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Ecological Considerations in Chemical Control1 Introduction By Department of Entomology RAY F. SMITH and Parasitology, Ulliversit'jI of California, Berkeley Traditional approaches to insect-pest problems have centered on the development of techniques which attempt to destroy pests as they appear in threatening numbers and hopefully to eliminate them. These techniques have, for the most part, involved the use of chemicals applied with little reference to the complexities of agricultural ecosystems. In general, these control measures have been successful. However, we must put certain qualifications on the success of these traditional approaches. Their inherent limitations become clearer each year. fundamental knowledge than does indiscriminate treatment with pesticides on specific dates or when the pests first appear. A sound background is needed on the following points: In recent months, it has been said many many times that we must turn to new methods-especially biological methods--()f pest control and a few say we must eliminate the use of pesticides. I disagree violently. \\Fe shouldn't eliminate any method that will help us in our struggle with the insects. The present need is new perspectives in pest control and these perspectives with their consequent new techniques are being developed. It is obvious that no one management technique, e.g. use of chemicals or introduction of biological control, will be capable of handling all pest problems. A multiple approach is needed which pursues all possible new leads and utilizes all techniques to their fullest in an integrated approach; this approach is of course what we have come to call integrated control. 1) Why does the insect achieve pest status 2) The complexities of the crop ecosystem 3) The effects of managment procedures on all pest species as well as the beneficial forms 4) An evaluation of the pest damage in terms of the economics of crop production I must integrated will be so incorporate ing as we These new approaches have as their objective the mallagement of the pest pop1dation in such a manner as to avoid economic damage rather than their complete elimination from a field or area. Of necessity, such pest population management requires a much broader base of ] A s~mposium presented at the St. Louis Meeting Entomological Society of America, December 3, 1963. of stress that chemical control is a mainstay of control. \Ve are dependent on chemicals and dependent indefinitely. \\That we must do is to modern ecological concepts into our thinkconsider chemical control procedures. It is my opinion that our ecological view of chemical control should be as broad as possible and our symposium panel will attempt to bring that broad ecological viewpoint to bear. the Ecological Considerations in Chemical Control Insects in the Human By Department MARSTON Ecosystem BATES of Zoology, University of Michigan, Ann Arbor Ecologists customarily divide the terrestrial environments of the earth into a series of major biomes: tundra, taiga, deciduous forest, grassland, desert, and the like. It seems to me that to this list should be added the "man-altered biome" or "man-altered landscape," for the environments in which human activity is the major determinant of the survival of organisms. The man-altered environments are, to be sure, extremely diverse, including such different things as cities, gardens, orchards, fields, rice paddies, managed forests, and abandoned clearings. But there is one important common factor to all of these: for survival, an animal or plant must be able to adapt to the consequences of the actions of man. habitats as little disturbed by man as possible. The result is a large area of neglect, which must be studicd if the system as a whole is to be understood. "Human ecosystem" is, I think, a better label than "man-altered landscape." The idea of ecosystcms involves awareness of the constant interaction, interrelation, between organisms and environments. '"\Then we use a term like "biome" or "biological community" we tend to think of a group of organisms in some particular enviromuental setting, while actually the organisms are constantly modifying the environment and vice-versa. To cite an obvious example: the character of a forest depends in part on the soil type in which it is growing; but the soil type is also, in part, a consequence of the kind of forest growth. This kind of transaction between organisms and the physical environment is constantly taking place. In the case of the biological environment, the vcry idea is based on the transactions between one This man-altered landscape is rapidly coming to dominate the land surface of the planet, yet it is far from receiving its fair share of attention from biologists. The agricultural scientists tend to concentrate their attention on cultivated areas, while ecologists like to find 67 organism or one kind of organism things present in the neighborhood. and other living forward in man's control of the ills that afflict him; but it is also a shift in ideas, a change in the conceptual environment. In a lecture given at Berkeley (Bates 1962), I attempted to analyze the human environment-the setting in which man's actions take place. A common approach is in terms of climate, inorganic resources, other organisms, and the like. Or we can use a quite different method of analysis. There is the perceptual environment-the elements in the world around us that are interpreted by our sense organs; the operational environment-the elements that influence us in some way, whether perceived or not; and the total environment-all the forces of the external world including those, like radio waves from distant stars, that have no detectable influence on our behavior. It is curious that insects are very generally regarded as pes:s, vermin, "enemies," by the peoples of Western civilization. Even the attempts of entomo!ogists to stress the "usefulness" of many kinds of insects meet with little success. Nowadays