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Introduction Deciduous orchard pests are increasingly targeted for
classical biological control in an overall integrated pest control effort. Even
partial reductions in the abundance of a pest in orchards can enhance the
effectiveness of cultural methods and bait and pheromone trapping. Principal
crops include almonds, apples, pears, peaches, cherries and walnuts (Minks
& Gruys 1980, Croft 1982, Hoyt et al. 1983, Legner 1983a, 1983b; Legner & Silveira-Guido 1983, Legner &
Goeden 1987, Legner et al. 1982a
, 1982b,; Croft &
AliNiazee 1996). For those pests in which almost no damage to the harvested
fruit is acceptable, control by natural enemies is not sufficient as the sole
source of control. Such pests include the codling moth, Cydia pomonella
L., apple maggot, Rhagoletis pomonella (Walsh),
cherry fruit flies, Rhagoletis cingulata Loew and Rhagoletis
indifferens Curran, oriental fruit moth, Grapholitha
molesta Busck, navel orangeworm, .Amyelois transitella (Walker)
(see Research). In A. transitella
a parasitoid, found in South America is being mass produced and released
annually in orchards. (See PHOTOS of Goniozus
legneri Gordh). Although
biological control can be effective against sporadic and some secondary pests
such as mites, aphids and leafhoppers, the application of nonselective
pesticides for control of a key pest may disrupt the balance in relationships
of secondary pests with natural enemies so that pesticides often are
implemented. At present there is reason to believe that some hasty
applications of pesticides to such apparent imbalances may actually worsen
the situations, and that no chemical treatments might have been the better
strategy. Spiders are general
predators that may well be preserved in orchards to combat phytophagous
pests (Legner 1964 ) In a recent review, Croft and AliNiazee (1996) discuss
biological control of deciduous orchard pests by dividing the effort into key,
sporadic and secondary types. Key pests may attack the fruit
directly and occur at damaging levels annually. Sporadic pests feed on either
the fruit or other part of the tree or both, but are less common; and
secondary pests are infrequently present unless disturbed by pesticides or
other phenomena, and they usually feed on foliage and other non-fruit parts
of the tree (Croft 1982). Biological control research has included various
combinations of classical importation of natural enemies, augmentation of
native and exotic species, conservation through cultural manipulation,
utilization of natural enemy food sprays and selective pesticide deployment. Natural enemies which have been directed against secondary
pests have been most effective in deciduous orchards. Most research has
involved endemic predators and using selective pesticides and improved
cultural methods. Some success has been achieved with genetic improvement of
pesticide resistant strains of predatory species (Hoy et al. 1983, Hoy 1985).
The greatest effort has been in the conservation of natural enemies of plant
feeding mites, aphids, leafminers and leafhoppers; however there has been
little effort to introduce new exotic natural enemies to control these pests.
This is partly because effective endemic species are thought to reside in
most areas which are capable of providing significant control if they are
properly managed. Similarly relatively little effort has been directed to
augmentation because of the high costs involved. Spider Mites.--Spider mites of the family Tetranychidae are important
plant feeders which cause damage to deciduous fruit trees (van de Vrie et al.
1972, van de Vrie 1986). In non commercial orchards where no pesticides or
poor horticultural practices are used, these mites are usually not
problematic because of the activity of predator species. However, in
commercial orchards which are fertilized, pruned and sprayed with pesticides,
these mites often rise to high population levels exceeding 100 per leaf,
resulting in fruit size and number reduction, and quality reduction. Trees
may even be killed after several successive years of attack by dense populations. Spider mites are controlled indirectly by implementing
chemical control that is selective to certain key pests. Several different
groups of predators are involved, such as Phytoseiidae and Stigmaeidae
predators mites, coccinellid beetles in the genus Stethorus,
hemipterans and neuropterans in the Chrysopidae and Hemerobiidae, and some
predaceous thrips, spiders, etc. The most widely used natural enemies are
phytoseiid mites and predatory Stethorus beetles. By monitoring
populations of phytophagous mites, alternate prey and predators, it is
possible to forecast predator/prey ratios early in the growth of pest
populations (Croft 1982). When prey levels exceed certain levels, selective
