Codling Moth Information Support System (CMISS)

Natural Enemies of Codling Moth and Leafrollers of Pome and Stone Fruits






1.Codling Moth

1.1- Principal characteristics of the pest.

The codling moth, Cydia pomonella (Linnaeus), is the most widely distributed pest of cultivated pome fruits and walnuts in the world, except in Japan (Barnes, 1991) and in the western part of Australia where it has been eliminated, being a key pest in most situations. Its origin is Eurasian. The distribution of the codling moth is especially affected by ecological factors: the northern boundary is defined by adequate heat summation above 10°C in summer of approximately 600 degree-days (thermal accumulation required for development of one generation). In both northern and southern hemispheres, its limits approaching the equator are approximately at the 25th parallels (Shel. deshova, 1967). The most important codling moth hosts are apple and pear, but it can also be a key pest in quince, walnut, apricot, plum, peach, nectarine, and even other Prunus species (like sweet cherry and almonds).

The life cycle of codling moth has two special features (Audemard, 1976):

  1. The number of annual generations varies from one to four according to the climate (varying with latitude and altitude), the year, and sometimes the host plant.
  2. It always overwinters as mature larvae diapausing in a cocoon-sheltered situation on a tree or in the ground. Alterations in the time of diapause (induced basically by the photoperiod, although temperature, food source, and some other variables can have some influence) allows the codling moth to adjust its life cycle to the climate and to host-plant phenology.
For a more complete description of its biological cycle and ecological aspects, see Bovey (1966), Shel-deshova (1967), and Audemard (1976; 1977).

Different methods of control have been used against the codling moth (Croft & Riedl, 1991). Chemical control has been the most extended method for a long time. After appearing resistant to DDT, the typical compounds used have been organophosphates (azinphosmethyl, basically, but also phosmet, diazinon, and phosalone) and carbamates (carbaryl). Now there are some reports from California and Washington about populations becoming resistant to azinphomethyl and other organophosphates.

In some areas, pyrethroids have been increasing used for codling moth control due to their effectiveness at low rates and development of resistance against organophosphates. Pyrethroids include compounds such as bifenthrin, cyfruthrin, cypermethrin, decamethrin, esfenvalerate, enpropathrin, fenvalerate, flucythrinate, fluvalinate, permethrin, and others. Insect growth regulators (IGRs) diflubenzuron, triflumuron, chlorfluazuron, and teflubenzuron have also shown efficacy against codling moth both in laboratory and field tests. Avermectin, a fermentation byproduct from Streptomyces avermitilis, also can be effective, especially in the early season against the neonates predominantly on pear (Croft & Riedl, 1991).

Other methods are the use of mechanical control with trap bands, cultural control (early varieties, resistant varieties), biotechnical control (releases of sterile males, mating disruption with pheromones, mass trapping with pheromones), along with biological control with natural enemies.
 
 

1.2.- Natural Enemies.

The natural enemies of the codling moth include birds, spiders, insects, nematodes, bacteria, fungi, protozoa, and viruses (Falcon & Huber, 1991). Natural enemies are most effective in reducing codling moth population at three times during its life cycle: egg stage, newly hatched larvae, and the wintering larvae (MacLellan, 1969).

Predators:

The most important predators of the codling moth are birds, bats, spiders, insects, and some mite species. All of them can feed on both codling moth eggs and young larvae (before entering the fruit).

Birds:

Birds are the most important group that prey on codling moth. Although there is evidence that larvae and pupae are the only stages attacked, it is also probable that birds capture and feed upon adult codling moth (Falcon & Huber, 1991).

MacLellan (1971) reported that two woodpecker species, Dendrocopus pubescens medianus (Swanson) and Dendrocopus villosus villosus (Linnaeus), are the most important natural enemies in Nova Scotia, because they effectively attack the codling moth when they are at very low density. These species attack the codling moth just as the larvae leave the fruit to spin cocoons, and even when they are in the cocoons on the trunk of the trees. Their activity starts in August, increases in early fall, and reaches a peak in November, following which the numbers of larvae available decline as the winter advances. In Ohio, the most common species observed preying on codling moth were the white breasted nuthatch, Sitta carolinensis Latham, and the brown creeper, Certhia familiaris americana Bonaparte, while the downy woodpecker, Picoides pubescens Linnaeus, seems to be less common.

In Quebec Province, Canada, different woodpeckers species, Dendrocopus spp., and the wilow-tit, Parus atricapillus Linnaeus, were found to be the most important birds attacking larvae and diapausing larvae (LeRoux & Perron, 1960; Mailloux & LeRoux, 1962), but LeRoux (1961) observed than only predation by woodpeckers is relatively important in winter in Canada.

The great tit, Parus major Linnaeus, was considered by Massee (1954) the most important species of bird attacking codling moth larvae in Britain. This species, along with the blue tits, Parus caeruleus Linnaeus, was also found to be the most important by Solomon et al.(1976) in different unsprayed apple orchards in Europe.

In other studies from New Zealand, Wearing (1975) found that the silvereye, Zosterops lateralis (Latham), was the principal species attacking codling moth larvae and pupae in cocoons on apple trees. The predation of this bird on the larvae in cocoons has been shown to be density dependent (Wearing, 1979).
 
 

Spiders:

Numerous generalist spider species feed on codling moth. Dondale et al. (1979) found 41 species preying on codling moth. The most important species regulating codling moth populations were Theridion muriarium Emerton, Araniella displicata (Hentz), and Philodromus rufus Walckenaer. They prey on all life stages of the codling moth.
 
 

Insects:

Codling moth can be attacked by insects from different orders. The most important codling moth predator species are Neuroptera, Thysanura, and Heteroptera, although some Coleoptera (like coccinelids and pentatomids) also can have an important impact on codling moth population levels. All these predators are very generalist, so the impact on codling moth population will depend greatly on the relative availability of other food sources.
 
 

Neuroptera:

In the U.S.A., generalist neuropters (Summerland & Steiner, 1943) were found to be the most important groups of predators, attacking both egg and first larval stages of the codling moth.
 
 

Thysanura:

Like Neuroptera, different species belonging to this order have been found to be very important as codling moth predators, attacking both egg and first larval stages of the codling moth (Jaynes & Marrucci, 1947).
 
 

Hymenoptera:

Different ants species [Formica fusca var. subsericea Say, Solenopis molesta (Say), Monomorium minimum (Buckley), Aphaenogaster fulva aquia (Buckley) and Tetramoriuim caespitum (Linnaeus)] have also been reported by Jaynes & Marucci (1947) to feed on first instar codling moth larvae.
 
 

Heteroptera (Hemyptera):

In Britain and Nova Scotia (Canada), heteropterans seem to be the most important codling moth predators (MacLellan, 1962, 1963; Glen, 1975). These predators feed on both eggs and young codling moth larvae, and can act as important codling moth population regulators (MacLellan 1963). The most important heteropterous species attacking the codling moth have been found to be Miridae and Anthocoridae. Within the mirid species, the most important are: Blepharidopterus angulatus (Fallen), Malacoris chlorizans (Panzer), Phytocoris tiliae (Fabricius), Deraeocoris fasciolus Knight, Deraeocoris nebulosus (Uhler), Deraeocoris brevis, Diaphnocoris spp, Hyaliodes harti Knight, Phytocoris perplexus (Denjis and Schiffermuller), and Plagiognathus obscurus Uhler. The most impotant anthocorid species are: Anthocorus nemorum (Linnaeus), Orius minutus (Linnaeus), Anthocorus nemoralis (Fabricius), and Orius insidiousus (Say).
 
 

Dermaptera:

The European earwig, Forficula auricularia (Linnaeus), can make an important contribution to codling moth mortality (Glen, 1975). It preys on codling moth eggs. Although it has relative high potential as a biological control agent, the fruit damage that it can cause makes it a natural enemy of little value from an IPM perspective.
 

Thysanoptera:

The species Haplothrips faurei Hood and Leptothrips mali (Fitch) can attack codling moth eggs and young larvae.
 

Mites:

The mite Anystis agilis Banks have been found (MacLellan, 1972) to attack both eggs and young codling moth larvae.

[Picture: http://www.ipm.ucdavis.edu/PMG/A/I-AC-AAGI-AD.001.html].
 
 

Parasitoids:

The most important codling parasitoids are hymenopterous, but some dipterous species have also been found to parasite the codling moth. They have been found to attack all codling moth life stages with the exception of the adult.

