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EDWARD F. KNIPLING March 20, I 909-March I 7, 2000 BY PERRY ADKISSON AND JAMES TUMLINSON EDWARD F. KNIPEING, RETIRED U.S. Department of Agricul ture entomologist en c! administrator slier! on March 17, 2000, in Arlington, Virginia, at the age of 91. He was best known for cleveloping the sterile insect technique, which was the principal technology used to eradicate the screw- worm fly from North America. Often callecl "Knip" by his friends en cl colleagues, he was born in Port Lavaca, Texas, on March 20, 1909, en c! grew up on his parents' small farm. His ciacl assigned many chores to him on the farm, one being the oclious task of doctoring baby calves that hac! screwworms burrowing in their navels en cl wounds. Another was picking cotton by hand in boll-weevil-ravagecl fielcis. From this experience young Knipling developed an interest in entomology and later said that he cleciclecl at an early age he wan tell to make a greater contribution to agriculture than treating screwworm-infestecl calves or pulling a sack down a cotton row. Thus, after leaving the insect-laclen farming area of the Texas gulf coast he decided that entomology was the logical field for him to enter. After graduating from high school Knip enrolled at Texas A&M College (now Texas A&M University), where he was 81

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82 B I O G RA P H I C A L EMOIRS awarclec! bachelor en c! master of science degrees in ento- mology in 1930 en cl 1932, respectively. In 1947 he was awarclecl a Ph.D. in entomology from Iowa State University. Knip's career with the U.S. Department of Agriculture (USDA) began in 1930 as a fielcl aicl in the former Bureau of Entomology en cl Plant Quarantine, where he assisted in field! studies in Mexico on the pink bollworm. In 1931 he was appointed a junior entomologist to concluct research at MenarcI, Texas, on the biology en cl control of the screw- worm. The screwworm is a subtropical fly that lays its eggs in the open wounds of warm-blooclecl animals. The flesh- eating larvae or maggots cause suffering, cleath, en cl untoIcl economic Tosses in cattle, other livestock, wiTcIlife, en c! even humans. In his screwworm research assignment Knipling's talents as a scientist became obvious, as clicl his keen intel- lect for looking at oic! problems in a new way. By 1937 he hacl teamed with a young colleague, R. C. BushIancI, to stucly the mating habits of screwworm flies. Observing that mate flies mater! repeatecITy while female flies mater! only once in their lifetime, Knip believecl they hacl founcl a weak link in the screwworm's life cycle that might be exploitecl for control. The question was: "How?" This was a question Knip pondered for several years before finding the answer. Knip's research in Texas on screwworms was interrupted intermittently when he was assignee! to concluct research on other pest problems of livestock in Illinois, Iowa, en cl Georgia. In 1940 he was placecl in charge of research on mosquitoes of the northwestern states with headquarters in PortIancI, Oregon. During World War II Knip was given the important as- signment of devising better ways for controlling the arthro- pocl vectors (flies, mosquitoes, lice, en cl other biting in- sects) of human diseases affecting our troops. He was macle director of the USDA research laboratory at Orlando, Florida,

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EDWARD F. KNIPLING 83 where he lee! the clevelopment of DDT en c! other insecti- cicles en cl repellents for use by our armecl forces en cl allies to control the vectors of malaria, typhus, plague, en cl other arthropocI-vectorec! diseases that hac! exacter! a tremendous toll on troops in previous wars. The laboratory was success- ful in its mission to clevelop effective control measures of the disease vectors, thus preventing infection, illness, en c! cleath of thousands of service personnel across the worIcI. The research concluctecl at the OrIanclo laboratory received national en c! international recognition. Many of the repel- lents en cl methods of control for the arthropod vectors of some of the most serious human diseases are still being user! throughout the woric! tociay. During this period Knip continual to think about the screwworm problem. With imagination en cl innovation he conceiver! the iclea of using sterile insects for population suppression en cl eradication. He reasoned that if male flies couIcl be proclucecl in large numbers, sterilizecI, en cl re- leasec! into the environment they might out-compete, on a simple probability basis, the wild fertile males in breeding with females. Because female screwworms mate only once, those that were brec! with sterile mates wouic! lay infertile eggs en cl thus not produce any progeny. Knip reasoned that if a sufficient number of sterile males couIcl be re- leasec! into the wiTc! population they wouic! essentially over- whelm and breed the screwworm population into extinc- tion. Knip was consumed by this iclea and soon began cleveloping simple mathematical moclels of the population dynamics of the screwworm fly. These moclels convinced him that the sterile insect concept shouIcl work according to laws of probability if methods conic! be clevelopec! for sterilizing the males en cl mass rearing the flies in sufficient numbers to out-compete the fertile males when releasecl into the fielcI. . .

