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Parasitoids as biological control agents – a fundamental approach

Published online by Cambridge University Press:  06 April 2009

J. K. Waage  and
M. P. Hassell
J. K. Waage
Affiliation:
Department of Pure and Applied Biology, Imperial College, London SW7 2AZ
M. P. Hassell
Affiliation:
Department of Pure and Applied Biology, Imperial College, London SW7 2AZ
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Summary

This review begins with a description of the parasitoid life-style and the ecological and evolutionary factors which generate the remarkable diversity of insect parasitoids. We then describe the various ways that parasitoids have been used in the biological control of insect pests, and survey their success to date. The use of parasitoids remains largely an art, aided by past experience of success and failure. A more fundamental approach, involving basic research and theory, has not as yet contributed significantly to practical biological control. We explore the potential for such a science of parasitoid use and review basic research on parasitoid ecology and evolution which is of particular relevance to biological control. Mathematical models are used to identify and examine those parasitoid and host attributes which lead to successful biological control. Factors such as parasitoid foraging behaviour, fecundity, larval survival and sex ratio are shown to be important in influencing the depression of host populations and/or the stability of host–parasitoid interactions after depression. Multiple release is discussed and a model for inundative release of parasitoids is explored.

Type
Research Article
Information
Parasitology , Volume 84 , Issue 4 , April 1982 , pp. 241 - 268
Copyright
Copyright © Cambridge University Press 1982

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References

Anonymous (1980). Biological Control Service. 25 Years of Achievement. Slough: Commonwealth Agricultural Bureaux.Google Scholar
Askew, R. R. (1971). Parasitic Insects. London: Heinemann Educational Books Ltd.Google Scholar
Askew, R. R. (1975). The organization of chalcid-dominated parasitoid communities centred upon endophytic hosts. In Evolutionary Strategies of Parasitic Insects and Mites (ed. Price, P. W.), pp. 130–53. New York: Plenum Press.Google Scholar
Beddington, J. R. (1975). Mutual interference between parasites or predators and its effects on searching efficiency. Journal of Animal Ecology 44, 331–40.Google Scholar
Beddington, J. R., Free, C. A. & Lawton, J. H. (1975). Dynamic complexity in predator–prey models framed in difference equations. Nature, London 225, 58–60.Google Scholar
Beddington, J. R., Free, C. A. & Lawton, J. H. (1978). Modelling biological control: on the characteristics of successful natural enemies. Nature, London 273, 513–19.Google Scholar
Beirne, B. P. (1975). Biological control attempts by introductions against pest insects in the field in Canada. Canadian Entomologist 107, 225–36.Google Scholar
Beirne, B. P. (1980). Biological control: benefits and opportunities. In Perspectives in World Agriculture, pp. 307–21. Slough: Commonwealth Agricultural Bureaux.Google Scholar
Bondarenko, N. V. (1978). Biologicheskaya Zashchita Rastenii. Leningrad: Isdatel'stvo Kolos (in Russian).Google Scholar
van den Bosch, R. & Messenger, P. S. (1973). Biological Control. London: Intertext Books.Google Scholar
Caltagirone, L. E. (1981). Landmark examples in classical biological control. Annual Review of Entomology 26, 213–32.Google Scholar
Caltagirone, L. E. & Huffaker, C. B. (1980). Benefits and risks of using predators and parasites for controlling pests. In Environmental Protection and Biological Forms of Control of Pest Organisms (ed. Lundholm, B. and Stackerud, M.), pp. 103–9. Stockholm: Swedish Natural Science Research Council.Google Scholar
