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.     

花曲柳窄吉丁的寄主植物范围、危害和防治对策

花曲柳窄吉丁(emerald ash borer, EAB) Agrilus planipennis Fairmaire是木犀科（Oleaceae）梣属Fraxinus植物树木的毁灭性蛀干害虫,目前发现其主要危害欧梣亚属Subgen. Fraxinus欧梣组(Section Fraxinus)和象蜡树组（Section Meliodes）的一些树木种类,如美国白蜡Fraxinus americana、洋白蜡F. pennsylvanica var subintegerrima、绒毛白蜡F. velutina、黑白蜡F. nigra和水曲柳F. mandshurica等。与早期文献上的记载不同,很少发现花曲柳窄吉丁对苦枥木亚属Subgen. Ornus白蜡树组(Section Ornaster)的白蜡树F. chinensis Roxb和花曲柳F. rhynchophylla造成危害。不同寄主白蜡树对花曲柳窄吉丁的敏感性不同,洋白蜡、绒毛白蜡等北美白蜡树种受其危害重于亚洲的水曲柳。这可能是由于长期的适应进化,分布在亚洲地区的梣属寄主树木发展了对花曲柳窄吉丁的抗虫性。中国部分地区发生的花曲柳窄吉丁灾害应该归属于引进的北美白蜡树种对本地害虫敏感所致。在天津、黑龙江、吉林、辽宁、河北、山东、四川、台湾、内蒙古和新疆等省市和自治区,都曾发现花曲柳窄吉丁的危害或分布。白蜡属树木广泛分布于我国除了青藏地区和海南省之外的大部分地区,其中大面积引种北美白蜡树种的北方和西北地区,以及近年来种植发展水曲柳的东北地区都是花曲柳窄吉丁这种毁灭性蛀干害虫的潜在发生地。花曲柳窄吉丁在我国部分地区为检疫性林业害虫,做好检疫是杜绝其扩散蔓延的首要方法。我国局部地区出现花曲柳窄吉丁灾害说明,北美白蜡树种在我国的适应性还需要进行更进一步科学的全面的风险评估,在我国生态建设中应重视利用我国本地的白蜡树种。在花曲柳窄吉丁灾害的治理中,可采用营造混交林以降低蔓延的速度和发挥自然控制能力,加强肥水管理以增强树木的抗虫能力,清除被害木消灭虫源,以及幼虫期树干注射、成虫期喷洒化学农药等防治措施。     

花曲柳窄吉丁的空间分布

2004年4月~9月,在天津市大港区官港森林公园对花曲柳窄吉丁Agrilus planipennis的空间分布和垂直分布进行了调查研究,应用扩散系数(C)等6种指数法分析测定,确定了该虫在林地中的空间分布呈聚集分布,分布的基本成分是个体群。扩散蔓延规律为聚集型扩散。花曲柳窄吉丁幼虫在树干的垂直分布主要区间在50~150 cm之间。对幼虫垂直分布与树干高度的关系进行分析,其符合立方曲线,曲线方程为:Y=-5.142 9+17.943 7X+21.662 3X2-14.545X3;相关系数R2=0.915,P=0.013。     

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.     

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.

     

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.

     

