1

Biological control history and development

STORY OF BIOLOGICAL PEST CONTROL

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
 2

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. Kollr <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
 3

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, Dactylopiushad 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
sphylloxera. 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
 4

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.

History of Biological Control 6

HISTORY AND DEVELOPMENT OF BIOLOGICAL CONTROL
Notes
I. The history of Biological Control may be divided into 3
periods:
A. The preliminary efforts when living agents were released rather
haphazardly with no scientific approach. Little precise information
exists on successes during this time. Roughly 200 A.D. to 1887 A.D.;
B. The intermediate period of more discrimminating BC which started with
 5

the introduction of the Vedalia beetle, Rodolia cardinalis Mulsant, for

control of the cottony cushion scale in 1888. Period extended from 1888
to ca. 1955; and
C. The modern period characterized by more careful planning and more
precise evaluation of natural enemies. Period from 1956 to the present.
II. Early History: 200 A.D. to 1887 A.D.
A. 200 A.D. to 1200 A.D: BC agents were used in augmentation
1. Chinese were the first to use natural enemies to control insect pests.
Nests of the ant Oecophylla smaragdina were sold near Canton in the
3rd century for use in control of citrus insect pests such as
Tesseratoma papillosa (Lepidoptera)
2. Ants were used in 1200 A.D. for control of date palm pests in Yemen
(south of Saudia Arabia). Nests were moved from surrounding hills
and placed in trees
3. Usefulness of ladybird beetles recognized in control of aphids and scales
in 1200 A.D.
B. 1300 A.D. to 1799 A.D.: BC was just beginning to be recognized.
1. Aldrovandi noted the cocoons of Apanteles glomeratus on a parasitized
Pieris rapae in 1602 A.D., but thought cocoons were insect eggs
2. Vallisnieri interpreted the phenomenon of insect parasitism (parasitoid)
in 1706 A.D. However the honor of being first to understand insect
parasitism may belong to the microbiologist Van Leeuwenhoek who
illustrated and discussed a parasite of a sawfly that feeds on willow in a
publication in 1701.
3. The first insect pathogen was recognized by de Reaumur in 1726. It
was a Cordyceps fungus on a noctuid
4. In 1734, de Reaumur suggested to collect the eggs of an "aphidivorous
fly" (actually a lacewing) and place them in greenhouses to control
aphids
5. The mynah bird, Acridotheres tristis, was successfully introduced from
India to Mauritius (off coast of Madagascar) for control of the red
locust, Nomadacris septemfasciata, in 1762
6. In the late 1700's, birds were transported internationally for insect
control
7. Control of the bedbug, Cimex lectularius, was successfully
accomplished by releases of the predatory pentatomid Picromerus
bidens in 1776 in Europe
History of Biological Control 7
Notes
C. 1800 A.D. to 1849 A.D. During this period advances were made in
Europe which were both applied and basic
1. In the 1800s, Darwin discussed "Ichneumonids" as natural control
factors for cabbage caterpillars
2. Malthus (in England) published Essays on the Principles of
Population in 1803
3. Hartig (Germany) suggested the rearing of parasites from parasitized
caterpillars for mass releases in 1827
4. Kollar (Austria) put forth the concept of "natural control" in 1837
5. Verhulst (1838) described the logistic growth equation but the idea
layed dormant until 1920 when rediscovered by Pearl. Expressed
 6

idea of "environmental resistance".