Charnov, E. L. (1976). Optimal foraging; the marginal value theorem. Theoretical Population Biology 9, 129–36.Google Scholar
Charnov, E. L. (1979). The genetical evolution of patterns of sexuality: Darwinian fitness. American Naturalist 113, 465–80.Google Scholar
Charnov, E. L. (1982). The Theory of Sex Allocation. Princeton: Princeton University Press. (In the Press.)Google Scholar
Charnov, E. L., Los-Den Hartogh, R. L., Jones, W. T. & van den Assem, J. (1981). Sex ratio evolution in a variable environment. Nature, London 289, 27–33.Google Scholar
Chew, F. & Robbins, R. K. (1982). Egg laying in butterflies. In The Biology of Butterflies. Proceedings of a Symposium of the Royal Entomological Society of London, No. 11 (ed. Vane-Wright, R. I. and Ackery, P. R.). (In the Press.)Google Scholar
Clausen, C. P. (1978). Introduced Parasites and Predators of Arthropod Pests and Weeds. A World Review. Washington: United States Department of Agriculture/Agricultural Research Service.Google Scholar
Cook, R. M. & Hubbard, S. F. (1977). Adaptive searching strategies in insect parasites. Journal of Animal Ecology 46, 115–25.Google Scholar
Cook, R. M. & Hubbard, S. F. (1980). Effect of host density on searching behaviour of Nemeritis canescens (Hymenoptera: Ichneumonidae). Entomologia experimentalis et applicata 27, 205–10.Google Scholar
Coppel, H. C. & Mertins, J. W. (1977). Biological Insect Suppression. Berlin: Springer-Verlag.Google Scholar
Corbet, S. A. (1973). Concentration effects and the response of Nemeritis canescens to a secretion of its host. Journal of Insect Physiology 19, 2119–28.Google Scholar
De Bach, P. (1971). The use of imported natural enemies in insect pest management ecology. Proceedings Tall Timbers Conference on Ecological Animal Control by Habitat Management 3, 211–33.Google Scholar
Debach, P. (1974). Biological Control by Natural Enemies. Cambridge: Cambridge University Press.Google Scholar
Delucchi, V. L. (Ed.) (1976). Studies in Biological Control. Cambridge: Cambridge University Press.Google Scholar
Dowell, R. V. & Horn, D. J. (1977). Adaptive strategies of larval parasitoids of the alfalfa weevil (Coleoptera: Curculionidae). Canadian Entomologist 109, 641–8.Google Scholar
Ehler, L. E. & Van Den Bosch, R. (1974). An analysis of the natural biological control of Trichoplusia ni (Lepidoptera: Noctuidae) on cotton in California. Canadian Entomologist 106, 1067–73.Google Scholar
Ehler, L. E. & Miller, J. C. (1978). Biological control in temporary agro-ecosystems. Entomorphaga 23, 207–12.Google Scholar
Embree, D. G. (1966). The role of introduced parasites in the control of the winter moth in Nova Scotia. Canadian Entomologist 98, 1159–68.Google Scholar
Flanders, S. E. (1953). Variations in susceptibility of citrus-infesting coccids to parasitization. Journal of Economic Entomology 46, 266–9.Google Scholar
Flanders, S. E. (1971). Multiple parasitism of armoured coccids (Homoptera) by host regulative aphelinids (Hymenoptera); ectoparasites versus endoparasites. Canadian Entomologist 103, 857–72.Google Scholar
Force, D. C. (1970). Competition among four hymenopterous parasites of an endemic host. Annals of the Entomological Society of America 63, 1675–8.Google Scholar
Force, D. C. (1972). r and K, strategies in endemic host–parasitoid communities. Bulletin of the Entomological Society of America 18, 135–7.Google Scholar
Force, D. C. (1975). Succession of r and K strategists in parasitoids. In Evolutionary Strategies of Parasitic Insects and Mites (ed. Price, P. W.), pp. 112–129. New York: Plenum Press.Google Scholar
Free, C. A., Beddington, J. R. & Lawton, J. H. (1977). On the inadequacy of simple models for mutual interference for parasitism and predation. Journal of Animal Ecology 46, 543–54.Google Scholar
Galis, F. & Van Alphen, J. J. M. (1981). Patch time allocation and search intensity of Asobara tabida Nees (Braconidae), a larval parasitoid of Drosophila. Netherlands Journal of Zoology 31, 596–611.Google Scholar
