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BIOLOGICAL PEST
CONTROL E. F. Legner Professor of Biological
Control University of California Biological Pest Control emphasizes the introduction of natural
enemies from distant areas. Usually this involves going to other geographic
areas to secure predators, parasitoids and/or pathogens to control the target
pest. The technique has met with numerous successes starting dramatically
with the control of the invaded cuttony-cushion scale. In that and other
dramatic successes the degree of suppression has remained at 99% or more,
with no need to apply any other control measure. There are numerous cases
where the success has been less dramatic, and for these it is necessary to
deploy other control measures, which should be done in harmony with the
controls that are being given by the natural enemies, albeit not to complete
satisfaction. Here considerable knowledge is required to do it properly,
which more often than not is not available. In the latter case we enter the
realm of Integrated Pest Management. When agricultural scientists working out
of colleges and universities are in control there is a greater likelihood of
success. The
biological control of pests with imported natural enemies involves the
addition of new biotic mortality factors to the pest's ecosystem. This
practice is often carefully scrutinized by regulatory agencies, which strive
to eliminate the establishment of potentially harmful organisms. Biological
control researchers continuously seek more effective guidelines for judging a
natural enemy's capabilities before importation in order to accelerate
biological control success rates and to reduce project costs (Coulson 1981).
The manner by which biological control is achieved varies considerably among
projects and the various countries utilizing the technique; and there is a
continuing debate on proper procedures for selection of natural enemies and
regulation of their importation (Legner & Bellows 1999). The
primary goal of federal, state or university importation programs is the
same, i.e., the collection, safe transport, and quarantine processing,
leading ultimately to the colonization in the field of candidate biological
control agents. However, there are differences in the methods, which are, or
can be, used by each entity. Perhaps the main factor in the United States is
that the U. S. Department of Agriculture (APHIS) either on its own initiative
or in concurrence with overriding dicta (as from the Environmental Protection
Agency) issues regulations regarding the importation and quarantine handling
of biological agents which the USDA (ARS), individual states and universities
are expected to follow. Of
mutual concern to the explorer/collector/shipper and government regulatory
agencies and quarantine personnel are the identification of target species
and their hosts, permits to import the material collected, packaging and
labeling, method of shipment, clearance at the port of entry by customs and
agricultural inspectors, and the quarantine facility itself. Most
of the technical and biological considerations relative to acquiring and
shipping biological agents remain much the same as those described for
entomophagous arthropods and /or weed feeders by Bartlett and van den Bosch
(1964), Boldt and Drea (1980), Coulson and Soper (1989), Klingman and Coulson
(1983), and for phytophagous (weed feeders) organisms by Schroeder and Goeden
(1986). In actual operation USDA (ARS) sponsored quarantine laboratories
receive shipments which usually originate from a USDA laboratory abroad where
the material has been screened for contaminants before being shipped to a
primary USDA quarantine facility in the United States, such as the laboratory
at Newark, Delaware, where further screening for unwanted organisms may occur
before the biological agent is forwarded to requestors in the field who may
or may not work out of a secondary quarantine facility where the biological
agent can be propagated or released directly into the field. State
departments of agriculture or universities usually send out members of their
staff as explorer/collectors, who typically do not have access to laboratory
facilities while in the field. As a consequence shipments sent to their
quarantine laboratories may contain more than one targeted pest species and
more than one natural enemy of each of these. They must then be segregated in
quarantine and studied through one generation (for newly introduced species)
before they can be released. Unsolicited extraneous material inadvertently
included may warrant further study in quarantine. If so, specific
arrangements must be made with APHIS PPQ regarding the handling of such
material. USDA collectors when abroad can utilize all available U.S.
