白蜡窄吉丁的生物学及其生物防治研究
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摘要
白蜡窄吉丁Agrilus planinennis Fairmaire,又名花曲柳窄吉丁、梣小吉丁,属鞘翅目
(Colcoptera)吉丁甲科(Buprestidae),异名A.marcopoli Obenberger,是木犀科(Oleaceae)
梣属(Fraxinus)树木的一种重要蛀干害虫。主要分布于亚洲部分国家(中国、同本、
朝鲜、蒙古)以及俄罗斯远东地区,在国内分布于黑龙江、吉林、辽宁、山东、内蒙古、
河北、天津、台湾等地。严重危害我国北方重要的用材树种水曲柳 F. mandshurica和园
林绿化树种绒毛白蜡 F. velutina。
白蜡窄吉丁以幼虫在树木的韧皮部、形成层和木质部浅层蛀食为害,因其隐蔽性强,
防治极为困难。20世纪60年代在我国东北地区发生较重,当年东北的美国白蜡 E
americana和洋白蜡 F. pennsylvanica var.lanceolata因受害严重而不得不逐渐被淘汰。目
前,该害虫在东北地区为害水曲柳 F. mandshurica,成为我国这种具有重要经济价值树
种的重要害虫。近年来白蜡窄吉丁在我国河北、天津部分地区的绒毛白蜡 F. velutina上
严重发生。2002年发现该虫己传入北美某些地区如美国的Michigan和Ohio州、加拿大
的Ontario省等,并已在当地大面积暴发成灾,严重危害美国白蜡F.americana、美洲红
梣F.pennsylvanica、黑白蜡 F,nigra,至今其分布范围仍在不断扩大。
目前有关白蜡窄吉丁的研究报道很有限,缺乏详细的生活史和生物学特性等方面的
资料,更没有有效的防治策略。为此,2003至2005年我们对白蜡窄吉丁进行了系统的
调查研究,基本弄清了该虫的年生活史和生态学习性,主要结果和结论如下。
(1)调查了白蜡窄吉丁在国内的地理分布和寄主范围,并采用林间定期踏查结合
室内饲养观察等方法,研究了白蜡窄吉丁的年生活史和生物学特性。结果表明,白蜡窄
吉丁在我国天津一年发生l代。以老熟幼虫蛀入木质部浅层越冬,每年7月下旬即有老
熟幼虫陆续蛀入木质部建造蛹室,一直持续到11月上旬基本全部进入越冬状态。次年4
月上旬至5月中旬为蛹期,蛹平均历期约20 d。4月下旬可调查到蛹穴内蛰居尚未出孔
的成虫,5月上旬成虫羽化出孔,出孔高峰期在5月中旬。成虫平均寿命约21 d,成虫
期一直持续到7月上旬。成虫取食白蜡树叶补充营养,在树冠层叶面或树干上完成交配。
5月中旬成虫产卵,卵期较长,平均为15.74 d。6月上旬出现初孵幼虫,整个幼虫期在
全时代中持续时间最长,共约300 d左右。
The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), (= A. marcopoli Obenberger), is an important wood-boring beetle attacking ash trees (Fraxinus spp.) (Oleceae). It mainly distributed in partial Asia countries (China, Japan, Korea, Mongolia) and Far East Russia, while in China it presented in Heilongjiang, Jilin, Liaoning, Shandong, Inner Mongolia, Hebei, Tianjin, Taiwan, etc. The important timber species F. mandshurica and gardening plantations F. velutina were damaged severely in northern China.EAB larvae feed on the phloem, cambium, and shallow sapwood under ash tree bark. It is, therefore, very difficult to detect and control because of its highly concealed life history. In the 1960s', EAB severely infested F. Americana and F. pennsylvanica var. lanceolata in the northeastern China so that the two species ash trees had to be substituted gradually by other trees. And now the EAB also could damage F. mandshurica. Recently, outbreaks of this pest arose on F. velutina in parts of Tianjin and Hebei, China. In June 2002, the EAB was reported to be seriously infesting the white ash (F. americana), green ash (F. pennsylvanica) and black ash (F. nigra) in parts of North America, including Michigan and Ohio of USA, and Ontario of Canada. Subsequently, a large number of ash trees died in those areas, and as yet the infestation areas were continuously enlarging.Because the information on EAB is extremely limited, the detailed biology and life cycle of this pest are scarce, needless to say the effective control strategies. Therefore, we investigated the EAB from 2003 to 2005 to obtain data about its biological and ecological habits, the main results and findings were as follows.(1) The life cycle and biology of the EAB were studied using methods of regular investigations in the forest and observations in laboratory. Results indicated that the EAB are mostly univoltine in Tianjin, northern China. They overwintered as full-grown larvae in shallow chamber excavated in the outer sapwood by themselves. In late July, some late stage larvae began to build overwintering chambers, and all larvae entered the sapwood for dormancy before early November. The EAB pupated in the overwintering chamber in early April to mid May the following year, and the average pupal duration was about 20 days. In
late April, some newly eclosed adults could be found in the pupal cells, but they had not emerged from the host tree. Adults began to emerge in early May, with a peak in mid May. The average longevity of adults was about 21 days and the adulthood lasted through early July. The adults fed on ash foliages as a source of nutrition. Usually mating was conducted and completed on the leaf or trunk surface of ash tree. Female adults oviposited in mid May and eggs hatched on average in 15.74 days. The first instar larvae appeared in early June. The larval stage lasted about 300 days to complete an entire generation.