Gauld, I. D. & Mound, L. A. (1982). Homoplasy and the delineation of holophyletic genera in some insect groups. Systematic Entomology 7 (in the Press).Google Scholar
Gross, H. R., Lewis, W. J., Jones, R. L. & Nordlund, D. A. (1975). Kairomones and their use for management of entomophagous insects: III. Stimulation of Trichogramma achaeae, T. pretiosum and Microplitis croceipes with host seeking stimuli at the time of release to improve efficiency. Journal of Chemical Ecology 1, 431–8.Google Scholar
Haeselbarth, E. (1979). Zur Parasitierung der Puppen von Forleule (Panolis flammea Schiff.), Kiefernspanner (Bupalus piniarius L.) und Heidelbeerspanner (Boarmia bistortata Goeze) in bayerischen Kiefernwäldern. Zeitschrift für angewandte Entomologie 87, 311–22.Google Scholar
Hall, R. W. & Ehler, L. E. (1979). Rate of establishment of natural enemies in classical biological control. Bulletin of the Entomological Society of America 25, 280–2.Google Scholar
Hamilton, W. D. (1967). Extraordinary sex ratios. Science 156, 477–88.Google Scholar
Hansberry, R. (1968). Prospects for non-chemical insect control – an industrial view. Bulletin of the Entomological Society of America 14, 229–35.Google Scholar
Hassan, S. A. (1981). Mass-production and utilization of Trichogramma: 2. Four years successful biological control of the European corn borer. Mededelingen der Faculteit Landbouwweten-schappan der Rijksuniversiteit Gent 46 (in the Press).Google Scholar
Hassell, M. P. (1978). The Dynamics of Arthropod Predator–Prey Systems. Monographs in Population Biology 13, Princeton University Press.Google Scholar
Hassell, M. P. (1979). The dynamics of predator-prey interactions: polyphagous predators, competing predators and hyperparasitoids. In Population Dynamics (ed. Anderson, R. M., Turner, B. D. and Taylor, L. R.), pp. 283–306. Oxford: Blackwell Scientific Publications.Google Scholar
Hassell, M. P. (1980). Foraging strategies, population models and biological control: a case study. Journal of Animal Ecology 49, 603–28.Google Scholar
Hassell, M. P. (1982). What is searching efficiency? Annals of Applied Biology (in the Press).Google Scholar
Hassell, M. P. & Comins, H. N. (1978). Sigmoid functional responses and population stability. Theoretical Population Biology 14, 62–7.Google Scholar
Hassell, M. P. & May, R. M. (1974). Aggregation in predators and insect parasites and its effect on stability. Journal of Animal Ecology 43, 567–94.Google Scholar
Hassell, M. P. & Moran, V. C. (1976). Equilibrium levels and biological control. Journal of the Entomological Society of South Africa 39, 357–66.Google Scholar
Hassell, M. P. & Varley, G. C. (1969). New inductive population model for insect parasites and its bearing on biological control. Nature, London 223, 1133–6.Google Scholar
Herting, B. (1972–1982). A Catalogue of Parasites and Predators of Terrestrial Arthropods. Slough: Commonwealth Agricultural Bureaux.Google Scholar
Hespenheide, H. A. (1979). Are there fewer parasitoids in the tropics? American Naturalist 113, 766–9.Google Scholar
Howell, J. & Fisher, R. C. (1977). Food conversion efficiency of a parasitic wasp, Nemeritis canescens. Ecological Entomology 2, 143–51.Google Scholar
Hoy, M. (1976). Establishment of gypsy moth parasitoids in North America: an evaluation of possible reasons for establishment or non-establishment. In Perspectives in Forest Entomology (ed. Anderson, J. F. and Kaya, H. K.), pp. 215–232. New York: Academic Press.Google Scholar
Hubbard, S. F. & Cook, R. M. (1978). Optimal foraging by parasitoid wasps. Journal of Animal Ecology 47, 593–604.Google Scholar
Huffacker, C. B. (Ed.) (1971). Biological Control. New York: Plenum Press.Google Scholar
Huffacker, C. B. & Messenger, P. S. (Eds.) (1976). Theory and Practice of Biological Control. New York: Academic Press.Google Scholar