6. During the 1840's releases of predators were used for control of the
gypsy moth and garden pests in Italy
D. 1850 to 1887. During this time the focus on BC switched to the United
States.
1. From 1850 to 1870 enormous plantings of many crops were being
grown in the United States (especially California) and were initially
free of pests. Later farmers saw their crops destroyed by hordes of
alien pests
2. Asa Fitch (New York) suggested importing parasites from Europe
to control the wheat midge, Contarinia tritici, in 1856. No action
was taken. In 1860 parasites were requested from Europe, but none
where received
3. During this time period, Benjamin Walsh (Illinois) actively worked
for the importation of natural enemies to control the exotic insects
in the United States but was unsuccessful. Fortunately, he
influenced Charles V. Riley greatly who was in Missouri during
Walsh's campaign
4. First practical attempt at BC of weeds occurred in 1863 when
segments of the prickly pear cactus, Opuntia vulgaris, infested with
the imported cochineal insect, Dactylopius ceylonicus, were
transported from northern to southern India
5. Riley conducted the 1st successful movement of parasites for
biological control when parasites were moved from Kirkwood,
Missouri, to other parts of the state for control of the weevil
Conotrachelus nenuphar in 1870
6. LeBaron transported apple branches infested with oyster-shell scale
parasitized by Aphytis mytilaspidis from Galena to Geneva,
Illinois in 1871
7. In 1873 Riley sent the predatory mite Tyroglyphus phylloxerae to
France to control the grape phylloxera. The mite was established
but did not exert control as hoped.
8. Trichogramma sp. (egg parasites) were shipped from the U.S. to
Canada for control of lepidopterous pests in 1882
9. In 1883 the USDA imported Apanteles glomeratus from England
for control of P. rapae (the imported cabbageworm). Parasites
were distributed in DC, Iowa, Nebraska, and Missouri. First
intercontinental shipment of parasites.
History of Biological Control 8
Notes
III. The Intermediate Period: 1888 to 1955
A. 1888 to 1889: The Cottony Cushion Scale Project
1. Cottony cushion scale, Icerya purchasi Maskell, was introduced into
California in ca. 1868 around the Menlo Park (CA) area (near San
Francisco)
2. It spread to southern California and by 1887 was threatening to destroy
the infant citrus industry
3. C. V. Riley (Chief of the Division of Entomology, USDA) employed
Albert Koebele and D. W. Coquillett in research on control of the
cottony cushion scale
 7

4. No method was working in 1887

5. Koebele was sent to Australia in 1888 to collect natural enemies of the
scale
6. He sent ca. 12,000 individuals of Cryptochaetum iceryae and 129
individuals of Rodolia cardinalis (the vedalia beetle)
7. Within the year, the cottony cushion scale ceased to be a substantial
pest
8. The vedalia beetle controls the scale mainly in the inland desert areas
and C. iceryae controls it in the coastal areas of California.
B. 1890 to 1899: Growing pains for BC
1. Following the success in 1889, California put pressure on Riley to
send Koebele back to Australia in search of parasites for other scale
parasites in California
2. Koebele went on foreign exploration, but on his return he was recalled
from California. Koebele resigned his position and went to work for
the Republic of Hawaii in 1893. He worked on BC projects in the
interest of Hawaii until 1912 when he retired due to ill health.
3. Due the success of the vedalia beetle, great emphasis was placed on
importation of coccinelids for BC initially in California and Hawaii. It
is believed that California was set back many years by promoting
mostly biological control projects and not researching alternative
control methodologies.
4. L. O. Howard replaced C. V. Riley as Chief of the Division of
Entomology, USDA in 1894. Howard was prejudice against BC due to
the problems he saw in California
5. George Compere began as a foreign explorer for California in 1899
C. 1900 to 1930: New faces and more BC projects
1. The Gypsy Moth Project in New England (1905-1911). W. F. Fiske
was in charge in Massachusetts. Howard conducted foreign exploration
in Europe and arranged for parasites to be imported to the U.S. Many
prominent entomologists employed on the project: Harry Scott Smith,
W. R. Thompson, P. H. Timberlake.
2. The Lantana Weed Project in Hawaii (1902) First published work on
BC of weeds. Koebele went to Mexico and Central America looking
for phytophagus insects which were sent to R. C. L. Perkins in
Hawaii.
3. The Sugar-cane Leafhopper Project in Hawaii (1904-1920). Hawaiian
Sugar Planters Association (HSPA) created a Division of Entomology
in 1904. R. C. L. Perkins was appointed superintendent. Staff
History of Biological Control 9
Notes
consisted of O. H. Swezey, G. W. Kirkaldy, F. W. Terry, Alexander
Craw, and Albert Koebele. Later Frederick Muir was employed due to
Koebele's health problems. Muir found the highly effective predator
Tytthus (= Cyrtorhinus) mundulus (Miridae) in Queensland, Australia,
in 1920.
4. Berliner described Bacillus thuringiensis in 1911 as causative agent of
bacterial disease of the Mediterranean flourmoth
5. Prof. H. S. Smith appointed superintendent of California state
insectary, Sacramento, CA, in 1913. Facility moved to the University
 8

of California's Citrus Experiment Station in 1923 (now UC Riverside).

