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T. H. C.(1955). Ann. appl. BioZ.

42, 190-196


Deputy-Director, Anti-Locust Research Centre, British Museum (Natural History),
London, S.W.7

The term biological control has been generally used during the last 50 years to
mean the control of an insect or plant pest by means of one or more parasitic or
predatory insects introduced deliberately, but it would be more appropriately used
in a much wider sense to cover all biological methods of control, as opposed to
chemical and physical methods.
One of the main objects of these Jubilee meetings is to survey the achievements of
the last half-century in each of the selected branches of applied biology and outline
present trends and probable future developments. It seems to me that the most
profitable course to pursue in the short time available to me is to consider first
whether the successes in this field during the period under review are such that
biological control, in the narrower sense of the term, can be regarded as a method
that is comparable in general value to chemical methods, in particular; and then to
consider whether biological methods in the widest sense should be developed for
use in the future, in conjunction with chemical measures or as alternatives to them.
It is quite usual, in generalizing on the control of pests, to speak or write as though
there are two main alternative methods available, the biological and the chemical ;
as though, in fact, these two are on a par in general usefulness. This occasion seems
a good one to consider whether this is true.
I do not propose to describe in detail the successful cases of control by introduced
natural enemies. I n the first half of the period under review, there were several
notable instances, and for the present purpose it is sufficient to remind you of the
control of Coccids on citrus in California and elsewhere by Coccinellids, the control
of a Delphacid and a weevil on sugar-cane in Hawaii, the former mainly by a Mirid
and the latter by a Tachinid, and the control of three pests of coconut palms in
Fiji, a Zygaenid moth, a Coccid and a Hispid beetle, by a Tachinid, a Coccinellid
and a Chalcid. In these cases, complete biological control was attained and it has
stood the test of time; the results were so dramatically satisfactory that they caused
entomologists and others of that period to regard the introduction of natural
enemies as the simplest and the most promising method of controlling insect pests
in all parts of the world.
Since that time there have been a few other similarly successful cases, I n the
Seychelles, Coccids on coconut palms have been controlled by Coccinellids, and the
Eucalyptus weevil in South Africa and Mauritius, the European wheat-stem sawfly
in Canada, the white butterfly and the diamond-backed moth on cruciferous crops
Biological control of insect pests 191
in parts of Australia and New Zealand, and the coffee Tortricid in Ceylon, have all
been controlled by Hymenopterous parasites. It must be stressed, however, that the
number of successes has been only a small proportion of the number of attempts
made, and instances of complete, rapid and permanent results such as those obtained
in Fiji and Hawaii many years ago must now be recognized as rare occurrences.
Moreover, experience of the last 30 or 40 years inevitably suggests that most of the
pest-control problems of that period that have been capable of entirely satisfactory
solution by this method have already been solved by it, so that similar successes will
be increasingly rare events in the future. Occasionally, insects will be introduced
accidentally and become established in countries where they did not previously
occur and in conditions such that they can thrive and become serious pests; but
experience indicates that whereas in some of these cases complete control by
natural enemies may be achieved, in most of them it will not.
It is often difficult to obtain from published accounts of the results of introduc-
tions of natural enemies a definite impression as to whether or not the desired
degree of control has been obtained and maintained. The literature of this subject
contains many statements to the effect that particular parasites have been introduced
and successfully established, as though the establishment of natural enemies were
in itself a satisfactory result. References to comparatively high percentages of
parasitization, without corresponding statements about the effect on the pest
populations, are also common, but knowledge of percentage parasitization without
reference to the actual numbers involved is of little or no meaning in relation to
control, since, in particular cases, even 99 yo destruction of a pest population is not
enough to reduce the damage done by the pest to the desired level. Sometimes, no
definite pronouncement is made as to whether the result attained is such as to make
other measures unnecessary, and we are merely informed that a useful degree of
control has been achieved. Such statements cause the reader to conclude that the
results may have been inadequate or merely temporary. I n general, the supporters
of biological control in the narrow sense of the term have unwittingly given the
impression in recent years that they are on the defensive against the supporters of
chemical control and are reluctant to admit the limitations of their method. This
impression may or may not be true, but its existence suggests that an attempt at an
unbiased assessment of the value of the method is worth while.