白蜡吉丁柄腹茧蜂的羽化和产卵与寄主之间的关系

白蜡窄吉丁是危害白蜡属树木的一种重要蛀干害虫,因其幼虫期高度的隐蔽性生活而极难检测和防治.白蜡吉丁柄腹茧蜂是新发现外寄生于白蜡窄吉丁幼虫的优势天敌种,对寄主害虫的控制作用较强,具有良好的生物防治利用前景.研究调查了白蜡吉丁柄腹茧蜂及其寄主白蜡窄吉丁越冬代成虫羽化的时序差异、不同时期林间的寄生率、寄主密度与寄生率的关系以及寄主幼虫大小与茧蜂产卵量的关系.结果表明,越冬后白蜡吉丁柄腹茧蜂的羽化时间比其寄主害虫白蜡窄吉丁的羽化要晚1个多月.2003年白蜡吉丁柄腹茧蜂越冬代成虫的羽化从6月中旬持续到8月中旬,羽化高峰期在7月份;寄主白蜡窄吉丁的羽化期在5月中下旬.2004年白蜡吉丁柄腹茧蜂越冬代成虫的羽化从5月下旬持续到7月下旬,羽化高峰期在6月下旬至7月上旬;寄主羽化期在4月中旬至5月中旬（室内）.越冬后天敌的羽化时间刚好适应最早寄主发育到可供寄生的龄期,这是二者长期协同进化的结果,也表明白蜡吉丁柄腹茧蜂很可能是一种专性寄生蜂.林间白蜡吉丁柄腹茧蜂对白蜡窄吉丁幼虫的自然寄生率随着时间的推移总体上是逐步上升的.自然条件下茧蜂决定是否产卵与寄主幼虫的大小即龄期有关,它仅在前胸宽和体宽1.5 mm、体长12 mm以上,即3～4龄的寄主幼虫体表产卵,但只要接受寄主并产卵,不同龄期寄主幼虫上的产卵量无显著的差异.     

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.     

Core RNAi machinery and gene knockdown in the emerald ash borer (Agrilus planipennis)

The RNA interference (RNAi) technology has been widely used in insect functional genomics research and provides an alternative approach for insect pest management. To understand whether the emerald ash borer (Agrilus planipennis), an invasive and destructive coleopteran insect pest of ash tree (Fraxinus spp.), possesses a strong RNAi machinery that is capable of degrading target mRNA as a response to exogenous double-stranded RNA (dsRNA) induction, we identified three RNAi pathway core component genes, Dicer-2, Argonaute-2 and R2D2, from the A. planipennis genome sequence. Characterization of these core components revealed that they contain conserved domains essential for the proteins to function in the RNAi pathway. Phylogenetic analyses showed that they are closely related to homologs derived from other coleopteran species. We also delivered the dsRNA fragment of AplaScrB-2, a β-fructofuranosidase-encoding gene horizontally acquired by A. planipennis as we reported previously, into A. planipennis adults through microinjection. Quantitative real-time PCR analysis on the dsRNA-treated beetles demonstrated a significantly decreased gene expression level of AplaScrB-2 appearing on day 2 and lasting until at least day 6. This study is the first record of RNAi applied in A. planipennis.     

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。     

Modelling local and long-distance dispersal of invasive emerald ash borer Agrilus planipennis (Coleoptera) in North America

Limiting the damage by non-indigenous species requires rapid determination of current and potential distributions and vectors of dispersal, and development of appropriate management measures. The emerald ash borer ( Agrilus planipennis ), a wood-boring beetle native to South-East Asia, was first reported in the Great Lakes region during summer 2002. The beetle poses an enormous threat to native ash ( Fraxinus ) species of North America, as untreated trees in infested areas of Ontario, Michigan and Ohio suffer high mortality. We demonstrate that the borer has spread in North America through a combination of diffusive range extension, associated with local flights, and by long-distance 'jump' dispersal associated with human movement of infested sapling or contaminated firewood. Probability of infestation was inversely related to distance from borer epicentres but positively related to the size of human population centres. At least 9 of 39 populations that were first reported in Michigan during 2004 cannot be accounted for by local diffusion, raising the possibility that other unidentified mechanisms may be contributing to the dispersal of the beetle. In the absence of quarantine, by 2005 all of Michigan's lower peninsula was contained within the boundaries of potential diffusive range expansion. Infested ash saplings also were introduced from Michigan to Maryland during 2003, and subsequently transplanted to five sites in Maryland and Virginia. Quarantine and eradication measures have had mixed results: in the south-central USA, the species appears on the brink of eradication, whereas its distribution has continued to spread during 2005 in the Great Lakes region despite extensive containment and quarantine measures. Quarantine success in the Great Lakes region is encumbered by multiple dispersal vectors, larger borer population sizes and by the more extensive geographical distribution that was achieved prior to implementation of control measures.     

