摘要
目前,国内蝗虫绿僵菌疾病流行病学上的研究还较少。本研究在内蒙古锡林郭勒盟多伦县进行了蝗虫绿僵菌疾病流行学试验,对蝗虫绿僵菌疾病的扩散、传播作了初步探讨。
试验地域共发现蝗虫18种,分属网翅蝗亚科、斑翅蝗亚科、斑腿蝗亚科、槌角蝗亚科、剑角蝗亚科、癞蝗亚科、蚱科共7个亚科。对不同植被类型中蝗虫群落多样性特征进行了分析,结果表明:毛足棒角蝗为早发优势种;白纹雏蝗为中期优势种;小翅雏蝗为晚发优势种。沙漠化草场蝗虫群落的Shannon-Wiener指数小于草甸草原向典型草原过渡类型草场和典型草原草场;沙漠化导致蝗虫群落多样性指数降低,沙化草场上大体型癞蝗科和斑翅蝗科所属种的数量较其在非退化草场上高。
采用模型N_t=α×t~B×e~(-c·t)对试验当年当地蝗虫的发生动态进行了拟合,结果显示:主要蝗虫种群数量都表现出季节性单峰波动;盛发期各不相同,致使混合种群高虫口密度持续时间长。根据各蝗虫种的盛发期前后次序,确定最佳防治时期应该选在6月20号以前。
采用罩笼饲养试验对个体间疾病的传播进行了模拟,结果显示:野外条件下病虫可以将疾病传播给健虫,在病、健虫混合比例分别为10:50、20:40、30:30、40:20时,同种蝗虫间疾病传播几率可达24.6%、31.0%、39.0%和52.0%;早期发生种对后期种的传播几率可达15.6%、23.5%、32.7%和40.0%;感染率及传播几率随病虫比例增大而提高,种间习性差异可能导致早发种对晚发种间传播率的降低。
对圆形施药区外不同距离、方向处绿僵菌疾病田间扩散情况进行了调查,结果显示:疾病可以扩散到施药区外400 m,药后40 d疾病感染率八个方向均值可达8.95%。药区外疾病感染率随时间的推移逐渐上升;不同方向疾病感染水平不同,西北、西南方向较高,西、北、南、东北次之,东、东南方向水平最低,这可能和蝗虫的移动趋向、风向有关。施药区外短距离内疾病的扩散在对虫口有控制作用,施药区外随时间的推移校正虫口减退率逐渐上升,处理后42 d距离施药区200 m处校正虫口减退率8个方向均值可达44.05%。校正虫口减退率高低水平在不同方向上也各不相同,与疾病感染率高低方向基本一致。
在施药方式上进行了条带间隔处理试验,结果表明:蝗虫移动扩散的习性可以有效地将病原传播到未施药区域;平行距离120 m的2施药条带间区域,药后49d校正虫口减退率达88.92%,防效良好,间隔施药的方式值得在现实防治中进行尝试。
采用2种模型和多个空间分布型指标,对药后药区内残存蝗虫和僵虫的空间分布型进行了研究。结果显示:施药后随蝗虫种群密度的减小,处理区残虫的分布型不同时间上呈现聚集-随机交错变动,僵虫的空间分布型为聚集分布;采用Taylor幂法则、改进的Iwao模型分析表明,整个调查时段处理区残虫、僵虫的空间分布均为聚集分布。药后蝗虫点片状死亡,部分地片相对密度较高,残虫分布型趋向聚集;自然消除作用使僵虫密度很低,低密度下取样产生大量空样本,僵虫的分布型产生聚集假象。蝗虫染病死亡前的迁飞移动、取食交配等行为是病原扩散传播的主要方式之一。染病未死蝗虫携病原运动可以促使疾病的空间扩散,僵虫是疾病后期潜在的传播源,理论上,后期流行中病原种群的分布型将于僵虫一致。
Epidemiology of Metarrhizium anisopliae was made in the Duolun Caimu Mountain Reservation Area in Inner Mongolian.There are 18 grasshopper species belonging to 7 subfamilies including Arcypteridae,Oedipodidae,Catantopidae,Gomphoceridae,Acrididae,Pamphagidae and Tetrigidae were recorded in the experiment area.The characteristics of grasshopper population diversity in different habitat environment was analyzed and the result showed that the predominant species was Dasyhippusbarbipes(F.-W.)in the early occurrence period,Chorthippus albonemus Cheng et Tu in the middle and Chrothgppus fOIlax(zuboysky)in the late.The Shannon-Wiener index of degraded grassland was less than that of typical steppe and transitional type between meadow steppe type and typical steppe,In addition,desertification of grassland results in the decrease of grasshoppers diversity index and the amount of Oedipodidae and Pamphagidae with bigger appetite and bigger somatotype in degraded grassland was higher than that in others habitat environment.