Smith started another facility at Albany, CA, in 1945. Riverside and
Albany (UC Berkeley) made up Department of Biological Control, UC.
6. USDA laboratory for biological control established in France in 1919.
7. The Imperial Bureau of Entomology created the Farnham House
Laboratory for BC work in England in 1927. This was later directed by
W. R. Thompson in 1928.
D. 1930 to 1955: Expansion and decline of BC
1. From 1930 to 1940 there was a peak in BC activity in the world with
57 different natural enemies established at various places.
2. World War II caused a sharp drop in BC activity.
3. BC did not regain popularity after WW II due to the production of
relatively inexpensive synthetic organic insecticides. Entomological
research switched predominantly to pesticide research.
4. In 1947 the Commonwealth Bureau of Biological Control was
established from the Imperial Parasite Service. In 1951 the name was
changed to the Commonwealth Institute for Biological Control (CIBC).
Headquarters are currently in Trinidad, West Indies.
5. In 1955 the Commission Internationale de Lutte Biologique contre les
Enemis des Cultures(CILB) was established. This is a worldwide
organization with headquarters in Zurich, Switzerland. In 1962 the
CILB changed its name to the Organisation Internationale de Lutte
Biologique contre les Animaux et les Plants Nuisibles. This
organization is also known as the International Organization for
Biological Control (IOBC). Initiated the publication of the journal
Entomophaga in 1956, a journal devoted to biological control of
arthropod pests and weed species.
IV. The Modern Period: 1957 to Present.
A. In 1959, Vern Stern et al. (1959) conceived the idea of economic injury
level and economic threshold which would permit growers to make
informed decisions on when they needed to apply control tactices in their
cropping systems and therefore eliminated the need for scheduled pesticide
treatments.
B. Interest developed nationwide in ecology and the environment after 1962
with the publishing of the Rachel Carsons book Silent Spring.
C. Silent Spring helped stimulate the implementation of the concept of
Integrated Pest Management (IPM) in the late 1960s, and biological
control was seen as a core component of IPM by some. More emphasis
was placed on conservation BC than classical BC.
D. In 1964, Paul DeBach and Evert I. Schliner (Division of Biological
Control, University of California, Riverside) publish an edited volume
titled Biological Control of Insect Pests and Weeds which becomes a
History of Biological Control 10
Notes
major reference source for the biological control community. This was
basically a California based book with international application.
E. In some areas in the USA (e.g., California, North Carolina, Kansas,
Texas), IPM scouting was commercialized in the 1970s and natural
enemies were relied upon to suppress pests in crops such as cotton,
alfalfa, citrus, soybeans, and other crops.
 9

F. During the 1970s and 1980s, Brian Croft and Marjorie Hoy made
impacts by using pesticide resistant natural enemies in cropping systems.
G. In 1983, Frank Howarth published his landmark paper titled Biological
Control: Panacea or Pandoras Box and significantly impacted classical
BC efforts by concluding that classical BC of arthropods significantly
contributed to extimction of desirable species (e.g., endemic).
1. This eventully forced a rethinking of legislative guidelines as well as
introduction methods which are still being changed today.
2. In Hawaii, BC efforts were diminished significantly and have not risen
to levels prior to 1985.
3. Research efforts into this area were stimulated with the general results
that many of Howarths claims were unjustified, but some impacts
were discovered. No species extinctions have been demonstrated to
have resulted from classical BC efforts to date.
H. In the 1990s, two additional biological control journals appeared,
Biological Control - Theory and Application in Pest Management
(Academic Press) and Biocontrol Science and Technology (Carfax
Publishing). Additionally, Entomophaga changed its name to
Biocontrol in 1997.
QUESTIONS
1. What are early examples of the use of natural enemies in agriculture?
2. What contribution did Vallisnieri make to biological control?
3. Who conducted the first successful movement of parasites for the purposes of biological
control?
4. Discuss in detail the biological control project on cottony cushion scale.
5. Who was Professor Harry Scott Smith?
6. What major change in control strategies in the late 1960s made biological control of
arthropods a more attractive
management tactic?
7. What did Frank Howarth contribute to the field of biological control?
REFERENCES
DeBach, P. 1974. Biological control by natural enemies. Cambridge University Press, London.
323 pp.
Doutt, R. L. 1964. The historical development of biological control. p. 21-42. In Biological
Control of
Insect Pests and Weeds (P. DeBach, editor). Chapman and Hall Ltd, London. 844 pp.
Hagen, K. S., and J. M. Franz. 1973. A history of biological control. p. 433-476. In A History
of
Entomology (R. F. Smith, T. E. Mittler, and C. N. Smith, editors). Annu. Rev. Inc., Palo Alto,
California.
517 pp.
Howard, L. O. 1930. A history of applied entomology. Smithsonian Misc. Coll., 84: 1-564.
History of Biological Control 11
Konishi, M., and Y. Ito. 1973. Early entomology in east Asia. pp. 1-20. In A History of
Entomology
(R. F. Smith, T. E. Mittler, and C. N. Smith, editors). Annu. Rev. Inc., Palo Alto, California.
517 pp.
Simmonds, F. J., J. M. Franz, and R. I. Sailer. 1976. History of biological control. In Theory
and
Practice of Biological Control (C. B. Huffaker and P. S. Messenger, editors). Academic Press,
 10