one can argue that insects transmit diseases, or that they are a major hazard for scientific agriculture. But I suspect the Western attitude long a:1tedates these discoveries. It would be interesting to try to trace the history of this dislike of insects, though I do not know how one would go about it. I do think our attitude is unusual, if not unique. For one thing, as far as I know ours is the only major culture with a. strong taboo against eating insects (if we except honey, as an insect product). We not only do not eat insects-we view the idea of eating them with horror. Yet we eat oysters, shrimp, lobster, and many other marine invertebrates. Why the taboo on insects? In any such analysis as this, we quickly come across the problem of how to deal with man's ideas. I first thought of this as an environmental problem in relation to the supernatural world that forms an important part of the setting of any primitive people. For such people, spirits are everwhere, in the bushes, the water, the clouds. They can cause great damage by bringing on disease, unleashing hurricanes, ruining crops; or if properly propitiated, they can be of great help in all sorts of human enterprises. We can laugh; we know that the spirits have no "real" existence, that they are products of the human imagination. But they are real enough to the people concerned-just as real as the sharks in the lagoon or the leopards in the forest. In terms of influence on human behavior, they are far more important than the visible animals. But let's get back to the human ecosystem. Man clearly is involved in a series of relationships with other organisms quite different from those of any other animal. He has become, in fact, a sort of geological force, moulding the landscape and determining the destinies of a host of other organisms. ("Man's Role in Changing the Face of the Earth" is the subject of a large symposium volume edited by Thomas (1956).) How did he get this way? And how can we classify the kinds of relationships he has developed? We can laugh; but we have our spirits, too, only we are likely to call them ideas. And these ideas, even though they are products of our minds, constantly influence our behavior. As an example, I have sometimes thought that any word ending in "-ism" in English tends to denote a malevolent spirit: Nazism, communism, Malthusianism, McCarthyism, scientistism, perhaps even humanism. With us, it is difficult to label such spirits as part of the supernatural world; it is better, I think, to call this world of ideas surrounding every man the "conceptual environment." This environment is the same thing as the "culture" of the anthropologists, and perhaps as the "superego" of the clinical psychologists. But we gain a different, and I think more useful, insight if we look at it as an aspect of the environment involved in thc total complex of the human ecosystem. I like to think of the protohominids, of the early primates in the human line, as social carnivores. Their ecologkal relationships would then be similar to those of the social canines, such as wolves. (Bartholomew and Birdsell (1953) have discussed the possible ecology of these protohominids.) As carnivores, they would have been predators on other animals including I am sure many ~:inds of insects-though insects were' probably not an important dietary element, at least after tools and the skills to get large game easily were acquired. Animals in the human line have also probably always had omnivorous tendencies, "scrounging" on all the possible food products that they could digest, such as berries, nuts, and fruits. Honey would fall in this "scrounging" category, and we have direct evidence of honey collecting, at least since the time of the Magdalenian cave paintings. This conceptual environment governs all om attitudes toward other aspects of the world around us, and very largely determines our behavior. The houses we live in are very real parts of our physical environment; but their structure is a consequence of our ideas. This fact applies equally to our clothing, the layout of our towns and cities, our methods of agriculture. Thes~ protohominids would not only get food, they would m turn serve as food for other animals in the community. They would be prey for such things as the big cats and crocodiles; and hosts for many kinds of parasites, including such insects as mosquitoes, lice, and fleas. Man learned early to cope quite successfully with large animals that might prey on him, but it is only re~entIy that he has managed. to avoid, in some degree, bemg food for these smaller msects. At first thought this concept seems remote from insects. But when we pause to consider, we realize that we cannot look at insects, or any other aspect of nature, except through the haze of our conceptual environment. Witchity-grubs are quite different things to the Australian aborigines who use them as a totem, and to the European farmers who regard them as a pest. Termites have one aspect if we look at them as a food delicacy, and another if we look at them as destroyers of wood constructions. House flies gain a new meaning when we see them as possible vectors of disease. This is quite rightly regarded as an advance in knowledge, as a step The aforementioned would about complete the list of basic relationships with other organisms until the time of the Neolithic revolution-until the development of agriculture and settled village life, when a whole new series of relationships started. Most obviously, we have the relations with cultivated plants and domesticated animals, organisms that can be grouped together as "cultigens." \;Yith these man took on a new role, becoming an agent of ~vo]utionary change. The cultigens often became so modified through 68 artificial selection that we can no longer be sure of the nature of their wild ancestors. Some soon became completely dependent on human care for survival. Maize is a classic example-it cannot reproduce without someone to husk the ears and plant the seeds. At least one