Mathematical models of population dynamic showed the amount of main populations were fluctuating in single peak,and different full incidence period of species resulted in the longest duration of high population density,Thus,the optimum control period should be before June 22 in current year.
Results of cage experiment showed that affected grasshopper could spread disease of Metarrhizium anisopliae to health.When the proportion of affected and health grasshopper was 10:50,20:40, 30:30 and 40:20,the transmission rate between individuals within same species was 24.6%,31.0%, 39.0%and 52.0%respectively,while the transmission rate from species occurring early in the season to those occurring late was 15.6%,23.5%,32.7%and 40.0%respectively,and this maybe resulted from interspecific difference.
Disease of Metarrhizium anisopliae can spread 400 meters far away from spraying area,and the average of infection rates from 8 directions around treatment area was 8.95%at 40 days after spraying, and the infection rate increased with time delaying.The level of infection rates from different directions was different,and it was the highest in northwest and southwest,east,north,south and northeast took second place,southeast and west were the lowest.The migratory tendency of grasshopper and wind direction may affect the spread of disease.Decrease of grasshopper population has been obtained with the spread of disease from spray area.At 200 metres distance to the treatment area,the average of corrected mortality from 8 directions was 44.05%at 42 days after spraying.The corrected mortality of different directions were different from each other,the level of corrected mortality keep consistent with infection rate in directions.
Metarrhizium anisopliae oil was sprayed on band pattern,and the results showed that moving and feeding behavior of grasshopper could spread disease,an effective control was obtained in non-spraying insecticide area between sprayed bands.
The spatial distribution of residual grasshoppers and cadavers after Metarhizium flavoviride oil spraying were analyzed by some indexes comprehensively in different sample interval and verified with Taylor's Power Law and improved Iwao's regression method during whole investigation period.The results showed that the spatial pattern of residual grasshoppers had the characteristics of regular aggregation and random distribution with the decrease of population density,and the cadavers were in aggregated pattern of distribution in different sample interval.The analysis of improved iwao's patchiness and Taylor's law showed that the spatial pattern of residual grasshoppers was aggregated within the whole investigation period which the cause was different reduced degree of densities in some plots after sprayed.Cadavers appeared to a false appearance of aggregative distribution with lower density and lesser sampling numbers.
引文
1.阿如旱,杨持.近50年内蒙古多伦县气候变化特征分析.内蒙古大学学报(自然科学版).2007.4(38):436-438.
2.阿如旱,杨持.内蒙古多伦县土地沙漠化景观格局变化特征.应用生态学报.2007,18(11):2520-2525.
3.白文明,李凌浩,宋世环.内蒙古多伦农牧交错区固定沙丘植被群落特征分析.草地学报.2003.3(11):223-227.
4.宝音陶格涛,刘丹.多伦县蔡木山自然保护区植物区系组成特征.内蒙古大学学报(自然科学版).2001,5(32):575-579.
5.陈永林.蝗虫再猖獗的控制与生态学治理.中国科学院院刊.2000,5:341-345.
6.丁岩饮.昆虫数量生态学.北京:科学出版社.1994,47-56.
7.樊美珍,李增智,王成树.感染球孢白僵菌的马尾松毛虫值虫产孢量的测定.安徽农业大学学报.1996,23(3):392-396.
8.冯光翰等.草地蝗虫种群数量消长数学模型研究.兰州大学学报(自然科学版).1994,30(1):100-103.
9.高丙利,吴东亮.马尾松毛虫白僵病流行在空间轴上的动态变化.中南林学院学报.1994,14:2 102-108.
