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.     

白蜡窄吉丁幼虫及其天敌的种群空间生态位研究

【目的】为了明确白蜡窄吉丁Agrilus planipennis Fairmaire幼虫及其天敌在寄主树上的分布规律和相互关系。【方法】对天津和北京两个地区绒毛白蜡Fraxinus velutina和辽宁地区水曲柳F mandshurica上白蜡窄吉丁幼虫及其天敌的种群生态位进行研究。【结果】白蜡窄吉丁幼虫及其天敌的数量均随着寄主树干高度的增加而逐渐减少,但在不同寄主树种上的分布范围不同。绒毛白蜡上的分布范围较广,而在水曲柳主干上的分布高度相对较低。天敌对白蜡窄吉丁幼虫在空间资源的利用上也存在差异。在天津地区,白蜡吉丁柄腹茧蜂Spathius agrili Yang(膜翅目:茧蜂科)和啄木鸟(■形目:啄木鸟科)的跟随作用最强,且白蜡吉丁柄腹茧蜂与啄木鸟之间对空间资源的利用存在激烈竞争。在北京地区,啄木鸟的跟随作用最强,病原微生物、扁胫旋小蜂、白蜡吉丁柄腹茧蜂之间对空间和营养资源利用的竞争较激烈。而在辽宁地区,白蜡窄吉丁幼虫和天敌的分布范围较窄,各类天敌在空间上对寄主幼虫的跟随作用不是很明显,但不同天敌种类之间对空间和营养资源的利用也存在竞争。【结论】研究结果为白蜡窄吉丁的综合治理和天敌资源的保护利用提供了理论依据。     

温度对白蜡吉丁柄腹茧蜂发育和繁殖的影响

为了提高白蜡窄吉丁Agrilus planipennis的重要天敌——白蜡吉丁柄腹茧蜂Spathius agrili Yang的繁育效率, 本研究调查了不同温度（22, 24, 26, 28, 30和32℃）对该蜂生长发育及繁殖的影响。结果表明, 在22~32℃范围内, 白蜡吉丁柄腹茧蜂的卵、茧蛹及世代发育速率均随着温度的升高而加快, 而幼虫的发育速率在26℃时最快. 卵、幼虫、茧蛹和世代的发育起点温度分别为14.34, 16.89, 14.16和13.84℃, 有效积温分别为24.59, 61.16, 166.27和276.80日·度。温度对白蜡吉丁柄腹茧蜂的寄生率、寄生成功率、产卵量、子代蜂数量和性比都有不同程度的影响。据此提出适于人工繁育白蜡吉丁柄腹茧蜂的温度范围为26~28℃。     

白蜡吉丁柄腹茧蜂的寄生和繁殖生物学

白蜡吉丁柄腹茧蜂Spathius agrili Yang是我国天津地区寄生白蜡窄吉丁幼虫的一种重要天敌。本研究通过林间调查并结合室内观察,测定了该寄生蜂在不同营养条件下的存活时间和不同时期的产卵量,调查了不同寄主大小对寄生蜂后代生长发育的影响,以及寄生蜂对不同深度树皮下的寄主利用情况。结果表明：不同营养条件下白蜡吉丁柄腹茧蜂的存活时间差异很大,补充20％蜂蜜水的成蜂存活时间明显延长,最长可达3个月。相同条件下雌蜂的存活时间比雄蜂略长。在补充营养的条件下,茧蜂的产卵时间最长可达8周,产卵的高蜂在羽化后第2周前后,平均每雌每周产卵量在整个产卵期内基本保持在8～12粒上下,没有大的波动。茧蜂低龄幼虫也可能分泌毒素主动攻击寄主,提高了寄生蜂后代的生存能力。寄主个体大,营养量也大,寄生其上的茧蜂幼虫成功完成发育的数量随之增加,表明寄主营养资源的多少直接影响到寄生蜂后代的生长发育和存活。白蜡吉丁柄腹茧蜂对1～6.5 mm树皮深度范围内的适龄寄主幼虫均能够寄生利用,约92％的被寄生白蜡窄吉丁幼虫个体分布于1～4 mm深度的树皮下,在3 mm厚的树皮下分布的数量最多。茧蜂数量与树皮深度大致上呈抛物线关系,理论上在树皮深度为3.97 mm处茧蜂分布数量最多。     