(2) The number of larval instars and stadium of each instar of the EAB were determined by sampling periodically in forests. Six variables including the body length, body width, width of the pronotum, width of the peristoma, length of the urogomphus, and anteriormost width of the urogomphus were examined under binocular microscope with an eyepiece micrometer for choosing the optimal morphological characters or index for the separation of larval instars. Results showed that the larvae of emerald ash borer have 4 instars. The urogomphus and its length were the best characteristic and variable for the separation of larval instars, and the width of peristoma was the second best according to the overlapping degree of variable. Body length, body width, width of the prothoracic plate, and anteriormost width of the urogomphus were not reliable due to high variation. Morphological characters on the inner side of larval urogomphus are also helped to separate the instars tersely and conveniently. The average duration of each instar measured by the median method of population in field was as follows: 18.14 days for the 1st instar, 11.23 days for the 2nd instar, 10.71 days for the 3rd instar, and 261.20 days for the 4th instar. The total larval stage duration was 301.28 days naturally.(3) The species of natural enemies attacked EAB in every stage have been investigated in Tianjin, Jilin, Liaoning, and Helongjiang with methods of forest investigation and laboratory rearing. A total of 16 species of natural enemies were discovered primarily, including 6 parasitoids, 2 woodpeckers, 4 ants, 1 spider, 1 mite, and 2 pathogenic microorganisms. The dominant parasitic natural enemies in the host larval stage were Spathius agrili Yang (Hymenoptera: Braconidae) (Tianjin) and Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) (northeastern China). Both the two parasitoids were new species. In addition, a Scleroderma sp. which parasitized EAB pupa found in Tianjin also showed better utilization prospects in biological control.
(4) Spathius agrili Yang is a new important obligatory natural enemy ectoparasitizing the larvae of emerald ash borer, Agrilus planipennis Fairmaire. The wasp generally occurs 3-4 generations, seldom 2 generations, one year in Tianjin, China, and overwinters in host gallery as a full-grown larva in the cocoon. The generations of S. agrili overlaps heavily as their so varied emergence time of overwintered adults. The sex ratios (female: male) are about 3:1. The adults can live 29.14 days for female and 23.64 days for male in average when fed with 20% honey solution. And the longevities decrease sharply without nutrition supplementation. Amphigony is the natural mode of reproduction, but female also can oviposit without mating under certain conditions and the progeny from parthenogenesis are all males. The mean oviposition is 23.25 eggs per female in laboratory. S. agrili has 5 larval instars and larvae feed on the host haemolymph gregariously. The developmental generation time from egg to adult wasp is 27-28 days at room temperature 22-26℃. The developmental threshod of the overwintering cocoon and thermal requirement of the whole generation are 14.12℃ and 386.57 degree-days respectively. The natural parasitism rates of this wasp can reach 40%-60%, which suggests that the parasitoid has excellent potentials for utilization on the biological control of emerald ash borer.(5) The spatial patterns of the larvae of EAB, the parasitoid Spathius agrili Yang, and pecks caused by woodpeckers preying EAB were investigated in forests. Results showed that the larva of EAB were distributed in a negative binomial fashion horizontally in forests, as was the parasitoid S. agrili and woodpecker attacks. In the vertical direction, the EAB larvae were distributed on ash trunks from above ground to 3.37 m in height, with a peak around 1.50 m. Natural enemies also favored searching in this range.(6) The differences of emergence date between overwintering S. agrili and its host, parasitism rates at different periods in forests, relations between parasitism rates and host densities, and relationships between ovipositions of braconid wasp and body sizes of host larvae were investigated using methods of regular surveys in forests and observations in laboratory. Results revealed that the emergence of S. agrili was more than one month later than that of its host, suggesting good synchrony between parasitoid emergence and host availability. The overwintering S. agrili emerged from mid June to mid August with the peak