insect, the silkworm, belongs in the category of "obligate cultigens." When man started to build shelters for himself, he also inadvertently provided shelter for a considerable number of other animals, including many insects. Cockroaches and silverfish moved in on him, along with various rats, mice, lizards, scorpions, spiders, and the like. Such animals can well be cal1ed human inquilinesanalogous with the inquilines of the nests of social insects. Whether things like bed bugs should be called inquilines or parasites is a nice problem of definition. Because they are associated more with the shelter than with an individual host, I think they are better considered to be inquilines. With the clearing of land for villages and fields, man created a kind of situation otherwise rare in nature, an open habitat. It is always surprising to get into a part of the world where agricultural man has had no impact. The common plants and insects of roadsides disappear or can be found only in such open-habitat situations as river sandbars. The weed and insect pests that get along so happily with man simply cannot maintain themselves in a closed community like that of the rain forest. I like to call such organisms "opportunists" because they have taken advantage of the opportunity offered them by human action. Their association with man is much looser than that of the inquilines of his shelters; but like the inquilines and unlike the cultigens, they are unwanted associates. A great many insects surely belong in this category. man's role in the extinction of insect species, since they attract less attention. The clearest case known to me is the disappearance of the butterfly Lycaella dispar from England, which seems to have been caused by the greed of collectors (Ford 1945). I would suspect, however, that many insect species have become extinct on oceanic islands like the Hawaiian archipelago since the advent of civilized man, simply through destruction of the babitat. In a few cases, man has successfully exterminated introduced insect species through planned action (Mediterranean fruit fly in Florida; AlloPheles gambiae in Brazil and Egypt). But I can think of no case in which he has succeeded in deliberately eliminating a native species. The most elaborate attempt has been with Allopheles labrallchiae in Sardinia (Logan 1953), but while tbe species was enormously reduced in numbers, a few specimens were found each year. Probably other kinds of relationships between men and insects could be found. Trying to work out such classifications can become an interesting and instructive game. But here, in closing, I would like to point out two quite different aspects of the human ecosystem-man's gradual withdrawal, with the development of civilization, from association with any particular biological community; and man's tendency to simplify energy exchange within the man-dominated system. Biological communities are ordinarily recognized in terms of food relations-who eats whom. \Vith man's development of trade, transport, and storage, however, this relationship has lost all meaning. If food differs in Saint Louis, New York, London, and Paris, it is for cultural, not biological reasons. Conversely, it is interesting to look at any meal, and speculate about the different parts of the world from which the materials have come. This trend is relatively new, dating at most from the urban revolution that started some 5,000 years ago. The Neolithic village was still associated with a particular biological community-grassland, forest or sea coast. The modern city has lost all such association; the human ecosystem has become a global network. In any general survey of the relations between man and other organisms, we would have to consider pets. These, by my definition, differ from cultigens in that they have not been modified by many generations of breeding under human supervision. This sort of definition has interesting consequences. Dogs and goldfish would have to be classed as cultigens, whereas cacti planted in rock gardens would be pets. Insects have had a limited role as pets in this sense. One thinks of children with rhinoceros beetles or dragonflies tied to strings, or of fireflies used for ornament or il1umination. The Meliponiine bees kept by Indians in tropical America (Bennett 1964) would be pets rather than cultigens; and one can argue about the extent to which the honey bee has become a cultigen rather than a pet. The simplification of energy exchange is more relevant to our present topic. With the development of cooking and especially with the invention of pottery containers for boiling things, man could utilize a wider variety of vegetable materials than before. His digestive system is not adapted for the breakdown of the cellulose walls of plant cel1s; but this breakdown can be effected by cooking, which thus becomes a sort of outside c.'Ctension of the digestive process. Man could then move from the relatively inefficient position of predator to that of firstorder consumer, living directly on plant products. With agriculture, he replaced vegetation he could not use or did not want with vegetation that he could use; he started restructuring the system. With the development of civilization, man has increasingly become an agent of dispersal for other organisms; and an agent of extinction. Charles Elton (1958) has written an excellent summary of man's actions as an agent of dispersal: and here insects are prominent. As every entomologist knows, a large proportion of the important crop pests in different parts of the world have come from somewhere else, inadvertently transported by man. And nowadays purposeful introductions are frequent-attempts to establish predators or parasites of the introduced pests. Where man has remained a predator, a carnivore, he has become a second-order consumer. He raises maize for hogs or grass for cows, and eats the hogs or cows. Only with fish is he a higher-order consumer; and fishing, curiously, has not changed much since the Neolithic; it