Ascogaster quadridentata Wesmaels, that belong to the genus Trichogramma (Braconidae) is the most important parasitoid of codling moth eggs and the natural enemy with a bigger potential in IPM programs. The adult female laid the eggs in the codling moth egg and the larva develops during the egg and larval stages of the host (Charmillot et al., 1997). It is also widely found in the center regions of Europe (France, Austria, Switzerland). With low populaton levels of the codling moth, it achieves parasitism levels up to 5%, that shows its efficiency in searching behavior by the adult female. Furthermore, as host levels increase, the percentage of parasitism also rises. MacLellan (1969) reported a maximum of 82.5% parasitism. The intensity of parasitism by this egg parasitoid depends on the stage of embryonic development of the eggs: Rupf & Russ (1976) observed, at 25°C and 70-80% RH, that the maximum degree of parasitism was when eggs were 2-4 days old.

Other Trichogramma species parasite codling moth eggs in different areas of the world, but their importance as biological control agents is minor. The most important of these species are: Trichogramma minutum Riley and Trichogramma platneri in South Ontario, Canada (Yu et al., 1984), Indiana, U.S.A. (Dolphin & Cleveland, 1966), Nova Scotia (MacLellan, 1963), and Australia (Wilson, 1962); Trichogramma evanescens Westwood in Turkey (Iren & Gurkan, 1971); Trichogramma embryophagum Hartig in Poland (Kot, 1962); and Trichogramma cacoeciae Marchal.

Larval parasitoids include Cryptus sexmaculatus Gravenhorst (Simmonds, 1944) and Elodia tragica (Meigen) (Turnbull & Chant, 1961). Ephialtes extensor Taschenberg and Ephialtes caudatus Ratzeburg are ectoparasitoids that laid their eggs on the interior of the cocoon (Coutin, 1960). Pimpla aquilonia Cresson and Pimpla turionellae (Linnaeus) can attack pupae. P.turionellae is an ichneumonid that also lays eggs in young codling moth larvae and its larvae develops as a solitaire endoparasitoid consuming practically all the intern body of the host. This parasitoid has many alternative hosts, so its abundance varies in response to relative changes in numbers of codling moth and alternative hosts from year to year (Glen et al., 1982). Notwithstanding none of these larval parasitoids, with the exception of P. turionellae in the southwest of England where it was one of the most important causes of codling moth mortality (Glen et al., 1982), have shown good enough potential as biological control agent (Glen & Curtis, 1978).

New imported parasitoid species from Central Asia, China and Europe have been studied to be introduced in pear orchards in California. Three parasitoid species are being maintained in culture: Liotryphon caudatus, Mastrus ridibundus and

Hyssopus pallidus - The larval ectoparasitoid Hyssopus pallidus functions as a potential antagonist against other larval parasitoids and so has not been field released. The solitary cocoon parasitoid, L. caudatus, has been field released at 39 locations in California since 1992 and the gregarious cocoon parasitoid, M. ridibundus, has been field released in 26 locations since 1995. Microdus rufipes, a solitary larval parasitoid that overwinters in the cocoon of the codling moth, has also been field released in small numbers since 1995. Both of the cocoon parasitoids have been recovered from a number of release sites, and M. ridibundus appear to be most active in pear orchards.

Liotryphon caudatus (Ichneumonidae) is a solitary ectoparasitoid of cocooned prepupal codling. It attacks codling moth prepupae under the bark by paralyzing them and laying an egg externally on the paralyzed host. Parasitoid development is completed in 3-4 weeks depending on temperature. An effective rearing procedure has been developed for the production of L. caudatus on diapausing codling moth cocoons. In 1994, a different Liotryphon species was obtained from Kazakhstan. This species is more difficult to rear on codling moth and it has not been able to obtain a release permit, as the identity of this species can not yet be determined.

Hyssopus pallidus (Eulophidae) is a gregarious ectoparasitoid of late instar codling moth. The small parasitoid adults enter infested fruit to find their hosts and will attack all later larval instars of the codling moth. The host larva is paralyzed and then a series of eggs are laid externally on the host. Parasitoid development is completed in 2-3 weeks depending on temperature. This parasitoid species can readily be reared on larger codling moth larvae (diapausing or non-diapausing stock) and does not require the presence of the host plant to secure host attack in captivity.

Trichomma enecator Rossi (Ichneumonidae) is a solitary larval-pupal endoparasitoid of the codling moth. The adult is normally 12 mm long and it has been found to attack more than fifteen hosts (Charmillot et al., 1997). It appears to be able to attack all larval instars of the codling moth but will only attack larvae inside of fruit. Development of the parasitoid is completed inside the host pupa under the bark. The female parasitoids are attracted to the exudations that accumulate on the surface of attacked fruit and that in the absence of these exudations the attack behavior of the parasitoids is disrupted. However, success in rearing T. enecator on thinning apples infested with codling moth larvae has so far been limited. It seems unlikely that this species can be reared in sufficient numbers to secure field establishment.

Microdus rufipes (Braconidae) is a solitary endoparasitoid that attacks young larvae of codling moth (1st and 2nd instar) and kills prepupae inside their cocoons. It is one of the most important parasitoids in Kazakhstan, where it regularly achieves levels of parasitism between 40-60%. Observations during the season indicate that this species will only attack host arvae in apples and that, like T. enecator, the exudates produced by codling moth larvae on the surface of the apple are essential for parasitoid attack.

Microdus conspicuus (Braconidae) is an endoparasitoid very similar to the last species in both morphology and biology. It is probably not a specialized codling moth parasitoid. It was previously released in the U.S. against the oriental fruit moth but apparently never established. In insectary rearing it does not do as well on codling moth as M. rufipes and we have not been able to maintain a continuous culture.

Pristomerus vulnerator Panz (Ichneumonidae) is a solitary larval-prepupal parasitoid. The adult parasitoids are 9 mm long normally. The adult female laid normally the eggs in young larvae just after they penetrate into the fruit (Charmillot et al., 1997). It seems probable that this species is not well adapted to the attack of codling moth and uses this host only incidentally. However, in 1995 this species was also collected from codling moth in Kazakhstan, a new record for this region suggesting that this species may be increasing its range eastward from Europe. It has been also found in Suisse (Charmillot et al., 1997).

Mastrus ridibundus is a gregarious ectoparasitoid of codling moth cocoons obtained from Kazakhstan for the first time in 1994. Its biology is similar to that of Liotryphon caudatus in that it attacks the prepupal stage of the codling moth. This is a poorly known parasitoid, but local sources of literature from Kazakhstan indicate that it is often responsible for relatively high levels of parasitism. In contrast to L. caudatus, however, M. ridibundus is gregarious producing 4-7 individuals on a single codling moth host. Furthermore, M. ridibundus is more likely to attack cocoons that spin up in the crotch of branches or at the base of the trunk, whereas L. caudatus is adapted to the attack of cocoons concealed beneath the bark of the main trunk.

Other parasitoids found in the center areas of Europe include the hymenopterous Dibrachys cavus Walk., and the dypterous Elodia tragica Meig. They have not been found in America and their introduction to America has not been tried.

D.cavus is a small chalcidid 2 mm long. It is an gregarious ectoparasitoid: normally from five to eight individuals develop together inside the codling moth. It can be either a primary parasitoid attacking the codling moth or a secondary parasitoid attacking other parasitoids like P. turionellae.

E.tragica is a tachinid totally black and shinny 6 mm long. Its biology is not well known, but it is though that it lays the eggs on the fruits and the larvae after eclosion are who search actively for the young codling moth larvae inside the fruit (Coutin, 1960). The larvae develops as a solitaire endoparasitoid inside codling moth larvae and pupae.
 
 

Pathogens:

Pathogens attacking codling moth include bacteria, fungi, viruses, protozoa, and nematodes.

Bacteria:

Experimental and commercial preparations of Bacillus thuringiensis Berliner var. kurstaki have been investigated and used as biological control agents for the codling moth (Roehrich, 1954; Falcon, 1971, Fedorintchik & Korostel, 1972; Vervelle, 1975; Lappa, 1978; De Reede et al., 1985). Due to its feeding behavior the larva hardly contacts deposits of the bacterium on fruit or foliage (Undorf & Huber, 1986), so B. thuringiensis has been combined with chemical pesticides in order to enhance its effectiveness against the codling moth (Galetenko et al., 1976; Videnova & Ismail, 1985). Notwithstanding, several years of testing commercial preparations of B. thuringiensis (like Dipel, Thuricide) in apple and pear orchards in California never resulted in acceptable control (Falcon & Berlowitz, 1986).