governmental facilities (embassies, agricultural attaches, commissary,
vehicles, communication facilities, etc.) to expedite their missions. Thus
far U.S. state and university collectors abroad have only rarely been able to
avail themselves of similar federal cooperation even though their missions
were financed by public funds and their efforts would potentially accrue to
the benefit of agricultural crop production on a regional if not national
scale in the U.S. International
geo-political and socio-economic unrest may impact heavily on the success of
failure of foreign exploration missions. Terrorism in its broadest sense has
become a major deterrent to the search for biological agents in many areas of
the world. Colleagues in such areas or intermediary organizations (i.e.,
charging a fee for service), such as the Commonwealth Institute For
Biological Control, Silwood, UK, may be able to supply the desired beneficial
organisms, but experience has shown that biological control workers who know
what they need and who physically participate in the collecting process tend
to make a better showing in terms of successful introductions (Legner &
Bellows 1999). A
highly important consideration is that during the last 25 years the number of
students trained in biological control and population ecology entomology
worldwide has been on the increase. The hope is that this expanding pool of
"applied ecologists" portends improved international cooperation
regarding greater use of the biological method of pest control. However, it
is anticipated that further legal constraints on biological control of pests
are, or will be, imposed by new and/or pending technical regulations
ostensibly aimed at protecting endangered species or the environment. These
regulations could severely hamper or preclude importation and field use of
new candidate natural enemies. The
purpose for exploration is to search for, import and colonize natural enemies
of our pests from areas where the pest is indigenous, or at least present in
low numbers because its natural enemies keep it in check. The need for
exploration is to protect our environment from needless or questionable use
of chemical pesticides, especially those with long half lives and/or broad
spectrum toxicity which can adversely affect non-target species and
beneficial organisms and ultimately the food chain within a wide range of
biologically diverse species. The
basic goal is to import species of strains presumed to be pre-adapted to
areas targeted for colonization of beneficial organisms. One tries for large
founder numbers in order to keep the gene pool as large as possible. Although
traditionally used for homopterous pests of perennial crops (DeBach 1964), it
is increasingly considered for non-homopterous and annual pests in
agricultural, urban and glasshouse environments. Extra agricultural uses in
medical, forest and household entomology are expanding. Environmental
concerns and laws, public opinion and resistance of arthropod and weed pests
to chemical pesticides are increasingly forcing a consideration and
implementation of non-chemical solutions of pest problems. Classical
biological control is a powerful and proven tool. The increasing threat that
federally mandated regulations may neutralize the importation and
colonization of new natural enemies by greatly slowing the process far beyond
sound biological protocols which have served applied biological control and
society for well over 100 years. Conclusions Natural
enemies for use in biological control may be categorized into separate risk
groups. Parasitic and predaceous arthropods fit into the lowest risk
category, but are the most difficult to study and to assess for potential
success. The policy of certain countries, e.g., Australia, of requiring
intensive studies on native organisms before allowing them to be exported is
especially devastating to the deployment of biological control. A recent case
of invading Australian wood borers that attack eucalyptus in America has
already caused the death of over half of the trees in California, while the
importation of effective natural enemies continues to move at a crawl. Yet
progress is being made with increased attention to basic ecological and
behavioral research. The rate of biological control successes may drop
initially as the style of "educated empiricism" (Coppell &
Mertins 1977) becomes more widely adopted, as has apparently already begun (Hall
& Ehler 1979, Hall et al. 1980). Success rates could be expected to
increase as the database enlarges and intercommunication possibilities
expand. Certainly the trend will ever more propel the activity of exotic
natural enemy importation into a solid scientific base. Andres, L. A. 1981. Conflicting
interests and the biological control of weeds. In: E. S. Del Fosse
(ed.), Proc. 5th Intern. Symp Biological Control of Weeds, 1980, Brisbane,
Qld., CSIRO, Melbourne, Vic., Australia: 11-120. Arthur, A. P. 1966. Associative learning
in Itoplectis conquisitor (Say) (Hymenoptera: Ichneumonidae).
Canad. Ent. 98: 213-23. Attique, M. R., A. I. Mohyuddin, C.
Inayatullah, A. A. Goraya & M. Mustaque. 1980. The present status of
biological control of Chilo partellus (Swinh.) (Lep.:
Pyralidae) by Apanteles flavipes (Cam.) (Hym.: Braconidae) in Pakistan. Proc. 1st
Pakistan Congr. Zool., B: 301-305. Bartlett, B.
R. & R. van den Bosch. 1964. Foreign exploration for beneficial
organisms. In: P. DeBach & E. I. Schlinger (eds.), Biological
Control of Insect Pests and Weeds. Chapman & Hall, London. Beddington, J. R., C. A. Free & J.
H. Lawton. 1978. Characteristics of successful natural enemies in models of
biological control of insect pests. Nature 273: 513-19. Beirne, B. P. 1980a. The human transport
of insect parasites of insects across the Northern Atlantic. Ent. Gen. 6: 267. Beirne, B.
B. 1980b. Biological control: benefits and opportunities. In:
"Perspectives in World Agriculture." Commonw. Agric. Bur., Slough,
England. 307. Bellows, T. S., Jr. & T. W. Fisher,
(eds) 1999. Handbook of Biological Control: Principles and Applications.
Academic Press, San Diego, CA. 1046 p. Birch, L. C.
1971. The role of environmental heterogeneity in determining distribution
and abundance, p. 109-28. In: P. J. den Boer & G. R. Gradwell
(eds.), Dynamics of Populations. Center Agr. Publ. Doc., Wageningen. Boldt, P. E. & J. J. Drea. 1980.
Packaging and shipping beneficial insects for biological control. FAO Plant
Protect. Bull., Vol. 28(2): 64-71. Bucher, G. E. & P. Harris. 1961.
Food-plant spectrum and elimination of disease of Cinnabar moth larvae, Hypocrita
jacobaeae L. (Lepidoptera: Arctiidae). Canad. Ent. 93: 931-36. Bustillo, A. E. & A. T. Drooz. 1977.
Cooperative establishment of a Virginia (USA) strain of Telenomus alsophilae
on Oxydia trychiata in Colombia. J. Econ. Ent. 70: 767-70. Carl, K. P.
1968. Thymelicus lineola (Lepidoptera:
Hesperidae) and its parasites in Europe. Canad. Ent. 100: 785-801. Carl, K. P. 1982. Biological control of
native pests by introduced natural enemies. Biocontrol News & Information
3: 191-200. Cheng, L. 1970. Timing of attack by Lypha
dubia Fall. (Diptera: Tachinidae) on the winter moth Operophtera
brumata (L.) (Lepidoptera: Geometridae) as a factor affecting parasite
success. J. Anim. Ecol. 39: 313-20. Clark, R. C., D. O. Greenbank, D. G.
