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The recorded history of biological control may be considered as dating
from Egyptian records of 4,000 years ago, where domestic cats were depicted
as useful in rodent control. Insect Predation was recognized at an early date, but the significance of
entomophagy and exploitation was lost except for a few early human populations
in Asia where a sophisticated agriculture had developed. The Chinese citrus growers placed nests of
predaceous ants, Oncophylla smaradina, in trees where the
ants fed on foliage-feeding insects.
Bamboo bridges were constructed to assist the ants in their movements
from tree to tree. Date growers in
Yemen went to North Africa to collect colonies of predaceous ants which they
colonized in date groves to control various pests. Insect Parasitoidism was not
recognized until the turn of the 17th Century. The first record is attributed to the Italian, Aldrovandi (1602). He observed the cocoons of Apanteles glomeratus being attached to larvae of Pieris rapae (the imported cabbageworm). He incorrectly thought that the cocoons
were insect eggs. Printed
illustrations of parasitoids are found in Metamorphosis by J. Goedart (1662) <PHOTO>. He described "small flies"
emerging from butterfly pupae. Antoni van Leeuwenhoek in 1700 (van Leeuwenhoek 1702) described
the phenomenon of parasitoidism in insects.
He drew a female parasitoid ovipositing in aphid hosts. Vallisnieri
(1706) <PHOTO> first
correctly interpreted this host-parasitoid association and probably became the
first to report the existence of parasitoids. Bodenheimer (1931),
however, noted that several earlier entomologists recognized the essence of
parasitoidism. Cestoni (1706) reported other parasitoids from eggs of
cruciferous insects. He called aphids,
"cabbage sheep," and their parasitoids, "wolf
mosquitoes." Erasmus Darwin (1800) discussed the
useful role of parasitoids and predators in regulating insect pests. During the remainder of the 18th Century an ever-increasing
number of references to entomophagous and entomogenous organisms appeared in
the literature, largely in the form of papers dealing with parasitoid
biologies. Diseases of silkworms were
recognized early in the 18th Century.
De Reamur (1726) <PHOTO> described
and illustrated Cordyceps
fungus infecting a noctuid larva. Biological Control Efforts in the 18th Century By 1762 the first successful importation of an organism from one country
to another for biological control took place with the introduction of the mynah bird from
India to the island of Mauritius, for locust control. Further development of modern biological control awaited the
recognition of the fact that insect pest problems were population
phenomena. The controversial
publications of Malthus
appeared toward the end of the 18th Century, and generated considerable
interest in the subject of populations.
Malthus' work will be discussed further in the next section on "Concepts
in Population Ecology." Biological Control Efforts in the Early 19th
Century A number of articles appeared during the first half of the 19th
Century that lauded the beneficial effects of entomophagous insects. Erasmus Darwin (1800) recommended protecting and encouraging
syrphid flies and ichneumonid wasps because they destroyed considerable
numbers of cabbage-feeding caterpillars.
Kirby & Spence (1815) [see <PHOTO>] showed that predaceous coccinellids controlled aphids. Hartig (1827) recommended the construction of large
rearing cages for parasitized caterpillars, with the ultimate aim of mass
release. Ratzeberg (ca. 1828) <PHOTO> called particular attention to the value of parasitic
insects with publication of a large volume on the parasitoids of forest
insects in Germany. He did not
believe that parasitic control could be augmented by humans. Agustino Bassi (1834) first demonstrated that a
microorganism, Beauvaria bassiana, caused an animal
disease, namely the muscardine
disease of silkworms. Kollär <PHOTO> (1837) writing an
article for farmers, foresters and gardeners pointed out the importance of entomophagous
insects in nature's economy; studied parasitoid biologies and was the first
to report the existence of egg parasitoids.
Boisgiraud
(1843) reported that he used the predaceous carabid beetle, Calasoma sycophanta, to successfully control gypsy moth larvae on
poplars growing near his home in rural France. He also reported that he had destroyed earwigs in his garden by
introducing predaceous staphylinid beetles. Biological Control in the Late 19th Century Beginning in 1850, events associated with the westward expansion
of agriculture in the United States paved the way for the further development
of the field of biological control.