10.高丙利,张景权.马尾松林昆虫群落与松毛虫白僵病流行的关系.中南林学院学报.1993,13:2 169-175.
11.巩爱岐,刘晓健,蒋湘,张龙.蝗虫微孢子虫疾病在青海草原蝗虫种群中的传播.中国生物防治.2003,(03):118-121.
12.胡加付,缪凯,柴守权,骆有庆,李增智.用酯酶同功酶分析白僵菌的林间扩散能力.中国生物防治.2005,21(4):238-241.
13.黄辉,朱恩林.我国蝗虫发生防治动态.大自然,2001.5:29-30.
14.蒋湘.病虫粪便在蝗虫微孢子虫病水平传播中的作用研究.青海畜牧兽医杂志.2001,31(2):4-5.
15.雷仲仁,问锦曾,谭正华,王音,谢志庚,李顺功,吴福海.绿僵菌油剂防治东亚飞蝗田间试验.植物保护.2003.29(1):11-19.
16.李宝玉.金龟子绿僵菌孢子制备、贮存及检测标准化研究[D].北京:中国农业科学院研究生院,2005.
17.李增智.昆虫真菌学的发展.见:中国虫生真菌研究与应用.第4卷,1997:1-5.
18.卢永洁.环境因子对白僵菌除治松毛虫效果的影响.河北林业科技.2005,5:4-5.
19.蒲蜇龙,李增智.昆虫真菌学.合肥:安徽科学技术出版社.1996,93:378.
20.王滨,丁德贵,樊美珍,李增智.白儒菌生物防治林中虫生真菌群落结构分析.应用生态学报,2005,16(12):2357-2360.
21.王振平,塔娜,严毓骅,石旺鹏.蝗虫微孢子虫病在优势蝗虫种类中的持续传播.华北农学报.2006,21(2):132-134.
22.吴福祯.中国的飞蝗.上海永祥印书馆.1951,3-7.
23.吴衍庆,史青茂,韩天文,王俊梅.蝗虫微孢子虫病在草原蝗虫中扩散传播的研究.草业科学.2000,17(4):25-26.
24.武觐文,夏乃斌.关于昆虫流行病的分析.北京林学院学报.1983,4:52-61.
25.徐汝梅,刘来福,丁岩饮.改进型Iwao m~*-m模型.生态学报.1984,4(2):111-118.
26.徐伟松等.昆虫病原真菌在害虫防治上的应用.广西农业科学.2003,(6):51-52.
27.严毓华.试论拓宽生物防治范围,发展虫害可持续治理.昆虫学报41(增刊):1998:1-4.
28.张龙.蝗虫微孢子病田间流行的初步调查.草地学报.1995,(3):223-229.
29.张泽华,高松,张刚应,王扬,杨宝东,张卓然,郑双悦,王梦龙.应用绿僵菌油剂防治内蒙草原蝗虫效果.中国生物防治.2000,16(2):49-52.
30.Alves RT,Bateman RP,Prior C,Leather SR.Effects of simulated solar radiation on conidial germination of Metarhiziumanisopliae in different formulations.Crop Protection.1998,17(8):675-679.
31.Anderson RM.Transmission dynamics and control of infectious diseases.In:Anderson RM,May MR,Ed.Population Biology of Infectious Diseases,Life Sciences Research Report 25,Dahlem Conference.Berlin:Springer.1982,149-176.
32.Arthurs SP,Thomas MB,Lawton JL.Seasonal patterns of persistence and infectivity of Metarhizium anisopliae var acridum in grasshopper cadavers in the Sahel.Entomologia Experimentalis et Applicata.2001,100(1):69-76.
33.Arthurs SP,Thomas MB.Factors affecting horizontal transmission of entomopathogenic fungi in locusts and grasshoppers.Asp Appl Biol.1999,53:89-98.
34.Balfour-Browne FL.The green muscardine disease of insects,with special reference to an epidemic in a swarm of locusts in Eritrea.Proc.R.Entomol.Soc.London Ser.A.1960,35:65-74.
35.Banlanca G.Visscher MN.Effects of very low doses of Fipronil on grasshoppers and nontarget insects following field trials for grasshopper control.Crop Protection.1997,16:553-564.
