落叶松毛虫对兴安落叶松6种挥发物的触角电位反应

利用触角电位技术(EAG)测定了落叶松毛虫未交尾雌蛾、已交尾雌蛾、未交尾雄蛾对R型α-蒎烯、S型α-蒎烯、罗勒烯、S型β-蒎烯、水芹烯、莰烯等6种兴安落叶松挥发物的触角电位反应.结果表明:不同生理状况的落叶松毛虫对6种挥发物的7个浓度的EAG反应中,除对0.1 μg·μL-1水芹烯的EAG反应值是未交尾雌虫＞已交尾雌虫＞未交尾雄虫外,对水芹烯的其它不同浓度以及其它挥发物的EAG反应值均是已交尾雌蛾＞未交尾雌蛾＞未交尾雄蛾.未交尾雌、雄蛾的EAG反应差异显著(P＜0.05),已交尾雌蛾与未交尾雌、雄蛾之间差异极显著(P＜0.01),表明已交尾雌蛾对寄主植物气味更为敏感.落叶松毛虫未交尾雌蛾、已交尾雌蛾、未交尾雄蛾对水芹烯的EAG反应均比较敏感.在暗周期内已交尾雌蛾与未交尾雄蛾的活跃期在20:00-4:00,与其羽化、交尾、产卵期一致.     

落叶松毛虫对兴安落叶松9种挥发性物质的行为反应

为弄清兴安落叶松挥发性物质及其变化对落叶松毛虫寄主选择行为的影响,将兴安落叶松苗进行剪叶损伤和落叶松毛虫幼虫取食25%针叶、50%针叶处理.以健康苗为对照,研究落叶松毛虫成虫对不同状况寄主的产卵选择情况.同时,研究落叶松毛虫雌雄成虫对兴安落叶松R型α-蒎烯、s型α-蒎烯、S型β-蒎烯、叶醇、月桂烯、3-蒈烯,β-罗勒烯、水芹烯和莰烯等9种挥发物的趋性反应.结果表明:已交尾雌虫优先在健康植株上产卵,其产卵量占总产卵量的86.1%～89.5%;在不同程度损伤中,优先在轻度损伤的植株上产卵;机械伤苗上的产卵量要大于虫伤苗上的产卵量;将落叶松毛虫成虫触角剪去后,发现92.4%的卵被产在地面和笼网上,只有7.6%的卵产在苗上.在健康植株与损伤植株中,幼虫偏嗜健康植株,这与成虫偏嗜选择健康植株产卵是一致的.雌成虫对于β-罗勒烯、月桂烯和3-蒈烯,雄成虫对于莰烯和0.01,0.1,0.2,0.5μL·μL-1的叶醇均表现出负趋性,β-蒎烯、S型α-蒎烯在低含量时对雌雄成虫均具有诱引作用,高含量时则表现出驱避作用.R型α-蒎烯、水芹烯、莰烯和3-蒈烯对幼虫有诱引作用,月桂烯则具有驱避作用.     

Dynamic feature of flavonoids content in different organs of larch （Larix gmelinii）

Flavonoids in plants is very important in its ecological role and economic value. The dynamic features of flavonoids content in different organs of larch (Larix gmelinii) at different light and temperature conditions were investigated in this study. Results showed that the order of flavonoids content in different organs from high to low was 7.78% (stem bark)>2.79% (leaves) >1.72% (branches) >1.19% (stem xylem)and different organs had a great seasonal variation in flavonoids content, but the change of flavonoids content at different temperature was not obvious in different organs., The content of flavonoids in barck had, a positive correlation with temperature (R ²=0.75), but that in other organs had slight variation with the change of temperature. For all the tested organs, the flavonoids content in summer and autumn was approximately 3–4 times higher than in spring and winter. This is attributed to the great stress from environmental physical variables such as UV radiation, high temperature that induce the accumulation of flavonoids. The flavonoid content of sun leaves was evidently higher than that of shade leaves, and leaves at upper part of canopy had a higher flavonoids content compared with that at other parts. This result indicates that sun radiation could improve flavonoids production in leaves (R ²=0.76). The flavonoids may actively evolve in plant defenses to environmental stress, protecting larch from the damage of high temperature and radiation, and its main function is different in different organs. Foundation item: This paper is supported by the National Natural Science Foundation of China (30300271) and the Key Project of Chinese Ministry of Education (104191). Biography: WANG Wen-jie (1974-), male, Lecturer in Key Laboratory of Forest Plant Ecology, Northeast Forestry University, Harbin 150040, P. R. China. Responsible editor: Zhu Hong     