白蜡吉丁肿腿蜂的生物学和生态学特性及繁殖技术研究

采用室内测定并结合林间实地调查的方法，研究了新发现的寄生于蛀干害虫白蜡窄吉丁的外寄生蜂——白蜡吉丁肿腿蜂Sclerodermus pupariae Yang et Yao的生物学特性及其人工繁殖技术。结果表明，白蜡吉丁肿腿蜂外寄生于白蜡窄吉丁的幼虫和蛹及预蛹，是自然控制该害虫的一种重要寄生蜂，自然寄生率为13.9%左右。该蜂在天津1年发生5代，世代重叠明显，寿命长。白蜡吉丁肿腿蜂的卵、幼虫、茧蛹期和全世代的发育起点温度分别为（16.89±0.79）℃，（17.03±1.42）℃，（16.90±1.68）℃和（15.31±0.47）℃。有效积温分别为（33.82±4.13）日·度、（49.11±7.93）日·度、（128.88±27.87）日·度和（277.00±14.15）日·度。雌性成蜂易于低温保存，对寄主有较强的搜索力和攻击力。繁蜂寄主容易大量获得，是生物防治白蜡窄吉丁的重要理想天敌之一。     

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。     

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.     

一种有效防控白蜡窄吉丁的阻隔网技术

[目的]白蜡窄吉丁Agrilus planipennis Fairmaire是一种危害极隐蔽、具有毁灭性的林木蛀干害虫.北京市延庆区栽植的洋白蜡Fraxinus pennsylvanica,受白蜡窄吉丁危害严重,当地林业保护站积极采取多种防控措施,其中阻隔网作为一种新的防控方法,可有效控制白蜡窄吉丁的危害.本研究调查不同防控管理措施林地内洋白蜡的受害情况,并估算不同管理措施所需的费用,以期评价阻隔网技术的防治效果.[方法]2017-2019年,在北京市延庆区选择使用阻隔网防治白蜡窄吉丁(样地1)、伐除受害严重的白蜡树后更植其它树种(样地2)和仅伐除受害严重的白蜡树(样地3)的样地3块,调查各样地内洋白蜡受害情况,调查指标包括:树冠枝枯梢率、树冠活力等级、羽化孔数量、萌蘖枝条数量和啄木鸟刻痕数.通过Tukey检验对各样地内地面调查指标进行差异性比较,并统计不同防控措施下每年单位面积样地的防治成本.[结果]样地1内白蜡保存率(100％)高于样地2(80％)和样地3(34％),且样地1的白蜡受害情况在树冠枝枯梢率、树冠活力等级、羽化孔数量、萌蘖枝条数量和啄木鸟刻痕数指标上显著低于样地2和样地3.样地1每年单位面积的防治成本为2150元/hm2,低于样地2(5625元/hm2)和样地3(5720元/hm2).[结论]使用阻隔网防治白蜡窄吉丁成效显著,且防治成本低,建议今后可与其它无公害防治技术相结合进行害虫综合治理.     

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。     

栗山天牛幼虫天敌白蜡吉丁肿腿蜂的转主寄主研究

在利用白蜡吉丁肿腿蜂Sclerodermus pupariae防治世代周期长、 发育整齐的害虫栗山天牛Massicus raddei中, 我们发现该天敌只在寄主1-3龄幼虫时具有良好的寄生效果, 而对3龄以上的中老龄幼虫基本不寄生。为了了解释放白蜡吉丁肿腿蜂后其种群能否在栎树林间延续下去, 达到持续控制靶标害虫的效果, 我们调查了其在栎树林间的转主寄主。结果表明： 白蜡吉丁肿腿蜂除了寄生栗山天牛外, 还能寄生双簇天牛Moechotypa diphysis、 四点象天牛Mesosa myops和栎块斑吉丁Lamprodila virgata幼虫。白蜡吉丁肿腿蜂寄生不同转主寄主幼虫的成功率差异显著, 对栎块斑吉丁和双簇天牛幼虫的寄生率最高, 分别为76.67%和73.33%。寄生上述4种寄主时, 其雌蜂在不同寄主上的产卵前期、 产卵量及第2代雌蜂出蜂量、 出蜂率、 体长和寿命等差异不显著, 只有幼虫期和蛹期历期及第2代雌蜂有翅率和雌雄性比略有差异。室内模拟林间寄生试验表明, 白蜡吉丁肿腿蜂对双簇天牛和四点象天牛幼虫的寄生率较高。对双簇天牛和四点象天牛野外生活史研究发现, 这2种天牛幼虫在白蜡吉丁肿腿蜂不适宜寄生的栗山天牛中老龄幼虫期、 蛹期和成虫期均大量存在, 表明双簇天牛和四点象天牛是白蜡吉丁肿腿蜂在栎树林中的主要转主寄主。本研究说明, 在利用白蜡吉丁肿腿蜂生物防治栗山天牛中, 白蜡吉丁肿腿蜂在林间寄生栗山天牛的1-3龄幼虫后, 可转移寄生双簇天牛等转主寄主, 以延续其种群, 待下一代栗山天牛幼龄幼虫出现时再转移寄生栗山天牛, 达到持续控制栗山天牛的防治效果。     

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.     