During and following the "Gold Rush" in California,
agriculture expanded tremendously in California especially. At first the new and expanded plantings
escaped the ravages of arthropod pests.
Predictably, however, crops soon began to suffer from destructive
arthropod outbreaks. Many of these
pests were found to be of foreign origin, and were observed to be far more
destructive in the newly colonized areas than in their native countries. Consequently, the notion grew that perhaps
these pests had escaped from some regulatory factor or factors during their
accidental introduction into America. Asa
Fitch <PHOTO> (1855) was the State
Entomologist of New York who is recorded as the first entomologist to
seriously consider the transfer of beneficial insects from one country to
another for the control of an agricultural pest. Fitch suggested that the European parasitoids of the wheat
midge, Sitydiplosis mesellana, be sent into the
eastern United States. Benjamin
Walsh <PHOTO> supported
Fitch's suggestion and in 1866 he became the first worker in the United
States to suggest that insects be employed in weed control. He proposed that insects feeding on
toadflax, Linaria vulgaris, be imported from
Europe to control invaded yellow toad flax plants. The first actual case of biological control of weeds was,
nevertheless, in Asia, where around 1865 the cochineal insect Dactylopius ceylonicus was introduced from
southern India into Ceylon for prickly pear cactus control (Opuntia vulgaris).
Originally, Dactylopius
had been imported to India from Argentina in 1795, in the mistaken belief
that it was the cochineal insect of commerce, D. cacti. Louis
Pasteur (1865-70) <PHOTO> studied silkworm diseases and saved the
silk industry in France from ruin [not really biological control]. Charles Valentine Riley <PHOTO> (1870) has been named the father of modern
biological control. He shipped
parasitoids of the plum curculio from Kirkwood, Missouri to other parts of
that state. In 1873 he became the
first person to successfully transfer a predator from one country to another
with the shipment of the American predatory mite, Tyroglyphus phylloxerae
to France for use against the destructive grapevine phylloxera. The results were not particularly
successful, however. In 1883, Riley
directed the first successful intercontinental transfer of an insect parasitoid,
Apanteles glomeratus, from England to the
United States for control of the imported cabbageworm. He was Chief Entomologist of the U. S.
Department of Agriculture. In 1872,
11 years before the importation of A.
glomeratus, Riley began his
interest in the cottony-cushion scale, Icerya
purchasi, which was
considered the most important citrus pest in California. He correctly located its point of origin
in Australia. [Doutt's account of
this biological control program on p. 31-38 of the DeBach (1964) text is
particularly colorful. Read this,
paying particular attention to the following: a. the roles played by
Riley, Albert Koebele and D. W. Coquillet. b. note the species of
insects involved (the vedalia beetle, Rodolia
cardinalis, and the
dipterous parasitoid, Cryptochaetum
iceryae), their source,
numbers imported, and their activities relative to the cottony-cushion scale. c. note the method of
colonization, and be able to describe the spectacular results of these
introductions, which changed the status of the pest to an insect of no
economic importance in only four years time. The successful biological control effort against the
cottony-cushion scale spirited many biological control attempts in many
countries, resulting in over 200 biological control successes (see Chapter 24
of the DeBach (1964) text and other hand-outs). The cottony-cushion scale success admittedly harmed overall pest
control in California for quite some time because growers thought that the
vedalia beetle would also control other insect pests. Consequently, they neglected other
mechanical and chemical control methods. George Compere (1899) became the first
state employee specifically hired for biological control work. He worked as a foreign collector until
1910, during which time he sent many shipments of beneficial insects to
California from many parts of the world.
Harold Compere <PHOTO>, his son, also devoted his entire career to the search for
and identification of natural enemies of scale insects. Harry Scott Smith (1913)
<PHOTO> was appointed
superintendent of the State Insectary in Sacramento. In 1923, biological control work was
transferred to the Citrus Experiment Station and Graduate School of
Subtropical Agriculture of the University of California, Riverside. Biological control work at Riverside was
first conducted in the Division of Beneficial Insect Investigations, and was
changed to the Division of Biological Control with Smith as chairman in
1947. Personnel were stationed at
Albany and Riverside. Under Smith,
importation of Chrysolina
beetles from Australia for Klamath weed control marked the beginning of
biological weed control in California in 1944. Edward Steinhaus (1947) <> established
the first laboratory and curriculum in insect pathology at the University of
California, Berkeley. Later he
transferred to the newly opened Irvine campus of the University and attempted
to further insect pathology there.
