest was found in Konakovo (Tver Province), Tula, Kaluga, Orel, and Voronezh. We have found that A. planipennis damages not only Fraxinus pennsylvanica (an American species which is commonly planted in cities), but also the aborigine European ash Fraxinus excelsior. Ashes in the European forests and in the protective forest belts are badly endangered.     

Susceptibility of two hymenopteran parasitoids of Agrilus planipennis (Coleoptera: Buprestidae) to the entomopathogenic fungus Beauveria bassiana (Ascomycota: Hypocreales)

Emerald ash borer (EAB), Agrilus planipennis Fairmaire, native to Asia, is killing ash trees (Fraxinus spp.) across 15 states and southeastern Canada. Integrated pest management using biological control is the only viable long-term approach for controlling the spread of EAB outside of host resistance. Three hymenopteran parasitoids, Spathius agrili Yang, Tetrastichus planipennisi Yang, and Oobius agrili Zhang and Huang were discovered attacking EAB in China and were approved for release in the United States in 2007. The objective of this study was to assess susceptibility of the larval parasitoid species S. agrili and T. planipennisi, relative to that of EAB, to Beauveria bassiana, an entomopathogenic fungus that infects and kills EAB adults when sprayed on ash bark or foliage. Adult EAB and parasitoids were exposed to B. bassiana inoculated ash twigs for 2 h and then monitored daily for death and signs of infection for up to 10 days. All EAB adults exposed to B. bassiana were fatally infected while mean survival for control EAB was 77%. Average survival in the treatment groups for T. planipennisi and S. agrili were 99% and 83%, respectively, indicating these parasitoids are relatively unaffected by exposure to B. bassiana. This research elucidates interactions between a fungal pathogen and two parasitoids of EAB, and provides data necessary to developing a successful multi-stage integrated management approach to control of EAB.     

Optimizing early detection strategies: defining the effective attraction radius of attractants for emerald ash borer Agrilus planipennis Fairmaire

1. Adult emerald ash borers are attracted to green prism traps baited with the ash host volatile (3Z)-hexenol and the sex pheromone of emerald ash borer (3Z)-dodecen-12-olide [(3Z)-lactone]. Quantifying the heretofore unknown range of attraction of such traps would help optimize deployment strategies for early detection.
2. Examining trap captures of traps deployed in pairs at variable distances offers insight into the range of attraction. Recent work has shown the range of attraction can be estimated as half the intertrap distance at which trap catch begins to decrease, which should occur when proximate traps overlap their respective attractive ranges.
3. We estimated these traps' attractive range for emerald ash borer using 98 baited dark green prism traps deployed in pairs, one trap per tree, in an urban park in Saint Paul, Minnesota, USA in summer 2020.
4. We estimate attractive range by fitting a logistic model to trap catch data using Bayesian inferential methods and describe advantages thereof.
5. The attractive range of these baited traps was estimated to be between 16 and 73 m, with a median of 28 m. We recommend that dark green prism traps baited with these semiochemicals be placed 25–35 m apart near high-risk entry points.

     

柑桔爆皮虫卵巢发育动态

本研究利用室内饲养成虫,对柑桔爆皮虫的蛹及正常取食和交配、正常取食但未交配以及正常交配但未取食三种处理的雌成虫的卵巢发育进行了系统观察。结果显示：该虫具有1对卵巢,每侧有5根卵巢管。前两种处理的雌虫卵巢管均能正常发育,成虫寿命在30天左右; 没有取食的雌虫寿命只有7天左右,在其卵巢管的生长区和成熟区均无卵形成。根据卵巢的形状、卵的产生过程、卵巢萼内有无卵粒以及卵黄沉积情况等将卵巢发育程度分为6个级别,即发育初期（0级）、卵黄沉积前期(Ⅰ级)、卵黄沉积期(Ⅱ级)、成熟待产期(Ⅲ级)、产卵盛期(Ⅳ级)和产卵末期(Ⅴ级)。每头雌虫最高怀卵量在140粒左右,根据雌虫怀卵量变化趋势,推测正常取食和交配的柑桔爆皮虫雌虫在出孔后10天左右开始产卵,产卵历期可达22天左右。据此提出该虫卵巢管发育到成熟待产期(Ⅲ级)之前（即羽化出孔后10天之内）为出孔成虫的防治适期。     