华山松大小蠹对9种植物挥发物的EAG和行为反应

选取5种非寄主挥发性化合物水杨醛、壬醛、三氯乙烯、二氯甲基甲醚和顺-3-己烯-1醇,4种寄主挥发性化合物β-蒎烯、(+)-3-蒈烯、β-石竹烯和(+)-α-蒎烯,应用触角电位仪和Y型嗅觉仪初步研究了华山松大小蠹成虫对这9种化合物的EAG反应和趋向反应。结果表明:雌虫对三氯乙烯、(+)-3-蒈烯和β-石竹烯,雄虫对壬醛、二氯甲基甲醚、(+)-3-蒈烯和(+)-α-蒎烯有明显的EAG反应;β-蒎烯、(+)-3-蒈烯和(+)-α-蒎烯对华山松大小蠹雌成虫有较强的引诱作用;三氯乙烯对雄成虫有较高的引诱率,且显著高于对雌成虫的引诱率。试验结果可为华山松大小蠹的行为控制技术研究提供借鉴。     

马尾松毛虫暴发机制分析

马尾松毛虫的暴发过程具有周期性和突发性相结合的特点,不同环境条件下,即不同的发生区,发生类型不一样,常灾区是典型的暴发发生类型,偶灾区和无灾区一般为渐进发生型、扩散发生型和低密度稳定发生型.暴发的原因一方面是马尾松毛虫具备了成为暴发种的遗传特性,另一方面是大面积马尾松纯林的存在.暴发的启动因子为高温干旱或由环境噪音引起的混沌动态,暴发的上升期为正密度相关作用驱使,下降期和潜伏期主要是由松树诱导抗性、严重失叶和天敌所引起的负密度相关的作用.松林和松林节肢动物群落结构的多样性是抑制马尾松毛虫暴发的主要因素.     

Reflectance features of water stressed Larix gmelinii needles

In recent years, Xing’an larch (Larix gmelinii) has been seriously infected by pests and drought. In order to improve the accuracy of monitoring the damage to larch by remote sensing (RS) and to predict the health of the larch, we studied the reflectance features of larch needles under different water conditions at the needle level by using the LIBERTY (Leaf Incorporating Biochemistry Exhibiting Reflectance and Transmittance Yields) model. Before applying the LIBERTY model, we recalibrated it for the needles of L. gmelinii based on ten field-measured spectral curves. After recalibration, LIBERTY can accurately model the needle reflectance spectra of L. gmelinii. Based on the recalibrated LIBERTY model, we extracted and analyzed the sensitive bands to needle water content by simulating the needle reflectance spectra under different drought conditions. Then, we established mathematical equations between the spectral indices (MSI, NDWI, and GVMI) and needle water content. Results show that the variations of larch needle water content can significantly change the needle spectra at the near-infrared and short-wave infrared bands. The higher the water content is, the higher the absorption peak is. We believe that our study will provide the theoretical basis and an optional method to investigate the forest water stress using multi-spectral or hyper-spectral remote sensing data.     

The influencen of stand state on the occurrence ofDendrolimus superans (Butler) in Daxing’an Mountains

Dendrolimus superans (Butler) occurred in the vast areas of Daxing’an Mountains forest, in Heilongjiang Province, in 1990. Stands damaged byI). Superans seriously, intermediately and lightly show patch shape distribution within damaged area. By ordination analysis and correlation analysis between ordination factors and coordinate axis. It was found that occurrence ofD Superans in different extent is owing to the effect of forest age, composition and exposure. The tended young pure forests grown on the sunward slopes are most seriously damaged and non-tended middle-age mixing stands grown on the sunless slopes are most lightly damaged. Therefore, in order to preventD. superans from seriously damaging larch forests, relatively large crown density (P > 0. 7) should be kept, and broadleaf trees should be also properly preserved. This research project was supported by Scientific and Techndogical Committee of Heilongjiang Province     

The study on site quality evaluation for dahurian larch (Larix gmelinii Rupr.)