中国白蜡窄吉丁研究进展

白蜡窄吉丁Agrilus planipennis Fairmaire是近年来发生和影响比较严重的国际性检疫害虫,主要危害白蜡属(Fraxinus spp.)树木。它以幼虫在树木的韧皮部、形成层和木质部浅层蛀食,从而切断树木的营养输导组织,导致树木衰弱死亡。为了深入了解该害虫,探索合理的治理策略,本文对国内外有关白蜡窄吉丁的生物学特性,包括生活习性、生活史、分布与危害、寄主范围等方面以及防治技术的研究进展进行了系统总结,并讨论了白蜡窄吉丁今后的研究方向和研究趋势。     

Progress in research on the emerald ash borer Agrilus planipennis in China

白蜡窄吉丁Agrilus planipennis Fairmaire是近年来发生和影响比较严重的国际性检疫害虫,主要危害白蜡属(Fraxinus sPP.)树木.它以幼虫在树木的韧皮部、形成层和木质部浅层蛀食,从而切断树木的营养输导组织,导致树木衰弱死亡.为了深入了解该害虫,探索合理的治理策略,本文对国内外有关白蜡窄吉丁的生物学特性,包括生活习性、生活史、分布与危害、寄主范围等方面以及防治技术的研究进展进行了系统总结,并讨论了白蜡窄吉丁今后的研究方向和研究趋势.     

High temperature tolerance and thermal plasticity in emerald ash borer Agrilus planipennis

• 1 The emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae) (EAB), an invasive wood‐boring beetle, has recently caused significant losses of native ash (Fraxinus spp.) trees in North America. Movement of wood products has facilitated EAB spread, and heat sanitation of wooden materials according to International Standards for Phytosanitary Measures No. 15 (ISPM 15) is used to prevent this.
• 2 In the present study, we assessed the thermal conditions experienced during a typical heat‐treatment at a facility using protocols for pallet wood treatment under policy PI‐07, as implemented in Canada. The basal high temperature tolerance of EAB larvae and pupae was determined, and the observed heating rates were used to investigate whether the heat shock response and expression of heat shock proteins occurred in fourth‐instar larvae.
• 3 The temperature regime during heat treatment greatly exceeded the ISPM 15 requirements of 56 °C for 30 min. Emerald ash borer larvae were highly tolerant of elevated temperatures, with some instars surviving exposure to 53 °C without any heat pre‐treatments. High temperature survival was increased by either slow warming or pre‐exposure to elevated temperatures and a recovery regime that was accompanied by up‐regulated hsp70 expression under some of these conditions.
• 4 Because EAB is highly heat tolerant and exhibits a fully functional heat shock response, we conclude that greater survival than measured in vitro is possible under industry treatment conditions (with the larvae still embedded in the wood). We propose that the phenotypic plasticity of EAB may lead to high temperature tolerance very close to conditions experienced in an ISPM 15 standard treatment.

     

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。     

The overwintering physiology of the emerald ash borer, Agrilus planipennis fairmaire (coleoptera: buprestidae)

Ability to survive cold is an important factor in determining northern range limits of insects. The emerald ash borer (Agrilus planipennis) is an invasive beetle introduced from Asia that is causing extensive damage to ash trees in North America, but little is known about its cold tolerance. Herein, the cold tolerance strategy and mechanisms involved in the cold tolerance of the emerald ash borer were investigated, and seasonal changes in these mechanisms monitored. The majority of emerald ash borers survive winter as freeze-intolerant prepupae. In winter, A. planipennis prepupae have low supercooling points (∼−30 °C), which they achieve by accumulating high concentrations of glycerol (∼4 M) in their body fluids and by the synthesis of antifreeze agents. Cuticular waxes reduce inoculation from external ice. This is the first comprehensive study of seasonal changes in cold tolerance in a buprestid beetle.