Quantifying the Impact of Woodpecker Predation on Population Dynamics of the Emerald Ash Borer (Agrilus planipennis)

The emerald ash borer (EAB), Agrilus planipennis, is an invasive beetle that has killed millions of ash trees (Fraxinus spp.) since it was accidentally introduced to North America in the 1990s. Understanding how predators such as woodpeckers (Picidae) affect the population dynamics of EAB should enable us to more effectively manage the spread of this beetle, and toward this end we combined two experimental approaches to elucidate the relative importance of woodpecker predation on EAB populations. First, we examined wild populations of EAB in ash trees in New York, with each tree having a section screened to exclude woodpeckers. Second, we established experimental cohorts of EAB in ash trees in Maryland, and the cohorts on half of these trees were caged to exclude woodpeckers. The following spring these trees were debarked and the fates of the EAB larvae were determined. We found that trees from which woodpeckers were excluded consistently had significantly lower levels of predation, and that woodpecker predation comprised a greater source of mortality at sites with a more established wild infestation of EAB. Additionally, there was a considerable difference between New York and Maryland in the effect that woodpecker predation had on EAB population growth, suggesting that predation alone may not be a substantial factor in controlling EAB. In our experimental cohorts we also observed that trees from which woodpeckers were excluded had a significantly higher level of parasitism. The lower level of parasitism on EAB larvae found when exposed to woodpeckers has implications for EAB biological control, suggesting that it might be prudent to exclude woodpeckers from trees when attempting to establish parasitoid populations. Future studies may include utilizing EAB larval cohorts with a range of densities to explore the functional response of woodpeckers.     

Effects of rearing conditions on reproduction of Spathius agrili (Hymenoptera: Braconidae), a parasitoid of the emerald ash borer (Coleoptera: Buprestidae)

Spathius agrili Yang (Hymenoptera: Braconidae) can be successfully reared on emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), larvae feeding in chambers drilled in small ash twigs that are wrapped with floral tape. Females maintained in groups with males for one week can receive enough sperm for production of female progeny throughout their lives. Volatiles released by emerald ash borer adults feeding on ash foliage increased parasitoid fecundity over ash foliage alone or no stimulus. The temperature at which the parasitoids were reared ranged from 20 to 25 degrees C in a daily cycle; however, raising the daily maximum temperature to 28 degrees C did not affect parasitoid longevity or fecundity. Adult females lived between 12 and 127 d, with an average of 60.8 +/- 4.5 d. Males lived slightly longer, with an average of 66 +/- 4.5 d. The first clutch of eggs was laid when the female was between 2 and 42 d old, with the average preoviposition period lasting 11.4 +/- 1.4 or 19.5 +/- 2.0 d in 2007 and 2009 trials, respectively. A higher proportion of the emerald ash borer larvae were feeding and thus attractive to parasitoids in the 2009 trial, and female S. agrili laid an average of 9.5 +/- 1.0 clutches containing 5.4 +/- 0.2 eggs, for an average of 51.2 eggs per female. Approximately three quarters of the progeny were female. The number of eggs per clutch was significantly greater when deposited on larger emerald ash borer larvae, further highlighting the need for quality larvae in rearing. Chilling S. agrili pupae at 10 degrees C to stockpile them for summer release was not successful; chilling resulted in lower survival and lower fecundity of emerging progeny. Female S. agrili proved capable of attacking emerald ash borer larvae through even the thickest bark of an ash tree that was 30-cm diameter at breast height. Even emerald ash borer larvae that were creating overwintering chambers in the outer sapwood of the tree were successfully attacked, suggesting that S. agrili could be reared on field collected logs infested with emerald ash borer.     

Temporal dynamics of woodpecker predation on emerald ash borer (Agrilus planipennis) in the northeastern U.S.A.