in July in 2003, while the emergence date lasted from late May through late July with peak in late June to early July in 2004. But the EAB emergence period ranged in mid-late May in 2003, and in mid April through mid May in 2004 (in laboratory). The emergence date of natural enemy asynchronously inosculated with the optimum developmental instars of the earliest host larvae, which was the results of a long-term co-evolution between the two species. It was also revealed that the parasitoid S. agrili could be a specialized natural enemy. The natural parasitism rates of EAB larvae attacked by S. agrili gradually increased on the whole at different periods in forests. The body sizes of host larvae, i.e. larval instar, affected the decision of parasitoid S. agrili to lay eggs or not. This wasp oviposited only on those host larvae with a prontum and body width more than 1.5 mm and a body length in excess of 12 mm, i.e. the 3rd or 4th instar larvae. Egg deposition by female wasps showed no significant relations with body sizes of host larvae only if the parasitoids accepted hosts. These findings would consequentially contribute to the further successfully biological control of the borer.(7) The mass rearing techniques of natural enemy Spathius agrili Yang parasitized the EAB, were researched in laboratory using 4 approaches. Also host ranges of the braconid were determined with methods of investigation in forests plus examination in laboratory. Primary results showed that the methods of 'branch segment' and 'paper package' succeeded in artificial production of the parasitoid under room conditions. S. agrili could exactly distinguish host larva which had been parasitized from healthy ones. At present, no any other insect was found to be the host of this braconid except the EAB. The possibilities for the rearing of parasitoid larvae using non-host larvae as substitute feed were also studied and discussed.Through these investigations and researches, the biology and ecology of the emerald ash borer and its important parasitic natural enemy Spathius agrili Yang have been solved primarily as well as the relationships between the two insects. These findings would consequentially contribute to the further successfully biological control of the bark beetle.
(Colcoptera)吉丁甲科(Buprestidae),异名A.marcopoli Obenberger,是木犀科(Oleaceae)
梣属(Fraxinus)树木的一种重要蛀干害虫。主要分布于亚洲部分国家(中国、同本、
朝鲜、蒙古)以及俄罗斯远东地区,在国内分布于黑龙江、吉林、辽宁、山东、内蒙古、
河北、天津、台湾等地。严重危害我国北方重要的用材树种水曲柳 F. mandshurica和园
林绿化树种绒毛白蜡 F. velutina。
白蜡窄吉丁以幼虫在树木的韧皮部、形成层和木质部浅层蛀食为害,因其隐蔽性强,
防治极为困难。20世纪60年代在我国东北地区发生较重,当年东北的美国白蜡 E
americana和洋白蜡 F. pennsylvanica var.lanceolata因受害严重而不得不逐渐被淘汰。目
前,该害虫在东北地区为害水曲柳 F. mandshurica,成为我国这种具有重要经济价值树
种的重要害虫。近年来白蜡窄吉丁在我国河北、天津部分地区的绒毛白蜡 F. velutina上
严重发生。2002年发现该虫己传入北美某些地区如美国的Michigan和Ohio州、加拿大
的Ontario省等,并已在当地大面积暴发成灾,严重危害美国白蜡F.americana、美洲红
梣F.pennsylvanica、黑白蜡 F,nigra,至今其分布范围仍在不断扩大。
目前有关白蜡窄吉丁的研究报道很有限,缺乏详细的生活史和生物学特性等方面的
资料,更没有有效的防治策略。为此,2003至2005年我们对白蜡窄吉丁进行了系统的
调查研究,基本弄清了该虫的年生活史和生态学习性,主要结果和结论如下。
(1)调查了白蜡窄吉丁在国内的地理分布和寄主范围,并采用林间定期踏查结合
室内饲养观察等方法,研究了白蜡窄吉丁的年生活史和生物学特性。结果表明,白蜡窄
吉丁在我国天津一年发生l代。以老熟幼虫蛀入木质部浅层越冬,每年7月下旬即有老
熟幼虫陆续蛀入木质部建造蛹室,一直持续到11月上旬基本全部进入越冬状态。次年4
月上旬至5月中旬为蛹期,蛹平均历期约20 d。4月下旬可调查到蛹穴内蛰居尚未出孔
的成虫,5月上旬成虫羽化出孔,出孔高峰期在5月中旬。成虫平均寿命约21 d,成虫
期一直持续到7月上旬。成虫取食白蜡树叶补充营养,在树冠层叶面或树干上完成交配。
5月中旬成虫产卵,卵期较长,平均为15.74 d。6月上旬出现初孵幼虫,整个幼虫期在
全时代中持续时间最长,共约300 d左右。
The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), (= A. marcopoli Obenberger), is an important wood-boring beetle attacking ash trees (Fraxinus spp.) (Oleceae). It mainly distributed in partial Asia countries (China, Japan, Korea, Mongolia) and Far East Russia, while in China it presented in Heilongjiang, Jilin, Liaoning, Shandong, Inner Mongolia, Hebei, Tianjin, Taiwan, etc. The important timber species F. mandshurica and gardening plantations F. velutina were damaged severely in northern China.EAB larvae feed on the phloem, cambium, and shallow sapwood under ash tree bark. It is, therefore, very difficult to detect and control because of its highly concealed life history. In the 1960s', EAB severely infested