The instances of biological control that I have so far mentioned are those in which
the control has been so complete that nothing further is needed. There have been
less than twenty cases in this category in our period of 50 years. There is a second
category of cases in which the suppression of the pest has not been so thorough or
extreme, but has nevertheless been so great in parts of the affected area that from
the practical and economic point of view the results can fairly be claimed to be quite
adequate in those parts. The case of the coffee mealy-bug in Kenya is probably in
this category, as is that of the wood wasp in New Zealand and that of the woolly
aphis in various countries. The next category is one in which the method begins to
break down in competition with chemical methods; it comprises the numerous
instances in which introduced enemies achieve some reduction in numbers of the
pest, but not enough to render other measures unnecessary. In most of these cases,
chemical measures which have been in use before the introduction of parasites or
predators must be continued afterwards, so that nothing has been gained by the
introductions. It is sometimes claimed in these circumstances that the parasites or
predators introduced are useful because they supplement other control measures,
but such a contention is unconvincing because modern insecticides, if used at all,
are usually completely effective by themselves so that any supplementary method
is superfluous. Farmers and other producers anxious to prevent or avoid damage
by insects are the first to welcome biological control when it gives the results they
desire because it costs them nothing, whereas chemical control involves a consider-
able outlay annually, but they will use insecticides, despite the annual cost, until
other methods can be shown to result in higher incomes.
We are thus faced with a curious situation. We have in biological control a method
of controlling pests which, at its best, is completely and permanently successful at
no cost whatever except initially, and even the initial cost is relatively small and is
usually borne by a government or a community. Yet, in general, it fails to compete
with chemical methods, even though these involve annual expense, because it only
succeeds in special circumstances which rarely occur. Unfortunately, these circum-
stances are still inadequately understood, so that the limitations of the method
cannot be clearly defined. There are, however, certain features of its past history
which are revealing and important in this connexion. Consideration of the success-
ful cases leads to the conclusions that the method is more likely to succeed in islands
than in continents, and in the tropics than in temperate countries; and conversely,
that it is most unlikely to give satisfactory results in countries like Britain and other
European countries, except in very special cases like that of white-fly in greenhouses.
Supporters of biological control have often argued and still argue that there is no
reason why it should not be as useful in continental conditions as in island condi-
tions. One would like to be able to support this view, but the evidence is too strongly
against it. The oustanding successes have been in islands and good results obtained
elsewhere have in the main been in areas which are comparable to islands in some
respects, notably in that they are isolated by natural barriers and are limited and
specialized in flora and fauna, as are parts of Australia, California and Kenya. The
principal reason suggested for the greater success of introduced natural enemies in
island conditions is that the relatively small fauna contains few or no species likely
to attack any beneficial insect that may be introduced. The greater degree of success
obtained in the conditions of tropical islands than in those of temperate countries
is almost certainly due at least partly to the equable climate of such islands, which
enables the introduced insects to breed continuously throughout the year and there-
fore to multiply very rapidly. Seasonal conditions unfavourable to breeding, such
as winters in temperate countries and long periods of hot, dry weather in the
Biological control of insect pests '93
continental tropics, seem to increase the hazards to which introduced species, not
yet adapted to the climate, are exposed.
While it is possible to suggest these reasons for the special suitability of tropical
islands for biological control, they are somewhat theoretical and are not sufficiently
precise. In most attempts at biological control, parasitic or predatory insects,
themselves little known ecologically, are introduced into conditions which are also
inadequately known. There are only a few cases in which efforts have been made,
by studying the pest in the country and in the conditions in which the control is
required, to obtain in advance an idea of the sort of parasite or predator that might
be expected to achieve satisfactory results, but in most cases the selection of natural
enemies to be introduced has been made largely without fundamental investigation.
The need for basic studies is probably greatest when the insect to be controlled has
become a pest in the country in which it is indigenous, through a modification of
its environment or the advent of some factor which renders the indigenous natural
enemies that have previously kept it in check unable to do so any longer. In these
circumstances, the parasites from which a choice is to be made must be obtained
from a foreign host species other than that which is to be controlled and most
parasite species could not succeed where several indigenous parasites 'have failed.