Hussey, N. W. & Bravenboer, L. (1971). Control of pests in glasshouse culture by the introduction of natural enemies. In Biological Control (ed. Huffaker, C. B. and Messenger, P. S.), pp. 195–216. New York: Plenum Press.Google Scholar
Janzen, D. H. & Pond, C. M. (1975). A comparison, by sweep sampling, of the arthropod fauna of secondary vegetation in Michigan, England and Costa Rica. Transactions of the Royal Entomological Society of London 127, 33–50.Google Scholar
Khorkhordin, E. G. (1976). Searching behaviour of females of natural populations of Pleolophus basizonus (Hymenoptera: Ichneumonidae). Zoologicheskii Zhurnal 55, 1006–17 (in Russian).Google Scholar
Khorkhordin, E. G. (1978). Some parameters of search behaviour in females of a natural population of Pleolophus basizonus (Hymenoptera: Ichneumonidae). Zoologicheskii Zhurnal 57, 1651–5 (in Russian).Google Scholar
Klomp, H. (1959). Infestations of forest insects and the role of parasites. Proceedings of the 15th International Congress of Zoology (1958), 797–802.Google Scholar
Knipling, E. F. & McGuire, J. O. (1968). Population models to appraise the limitations and potentialities of Trichogramma in managing host insect populations. United States Department of Agricultural Technical Bulletin, No. 1387, 44 pp.Google Scholar
Kochetova, N. I. (1978). Factors determining the sex ratio in some entomophagous Hymenoptera. Entomological Reviews 60, 1–5.Google Scholar
Laing, J. E. & Hamai, J. (1976). Biological control of insect pests and weeds by imported parasites, predators and pathogens. In: Theory and Practice of Biological Control (ed. Huffaker, C. B. and Messenger, P. S.), pp. 685–743. New York: Academic Press.Google Scholar
Lawton, J. H. & Hassell, M. P. (1982). Interspecific competition in insects. In Ecological Entomology (ed. Huffaker, C. B. and Rabb, R. L.). New York:Wiley Interscience. (In the Press.)Google Scholar
Lawton, J. H. & Price, P. W. (1979). Species richness of parasites on hosts: Agromyzid flies on the British Umbelliferae. Journal of Animal Ecology 48, 619–37.Google Scholar
van Lenteren, J. C. (1980). Evaluation of control capabilities of natural enemies: does art have to become science? Netherlands Journal of Zoology 30, 369–81.Google Scholar
van Lenteren, J. C. (1981). Host discrimination by parasitoids. In: Semiochemicals. Their Role in Pest Control (ed. Nordlund, D. A., Jones, R. L., and Lewis, W. J.), pp. 153–79. New York: John Wiley & Sons.Google Scholar
van Lenteren, J. C. & Bakker, K. (1978). Behavioural aspects of the functional response of a parasite (Pseudeucoila bochei Weld) to its host (Drosophila melanogaster). Netherlands Journal of Zoology 28, 213–33.Google Scholar
Lewis, C. J. (1977). The economics of pesticide research. In Origins of Pest, Parasite, Disease and Weed Problems (ed. Cherret, J. M. and Sagar, G. R.), pp. 237–45. Oxford: Blackwell Scientific Publications.Google Scholar
Lewis, W. J., Jones, R. L., Gross, H. R. & Nordlund, D. A. (1976). The role of kairomones and other behavioural chemicals in host-finding by parasitic insects. Behavioral Biology 16, 267–89.Google Scholar
Lewis, W. J., Jones, R. L., Nordlund, D. A. & Sparks, A. N. (1975). Kairomones and their use for management of entomophagous insects: I. Evaluation for increasing rates of parasitization by Trichogramma spp. in the field. Journal of Chemical Ecology 1, 343–7.Google Scholar
Luck, R. F., van Lenteren, J. C., Twine, P. H., Kuenen, L. & Unruh, T. (1979). Prey or host searching behaviour that leads to a sigmoid functional response in invertebrate predators and parasitoids. Researches on Population Ecology 20, 257–64.Google Scholar
Malyshev, S. I. (1968). Genesis of the Hymenoptera and the Phases of Their Evolution. London: Methuen & Co. Ltd.Google Scholar