F. Americana and F. pennsylvanica var. lanceolata in the northeastern China so that the two species ash trees had to be substituted gradually by other trees. And now the EAB also could damage F. mandshurica. Recently, outbreaks of this pest arose on F. velutina in parts of Tianjin and Hebei, China. In June 2002, the EAB was reported to be seriously infesting the white ash (F. americana), green ash (F. pennsylvanica) and black ash (F. nigra) in parts of North America, including Michigan and Ohio of USA, and Ontario of Canada. Subsequently, a large number of ash trees died in those areas, and as yet the infestation areas were continuously enlarging.Because the information on EAB is extremely limited, the detailed biology and life cycle of this pest are scarce, needless to say the effective control strategies. Therefore, we investigated the EAB from 2003 to 2005 to obtain data about its biological and ecological habits, the main results and findings were as follows.(1) The life cycle and biology of the EAB were studied using methods of regular investigations in the forest and observations in laboratory. Results indicated that the EAB are mostly univoltine in Tianjin, northern China. They overwintered as full-grown larvae in shallow chamber excavated in the outer sapwood by themselves. In late July, some late stage larvae began to build overwintering chambers, and all larvae entered the sapwood for dormancy before early November. The EAB pupated in the overwintering chamber in early April to mid May the following year, and the average pupal duration was about 20 days. In
late April, some newly eclosed adults could be found in the pupal cells, but they had not emerged from the host tree. Adults began to emerge in early May, with a peak in mid May. The average longevity of adults was about 21 days and the adulthood lasted through early July. The adults fed on ash foliages as a source of nutrition. Usually mating was conducted and completed on the leaf or trunk surface of ash tree. Female adults oviposited in mid May and eggs hatched on average in 15.74 days. The first instar larvae appeared in early June. The larval stage lasted about 300 days to complete an entire generation.(2) The number of larval instars and stadium of each instar of the EAB were determined by sampling periodically in forests. Six variables including the body length, body width, width of the pronotum, width of the peristoma, length of the urogomphus, and anteriormost width of the urogomphus were examined under binocular microscope with an eyepiece micrometer for choosing the optimal morphological characters or index for the separation of larval instars. Results showed that the larvae of emerald ash borer have 4 instars. The urogomphus and its length were the best characteristic and variable for the separation of larval instars, and the width of peristoma was the second best according to the overlapping degree of variable. Body length, body width, width of the prothoracic plate, and anteriormost width of the urogomphus were not reliable due to high variation. Morphological characters on the inner side of larval urogomphus are also helped to separate the instars tersely and conveniently. The average duration of each instar measured by the median method of population in field was as follows: 18.14 days for the 1st instar, 11.23 days for the 2nd instar, 10.71 days for the 3rd instar, and 261.20 days for the 4th instar. The total larval stage duration was 301.28 days naturally.(3) The species of natural enemies attacked EAB in every stage have been investigated in Tianjin, Jilin, Liaoning, and Helongjiang with methods of forest investigation and laboratory rearing. A total of 16 species of natural enemies were discovered primarily, including 6 parasitoids, 2 woodpeckers, 4 ants, 1 spider, 1 mite, and 2 pathogenic microorganisms. The dominant parasitic natural enemies in the host larval stage were Spathius agrili Yang (Hymenoptera: Braconidae) (Tianjin) and Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) (northeastern China). Both the two parasitoids were new species. In addition, a Scleroderma sp. which parasitized EAB pupa found in Tianjin also showed better utilization prospects in biological control.
