螟黄足盘绒茧蜂复合群的生物系统学研究（膜翅目：茧蜂科：小腹茧蜂亚科）
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摘要
螟黄足盘绒茧蜂复合群由五个种所组成，即螟黄足盘绒茧蜂(Cotesia flavipes)，印度—澳洲地区；大螟盘绒茧蜂(C.sesamiae)，非洲；二化螟盘绒茧蜂(C.chilonis)，日本、中国；芦螟盘绒茧蜂(C.chiloluteelli)，中国；汉寿盘绒茧蜂(C.hanshouensis)，中国。该复合群都是禾本科主要经济作物鳞翅目蛀茎害虫幼虫的聚集性内寄生蜂，在生物防治上具有重要意义。
本文对螟黄足盘绒茧蜂复合群的分类沿革、研究历史、形态特性、生物学特性及其在古典生物防治计划中的应用进行了简要而系统的介绍。特别是对其形态学特征，在前人研究的基础上，对该复合群的雌雄外生殖器进行了深入的研究和比较，并选择了螟黄足盘绒茧蜂复合群5个种和外群侧沟茧蜂Microplitis及荻茧蜂Diolcogaster的25个性状演变系列或复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂C.kariyai、粉蝶盘绒茧蜂C.glomerate的24个性状演变系列，通过外群比较法对其进行性状极化，得到两个不同的性状数据矩阵，然后利用Hennig86支序分析软件，按照支序分析的一般原则和方法对性状数据矩阵进行处理，以此来明确螟黄足盘绒茧蜂复合群内各分类单元之间的亲缘关系，并提出了该复合群系统发育关系的假说。研究结果表明：螟黄足盘绒茧蜂复合群是一个单系群；大螟盘绒茧蜂和二化螟盘绒茧蜂亲缘关系靠近，具有共同的假想祖先；芦螟盘绒茧蜂和汉寿盘绒茧蜂亲缘关系靠近，也具有共同的假想祖先；而螟黄足盘绒茧蜂相对独立。在该复合群5个种之间，螟黄足盘绒茧蜂是比较原始的类群，大螟盘绒茧蜂和二化螟盘绒茧蜂次之，芦螟盘绒茧蜂和汉寿盘绒茧蜂是较为进化的类群。
The Cotesia flavipes complex is currently considered to include five species: Cotesia flavipes (Cammeron) which occurs in the Indo-Australian region, C.sesamiae (Cammeron) in Africa, C.chilonis [Munakata] in Japan and China, C.chiloluteelli (You et Xiong) in China and C.hanshouensis (You et Xiong) in China. The species of the complex are all gregarious endoparasitoids of lepidopteran stemborers of gramineous plants, and they are very important in classical biological control programs against various cereal stemborers.
In the present paper, historical review, morphology, biology and applications of the Cotesia flavipes complex in classical biological control programmes are introduced briefly and systematically. Based on morphological studies (include male genitalia), 25 character transformation series of 5 species of the complex (as ingroup) and 1 species of Microplitis and Dilolcogaster (as outgroup) respectively or 24 character transformation series of 5 species of this complex (as ingroup) and 3 species of the same genera (as outgroup) were selected. Through analysis of character polarity using the method of outgroup analysis, two character data matrixes were obtained. Then through analysis of Hennig86 software based on the method of cladistics, two consensus cladograms were constructed at last. The results of the cladograms indicated that the Cotesia flavipes complex is a monophyletic group and that C.sesamiae and C.chilonis are more closely related, C.chiloluteelli and C.hanshouensis are more closely related, too, C.flavipes is a species to be independed upon above 4 species. It was also founded that among the 5 species of Cotesia flavipes complex, C.flavipes is primitive, C.sesamiae and C.chilonis are primitive secondly, C.chiloluteelli and C.hanshouensis are the relatively evolutive species. In conclution, wo do whatever the relative genus as outgroup or the relative species as outgoup, the results can correctly reflect the relationships among the species of Cotesia flavipes complex.