Site quality of larch (Larix gmelinii Rupr.) in Da Xingan Mountain in Inner Mongolia was studied systematically with about 1000 temporary plots and computer software. The nature of site class and site index curve was discussed in view of general mathematical model of height growth. The height growth of dominant tree of four divisions of forest stands were studied and site index tables were constructed respectively. In order to unify the comparing standard and convenience for management in this region, site index table was constucted too. Based on discussing the action and accuracy of forest type, a series of site quality evaluation (forest type-site index class-site index) was suggested. This subject of study is supported by the National Natural Scientific Fund (No. 39270549).     

New cell line establishment of a cell line from embryos ofDendrolimus superans

A new cell line was established from 5-day-old embryonated eggs ofDendrolimus superans and has been designated NEAU-Ds-950821. The cell line consists of mixture of cell types, including majority of spherical shaped cells and a few of spindle shaped cells. The cell line has a population of doubling time of 52.6 h. Chromosome analysis revealed typical lepidopteran chromosomes. Isozyme characterization of Esterase showed the patterns were different from other three cell lines (Ms-927311, Xc-920730, and SF-21AE). Virus infectivity tests revealed the cell line can supportD. superans cytoplasmic polyhedrosis virus. The project is supported by National Science Foundation of China Responsible editor: Chai Ruihai     

基于加权信息量模型的油松毛虫灾害发生危险性评价

【目的】充分考虑影响灾害发生及灾害等级的气象、地形地貌等多种因素,实现多因子灾害发生危险性评价和制图,以期为灾前的防控和预警提供依据。【方法】以辽宁省西部的油松人工林为研究区,基于Landsat影像提取油松分布范围,以高程、坡向、坡度、降雨量、活动积温、日照时数、上一年灾害程度和距离上一年重灾区远近8个危险性评价因子,采用层次分析法确定各因子权重,建立加权信息量模型,结合GIS空间分析方法,将油松的受灾危险性划分为5个等级:极低危险区、低危险区、中危险区、高危险区和极高危险区,实现危险区划制图,并与实际灾害程度监测结果对比进行精度验证。【结果】1)根据信息量法原理,信息量值越大代表发生灾害的危险性越大。本文计算得到的各因子类别信息量值均与油松毛虫的生物学特性相吻合。2)研究区2017年虫灾极高和高危险区主要分布在建平县北部,中危险区主要分布在北部部分地区和建平县与凌源市交汇处,其他地区发生虫灾危险性较低,与实际调查结果相吻合。3)最终划分等级中的中低危险区和实际受灾油松失叶率大小对应关系明显,实际成灾油松林地中有90.32%被划分至高危险区和极高危险区。【结论】基于加权信息量模型的油松毛虫灾害发生危险性评价充分考虑了不同评价因子对灾害发生影响程度的差异,得到的风险区划结果较为准确,具有应用价值,可为大区域的森林病虫害危险性评价及风险评估提供技术依据。     

思茅松毛虫3个地理种群性信息素成分差异分析

[目的]分析思茅松毛虫雌成虫不同地理种群间性信息素成分、含量和比例差异,比较雄成虫触角敏感性和不同配比诱芯诱捕效果差异,开发不同地域有针对性的思茅松毛虫种群监测方法和诱捕防控技术。[方法]运用气相色谱(GC)分析思茅松毛虫湖南、云南和江西地理种群雌虫性信息素腺体提取物,利用触角电位仪(EAG)测定思茅松毛虫各地理种群雄蛾触角对性信息素标准品及其不同配比组分的电生理敏感性,然后进行林间生物测定,比较不同配比的性信息素成分诱蛾效果。[结果]思茅松毛虫性信息素腺体中含有顺5,反7-十二碳二烯乙酸酯(Z5,E7-12:OAc)、顺5,反7-十二碳二烯醇(Z5,E7-12:OH)和顺5-十二碳烯乙酸酯(Z5-12:OAc)3种成分,3个地理种群间这3种腺体成分含量和比例存在微小差异。3种成分以100:10:25或者100:10:10的比例配比,不同地理种群间均具有很好的触角电位活性和林间诱蛾活性。[结论]思茅松毛虫性信息素组分还没有发生明显的种下分化现象,3个地理种群间性信息素含量和比例的细微差异暗示着与地域差异、寄主植物差异和人为化学防控干扰等因素相关。     

Climatic responses of tree-ring widths of <Emphasis Type="Italic">Larix gmelinii</Emphasis> on contrasting north-facing and south-facing slopes in central Siberia