Bryant and J. W. E. Harris. 1971. Adelges piceae (Ratz.),
balsam woolly aphid (Homoptera: Adelgidae). Tech. Commun. Commonw. Inst.
Biol. Contr. 4: 113-27. Clausen, C.
P. 1956. Biological control of insect pests in the continental United
States. U. S. Dept. Agric. Tech. Bull. No. 1139. 151 p. Clausen, C.
P. (ed.) 1978. Introduced parasites and predators of arthropod pests and weeds:
a world review. Agric. Handb. No. 48, U. S. Dept. Agric., Wash., D.C. 545 p. Cock, M. J. W. 1986. Requirements for
biological control: an ecological perspective. Biocontrol News &
Information 7: 7-16. Common, I. F. B. 1958. A revision of the
pink bollworms of cotton [Pectinophora Busck (Lepidoptera:
Gelechiidae)] and related genera in Australia. Aust. J. Zool. 6(3): 268-306. Coppel, H.
C. & J. W. Mertins. 1977. Biological Insect Pest
Suppression. Springer-Verlag, Berlin, Heidelberg,
New York. 314 p. Coulson, J. R. (ed.). 1981. Use of
beneficial organisms in the control of crop pests. Entomol. Soc. Amer. Publ. Proc.
Joint American-Soviet Conf., Wash., D.C., Aug 13-14, 1979: 62 p. Coulson, J. R. & R. S. Soper. 1988.
Protocols for the introduction of biological control agents in the United
States. p. 1-35. In: R. Kahn (ed.), Plant Quarantine. CRC Press, Boca
Raton, Florida. Croft, B. A. 1970. Comparative studies
on four strains of Typhlodromus occidentalis Nesbitt (Acarina:
Phytoseiidae). Ph.D. Thesis, Univ. of Calif., Riverside. 92 p. Croft, B. A. & M. T. AliNiazee.
1983. Differential tolerance or resistance to insecticides in Typhlodromus
arboreus Chant and associated phytoseiid mites from apple in the
Willamette Valley, Oregon. J. Econ. Ent. 12: 1420-23. Davis, C. J. 1967. Progress in the
biological control of the southern green stink bug, Nezara viridula
smaragdula, in Hawaii. Muschi
(Suppl.): 9-16. DeBach, P.
1964. Successes, trends, and future possibilities (p. 673-713). In:
"Biological Control of Insect Pests and Weeds," P. DeBach (ed.).
Reinhold Publ. Co., New York. 844 p. DeBach, P. 1974. Biological Control by
Natural Enemies. Cambridge Univ. Press, London-New York. 323 p. Drooz, A. T, A. E. Bustillo, G. F. Fedde
& V. H. Fedde. 1977. North American egg parasite successfully controls a
different host in South America. Science 197: 390-91. Ehler, L. E. 1976. The relationship
between theory and practice in biological control. Bull. Ent. Soc. Amer. 22: 319-21. Ehler, L. E.
1979. Assessing competitive interactions in parasite guilds prior to
introduction. Environ. Ent. 8: 558-60. Ehler, L. E. 1982. Foreign exploration
in California. Environ. Ent. 11: 525-30. Ehler, L. E. 1989. Environmental impact
of introduced biological-control agents: implications for agricultural
biotechnology. In: "Risk Assessment in Agricultural
Biotechnology," J. J. Marois & G. Bruening (eds.). Univ. of Calif.,
Div. Agr. & Nat. Res., Oakland, CA. Ehler, L. E.
1990. Introduction strategies in biological control of insects. Crit.
Issues in Biol. Contr., Chap. 6. 1990: 111-134. Ehler, L. E. & L. A. Andres. 1983.
Biological control: exotic natural enemies to control exotic pests (p.
395-418). In: "Exotic Plant Pests and North American
Agriculture," C. L. Wilson & C. L. Graham (eds.). Academic Press,
New York. 522 p. Ehler, L. E.
& R. W. Hall. 1982. Evidence for competitive exclusion of
introduced natural enemies in biological control. Environ. Ent. 11: 1-4. Ehler, L. E.
& J. C. Miller. 1978. Biological control in temporary agroecosystems. Entomophaga 23: 207-212. Eichhorn, O.
1969. Natürliche Verbreitungsareale und
Einschleppungsgebiete der Weisstannen Wolläuse (Gattung Dreyfusia) und
die Möglichkeiten ihrer biologischen Bekämpfung. Z. angew. Ent. 63: 113-31. Eikenbary, R. D. & C. E. Rogers.
1973. Importance of alternate hosts in establishment of introduced parasites.
Proc. Tall Timbers Conf. Ecol. Anim. Control Habitat Management 5: 119-33. Embree, D. G. 1971. The biological
control of the winter moth in eastern Canada by introduced parasites (p.
217-26). In: "Biological Control", C. B. Huffaker (ed.).
Plenum Press, New York. 511 p. Ervin, R. T., L. J. Moffitt, & D. E.
Meyerdirk. 1983. Comstock mealybug (Homoptera: Pseudococcidae): cost analysis
of a biological control program in California. J. Econ. Ent. 76: 605-609. Fedde, G. F, V. H. Fedde & A. T.