New York. 788

pp.
van den Bosch, R., P. S. Messenger, and A. P. Gutierrez. 1982. An introduction to
biological
control. Plenum Press, New York and London. 247 pp.
READING ASSIGNMENT:
Chapter 1: pp. 320, Van Driesche, R. G. and T. S. Bellows, Jr. 1996. Biological control.
Chapman
and Hall, New York. 539 pp.

he 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.

[For a more complete outline of the history of biological control, visit this website].

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). 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 laying eggs in aphid
hosts. Vallisnieri (1706) 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)
described and illustrated Cordyceps fungus infecting a noctuid larva.

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.

Historical Developments in Biological Control - Middle History

to 1940

The knowledge about parasitoids, predators and diseases were accumulating and
the struggle for existence was taking place an early Ecologist George Russel
 11

Wallace and Charles Darwin put forth general idea called Balance of Nature.

John Curtis in his book on Farm Insect (1860) includes ecology of various
agricultural pests and emphasized functions of parasitoids and predators in crop
protection. Riley (1873) arranged first international shipment of natural enemies in
the transfer of predatory mite Tyroglyphus phyllaxerae to France from North
America for possible control of grapevine phylloxera. Riley (1883) directed import
of internal parasite of cabbage butterfly from England to America and A.
glomeratus eventually, become successful and well distributed in Eastern and mid
western state as larval parasitoid. This successful importation of A. glomeratus by
Riley was the first international transfer of parasitoid for biological control.

The Italian Microbiologist Agastina Bassi is the first worker to suggest use of
microbes for insect pest suppression in 1836. Lewis pasture from France was more
confident for his suggestion of use of protozoa causing pebrine disease in bees.
Russian Zoologist Metchnikiff recorded large fluctuation in pest population of wheat
cock chafer beetle and one of the causes was green muscardine fungus,
Meterrhizium anisopliae.

The successful introduction of coccinellid beetle Radalia cardinalis from Australia to

California in 1888, was the first spectacular success controlling cottony cushion
scale (Icerya puchasi) a serious pest of citrus and become a first classical example
of biological insect pest suppression and it was later acclaimed as miracle
Cryptolaemns montrouzieri. Mulsent, a mealy bug predator approached usefulness
as like that of R. cardinalis (vedalia beetle). However Smith H. S. (1912) put
quarantine measures to stop importation of beneficial organisms into California. In
1919, he was first to propose the term Biological Control and wrote voluminously
in 1935-1939 on theoretical aspects of biological control.

In Europe, Metchnikoff tried Beauveria sp against nunmoth caterpillar, Gypsy moth

and Melolontha beetles. In mid western U. S. the fungus, B. basiana was used for
control Chinch bug, Blesses leucopterus. In 1920, Albert koebele used successful
biological programs against leaf hoppers by introduction of egg parasitoids from
Australia and a predacious mired bug.