本文对螟黄足盘绒茧蜂复合群的分类沿革、研究历史、形态特性、生物学特性及其在古典生物防治计划中的应用进行了简要而系统的介绍。特别是对其形态学特征，在前人研究的基础上，对该复合群的雌雄外生殖器进行了深入的研究和比较，并选择了螟黄足盘绒茧蜂复合群5个种和外群侧沟茧蜂Microplitis及荻茧蜂Diolcogaster的25个性状演变系列或复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂C.kariyai、粉蝶盘绒茧蜂C.glomerate的24个性状演变系列，通过外群比较法对其进行性状极化，得到两个不同的性状数据矩阵，然后利用Hennig86支序分析软件，按照支序分析的一般原则和方法对性状数据矩阵进行处理，以此来明确螟黄足盘绒茧蜂复合群内各分类单元之间的亲缘关系，并提出了该复合群系统发育关系的假说。研究结果表明：螟黄足盘绒茧蜂复合群是一个单系群；大螟盘绒茧蜂和二化螟盘绒茧蜂亲缘关系靠近，具有共同的假想祖先；芦螟盘绒茧蜂和汉寿盘绒茧蜂亲缘关系靠近，也具有共同的假想祖先；而螟黄足盘绒茧蜂相对独立。在该复合群5个种之间，螟黄足盘绒茧蜂是比较原始的类群，大螟盘绒茧蜂和二化螟盘绒茧蜂次之，芦螟盘绒茧蜂和汉寿盘绒茧蜂是较为进化的类群。
The Cotesia flavipes complex is currently considered to include five species: Cotesia flavipes (Cammeron) which occurs in the Indo-Australian region, C.sesamiae (Cammeron) in Africa, C.chilonis [Munakata] in Japan and China, C.chiloluteelli (You et Xiong) in China and C.hanshouensis (You et Xiong) in China. The species of the complex are all gregarious endoparasitoids of lepidopteran stemborers of gramineous plants, and they are very important in classical biological control programs against various cereal stemborers.
In the present paper, historical review, morphology, biology and applications of the Cotesia flavipes complex in classical biological control programmes are introduced briefly and systematically. Based on morphological studies (include male genitalia), 25 character transformation series of 5 species of the complex (as ingroup) and 1 species of Microplitis and Dilolcogaster (as outgroup) respectively or 24 character transformation series of 5 species of this complex (as ingroup) and 3 species of the same genera (as outgroup) were selected. Through analysis of character polarity using the method of outgroup analysis, two character data matrixes were obtained. Then through analysis of Hennig86 software based on the method of cladistics, two consensus cladograms were constructed at last. The results of the cladograms indicated that the Cotesia flavipes complex is a monophyletic group and that C.sesamiae and C.chilonis are more closely related, C.chiloluteelli and C.hanshouensis are more closely related, too, C.flavipes is a species to be independed upon above 4 species. It was also founded that among the 5 species of Cotesia flavipes complex, C.flavipes is primitive, C.sesamiae and C.chilonis are primitive secondly, C.chiloluteelli and C.hanshouensis are the relatively evolutive species. In conclution, wo do whatever the relative genus as outgroup or the relative species as outgoup, the results can correctly reflect the relationships among the species of Cotesia flavipes complex.
引文
1 Alam M.M., 1972. Technical Institute of Biological Control. 15: 1-10.
2 Anon.,1980. planalsUcar, Programa nacional de melboramento da cana—de-acucar. Relatorio anual 1980. Instituto do Acucar e do Alcool. pp. 86-87.
3 Betbeder-Matibet M and Malinge P. 1967..Un succes de la lutte biologique: controle de Proceras sacchariphagus Boj. Borer ponctue de la canne a sucre a Madagascar par un parasite introduit: Apanteles flavipes Cam. Agronomie Tropical 22, 1196-1220.
4 Bordat D et al, 1977. Agrononis Tropicate 32(4): 391-399.
5 Bordat D. 1983. Mise au point de l'elevage de masse d'Apanteles chilonis Matsumura, 1912 et d'ApanteJes flavipes (Cameron, 1891) (Hymenopteres Braconidae) sur trois lepidopteres pyralidae foreurs des graminees (Chilo zacconius Bleszynski, 1970; Chilo partellus (Swinhoe, 1884) et Diatraea saccharalis (Fabricius, 1794)) dans un objectif de lutte biologique. These. Universite des Sciences et Techniques du Langeudoc, Montpellier, France. 184 p.
6 Breniere J. 1982. Agronomic Tropical [J], 37(2):203-206.
7 Chang Y D 1978. Korean Joural of Plant Protection [J], 17(1):65-69.
8 Chu Y I 1979. Phytopathologist and Entomologist [J], 6:38-43.
9 Cracraft, J.,1989. Speciation and its ontology: the empirical consequences of alternative species concepts for understanding patterns and processes of differentiation, pp. 28-59 in Otte, D. & Endler, J.A. (eds). Speciation and its consequences.
10 FAO.1995. FAO yearbook 1994 production. Food and agricultural organization of the United Nations. Rome.
11 Fuchs, T. W., F. R. Huffman & J. W. Smith, Jr. 1979. Introduction and establishment of Apanteles flavipes (Hym.: Braconidae) on Diatraea saccharalis (Lep.: Pyralidae) in Texas. Entomophaga 24: 109-114.