(4) Spathius agrili Yang is a new important obligatory natural enemy ectoparasitizing the larvae of emerald ash borer, Agrilus planipennis Fairmaire. The wasp generally occurs 3-4 generations, seldom 2 generations, one year in Tianjin, China, and overwinters in host gallery as a full-grown larva in the cocoon. The generations of S. agrili overlaps heavily as their so varied emergence time of overwintered adults. The sex ratios (female: male) are about 3:1. The adults can live 29.14 days for female and 23.64 days for male in average when fed with 20% honey solution. And the longevities decrease sharply without nutrition supplementation. Amphigony is the natural mode of reproduction, but female also can oviposit without mating under certain conditions and the progeny from parthenogenesis are all males. The mean oviposition is 23.25 eggs per female in laboratory. S. agrili has 5 larval instars and larvae feed on the host haemolymph gregariously. The developmental generation time from egg to adult wasp is 27-28 days at room temperature 22-26℃. The developmental threshod of the overwintering cocoon and thermal requirement of the whole generation are 14.12℃ and 386.57 degree-days respectively. The natural parasitism rates of this wasp can reach 40%-60%, which suggests that the parasitoid has excellent potentials for utilization on the biological control of emerald ash borer.(5) The spatial patterns of the larvae of EAB, the parasitoid Spathius agrili Yang, and pecks caused by woodpeckers preying EAB were investigated in forests. Results showed that the larva of EAB were distributed in a negative binomial fashion horizontally in forests, as was the parasitoid S. agrili and woodpecker attacks. In the vertical direction, the EAB larvae were distributed on ash trunks from above ground to 3.37 m in height, with a peak around 1.50 m. Natural enemies also favored searching in this range.(6) The differences of emergence date between overwintering S. agrili and its host, parasitism rates at different periods in forests, relations between parasitism rates and host densities, and relationships between ovipositions of braconid wasp and body sizes of host larvae were investigated using methods of regular surveys in forests and observations in laboratory. Results revealed that the emergence of S. agrili was more than one month later than that of its host, suggesting good synchrony between parasitoid emergence and host availability. The overwintering S. agrili emerged from mid June to mid August with the peak
in July in 2003, while the emergence date lasted from late May through late July with peak in late June to early July in 2004. But the EAB emergence period ranged in mid-late May in 2003, and in mid April through mid May in 2004 (in laboratory). The emergence date of natural enemy asynchronously inosculated with the optimum developmental instars of the earliest host larvae, which was the results of a long-term co-evolution between the two species. It was also revealed that the parasitoid S. agrili could be a specialized natural enemy. The natural parasitism rates of EAB larvae attacked by S. agrili gradually increased on the whole at different periods in forests. The body sizes of host larvae, i.e. larval instar, affected the decision of parasitoid S. agrili to lay eggs or not. This wasp oviposited only on those host larvae with a prontum and body width more than 1.5 mm and a body length in excess of 12 mm, i.e. the 3rd or 4th instar larvae. Egg deposition by female wasps showed no significant relations with body sizes of host larvae only if the parasitoids accepted hosts. These findings would consequentially contribute to the further successfully biological control of the borer.(7) The mass rearing techniques of natural enemy Spathius agrili Yang parasitized the EAB, were researched in laboratory using 4 approaches. Also host ranges of the braconid were determined with methods of investigation in forests plus examination in laboratory. Primary results showed that the methods of 'branch segment' and 'paper package' succeeded in artificial production of the parasitoid under room conditions. S. agrili could exactly distinguish host larva which had been parasitized from healthy ones. At present, no any other insect was found to be the host of this braconid except the EAB. The possibilities for the rearing of parasitoid larvae using non-host larvae as substitute feed were also studied and discussed.Through these investigations and researches, the biology and ecology of the emerald ash borer and its important parasitic natural enemy Spathius agrili Yang have been solved primarily as well as the relationships between the two insects. These findings would consequentially contribute to the further successfully biological control of the bark beetle.