Drooz. 1979. Biological control prospects of an egg parasite, Telenomus
alsophilae Viereck, p. 123-27. In: Current Topics in Forest
Entomology. Selected papers from XV Intern. Congr. Entomol., U. S. Dept. Agric. For. Serv. Gen. Tech. Rep. WO-8.
174 p. Fisher, T. W. & G. L. Finney. 1964.
Insectary facilities and equipment, p. 381-401. In: DeBach, P. (ed.),
Biological Control of Insect Pests and Weeds. Reinhold, New York. Force, D. C. 1970. Competition among
four hymenopterous parasites of an endemic insect host. Ann. Ent. Soc. Amer. 63: 1675-88. Force, D. C. 1974. Ecology of insect
host-parasitoid communities. Science 184:
625-32. Franz, J. M.
1961a. Biologische Schädlingsbekämpfung, p. 1-302. In: P. Sorauer
(ed.), "Handbuch der Pflanzenkrankheiten," Band VI. Paul Parey
Verlag, Berlin-Hamburg. 627 p. Franz, J. M.
1961b. Biological control of pest insects in Europe. Ann. Eve. Ent. 6: 183-200. Franz, J. M.
1973a. Quantitative evaluation of natural enemy effectiveness.
Introductory review of the need for evaluation studies in relation to
integrated control. J. Appl. Ecol. 10: 321-23. Franz, J. M.
1973b. The role of biological control in pest management. Bull. Lab.
Ent. Agraria 30: 235-43. Franz, J. M.
& A. Krieg. 1982. Biologische Schädlingsbekämpfung, 3 Auflage. Verlag
Paul Parey, Berlin-Hamburg. 252 p. Ghani, M. A. 1969. natural enemies of
forage and grain legume aphids in Pakistan. Ann. Rep. Commonw. Inst. Biol.
Contr. Pakistan Sta. Rept. (unpub.) Goeden, R.
D. 1971. Insect ecology of silverleaf nightshade. Weed Sci. 19: 45-51. Goeden, R. D. 1983. Critique and
revision of Harris' scoring system for selection of insect agents in
biological control of weeds. Prot. Ecol.
5: 287-301. Goeden, R.
D. 1988. A capsule history of biological control of weeds. Biocontrol News
& Information. 9(2): 55-61. Goeden, R.
D. & L. T. Kok. 1986. Comments on a proposed "new"
approach for selecting agents for the biological control of weeds. Canad.
Ent. 118: 51-58. Greathead, D. J. 1971. A Review of
Biological Control in the Ethiopian Region. Commonw. Inst. Biol. Contr. Tech.
Commun 5. 162 p. Greathead, D. J. 1973. Progress in the
biological control of Lantana camara in East Africa and
discussion of problems raised by the unexpected reaction of some of the more
promising insects to Seasamum indicum (p. 89-92). In:
Proc. 2nd Int. Symp. Biol. Contr. Weeds, P. H. Dunn (ed.). Commonwealth Inst.
Biol. Control Misc. Publ. 6. Gruys, P. 1971. Mutual interference in Bupalus
pinarius, p. 199-207. In: P. J. den Boer & G. R. Gradwell
(eds.), Dynamics of Populations. Center Agr. Publ. Doc., Wageningen. Hagen, K. S.
& J. M. Franz. 1973. A history of biological control. Ann. Rev. Ent. 18: 433-76. Hall, R. W.
& L. E. Ehler. 1979. Rate of establishment of natural enemies in classical
biological control. Bull. Ent. Soc. Amer. 25: 280-282. Hall, R. W.,
L. E. Ehler & B. Bisabri-Ershadi. 1980. Rate of success in
classical biological control of arthropods. Bull. Ent. Soc. Amer. 26: 111-14. Harris, P. 1973a. Selection of effective
agents for the biological control of weeds, p. 29-34. In: Proc. 2nd Int.
Symp. Biol. Contr. of Weeds., Misc. Publ. Commonw. Inst. Biol. Contr. 6. Harris, P. 1973b. The selection of
effective agents of the biological control of weeds. Canad. Ent. 105:
1495-1503. Harris, P. & H. Zwolfer. 1968.
Screening of phytophagous insects for biological control of weeds. Canad.
Ent. 100: 295-303. Harris, P., D. Peschken & J. Milroy.
1969. The status of biological control of the weed Hypericum perforatum
in British Columbia. Canad. Ent. 101: 1-15. Hassan, S. 1970. The possible control of
skeleton weed, Chondrilla juncea L., using Puccinia chondrillina
Bubak & Syd. Proc. 1st Int. Symp. Biol. Contr. of Weeds, p. 11-14. Misc.
Publ. Commonw. Inst. Biol. Contr. 1. Hassell, M.
P. 1969a. A study of the mortality factors acting upon Cyzenis
albicans (Fall.), a tachinid parasite of the winter moth, Operophtera
brumata (L.). J. Anim. Ecol. 38: 329-39. Hassell, M. P. 1969b. A population model
for the interaction between Cyzenis albicans (Fall.)
(Tachinidae) and Operophtera brumata (L.) (Geometridae) at
Wytham, Berkshire. J. Anim. Ecol. 38:
567-76. Hassell, M.