12 Galichet P. F., 1979. Entomophaga 24(2): 119-130.
13 Holling C.S., 1973. Indian Joural of Entomology. 35(4): 281-284.
14 Huang, D W, 1996 An Introduction to Cladistics. China Agriculture Press, Beijing. [黄大卫，1996，支序系统学概论，农业出版社，北京，189页]
15 Kazuo I.1986. Bulletin of Fukui Agri. Exp. Sta. [J],23:19-25.
16 Kimani S. W. and Overholt W. A. 1995. Biosystematics of the Cotesia flavipes complex (Hymenoptera: Braconidae): interspecific hybridization, sex pheromone and mating behaviour studies. Bulletin of Entomological Research 85: 379-386.
17 Kimani-Njogu, S. W., W. A. Overholt, J. Woolley and A. Walker. 1997. Biosystematics of the Cotesia flavipes species complex (Hymenoptera: Braconidae)-morphometrics of selected allopatric populations.
Bulletin of Entomological Research 87: 61-66.
18 Maddison, W.P.,M.J. Donoghue and D.R. Maddison. 1984. Outgroup analysis and parsimony. Syst. Zool. 33: 83-103.
19 Mason, W.R.M., 1981. The polyphyletis nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microfastrinae, Mem. Ent. Soc. Can. 115: 1-147.
20 Mbapila J. 1994. Laboratory life tables of Cotesia spp. reared at difference temperatures on Chilo partellus, pg. 31 In 1994 ICIPE Annual Report. ICIPE Science Press. 199 p.
21 Meng Kaibaryer, 2002. Phygenetic Analysis Using Both PAUP and MacClade. Chinese Journal of Zoology. 37 (6): 59-62. [孟凯巴依尔等，2002，联合使用PAUP和MacClade程序进行支序分析，37(6)：59—62．]
22 Mohyuddin A. I.,Inayatullah C., King E.G. 1981. Host selecton and strain occurrence in Apanteles flavipes (Cameron) (Hymenoptera: Braconidae) and its bearing on biological control of graminaeous stem-borers (Lepidoptera: Pyralidae. Bulletin of Entomological Reseach, 71: 575-581.
23 Nagaraja, H. 1971. Morphological difference between Apanteles chilonis [Munakata] and A. sesamiae Cameron (Hym.: Braconidae), parasites on Graminaceous moth borers. Technical Bulletin, Commonwealth Institute Biological Control, 14:59-61.
24 Nagarkatti & Fazaluddin, 1973. Biosystematic studies on Trichogramma species (Hymenoptera: Trichogramatidae) Ⅱ. Experimental hybridistion between some Trichogramma app. From the New Would. Systematic Zoology 22:103-117.
25 Ngi-Song A. J. 1995. Parasitization of selected African stemborers by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae), with emphasis on host selection and host suitability. PhD Thesis. University of Ghana, Legon. 198 p.
26 Ngi-Song A. J., Overholt W. A., Njagi P. O. N., Dicke M., Ayertey J. A. and Lwande W. 1996. Volatile infochemicals used in host and host habitat location by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae), larval parasitoids of stemborers on graminae. Journal of Chemical Ecology 22(2): 307-323.
27 Ngi-Song, A.J., Kimani-Njogu, S. and Overholt, W. A., 2001. Multiple parasitism by Cotesia sesamiae and Cotesia flavipes (Hymenoptera: Braconidae) on Busseola fusca (Lepidoptera: Noctuidae). Biocontrol Science and Technology. 11(3): 381-390.
28 Nixon, O.E..J. 1965. A reclassification of the tribe Microgasterini (Hymenoptera: Braconidae). Bull. Brit. Mus. Nat. Hist. (Ent 2(1): 284
29 Nixon, O.E.J. 1973. A revision of the north-western European species od the vitripennis, pallipes, octonrius, triangulator, fraternus, formisus, parasitellae, metacarpalits and circumscriptus -groups of
Apanteles Foerster (Hymenoptera: Braconidae). Bull. Ent. Res. 63: 169-230.
30 Oatman et al., 1970. Reproductive differentiation of Trichogramma pretiosum, T. semifumarum, T. minutum and T. evanescens, with notes on the geographic distribution of T. pretiosum in the southwestern United States and in Mexico (Hymenoptera: Trichogrammatidae). Annals of the Entomological Society of America 61:956-959.