Marsden, J. S., Martin, G. E., Parnham, D. J., Ridsdill Smith, T. J. & Johnston, B. G. (1980). Returns on Australian Agricultural Research. Canberra: Commonwealth Scientific and Industrial Research Organization.Google Scholar
May, R. M. (1978). Host-parasitoid systems in patchy environments: a phenomenological model. Journal of Animal Ecology 47, 833–43.Google Scholar
May, R. M. & Hassell, M. P. (1981). The dynamics of multiparasitoid-host interactions. American Naturalist 117, 234–61.Google Scholar
May, R. M., Hassell, M. P., Anderson, R. M. & Tonkyn, D. W. (1981). Density dependence in host-parasitoid models. Journal of Animal Ecology 50, 855–65.Google Scholar
Messenger, P. S. (1976). Theory underlying introduction of exotic parasitoids. In Perspectives in Forest Entomology (ed. Anderson, J. F. and Kaya, H. K.), pp. 191–214. New York: Academic Press.Google Scholar
Miller, J. C. (1977). Ecological relationships among parasites of Spodoptera praefica. Environmental Entomology 6, 581–5.Google Scholar
Mills, N. J. (1982). Voracity, cannibalism and coccinellid predation. Annals of Applied Entomology (in the Press).Google Scholar
Moran, V. C. (1981). Belated kudos for cochineal insects. Bulletin of the Royal Entomological Society of London 5, 54–8.Google Scholar
Morrison, G., Auerbach, M. & McCov, E. D. (1979). Anomalous diversity of tropical parasitoids: a general phenomenon? American Naturalist 114, 303–7.Google Scholar
Morrison, G., Lewis, W. J. & Nordlund, D. A. (1980). Spatial differences in Heliothis zea egg density and the intensity of parasitism by Trichogramma spp.: an experimental analysis. Environmental Entomology 9, 79–85.Google Scholar
Munroe, E. G. (1971). Status and potential of biological control in Canada. In Biological Control Programmes Against Insects and Weeds in Canada, pp. 215–54. Slough: Commonwealth Agricultural Bureaux.Google Scholar
Murdoch, W. W. (1977). Stabilizing effect of spatial heterogeneity in predator–prey system. Theoretical Population Biology 11, 252–73.Google Scholar
Murdoch, W. W. & Oaten, A. (1975). Predation and population stability. Advances in Ecological Research 9, 2–131.Google Scholar
Nicholson, A. J. & Bailey, V. A. (1935). The balance of animal populations, Part 1. Proceedings of the Zoological Society of London 3, 551–98.Google Scholar
Patton, S., Craig, I. & Conway, G. R. (1980). Insecticide and acaricide resistance. In: Pesticide Resistance and World Food Production (ed. Conway, G. R.), pp. 14–28. London: Imperial College Centre for Environmental Technology, Series E, No. 2.Google Scholar
Pickering, J. (1980). Larval competition and brood sex ratios in the gregarious parasitoid Pachysomoides stupidus. Nature, London 283, 291–2.Google Scholar
Pimentel, D. (1961). Animal population regulation by the genetic feedback mechanism. American Naturalist 95, 65–79.Google Scholar
Pimentel, D. (1963). Introducing parasites and predators to control native pests. Canadian Entomologist 95, 785–92.Google Scholar
Price, P. W. (1973). Parsitoid strategies and community organization. Environmental Entomology 2, 623–6.Google Scholar
Price, P. W. (1975 a). Reproductive strategies of parasitoids. In: Evolutionary Strategies of Parasitic Insects and Mites (ed. Price, P. W.), pp. 87–111. New York: Plenum Press.Google Scholar
Price, P. W. (Ed.) (1975 b). Evolutionary Strategies of Parasitic Insects and Mites. New York: Plenum Press.Google Scholar
Pu, C. L. (1978). Principles and Methods of Biological Control of Insect Pests. Peking: Ko-hsueh Press (in Chinese).Google Scholar
Pyle, R., Bentzien, M. & Opler, P. (1981). Insect conservation. Annual Review of Entomology 26, 233–58.Google Scholar
Rabinovich, J. E. (1970). Population dynamics of Telenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas disease vectors. IV. Effect of female size on progeny. Journal of Medical Entomology 7, 561–7.Google Scholar
Rabinovich, J. E. (1971). Population dynamics of Telenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas disease vectors. V. Parasite size and vital space. Revista de Biologia Tropical 19, 109–20.Google Scholar