A strain of Serratia marcescens Bizio and different strains of Bacillus cereus Frankland and Frankland were considered to be potential biological control agents of the codling moth by Stephens (1952), but the application of this last one in the Niagra Peninsula, Ontario, and other areas in Canada failed to its control (Phillips et al., 1962).

Fungi:

Different entomogenous fungi attack the coding moth; the most infective are Spicaria farinosa (Fries) Vuillemin, Metarrhizium anisopliae (Metchnikoff) Sorokin (Pristavko et al., 1975), Aspergillus flavus Link (Pristavko et al., 1975),Verticillium lecanii Zimmerman, Fusarium oxysporum Schlecht (Pristavko et al., 1975), Beauveria brongniartii (Saccardo) Petch ( Pristavko et al., 1975), Beauveria globulifera (Spegazzini) Picard, and Beauveria bassiana. This last one is one of the most common species found infecting codling moth in nature (Ferreira, 1947; Russ, 1964; Jaques & MacLellan, 1965; Hagley, 1971), and it also attacks other pests like aphids and Lygus spp. Preparations based on B. bassiana (i.e., Boverin) or Paecilomyces farinosus (Dickson ex Fries) (i.e., Pecilomin) mixed with reduced doses of chemical pesticides (one-fifth to one-tenth of the normal concentration) are recommended for codling moth control, both by killing the larvae or by reducing fertility of adults surviving (Falcon & Huber, 1991).

Viruses:

A granulosis virus (CpGV), isolated from dead codling moth larvae in 1963, has been considered the most effective biological control agent ever tested against the codling moth: its virulence for the neonate larvae is more than 10,000 times higher than that of the bacterium B. thuringiensis (Undorf & Huber, 1986). Furthermore, CpGV can be produced efficiently and effectively under laboratory conditions, it can be applied to fruit trees with the same equipment as chemical insecticides, it doesn. t interfere with population levels of other natural enemies, and it reduces codling moth damage and the abundance of surviving larvae (getting a bigger effect when a larger and uniformly area is sprayed).

The first experimental-commercial production of CpGV, labelled SAN 406, has been very successful, specially in America. In Europe, a major difficulty in the field use of CpGV is timing treatments for maximum protection of the fruit from codling moth damage, because pheromone trap data are difficult to interpret, particularly under the fluctuating weather conditions of northern Europe. In order to avoid this problem, weekly sprays of reduced concentrations of CpGV during the entire period of codling moth activity are applied. In fact, nine applications of one-tenth of the CpGV regular dose, applied at weekly intervals, have been shown to be as four treatments applied at full dose (Dickler & Huber, 1986).

The nuclear polyhedrosis virus (NPV) can also attack the codling moth, but its importance as biocontrol agent is reduced.
 

Protozoa:

Although they don. t represent an important factor in the natural control of codling moth populations, some protozoa have been found attacking the codling moth. Nosema carpocapsae Paillot, first described from France, is a common pathogen of codling moth in many areas of the world (Malone & Wigley, 1980). Other species of microsporidia, like Pleistophora carpocapsae Simchuk and Issi, a parasite of larvae and pupae that has been found in Moldavia (Simchuk & Issi, 1975), Pleistophora californica Steinhaus and Hughes, Nosema destructor Steinhaus and Hughes, and Vairimorpha plodiae Kellen and Lindegren (Weiser and Hostounsky, 1971), have been found attacking the codling moth.
 

Nematodes:

The entomogenous nematode Steinernma feltiae Filipjev (=Neoaplectana carpocapsae Weiser) attack immature stages of the codling moth. It has been foun in the state of Virginia, U.S.A. (Dutky & Hough, 1955), Czech Republic (Weiser, 1955), and Mexico (Poinar, 1979). The use of this nematode for suppression of overwintering codling moths appears promising during the winter moths: its application to the trunks and main branches of apple trees can cause from 60% (Dutky, 1959) up to 95% (Kaya et al., 1984) mortality in prepupae.
 
 

2.-LEAFROLLERS

Leafrollers are attacked by a large number of mostly unspecific predators and a big range of parasitoids, comprising specialists and generalists, as by different pathogens, including viruses, bacteria, fungi, and protozoa.

Leafrollers predators include insectivorous birds, small mamals and different groups of arthropods (including almost all orders of predatory insects).

Parasitoids include species attacking to all different life stages:

-Egg parasitoids of leafrollers are confined to the chalcidoid family Trichogrammatidae.

-Egg-larval parasitoids include species of the genus Ascogaster spp.

-Larval parasitoids include braconid, ichneumonid, chalcidid, bethylid, tachinid and sarcophagid species.

-Pupal parasitoids include basically ichneumonid and chalcidid species.

A big number of parasitoid species have been observed attacking the codling moth, but the problem is that it only exists the reference, but not additional information about their biological cycle, range of hosts and potential for codling moth control has been obtained.
 
 

Pathogens attacking leafrollers include viruses (specially granulosis virus and nuclear polyhedrosis virus), bacteria (all leafrollers can be attacked with different degrees of successful control by B. thuringiensis), different genera of fungi, and different protozoa.
 
 

2.1.-Acleris minuta (Robinson): yellowheaded fireworm or lesser apple leaf-folder.

2.1.1.-Principal characteristics of the pest.

The yellowheaded fireworm exhibits more or less the typical leafrolling habit in the larval stage. It occurs over most of the U.S.A. east of the 100th meridian, in the southern regions of the adjoining Canadian provinces, but not, with the exception of Texas, in the more southern states. It has a large range of plant hosts, including apple, plum, canberry, pear, blueberry, willlow, sweet gale, huckleberry and wild rose (Chapman & Lienk, 1971), but currently it is not commonly found on apple or other tree fruits (Schwarz et al., 1983). For more aspects about its biology and ecology, see Weatherby & Hart (1984).
 
 

2.1.2.-Natural enemies.

In addition to the generalist predators, other natural enemies of A. minuta are:

Parasitoids:

The hymenopterous ichneumonid Itoplectis conquisitor Say has been found to be the most important natural enemy of A. minuta in North-America (Townes & Townes, 1962). It attacks A. minuta in its pupal stage.

Pathogens:

The fungus Conidiobolus thromboides Dreschsler (=Entomophthora virulenta Hall and Dunn) is the most important attacking A. minuta.
 

2.2.-Archips argyrospila (Walker): fruittree leafroller.

2.2.1.-Principal characteristics of the pest.

A. argyrospila (Lepidoptera: Tortricidae, Tortricinae) is a native species from North America (Weires & Riedl, 1991), but it is not found in the arid areas of the Southwest or at high elevations (Freeman, 1938). The feeding habits of the fruittree leafroller are not very specific; a large number of host species have been reported, including apple, pear, plum, cherry, apricot, peach, currant, raspberry, rose, black walnut, soft maple, hickory, oak, elm, ash, boxelder, hazelnut, poplar, lilac, oats, wheat, alfalfa, onions, beans, and others. The fruittree leafroller is univoltine throughout its range and overwinters in the stage of egg. Larvae can cause heavy defoliation and lead to important losses of the crop. See Powell (1964) for a more extensive description of fruittree leafroller biology.

The usual control method for the fruittree leafroller has been the application of organophosphate insecticides, but the pest has developed resistance in areas where the use of these compounds has been too abusive (Madsen & Carty, 1997; Vakenti et al., 1984). Now the use of Bacillus thuringiensis Berliner is recommended for control of fruittree leafroller in place of organophosphate insecticides (Vakenti et al., 1984). Identification of the sex pheromone components (Roelofs et al., 1974) has allowed a very specific monitoring method, and the practical value of pheromone traps in a management program was demonstrated by Madsen & Peters (1976).
 
 

2.2.2.-Natural enemies.

Predators:

Different generalist predators attack the fruittree leafroller. The most important are spiders and insects belonging to the order of the Heteroptera and Neuroptera (Herting, 1975).

Spiders:

Spiders from four different families have been shown to be the most efficient predators on young stages of A. argyrospilla in Canada (LeRoux, 1961):

Araniella displicata Hentz (Argiopidae)

Metaphidippus marginatus Walchenaer (Salticidae)

Tetragnatha versicolor Walck. (Tetragnathidae)

Misumena vatia Clerk (Thomisidae)

Misumenops asperatus Clerk (Thomisidae).

