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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
that 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
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of the apple aphid (Homoptera: Aphididae) in Massachusetts. Prot. Ecol. 2:
27-39. AliNiazee, M. T. 1974. Role of predatory mite, Typhlodromus
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