引文
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张喆.国外花曲柳窄吉丁危害与防治现状简介.天津农林科技,2004,(2):27～28
张良玉,陈周羡,杨国栋,等.绒毛白蜡窄吉丁虫防治技术的研究.园林科技通讯,1995,(4):1～7
张连芹,宋世涵,范景祥.斑头陡盾茧蜂防治粗鞘双条杉天牛的研究.林业科学,1987,23(3):306～313
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张俊文.杨树吉丁虫幼虫空间分布型及其应用.吉林林业科技,1994,(3):21～22,29
李磊,邹运鼎,毕守东,等.棉蚜和草间小黑蛛种群空间格局的地统计学研究.应用生态学报,2004,15(6):1043～1046
李中庭.粗鞘双条杉天牛生物防治初步研究.湖南林业科技,1986,(1):21
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邵崇斌,徐振武,韩明耀.杨树蛀干害虫空间格局的研究.西北林学院学报,1997,12(增刊):33～36
邹运鼎,毕守东,王祥胜,等.麦长管蚜及蚜茧蜂空间格局的地统计学研究.应用生态学报,2001,12(6):887～891
邹运鼎.毕守东,周夏芝,等.桃—点叶蝉及草间小黑蛛空间格局的地统计学研究.应用生态学报,2002,13(12):1645～1648
邹运鼎.周夏芝,毕守东,等.桃—点叶蝉及其天敌草间小黑蛛种群三维空间格局动态.应用生态学报,2003,14(9):1485～1488
邹运鼎.害虫管理中的天敌评价理论与应用.北京:中国林业出版社,1997,50～64
陈子星,徐夕水.生物统计SAS程序题解.北京:中国农业科学院计算中心,1997,38～40,117～123
陈永年.潘桐.也谈昆虫幼虫头宽及摄食量的增长规律.昆虫知识,1988,25(4):235～240
周明华,冉俊祥,杜国兴,等.绿梣树吉丁虫.植物检疫,2004,18(2):88～90
周祖基,曾垂惠,杨伟,等.川硬皮肿腿蜂环境适应性的初步研究.森林病虫通讯,1996,(3):4～6
郝树广,张孝羲,程遐年.稻田节肢动物群落优势功能集团的垂直分布,数量动态及天敌作用估计.应用生态学报,2000,11(1):103～107
彩万志,庞雄飞,花保祯,等编.普通昆虫学.北京:中国农业大学出版社,2001,236
黄保宏,邹运鼎,毕守东,等.朝鲜球坚蚧及黑缘红瓢虫空间格局的地统计学研究.应用生态学报,2003,14(3):413～417
黄琼,周祖基,杨伟,等.川硬皮肿腿蜂离体培养初探.昆虫天敌,2005,27(1):43-48
曾垂惠,杨德敏,叶伟军,等.两色刺足茧蜂研究初报.四川林业科技,1997,18(3):46～50
程惠珍,孟宪华,陈君,等.肿腿蜂对蛀干害虫控制效能的评估.中药材,2003,26(1):1～3
谢中能,李理,谢以权.白虫小茧蜂的离体培养.昆虫学报,1989,32(4):433～437
潘志刚,游应天编著.中国主要外来树种引种栽培.北京:北京科学技术出版社,1994,32～643
戴志一.昆虫幼虫种群众数龄期的测定法.昆虫知识,1980,17(4):185～187
Bjφrnstad O N, Sait S M, Stenseth N C, et al. The impact of specialized enemies on the dimensionality of host dynamics. Nature, 2001, 409: 1001～1006
Cassie R M. Frequency distribution models in the ecology of plankton and other organisms. Journal of Animal Ecology, 1962, 31: 65～92
Haack R A, Benjamin D M. The biology and ecology of the twolined chestnut borer, Agrilus bilineatus (Coleoptera: Buprestidae). The Canadian Entomologist, 1982, 114: 385～396
Haack R A, Jendek E, Liu H P, et al. The emerald ash borer: a new exotic pest in North America. Newsletter of the Michigan Entomological Society, 2002, 47(3&4): 1～5
Hawkins B A, Comell H V. Maximum parasitism rates and successful biological control Science, 1994, 266: 1886
http://www.inspection.gc.ca/english/plaveg/protect/dir/d-03-08e.shtml "Phytosanitary Requirements to Prevent the Introduction into and Spread within Canada of the Emerald Ash Borer, Agrilus planipennis (Fairmaire), into Canada"
http://www.michigan.gov/mda/Emerald Ash Borer "Emerald Ash Borer Michigan Eradication Strategy"-PDF
http://www.ncrs.fs.fed.us/4501/eab/downloads/statusreportAugust2002.pdf "Status Report-Emerald Ash Borer: A New Threat to North American Forests"-PDF
http://www.ohioagriculture.gov/eab/eab-nr-workwithpatrol-102504.stm "Agriculture Officials Work With State Highway Patrol to Enforce Emerald Ash Borer Quarantines"
Jendek E. Studies in the East Palaearctic species of the genus Agrilus Dahl, 1823 (Coleptera: Buprestidae) Part Ⅰ. Entomological Problems, 1994, 25(1): 9～25
Lee H J. Ovipositional mechanism of an ecto-parasitoid, Goniozus legneri Gordh (Hymenoptera: Bethylidae). Chinese Journal of Entomology, 1992, 12(3): 193～199