His untimely death in 1968 precluded this goal. The Division of Biological Control became the Department of
Biological Control at UC Riverside and Berkeley in 1954. In 1969 Biological Control was dropped as
a department, becoming a Division of Biological Control within the Department
of Entomology, against the wishes of the entire biological control faculty,
numbering over 24 academics at Riverside and Berkeley at that time. The Berkeley faculty created their own
separate Division of Biological Control with guaranteed privileges and
minimum control by the Department of Entomology. At Riverside, the Division of Biological Control gradually
became dominated by chemical control oriented faculty in the Department of
Entomology. In 1989 the Division was
abolished, against the wishes of 85% of the faculty in the Division. Ignorance and pecuniary control among the
ranks of University of California bureaucrats is believed to be the principal
cause. Although the dissenting
faculty in the Division each wrote a personal plea to the then Chancellor
Rosemary S. J. Schraer to discuss the matter, in not one case was a reply
received. Breakdown of the University of California Biological Control Department The biological control unit at the new campus of the University
of California in Riverside was by 1962 the most renouned research entity for that discipline in the world. It served as a mecca for students and
scientists devoted to the practice of classical biological control, where
natural enemies were sought worldwide for importation and establishment. Although this unit's headquarters was at
Riverside, about 1/3rd of the faculty worked out of a facility in Central
California at Albany, just five miles from the UC-Berkeley campus. By 1961, UC-Riverside and the Albany
facility had a total of about 18 full-time professional biological control
faculty plus several emeriti; about 10 research associates, and graduate
students that varied from 10-20 until the 1980's. Members of this statewide department interacted with other
similar organizations in various parts of the world, especially the
Commonwealth Institute of Biological Control, that had established
laboratories worldwide, and the U. S. Department of Agriculture. On the world
scene, it is estimated that there were over 300 entomologists engaged in
classical practical biological control work.
This does not include scientists engaged only in fundamental
research. The harmony amongst these
scientists was exceptional and admired, and was spirited most likely from a
realization that cooperation accelerated achievements in a field that
required extensive knowledge of
arthropod biologies and breeding habits. However,
a feud developed among some of the top administrators in the University of
California and within the Department of Biological Control itself that
ultimately contributed to the demise of this outstanding organization. The basis was involved, but especially
referred to unprofessional conduct, the hiring of new faculty that was not
supported by a majority of the Department, and animosities developed in
previous years when current administrators had previously served as technical
staff. The then Dean of Agriculture,
Dr. Alfred Boyce, operating through departmental administrators, organized a
voting block among the younger naive faculty against one Dr. Robert van den
Bosch <PHOTO>,
who was very vociferous in denouncing what he perceived to be administrative
sleaze. This ultimately led to van
den Bosch being asked to leave the unit, especially after his rebellion
extended to a denunciation of the scientific integrity of several active
faculty. This produced a general
gloomy atmosphere, especially among the newly hired faculty. Van den Bosch left the Riverside campus
for the branch laboratory in Albany, California, taking with him some highly
skilled technical personnel, and a while later a new faculty member, Dr.
George Poinar, Jr. Other faculty and
staff at Riverside that were disturbed by the politics of these events then
joined the separate Department of Entomology. Dr. Boyce later expounded on matters that further revealed more
of the nature of the feud (Boyce 1997/98 and personal communications). He was especially distraught when Dr. Paul
DeBach <PHOTO>
and associates at Albany ignored his contribution to the discovery of the
citrus red scale parasitoids, Aphytis maculicornis and Coccophagoides
utilis parasitizing olive scale in Pakistan and Iran. Boyce had obtained the original cultures
himself from that region that was typically undergoing intense political
unrest. He was able to bring out
living cultures "in a hail of bullets" as he once described it to
Dr. E. F. Legner. Yet, not one
mention of his involvement in the acquisition of this important parasitoid
was ever made by DeBach or his associates.