36.Bateman RP,et al.Screening for virulent isolates of entomopathogenic fungi against the desert locust Schistocerca gregaria(Forsk(?)l).Biocontrol Sci.Technol.1996,6:549-560.
37.Bennett LV,Symmons PM.A review of estimates of the effectiveness of certain control techniques and insecticides against the desert locust. Anti-Locust Bull. 50. London: Cent. Overseas Pest Control. 1972, 1-15.
38. Blanford S, Thomas MB, Langewald J. Behavioural fever in a population of the Senegalese grasshopper Oedaleus senegalensis and its implications for biological control using pathogens. Ecol. Entomol. 1998,23:9-14.
39. Blanford S, Thomas MB, Langewald J. Thermal ecology of Zonocerus variegates and its effect on biocontrol using pathogens. Agric For Entomol. 2000, 2: 3 - 10.
40. Blanford S, Thomas MB. Host thermal biology: the key to understanding insect-pathogen interactions and microbial pest control ? Agric For Entomol. 1999, 1: 195 - 202.
41. Carruthers R, Larkin TS, Firstencel H. Influence of thermal ecology on the mycosis of a rangeland grasshopper. Ecology. 1992, 73: 190 - 204 .
42. Cigliano MM, De Wysiecki ML, Lange CC. Grasshopper species diversity in the pampas grassland of Argentina. Diversity and Distribution. 2000, 6: 81-91.
43. Curriuthers RI, Feng Z. In vivo temperature dependent of Beauveria bassiana Mycosis of the European corn borer, ostrinia nubilis. Invertebr pathol. 1985, 46: 305 -311.
44. Driver F, Milner RJ, Trueman WH. A taxonomic revision of Metarhizium based on a phylogenetic analysis of ribosomal DNA sequence data. Mycol Res. 2000, 104: 135 - 151.
45. Food and Agriculture Organization. Evaluation of field trial data on the efficacy and selectivity of insecticides on locusts and grasshoppers. Reported by the FAO Locust Pesticide Referee Group. Locust Pesticide Referee Group Meet. 7~(th). Rome, Italy. 1998, 1 -24.
46. Fuxa JR,Yoshinori Tanada. 昆虫疾病流行学[M]. 北京:北京农业大学出版社, 1991.
47. Fuxa JR. Dispersion and spread of the entomopathogenic. fungus, Nomuraea rileyi in a soybean field. Environ Entomol. 1984, 13: 252 - 258.
48. Gapparov F. Locust and grasshopper outbreaks become increasingly serious in Uzbekistan. Adv. Appl.Acridology. 2001. 17.
49. Goettel MS, Johnson DL. Microbial ontrol of grasshoppers and locusts. em. Entomol Soc Can, 1997,400.
50. Hajek AE, et al. Detection of Entomophaga maimaiga (Zygomycetes:Entomophthorales) using enzyme-linked immunosorbent assay (ELISA). Invertebr Pathol. 1991, 58: 1 - 9.
51. Hooper GHS, Milner RJ, Spurgin PA, Prior C. Initial field assessment of Metarhizium flavoviride for control of Chortoicetes terminifera (Walker) (Orthoptera: Acrididae). J Aust Entomol Soc. 1995, 34: 83 - 84.
52. Hunter D, Surgin P. Fipronil and fenitrothion for locust control in Au2stralia: less is more. Adv. Appl.Acridology. 2001. 14.
53. Inglis DG, Johnson DL, Goettel MS. Effects of temperature and thermoregulation on mycosis by Beauveria bassiana in grasshoppers. Biol Control. 1996, 7: 131 - 139.
54. Johnson DL. Nosematidae and other Protozoa as agents for control of grasshoppers and locusts: current status and prospects. Mem Entomol Soc Can. 1997, 171: 375 - 389.
55. Kassa A, Stephan D. Laboratory and field evaluation of different formulations of Metarhizium anisopliae var. acridum submerged spores and aerial conidia for the control of locusts and grasshoppers. BioControl. 2004, 49 (1): 63 - 81.