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84 B I O G RA P H I C A L EMOIRS In 1946 Knip was placer! in charge of all USDA research on insects affecting livestock, man, househoIcis, en cl stored products en cl was transferred to headquarters in Washing- ton, D.C. From this position he encourages! Bushianc! who was still in Texas to pursue this line of research with the screwworm fly. They maintained frequent communication to exchange icleas en c! discuss new research approaches to the problem. In the January 1950 issue of American Scientist Profes- sor H. l. Muller of Indiana University in Bloomington re- portecl that fruit flies couIcl be sterilizecl by exposure to X rays. This report excited Knipling en cl BushIancl en cl they cleciclec! to try this procedure on screwworm flies, however, BushIancl's laboratory clicl not have the neeclecl equipment, en cl funcis were not available to purchase what was neeclecI. Not being one to give up, Bushianc! smuggler! insects into the X-ray laboratory of an army hospital in San Antonio, where he was a friend with some of the staff members. There he was able to use their equipment on clays when the unit was not busy. He tested various dosages of X rays on the aclult, larval, en cl pupal stages of the screwworm. He cliscoverec! that screwworm flies subjected! to an appropr~- ate dosage of X rays in the pupal stage not only survivecl but also emerged into healthy aclults that were sexually sterile. Cage studies using both sterile en c! normal flies in various ratios confirmed the theory that reproduction of the screw- worm couIcl be inhibited at levels consistent with the math- ematical probability moclels. Part of the problem was solvecI. There was a way to sexu- ally sterilize the screwworm without any serious adverse af- fect on their health or their ability to compete with wiTc! males in mating with females. The other parts of the prob- lem were: "How clo you mass rear large numbers of screw- worm flies en c! how many will be neeclec! to suppress a field! .

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EDWARD F. KNIPLING 85 population over a large area?" Bushianc! was given the prob- lem of cleveloping methods for mass rearing the insects en cl improving the sterilization technology, while Knipling con- tinucc! working on mathematical moclels to answer the ques- tion of how many flies wouIcl be neeclecl to obtain success with sterile male releases. In 1954 Knip was given the opportunity to test his theo- ries when the Dutch government askocl the USDA for help in controlling screwworms that were clecimating the goat en c! ciairy calf populations on the island! of Curacao off the coast of Venezuela. From a rearing facility in Floricia 170,000 flies a week were proclucecI, sterilizecl with a gamma raclia- tion source, en c! transporter! to en c! releaser! across the is- lancI. Very soon the number of wilcl flies in the population was reduced, after several months and about three fly gen- erations the population was eraclicatec! from the isTancI, which has remained free of infestation since 1954. This successful demonstration that the sterile insect re- lease methoc! conic! be user! to eradicate insect pest popula- tions not only excited Knipling en cl BushIancl but also gave their work great impetus as it excited the livestock procluc- ers in screwworm-infestec! areas of the Uniter! States. . During the late 1950s a much larger en cl more clifficult test of the sterile insect release technique was macle. Live- stock producers in Floricia gainer! support of fecleral en c! state agencies to concluct an eradication program in their state. By 1959 the screwworm was eraclicatecl from Floricia and all of the southeastern United States. This success led livestock producers in Texas and Okla- homa to organize en cl gain support for a similar program. The producers raiser! a substantial amount of money for the program en cl were assisted by two prominent Texas ranch- ers, President Lynclon Johnson en cl Governor Dolph Briscoe of Texas, in obtaining federal and state funds and technical

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86 B I O G RA P H I C A L EMOIRS assistance. By 1966 the screwworm was eraclicatec! from the Unitecl States. In 1972 the program was expanclecl to Mexico, where eradication was achieved in 1991. The program has since mover! through all of Central America en c! a barrier zone is now being maintained at the Panama-Colombia bor- cler. The eradication of the screwworm from North America was truly a remarkable achievement, both technically en cl economically. The benefits to livestock producers through- out the eradication zone are well over $1 billion per year. The cumulative benefits over more than 50 years, inclucling all the economic multipliers, environmental quality, en cl avoidance of animal en c! human suffering, are too large en cl staggering to even estimate. The success of the sterile insect release technique for eradicating the screwworm fly was the first successful clem- onstration that a pest species can be eliminatecl from large geographical areas with appropriate technology. Further- more, the sterile insect technique is creclitec! as being one of the most significant peaceful applications of nuclear ra- cliation for the benefit of mankind. This technique has since been user! to eradicate en c! control other pests, such as the Mediterranean en cl other fruit flies in California, Floricia, en cl other parts of the worIcI. In 1988 it was also cliscoverecl that the screwworm had been accidentally introduced into Libya en cl soon thousands of animals were infested. The greater threat was that the screwworm fly might infest most of Africa, causing severe Tosses to foot! animals en c! wiTcIlife. The sterile insect release technique was employocI, en cl the fly was successfully eradicated from Africa. From 1953 to 1971 Knip was director of the USDA's Entomology Research Division, where he was in charge of all arthropod research concluctecl by the Agricultural Re- search Service. During this perioc! great advances were macle