31 Overholt W. A. 1993. Release of beneficial insects in Kenya. Discovery and Innovation 5(3): 199-200.
32 Overholt W. A., Ngi-Song A. J., Kimani S. W, Mbapila J., Lammers P. M. and. Kioko E. (1994a) Ecological considerations of the introduction of Cotesia flavipes Cameron (Hymenoptera: Braconidae) for biological control of Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae), in Africa. Biocontrol news and information 15(2): 19N-24N.
33 Overholt, W. A. and Ngi-Song, A. J. 1996. An Ecological Approach toBiological Control of Gramineous temborers in Africa:The Introduction and Establishment of Cotesia flavipes Cameron (Hymenoptera: Braconidae) National IPM Network of Minnesota University.
34 Pinto et al., 1991. Variation in reproductive comparibility in Trichogramma and its taxonomic significance (Hymenoptera: Trichogrammatidae). Annals of the Entomological Society of America 84: 37-46.
35 Polaszek, A. and A. K. Walker. 1991. The Cotesia: flavipes species-complex: parasitoids of cereal stem borers in the tropics. Redia 74: 335-341.
36 Potting R P J, 1995. Hunting for Hiding Hosts: The Behavioral Ecology of the Stenborer Parasitoid Cotesia flavipes. The Netherlands: Wageningen Agricultural University.
37 Potting R. P. J., Vet L. E. M. and Dicke M. 1995. Host microhabitat location by stem-borer parasitoid Cotesia flavipes: the role of herbivore volatiles and locally and systematically induced plant volatiles. Journal of Chemical Ecology 21(5): 525-539.
38 Qi Bao-ying 2001 A Cladistic Analysis on the Genera of Tinginae (Hym.: Heteroptera: Tingidae) from Northen China Zoological Research 22(1):1-8. [齐宝瑛，2001，中国北方网蝽亚科的支序分析(半翅目：异翅亚目：网蝽科)，动物学研究，22(1)：1—8．]
39 Rao, V. P., Nagaraja, H. 1967. Validity of specific states if A. chilonis [Muna- kata] technical Bulletin, Commonwealth Institute Biological Control 9:45-52.
40 Sallam and Overholt, 2002. Intraspecific and Interspecific Competition Between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), Gregarious Larval Endoparasitoids of
Lepidopteran Stemborers. Biocontrol Science and Technology. 12(4): 493-506.
41 Sallam, Mohamed N. and Overholt, W.A., and Kairu, E. and Guofa, Z. 2001. Will Cotesia flavipes drive its African homelogue, Cotesia sesamiae to extinction. Insect Science and its Application 21(4): 397-302.
42 Sigwalt, B., Pointel, J.G., 1980. statut specifique et separation des Apanteles du sous-group flavipes (Hym.: Braconidae) utilisees en lutte biologique. Annales de la Societe Entomologique de France, 16: 109-128.
43 Smith, P. T. and S. Kambhampati. 1999. Status of the Cotesia flavipes species complex (Braconidae: Microgastrinae) based on mitochondrial 16S rRNA and NADH1 dehydrogenase gene sequence. Journal of the Kansas Entomological Society 72: 306-314.
44 Varina G C 1974. Indian Journal of Entomology [J],36(1):34-37.
45 Walker, A K, 1994. Species of Microgastrinae (Hymenoptera: Braconidae) parasitizing Lepidopterous cereal stem borers in Africa. Bull. Entomol. Res. 84, 421-434.
46 Watanabe C., 1965. Notes on the Apanteles flavipe Complex (Hymenoptera, Braconidae) Mushi vol. 38 pars 13:111-116.
47 Williams, J.G.K.,Kbelik, A.R.,et al,1990. DNA polymorphism amplified by arbitrary primers are useful as genetic markers. Nucl. Acids Res. 18(22): 6531-6535.