Examples of indigenous insects which became pests in their own country and have
been completely controlled are the coconut Hispid beetle and the coconut Zygaenid
moth, both in Fiji; the former was controlled by a Chalcid which was selected on
the basis of considerable knowledge, and the coconut moth (which, incidentally,
was thought at the time to have been accidentally introduced into Fiji from an
unknown overseas source but is now believed to be indigenous) was controlled
by a Tachinid, the selection of which involved a considerable element of good
The control in tropical islands of introduced pests is less dependent as a rule on
thorough fundamental studies, the requirements which the natural enemy that is to
be introduced must fulfil being less rigid, and the choice therefore wider. However,
even in tropical islands, special circumstances sometimes exist which make the
control of particular introduced pests by this method difficult or impossible.
A recent example is the case of the Coccids that have threatened to destroy the
Bermuda Cedar in Bermuda. Thorough investigation of this problem has led to the
conclusion that the tree is so susceptible to the Coccids that even a very low popula-
tion of them, such as would normally be regarded as negligible, is very harmful, and
the natural enemies introduced cannot suppress the Coccids to the still lower level
required. A similar difficulty arises in the control of insects that are harmful as
virus vectors, the degree of suppression needed in such cases being as a rule so
great as to be beyond the capabilities of natural enemies; an example is afforded by
the mealy-bug vectors of swollen-shoot disease of cacao in West Africa.
I n the least complicated environments, such as those of tropical islands, careful
study of a problem sometimes leads to successful control of the pest by means of
13 App. Biol. 42
a parasite that would probably not otherwise have been selected, but it must be
admitted that it is never possible to predict with certainty the result of any particular
introduction. In continental areas, the factors concerned in the establishing and
multiplication of insects are so numerous and complex that prediction of results is
useless and any attempt at the utilization of natural enemies can be little more than
a process of trial and error. Numerous attempts to put biological control work of
this kind on a mathematical basis have been made from time to time, but except in
the simplest and most straightforward instances they have failed. I n this con-
nexion, it is worth noting that the late G. C. Ullyett, who made a special study of
insect populations in relation to biological control by natural enemies, stated in
a recently published paper entitled Biomathematics and Insect Population
Problems that mathematics has not provided us with a more exact method of
expressing our major problems and that its use in biology is, in general, extremely
limited. He went on to say that Present developments in mathematics are not
adequate to deal with the complex situations found in Nature. . . . These statements
are perhaps a little too severe when removed from their context, but in the main
they are true when applied to natural enemies, and they serve to emphasize the
difficulties with which attempts to make biological control more precise are faced.
I can best summarize my views on this sort of biological control by saying that
I know it to be the best of all methods of controlling pests when it works, but that
it seldom works and that there is little future for it in continental areas. I think that
the present tendency to organize the moving of parasites and predators about the
world on an ever-increasing scale, despite decreasing results, is unsound and is,
therefore, to be regretted. The limitations of the method are inherent in it and will
not be changed by organization. Moreover, attempts to use it in combination with
chemical methods are, in general, foredoomed to failure, biological and chemical
methods being incompatible. At the same time, being, I suspect, one of the few
people in this country who have witnessed the spectacularly rapid, thorough and
permanent control that can result from the introduction of natural enemies, with
little initial cost and no subsequent cost, I am a strong supporter of it, and if my
criticism of the present trends seems destructive, it is only because I am anxious to
give a true picture of the situation as I see it.
If we admit, as I think we must, that biological control, in the present narrow
sense of the term, is not going to solve many of our problems, then we must con-
sider whether any other methods that can be termed biological are likely to be used
to better advantage. This brings us into the field of ecological control, or control
based on thorough knowledge of the ecology and population dynamics of the insects
to be controlled. Here we are faced with the inescapable difficulty that an under-
standing of the ecology of any one species can be gained only by prolonged and
intensive quantitative studies over a long period of years, and this is hardly practicable
for all the species of insects that we wish to suppress. Nevertheless, there can be no
Mem. ent. Sot. S.Afr. no. 2, 1953.