Liu H P, Bauer L S, Gao R T, et al. Exploratory survey for the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae), and its natural enemies in China. The Great Lakes Entomologist, 2003, 36(3&4): 191～204
Lloyd M. Mean crowding. Journal of Animal Ecology, 1967, 36: 1～30
Loerch C R, Cameron E A. Determination of larval instars of the bronze birch borer, Agrilus anxius (Coleoptera: Buprestidae). Annals of the Entomological Society of America, 1983, 76: 948～952
Rivero A. The relationship between host selection behaviour and offspring fitness in a koinobiont parasitoid. Ecological Entomology, 2000, 25: 467～472
Roland J, Taylor P D. Insect parasitoid species respond to forest structure at different spatial scales. Nature, 1997, 386: 710～713
Southwood T R E. Ecological methods (2nd Edition). London: Chapman and Hall, 1978
Wang K W, Jiang J X, You L S, et al. Preference of Microplitis sp. to host age. Journal of Hunan Agricultural University (Natural Sciences), 2001, 27(5): 367～369
Waters W E. A quantitative measure of aggregation in insects. Journal of Economic Entomology, 1959, 52: 1180～1184
Wei X, Reardon D, Wu Y, et al. Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in China: a review and distribution survey. 昆虫学报,2004,47(5):679～685
Yang Z Q, Strazanac J, Marsh P, et al. A new important braconid species parasitizing larva of emerald ash borer from China. Annals of the Entomological Society of America, 2005, 98(1): (in press)
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邬恤民.柑桔爆皮虫(Agrilus auriventris Saunders)及其防治.植物保护学报,1964,3(1):61～71
邬祥光.昆虫生态学的常用数学分析方法(修订版).北京:中国农业出版社,1995
张喆.国外花曲柳窄吉丁危害与防治现状简介.天津农林科技,2004,(2):27～28
张良玉,陈周羡,杨国栋,等.绒毛白蜡窄吉丁虫防治技术的研究.园林科技通讯,1995,(4):1～7
张连芹,宋世涵,范景祥.斑头陡盾茧蜂防治粗鞘双条杉天牛的研究.林业科学,1987,23(3):306～313
张连芹.利用天牛肿腿蜂防治粗鞘双条杉天牛试验.林业科技通讯,1982,(9):10
张俊文.杨树吉丁虫幼虫空间分布型及其应用.吉林林业科技,1994,(3):21～22,29
李磊,邹运鼎,毕守东,等.棉蚜和草间小黑蛛种群空间格局的地统计学研究.应用生态学报,2004,15(6):1043～1046
李中庭.粗鞘双条杉天牛生物防治初步研究.湖南林业科技,1986,(1):21
李吕丕.人工繁殖管氏肿腿蜂防治双条杉天牛的试验.林业科技通讯,1993,(4):13
邵崇斌,徐振武,韩明耀.杨树蛀干害虫空间格局的研究.西北林学院学报,1997,12(增刊):33～36
邹运鼎,毕守东,王祥胜,等.麦长管蚜及蚜茧蜂空间格局的地统计学研究.应用生态学报,2001,12(6):887～891
邹运鼎.毕守东,周夏芝,等.桃—点叶蝉及草间小黑蛛空间格局的地统计学研究.应用生态学报,2002,13(12):1645～1648
邹运鼎.周夏芝,毕守东,等.桃—点叶蝉及其天敌草间小黑蛛种群三维空间格局动态.应用生态学报,2003,14(9):1485～1488
邹运鼎.害虫管理中的天敌评价理论与应用.北京:中国林业出版社,1997,50～64
陈子星,徐夕水.生物统计SAS程序题解.北京:中国农业科学院计算中心,1997,38～40,117～123
陈永年.潘桐.也谈昆虫幼虫头宽及摄食量的增长规律.昆虫知识,1988,25(4):235～240
周明华,冉俊祥,杜国兴,等.绿梣树吉丁虫.植物检疫,2004,18(2):88～90
周祖基,曾垂惠,杨伟,等.川硬皮肿腿蜂环境适应性的初步研究.森林病虫通讯,1996,(3):4～6
郝树广,张孝羲,程遐年.稻田节肢动物群落优势功能集团的垂直分布,数量动态及天敌作用估计.应用生态学报,2000,11(1):103～107
彩万志,庞雄飞,花保祯,等编.普通昆虫学.北京:中国农业大学出版社,2001,236
黄保宏,邹运鼎,毕守东,等.朝鲜球坚蚧及黑缘红瓢虫空间格局的地统计学研究.应用生态学报,2003,14(3):413～417
黄琼,周祖基,杨伟,等.川硬皮肿腿蜂离体培养初探.昆虫天敌,2005,27(1):43-48
曾垂惠,杨德敏,叶伟军,等.两色刺足茧蜂研究初报.四川林业科技,1997,18(3):46～50
程惠珍,孟宪华,陈君,等.肿腿蜂对蛀干害虫控制效能的评估.中药材,2003,26(1):1～3
谢中能,李理,谢以权.白虫小茧蜂的离体培养.昆虫学报,1989,32(4):433～437
潘志刚,游应天编著.中国主要外来树种引种栽培.北京:北京科学技术出版社,1994,32～643
戴志一.昆虫幼虫种群众数龄期的测定法.昆虫知识,1980,17(4):185～187
Bjφrnstad O N, Sait S M, Stenseth N C, et al. The impact of specialized enemies on the dimensionality of host dynamics. Nature, 2001, 409: 1001～1006