48 You, L.S. and Xiong, S.L.,1983. Two new species of Apanteles Foerster from China. (Hymenoptera: Braconidae: Microgastrinae) Entomotaxonomia 5(3): 225-229. [游兰韶，熊漱琳，1983，中国绒茧蜂属两新种记述，昆虫分类学报，5(3)：225—229．]
49 You, L.S., Xiong, S.L. and Wang, Z.D.,1985. A new species of Apanteles Foerster from China (Hymenoptera:Braconidae:Microgasterinae) Acta Zootaxo. Sinica 10(4):421-423. [游兰韶，熊漱琳，王宗典，1985，中国绒茧蜂属一新种(膜翅目：茧蜂科)，动物分类学报，10(4)：421-423．]
50 何俊华，1979，浙江省水稻害虫天敌图册[M]，杭州：浙江科技出版社。
51 祁彪，1983，农垦生防[J]，10：73—74．
52 郑乐怡，1987，动物分类原理与方法，高等教育出版社，1—185．
53 杭三保，陈培红等，1989，二化螟绒茧蜂生态学特性研究，江苏农学院学报，10(1)：23—26．
54 杭三保，1989，二化螟绒茧蜂对寄主二化螟幼虫取食量和生长发育的影响，江苏农学院学报，10(3)：33—36．
55 杭杉葆，1989，二化螟绒茧蜂生物学特性研究，生物防治通报，1(1)：16—18．
56 孙修勤等，1993，酵母菌支序分类研究，黄渤海海洋，11(2)：28—32．
57 杨宗岱，1993，支序分类在海草分类划分中的应用，黄渤海海洋，11(2)：33—37．
58 徐志强，1993，蝽总科科间支序分析及系统发育，北京农业大学学报，19(2)：49—53．
59 李正新，1994，支序分类学简介Ⅰ，海洋科学(?)：26—28．
60 杭三保，1994，一种室内累代饲养二化螟绒茧蜂的方法，昆虫学报，37(4)：504—505．
61 戴灼华，1994，Drosophila auraria复合种的进化遗传学研究——支序分析与数值分析(上)，21(5)：362—372．
62 李文凤，1995，螟黄足绒茧蜂的初步观察，昆虫天敌，17(1)：7—8．
63 鲁亮等，1995，RAPD技术的特点及其在昆虫分类中的应用，昆虫学报，38(1)：117-121．
64 蒋明星，1999，不同生境中水稻二化螟的自然寄生情况，中国生物防治[J]，15(4)：145—149．
65 刘宇晖，2000，中国库蚊属9个亚属的支序分析，寄生虫与医学昆虫学报，7(4)：218—224．
66 金道超，2000，水螨群总科阶元系统发育的支序分析，昆虫学报，43(3)：310—317．
67 郭鹏等，2000，亚洲腹亚科蛇属间系统发生支序分析，生命科学研究，4(3)：262—266．
68 马雅军等，2001，我国中华按蚊群体分子遗传多态研究，昆虫学报，44(1)：33-39．
69 左正宏，桂慕燕，王学民，陈元霖，2001，RAPD分析在绢丝昆虫亲缘关系研究中的应用Ⅰ．蓖麻蚕品种间的遗传差异，遗传，23(2)：128—130．
70 江世宏，2001，中国叩甲科昆虫12亚科的支序分类研究，华中农业大学学报，20(4)：229—332．
71 李莉，2001，我国北方麦区麦长管蚜种群的RAPD分析，植物保护学报，28(1)：89—90．
72 桂慕燕，左正宏，陈元霖，2001，五种绢丝昆虫随机扩增多态性DNA分析，遗传，23(1)：25—28．
73 王文凯，2002，天牛总科高级分类阶元系统发育关系的研究，河南农业大学学报，36(1)：11—14．
74 刘彦群，鲁成，向仲怀，2002，中国柞蚕DNA多态性的RAPD分析，蚕业科学，28(4)：283—288．
75 陈华才，娄永根等，2002，二化螟绒茧蜂对二化螟及其寄主植物挥发物的趋性反应，昆虫学报，45(5)：617—622．
76 陈华才等，2002，二化螟绒茧蜂研究与应用概况，中国生物防治，18(2)：90—93．
77 庞乾林 2002首届国际水稻大会在北京召开——江泽民出席并作重要讲话 中国稻米 (6)：14—15．
78 程道军，鲁成，周泽扬，向仲怀，2002，几种绢丝昆虫遗传多样性的RAPD研究 蚕业科学，28(4)：277—282．
2 Anon.,1980. planalsUcar, Programa nacional de melboramento da cana—de-acucar. Relatorio anual 1980. Instituto do Acucar e do Alcool. pp. 86-87.
3 Betbeder-Matibet M and Malinge P. 1967..Un succes de la lutte biologique: controle de Proceras sacchariphagus Boj. Borer ponctue de la canne a sucre a Madagascar par un parasite introduit: Apanteles flavipes Cam. Agronomie Tropical 22, 1196-1220.
4 Bordat D et al, 1977. Agrononis Tropicate 32(4): 391-399.
5 Bordat D. 1983. Mise au point de l'elevage de masse d'Apanteles chilonis Matsumura, 1912 et d'ApanteJes flavipes (Cameron, 1891) (Hymenopteres Braconidae) sur trois lepidopteres pyralidae foreurs des graminees (Chilo zacconius Bleszynski, 1970; Chilo partellus (Swinhoe, 1884) et Diatraea saccharalis (Fabricius, 1794)) dans un objectif de lutte biologique. These. Universite des Sciences et Techniques du Langeudoc, Montpellier, France. 184 p.