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Effects of learning experience on behaviour of the generalist parasitoid Sclerodermus pupariae to novel hosts

Massicus raddei Blessig (Coleoptera: Cerambycidae), also referred to as the oak long‐horned beetle (OLB), is a non‐natural host for the generalist parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae). To determine whether this generalist parasitoid might be a suitable agent for the control of OLB, the adaptive learning experience of adult female parasitoids to OLB larvae was investigated in the laboratory. A Y‐tube olfactometer bioassay was used to examine the effects of adaptive learning experience on the foraging ability of parasitoids for OLB larvae. The results indicated that parasitoids were significantly attracted by the volatiles of ash bark, Fraxinus velutina, with emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae and larval frass, after exposure to ash bark mixed with EAB larval frass (learning condition A). In contrast, after exposure to oak bark, Quercus liaotungensis, mixed with OLB larval frass (learning condition C), parasitoids showed significant preference for the volatiles of oak bark with OLB larvae and larval frass. On the basis of the results of no‐choice tests, we found that parasitoids exposed to learning condition C had greater paralysis efficiency and higher OLB larvae parasitism rates than those exposed to learning condition A or no experience. Furthermore, parasitoids fed on OLB larvae in learning condition C had significantly greater paralysis efficiency and higher OLB larvae parasitism rates than other parasitoids tested. Parasitoids fed on EAB larvae in learning condition A had the lowest paralysis efficiency and OLB larvae parasitism rates among the parasitoids tested. These findings suggested that adaptive learning significantly enhanced the ability of a generalist parasitoid to utilize a novel host. This may provide a new approach to controlling non‐natural hosts using generalist parasitoids.     

Differential utilization of ash phloem by emerald ash borer larvae: ash species and larval stage effects

• 1 Two experiments were performed to determine the extent to which ash species (black, green and white) and larval developmental stage (second, third and fourth instar) affect the efficiency of phloem amino acid utilization by emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae.
• 2 EAB larvae generally utilized green ash amino acids more efficiently than those of the other two species. For example, the concentrations of only six (two essential) and seven (two essential) amino acids were lower in frass from EAB that fed upon black and white ash than in the corresponding phloem, respectively. By contrast, concentrations of 16 (eight essential) amino acids were lower in the frass from EAB that fed upon green ash than in the phloem. In addition, in green ash, the frass : phloem ratios of 13 amino acids were lower than their counterparts in black and white ash.
• 3 The concentrations of non‐essential amino acids glycine and hydroxylproline were greater in frass than in phloem when EAB fed on black ash, although not when EAB fed on green or white ash.
• 4 The concentration of total phenolics (a group of putative defensive compounds to EAB, expressed as antioxidant activity of acetone extraction) was high in EAB frass but even higher in the phloem samples when the data were pooled across ash species and EAB larval stages. This suggests EAB larvae may eliminate phenolics through a combination of direct excretion and enzymatic conversion of phenolics to nonphenolics before excretion. Because the ratio of frass total phenolics to phloem total phenolics in white ash was lower than the ratios in black and green ash, the ability to destroy phenolics or convert them to nonphenolics was greater when EAB larvae fed on white ash.
• 5 Fourth‐instar EAB extracted phloem amino acids, including threonine, more efficiently than third‐instar EAB. The different larval developmental stages of EAB did not differ in their apparent ability to destroy phenolics or convert them to nonphenolics.

     

The effect of bark thickness on host partitioning between Tetrastichus planipennisi (Hymen: Eulophidae) and Atanycolus spp. (Hymen: Braconidae), two parasitoids of emerald ash borer (Coleop: Buprestidae)

Parasitoids have recently been introduced from Asia to aid in biological control in the United States of the invasive, highly damaging, emerald ash borer, Agrilus planipennis. Three introduced parasitoids have established and field biological studies are underway to improve our understanding of niche partitioning among them. Here we report one such investigation, a field experiment conducted to determine how outer bark thickness of ash trees might affect parasitism by one introduced (Tetrastichus planipennisi) and one native parasitoid (Atanycolus spp.). We found that T. planipennisi was unable to parasitize EAB larvae in trees with outer bark thicker than 3.2 mm (>11.2-cm DBH) whereas Atanycolus spp. parasitized EAB larvae in ash trees with outer bark up to 8.8 mm thick (>57.4-cm DBH). These results suggest that establishment of, and control by T. planipennisi at release sites with only large diameter trees is less likely, and that T. planipennisi will be more effective in stands with younger trees (<12-cm DBH). Releasing T. planipennisi near the leading edge of EAB invasion may have little impact on EAB populations if many ash trees are too large. We recommend releasing T. planipennisi in stands dominated by small, early successional or regenerating ash trees. This may maximize the establishment and effectiveness of this species. This limitation of T. planipennisi for biological control of emerald ash borer suggests that other EAB parasitoids from its native range with longer ovipositors, such as Spathius galinae (Hymenoptera: Braconidae), should be sought and evaluated for possible use as EAB biocontrol agents in the US. The results of this study also suggest the importance of parasitoid guild introduction for biological control in general, and hint at possible broader implications relating to resource partitioning among native and introduced parasitoids.