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The Potential of Entomophilic Nematodes in Insect Management 1 John M 270 Journal of Nematology, Volume 12, No. 4, October 1980 terial parasite of a plant nematode: morphology and Pasteuria-Blastobacter group. Can. J. Microbiol. 19: ultrastrueture. J. Bacteriol. 129:1091-1101. 609-614. 26. Sayre, R. M., W. P. Wergin, and R. E. Davis. 30. Thorne, G. 1940. Duboscqia penetrans n. sp. 1977. Occurrence of Moina rectirostris (Cladocera: (Sporozoa, Microsporidia, Nosematidae), a parasite Daphnidae) of a plant morphologically similar to of the nematode Pratylenchus pratensis (de Man) Pasteuria ramosa (Metchnikoff, 1888). Can. J. Filipjev. Proc. Hehn. Soc. Wash. 7:52-53. Microbiol. 23:1573-1579. 31. Walker. J. T. 1969. Pratylenchus penetrans 27. Shepherd, A. M., S. A. Clark, and A. Kemp- (Cobb) populations as influenced by microorganisms ton. 1973. An intracellular microorganism associated and soil amendments. J. Nematol. 1:260-264. with tissues of Heterodera spp. Nematologica 19: 32. Williams, J. R. 1960. Studies on the nema- 31-34. tode soil fauna of sugarcane fields in Mauritius. 28. Slack, J. M., and M. A. Gerencser. 1975. Nematologica 5:37-42. Actinomyces, filamentous bacteria; biology and 33. Williams, J. R. 1967. Observations on para- pathogenicity. Burgess Publishing Co., Minneapolis. sitic protozoa in plant-parasitic and free-living nem- p. 175. atodes. Nematologica 13:336-342. '29. Staley, J. T. 1973. Budding bacteria of the Biocontroh The Potential of Entomophilic Nematodes in Insect Management 1 John M. Webster 2 Abstract: A review of the development of entomophilic nematology and a commentary on the potential of entomophilic nematodes in controlling insect pests. The paper considers some of the major contributions to our knowledge of entomophilic nematology; factors involved in insect pest management and how they are applicable to the use of nematodes; nematodes which are most promising as biological control agents; and problems to be solved to facilitate the use of entomophilic nematodes in insect management. Key Words: Mermis nigrescens, Neoaplectana spp., Romanomermis culicivorax, Deladenus siricidicola, Tetradonema plicans. "A modern endeavour is to bring insect realized. Why? In the following discussion pests under natural control ..... The I will give some of the reasons. 'natural control' of insects will be most effective if all possible agencies and fac- MILEPOSTS tors are utilized; among these agencies nemas are by no means negligible." Mermis nigrescens, though atypical of the family, is often considered the type or- This is a quotation from Nathan Cobb ganism for entomophilic nematology and (12) in the 1927 USDA Yearbook of Agri- was described by Dujardin in France 137 culture. In recent years, pest management years ago (16). About 20 years later, Sir philosophy has developed in ways that John Lubbock, in Edinburgh (34), pro- avoid harm to useful organisms. As nema- vided biological and taxonomic informa- todes that kill pest insects are usually harm- tion on Sphaerularia bombi that widened less to other organisms, they have enormous contemporary perspectives on the nature of potential for use in place of chemical pesti- nematode-insect associations. Numerous cides in pest management systems. Never- publications in entomophilic nematology theless, more than 50 years after Cobb's followed, especially in Germany, and this statement, this potential has yet to be fully was crowned in the early part of the 20th century by a detailed taxonomic account of Received for publication18 February 1980. the mermithids by Hagmeier (27) of the 1Symposium paper presented at the annual meeting of University of Heidelberg. Cobb (10) entered the Society of Nematologists, Salt Lake City, Utah, U.S.A., 23-26 July 1979. tile arena in 1920 with descriptions of sev- 2Pestology Centre, Department of Biological Sciences, eral new species of entomophilic nematodes Simon Fraser University,Burnaby, Vancouver, Canada. VSA lS6. and followed by his famous observations on I thank Dr. B. P. Beirne of Simon Fraser University the biology and taxonomy of mermithids and Dr. J. J. Petersen of the United States Department of Agriculture for comments on the manuscript. (I 1,13). During the first half of the 20th cen- Entomophilic Nematodes: Webster 271 tury Cobb in the United States, Filipjev in of research stepped Rachael Carson, whose Russia, Steiner in Germany, and Bovien in book (8) stimulated major shifts in the Denmark made significant contributions to agricultural and social philosophies of the our understanding of the taxonomy of en- industrialized world. Since that time we tomophilic nematodes, and Tom Goodey's have witnessed a plethora of publications in presentation (22) to the Royal Society in entomophilic nematology and this time, 1930 on the biology and taxonomy