Insects:

*Heteroptera:

Other important predators are heteropterous belonging to the family Pentatomidae, concretely to the genus Podisus spp. (Paradis, 1961). The most important predator species are P. maculiventris Say, P. modestus Dallas, and P. placidus (Uhler).

*Neuroptera:

The only neuropterous predators that have been seen successful in regulating fruittree leafroller populations are Chrysopidae, belonging to the genus Chrysopa spp.
 
 

Parasitoids:

Numerous insect parasitoids attack the fruittree leafroller. They are lepidopterous and dipterous pertaining basically, but not exclusively, to the families Ichneumonidae, Braconidae, and Tachinidae. Mayer & Beirne (1974) and Powell (1964) give references containing information on the most important parasitoids in various regions of British Columbia and United States, respectively. The parasitoid species found attacking and controlling more or less successfully the fruittree leafroller are:

*Hymenoptera:

-Ichneumonidae:

Angitia cacoeciae Viereck, in Canada (Paradis, 1961; Paradis & LeRoux, 1962).

Angitia eureka Ashmed, in United States (Newcomer, 1958).

Campoplex nr. atridens Townes, in California (Powell, 1962).

Camposcopus (Gravenhorstia) spp., in Canada (Paradis, 1961; Paradis & LeRoux, 1962).

Exochus nigripalpis Thompson, in Canada (Paradis, 1961) and in California (1962).

Exochus pallipes Cresson, in United States (Schaffner, 1959).

Ischnus inquisitorius Muller, in United States (Townes & Townes, 1962).

Ischnus minor Townes, in Canada (Townes & Townes, 1962).

Itoplectis conquisitor Say, in Canada (Paradis, 1957b) and United States (Townes & Townes, 1962).

Itoplectis quadricingulatus Provander, in North America (Townes & Townes, 1962).

Phytodietus vulgaris Cresson, in Canada (Paradis, 1961; Paradis & LeRoux, 1957b).

Pimpla pedalis Cresson, in North America (Townes & Townes, 1962).

Tranosema pterophorae Ashmead, in United States (Newcomer, 1958).

Triclistus emarginatus Say, in California (Powell, 1962).

-Braconidae:

Apanteles sp., in Canada (Raizenne, 1952; Paradis, 1961).

Macrocentrus iridescens French, in Canada (Raizenne, 1952).

Microgaster canadensis Muesebeck, in Canada (Paradis, 1961; Paradis & LeRoux, 1962).

Microgaster peroneae Walley, in Canada (Paradis, 1961).

-Chalcididae:

Brachymeria ovata Say, in California (Powell, 1962).

Spilochalcis leptis Burks, in United States (Burks, 1940).

-Eulophidae:

Elachertus proteoteratis Howard, in Canada (LeRoux, 1961).

-Trichogrammidae:

Trichogrammatomyia tortricis Girault, in United States (Chapman & Lienk, 1971).

-Vespidae:

Ancistrocerus waldeni Vieric, in California (Richards, 1962).

*Dyptera

-Tachinidae:

Actia interrupta Curran, in California (Powell, 1962).

Compsilura concinnata Meigen, in Canada (Paradis, 1961).

Eumea caesar Aldrich, in Canada (Paradis, 1961) and the U.S.A. (Newcomer, 1958).

Nemorilla pyste Walker, in the U.S.A. (Schaffner, 1959).

Paralispe infernalis Townsend, in the U.S.A. (Newcomer, 1958).

Phryxe pecosensis Tns., in Canada (LeRoux, 1961) and in the U.S.A. (Sellers, 1953).

Phryxe vulgaris Fallen, in the U.S.A. (Schaffner, 1959).

Pseudoperichaeta erecta Coq., in the U.S.A. (Schaffner, 1959).
 
 

Pathogens:

The granulosis virus (GV) has been found to attack naturally the fruittree leafroller.
 
 

2.3.-Archips rosana (Linnaeus): European leafroller.

2.3.1.-Principal characteristics of the pest.

The European leafroller is a general feeder: its principal hosts are apple, pear, hawthron, currant, filbert, and species of Ligustrum (Chapman, & Lienk, 1971). Although it has been reported as a pest in North America, specially in British Columbia, it has not achieved the pest status it has in Europe, its original area (Weires & Riedl, 1991). The seasonal biology and feeding habits of this leafroller are very similar to those of the fruittree leafroller. It is also a univoltine species and overwinters in the egg stage. See Chapman & Lienk (1971) for a more complete description of its biological and ecological characteristics.

Control of A. rosana in commercial orchards is similar to control of the fruittree leafroller (Madsen et al., 1977); organophosphate insecticides applied at the pink bud stage are effective. B. thuringiensis has been used in filbert orchards as part of integrated control programs (AliNiazee, 1974). Its sex pheromone components have been also identified (Roelofs et al., 1976b).
 
 

2.3.2.-Natural enemies.

Predators:

Insects:

The most important predators of this leafroller belong to a diversity of insect families. Most of them are generalist predators as the coccinellids, syrphids, and mirids and anthocoridis, and they may feed on overwintering eggs and young larvae (AlliNiazee, 1983).

Mites:

Mites belonging to the family Trombidiidae can feed in young stages of the European leafroller, although its importance as natural enemies is reduced specially in North America. In France, the species Allothrombium fuliginosum Hermann was found to be relatively successful in controlling European leafroller populations (Guennelon & Tort, 1958).
 
 

Parasitoids:

Most of the significant impact on population reduction is caused by a number of generalist parasitoids. Mayer & Beirne (1974) distinguished 28 different parasitoid species attacking larvae and pupae collected from apple in British Columbia. AliNiazee (1977) also identified in filberts and hazelnuts different major parasitoids:

*Dyptera:

-Tachinidae:

The most important species found in the U.S.A., most of them in hazelnut, are:

Hemisturmia tortricis (Coquerel).

Erynnia tortricis Coquillet.

Eulasiona nigar.

Aplomya caesar (Ald).

Other species found in other areas are:

Actia crassicornis Meigen, in the Czech Republic (Capek, 1961).

Bessa parallela Meigen, in Belgium (Brande & Verbeke, 1949).

Blondelia nigripes Fallen, in Belgium (Brande & Verbeke, 1949) and Russia (Markelova, 1957).

Clemelis pullata Mg., in Moldavia (Bichina & Talitskii, 1955).

Elodia morio Fall., in Belgium (Brande & Verbeke, 1949) and Moldavia (Talitskii, 1961).

Eumea mitis Mg., in Belgium (Brande & Verbeke, 1949) and Poland (Wiackowski, 1957).

Eumea westermanni Zetterstedt, in Belgium (Brande & Verbeke, 1949) and Britain (Emden, 1954).

Eurysthaea scutellaris R.D., in Belgium (Brande & Verbeke, 1949), the Czech Republic (Capek, 1961), and areas of Central Europe (Herting, 1957).

Exorista spp., in Russia (Markelova, 1957).

Nemorilla floralis Fall., in the Czech Republic (Kolubajiv, 1962).

Pseudoperichaeta erecta Coq., in Canada (Putman, 1942).

-Species belonging to other dypterous families have been found parasitizing A. rosana in other areas of the world, like:

Syrphidae: Xanthandrus comtus Harris, in Belgium (Brande & Verbeke, 1949); and,

Sarcophagidae: Agria affinis Fallen, in Moldavia (Talitskii, 1961).

*Hymenoptera:

-Ichneumonidae:

The most important species found in the U.S.A. are:

Scambus trangressus.

Itoplectis quadriccingulata (Prov.).

Ischnus inquisitorius atriceps (Cresson).

Glypta simplicipes Cresson.

Iseropus conquisitor Say

Iseropus quadricingulata Provancher

Phytodietus vulgaris Cresson

Scambus hispae Harris

In other areas of the world other species have been found attacking the European leafroller:

Agrypon flaveolatum Ratzeburg, in Germany (Starke, 1940).

Angitia areolaris Holmgren, in Moldavia (Talitskii, 1961).

Angitia armillata Grav., in Belgium (Brande & Verbeke, 1949) and Moldavia (Talitskii, 1961).

Angitia contracta Brischke, in Moldavia (Talitskii, 1961).

Angitia exareolata Ratzeburg, in Belgium (Brande & Verbeke, 1949) and Moldavia (Talitskii, 1961).

Angitia fenestralis Holmgren, in Moldavia (Talitskii, 1961).

Angitia interrupta Holmgren, in Belgium (Brande & Verbeke, 1949).

Angitia rufipes Grav., in Belgium (Brande & Verbeke, 1949) and Germany (Starke, 1940).