56. Kish LP, Allen GE. The biology and ecology of. Nomuraea rileyi and a program for predicting its incidence, on Anticarsia gemmatalis in soybean. Fl Agric Exp Stn Bull. 1978, 795: 1 - 58.
57. Kooyman C, et al. Operational-scale application of entomopathogenic fungi for control of Sahelian grasshoppers. Proc R Soc London Ser B. 1997, 264: 541 - 46.
58. Krall S, Peveling R, Ba-Diallo D eds. New Strategies in Locust Control. 1997, Basel: Birkhauser. 522.
59. Lactin DJ, Johnson DL. Effects of insolation and body orientation on internal thoracic temperature of nymphal Melanoplus packardii (Orthoptera: Acrididae.). Environ Entomol 1996,25:423-429.
60. Lactin DJ, Johnson DL. Environmental, physical, and behavioural determinants of body temperature in grasshopper nymphs (Orthoptera: Acrididae). Can Entomol. 1998, 130: 551 -577.
61. Lange CE, Sánchoz, Wittenstein E. Effect of the pathogen Nosema locustae (Protozoa: Microspora) on mortality and development of nymphs of the South American locust , Schistocera cancella (Orthoptera : Acrididae). J. Orthoptera Res. 2000, 9: 77 - 80.
62. Langewald J, et al. Comparison of an organophosphate insecticide with a mycoinsecticide for the control of Oedaleus senegalensis (Orthoptera: Acrididae) and other Sahelian grasshoppers at an operational scale. Biocontrol Sci. Technol. 1999, 9: 199-214.
63. Langewald J, et al.. Use of Metarhizium anisopliae (flavoviride) var. acridum for control of Zonocerus variegatus: a model linking dispersal and secondary infection from the spray residue with mortality in caged field samples. Entomol Exper Applic. 1997, 82: 1 - 8.
64. Langewald J, Ouambama Z, Mamadou A, Peveling R, Stolz 1, Bateman R, Attignon S, Blanford S, Arthurs S, Lomer C. Comparison of an organophosphate insecticide with a mycoinsecticide for the control of Oedaleus senegalensis (Orthoptera : Acrididae) and other Sahelian grasshoppers at an operational scale. Biocontrol Sci. Technol. 1999, 9 (2): 199-214.
65. Latchininsky A. Problems and progress emerge from the acridid outbreak in Kazakhstan. Adv. Appe. Acridology. 2001, 15 - 16.
66. Lockwood JA, Bomer CR, Ewen AB. The history of biological control with Nosema locustae: lessons for locust management. Insect Science and Its Application. 1999a, 19: 1— 17.
67. Lockwood JA, Schell SP, Forster RN, Reuter C, Rachadi T. Reduced agentarea treatments (RAATs) for management of rangeland grasshoppers: efficacy and economics under operational conditions. International Journal of Pest Management. 1999b, 46: 29 -42.
68. Lomer CJ, Bateman RP, De Groote H, Dent D, Kooyman C, Langewald J, Peveling R, Thomas M. Development of strategies for the incorporation of microbial pesticides into the integrated management of locusts and grasshoppers. J. Agri. Forest Entomol. 1999, 1:71 -88.
69. Lomer CJ, Bateman RP, Johnson DL, Langewald J, Thomas MB. Biological control of locusts and grasshoppers. Annual Review of Entomology. 2001, 46: 667 - 702.
70. Lomer CJ, et al. Field infection of Zonocerus variegates following application of an oil based formulation of Metarhizium flavoviride conidia. Biocontrol Sci. Technol. 1993, 3: 337 - 346.
71. Lomer CJ, Prior C, Kooyman C. Development of Metarhizium spp. For the control of locusts and grasshoppers. Mem Entomol Soc Can. 1997, 171: 265 - 286.
72. Magalhaes BP, Lecoq M, Faria MR, Schmidt FGV, Guerra WD. Field trial with the entomopathogenic fungus Metarhizium anisopliae var. acridum against bands of the grasshopper Rhammatocerus schistocercoides in Brazil. Biocontrol Sci. Technol. 2000, 10(4): 427-441.
73. Millstein et al. Microclimatic humidity influence on conidial discharge in Erynia sp. (Entomopthorales: Entomophthoraceae), an entomophthogenic fungus of the. alfalfa weevil (Coleoptera: Curculionidae). Environ Entomol. 1982, 11:1166 — 1169.