P. 1971. Parasite behaviour as a factor contributing to the stability of
insect host-parasite interactions, p. 366-79. In: P. J. den Boer &
G. R. Gradwell (eds.), Dynamics of Populations. Center Agr. Publ. Doc.,
Wageningen. Hassell, M.
P. 1978. The Dynamics of Arthropod Predator-Prey Systems. Princeton Univ.
Press, Princeton, New Jersey. Hassell, M.
P. 1980. Foraging strategies, population models and biological control: a
case study. J. Anim. Ecol. 49: 603-28. Hassell, M. P. & H. N. Comins. 1978.
Sigmoid functional response and population stability. Theor. Pop. Biol. 14:
62-67. Hassell, M. P. & R. M. May. 1973.
Stability in insect host-parasite models. J. Anim. Ecol. 42: 693-726. Hokkanen, H.
1985a. Exploiter-victim relationships of major plant diseases:
implications for biological weed control. Agriculture Ecosystems &
Environment 14: 63-76. Hokkanen, H. M. T. 1985b. Success in
classical biological controls. CRC Crit. Rev. in Plant Sci., Vol 3(1): 35-72. Hokkanen, H.
& D. Pimentel. 1984. New approach for selecting biological control agents.
Canad. Ent. 116: 1109-1121. Hokkanen, H. M. T. & D. Pimentel.
1989. New associations in biological control: theory and practice. Canad.
Ent. 121: 829-40. Howarth, F. G. 1985. Impacts of alien
land arthropods and mollusks on native plants and animals in Hawaii. In:
"Hawaii's Terrestrial Ecosystems: Preservation and Management," C.
P. Stone & J. M. Scott (eds.). pp. 149-178. Univ. of Hawaii Press,
Honolulu. Hoy, M. A.
1985. Improving establishment of arthropod natural enemies. In:
"Biological Control in Agricultural IPM Systems," M. A. Hoy &
D. C. Herzog (eds.). pp. 151-166. Academic Press, New York. Hoy, M. A.,
D. Castro & D. Cahn. 1982. Two methods for large-scale production
of pesticide-resistant strains of the spider mite predator Metaseiulus
occidentalis. Z. angew. Ent. 94:
1-9. Huettel, M.
D. & G. L. Bush. 1972. The genetics of host selection and its
bearing on sympatric speciation in Procecidochares (Dipt.: Tephritidae). Ent.
Exp. Appl. 15: 465-80. Hughes, R. D. 1973. Quantitative
evaluation of natural enemy effectiveness. J. Appl. Ecol. 10: 321-51. Huber, R. D.
1973. Quantitative evaluation of natural enemy effectiveness. J. Appl.
Ecol. 10: 321-51. Huber, P. 1983. Exorcists vs gatekeepers
in risk regulation. Regulation 7(6): 23-32. Huffaker, C. B. 1957. Fundamentals of
biological control of weeds. Hilgardia
27: 101-157. Huffaker, C.
B., P. S. Messenger & P. DeBach. 1971. The natural enemy component
in natural control and the theory of biological control (p. 16-17). In:
"Biological Control," C. B. Huffaker (ed.). Plenum Press, New York.
511 p. Huffaker, C. B. 1954. Introduction of
egg parasites of the beet leafhopper. J. Econ. Ent. 47: 785-89. Huffaker, C.
B., C. E. Kennett, B. Matsumoto & E. G. White. 1968. Some
parameters in the role of enemies in the natural control of insect abundance,
p. 59-75. In: T. R. E. Southwood (ed.), Insect Abundance. Blackwell,
Oxford. Huffaker, C. B., P. S. Messenger &
P. DeBach. 1971. The natural enemy component in natural control and the
theory of biological control, p. 16-67. In: C. B. Huffaker (ed.),
Biological Control. Plenum Press, New York. Inman, R. E.
1970a. Problems in searching for and collecting control organisms. Proc.
1st Int. Symp. Biol. Contr. Weeds, p. 105-08. Misc. Publ. Commonw. Inst. Biol. Contr. 1. Inman, R. E.
1970b. Host resistance and biological weed control. Proc. 1st Int. Symp.
Biol. Contr. Weeds, p. 41-5. Misc. Publ. Commonw. Inst. Biol. Contr. 1. Julien, M.
M. (ed.). 1982. Biological control of weeds: a world catalogue of
agents and their target weeds. Commonw. Agric. Bur., Farnham Royal, Slough,
U.K. 197 p. Klingman, D. L. & J. R. Coulson. 1982.
Guidelines for introducing foreign organisms into the United States for
biological control of weeds. Weed Sci. 30: 661-67. Legner, E. F. 1978. Natural enemies imported in California
for the biological control of face fly, Musca
autumnalis DeGeer, and horn fly, Haematobia irritans
(L.). Proc. Calif. Mosq. & Vector Contr. Assoc., Inc. 46: 77-79. Legner, E. F. 1978. Part I: Parasites and predators
introduced against arthropod pests. Diptera. In: Introduced Parasites and Predators of
Arthropod Pests and Weeds: a World Review (C. P. Clausen, ed.), pp. 335-39;
346-55. Agric. Handbk. No. 480, ARS, USDA, U. S.