First modest biological control laboratory in Canada was established in the

University of New Brunswick and by 1960 when importation was stopped. Three
important parasitoids had been established as a footing stone for future
development. Parker and Thompson set laboratory in France and during 1927-40
imported several parasitoids for biological control of European corn borer, Ostrilia
nubilalis. Baird (1923-56) carried out corn borer suppression program in Canada
for alfa weevil. Japanese beetle and European Ear wig with the help of USDA,
Bureau of Entomology. In 1929, Canadian Entomologist established Dominion
parasite Laboratory at Belleville and then new era of biological control was dawned
by 1933. In 1940, because of World War II, the Fernharn laboratory (U. S.) was
closed and Thompson service later recognized as CIBC i.e. Common wealth
Institute of Biological Control.

17: BIOLOGICAL CONTROL - DEFINITION - HISTORY AND

DEVELOPMENT - CLASSICAL EXAMPLES - FACTORS GOVERNING
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BIOLOGICAL CONTROL
Biological control
Definition
The study and utilization of parasitoids, predators and pathogens for the
regulation of pest population densities.
Biological control can also be defined as the utilization of natural enemies to
reduce the damage caused by noxious organisms to tolerable levels.
Biological control is often shortened to biocontrol.
History and development of biological control and classical examples of biological
control
Antient times - In China Pharoahs ant Monomorium pharaonis was used to control
stored grain pest. Red ant Oecophylla spp. used to control foliage
feeding caterpillar.
Year 1762 - Mynah bird imported from India to Mauritius to control locust.
1770 - Bamboo runways between citrus trees for ants to control
caterpillars.
1888 - First well planned and successful biological control attempt
made
- During 1888 citrus industry in California (USA) seriously threatened by cottony
cushion scale, Icerya purdian
- Chemical treatments not known at that time
- Mr. C.V. Riley, a prominent entomologist suggested that the scale inset originated
from Australia and natural enemy for the scale from Australia should be
introduced into USA
- Mr. Albert Koebele was sent to Australia
- He found a beetle called Vedalia (Rodolia cardinalis) attacking and feeding on
seeds
- Vedalia beetle (Rodolia cardinalis) was imported in November 1888 into USA
and allowed on scale infested trees
- Within a year spectacular control of scale insect achieved
- Even till date this beetle controls the scale insect
- After this successful attempt of biological control many such introduction of
natural enemies were tried.
1898 - First introduction of natural enemy into India
1898 - A coccinellid beetle, Cryptolaemus montrouzieri was imported into India
from Australia and released against coffee green scale, Cocus viridis.
Even today it is effective against mealybugs in South India.
1920 - A parasitoid Aphelinus mali introduced from England into India to control
Woolly aphid on Apple, Eriosoma lanigerum.
1929-31 - Fodolia cardinalis imported into India (from USA) to control cottony
cushion scale Icerya purchasi on Wattle trees.
1958-60 - Parasitoid Prospatella perniciosus imported from China
1960 - Parasitoid Aphytis diaspidis imported from USA
Both parasitoids used to control Apple Sanjose scale Quadraspidiotus
perniciosus
1964 - Egg parasitoid Telenomus sp. imported from New Guinea to control Castor
semilooper Achaea janata
1965 - Predator Platymeris laevicollis introduced from Zanzibar to control coconut
Rhinoceros beetle, Oryctes rhinoceros
History, development, classical examples of biocontrol
 13