Rathke, B. J. & Price, P. W. (1976). Anomalous diversity of tropical ichneumonid parasitoids: a predation hypothesis. American Naturalist 110, 889–93.Google Scholar
Reuter, O. M. (1913). Lebensgewohnheiten und Instinkte der Insekten. Berlin: Friedlander.Google Scholar
Rogers, D. J. (1972). Random search and insect population models. Journal of Animal Ecology 41, 369–83.Google Scholar
Rogers, D. J. & Hassell, M. P. (1974). General models for insect parasite and predator searching behaviour: interference. Journal of Animal Ecology 43, 239–53.Google Scholar
Rogers, D. J. & Hubbard, S. F. (1974). How the behaviour of parasites and predators promotes population stability. In Ecological Stability (ed.Usher, M. B. and Williamson, M. H.), pp. 99–119. London: Chapman and Hall Ltd.Google Scholar
Rosen, D. (Ed.) (1981). The Role of Hyperparasitism in Biological Control – A Symposium. Berkeley: Division of Agricultural Sciences, University of California.Google Scholar
Sandlan, K. (1979). Sex ratio regulation in Coccygomimus turionella Linnaeus (Hymenoptera: Ichneumonidae) and its ecological implications. Ecological Entomology 4, 365–8.Google Scholar
Salt, G. (1935). Experimental studies in insect parasitism. III. Host selection. Proceedings of the Royal Society of London, B. 117, 413–35.Google Scholar
Salt, G. (1963). The defence reactions of insects to metazoan parasites. Parasitology 53, 527–642.Google Scholar
Salt, G. (1968). The resistance of insect parasitoids to the defence reactions of their hosts. Biological Reviews 43, 200–32.Google Scholar
Sharov, A. A. (1979). Effect of the spatial structure of interacting populations of Neodiprion sertifer and its parasite, Exenterus abruptorius, on the dynamics of their numbers. Zoologicheskii Zhurnal 58, 356–65 (in Russian).Google Scholar
Simmonds, F. J. & Bennett, F. D. (1977). Biological control of agricultural pests. Proceedings of the 15th International Congress of Entomology, 1976, pp. 464–472. Washington: Entomological Society of America.Google Scholar
Slansky, F. (1978). Utilization of energy and nitrogen by larvae of the imported cabbageworm, Pieris rapae, as affected by parasitism by Apanteles glomeratus. Environmental Entomology 7, 179–85.Google Scholar
Southwood, T. R. E. (1977). The relevance of population dynamic theory to pest status. In Origins of Pest, Parasite, Disease and Weed Problems (ed. Cherrett, J. M. and Sagar, G. R.), pp. 35–54. Oxford: Blackwell Scientific Publications.Google Scholar
Southwood, T. R. E. (1981). Bionomic strategies and population parameters. In: Theoretical Ecology (ed. May, R. M.), pp. 30–52. Oxford: Blackwell Scientific Publications.Google Scholar
Starý, P. & Rejmánek, M. (1981). Number of parasitoids per host in different systematic groups of aphids: the implications for introduction strategy in biological control (Homoptera: Aphidoidea; Hymenoptera: Aphidiidae). Entomologica Scandinavica Suppl. 15, 341–51.Google Scholar
Stinner, R. E. (1977). Efficacy of inundative release. Annual Review of Entomology 22, 515–31.Google Scholar
Stinner, R. E., Ridgway, R. L. & Morrison, R. K. (1974). Longevity, fecundity, and searching ability of Trichogramma pretiosum reared by three methods. Environmental Entomology 3, 558–60.Google Scholar
Suzuki, Y. & Iwasa, Y. (1980). A sex ratio theory of gregarious parasitoids. Researches on Population Ecology 22, 366–82.Google Scholar
Sweetman, H. L. (1936). The Biological Control of Insects. Ithaca, New York: Comstock Publishing Company Incorporated.Google Scholar
Taylor, J. C. (1979). Sensoriaphis in Western Australia. Bulletin of the Royal Entomological Society of London 3, 106.Google Scholar
Taylor, T. H. C. (1955). Biological control of insect pests. Annals of Applied Biology 42, 190–6.Google Scholar