An analysis was performed on the climatic responses of the radial growth of Larix gmelinii (Rupr.) Rupr. on contrasting north-facing and south-facing slopes in Tura, central Siberia. We developed chronologies of tree-ring width for four plots, designated as north-upper, northlower, south-upper, and south-lower. Both residual and standard chronologies of tree-ring widths exhibited a significant positive correlation with temperature from the end of May until early June in all four plots. The chronologies of ring width did not reveal any major difference in the response to temperature among the four plots. The standard chronologies of ring widths on the north-facing slope were negatively correlated with precipitation during the winter (October–April) and in early and mid-May, whereas the residual chronologies did not reveal clear relationships with precipitation during the winter and May. The significant correlation between ring width and temperature from the end of May until early June indicates that temperatures in springtime play a significant role in the radial growth of L. gmelinii. The negative correlations between standard chronologies of tree-ring width and precipitation in the winter and in May on the north-facing slope indicate that lowfrequency fluctuations in snowfall have negative effects on the radial growth. However, these effects vary and depend on the microscale topography. Part of this report was presented at the 55th Annual Meeting of the Japan Wood Society, March 2005, Kyoto     

油松毛虫亚居群遗传结构的SSR分析

采用SSR分子标记技术对平泉县5个油松毛虫亚居群进行遗传多样性和遗传分化研究.结果显示:8对SSR引物对5个油松毛虫亚居群的扩增片段长度范围为78～430 bp,检测到的等位基因数为1～6个;种群总体水平多态位点比率P=87.50%,平均有效基因数A＝3.1250,平均期望杂合度He＝0.474 7,种群平均遗传距离为0.070 3～0.419 7;遗传分化度Fst=0.215 9,基因流Nm=0.908 1.由此可以得出在5个天然油松毛虫亚居群中,天然油松纯林居群内杂合度要比其他居群高;油松毛虫亚居群间已发生明显的遗传分化,基因交流较少,遗传漂变已经成为导致该物种种群分化的主要原因之一;油松毛虫亚居群遗传结构产生变异推测是由于常年化学防治和单一寄主植物造成的选择压力的影响.     

Comparison of host volatiles under different physiological states and electroantennogram responses of Semanotus bifasciatus （Motschulsky） to these volatiles

Based on an analytical TCT-GC/MS method, volatile compounds of branches of Platycladus orientalis under three physiological states （healthy, weak and baited trees） were analyzed. The results showed that α-pinene, β-phellandrene, 3-careen, limonene and thujopsene were released under all three physiological states. There were other kinds of volatiles released from branches of weak P. orientalis trees, such as camphene, γ-terpinene, （＋）-4-carene and cedrene, but these four volatiles compounds were not released from the branches of healthy P. orientalis. Volatile components from the physically damaged branches of P. orientalis had changed. β-terpinene, mycrene, isocaryophillene and caryophyllene were identified in both healthy and weak branches of P. orientalis. Only cedrene was identified in the baited wood. Electroantennogram （EAG） tests were conducted with nine volatile compounds from the branches of P. orientalis. Antennnae of male and female Semanotus bifasciatus （Motschulsky） were highly sensitive to these nine volatile compounds. Cedrene elicited the strongest response, whereas β-terpinene the lowest. Males and females showed no signifi-cant difference in their EAG responses to the nine volatile compounds.     

阿尔山地区兴安落叶松林下植物种群生态位

将由林龄(AG)、海拔(AL)以及物种数目(IS)组合而成的综合资源位指数(RGI)作为资源轴的划分标准,应用Levins(B_(sw)i)和Shannon-Wiener(B_(L)i)生态位宽度指数,生态位重叠指数(L_ih和L_hi)以及生态位相似性比例指数(C_ih)等将阿尔山地区不同起源的兴安落叶松林(天然林、人工林及火烧迹地恢复林)划分为6个资源位,在此基础上进行群落调查,对林下植物种群的生态位进行了定量分析。结果表明:兴安苔草和黑麦草的重要值最大,不同资源位重要值之和达到114.924 8%和70.911 1%。生态位宽度较大的有北方拉拉藤、粗根老鹳草、黑麦草及兴安苔草,这些物种适应能力强,分布范围大,倾向于泛化种。生态位狭小的植物种群则可能成为林区进行植物多样性保护的重点,需要引起特别关注。本研究证实生态位宽度较大种群与其他种群存在较大的生态位重叠和相似性,而这在某些重要值较低且生态位宽度较窄的种群也会出现(如铁杆蒿、黄花委陵菜及巴天酸模),生态位重叠和相似性并非仅仅取决于生态位宽度。生态位重叠既能反映两种群在共享资源时的竞争关系,也能反映两者在利用资源时相互促进的关系。生态位宽度较大的植物种可作为植被恢复阶段的先锋种,而与其生态位重叠较大的物种以及与先锋种生态位相似性比例较小的物种则可作为伴生种,以便于充分利用环境资源,逐步恢复受损地区的植被。     

