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Biological Control of Diamondback Moth Guan-Soon Lim Malaysian Agricultural Research and Development Institute, G

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Biological Control of Diamondback Moth Guan-Soon Lim Malaysian Agricultural Research and Development Institute, G 16 Biological Control of Diamondback Moth Guan-Soon Lim Malaysian Agricultural Research and Development Institute, G. P. 0. 12301, Kuala Lumpur 01-02, Malaysia Abstract The diamondback moth, Plutella xylostella (L), is recognized as a widely distributed and serious pest of crucifers in many countries. In some countries, however, it is effectively suppressed by parasites. These are critically examined as are the various attempts to import parasites and to release them. The appraisal reveals that the parasites of P. xylostella are a valuable control component and resource. Though numerous parasite species exist, not all are found to be effective. On the contrary, the key ones are restricted to only a few species and these belong largely to the genera Diadegma, Apanteles, and Microplitis. Except for Diadegma eucerophaga and possibly Diadegma fenestralis, none appears capable of exerting full control by itself. Many countries are still continuously plagued by P. Xylostella and there is strong evidence that this is due to the lack of crucially important parasites. Unless the relevant key species are made a part of the host-parasite complex, these problems are likely to persist. On the other hand, abandoning them will condemn crucifer cultivation to continued drenching in ever-increasing amounts of insecticides. These conclusions seem inescapable as all known cases of successful P. xylostella control have been obtained only when the basic control component constitutes either parasites or chemical insecticides. Introduction The diamondback moth (DBM), Plutella xylostella (L) (Lepidoptera: Yponomeu- tidae), has been recorded as long ago as 1746 (Harcourt 1962). Since then there have been many accounts of its importance throughout literature on economic entomology. In some countries, such as Argentina, Australia, New Zealand, and South Africa, its depredations were the cause of serious economic losses to cruciferous crops well before 1930 (Muggeridge 1930). Since then and until 1939, Robertson (1939) noted that although little detailed work appeared to have been done, the moth was recorded as a pest of crucifers in many other parts of the world: Tanganyika, Morocco, Chile, the Bahamas, British Columbia, Jamaica, Montana, Vancouver, Cyprus, Kona, Northern Caucasus, Lower Volga, Pulawy and Lubin, Latvia, Poland, Finland, Denmark and Britain. Many more localities were added in later years, viz Brazil (Becker M. 1981, personal com- munication), Bulgaria (Khristova 1957), Egypt (Hassanein 1958), Hong Kong (Lee 1968), India (Abraham and Padamanabhan 1968), Indonesia (Ankersmit 1953, Vos 1953), Malaysia (Henderson 1957, Ho 1965), the Philippines (Capco 1959), Singapore (Chuo 1973) and Taiwan (Hsu and Wang 1971, Chin 1974). Altogether, up to 1972 at least 128 countries or territories had reported the occurrence of this pest (CIE 1967, Salinas 1972). In general, the level of DBM infestation varies considerably according to the locality. For example, it is particularly serious in most of the countries in south and southeast Asia while only moderately so in several other Asian regions. On the other hand, in many parts of Canada, Europe and the United States (Muggeridge 1930, Hardy 160 Lim 1938, Sutherland 1966, Oatman and Platner 1969) it is generally of minor concern. In England, only very occasionally were large economic losses involved, resulting from mass immigration (Hardy 1938, French and White 1960, French 1965). In Germany, France and Italy, this insect is not present in sufficient numbers to be a serious pest, and in the last-named country it is comparatively rare (Muggeridge 1930). Nevertheless, it is very widely distributed, and the true distribution is undoubtedly greater than indicated by CIE (1967). Hardy (1938) believed that it could survive wherever crucifers are cultivated. In spite of the wide adaptability of DBM, the insect appears to be held in check in some regions. Marsh (1917), in outlining the situation in the United States, pointed out that DBM was a striking example of a potentially serious pest normally held in repression naturally by parasites. The occurrence of such natural control of DBM is, however, also recorded in many other places where the moth has remained relatively scarce (Muggeridge 1930, ardy 1938, Robertson 1939, Ullyett 1947, Kopvillem 1960). Such records suggest that D may have become a serious pest only in those countries in which natural enemies are absent or ineffective. Whether this is true requires critical examination. To date, investigations into the importance of natural enemies have been limited. This was clearly revealed in an analysis by