Angitia trochanterata Thomson, in Moldavia (Talitskii, 1961).

Apechthis rufata Gmelin, in the Czech Republic (Capek, 1956), Russia (Markelova, 1957), and Moldavia (Talitskii, 1961).

Campoletis latrator Grav., in U.S.S.R. (Tomilova, 1962).

Campoletis zonata Grav, in Germany (Starke, 1940; Hedwig, 1950).

Campoplex argentator Aubert, in France (Aubert, 1961).

Camposcopus nigricornis Wesmael, in Germany (Hedwig, 1950).

Ephialtes punctulatus Ratz., in Belgium (Brande & Verbeke, 1949).

Exochus nigripalpis Thomson, in Canada (Townes & Townes, 1959).

Glypta bipunctoria Thunberg, in Belgium (Brande & Verbeke, 1949).

Hybophanes scabriculus Grav., in Belgium (Brande & Verbeke, 1949).

Iseropus inquisitor Scopoli, in Belgium (Brande & Verbeke, 1949).

Iseropus maculator Fabr., in Belgium (Brande & Verbeke, 1949), Czech Republic (Capek, 1956), Switzerland (Baggiolini, 1958), other areas of Central Europe (Zwolfer, 1956), and Moldavia (Bichina & Talitskii, 1955).

Iserotus viduata Grav., in Russia (Markelova, 1957).

Lissonota spp., in Switzerland (Baggiolini, 1958).

Lycorina triangulifera Holmgren, in Moldavia (Talitskii, 1961).

Perilissus pallidus Grav., in Germany (Starke, 1940).

Phaeogenes semivulpinus Grav., in Russia (Markelova, 1957).

Phytodietus polyzonias Forster, in Belgium (Brande & Verbeke, 1949), Czech Republic (Capek, 1956), Poland (Wiackowski, 1957), Moldavia (Bichina & Talitskii, 1955), Russia (Markelova, 1957), and Latvia (Tsimdin, 1959).

Pimpla instigator Fabr., in Belgium (Brande & Verbeke, 1949) and Russia (Markelova, 1957).

Pimpla turionellae L., in Poland (Wiackowski, 1957), Switzerland (Baggiolini, 1958), Russia (Markelova, 1957), and Latvia (Tsimdin, 1959).

Prionopoda stictica Fabr., in Germany (Hedwig, 1950).

Scambus brevicornis Grav., in Moldavia (Talitskii, 1961).

Scambus sagax Hartig, in Moldavia (Talitskii, 1961).

Scambus vesicarius Ratzeburg, in Moldavia (Bichina & Talitskii, 1955).

Tranosema arenicola Thomson, in Central Europe (Zwolfer, 1956).

Trichomma enecator Rossi, in Germany (Hedwig, 1955) and Moldavia (Talitskii, 1961).

Triclistus globulipes Desvignes, in Moldavia (Talitskii, 1961).

-Braconidae:

The most important species found in hazelnut orchards in the U.S.A. are:

Bracon politiventris.

Macrocentrus iridescens.

Apanteles polychrosidis.

Meteorus trachynotus.

Microgaster epagoges Gahan.

Ascogaster quadridentata Wesm.

Oncophanes spp. (O. americanus Weed and O. lanceolatus Nees, specially).

Other species have been found attacking European leafroller in other areas:

Agathis dimidiator Nees, in Latvia (Tsimdin, 1959).

Apanteles ater Ratzeburg, in Belgium (Brande & Verbeke, 1949) and Czech Republic (Capek, 1959).

Apanteles dilectus Haliday, in Belgium (Brande & Verbeke, 1949).

Apanteles lacteicolor Vierek, in Moldavia (Talitskii, 1961).

Apanteles xanthostigmus Hal., in Poland (Wiackowski, 1957) and Moldavia (Talitskii, 1961).

Ascogaster rufipes Latreille, in Moldavia (Zhigaltseva, 1959).

Bracon brevicornis Wesmael, in Belgium (Brande & Verbeke, 1949) and Moldavia (Talitskii, 1961).

Eubadizon extensor L., in the Czech Republic (Capek, 1956).

Macrocentrus abdominalis Fabr., in Belgium (Brande & Verbeke, 1949) and the Czech Republic (Capek, 1961).

Macrocentrus thoracicus Nees, in Belgium (Brande & Verbeke, 1949), the Czech Republic (Capek, 1961), and Moldavia (Talitskii, 1961).

Meteorus confinis Ruthe, in Moldavia (Talitskii, 1961).

Meteorus ictericus Nees, in Belgium (Brande & Verbeke, 1949), Germany (Starke, 1940), Switzerland (Baggiolini, 1958), and Moldavia (Talitskii, 1961).

Microgaster abdominalis Nees, in Belgium (Brande & Verbeke, 1949).

Microgaster canadensis Muesebek, in Canada (Putman, 1942)

Microgaster crassicornis Ruthe, in Modavia (Talitskii, 1961).

Microgaster globata L., in Belgium (Brande & Verbeke) and the Czech Republic (1956).

Microgaster meridiana Haliday, in Belgium (Brande & Verbeke) and Moldavia (Talitskii, 1961).

Orthostigma spp., in Switzerland (Baggiolini, 1958).

-Chalcididae:

The most important species found in the U.S.A. is Brachymeria ovata (Say).

Other Brachymeria species have been found in Russia and Moldavia (Brachymeria intermedia Nees), Switzerland (Brachymeria pseudorugosa Masi) (Baggiolini, 1956), and France (Brachymeria rugulosa Forster) (Steffan, 1959).

-Pteromalidae:

The most abundant species found in the U.S.A., in hazelnut orchards, is Habrocytus phycidis Ashmead.

Other ptermoalids found in other areas are:

Cyclogastrella deplanata Nees, in Zwitzerland (Baggiolini, 1958).

Dibrachys cavus Walker, in Belgium (Brande & Verbeke, 1949) and Moldavia (Talitskii, 1961).

Pteromalus puparum L., in Moldavia (Talitskii, 1961).

-Elasmidae:

There is only one species with some importance as biological control agent of A. rosana:

Elasmus albipennis Thomson, found in Moldavia (Talitskii, 1961).

-Eulophidae:

The most important parasitoid species of A.rosana belonging to this family have been found in Europe:

Colpoclypeus florus Walk., in Moldavia (Talitskii, 1961).

Euplectrus cacoeciae Ferriere, in Bulgaria (Ferriere, 1940).

Pediobius cassidae Erdos, a primary and secondary parasitoid found in Moldavia by Talitskii (1961).

Pediobius crassicornis Thomson, in Moldavia (Bichina & Talitskii, 1955).

Pediobius facialis Giraud, another primary and secondary parasitoid found in Moldavaia by Talitskii (1961).

Pediobius pyrgo Walker, in Moldavia (Talitskii, 1961).

-Torymidae:

The only species of this family with some importance as natural enemy of A. rosana is Monodontomerus aereus Walker. It was foun in Switzerland (Baggiolini, 1958) and Moldavia (Talitskii, 1961), but it is not present in North America.

-Trichogrammidae:

The most important parasitoid species of A. rosana belonging to this family have been found in Europe:

Trichogramma cacoeciae Marchal, in France (Guennelon & Tort, 1958), Switzerland (Geier, 1946; Baggiolini, 1956; Ferriere & Geier, 1956), and other areas of Europe (Quednau, 1956).

Trichogramma embryophagum Hartig, in Russia (Markelova, 1957), Latvia (Tsimdin, 1959), and other ex-soviet republics (Volkov, 1959).

Trichogramma evanescens Westwood, in Russia (Markelova, 1957).

The importance of the parasitoids found in the United States increase as the season progresses. Coop (1982) also found Meteorus argyrotaenia Johanson, Meloboris spp., and Glypta spp. attacking A. rosana.
 

2.4.-Archips semiferana (Walker): oak leafroller.

2.4.1.-Principal characteristics of the pest.

Although the oak leafroller has been reported as a incidentally important apple pest, it is a major pest of oak (Rexrode, 1971; Wilson, 1972). In fact, it has not become a serious problem on deciduous tree fruits outside of Ontario (Weires & Riedl, 1991). The oak leafroller is univoltine and resembles the fruittree and European leafrollers in its seasonal biology and feeding habits. See Chapman & Lienk (1971) for a more complete description of its biological and ecological characteristics.
 
 

2.4.2.-Natural enemies.