Till 1988
At global level 384 importations made against 416 species of insect pests. Out of
them
164 species (39.4%) - Completely controlled
75 species - Substantially controlled
15 species - Partially controlled
- Regional Station of Commonwealth Institute of Biological Control (CIBC)
established at Bangalore in 1957
- Presently Project Directorate of Biological Control (PDBC) Bangalore looks after
Biocontrol in India.
Factors affecting biological control
1. Tolerance limit of crop to insect injury - Successful in crops with high tolerance
limit
2. Crop value - Successful in crops with high economic value
3. Crop duration - Long duration crops highly suitable
4. Indigenous or Exotic pest - Imported NE more effective against introduced pest
5. If alternate host available for NE, control of target pest is less
6. If unfavourable season occurs, reintroduction of NE required
7. Presence of hyperparasites reduces effectiveness of biocontrol
8. Tritrophic interaction of Plant-Pest-Natural enemy affects success of biocontrol,
e.g. Helicoverpa parasitization by Trichogramma more in timato than corn
9. Use of pesticides affect natural enemies
10. Selective insecticides (less toxic to NE required)
11. Identical situation for successful control does not occur
Qualities of an effective natural enemy
1. Adaptable to the environmental condition
2. Host specific (or narrow host range)
3. Multiply faster than the host (with high fecundity)
4. Short life cycle and high female : male ratio
5. High host searching capacity
6. Amenable for easy culturing in laboratory
7. Dispersal capacity
8. Free from hyper parasites
9. Synchronise life cycle with host
Three major techniques of biological control
1. Conservation and encouragement of indigenous NE
Defined as actions that preserve and increase NE by environmental
manipulation. e.g. Use of selective insecticides, provide alternate host and refugia for
NE.
2. Importation or Introduction
Importing or introducing NE into a new locality (mainly to control introduced
pests).
3. Augmentation
Propagation (mass culturing) and release of NE to increase its population.
Two types,
(i) Inoculative release: Control expected from the progeny and subsequent
generations only.
(ii) Inundative release: NE mass cultured and released to suppress pest directly
e.g. Trichogramma sp. egg parasitoid, Chrysoperla carnia predator
ROLE OF PARASITOIDS AND PREDATORS IN IPM
 14

- Parasitoids and predators may be used in Agriculture and IPM in three ways.
They are
i) Conservation
ii) Introduction
iii) Augmentation - (a) Inoculative release, (b) Inundative release
- Since biological control is safe to environment, it should be adopted as an
important component of IPM.
- Biological control method can be integrated well with other methods namely
cultural, chemical methods and host plant resistance (except use of broad
spectrum insecticides)
- Biological control is self propagating and self perpetuating
- Pest resistance to NE is not known
- No harmful effects on humans, livestock and other organisms
- Biological control is virtually permanent
- Biological agents search and kills the target pest
MICROBIAL CONTROL
- It is a branch of biological control
- Defined as control of pests by use of microorganisms like viruses, bacteria,
protozoa, fungi, rickettsia and nematodes.
I. VIRUSES
Viruses coming under family Baculoviridae cause disease in lepidoptera
larvae. Two types of viruses are common.
NPV (Nucleopolyhedro virus) e.g. HaNPV, SlNPV
GV (Granulovirus) e.g. CiGV
Symptoms
Lepidopteran larva become sluggish, pinkish in colour, lose appetite, body
becomes fragile and rupture to release polyhedra (virus occlusion bodies). Dead larva
hang from top of plant with prolegs attached (Tree top disease or Wipfelkrankeit)
II. BACTERIA
Spore forming (Facultative - Crystalliferous)
2 types of bacteria Spore forming (Obligate)
Non spore forming
i. Spore forming (Facultative, Crystelliferous)
The produce spores and also toxin (endotoxin). The endotoxin paralyses gut
when ingested e.g. Bacillus thuringiensis effective against lepidopteran. Commercial
products - Delfin, Dipel, Thuricide
ii. Spore-forming (Obligate)
e.g. Bacillus popilliae attacking beetles, produce milky disease
Commercial product - Doom against white grubs
iii. Non-spore forming
e.g. Serratia entomophila on grubs
III. FUNGI
i. Green muscardine fungus - Metarhizium anisopliae attack coconut rhinoceros
beetle
ii. White muscardine fungus - Beaveria bassiana against lepidopteran larvae iii.
White halo fungus - Verticillium lecanii on coffee green scale.
Other Microbs: Protoza, Nematodes
Limitations of biocontrol technique
- Complete control not achieved - Slow process
- Subsequent pesticide use restricted
 15