Effect of fluoride pollution on genetic variability of <Emphasis Type="Italic">Larix gmelinii</Emphasis> (Pinaceae) in East Siberia

The genetic variability of four natural populations of Larix gmelinii from East Siberia was determined and compared by RAPD analysis. Comparison of the RAPD profiles provided an estimation of variability in 193 RAPD fragments. More than 89% of these fragments were found to be polymorphic. The main genetic variability parameters of the two populations from Central Yakutia, a region free of fluoride pollution, had considerably higher values than those from East Transbaikalia, a region potentially affected by fluoride pollution (FLU, near a fluorite quarry growing on soils with a high natural content of fluorides). AMOVA revealed that 72.94% of the variation was within populations, while only 7.05% of the variation was between populations within geographical regions. The genetic diversity of the FLU fluoride-tolerant population was the lowest, but only slightly lower than that of a fluoride non-tolerant population from Chita, 50 km distant from FLU. Although this study demonstrates the absence of fundamental alterations of genetic structure within the populations of L. gmelinii growing on soils with a high content of fluorides, it is presumed that the reduction of genetic diversity was the genetic response of the FLU population to such an environmental stress as a constantly high concentration of fluorides within the soil.     

大兴安岭兴安落叶松林树木生物量对氮沉降的响应

[目的]研究兴安落叶松生物量增长率对氮沉降是否存在响应,年际间是否存在差异;不同径级是否会影响兴安落叶松生物量增长率对氮沉降的响应。[方法]通过人工氮添加的方法模拟大气氮沉降变化,探究兴安落叶松林连续3年(2014—2016)树木生物量的变化对氮沉降的响应。从2012年开始,共设置4个氮添加水平,分别为对照(CK,0 g·m-2·a-1)、低氮(TL,2.5 g·m-2·a-1)、中氮(TM,5 g·m-2·a-1)和高氮(TH,7.5 g·m-2·a-1),每年的生长季(5—10月)按月将NH4NO3溶于32 L蒸馏水,利用背喷式喷雾器均匀喷洒于样方内,对照样地喷洒等量纯净水。[结果](1)3年间各施氮处理生物量增长率均显著高于对照组(P0.05)、各年份生物量增长率均存在显著差异(P0.05),而且2016年生物量增长率明显高于前两年(P0.05)。(2)2014年各处理间无显著差异;2015年TL处理的生物量增长率显著高于TM和TH处理(P0.05),2016年TL处理的生物量增长率显著高于TH处理(P0.05)。(3)不同处理对年际间不同径级生物量增长率也产生不同影响。TL处理对2014和2016年的10 15cm径级的兴安落叶松生物量增长率有显著促进作用(P0.05);TM处理对所有年份10 15 cm和15 20 cm径级兴安落叶松生物量增长率均有显著促进作用(P0.05);TH处理对所有年份胸径大于10 cm的3个径级(1015、15 20、20 cm)兴安落叶松生物量增长率均有显著促进作用(P0.05)。[结论](1)长期氮添加有可能导致树木生长率的增量逐渐减少甚至出现抑制,将改变现有北方森林生态系统碳源汇动态;(2)由于不同径级树木对氮沉降的响应不同,为提高氮沉降对北方森林碳库影响的模型预测精度,应按不同径级分别模拟。     

Impact of defoliation caused by the sawfly Cephalcia lariciphila (Hymenoptera: Pamphilidae) on radial growth of larch (Larix decidua Mill.)