Predators:

No specific predators of A. semiferana have been found, but all generalist predators can attack it. The potential of this predators as natural enemies of A. semiferana is high, but the success in keeping the pest population densities at low levels depends on the relative abundance of alternative prey.

Parasitoids:

Larvae and pupae of the oak leafroller are attacked by dipterous as well as hymenopterous parasitoids (Mumma et al., 1974); Mumma & Zettle, 1977b), but only two species have shown some potential to control A. semiferana populations in the U.S.A.

*Hymenoptera:

-Ichneumonidae:

Itoplectis conquisitor Say (Sellers, 1943).

*Diptera:

-Tachinidae:

Eumea caesar Aldrich (Sellers, 1953; Schaffner, 1959).
 
 
 

2.5.-Argyrotaenia citrana (Fernald): orange tortrix or apple skinworm.

2.5.1.-Principal characteristics of the pest.

The orange tortrix is one of the most polyphagous tortricids in North America, attacking woody plants (including apples, apricots, citrus), weeds, vegetables and ornamentals. Although as economic problem it is primarily important on fruit, its larva also exhibits the typical leafrolling behavior. For an extensive explanation about biological and ecological aspects of this pest, see Borden (1953) and Madsen & Falcon (1960).

Control in commercial apple orchards is primarily achieved by the use of protective residual insecticides (Weires & Riedl, 1991), but timing of treatments is difficult. The identification of its female sex pheromone by Hill et al. (1975) and studies on the thermal requirements of immatures stages and their physiological thresholds for development (Basinger, 1938; Coop, 1982; Knight, 1986) have helped with the management of this pest.
 
 

2.5.2.-Natural enemies.

Predators:

Like in the case of A. semiferana, only generalist predators have been found attacking A. citrana, so success in keeping the pest population densities at low levels depends on the relative abundance of alternative prey.

Parasitoids:

Many mostly generalist insect parasitoids attack the orange tortrix. Coop (1982) collected 18 species from this tortricid in cultivated Rubus spp., in Oregon, of which five accounted for 90% of the parasitism, which ranged from 4 to 100%. The hymenopterous and dypterous parasitoids attacking A. citrana, by order of importance, belong to the families Braconidae, Ichneumonidae, Tachinidae, Eulophidae, and Trichogrammatidae:

*Hymenoptera:

-Braconidae:

Apanteles aristoteliae Viereck, univoltine solitary larval endoparasitic, attacking the second, third, and fourth larval instars, and small fifth instar individuals. It is able of overwintering inside the host overwintering larvae and to attack it during all year long, but specially during the three first generations. It seems to be the only specific parasitoid of the orange tortrix (Coop, 1982).

Meteorus argyrotaeniae Johanson, univoltine solitary larval endoparasitic that normally attacks fouth, fifth, and sixth instar larvae and emerges from fifth and sixth instars. Like A. aristoteliae it is able to overwinter inside the host larvae, and although can attack them from March to November it its specially important during the first two generations of this host (Coop, 1982).

Oncophanes americanus (Weed), gregarious larval ectoparasitic that produces a nearly complete paralysis of the larval host, thus stopping its development after the adult parasitize nearly mature larvae (fourth and fifth instars). It overwinters as mature larvae inside cocoons and its importance as biological control agent of the orange tortrix is mostly during Summer and Fall (from June to September) (Coop, 1982).

Meteorus dimidiatus (Cresson), solitary larval endoparasitic that predominantly attacks fourth and fifth instar larvae (Coop, 1992).

Meteorus trachynotus Viereck, solitary larval endoparsitic (Coop, 1992).

Another braconid found attacking A. citrana in California (Basinger, 1938; Powell, 1962) was Hormius basalis Provancher.

-Ichneumonidae:

Phytodietus vulgaris Cresson, univoltine solitary larval ectoparasitic that attacks preferently mature larvae although hosts as small as third instars can also be utilized. Like O. americanus, overwintering normally occurs as a mature larva in diapause within a cocoon formed at the site where the host had prepared to pupate. It is more important during the Summer and Fall, specially the second and third generation of the orange tortrix (Coop, 1982).

Enytus eureka (Ashmead), univoltine solitary larval endoparasitic attacking predominantly fourth and fifth larval instars. It can overwinter inside orange tortrix larvae (Coop, 1982).

Diadegma spp. (Ichneumonidae), solitary larval endoparasitic attacking mature larvae (Coop, 1982).

Parania geniculata (Holmgren), solitary larval-pupal endoparastic (Coop, 1982).

Meloboris sp.(Ichneumonidae), solitary larval endoparasitic (Coop, 1982).

Itoplectis quadricingulata (Provancher), solitary pupal endoparasitic (Coop, 1982).

Other ichneumonids attacking A. citrana in North America, most of them generalist parasitoids and with lower potential as biological control agents, are:

Angitia eureka Ashmead (Rosenstiel, 1949; Johansen & Breakey, 1938; Breakey, 1951).

Campoletis spp. (Johansen & Breakey, 1938; Breakey, 1951).

Campoplex (Omorgus) spp. (Basinger, 1938).

Exochus nigripalpis Thomson (Townes & Townes, 1959; Powell, 1962).

Glypta spp. (Powell, 1962).

Ischnus inquisitorius Mueller (Johansen & Breakey, 1938; Breakey, 1951).

-Eulophidae:

Elachertus spp., solitary larval ectoparasitic.

-Trichogrammatidae:

Trichogramma spp., solitary egg endoparasitoid.

*Diptera:

-Tachinidae:

Pseudoperichaeta erecta (Coquillet), solitary larval or larval/pupal endoparasitic.
 
 

Of all larval parasitoids, A. aristoteliae and M. argyrotaeniae seem to be the ones with more potential to control orange tortrix populations. Coop (1982) found that these two species accounted for 63% of all parasitoids found in this pest, being both the most widespread as well as most abundant species. P. vulgaris, Diadegma spp., E. eureka, and O. americanus had a relative importance. Both E. eureka and P. vulgaris were found relatively widespread but with low levels of attack on orange tortrix larvae. P. geniculata and M. dimidiatus were rarely found.

Of the pupal parasitoids, I. quadricingulata, a cosmopolitan and non host-specific parasitoid, was the most abundant. Two others, P. geniculata and P. erecta, parasite both larvae and pupae.
 

2.6.-Argyrotaenia quadrifasciana (Fernald): fourbanded or fourlined leafroller, or lesser all-green leafroller.

2.6.1.-Principal characteristics of the pest.

The fourlined leafroller is a native species which most important host is apple trees, although its economic impact this host has been limited. Other known hosts are hawthorn (Chapman & Lienk, 1971), shadbush and cherry (Freeman, 1938). In North America it can be found in commercial orchards from Nova Scotia west to central Manitoba and as far south as Arkansas (Chapman & Lienk, 1971). It also has been frequently encountered in abandoned orchards in West Virginia (Brown et al., 1988) and Michigan (Strickler & Whalon, 1985).

From information from New York state, the fourlined leafroller is univoltine and overwinters as a third instar larva. Its feeding habits concentrate on terminal shoot growth, while feeding on young fruit has not been observed (Weires & Riedl, 1991).
 
 

2.6.2.-Natural enemies.

Neither specific predators nor parasitoids have been found attacking the fourlined leafroller, so its range of natural enemies keeps restrict to generalist predators and parasitoids, which are more or less successful in controlling pest populations levels depending on the relative abundance of alternative prey or hosts.
 
 

2.7.-Argyrotaenia velutinana (Walker): redbanded leafroller.

2.7.1.-Principal characteristics of the pest.

The redbanded leafroller has a very wide list of hosts (McCabe & Sheviak, 1980). Undoubtedly apple has now to be considered one of the most important hosts. It doesn. t exhibit a preference for specific apple cultivars (Goonewardene et al., 1979). As a pest of decidious fruits it has been important specially in the easthern part of North America.

The redbanded leafroller overwinters in the pupal stage. The number of generations goes from two in western New York to four in southern areas. For more information about its biological and ecological characteristics see Weires & Riedl (1991).

The control measures of this leafroller were based in pesticide applications (DDT-related and TDE compounds first and organophosphates later) until it became resistant to those compounds. Lately, some attempts to achieve commercial control through mass trapping of the males with pheromone traps have been done in apples (Trammel et al., 1974) and grapes (Taschenberg et al., 1974), but the use of the pheromone for mating disruption may have more potential (Reissig et al., 1978; Taschenberg & Roelofs, 1978).
 