Govt. Printing Off., Wash., D. C. 545 pp Legner, E. F. 1986. The requirement for reassessment of interactions
among dung beetles, symbovine flies and natural enemies. Entomol. Soc. Amer. Misc. Publ. 61: 120-131. Legner, E. F.
1986. Importation of exotic natural enemies. In: pp. 19-30, "Biological Control of Plant Pests and of
Vectors of Human and Animal
Diseases." Fortschritte der
Zool. Bd. 32: 341 pp. Legner, E. F.
& T. S. Bellows, Jr.. 1999. Exploration for natural enemies. In: T. W. Fisher & T. S.
Bellows (eds.), Chapter 15, p. 87- 101., Handbook of Biological Control: Principles
and Applications. Academic Press, San Diego, CA 1046 p. Legner, E. F. & R. A. Medved. 1973. Influence of Tilapia mossambica (Peters), T.
zillii (Gervais) (Cichlidae) and Mollienesia latipinna LeSueur
(Poeciliidae) on pond populations of Culex
mosquitoes and chironomid midges. J. Amer. Mosq. Contr. Assoc. 33(3): 354-364. Legner, E. F. & R. A. Medved. 1979. Influence of parasitic
Hymenoptera on the regulation of pink bollworm, Pectinophora gossypiella, on cotton in the lower Colorado
Desert. Environ. Entomol. 8(5): 922-930. Legner, E. F.
& R. W. Warkentin. 1983. Questions concerning the dynamics of Onthophagus gazella (Coleoptera: Scarabaeidae) with symbovine flies
in the lower Colorado Desert of California. Proc. Calif. Mosq. & Vector
Contr. Assoc., Inc. 51: 99-101. Legner, E. F. & R. D. Sjogren. 1984. Biological mosquito
control furthered by advances in technology and research. J. Amer. Mosq. Contr. Assoc. 44(4): 449-456. Legner, E. F., R. D. Sjogren & I. M. Hall. 1974. The biological
control of medically important arthropods. Critical Reviews in Environmental
Control 4(1): 85-113. Legner, E. F., R. A. Medved & W. J. Hauser. 1975. Predation
by the desert pupfish, Cyprinodon macularius on Culex mosquitoes and benthic
chironomid midges. Entomophaga 20(1): 23-30. Legner, E. F.,
R. A. Medved & F. Pelsue. 1980. Changes in chironomid breeding patterns
in a paved river channel following adaptation of cichlids of
the Tilapia mossambica-hornorum
complex. Ann. Entomol. Soc. Amer. 73(1):
293-299. Lucas, A. M.
1969. The effect of population structure on the success of insect
introductions. Heredity 24: 151-57. Luck, R. F. 1982. Parasitic insects
introduced as biological control agents for arthropod pests (p. 125-284). In:
CRC Handb. Pest Management in Agriculture, D. Pimentel (ed.). Vol II, CRC
Press, Boca Raton, Florida. Luck, R. F., P. S. Messenger & J.
Barbieri. 1981. The influence of hyperparasitism on the performance of
biological control agents. p. 33-42. In: D. Rosen (ed.), The Role of
Hyperparasitism in Biological Control: a Symposium. Univ. of Calif. Div.
Agric. Sci. Macqueen, A. 1975. Dung as an insect
food source: dung beetles as competitors of other coprophagous fauna and as
targets for predators. J. Appl. Ecol. 12: 821-27. May, R. M. & M. P. Hassell. 1981.
The dynamics of multiparasitoid-host interactions. Amer. Nat. 117: 234-261. May, R. M. & M. P. Hassell. 1988.
Population dynamics and biological control. Phil. Trans. Roy. Soc. London B
318: 129-169. McMurtry, J. R., E. R. Oatman, P. A.
Phillips and C. W. Wood. 1978. Establishment of Phytoseiulus persimilis
(Acari: Phytoseiidae) in southern California. Entomophaga 23: 175-79. Messenger,
P. S. 1971. Climatic limitations to biological controls (p. 97-114). In:
Proc. Tall Timbers Conf. Ecol. Anim. Contr.
Habitat Manag. 3. Tallahassee, Florida. Meyerdirk, D. E. & I. M. Newell.
1979. Importation, colonization and establishment of natural enemies on the
Comstock mealybug in California. J. Econ. Ent. 72: 70-73. Meyerdirk, D. E., I. M. Newell & R.
W. Warkentin. 1981. Biological control of Comstock mealybug. J. Econ. Ent. 74: 79-84. Miller, J.
C. 1983. Ecological relationships among parasites and the practice of biological
control. Environ. Ent. 74: 79-84. Mohyuddin, A. I. 1971. Comparative
biology and ecology of Apanteles flavipes and A. sesamiae
as parasites of graminaceous borers. Bull. Ent. Res. 61: 33-9. Mohyuddin, A. I., C. Inayatullah &
E. G. King. 1981. Host selection and strain occurrence in Apanteles flavipes
(Cameron) (Hymenoptera: Braconidae) and its bearing on biological control of
graminaceous stem-borers (Lepidoptera: Pyralidae). Bull. Ent. Res. 71:
575-581. Murakami, Y. 1966. Studies on the natural
enemies of the Comstock mealybug. II. Comparative biology on two types of
internal parasites, Clausenia purpurea and Pseudaphycus melinus
(Hymenoptera, Encyrtidae). Bull. Hort. Res. Sta. Ser. A, 5: 139-163. Murakami, Y. & H.-B. Ao. 1980.
