History of Biological Control

Classical biological control is long lasting and inexpensive. Other than the initial costs of collection, importation, and rearing, little expense is incurred. When a natural enemy is successfully established it rarely requires additional input and it continues to kill the pest with no direct help from humans and at no cost. Classical biological control does not always work. It is usually most effective against exotic pests and less so against native insect pests. 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 He

glomeratus being attached to larvae of Pieris rapae (the imported cabbageworm).

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 ovipositing in aphid hosts. Vallisnieri (1706) first correctly interpreted this host-parasitoid association and Bodenheimer (1931),

probably became the first to report the existence of parasitoids.

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. 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." 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) 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) 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 (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. 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 (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 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) studied silkworm diseases and saved the silk industry in France from ruin [not really biological control]. Charles Valentine Riley (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, his son, also devoted his entire career to the search for and identification of natural enemies of scale insects. Harry Scott Smith (1913) 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.

Examples of Successful Biological Control

There are many examples of successful classical biological control programs. Joseph Needham noted a Chinese text dating from 304AD, Records of the Plants and Trees of the Southern Regions, by Hsi Han, which describes mandarin oranges protected by biological pest control techniques that are still in use today. One of the earliest successes in the west was in controlling Icerya purchasi, the cottony cushion scale, a pest that was devastating the California citrus industry in the late 19th century. A predatory insect Rodolia cardinalis (the Vedalia Beetle), and a parasitoid fly were introduced from Australia by Charles Valentine Riley. Within a few years the cottony cushion scale was completely controlled by these introduced natural enemies.

Damage from Hypera postica Gyllenhal, the alfalfa weevil, a serious introduced pest of forage, was substantially reduced by the introduction of several natural enemies. 20 years after their introduction the population of weevils in the alfalfa area treated for alfalfa weevil in the Northeastern United States was reduced by 75 percent. A small wasp, Trichogramma ostriniae, was introduced from China to help control the European corn borer making it a recent example of a long history of classical biological control efforts for this major pest. Many classical biological control programs for insect pests and weeds are under way across the United States and Canada. The population of Levuana irridescens (the Levuana moth), a serious coconut pest in Fiji, was brought under control by a classical biological control program in the 1920s.

Examples of Unsuccessful Biological Control Programs

An example of an invasive species is the alligator weed. This plant was introduced to the United States from South America. This aquatic weed spreads rapidly and causes many problems in lakes and rivers. The weed takes root in shallow water causing major problems for navigation, irrigation, and flood control. The alligator weed flea beetle and two other biological controls were released in Florida. Because of their success, Florida banned the use of herbicides to control alligator weed three years after the controls were introduced (Cofrancesco 2007). The cane toad, Bufo marinus, was introduced as a biological control and had significant negative impact on biodiversity. The cane toad was intentionally introduced to Australia to control the cane beetle. When introduced, the cane toad thrived very well and did not only feed on cane beetles but other insects as well. The cane toad soon spread very rapidly, thus taking over native amphibian habitat. The introduction of the cane toad also brought foreign disease to native reptiles. This drastically reduced the population of native toads and frogs.

The cane toad also exudes and can squirt poison from the parotid glands on their shoulders when threatened or handled. This toxin contains a cocktail of chemicals that can kill animals that eat it. Freshwater crocodiles, goannas, tiger snakes, dingos and northern quolls have all died after eating cane toads, as have pet dogs (Cane toad, 2003). The reasons for failure are not often known but may include the release of too few individuals, poor adaptation of the natural enemy to environmental conditions at the release location, and lack of synchrony between the life cycle of the natural enemy and host pest. A second example of a biological control agent that subsequently crossed over to native species is the Rhinocyllus conicus. The seed feeding weevil was introduced to North America to control exotic thistles (Musk and Canadian). However, the weevil did not target only the exotic thistles, it also targeted native thistles that are essential to various native insects. The native insects rely solely on native thistles and do not adapt to other plant species. Therefore, they cannot survive. Biological controls do not always have negative impacts on biodiversity (Corry 2000).

References:
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