The period between 2000 and 2002 repeated outbreaks of the web spinning sawfly Cephalcia lariciphila (Wachtl 1898) around the village of Větrny Jeníkov, Czech Republic. The impact of defoliation, caused by C. lariciphila feeding, on tree-ring formation of European larch (Larix decidua Mill.) was studied using dendrochronological methods. Heavy defoliation resulted in much less growth in the years of insect attack, the average incremental loss being 67% for the 2000–2002 period. Also, defoliation resulted in the formation of latewood with fewer cells and reduced cell-wall thickness.     

不同寄主植物对云南松毛虫生长发育、繁殖及生理酶活性的影响

目的 基于不同寄主植物对云南松毛虫（Dendrolimus houi Lajonquiere）生长发育、繁殖、解毒酶和消化酶活性的影响,解析其对不同寄主植物的生理适应机制,为营造新型混交林开展生态调控提供理论依据。 方法 以柳杉、侧柏、柏木、云南松和马尾松为寄主植物进行室内饲养,分别比较不同寄主植物条件下云南松毛虫幼虫、蛹的发育历期和成虫繁殖力,并测定云南松毛虫5龄幼虫的解毒酶（谷胱甘肽-S-转移酶（GSTs）、细胞色素P450酶（CYP450）、羧酸酯酶（CarE））和消化酶（蛋白酶、淀粉酶（AMS）、脂肪酶（LPS））的活性。 结果 取食柳杉、柏木、侧柏和云南松的云南松毛虫幼虫具完整的生活史,而取食马尾松的幼虫只存活至5龄,不具有完整的生活史。同时,取食柳杉和柏木的云南松毛虫幼虫和蛹发育历期短、雌蛹重且成虫产卵多。取食柳杉和柏木的云南松毛虫5龄幼虫体内谷胱甘肽-S-转移酶和细胞色素P450酶活性显著高于取食云南松、侧柏和马尾松的幼虫（P<0.05）。在3种消化酶中,取食柳杉与柏木的云南松毛虫幼虫体内蛋白酶活性均处于较高水平。 结论 云南松毛虫对柳杉和柏木具较好的适应性,虫体内解毒酶和蛋白酶可能是其对柳杉和柏木有较好适应性的主要原因。     

云南松毛虫线粒体基因组全序列测定和分析

目的 测定和分析了云南松毛虫线粒体基因组特征,从线粒体基因组水平探究鳞翅目蛾类昆虫高级阶元的系统发育关系。 方法 采用Illumina HiSeq测序方法测定了云南松毛虫线粒体全基因组序列,参考鳞翅目昆虫已知线粒体基因组的全序列对其各基因进行定位和注释。采用tRNA Scan-SE 2.0在线预测云南松毛虫线粒体基因组tRNA基因的二级结构。基于线粒体全基因组的蛋白编码序列构建了鳞翅目13个科32种蛾类昆虫的系统发育树和松毛虫属近缘种间的系统发育树。 结果 结果显示云南松毛虫线粒体基因组全长15 443 bp,包括13个蛋白质编码基因,22个tRNA基因,2个rRNA基因和一段长度为321 bp的A+T富含区,无基因重排,存在较高的A+T含量（80.0%）。13个蛋白编码基因中除了 ND2和 COX1,其余均以ATN做为起始密码子。9个蛋白编码基因共享相同的终止密码子TAA（ ND2、 ATP8、 ATP6、 COX3、 ND5、 ND4L、ND6、CYTB和 ND1）,其他4个基因的终止密码子都是残缺的, COX1、 COX2、 ND4以T为终止密码子, ND3以TA为终止密码子。22个tRNA基因中,除了tRNASer（AGN）由于缺少DHU臂无法构成三叶草结构,其余T均为典型的三叶草结构。整个线粒体结构与鳞翅目中目前已得到的其他昆虫线粒体基因组结构一致。 结论 系统发育分析结果显示,云南松毛虫与思茅松毛虫是完全不同的近缘种,与其他松毛虫亲缘关系也较远,云南松毛虫与6种近缘种的系统发育关系为：（（（（（油松毛虫+文山松毛虫）+赤松毛虫）+落叶松毛虫）+（思茅松毛虫+云南松毛虫））+家蚕）。鳞翅目蛾类各科之间的系统发育关系为：（（（（（（（（毒蛾科+灯蛾科）+夜蛾科）+舟蛾科）+（尺蛾科+（蚕蛾科+（天蛾科+大蚕蛾科））））+枯叶蛾科）+（螟蛾科+草螟科））+卷蛾科）+蝙蝠蛾科）。