 

2.7.2.-Natural enemies.

Predators:

Like in the case of the fruittree leafroller, the most important predators of A. velutinana are spiders and heteropterous and neuropterous insects.

Spiders:

The most effective spider predators of A. velutinana in North America are the same generalist predators that attack A. argyrospila (LeRoux, 1960), this is to say:

Araniella displicata Hentz (Argiopidae).

Metaphidippus marginatus Walchenaer (Salticidae).

Tetragnatha versicolor Walck. (Tetragnathidae).

Misumena vatia Clerk (Thomisidae).

Misumenops asperatus Clerk (Thomisidae).

Insects:

*Hemyptera:

-Pentatomidae:

The most important insects regulating A. velutinana populations are two pentatomids belonging to the same genus (Paradis, 1957a):

Podisus maculiventris Say.

Podisus modestus Dallas.

*Neuroptera:

-Chrysopidae:

These chrysopid predators are very generalist and they attack all different leafrollers. The ones that have been found to control more effectively A. velutinana populations are:

Chrysopa spp., in the U.S.A. (Clancy & Pollard, 1948).

Chrysopa oculata Say, in Canada (Paradis, 1957a)
 
 

Parasitoids:

*Hymenoptera:

Different generalist parasitoids that attack other leafrollers can also regulate populations of A. velutinana with different levels of success. The most important are hymenopterous belonging to different families.

-Ichneumonidae:

Angitia obliterata Cresson, found in Canada (Paradis, 1957a).

Iseropus spp., also found in Canada (Paradis, 1957a).

Itoplectis conquisitor Say, found in Canada (Paradis, 1957a) and in the U.S.A. (Summerland & Hamilton, 1954).

Phytodietus vulgaris Cresson, found in the U.S.A. (Clancy & Pollard, 1948).

Pimpla aequalis Provancher, found in the U.S.A. (Summerland & Hamilton, 1954).

Triclistus emarginatus Say, found in the U.S.A. (Clancy & Pollard, 1948; Townes & Townes, 1959).

-Braconidae:

Agathis laticincta Cresson, found in Canada (LeRoux, 1961).

Microgaster epagoges Gahan, found in the U.S.A. (Clancy & Pollard, 1948).

-Trichogrammidae (Paradis, 1957a):

Trichogramma minutum Riley, found in Canada.

Trichogramma pretiosum Ril., found in the U.S.A.

-Bethylidae:

Goniozus platynotae Ashmead, found in the U.S.A. (Clancy & Pollard, 1948).
 
 

Pathogens:

The granulosis virus can attack the redbanded leafroller.
 
 

2.8.-Choristoneura rosaceana (Harris): obliquebanded leafroller.

2.8.1.-Principal characteristics of the pest.

The obliquebanded leafroller is indigenous to North America and feeds on the foliage and occasionally fruit hosts, being considered the family Rosaceae its primary hosts (Chapman & Lienk, 1971). Economic problems have been detected on rose (Sanderson & Jackson, 1909), dewberry (Knowlton & Allen, 1937), red rapsberry (Schuh & Mote, 1948), filberts (AliNiazee, 1986), and apple (Chapman & Lienk, 1971). Increasing problems have been detected in apple (Reissig, 1978; Madsen & Madsen, 1980) and filbert orchards (AliNiazee, 1986). It is distributed throughout North America, except for the arid areas on the Southwest (Freeman, 1938; Powell, 1964; Prentice, 1955). It usually has two generations (Chapman & Lienk, 1971), except on its northern limit of distribution or at higher elevations, where it is univoltine. It overwinters as a third instar diapausing larva. For a more complete description of its biological and ecological characteristics, see Knowlton & Allen (1937), Reissig (1978), Onstad & Reissig (1986). The economic damage of the obliquebanded leafroller is primarily on the fruit due to larvae feeding habits. Leaf damage is normally minor, except to growing tips on young trees, and even there is not, like in peaches.

Normal cover spray programs used to keep populations of these the obliquebanded leafroller at low levels, but the development of resistance to organophosphate insecticides has lead to the need of developing more specific control strategies. The identification of the sex pheromone components (Roelofs & Hill, 1979) along with more information on temperature effect on the development of the different stages, more tools became available to time and target control measures against specific life stages (Reisig, 1978; Onstad et al., 1985; AliNiazee, 1986).
 
 

2.8.2.-Natural enemies.

Predators:

The obliquebanded leafroller pupate in the litter or soil. That makes it specially accessible to predation by small mammals and arthropods such as carabids and staphylinids, that can lead to daily predation rates of 66-91%, which would eliminate almost all individuals pupating on the floor (Kelly & Regniere, 1985).

Parasitoids:

Different tachinid, ichneumonid and braconid parasitoid species attack the obliquebanded leafroller (Powell, 1964), some of them 71 and 33% have been reported on caneberries (Schuh & Mote, 1948) and maple (Simmons, 1973), respectively.

Different larval parasitoids, including Diadegma spp. (that attacks second, third, fourth and fith instars), M. argyrotaeniae (attacking the second, third, and fourth instars), O. americanus (that attacks second, third, fourth, fifth and sixth instars), E. eureka (that attacks second, third, and fourth instars), Glypta spp. (attacking third and fourth instars), Macrocentrus irridescens French (that attacks third, fourth, and fifth instars), Apanteles spp. (that attacks the second larval instar), P. erecta (attacking the fourth, fifth, and sixth instars), Elachertus spp. (attacking the second instar), Microgaster epigoges Gahan (that attacks the fourth instar), Charman extensor (Linnaeus) (attacking the fourth instar), M. dimidiatus (attacking the fifth instar), and Bracon spp. (attacking the sixth instar) have been reported (Coop, 1982).

Four of the six most important parasitoids of the orange tortrix are also of the greatest abundance on the obliquebanded leafroller, accounting for 61% of total parasitism (Coop, 1982). These were Diadegma spp., M. argyrotaeniae, O. americanus, and E. eureka. Notwithstanding it seems clear that the obliquebanded leafroller is an unsuitable host for two other common orange tortrix parasitoids, A. aristoteniae and P. vulgaris.

Some pupal parasitoids, including Itoplectis viduata (Gravenhorst) and Brachymeria ovata (Say), were found to attack the obliquebanded leafroller, but their potential as biological control agents is reduced.

Pathogens:

It has been observed that the nuclear polyhedrosis virus can attack C. rosaceana (Martignoni & Iwai, 1986).
 

2.9.-Pandemis limitata (Robinson): threelined leafroller.

2.9.1.-Principal characteristics of the pest.

The threelined leafroller attacks a large range of hosts, including willow, birch, maple, elm and apple. It can be found anywhere in North America except the Southeast area (Weires & Riedl, 1991). The seasonal biology and feeding habits of P.limitata are very similar to those of the obliquebanded leafroller. Throughout most of its distribution range it is bivoltine, but at higher elevations and to northern limits of distribution it has only one generation (Madsen et al., 1984). Its importance in apple orchards had become higher in the last years and it is now considered a major pest of apple in British Columbia (Madsen & Madsen, 1980). Chemical control and timing of control measures are similar to those for the obliquebanded leafroller (Weires & Riedl, 1991).
 
 

2.9.2.-Natural enemies.

Predators:

Like in the case of some other leafrollers, there are not references about specific predators attacking P. limitata, and only the generalist one can have more or less success in regulating its population levels depending on the relative abundance both of predator and prey and alternative preys.

Parasitoids:

The most important natural enemies attacking P. limitata in North America are insect parasitoids.

*Hymenoptera:

-Ichneumonidae:

Scambus alborictus Cresson (Townes & Townes, 1962).

*Diptera:

-Tachinidae:

Blepharomya spinosa Coquillet, specially abundant in Canada (Raizenne, 1952).
 

2.10.-Platynota idaeusalis (Walker): tufted apple budmoth.

2.10.1.-Principal characteristics of the pest.

The variegated leafroller is widely found in the northern USA and in southern Canada (MacKay, 1962). However, it is only important as a fruit pest in the eastern and mid-Atlantic regions (Bode, 1975; David, 1985; Shaffer & Rock, 1983a).

It is a polyphagous feeder. The more important hosts of this leafroller are black haw, blackberry, goldenrod, different species of Solanum, clover, several species of Vaccinium, apple (Chapman & Lienk, 1971; Bode, 1975), and some sweet cherry cultivars (Hogmire & Howitt, 1979).