Natural enemies of the chestnut gall wasp in Hopei Province, China
(Hymenoptera: Chalcidoidea). Appl. Ent. Zool. 15: 184-86. Murakami, Y., K. Umeya & N. Oho.
1977. A preliminary introduction and release of a parasitoid (Chalcidoidea,
Torymidae) of the chestnut gall wasp, Dryocosmos kuriphilus
Yasumatsu (Cynipidae) from China. Japan J. Appl. Ent. 21: 197-203. Naumann, I.
D. & D. P. A. Sands. 1984. Two Australian Elasmus spp.
(Hymenoptera: Elasmidae), parasitoids of Pectinophora gossypiella
(Saunders) (Lepidoptera: Gelechiidae): their taxonomy and biology. J. Aust.
Ent. Soc. 23: 25-32. Oatman, E. R. and G. R. Platner. 1974.
Parasitization of the potato tuberworm in southern California. Environ. Ent. 3: 262-64. Oman, P. W.
1948. Notes on the beet leafhopper Circulifer tenellus
(Baker) and its relatives. J. Kansas Ent. Soc. 21: 10-14. Osborne, J. A. 1982. Efficacy of Hydrilla
control and a stocking model for hybrid grass carp in freshwater lakes. Off.
of Exploratory Res. (RD-675), U. S.
Environ. Prot. Agency, Wash., D.C. 143 p. Pimentel, D. 1963. Introducing parasites
and predators to control native pests. Canad. Ent. 92: 785-92. Pimentel, D. 1973. Comments on present
status of biological agents. WHO/VBC/73.445. In: Conf. on the Safety
of Biological Agents for Arthropod Control. p. 9. Pimentel, D. 1980. Environmental risks
associated with biological controls, p. 11-24. In: B. Lundholm &
M. Stackerud (eds.), Environmental Protection and Biological Forms of Control
of Pest Organisms. Ecol. Bull. 31, Stockholm. Pimentel, D.
1988a. Improved success in biological control, p. 1-3. In:
International Conference "Biological Control of Vectors with Predaceous
Arthropods." 7-10 Jan. Loyola College, Madras, India. Pimentel, D. 1988b. Improved success in
biological control. Bicovas 1: 90--3. Pimentel, D.
& H. Hokkanen. 1989. Alternative for successful biological control in
theory and practice, p. 21-51. In: E. L. Kulhavy & M. C. Miller
(eds.), Potential for Biological Control of Dendroctonus and Ips
Bark Beetles. Center for Applied Studies, School of Forestry, Stephen F.
Austin St. Univ., Nacogdoches, Texas. Pimentel,
D., C. Glenister, S. Fast & D. Gallahan. 1983. An
environmental risk assessment of biological and cultural controls for organic
agriculture, p. 73-90. In: W. Lockeretz (ed.), Environmentally Sound
Agriculture. Praeger Special Studies, N.Y. Pimentel, D., C. Glenister, S. Fast
& D. Gallahan. 1984. Environmental risks of biological pest controls.
Oikos 42: 283-90. Price, P. W. 1972. Methods of sampling
and analysis for predictive results in the introduction of entomophagous
insects. Entomophaga 17: 211-22. Pschorn-Walcher,
H. 1973. Die Parasiten der gesellig lebenden Kiefern-Buschhornblattwespen (Fam.
Diprionidae) als Beispiel für Koexistenz und Konkurrenz in multiplen
Parasit-Wirt-Komplexen. Verh. deut. Zool. Ges. (Jahresversammlung) 66:
136-45. Pschorn-Walcher,
H. & H. Zwölfer. 1956. The predator complex of the white-fir woolly aphids
(Gen. Dreyfusia, Adelgidae). Z. angew. Ent. 39: 63-75. Pschorn-Walcher,
H., D. Schröder & O. Eichhorn. 1969. Recent attempts at biological
control of some Canadian forest insect pests. Tech. Bull. Commonw. Inst.
Biol. Contr. 11: 1-18. Quezada, J.
R. & P. DeBach. 1973. Bioecological and population studies of the
cottony-cushion scale, Icerya purchasi Mask., and its natural
enemies, Rodolia cardinalis Muls., and Cryptochaetum iceryae
Will., in southern California. Hilgardia 41: 631-88. Ratcliffe, F. N. 1966. Biological
control. Aust. J. Sci. 28: 237-40. Remington, C. L. 1968. The population
genetics of insect introduction. Ann. Rev. Ent. 13: 415-26. Ross, H. H. 1953. On the origin and
composition of the Nearctic insect fauna. Evolution 7: 145-58. Roth, J. P., G. T. Fincher & J. W.
Summerlin. 1983. Competition and predation as mortality factors of the horn
fly, Haematobia irritans (L.) (Diptera: Muscidae) in a central
Texas pasture habitat. Environ. Ent. 12: 106-109. Sailer, R. I. 1981. Elements of
opportunity in biological control. In: "Biological Control in
Crop Production," G. C. Papavizas (ed.). Granada Publ. Co., London. 419. Sands, D. P. A. & A. R. Hill. 1982.
