I. 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.
II. Insect Predation
A. 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.
B. The Chinese citrus growers since at least 400 BC 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.
C. Date growers in Yemen went to North Africa to collect colonies of predaceous ants which they
colonized in date groves to control various pests.
III. Insect Parasitoidism
A. Not recognized until the turn of the 17th Century.
B. First record 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.
C. Printed illustrations of parasitoids are found in Metamorphosis by J. Goedart (1662) <PHOTO>.
He described "small flies" emerging from butterfly pupae.
D. Antoni van Leeuwenhoek in 1700 (van Leeuwenhoek 1702) described the phenomenon of parasitoidism
in insects. He drew a female parasitoid laying eggs in aphid hosts.
E. Vallesnieri (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.
F. Cestoni (1706) reported other parasitoids from eggs of cruciferous insects. He called aphids, "cabbage
sheep," and their parasitoids, "wolf mosquitoes."
G. Erasmus Darwin (1800) discussed the useful role of parasitoids and predators in regulating insect pests.
IV. 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.
A. Diseases of silkworms were recognized early in the 18th Century.
B. De Reamur (1726) described and illustrated Cordyceps fungus infecting a noctuid larva.
V. Biological Control Efforts in the 18th Century
A. 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
B. 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."
VI. 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.
A. Erasmus Darwin (1800) recommended protecting and encouraging syrphid flies and ichneumonid
wasps because they destroyed considerable numbers of cabbage-feeding caterpillars.
B. Kirby & Spence (1815) <PHOTO> showed that predaceous coccinellids controlled aphids.
C. Hartig (1827) recommended the construction of large rearing cages for parasitized caterpillars, with the
ultimate aim of mass release.
D. 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.
E. Agustino Bassi (1834) first demonstrated that a microorganism, Beauvaria bassiana, caused an animal
disease, namely the muscardine disease of silkworms.
F. Kollär (1837) <PHOTO> 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.
G. 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.
VII. 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.
A. Asa Fitch (1855) <PHOTO> 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.
B. 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 toad flax, 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,
C. Louis Pasteur (1865-70) <PHOTO> studied silkworm diseases and saved the silk industry in France from
ruin [not really biological control].
D. Charles Valentine Riley (1870) <PHOTO>.
1. The father of modern biological control.
2. He shipped parasitoids of the plum curculio from Kirkwood, Missouri to other parts of that state.
3. 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.
4. 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.
5. 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:
b. note the species of insects involved (the vedalia beetle, Rodolia cardinalis <PHOTO>, and the dipterous
parasitoid, Cryptochaetum iceryae), their source, numbers imported, and their activities relative to the cottony-
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.
VIII. 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
A. 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.
B. 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.
C. Harry Scott Smith (1913) <PHOTO>.
1. He was appointed superintendent of the State Insectary in Sacramento.
2. In 1923, biological control work was transferred to the Citrus Experiment Station and Graduate School of
Subtropical Agriculture of the University of California, Riverside.
3. 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.
4. Importation of Chrysolina beetles from Australia for Klamath weed control marked the beginning of biological
weed control in California in 1944.
5. Edward Steinhaus (1947) <PHOTO> 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.
6. The Division of Biological Control became the Department of Biological Control at UC Riverside and Berkeley
7. 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.
8. 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.
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.
Bodenheimer, F. S. 1931. Der Massenwechsel in der Tierwelt. Grundriss einer allgemeinen tierischen Bevölkerungslehre. Arch. Zool. Ital. (Napoli) 16: 98-111.
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 relation to biological control. Calif. Citrog. 6: 197.
Darwin, E. 1800. Phytologia. Publ., London.
DeBach, P. 1974. Biological control by natural enemies. Combridge Univ. Press.
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. 1837. In: London's Gardner's Magazine. 1840. [English translation].
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. 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 6: 130-41.
Riley, W. A. 1931. Erasmus Darwin and the biologic control of 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. 12: 288-92.
Smith, H. S. 1929. The utilization of entomophagous insects in the control of citrus pests. Trans. 4th Internatl. Congr. Ent. 2: 191-8.
Steinhaus, E. A. 1946. Insect Microbiology. Comstock Publ. Co., Inc., Ithaca, New York. 763 p.
Steinhaus, E. A. 1949. Principles of Insect Pathology. McGraw-Hill Book Co., Inc., New York. 757 p.
van Leeuwenhoek, A. 1702. Letter in Nr. 266 of the Philosophical Transaction 1700-1701, Vol. 22, p. 659-72. Smith & Walford, London.
van Lenteren, J. C. 1983. Biological pest control: passing fashion or here to stay? Organorama (Netherlands) 20: 1-9.
Walsh, B. D. 1866. Practical Entomologist. June 1866. p. 101.