6 Breniere J. 1982. Agronomic Tropical [J], 37(2):203-206.
7 Chang Y D 1978. Korean Joural of Plant Protection [J], 17(1):65-69.
8 Chu Y I 1979. Phytopathologist and Entomologist [J], 6:38-43.
9 Cracraft, J.,1989. Speciation and its ontology: the empirical consequences of alternative species concepts for understanding patterns and processes of differentiation, pp. 28-59 in Otte, D. & Endler, J.A. (eds). Speciation and its consequences.
10 FAO.1995. FAO yearbook 1994 production. Food and agricultural organization of the United Nations. Rome.
11 Fuchs, T. W., F. R. Huffman & J. W. Smith, Jr. 1979. Introduction and establishment of Apanteles flavipes (Hym.: Braconidae) on Diatraea saccharalis (Lep.: Pyralidae) in Texas. Entomophaga 24: 109-114.
12 Galichet P. F., 1979. Entomophaga 24(2): 119-130.
13 Holling C.S., 1973. Indian Joural of Entomology. 35(4): 281-284.
14 Huang, D W, 1996 An Introduction to Cladistics. China Agriculture Press, Beijing. [黄大卫，1996，支序系统学概论，农业出版社，北京，189页]
15 Kazuo I.1986. Bulletin of Fukui Agri. Exp. Sta. [J],23:19-25.
16 Kimani S. W. and Overholt W. A. 1995. Biosystematics of the Cotesia flavipes complex (Hymenoptera: Braconidae): interspecific hybridization, sex pheromone and mating behaviour studies. Bulletin of Entomological Research 85: 379-386.
17 Kimani-Njogu, S. W., W. A. Overholt, J. Woolley and A. Walker. 1997. Biosystematics of the Cotesia flavipes species complex (Hymenoptera: Braconidae)-morphometrics of selected allopatric populations.
Bulletin of Entomological Research 87: 61-66.
18 Maddison, W.P.,M.J. Donoghue and D.R. Maddison. 1984. Outgroup analysis and parsimony. Syst. Zool. 33: 83-103.
19 Mason, W.R.M., 1981. The polyphyletis nature of Apanteles Foerster (Hymenoptera: Braconidae): a phylogeny and reclassification of Microfastrinae, Mem. Ent. Soc. Can. 115: 1-147.
20 Mbapila J. 1994. Laboratory life tables of Cotesia spp. reared at difference temperatures on Chilo partellus, pg. 31 In 1994 ICIPE Annual Report. ICIPE Science Press. 199 p.
21 Meng Kaibaryer, 2002. Phygenetic Analysis Using Both PAUP and MacClade. Chinese Journal of Zoology. 37 (6): 59-62. [孟凯巴依尔等，2002，联合使用PAUP和MacClade程序进行支序分析，37(6)：59—62．]
22 Mohyuddin A. I.,Inayatullah C., King E.G. 1981. Host selecton and strain occurrence in Apanteles flavipes (Cameron) (Hymenoptera: Braconidae) and its bearing on biological control of graminaeous stem-borers (Lepidoptera: Pyralidae. Bulletin of Entomological Reseach, 71: 575-581.
23 Nagaraja, H. 1971. Morphological difference between Apanteles chilonis [Munakata] and A. sesamiae Cameron (Hym.: Braconidae), parasites on Graminaceous moth borers. Technical Bulletin, Commonwealth Institute Biological Control, 14:59-61.
24 Nagarkatti & Fazaluddin, 1973. Biosystematic studies on Trichogramma species (Hymenoptera: Trichogramatidae) Ⅱ. Experimental hybridistion between some Trichogramma app. From the New Would. Systematic Zoology 22:103-117.
25 Ngi-Song A. J. 1995. Parasitization of selected African stemborers by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae), with emphasis on host selection and host suitability. PhD Thesis. University of Ghana, Legon. 198 p.
26 Ngi-Song A. J., Overholt W. A., Njagi P. O. N., Dicke M., Ayertey J. A. and Lwande W. 1996. Volatile infochemicals used in host and host habitat location by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae), larval parasitoids of stemborers on graminae. Journal of Chemical Ecology 22(2): 307-323.
27 Ngi-Song, A.J., Kimani-Njogu, S. and Overholt, W. A., 2001. Multiple parasitism by Cotesia sesamiae and Cotesia flavipes (Hymenoptera: Braconidae) on Busseola fusca (Lepidoptera: Noctuidae). Biocontrol Science and Technology. 11(3): 381-390.