A disregard of the honor process among scientists in recognizing each
others contributions may have far reaching effects. Yet these failures continue and may be widespread as shown by
the recent description of Biosteres
sublaevus Wharton that ignored mention of original specimens donated
from years of effort in securing them from the wild (Legner & Goeden 1987 ). Being weakened as it was, by the loss of highly capable and
productive scientists, and lacking in political adeptness, the Riverside unit
fell victim to the one politician who had the power to do a coup de graz. It may be debated that his professional background
certainly did not justify his making unilateral decisions concerning the
Biological Control discipline. By this time, DeBach had become
dismayed at the politics and rather accepted the final triumph of the Al
Boyce lobby. DeBach, because of his
international renown in the field of Biological Control, should have been the
logical choice to lead the Department as Chairman. However, against the wishes of most of the faculty, Boyce
hired Dr. Don Chant of Ontario, Canada to head the Department. Chant had a very positive influence on the
younger faculty especially by helping them to attain the research funds that
are needed to do this work. However,
he then gradually became increasingly dismayed at the politics of the higher
administration and after three years returned to Canada to head the
Department of Zoology at the University of Toronto. Boyce then, against the wishes of the entire faculty, unilaterally abolished the Department of
Biological Control, and forced it to reorganize as a subsidiary Division of
Biological Control within the Department of Entomology, that was on the whole
devoted to the use of pesticides.
Later the Division itself was abolished through the intense efforts of
Boyce and against the objection of 90% of its faculty. In the meantime, the Albany faculty
continued relatively autonomous from the pesticide-oriented fraction, but
ultimately lost critical numbers who were devoted solely to the classical
biological control approach. Another contributing factor to
Riverside’s decrease in classical biological control activity is related to a
reduced ability to interact with professionals overseas. To illustrate this it should be considered
that classical biological control successes have relied heavily on the
interaction with other international organizations, especially the
Commonwealth Institute of Biological Control with headquarters in Curepe,
Trinidad. Various permanent and
temporary laboratories of this organization existed in all parts of the
world. Researchers there would host,
assist and otherwise interact with those of the United States Department of
Agriculture and the University of California to obtain beneficial
species. As independence from the
British Commonwealth developed among the different countries that maintained
laboratories, local support for their continuance diminished, and in many
cases ceased entirely. This has
resulted in a greater than 90% decrease in classical biological control
activity worldwide. Strange Mortalities of Biological Control Specialists There have been four known suicides among
the ranks of biological control scientists.
These were Owen Smith and Irv Newell of the Untied States, Giuseppii
Zinna of Italy, and David Annecke
<PHOTO>
of South Africa. Smith was found by
technician Louis Dawson, hanging from a tree in the biological control orange
grove on the University of California, Riverside campus. This just after his success in classical
biological control of the grape leaf skeletonizer. The caterpillars possess urticating hairs that can interfere
with the health of persons in close contact with them. Zinna had just been hired by the Division
of Biological Control in Riverside as chief systemiatist: the position that was later filled by Gordon
Gordh. Zinna returned to Italy,
presumably to gather his personal effects, when he, unprovoked, jumped from an eight-story building. Annecke killed himself in South Africa,
also without known provocation.
Newell killed himself with a shotgun at his home in Riverside. Rumors were that he suffered from cancer,
but he also was known to suffer constant severe pain in the facial area,
which may have been an allergic reaction to the mites with which he so diligently
worked. Robert van den Bosch died
from a heart attack while jogging in the Berkeley, California area. He had been ardently pursuing the
Pesticide Industry (van den Bosch 1978) for unscrupulous activities in pest
control, gaining the animosity of many dedicated to chemical pest
control. There seemed to be no
generally known history of cardiac illness.
Paul Messenger, who took up the struggle against pesticides after van
den Bosch's death, also died mysteriously from a heart attack. Blair Bartlett died in his hospital bed
immediately after having heart bypass surgery in San Bernardino,
California. He had been studying the
effects of pesticides on beneficial organisms, and just concluded that almost
all available materials had severe detrimental effects on an array of species
in many important families (Bartlett 1964*-- 1966). Harry Shorey, working with insect pheromones as a substitute
for chemical insecticides in the Coachella Valley, was killed when the
automobile that his student was driving collided with a truck transporting
produce from Mexico. Exercises
Exercise 2.1-- When was the importance of insect
predation first recognized? Insect
parasitoidism? Exercise 2.2-- Trace the development of biological
control in the 18th, 19th and 20th centuries. Exercise 2.3-- Give the past and present organization
of biological control research in the University of California. REFERENCES: [Additional references may be found at MELVYL
Library ] Bartlett, B. R. 1964a.
