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Introduction 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. Decline of Biological Control in the University of California The biological control unit at the new campus of the University
of California in Riverside was by 1962 the most renowned research entity for that discipline in the world. It served as a World Center 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 resided at facility in Central
California at Albany, just five miles north of 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 more than 300 scientists engaged in Classical
Biological Control (= The search for, importation and propagation of
new species of natural enemies). This
does not include investigators 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 biology 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
unit. 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 faculty against one Dr. Robert van
den Bosch <PHOTO>, who was very vociferous in denouncing what he
perceived to be administrative inadequacy.
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. A pervasive gloomy atmosphere followed, 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. The feud had far reaching consequences in the University of
California that persisted into the latter part of the 20th Century. One Riverside professor who had sympathy
for the Albany group tried on two occasions to have a faculty member fired
whom the Riverside group had supported.
This even though his first attempt presumably banned him from the
fellow's promotion committee. Another
especially malevolent incident involved a Korean graduate student, where a
junior member of the Qualifying Committee who had been a student at Albany
contrived a scheme to deny the Korean student his PhD Degree. Teaming up with another faculty of the
Biology Department the two failed the student and refused to grant him a
second Qualifying Examination even though other members of the committee
deemed his performance on the examination excellent. The incident was especially illogical and
sordid because the two dissenting faculty had given the student high passing
grades in their courses, and no indication of inadequacy was ever made to the
major professor whom they obviously despised for his support of Dean Al Boyce
in the earlier Interdepartmental conflicts.
The Korean student had gained high grades in all his courses and
established an excellent rapport among Public Health organizations in
California while performing his thesis research. As a credit to the integrity of the University of California,
wisdom prevailed as the two dissenters were removed from the student's
committee. He gained the PhD Degree
following a successful reexamination and later became Head of the Department
of Public Health in Seoul, Korea. 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 that parasitized
olive scale in Pakistan and Iran. He
also maintained that he had made the original discovery of Aphytis melinus
that attacks red scale (Aoidinella auranti) in Pakistan, although it
is unclear whether he was able to send a viable culture to California
[Personal communication to Dr. E. F. Legner]. The living cultures that he did
obtain from that region that was typically undergoing intense political unrest
exposed him to "a hail of bullets" as he once described to Dr. E.
F. Legner. Yet, not one mention of
his involvement in the discovery or acquisitions of these parasitoids was
ever made by DeBach or his associates who later were credited with their
discovery. A disregard of the honor
process among scientists in recognizing each other's 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 Riverside Campus President who had the power to do a coup de graz. It may be debated that his professional
background in Political Science 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, Dr. 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 highly
sophisticated and time-consuming research.
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 to control
agricultural pests. Later the
Division itself was abolished through the intense efforts of Dr. 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
on the noxious, beneficial and other insects of the state of New York. Albany, New York. 259 p. Goedaert, J. 1662. Metamorphosis et Historia Naturalis
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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western Texas and southeastern New Mexico.
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739-743. Malthus, T. R. 1803. An Essay on the Principle of Population as
It Affects the Future Improvement of Society. J. Johnson, London, 2nd ed.
610 p. Pasteur, L. 1870 Etudes dur la maladie des vers a
soie. Gautherie-Villars, Paris,
I: 322 p.; II: 327 p. Ratzeburg, J. T. C.
1944a. Die Ichneumonen der
Forstinsekten in forstlicher und entomologischer Beziehung; ein Anhang zur
Abbildung und Beschreibung der Forstinsekten. Theile, Berlin. 3 vol. Ratzeburg, J. T. C.
1944b. Die Ichneumonen der
Forstinsekten, Vol. I. Berlin. Réaumur, M. de.
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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
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