28 Nixon, O.E..J. 1965. A reclassification of the tribe Microgasterini (Hymenoptera: Braconidae). Bull. Brit. Mus. Nat. Hist. (Ent 2(1): 284
29 Nixon, O.E.J. 1973. A revision of the north-western European species od the vitripennis, pallipes, octonrius, triangulator, fraternus, formisus, parasitellae, metacarpalits and circumscriptus -groups of
Apanteles Foerster (Hymenoptera: Braconidae). Bull. Ent. Res. 63: 169-230.
30 Oatman et al., 1970. Reproductive differentiation of Trichogramma pretiosum, T. semifumarum, T. minutum and T. evanescens, with notes on the geographic distribution of T. pretiosum in the southwestern United States and in Mexico (Hymenoptera: Trichogrammatidae). Annals of the Entomological Society of America 61:956-959.
31 Overholt W. A. 1993. Release of beneficial insects in Kenya. Discovery and Innovation 5(3): 199-200.
32 Overholt W. A., Ngi-Song A. J., Kimani S. W, Mbapila J., Lammers P. M. and. Kioko E. (1994a) Ecological considerations of the introduction of Cotesia flavipes Cameron (Hymenoptera: Braconidae) for biological control of Chilo partellus (Swinhoe) (Lepidoptera: Pyralidae), in Africa. Biocontrol news and information 15(2): 19N-24N.
33 Overholt, W. A. and Ngi-Song, A. J. 1996. An Ecological Approach toBiological Control of Gramineous temborers in Africa:The Introduction and Establishment of Cotesia flavipes Cameron (Hymenoptera: Braconidae) National IPM Network of Minnesota University.
34 Pinto et al., 1991. Variation in reproductive comparibility in Trichogramma and its taxonomic significance (Hymenoptera: Trichogrammatidae). Annals of the Entomological Society of America 84: 37-46.
35 Polaszek, A. and A. K. Walker. 1991. The Cotesia: flavipes species-complex: parasitoids of cereal stem borers in the tropics. Redia 74: 335-341.
36 Potting R P J, 1995. Hunting for Hiding Hosts: The Behavioral Ecology of the Stenborer Parasitoid Cotesia flavipes. The Netherlands: Wageningen Agricultural University.
37 Potting R. P. J., Vet L. E. M. and Dicke M. 1995. Host microhabitat location by stem-borer parasitoid Cotesia flavipes: the role of herbivore volatiles and locally and systematically induced plant volatiles. Journal of Chemical Ecology 21(5): 525-539.
38 Qi Bao-ying 2001 A Cladistic Analysis on the Genera of Tinginae (Hym.: Heteroptera: Tingidae) from Northen China Zoological Research 22(1):1-8. [齐宝瑛，2001，中国北方网蝽亚科的支序分析(半翅目：异翅亚目：网蝽科)，动物学研究，22(1)：1—8．]
39 Rao, V. P., Nagaraja, H. 1967. Validity of specific states if A. chilonis [Muna- kata] technical Bulletin, Commonwealth Institute Biological Control 9:45-52.
40 Sallam and Overholt, 2002. Intraspecific and Interspecific Competition Between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), Gregarious Larval Endoparasitoids of
Lepidopteran Stemborers. Biocontrol Science and Technology. 12(4): 493-506.
41 Sallam, Mohamed N. and Overholt, W.A., and Kairu, E. and Guofa, Z. 2001. Will Cotesia flavipes drive its African homelogue, Cotesia sesamiae to extinction. Insect Science and its Application 21(4): 397-302.
42 Sigwalt, B., Pointel, J.G., 1980. statut specifique et separation des Apanteles du sous-group flavipes (Hym.: Braconidae) utilisees en lutte biologique. Annales de la Societe Entomologique de France, 16: 109-128.
43 Smith, P. T. and S. Kambhampati. 1999. Status of the Cotesia flavipes species complex (Braconidae: Microgastrinae) based on mitochondrial 16S rRNA and NADH1 dehydrogenase gene sequence. Journal of the Kansas Entomological Society 72: 306-314.
44 Varina G C 1974. Indian Journal of Entomology [J],36(1):34-37.
45 Walker, A K, 1994. Species of Microgastrinae (Hymenoptera: Braconidae) parasitizing Lepidopterous cereal stem borers in Africa. Bull. Entomol. Res. 84, 421-434.
46 Watanabe C., 1965. Notes on the Apanteles flavipe Complex (Hymenoptera, Braconidae) Mushi vol. 38 pars 13:111-116.
47 Williams, J.G.K.,Kbelik, A.R.,et al,1990. DNA polymorphism amplified by arbitrary primers are useful as genetic markers. Nucl. Acids Res. 18(22): 6531-6535.