acaricides may be used to reduce phytophagous mites without influencing
predators (Croft & McGroarty 1977, Mowery et al. 1977). The use of
combinations of predators and selective acaricides can reduce acaricide usage
50-90%, and pesticide resistance development in spider mites can be slowed
(Croft et al. 1987). Alternative prey species of predatory mites may be
enhanced. Examples include Aculus schlechtendali
(Nalepa) as prey for Metaseiulus occidentalis (Nesbitt)
in apples (Madsen 1968, Hoyt 1969), ground cover manipulations to enhance
overwinter survivorship and early dispersal of Amblyseius fallacis
(Garman) into trees (Croft & McGroarty 1977), alternate middle row
application of pesticides to conserve Stethorus punctum
(LeConte) (Hull & Beers 1985), and transferring foliage cuttings from
trees infested with predators to areas lacking Typhlodromus pyri
(Schenten) (Solomon & Easterbrook 1984). Insecticide resistant predatory mites have been deployed
in management with both resistance developed in the field and that resulting
from genetic improvement through hybridization and artificial selection. In
North America, and to some extent in Australia, New Zealand and Europe,
resistant strains of Amblyseius fallacis, Typhlodromus
pyri and Stethorus punctum have been used,
which usually involved inter orchard movement of resistant strains. Successes
with populations of Metaseiulus occidentalis having
resistance to two or more pesticides have been accomplished (Croft &
Strickler 1983, Hoy 1985, Croft & Roush 1988). When resistant natural
enemies are selected in the laboratory, relatively low levels of recessive
and polygenically determined resistance traits are maintained. However, this
resistance is not stable under field conditions and might be lost by
hybridization with susceptible native strains, thus making them difficult to
manage (Croft 1983, Croft & Strickler 1983, Hoy 1985). Aphids.--Several aphid species are major pests in deciduous
orchards, including the apple aphid, Aphis pomi DeGeer,
wooly apple aphid, Eriosoma lanigerum Hausmann), rosy
apple aphid, Dysaphis plantaginea (Passerini) and green
peach aphid, Myzus persicae (Sulzer). Myzus persicae
is a vector of crop pathogens and thus the economic threshold is so low that
biological control is usually not feasible; but possibilities exist with the
other aphid species. The walnut aphid, Chromaphis juglandicola
Kaltenbach and filbert aphid, Myzocallis coryli (Goetze)
have been successfully attacked with classical biological control. Classical biological control of aphids in deciduous
orchards was emphasized for many years. For example, the woolly apple aphid
parasitoid, Aphelinus mali (Hald.), originally found in
eastern North America, was distributed to other parts of the this continent
during 1921-1939 (DeBach 1964). This parasitoids has been introduced to over
50 different countries with successful establishment in about 42 (Clausen
1978). Results of parasitoid establishment were variable, but excellent
control was reported in the Northwestern United States, Australia, Tasmania,
British Columbia, New Zealand, Uruguay and Chile, while moderate control was
reported from Europe and Asia. Attention has recently focused on Aphis
pomi in Massachusetts with an examination of a cecidomyiid
predator, Aphidoletes aphidimyza (Rondani). Adams &
Prokopy (1977, 1980) documented resistance in this predatory fly to some
organophosphate insecticides, and a number of selective pesticides were
deployed. An aphid/predator ratio of 40:1 or lower was necessary to preclude
selective pesticide treatment. Warner & Croft (1982), Morse (1981) and Morse &
Croft (1987) identified selective pesticides and examined predator/prey
relationships of A. aphidimyza and A. pomi
in Michigan apple orchards. Warner & Croft (1984) reported >12-fold
resistance to azinphosmethyl in field populations and found phosalone, phosphamidon,
carbophenothion, pirimicarb and several fungicides and acaricides to be
relatively harmless to this dipteran predator. Morse & Croft (1987) found
that a critical prey/predator ratio of 70-90:1 would provide adequate
biological control of the aphid before an action threshold level of from
40-60 per terminal was reached. Predators such as Forficula auricularia
L. and aphidiid braconid parasitoids have been investigated in Washington
state (Carroll & Hoyt 1984a,b, 1985, 1986). Praon unicum
Smith and Lysiphlebus testaceipes (Cresson) were
significant biological control agents of the aphid early in the season in
some orchards, even though they did not successfully develop through their
entire life cycle while feeding on A. pomi. Alternate
winter and summer aphid hosts of these parasitoids occurred on weeds and
grasses in nearby peach orchards, with Myzus persicae
being one of the major hosts on peaches. An insecticide tolerant earwig, Forficula