Salinas (1972), wherein most of the published work on DBM between 1947-71 was found to be related to chemical control while studies on its general biology (life history, population dynamics, distribution and host plant records) ranked second. In contrast, papers mentioning natural enemies comprised barely 23% of the total literature, with those containing detailed studies on their biology, ecology and actual utilization even scarer. To a large extent this reflects how little importance has been attached to the biological control of DBM. Whether this is justified can only be gauged from an overview consideration and assessment of the status of the biological agents involved. Evidently, in such an appraisal it is recognized that not all the species of natural enemies are of equal significance and importance. Their contributions vary considerably, depending on the kinds of natural enemy agents, their interrelations and also their relationships with the environment. In view of this, it is essential to examine and estimate closely the real potential and/or limitations of these agents individually. Such knowledge is deemed necessary not only for formulating sound biological control programs but also for ensuring a better chance of success in their eventual utilization. Considering the agents individually or in small distinct groups is essential since biological systems are often complex. Besides, broad generalization would be difficult without running the risk of overlooking any specific and vital features that may exist. Such an analysis is thus attempted for all the known parasites of DBM in this paper. Other natural enemies, such as predators, are not considered as investigations into these have been scanty and incomplete. It is hoped that this appraisal will contribute to the attempts to realize the potential of some of these parasites in the control of DBM. Attempts at Parasite Importation and Release Biological control, which involves principally the introduction, augmentation, and conservation of natural enemies, has already proven itself to be a valuable weapon in pest control on a number of crops. Substantial numbers of successes have been achieved to date (DeBach 1964, Huffaker 1971, Pemberton 1948, Sweetman 1958). However, against DBM, it has been attempted only on a few occasions and the effort concerned largely the use of parasites, both indigenous and introduced. One of the earliest attempts was in New Zealand. Initially, work on biological control was undertaken to ascertain what natural enemies were present (Muggeridge 1930). From 1768 hosts collected 1582 DBM Biological Control 161 moths and 120 parasites were reared, revealing a very low degree (about 7%) of parasitic control. This was effected by the only larval parasite, Angitia laterallis Grav (Hymenop- tera: Ichneumonidae). However, follow-up studies in 1935-37 by Robertson (1939) recorded three parasitic species. Among these, only one species (Angitia sp) was con- firmed as a primary larval parasite, the second (Eupteromalus s~.)a hyperparasite, while in the case of the third (Diadromus sp) researchers were uncertain as to whether it was a parasite or a hyperparasite. In any case, none was able to provide effective control of DBM. Introductions were subsequently made (Muggeridge 1939) with effective suppression achieved by Diadegma eucerophaga Horstm (Angitia cerohaga) (Hymenop- tera: Ichneumonidae) and Thyraeella (Diadromus) collaris (Grav) (Hymenoptera: Ichneumonidae). In Australia a somewhat similar situation existed. Here, DBM was a major pest of crucifers throughout Australia until many parasitic species were introduced (Wilson 1960). Among these the more important ones were D. (Horogenes) eucerophaga, Diadegma (Hymenobosmina) rapi (Cam), T. collaris, Apanteles ippeux Nixon (Hymenop tera: Braconidae), and Apanteles plutellae Kurdj (Hymenoptera: Braconidae) (Wilson 1960, Hamilton 1978a, Goodwin 1979). D. eucerophaga was widely established both on the mainland of Australia and in Tasmania, and was extremely abundant in many areas. However, T. collaris was established mainly in Queensland, New South Wales (NSW), Victoria, and Tasmania, and A. plutellae in the Australian Capital Territory, New South Wales, and Queensland. Following the introductions there were subsequent reports of heavy parasitism with marked reduction in the abundance of DBM (Wilson 1960, Hamilton 1978b, Goodwin 1979). For example, in Richmond (NSW) in 1971-74 Diadegma spp, T. collaris and A. ippeus were responsible for parasitizing an average of 72% of DBM pupae collected, while in Victoria D. eucerophaga, T. collaris and D. rapi effected an average of 93% from 1972 to 1974. A highly successful introduction was also observed in Indonesia (Vos 1953). Efforts at biological control of DBM by parasite introductions had been initiated in 1928 (Eveleens and Vermeulen 1976). But these did not result in success until the
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