Biological control of insect pests I95
doubt that the ecological approach is the right one and that chemical methods should,
in general, be regarded as stop-gaps which give control during the time needed for
ecological research. If the major problems in applied entomolbgy can be shown to
be solvable by ecological means, then some at least of the others will be similarly
solved in due course. The striking results obtained in recent years with viruses for
the control of the Alfalfa Butterfly and the Japanese Beetle in the United States are
an encouraging example of what can be achieved by patient, fundamental research
over a long period.
Most of you will have read the recent excellent paper by Solomon" entitled
'Insect Population Balance and Chemical Control of Pests'. He concludes that
entomologists should do all they can 'to foster the development of research and
control measures which have an adequate ecological basis '. There is no need for me
to repeat his arguments here. My object is to emphasize that the days of empirical
biological control are past and that a new approach, based on fundamental biological
studies of the insects themselves and the factors that cause their numbers to
fluctuate, is essential. Natural enemies constitute just one of the many ecological
factors that affect insect numbers and they are not necessarily the most important;
in those cases in which the natural enemies are not known to be important in the
normal ecology, it is hardly reasonable to move them about the world for control
There is already a widespread awareness of the need for more basic studies of
insect populations in relation to the control of harmful species, and this has arisen
not only from the discoveries that strains of insects resistant to insecticides can
arise and that chemicals applied for the control of one pest can cause outbreaks of
others, but also from a general feeling of uneasiness about the ultimate, accumulated
consequences of the general application to the soil and to plants of highly toxic
insecticidal chemicals. We are at present passing through a period of confusion and
conflicting interests in which most of us are eager to be able to advocate ecological
rather than chemical measures, but in the present state of knowledge the supporters
of ecological measures often have none that they can advocate with sufficient
conviction. This is an inevitable interim condition which only research and time
can end. Some of the most formidable of the world's problems in applied ento-
mology, such as those of locusts and tsetse flies, are at present being tackled by a
combination of chemical and ecological measures which is to be regarded as only
a means to a more satisfactory end. The outbreaks of mites and Coccids that have
resulted from the general use of modern insecticides on fruit trees have had the
advantage that they have caused intensive studies to be made in Canada, the United
States and Britain of the whole complex of animal species inhabiting the trees, and
while growers will continue to use insecticides and acaricides so long as they obtain
a bigger financial return by doing so, ecological information which will probably be
of great value ultimately is being obtained meanwhile. How such information will
* Chemistry & Industry, 1953, pp. 1 1 4 3 7 .
196 T. H. C . TAYLOR
eventually be used we cannot yet judge, but the difficulties that have already arisen
from the use of modern insecticides suggest that the time may come when it is
generally felt that it is better in the long run to use ecological methods to maintain
all the species in a particular association at a reasonably low population level than
to use chemical methods to reduce the most harmful species to a much lower level.
General acceptance of the advantages of such a policy cannot come until the funda-
mental knowledge needed to support it is available. Our aim should be to ensure
that the ecological conditions that we create are such that potentially harmful
animal species are not unduly favoured and that we have the knowledge necessary
for modification of the environment of insects wherever this is desirable for their
control in the interests of our health or economy. When this happy state of affairs
is reached, insecticidal chemicals will probably still be needed occasionally, but
they will be used, not as part of a seasonal routine, but as a means of reducing
particular populations that might arise in exceptional circumstances.
Meanwhile, we in Britain can derive consolation from the thought that so long
as we maintain our present agricultural and horticultural practice of growing a great
variety of crops in a mosaic of fields and small plots separated by hedgerows and
uncultivated areas, our entomological problems will rarely be severe in comparison
with those of tropical countries and vast continental areas. We already have a
system which gives us a relatively high degree of ecological control of insects, even
though we do not yet fully understand how it works. In the still developing regions
of Africa and Asia, on the other hand, development will almost certainly continue
to create environments in which grave new entomological problems will arise, and
it is in these continents particularly that control of insect populations by methods
based on thorough knowledge of their biology and ecology is likely to be most