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EDWARD F. KNIPLING 87 in the field of entomology and pest management, attrib- uted to his vision and leadership in developing and pro- moting principles and strategies for suppressing insect pests by such techniques as sterile insect releases, pheromone traps, biological control agents, and cultural practices, with a minimal use of insecticides. During the latter part of his career Knip became con- vinced that it was also possible to eradicate the boll weevil, the second scourge of his youth, from the United States. He was able to gain support for his ideas within the USDA and with cotton producer groups. A large research project was mounted in the early 1960s to develop the technology needed to eradicate the weevil, or at least eliminate it as an economic pest of cotton in the United States. During the ~960s and ~970s Knipling developed a conceptual frame- work for an eradication program, using a variety of tech- niques. His concepts were field tested in Mississippi and later fine tuned in the cotton fields of Virginia and North Carolina, where the boll weevil was successfully eradicated in 1987. Following this, eradication programs were conducted throughout the southeastern United States. The program has moved westward across the Cotton Belt, with eradica- tion efforts currently underway in the mid-south, Texas, and Oklahoma. Eradication of the boll weevil from the United States should be accomplished relatively soon. When this is done, the use of chemical peshc~des on cotton, and the consequential environmental impact, will be greatly reduced. After retirement in 1973 and until his death Knipling remained professionally active, including serving as an un- paid consultant to the USDA's Agricultural Research Ser- vice and the Animal and Plant Health Inspection Service, where he advised on pest management programs. Through- out his 28 retirement years he continued to publish exten- sively, and was considered a leading authority on insect ~ ~ . ~ . . ~ ~ ~

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88 B I O G RA P H I C A L EMOIRS population dynamics en c! control. In 1979 he wrote a book on the basic principles of insect population suppression. In 1992 he wrote another book on insect parasitism from new perspectives. In these en c! many other publications, as well ~ . ~ .. .. as In seminars en cl lectures on insect pest management, Knipling constructively questioned en cl challengecl many conventional insect control strategies that are baser! on small areas, farm-to-farm applications, continual heavy reliance on insecticides, en cl reactive treatments after pest popula- tions reach high levels en c! damage occurs. He was a strong proponent of the area-wicle management of pest popula- tions by a variety of proactive technologies. His primary theme was to prevent insect pests from reaching damaging levels by using biological and other nonpesticidal suppres- sion methods that wouIcl not aciversely impact the environ- ment en c! nontarget organisms over large geographic ar- eas. . In aciclition to his many professional achievements Knipling was the patriarch of a large en c! active family that shared many common interests en cl boncis. His wife of 66 years, Phoebe, was also an Iowa State University Ph.D. graclu- ate en c! was an accomplishes! educator in the Arlington County, Virginia, public school system. Together they hacl 5 chilciren, 14 grancichilciren, en cl 9 great-grancichilciren. Knip's professional interest in entomology en c! nature In general greatly influenced ciay-to-ciay life en cl activities of the family. For example. all of their nets were named after insects, either their common or scientific names: Siamese cats Anthonomus and Culex were named after the cotton boll weevil en cl a type of mosquito, respectively. Knip was an avic! outdoorsman en c! naturalist with a strong conservation ethic. The family owned several large proper- ties in the mountain regions of Virginia, West Virginia, and Vermont. These properties were managed for recreation

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EDWARD F. KNIPLING 89 en c! timber production. Knip was an accomplishes! archer, hunter, en cl fisherman. He almost always caught more fish than anybody else, often self-attributecl to being able to "think like a fish" en c! outsmart them. He even carver! en c! pain tell his own fishing lures, making them look like in- sects, of course. His favorite was a lure that lookocl like a cicada, he caller! it "Humbug" en c! caught Tots of fish with it. In summary, Knipling hacl three main themes in his life: his family, his profession, en c! his great reverence for na- ture en cl love of the outdoors. In his roles as a clistinguishecl scientist en cl administrator Knip significantly acivancecl the woricl's knowlecige of insect pest management en c! allevi- atecl some of the most important insect pest problems of agriculture across the worIcI, in an environmentally sound manner. The scientific principles en c! strategies he promoter! en cl clocumentecl are sure to continue to guicle new clevel- opments in insect population management well into the future. We thank Ec~warcl B. Knipling for helpful comments en cl information and Susan H. Fugate of the National Agricul- tural Library for assistance in locating and verifying infor- mation. SIGNIFICANT HONORS AND DISTINCTIONS 1947 Presidential Medal for Merit 1948 King's Medal for Service in the Cause of Freedom, Great Britain 1952 President, Entomological Society of America 1966 Membership, National Academy of Sciences Rockefeller Public Service Award 1967 National Medal of Science Honorary doctorate, North Dakota State University