auricularia was one important predator that prevented resurgences
of A. pomi following treatments with selective
pesticides. Augmentive releases of the earwig early in the growing season
maintained aphids below 50/tree compared to 2-3,000/tree in orchard plots
where earwigs were excluded or where releases were not made (Carroll &
Hoyt 1986). Effective natural enemies of the woolly apple aphid
include a host specific aphelinid endoparasitoid Aphelinus mali
and a complex of generalist predators in Washington State (Walker 1985). With
the generalist predators excluded, A. mali was incapable of
preventing aphids from soaring to unacceptable levels. The most important
predator was Coccinella transversoguttata Fald in
midsummer, the neuropteran Chrysoperla nigricornis
(Burm.) and the mirid Dareaocoris brevis (Uhler)
throughout the middle and later summer. Other generalist predators such as Adalia
bipunctata (L.) were observed to be important natural enemies in
Oregon (Croft & AliNiazee 1996). French and Spanish strains of Trioxys pallidus
Haliday have been used effectively in Oregon against the filbert aphid, Myzocallis
coryli Goetze (Messing & AliNiazee 1988, Messing 1986),
whereas the French and Iranian strain of T. pallidus
successfully controlled walnut aphid, Chromaphis juglandicola
(Kalt.) in California walnut growing areas, with different strains active in
different areas, as previously discussed (Schlinger et al. 1960, van den
Bosch et al. 1970). Leafminers.--Leafminers in the genus Phyllonorycter
sustain significant parasitism by parasitic Hymenoptera in the families
Braconidae, Pteromalidae, Eulophidae and Ichneumonidae (Hagley et al. 1981.
Wieres et al. 1982, van Driesche et al. 1985, Hagley 1985). There have been
leafminer outbreaks reported from almost all parts of North America
coincident with the development of resistance to pesticides and disruption of
natural enemies (Pree et al. 1980, 1986; Wieres et al. 1982, Maier 1983, van
Driesche et al. 1985, Trimble & Pree 1987). Impacts by the various
leafminer natural enemies have been measured by Johnson et al. (1976), Hagley
(1985), Ridgeway & Mahr (1985) and Barrett & Jorgensen (1986).
Effects of pesticides on them were reported by Hagley et al. (1981), Weires
et al. (1982) and van Driesche et al. (1985). Deployment of pesticides during
periods when natural enemies were least vulnerable has been undertaken
(Hagley et al. 1981, Weires et al. 1982, van Driesche et al. 1985).
Temperature dependent phenological models of the emergence of pest and
natural enemies, such as Sympiesis marylandensis Girault
and Photesor ornigis (Weed), show rather narrow
biological windows of invulnerability for the parasitoids (Drummond et al.
1985). Leafhoppers.--The white apple leafhopper, Typhlocyba pomaria
McAtee and several other less specific species, Empoasca fabae,
Edwardsiana rosae and Erythroneura spp.
attack deciduous orchards in North America (Sayedoleslami 1978, Elsner &
Beers 1987). Resistance to organophosphate insecticides has gradually
increased problems with leafhopper control (Croft & Hoyt 1983).
Biological control has stressed egg and larval parasitoids. In Michigan,
overwintering eggs of T. pomaria are attacked primarily
by the mymirid Anagrus epos Girault, with parasitism
ranging from 20-50% to 100% (Sayedoleslami & Croft 1980). Research has
shown good synchrony between pest and natural enemy throughout orchard tree
scaffolds. Parasitism by the nymphal-adult parasitoid Aphelopus typhlocyba
was lower than egg parasitoids (Sayedoleslami 1978). Parasitoids seem to
tolerate organophosphate insecticides such as azinphosmethyl which has been
in use for a long time, so that field resistance is suspected (Croft 1982).
In Washington, the white apple leafhopper eggs are heavily parasitized by Anagrus
epos with a high degree of synchronization occurring between the
host and parasitoid (Beers & Elsner 1988). Croft and AliNiazee (1996) give some examples of classical
biological control in deciduous orchard environments, such as spider mites,
pear psylla, Psylla pyricola Foerster, codling moth, Cydia
pomonella, apple maggot, Rhagoletis pomonella and other
fruitflies, etc. However, detailed discussions of classical biological
control of these and other orchard pests may be found under each pest's name
separately in the section on case histories (CASEHIST.*). REFERENCES: [ Additional
references may be found at MELVYL Library ] Adams, R. G., Jr. & R. J. Prokopy. 1977. Apple aphid control
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arboreus in biological control of spider mites on apple in western
Oregon. In: Proc. 4th Intern. Congress of Acarology, Saalfeldon,
Austria. AliNiazee, M. T. 1977. Bionomics and life history of a filbert
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