remains a gathering process. As an agent of extinction, man's actions are easy enough to document for birds and mammals. For each of these he has been the direct or indirect cause of the disappearance, on the average, of one species every year during the present century. It is less easy to document The ordinary biological community is a complex network of organisms using each other for food in all sorts of ways. Any time an ecologist tries to diagram this complex by drawing arrows from the eaters to the eaten, 69 he ends with a blur of criss-crossed lines. The idea of "food chains" in nature has long been abandoned-they are "food webs." But man has reached the point where he will not willingly share his food with anything else; he will not tolerate "worms" in his apples or foxes in his chicken runs. He has set out, in other words, to eliminate competition, to establish short and simple food chains ending with his consumption. He even tries to defeat the decomposing system by burying his dead in metal-lined caskets. ethic. As for esthetics, variety is generally more pleasing than uniformity, and for our own pleasure and edification, we need to preserve as much of the wildlife of the planet as we can. But this is no place to elaborate the arguments for conservation. We cannot go back to the Neolithic, and I doubt whether any of us would want to if we could. Vie are stuck with a very large population and with the need for efficient energy exchange to support this population. But in the long run it is likely that we would be most efficient if we could work in accord with ecological principles. As Francis Bacon remarked a long time ago, "you cannot command nature except by obeying her." We have become so dazzled by our accomplishments that we tend to forget this truism. But nature has been arounc. for a long time, and it may even manage to survive man. It would be fine, though, if man could survive too. This simplification of energy exchange is not only efficient, from the human point of view, but it is necessary, to support the present enormous world population. But it also has dangers. The complex system of the natural community is a stable system, while the simplified human system is much more liable to catastrophe. Insect activities illustrate this fact very well. A disastrous outbreak of a particular species is very rare in a natural community. The whole system of checks and balances operates to keep the population of anyone species well under control. But in the human ecosystem, the large areas of single crops are often liable to destruction by some herbivorous insect multiplying in the absence of the parasites and predators that would ordinarily keep its population in check. Hence the need to be ever alert and to resort to chemical measures of control. The drawbacks and dangers are brought out in other papers of this symposium. REFERENCES 481·-98. Bates, Marston. 1962. The human environment. The HQrace M. Albright Conservation Lectureship, II. Uuiv. California, School of Forestry. Berkeley. 22 pp. Bennett, C. F., Jr. 1964. Stingless-bee keeping in western Mexico. Geographical Rev. 54: 85-92. Elton, C. S. 1958. The Ecology of Invasions by Animals and Plants. John Wiley & Sons. New York. 18: pp., illus. Ford, E. B. 1945. Butterflies. Collins. London. 368 pp. illus. Logan. J. A. 1953. The Sardinian Project. Johns Hopkins Press. Baltimore. 415 pp., illus. Thomas, W. L., Jr. 1956. Man's Role in Changing the Face of the Earth. Univ. Chicago Press. 1193 pp., illus. Elton (1958) has reviewed the evidence that the diversity of species in the natural community tends to promote population equilibrium in each species. This is the utilitarian argument for maintaining as much diversity as possible in the human ecosystem. There are also, as Elton points out, ethical and esthetic arguments. What gives Homo sapie1ls, here for perhaps 50,000 years, and a major force operating on nature for 5,000, the right to destroy a system that has been building up over many millions of years? It can be said that he has the right beca1>lsehe has the power; but in other situations most of us do not consider "might makes right" to be a valid Ecological Considerations Insect Population By E. Associate Professor CITED Bartholomew, G. A., and J. B. Birdsell. 1953. Ecology and the protohominids. Am. Anthropologist 55: in Chemical Control Problems J. LERoux of Entomology, Depart'ment of Entomology and Plant Pathology Macdonald College of JIIIcGill University, Quebec first biologists, forgot they were dealing with a class of animals subject to the general laws of variability and complex interrelations, and forgot their scientific obligation to seek explanations in terms of general conceptsto ask ~ow and why, as well as what and where. The most striking support for this contention lies in the failure of entomologists generally to anticipate the development of resistance to insecticides as a consequence of one of the basic tenets of biology-the principle of natural selection." After 20 years of exposure to the pitfalls attendant on the use of chemicals for pest control, without prior knowledge of their effects on populations, we must as biologists charged with the solution of such problem, make a complete reappraisal of our objectives, The need for a fundamental approach to the study of chemical control of crop pests is more urgently upon us now than at any time in the past. For there is now a great awareness on the part of everyone that the large gaps that exist in our knowledge of pest problems cannot be resolved solely on the basis of the methods of experimentation that have been employed to date. Entomologists, generally, recognize the need for basic investigations but for the most part still pay lip service to this need. ''Vhy is this so? A recent examination of the problem in Canada has revealed, in the words of the reporting chairman (Smallman 1964), that "During the period of preoccupation with the application of control methods, applied entomologists forgot that they were 70