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go BIOGRAPHICAL MEMOIRS 1970 Membership, American Academy of Arts and Sciences Honorary doctorate, Clemson University 1971 President's Award for Distinguished Federal Service 1975 Honorary doctorate, University of Florida 1986 Agricultural Research Service Science Hall of Fame 1991 FAO Medal for Agricultural Science 1992 World Food Prize 1995 Japan Prize 1996 Honorary doctorate, Texas A&M University

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EDWARD F. KNIPLING 91 SELECTED BIBLIOGRAPHY 1949 Insect control investigations of the Orlando, Fla., laboratory during World War II. Smithsonian Annual Report for 1948. Publication 3968, pp. 331-48. 1955 Possibilities of insect control or eradication through the use of sexually sterile males. 7. Econ. Entomol. 48~4~:459-62. 1957 Control of screw-worm eradication fly by atomic radiation. Sci. Mon. 85 (4~: 195-202. 1959 Screw-worm eradication: Concepts and research leading to the ster- ile male method. Smithsonian Annual Report for 1958. Publica- tion 4365, pp. 409-18. Sterile-male method of population control. Science 130~3380~:415- 20. 1960 Use of insects for their own destruction. 7. Econ. Entomol. 53~3~:415- 20. Plans for a comprehensive research program on the boll weevil. Summary-Proceedings: The Cotton Gin and Oil Mill Press 61 (2) :43- 44. The eradication of the screw-worm fly. Sci. Am. 103~4~:54-61. 1962 With L. E. LaChance. Control of populations through genetic ma- nipulations. Ann. Entomol. Soc. Am. 55 (5 ~ :515-20. 1963 A new era in pest control: The sterility principle. Agric. Sci. Rev. 1~1~:2-12.

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92 B I O G RA P H I C A L EMOIRS Opportunities for the development of specific methods of insect control. Proceedings of the XVI International Congress of Zoology 7:14- 26. 1966 Some basic principles in insect population suppression. Bull. Entomol. Soc. Am. 12~1~:7-15. 1968 With T. U. McGuire. Population models to appraise the limitations and potentialities of Trichogramma in managing host insect popu- lations. TB-1387. U.S. Department of Agriculture. Technically feasible approaches to boll weevil eradication. Summary Proceedings. 1968 Beltwide Cotton Production Mechanization Conference, pp. 14-18. 1969 Concept and value of eradication or continuous suppression of in- sect populations. IAEA/FAD panel meeting, sterile-male tech- nique for eradication or control of harmful insects. Vienna, Aus- tria, pp. 19-32. 1970 Suppression of pest Lepidoptera by releasing partially sterile males- A theoretical appraisal. Bioscience, April 15, pp. 495-70. 1972 Use of population models to appraise the role of larval parasites in suppressing Heliothis populations. Technical Bulletin 1434. U.S. Department of Agriculture. Sterilization and other genetic techniques. In Proceedings, Symposium of Pest Control: Strategies for the Future. Washington, D.C.: National Academy of Sciences. Entomology and the management of man's environment. 7. Aust. Entomol. Soc. 2:153-67. 1979 The basic principles of insect population suppression and manage- ment. ESA Agriculture Handbook No. 512. U.S. Department of Agriculture.

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EDWARD F. KNIPLING 93 1983 With E. A. Stadelbacher. The rationale for areawide management of Heliothis (Lepidoptera: Noctuidae) populations. Bull. Entomol. Soc. Am. 29 (4) :29-37. 1984 With R. L. Ridgeway, E. P. Lloyd, and W. H. Cross. Analysis of technology available for eradication of the boll weevil. Agricul- tural Handbook No. 589, pp. 409-435. U.S. Department of Agri- culture. 1 985 Sterile insect technique for screwworm suppression The concept and its development. ESA Miscellaneous Publication No. 62, pp. 4-7. U.S. Department of Agriculture. 1992 Principles of insect parasitism analyzed from new perspectives: Practical implications for regulating insect populations by biological means. Agricultural Research Service Handbook No. 693. U.S. Depart- ment of Agriculture. 1998 Sterile insect and parasite augmentation techniques: Unexploited solutions for many insect problems. Fla. Entomol. 81~1~:134-60.