- Expensive to culture many NE

- Historical Perspective of Biological Control
The first accounts of predatory insects being used as insect management tools
date back as early as 300 AD when Chinese citrus growers placed paper nests of
ants (Oecophylla smaragdina F.) on trees to protect them from other insects (van
Lenteren, 2005). These early augmentation efforts were apparently helped along by
the conservation biological control practice of aiding inter-tree movement of the ants
by placing bamboo rods as runways or bridges between trees (DeBach, 1974). These
ants reportedly were still available for purchase up to at least the 1970s (DeBach,
1974).
While the predatory behavior of some insects was recognized long ago and taken
advantage of for pest management, the recognition and utilization of the less obvious
parasitic insects did not occur until much later. Parasitism by tachinid flies
was first correctly interpreted in China in the 11th century, while ichneumonoid
parasitism was correctly interpreted in Europe in the 17th century (Cai et al., 2005;
van Lenteren and Godfray, 2005). The difference in time between these two events
was likely the more complex life history of the latter group.
The first deliberate movement of parasitoids from one location to another was
conducted by C.V. Riley, who distributed parasitoids of the weevil Conotrachelus
nenuphar (Herbst) around the state of Missouri in 1870 (Doutt, 1964). The first parasitoid
successfully moved and established from one continent to another, however,
was Cotesia (=Apanteles) glomeratus (L.), which was shipped from England to the
United States for suppression of Pieris rapae (L.) by the U.S. Dept. of Agriculture in
1883 (Riley, 1885; Riley, 1893). Transcontinental shipment of a predatory arthropod
soon followed with the transport of the predatory mite, Tyroglyphus phylloxerae
Riley &. Plancon, from the United States to France in 1873 for suppression of
the grape phylloxera, Daktulosphaira vitifoliae (Fitch) which it did not suppress
(Fleschner, 1960; Doutt, 1964). While a variety of international movements of insects
for pest control occurred in the late 1800s, none of them achieved complete
economic control (Fleschner, 1960).
It is generally accepted that the first case of complete and sustained economic
control of an insect pest by another insect was control of the cottony cushion
scale, Icerya purchasi Maskell, in California during the late 1800s (Fleschner, 1960;
Doutt, 1964; Debach, 1974; van den Bosch et al., 1982). Icerya was introduced
into Californiai in 1869, and by 1886 it threatened to destroy the entire southern
California citrus industry (DeBach, 1974). Two insects, the vedalia beetle, Rodolia
cardinalis Mulsant (Coleoptera: Coccinellidae), and a parasitic fly, Cryptochetum
9 Biological Control and Integrated Pest Management 215
iceryae (Williston) (Diptera: Cryptoch`etidae), were imported to California from
Australia in 1877 and 1888. Within two years, I. purchasi was under complete biological
control throughout the state. Although the vedalia beetle is mostly credited
for controlling the cottony cushion scale, once established, the parasitic fly became
the major control factor of the pest in the coastal areas of the state (Van Driesche
and Bellows, 1996). This classic example is presented in many books dealing with
insect biological control (e.g. DeBach, 1964, 1974; van den Bosch et al., 1982; Van
Driesche and Bellows, 1996), and set the stage for future biological control programs.
Probably because I. purchasi provides suppression of C. iceryae only over
a limited portion of the pests range, Greathead (1986) considered the importation
of Encarsia berlesi (Howard) into Italy from USA in 1906 for control of the mulberry
scale, Pseudaulacaspis pentagona Targioni-Tozzetti to be the first successful
introduction of a parasitoid from one country to another for insect pest control.
Following the success of the cottony cushion scale project, numerous biological
control efforts ensued worldwide (Clausen, 1978; Luck, 1981; van den Bosch
et al., 1982; Greathead, 1986; Greathead and Greathead, 1992) some of which were
 16

just as successful. Although the primary focus of early efforts in biological control
was importation of natural enemies, other methods of manipulating parasitoids and
predators were also considered.While the concept of mass rearing insects for future
releases was proposed as early as 1826 by Hartig, the first practical attempt towards
augmentation of natural enemies in western Europe was probably made in 1899 by
Decaux who devised a complete management program for apple orchards, including
releases of field-collected inchneumonid wasps (Biliotti, 1977). The first sustained,
large-scale, and successful augmentation biological control project involved
mass-production of the ladybeetle Cryptolaemus montrouzieri Mulsant, targeting
the citrophilus mealybug, Pseudococcus calceolariae Fernald (= gahani Green), a
pest of citrus in southern California (Luck and Forster, 2003). Large-scale releases
began in the early 1920s, and continued for decades, with as many as 40 million
beetles being produced annually. This beetle is still available through commercial
insectaries in both the United States and Europe (van Lenteren, 2003b).
The history of conservation biological control has been one of mainly potential
practices developed by researchers that do not appear to have become widely
adopted (Ehler, 1998). However, organic and sustainable farming systems have tried
to take advantage of these practices to some degree (Altieri et al., 2005).