Integration of chemical and biological control, p. 489-511. In: P. DeBach (ed.), Biological Control of
Insect Pests and Weeds. Reinhold, New
York. Bartlett, B. R. 1964b.
The toxicity of some pesticides to eggs, larvae, and adults of the
green lacewing, Chrysopa carnea. J. Econ. Ent. 57: 366-9. Bartlett, B. R. 1964c.
The toxicity of some pesticide residues to adult Amblyseius hibisci,
with a compilation of the effects of pesticides upon phytoseiid mites. J. Econ. Ent. 57: 559-63. Bartlett, B. R. 1965.
The repellent effects of some pesticides to hymenopteraous parasites
and coccinellid predators. J. Econ.
Ent. 58: 294-96. Bartlett, B. R. 1966.
Toxicity and acceptance of some pesticides fed to parasitic
Hymenoptera and predatory coccinellids.
J. Econ. Ent. 59: 1142-49. Bassi, A. 1935. Del mal del
segno, calcinaccio o moscardino, mallatia che affigge i bachi da seta e sul
modo di liberarne le bigattaie anche le piu infestate. Part I:
Theoria. Orcesi, Lodi. p. 1-9,
1-67. Bellows, T. S., Jr. & T. W.
Fisher, (eds) 1999. Handbook of Biological Control: Principles and Applications. Academic Press, San Diego, CA. 1046 p. Bodenheimer, F. S. 1931.
Der Massenwechsel in der Tierwelt.
Grundriss einer allgemeinen tierischen Bevölkerungslehre. Arch. Zool. Ital. (Napoli) 16: 98-111. Boyce, A. M. 1987. Odyssey of an Entomologist. Kingsport Press, Kingsport Tenn. pg 213-219.
Boyce, Alfred Mullikin.
1997/1998. Odyssey of an
Entomologist: adventures on the farm, at sea, and in the university / by
Alfred M. Boyce ; based on taped conversations with John G Gabbert ; edited by Elizabeth Lang and
Robert Lang. Riverside, Calif. : UC
Riverside Foundation, 1987, c1986. Compere, G. 1902. Entomologist's Report. Introduction of Parasites. West. Austral. Dept. Agric. J. 6: 237-40. Compere, G. 1904. Black scale parasite (Scutellista cyanea). West Austral. Dept. Agric. J. 10: 94. Compere, G. 1921. Seasonal history of black scale and
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Calif. Citrog. 6: 197. Darwin, E. 1800. Phytologia. Publ., London. Doutt, R. L. 1964. The historical Development of biological
control. In: P. DeBach (ed.),
Biological Control of Insect Pests and Weeds. Reinhold Publ. Corp., New York. 844 p. Fitch, Asa. 1954. Sixth, seventh, eighth and ninth reports
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Insectorum. Jacques Fierens,
Middelburgh. Kirby, W. &
W. Spence. 1815. An Introduction to Entomology. Longman, Brown, Green & Longmans,
London. 285 p. Kollär, Vincent.
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It Affects the Future Improvement of Society. J. Johnson, London, 2nd ed.
610 p. Pasteur, L. 1870 Etudes dur la maladie des vers a
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1726. Remarques sur la plante
appellée a la Chine Hia Tsao Tom Tchom, ou plante ver. Mem. Acad. Roy. Sci. (21 Aug 1726). p. 302-5. Riley, C. V. 1893. Parasitic and predaceous insects in
applied entomology. Insect Life
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insects. Science 73: 475-6. Smith, H. S. 1916. An attempt to redefine the host
relationships exhibited by entomophagous insects. J. Econ. Ent. 9:
477-86. Smith, H. S. 1919. On some phases of insect control by the
biological method. J. Econ. Ent.
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