48 You, L.S. and Xiong, S.L.,1983. Two new species of Apanteles Foerster from China. (Hymenoptera: Braconidae: Microgastrinae) Entomotaxonomia 5(3): 225-229. [游兰韶，熊漱琳，1983，中国绒茧蜂属两新种记述，昆虫分类学报，5(3)：225—229．]
49 You, L.S., Xiong, S.L. and Wang, Z.D.,1985. A new species of Apanteles Foerster from China (Hymenoptera:Braconidae:Microgasterinae) Acta Zootaxo. Sinica 10(4):421-423. [游兰韶，熊漱琳，王宗典，1985，中国绒茧蜂属一新种(膜翅目：茧蜂科)，动物分类学报，10(4)：421-423．]
50 何俊华，1979，浙江省水稻害虫天敌图册[M]，杭州：浙江科技出版社。
51 祁彪，1983，农垦生防[J]，10：73—74．
52 郑乐怡，1987，动物分类原理与方法，高等教育出版社，1—185．
53 杭三保，陈培红等，1989，二化螟绒茧蜂生态学特性研究，江苏农学院学报，10(1)：23—26．
54 杭三保，1989，二化螟绒茧蜂对寄主二化螟幼虫取食量和生长发育的影响，江苏农学院学报，10(3)：33—36．
55 杭杉葆，1989，二化螟绒茧蜂生物学特性研究，生物防治通报，1(1)：16—18．
56 孙修勤等，1993，酵母菌支序分类研究，黄渤海海洋，11(2)：28—32．
57 杨宗岱，1993，支序分类在海草分类划分中的应用，黄渤海海洋，11(2)：33—37．
58 徐志强，1993，蝽总科科间支序分析及系统发育，北京农业大学学报，19(2)：49—53．
59 李正新，1994，支序分类学简介Ⅰ，海洋科学(?)：26—28．
60 杭三保，1994，一种室内累代饲养二化螟绒茧蜂的方法，昆虫学报，37(4)：504—505．
61 戴灼华，1994，Drosophila auraria复合种的进化遗传学研究——支序分析与数值分析(上)，21(5)：362—372．
62 李文凤，1995，螟黄足绒茧蜂的初步观察，昆虫天敌，17(1)：7—8．
63 鲁亮等，1995，RAPD技术的特点及其在昆虫分类中的应用，昆虫学报，38(1)：117-121．
64 蒋明星，1999，不同生境中水稻二化螟的自然寄生情况，中国生物防治[J]，15(4)：145—149．
65 刘宇晖，2000，中国库蚊属9个亚属的支序分析，寄生虫与医学昆虫学报，7(4)：218—224．
66 金道超，2000，水螨群总科阶元系统发育的支序分析，昆虫学报，43(3)：310—317．
67 郭鹏等，2000，亚洲腹亚科蛇属间系统发生支序分析，生命科学研究，4(3)：262—266．
68 马雅军等，2001，我国中华按蚊群体分子遗传多态研究，昆虫学报，44(1)：33-39．
69 左正宏，桂慕燕，王学民，陈元霖，2001，RAPD分析在绢丝昆虫亲缘关系研究中的应用Ⅰ．蓖麻蚕品种间的遗传差异，遗传，23(2)：128—130．
70 江世宏，2001，中国叩甲科昆虫12亚科的支序分类研究，华中农业大学学报，20(4)：229—332．
71 李莉，2001，我国北方麦区麦长管蚜种群的RAPD分析，植物保护学报，28(1)：89—90．
72 桂慕燕，左正宏，陈元霖，2001，五种绢丝昆虫随机扩增多态性DNA分析，遗传，23(1)：25—28．
73 王文凯，2002，天牛总科高级分类阶元系统发育关系的研究，河南农业大学学报，36(1)：11—14．
74 刘彦群，鲁成，向仲怀，2002，中国柞蚕DNA多态性的RAPD分析，蚕业科学，28(4)：283—288．
75 陈华才，娄永根等，2002，二化螟绒茧蜂对二化螟及其寄主植物挥发物的趋性反应，昆虫学报，45(5)：617—622．
76 陈华才等，2002，二化螟绒茧蜂研究与应用概况，中国生物防治，18(2)：90—93．
77 庞乾林 2002首届国际水稻大会在北京召开——江泽民出席并作重要讲话 中国稻米 (6)：14—15．
78 程道军，鲁成，周泽扬，向仲怀，2002，几种绢丝昆虫遗传多样性的RAPD研究 蚕业科学，28(4)：277—282．