74. Milner RJ, Soper RS. Bioassay of Entomophthora against the spotted alfalfa aphid, Therioaphis trifolii f. maculate. J. Invertebr Pathol. 1981, 37: 163 - 173.
75. Missonier J, Robert Y, Thoizon G. Epidemiological circumstances which seem to promote Entomophthorosis epizootics in three aphids. Entomophaga. 1970, 15: 169.
76. Paraiso A, Lomer CJ, Godonou I, Kpindou D. Preliminary studies on the ecology of Zonocerus variegatus in the Republic of Benin. 1992, See Ref. 116a, 133 - 141.
77. Peveling R, Attignon S, Langewald J, Ouambama Z. An assessment of the impact of biological and chemical grasshopper control agents on ground dwelling arthropods in Niger , based on presence/absence sampling. Crop Protection. 1999, 18(5): 323 - 329.
78. Price RE, Bateman RP, Browm HD, Butlera ET, Muller EJ. Aerial spray trials against brown locust (Loustana pardalina, Walker) nymphs in South Africa using oil-based formulations of Metarhizium flavoviride. Crop Protection. 1997, 16(4): 345 - 357.
79. Rooke HGD. Note on locusts in Iraq and the control measures adopted. Dept Agric Iraq Mem Baghdad RAE. 1930, 19: 1 - 84.
80. Shah PA, Gbongboui C, Godonou I, Houssou A, Lomer CJ. Survival and mortality of grasshopper egg pods from semi-arid cereal cropping areas of northern Benin. Bull Entomol Res. 2000. In press.
81. Shah PA, Godonou I, Gbongboui C, Lomer CJ. Natural levels of fungal infections in grasshoppers in Northern Benin. Biocontrol Sci. Technol. 1994, 4: 331 - 342.
82. Shah PA, Kooyman C, Paraiso A. Surveys for fungal pathogens of locusts and grasshoppers in Africa and the NearEast. Mem. Entomol. Soc. Can. 1997, 171: 27 - 35.
83. Siegel JP. Testing the pathogenicity and infectivity of entomopathogens to mammmls New York: Academic. 1997, 325 - 336.
84. Smits JE, Johnson DL, Lomer CJ. Avian pathological and physiological responses to dietary exposure to the fungus Metarhizium flavoviride, an agent for control of grasshoppers and locusts in Africa. Journal of Wildlife Diseases. 1999, 35 (2): 194 - 203.
85. Thomas MB, Gbongboui C, Lomer CJ. Between-season survival of the grasshopper pathogen Metarhizium flavoviride in the Sahel. Biocontrol Sci. Technol. 1996, 6: 569 - 573.
86. Thomas MB, Wood SN, Langewald J, Lomer CJ. Persistence of biopesticides and consequences for biological control of grasshoppers and locusts. Pestic Sci. 1997, 49: 47 - 55.
87. Thomas MB, Wood SN, Lomer CJ. Biological control of locusts and grasshoppers using a fungal pathogen: the importance of secondary cycling. Trans R Soc London Ser B. 1995, 259: 265 -270.
88. Thomas MB, Wood SN, Solorzano V. Application of insect-pathogen models to biological control. In Theoretical Approaches to Biological Control. London: Cambridge University Press. 1999,368-384.
89. Van der Valk H, Niassy A, Beye AB. Does grasshopper control create grasshopper problem? Monitoring side-effects of fenitrothion applications in the western Sahel. Crop Protection. 1999,18: 139-149.
90. Vandenberg JD, Soper RS. Prevalence of entomophthora mycoses in populations of spruce budworm, Choristoneura fumiferana. Environ Entomol. 1978, 7: 847 - 853.
91. Voronina EG. Entomophthorosis epizootics of the pea. aphid. Acyrthosiphon pisum Harris (Homoptera, Aphidoidea). Entomol. Rev. 1971, 4: 444 - 453.
92. Wright DE. Economic assessment of actual and potential damage to crops caused by the 1984 locust plague in south-eastern Australia. Journal of Environment Management. 1986, 23: 293 -308.