Surveys for parasitoids of Pectinophora gossypiella (Saunders) (Lepidoptera:
Gelechiidae) in Australia. Commonw. Scien. Ind. Res. Org., Div. Ent. Rept.
No. 29: 1-18. Schröder. D. 1974. A study of the
interactions between the internal larval parasites of Rhyacionia buoliana
(Lep. Olethreutidae). Entomophaga 19: 145-71. Schroeder, D. & R. D. Goeden. 1986.
The search for arthropod natural enemies of introduced weeds for biological
control--in theory and practice. Biocontrol News and Information 7(3):
147-155. Sechser, B.
1970. Der Parasitenkomplex des Kleinen Frostspanners (Operophthera brumata
L.) (Lep., Geometr.) under besonderer Berücksichtigung der Kokonparasiten.
Teil I und II. Z. angew. Ent. 66: 1-35, 144-60. Sheldeshova, G. G. 1967. Ecological
factors determining distribution of the codling moth, Laspeyresia pomonella
L. (Lepidoptera: Tortricidae) in the northern and southern hemispheres. Ent.
Rev. 46: 349-61. Simmonds, F. J. 1949. Insects attacking Cordia
macrostachya (Jacq.) Roem & Schult in the West Indies. 1. Physonota
alutacea Boh. (Col., Cassididae). Canad. Ent. 81: 185-99. Simmonds, F. J. 1969. Commonwealth
Institute of Biological Control. Brief Resume of Activities and Recent
Successes Achieved. Commonw. Agr. Bureaux Publ. Ferozsons Ltd., Rawalpindi.
16 p. Smith, H. S. 1939. Insect populations in
relation to biological control. Ecol. Monogr. 9: 311-20. Taylor, T. H. C. 1937. The Biological
Control of an Insect in Fiji. An Account of the Coconut Leaf-Mining Beetle
and its Parasite Complex. Imp. Inst. Ent., London. 239 p. Turnbull, A. L. 1967. Population
dynamics of exotic insects. Bull. Ent. Soc. Amer. 13: 333-37. Turnbull, A. L. & D. A. Chant. 1961.
The practice and theory of biological control of insects in Canada. Canad. J.
Zool. 39: 697-753. Turnock, W. J. & J. A. Muldrew.
1971. Pristiphora erichsonii (Hartig), larch sawfly
(Hymenoptera: Tenthredinidae), p. 175-94. In: Biological Control
Programs Against Insects and Weeds in Canada 1959-1968. Tech. Commun.
Commonw. Inst. Biol. Contr. 4. van den
Bosch, R. 1968. Comments on population dynamics of exotic insects. Bull.
Ent. Soc. Amer. 14: 112-115. van den
Bosch, R. 1971. Biological control of insects. Ann. Rev. Ecol. Syst. 2:
45-66. van Lenteren, J. C. 1980. Evaluation of
control capabilities of natural enemies: Does art have to become science?
Neth. J. Zool. 30: 369-381. Waage, J. K.
& M. P. Hassell. 1982. Parasitoids as biological control agents--a
fundamental approach. Parasitol. 82: 241-68. Walters, L. L.
& E. F. Legner. 1980. Impact of the desert pupfish, Cyprinodon macularius,
and Gambusia affinis on fauna in pond ecosystems.
Hilgardia 48(3): 1-18. Wapshere, A. J. 1970. The assessment of
the biological control of organisms for controlling weeds. Proc. 1st Int.
Symp. Biol. Contr. Weeds, p. 79-89. Misc. Publ. Commonw. Inst. Biol. Contr.
1. Waterhouse, D. F. 1974. The biological
control of dung. Scien. Amer. 230: 100-109. Watt, K. E. F. 1965. Community stability
and the strategy of biological control. Canad. Ent. 97: 887-895. Wilson, A. G. L. 1972. Distribution of
pink bollworm, Pectinophora gossypiella (Saund.), in Australia
and its status as a pest in the Ord irrigation area. J. Aust. Inst. Agric. Sci. 38: 95-9. Zwölfer, H. 1961. A comparative analysis
of the parasite complexes of the European fir budworm, Choristoneura murinana
(Hb.), and the North American spruce budworm, C. fumiferana
(Clem.). Tech. Bull. Commonw. Inst. Biol. Contr. 1: 1-162. Zwölfer, H. 1971. The structure and
effect of parasite complexes attacking phytophagous host insects (p. 405-18).
In: "Dynamics of Populations," P. J. den Boer & G. R.
Gradwell (eds.). Cent. Agric. Publ. Doc., Wageningen. Zwölfer, H. & P. Harris. 1971. Host
specificity determination of insects for biological control of weeds. Ann.
Rev. Ent. 16: 159-78. Zwölfer, H., M. A. Ghani & V. P. Rao. 1976. Foreign
exploration and importation of natural enemies (p. 189-207). In:
"Theory and Practice of Biological Control," C. B. Huffaker & P. S. Messenger
(eds.). Academic Press, New York. 788 p. |