of the nematologists have focused more on the use tylenchid Tylenchinema oscinellae, a para- of nematodes in insect control rather than site of the flit fly, was, and still is, a fine ex- on their taxonomy. ample of detailed biological observation. Many claim that tylenchids are possible Superimposed on these were Oldham's biological control agents (37,45). However, perspicacious comments in his paper "Hel- the only major success has been that of minths in the biological control of insect Bedding (1,2,4,5), with the Commonwealth pests" published by the former Imperial Scientific and Industrial Research Organiza- Bureau of Agricultural Parasitology (39). tion in Australia, using Deladenus, a Thereafter, interest in the subject waned for tylenchid parasite of the Sirex woodwasp. 20 years, except for occasional curiosity re- The mermithid parasites of insects have cordings such as that of Muspratt (35), who emerged as the most favoured nematological described mermithids in tree-hole breeding tool for insect control, and significant ad- Culicidae. vances have been made in all aspects of our Despite the mental perambulations of knowledge of this group--on their taxonomy Cobb in 1927, and the enthusiasm and suc- by Rubtsov (48,57) and Welch (55,57), on cesses of the 1930s, little progress was made their biology and ecology by Petersen (40, toward the application of entomophilic 43), and on the physiological and biochem- nematodes in the control of pest insects. An ical relationships with their hosts by Gordon exception to this generalization was the and Webster (23,24), Rutherford (49), and work of Glaser and his colleagues, of the Wtilker (59). However, only one species is New Jersey Department of Agriculture on the verge of commercial application, (United States),who steadily pursued a line namely Romanomermis culicivorax on mos- of research with rhabditid nematodes of the quitoes. Our knowledge of this nematode genus Neoaplectana. This was to provide a comes largely from 10 years of observations valuable base for subsequent applied stud- by Petersen (40-44) and colleagues at the ies. They spent several years examining the United States Department of Agriculture association of N. glaseri with the Japanese Laboratory at Lake Charles. Laird, of the beetle; subsequently they developed in vitro Research Unit on Vector Pathology in culture techniques for this nematode and Newfoundland, Canada has been a strong applied a similar species experimentally in advocate of the use of mermithids to control the field (21). In 1955 Dutky (17), at the biting flies in temperate and tropical cli- USDA Laboratory at Beltsville, pioneered mates (32). the use of the DD-136 strain of N. carpo- capsae for control of pest insects of orchards (17). This was followed in great anticipation INSECT PEST MANAGEMENT by applications of this species to pests of Despite tile long history of agriculture vegetable and field crops, forest crops, and and its success in progressively increasing to turf crops. crop productivity, the goal of optimum eco- During the same period Welch com- nomic production of a crop of high quality pleted his studies, at Rothamsted Experi- remains elusive. To achieve optimum qual- mental Station, England on entomophilic ity and economic production, the develop- nematodes of fruit flies and subsequently, ment of a sound pest management scheme at the Canada Department of Agriculture, is essential for each crop. To do this, a Belleville Laboratory, published a sequence systems approach constructed on a large of papers (eg. 54, 56) that demonstrated the data base should be used to enable simula- potential of nematodes for the control of tion and forecasting of the various biolog- pest insects. Into this gathering momentum ical, economic, and environmental events 272 Journal of Nematology, Volume 12, No. 4, October 1980 that provide information upon which the ogy, the fundamental obstacle to inte- pest manager can make his decision. grated control is epitomized in this Entomologists have been prolific in their conflict of short- and long-term goals. studies of pest biology, but there is a danger that research on modeling insect pest prob- We are now conscious of the energy lems might terminate in the publication of problem but insufficiently responsive to its a set of monographs rather than in the utili- solution. An energy shortage will hamper zation of the results in an interactive, ap- our ability to provide adequate nutrition plied programme. In nematology we have for the world's growing population. The just begun the modeling approach (30) on greater the energy shortage, the greater the plant-parasitic nematodes. Nothing has destruction of arable land and the higher been done using this technique with en- the costs of chemical pesticides. This pro- tomophilic
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