Effects of grafting on host plant resistance in ash (Fraxinus spp.) to emerald ash borer (Agrilus planipennis Fairmaire)

1. The invasive emerald ash borer (EAB) (Agrilus planipennis Fairmaire Coleoptera: Buprestidae) is a phloem‐boring pest from Asia that has killed hundreds of millions of North American ash trees (Fraxinus spp.). Propagation of trees resistant to EAB must be explored as a tool to maintain the North American ash resource.
2. Interactions between the root and shoot of a tree may be critical to its ability to resist attack by insects and pathogens because many secondary metabolites critical to plant defence are synthesized in the roots.
3. In the present study, we determined the extent to which grafting susceptible green ash (Fraxinus pennsylvanica Marshall) and resistant Manchurian ash (Fraxinus mandshurica Rupr.) affects EAB resistance in ash trees. We measured adult EAB survivorship, herbivory, fecundity and morphological tree characteristics on grafted and nongrafted green and Manchurian ash.
4. Among grafted trees, beetles caged on grafted green ash lived four days longer and laid at least three times more eggs than beetles caged on trees with a Manchurian ash scion or rootstock. Herbivory was unaffected by graft combination.
5. Our findings suggest that grafted trees with Manchurian ash rootstocks or scions will be more resistant to EAB than nongrafted green ash trees.

     

Biology and damage traits of emerald ash borer planipennis Fairmaire）in China

Emerald ash borer （Agrilus planipennis Fairmaire） （Coleoptera： Buprestidae） is a major stem borer of ash （Fraxinus spp.）. It is univoltine in Tianjin, while it is semivoltine in Heilongjiang Province, and both univoltine and semivoltine in Changchun, Jilin Province, where the majority is univoltine. The longevity of emerald ash borer adults is 17.2 ± 4.6 days （n = 45）, eggs 9.0 5：1.1 days （n = 103）, univoltine larvae 308 days, semivoltine larvae 673 days, and pupae 61.2 ± 1.6 days （n = 45）. It takes about 100 days from the time larvae bore into the phloem to when they complete the pupal cell. In a 10-year-old velvet ash （Fraxinus velutina Tort.） plantation in Tianjin, emerald ash borer preferred to oviposit on the regions of boles from 50-150 cm above ground, accounting for 76.7% of the total girdling. Girdling on the south side of the tree boles accounted for 43.40% of the total girdling. The emerald ash borer population density is higher at the edge of the plantation compared with the center.     

柑桔爆皮虫羽化孔的空间分布

通过调查柑桔爆皮虫成虫羽化孔的分布,经空间分布型检验和聚集度指标的测定以及Taylor幂法则和m*- m（Iwao）回归分析,结果表明,柑桔爆皮虫的羽化孔在柑桔树树体内和桔园内均呈聚集分布: 在枯树内呈负二项分布,在半枯树内呈负二项分布或奈曼A型分布,在桔园内呈负二项分布,在树体内和桔园内分布的基本成分均为为个体群,个体间相互吸引。树势较差的柑桔树容易受柑桔爆皮虫危害,在树势强的树体内不能完成世代发育。因此,柑桔爆皮虫采用多次聚集攻击的策略攻击寄主。在桔园内聚集危害部分寄主,在树体内聚集危害寄主主枝的中下部,主枝的死亡导致整株树势下降,下一代虫源进而再次攻击寄主,在树势下降的树体内柑桔爆皮虫完成世代发育并羽化出孔,成虫继续为害健康的柑桔树。