Cassie R M. Frequency distribution models in the ecology of plankton and other organisms. Journal of Animal Ecology, 1962, 31: 65～92
Haack R A, Benjamin D M. The biology and ecology of the twolined chestnut borer, Agrilus bilineatus (Coleoptera: Buprestidae). The Canadian Entomologist, 1982, 114: 385～396
Haack R A, Jendek E, Liu H P, et al. The emerald ash borer: a new exotic pest in North America. Newsletter of the Michigan Entomological Society, 2002, 47(3&4): 1～5
Hawkins B A, Comell H V. Maximum parasitism rates and successful biological control Science, 1994, 266: 1886
http://www.inspection.gc.ca/english/plaveg/protect/dir/d-03-08e.shtml "Phytosanitary Requirements to Prevent the Introduction into and Spread within Canada of the Emerald Ash Borer, Agrilus planipennis (Fairmaire), into Canada"
http://www.michigan.gov/mda/Emerald Ash Borer "Emerald Ash Borer Michigan Eradication Strategy"-PDF
http://www.ncrs.fs.fed.us/4501/eab/downloads/statusreportAugust2002.pdf "Status Report-Emerald Ash Borer: A New Threat to North American Forests"-PDF
http://www.ohioagriculture.gov/eab/eab-nr-workwithpatrol-102504.stm "Agriculture Officials Work With State Highway Patrol to Enforce Emerald Ash Borer Quarantines"
Jendek E. Studies in the East Palaearctic species of the genus Agrilus Dahl, 1823 (Coleptera: Buprestidae) Part Ⅰ. Entomological Problems, 1994, 25(1): 9～25
Lee H J. Ovipositional mechanism of an ecto-parasitoid, Goniozus legneri Gordh (Hymenoptera: Bethylidae). Chinese Journal of Entomology, 1992, 12(3): 193～199
Liu H P, Bauer L S, Gao R T, et al. Exploratory survey for the emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae), and its natural enemies in China. The Great Lakes Entomologist, 2003, 36(3&4): 191～204
Lloyd M. Mean crowding. Journal of Animal Ecology, 1967, 36: 1～30
Loerch C R, Cameron E A. Determination of larval instars of the bronze birch borer, Agrilus anxius (Coleoptera: Buprestidae). Annals of the Entomological Society of America, 1983, 76: 948～952
Rivero A. The relationship between host selection behaviour and offspring fitness in a koinobiont parasitoid. Ecological Entomology, 2000, 25: 467～472
Roland J, Taylor P D. Insect parasitoid species respond to forest structure at different spatial scales. Nature, 1997, 386: 710～713
Southwood T R E. Ecological methods (2nd Edition). London: Chapman and Hall, 1978
Wang K W, Jiang J X, You L S, et al. Preference of Microplitis sp. to host age. Journal of Hunan Agricultural University (Natural Sciences), 2001, 27(5): 367～369
Waters W E. A quantitative measure of aggregation in insects. Journal of Economic Entomology, 1959, 52: 1180～1184
Wei X, Reardon D, Wu Y, et al. Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in China: a review and distribution survey. 昆虫学报,2004,47(5):679～685
Yang Z Q, Strazanac J, Marsh P, et al. A new important braconid species parasitizing larva of emerald ash borer from China. Annals of the Entomological Society of America, 2005, 98(1): (in press)