Thompson, W. R. (1924). Le théorie mathématique de l'action des parasites entomophages et le facteur du hassard. Annates de la Faculté des sciences de Marseille 2, 69–89.Google Scholar
Thompson, W. R. (1939). Biological control and the theories of the interactions of populations. Parasitology 31, 299–388.Google Scholar
Thompson, W. R. (1943–1958). A Catalogue of the Parasites and Predators of Insect Pests. Belleville: Commonwealth Agricultural Bureaux.Google Scholar
Townes, H. (1969). The Genera ofIchneumonidae. Part 1. Memoirs of the American Entomological Institute. Number 11.Google Scholar
Turnbull, A. L. & Chant, D. A. (1961). The practice and theory of biological control of insects in Canada. Canadian Journal of Zoology 39, 697–753.Google Scholar
Vet, L. E. M., van Lenteren, J. C. & Woets, J. (1980). The parasite–host relationship between Encarsia formosa (Hymenoptera: Aphelinidae) and Trialeurodes vaporariorum (Homoptera: Aleyrodidae). IX. A review of the biological control of the greenhouse whitefly with suggestions for future research. Zeitschrift für angewandte Entomologie 90, 26–51.Google Scholar
Viktorov, G. A. (1976). The Ecology of Entomophagous Parasites. Moscow: Izdatel'stvo Nauka. (In Russian).Google Scholar
Vinson, S. B. (1976). Host selection by insect parasitoids. Annual Review of Entomology 21, 109–33.Google Scholar
Vinson, S. B. (1981). Habitat location. In Semiochemicals. Their Role in Pest Control (ed. Nordlund, D. A., Jones, R. L. and Lewis, W. J.), pp. 51–77. New York: John Wiley & Sons.Google Scholar
Vinson, S. B. & Iwantsch, G. F. (1980). Host regulation by insect parasitoids. Quarterly Review of Biology 55, 143–65.Google Scholar
Waage, J. K. (1978). Arrestment responses of the parasitoid, Nemeritis canescens, to a contact chemical produced by its host, Plodia interpunctella. Physiological Entomology 3, 135–46.Google Scholar
Waage, J. K. (1979). Foraging for patchily-distributed hosts by the parasitoid, Nemeritis canescens. Journal of Animal Ecology 48, 353–71.Google Scholar
Waage, J. K. (1982 a). Sib-mating and sex ratio strategies in scelionid wasps. Ecological Entomology 7 (in the Press).Google Scholar
Waage, J. K. (1982 b). Sex ratio and population dynamics in natural enemies – some possible interactions. Annals of Applied Biology (in the Press).Google Scholar
Watt, K. E. F. (1965). Community stability and the strategy of biological control. Canadian Entomologist 97, 887–95.Google Scholar
Way, M. J. (1973). Objectives, methods and scope of integrated control. In: Insects: Studies in Population Management (ed.Geier, P. W., Clark, L. R., Anderson, D. J. and Nix, H. A.), pp. 1–17. Ecological Society of Australia, memoirs 1.Google Scholar
Werren, J. H. (1980). Sex ratio adaptations to local mate competition in a parasitic wasp. Science 208, 1157–9.Google Scholar
Weseloh, R. M. (1976). Behaviour of forest insect parasitoids. In Perspectives in Forest Entomology (ed. Anderson, J. F. and Kaya, H. K.), pp. 99–110. New York: Academic Press.Google Scholar
Weseloh, R. M. (1981). Host location by parasitoids. In Semiochemicals: Their Role in Pest Control (ed. Nordlund, D. A., Jones, R. L. and Lewis, W. J.), pp. 79–95. New York: John Wiley & Sons.Google Scholar
Wylie, H. G. (1976). Interference among females of Nasonia vitripennis (Hymenoptera: Pteromalidae). Canadian Entomologist 108, 655–61.Google Scholar
Wylie, H. G. (1979). Sex ratio variability of Muscidifurax zaraptor (Hymenoptera: Pteromalidae). Canadian Entomologist 111, 105–9.Google Scholar
Yano, E. (1978). A simulation model of searching behaviour of a parasite. Researches on Population Ecology 20, 105–22.Google Scholar
Zwölfer, H. (1971). The structure and effect of parasitoid complexes attacking phytophagous host insects. In: Dynamics of Populations (ed. den Boer, P. J. and Gradwell, G. R.), pp. 405–418. Wageningen: Centre for Agricultural Publishing and Documentation.Google Scholar