Generally P. idaeusalis has two generations, and its life cycle and feeding habits are basically the same as P. flavedana. A more complete information about its biology and ecological characteristics can be found in Berkett et al. (1976), Shaffer & Rock (1983b), and Rock & Shaffer (1983).
 
 

2.10.2.-Natural enemies.

Predators and parasitoids:

Neither specific predator nor parasitoid species have been found attacking P. idaeusalis, so the control of its population levels is restricted to generalist natural enemies, which effectiveness depend on the relative density of predator and prey populations and relative abundance of alternative prey.
 
 

Pathogens:

Both the nuclear polyhedrosis and the cytoplasmic polyhedrosis virus can attack P. idaeusalis (Martignoni & Iwa, 1986).
 

2.11.-Platynota flavedana Clemens.

2.11.1.-Principal characteristics of the pest.

Like most of leafroller species P. flavedana is very generalist in its feeding habits. The most important plant host are apples, but they have been found also feeding in strawberry, roses (Hamilton, 1940), peach (Summerland & Hamilton, 1954), cotton (Harding, 1976), azalea, Helianthus species, maple (Chapman & Lienk, 1971), and, in laboratory condition, fava bean (Weires, 1985).

It is more commonly found in southern areas of North America. It has two to three generations, depending on the region. It overwinters as a larva. More extensively discussed aspects about its biological cycle and ecology can be found in Bobb (1972) and Thomas & Hill (1975).
 
 

2.11.2.-Natural enemies.

Predators:

Like in the case of P. idaeusalis, only generalist predators attack P. flavedana.

Parasitoids:

*Diptera:

-Tachinid:

The most important natural enemy of P. flavedana in the U.S.A. is the tachinid Erynnia tortricis Coquillet (Summerland & Hamilton, 1955).
 

2.12.-Sparganothis sulfureana (Clemens): sparganothis fruitworm.

2.12.1.-Principal characteristics of the pest.

S. sulfureana is mostly found in the eastern part of North America (Chapman & Lienk, 1971). Although its importance as a pest is only in cranberries orchards, it can also feed on apple, alfalfa, hawthorn, blueberry and certain Compositae (Chapman & Lienk, 1971).

The life cycle and feeding habits of this leafroller on apple are like the ones of P. flavedana.

The traditional method of control of this pest has been being the use of organophosphate insecticides, but the appearance of resitance lead to look for other alternatives (Weires, 1985).

2.12.2.-Natural enemies.

Predators:

There are not references of specific predators attacking S. sulfureana. Notwithstanding, the role of generalist predators in regulating its population levels can be important, depending on the relative abundance of other potential prey.

Parasitoids:

Different insect parasitoids belonging to the Hymenoptera and Diptera orders, most of them generalists, have been found attacking S. sulfureana in North America.

*Hymenoptera:

-Ichneumonidae:

Angitia compressa Cresson, in the U.S.A. (Beckwith, 1938).

Exochus spp., in Canada (Raizenne, 1952).

Scambus tecumseh Viereck (Townes & Townes, 1962).

-Braconidae:

Bracon gelechiae Ashmead, in the U.S.A. (Beckwith, 1938).

Bracon mellitor Say, in the U.S.A. (Beckwith, 1938).

Meteorus trachynotus Viereck, in the U.S.A. (Beckwith, 1938).

Oncophanes americanus Weed, in the U.S.A. (Beckwith, 1938).

-Eulophidae:

Sympiesis ancylae Girault, in the U.S.A. (Beckwith, 1938).
 
 

*Diptera:

-Tachinidae:

Erynnia tortricis Coquillet, in the U.S.A. (Beckwith, 1938).

Nemorilla pyste Walker, in the U.S.A. (Beckwith, 1938).
 
 
 
 
 
 
 
 

2.13.-Adoxophyes orana (Fisher von Röslerstamm): summer fruit tortrix.

2.13.1-Principal characteristics of the pest.

This species is native from the Eurasian area. Its range extends over the palaearctic region. It is polyphagous, feeding on all types of pome and stone fruits, and on numerous deciduous trees in hedgerows and woods. It is the most important member of a complex of several leafrolling species, and it is the main pest in several European apple-growing regions (center and eastern Europe).

It is a bivoltine species, but it can have a third generation under favorable climatic conditions. For a complete description of its biological cycle and feeding habits see Dickler (1991).

In commercial orchards, its control is normally carried out with broad-spectrum organophosphorous insecticides or synthetic pyretroids. Selective procedures such as IGRs and baculoviruses are still in the experimental phase.
 
 
 
 

2.13.2.-Natural enemies.

Predators:

There are not references of specific natural enemies of A. orana, but generalist leafroller predators (insectivorous birds, small mamals and different arthropod groups, including all orders of predatory insects) can have, depending on relative predator/prey density and abundance of alternative prey, an important impact in controlling A. orana populations levels.
 
 
 
 
 
 
 
 

2.14.-Pandemis heparana (Denis and Schiffermüller).

2.14.1-Principal characteristics of the pest.

P. heparana is also an Eurasian native species. It is widely distributed in all Europe, specially in the Mediterranean area. In the American continent it was observed for the first time in 1978 (Mutuura, 1980). It is a polyphagous species, attacking apple, pear and plum fruits, but it is also considered an important member of the oak pest complex (Bogenschütz, 1978).

Like A. orana, it is a bivoltine species. For more information about its biological cycle and feeding habits, see Dickler (1991).

The usual control practice is the application of broad-spectrum insecticides (Injac and Dulic, 1983). In IPM programs, diflubenzuron is recommended. IGR. s are effective, but only available in a few countries.
 
 

2.14.2.-Naturals enemies.

Parasitoids:

Different insect hymenopterous and dipterous parasitoids have been shown to attack P. heparana in different regions of Europe.

*Hymenoptera:

-Ichneumonidae:

Angitia exareolata Ratzeburg, in Germany (Janssen, 1958).

Apechthis rufata Gmelin, in Latvia (Tsimdin, 1959).

Campoplex multicinctus Grav., in Germany (Janssen, 1958).

Glypta bipunctoria Thunberg, in Germany (Janssen, 1958).

Ischnus inquisitorius Muller, in Poland (Glowacki, 1953).

Itoplectis alternans Gravenhorst, in Germany (Janssen, 1958).

Scambus buolianae Hartig, in Germany (Janssen, 1958).

Teleutaea striata Grav., in Germany (Janssen, 1958).

Tranosema arenicola Thomson, in Germany (Janssen, 1958).
 
 

-Braconidae:

Apanteles ater Ratzeburg, in the Czech Republic (Capek, 1961) and Germany (Janssen, 1958).

Ascogaster spp., in Latvia (Tsimdin, 1959).

Ascogaster rufidens Wesmael, in the Czech Republic (Capek, 1961).

Macrocentrus nitidus Wesmael, in Latvia (Tsimdin, 1959).

Macrocentrus pallipes Nees, in the Czech Republic (Capek, 1961).

Meteorus spp., in Latvia (Tsimdin, 1959).

Meteorus ictericus Nees, in Germany (Janssen, 1958).

-Eulophidae:

Colpoclypeus florus Walker, in Germany (Janssen, 1958).

-Trichogrammidae:

Trichogramma embyophagum Htg., in Latvia (Tsimdin, 1959) and different ex-soviet republics (Volkov, 1959).
 
 

*Diptera:

-Tachinidae:

Clemelis pullata Meigen, in the Czech Republic (Cepelak, 1952).

Nemorilla floralis Fallen, in Austria (Herting, 1957).

Pseudoperichaeta insidiosa R.D., in the Czech Republic (Capek, 1961).

Zenillia libatrix Panzer, in Germany (Herting, 1957).
 
 

Pathogens:

The nuclear polyhedrosis virus attacks P. heparana (Martignoni & Iwai, 1986; Amargier et al., 1981).

Currently available strains of B. thuringiensis are too weak to be effective (Grus, 1982; Undorf and Huber, 1986).
 
 
 
 
 
 
 
 

2.15.-Archips podana (Scopoli): fruit tree tortrix.

2.15.1.-Principal characteristics of the pest.

This native Eurasian species is widely distributed, and, although polyphagous, it is considered a major pest of cultivated apple specially in Italy, England and the Netherlands (De Reede et al., 1985; Audemard, 1986).

It has only one single generation in northern and central Europe, but sometimes also a second one when climatic conditions are favorable. In southern areas it is bivoltine (Audemard, 1986). For a more complete description of its biological cycle and feeding habits, see Dickler (1991).

The normal practice for its control is the use of broad-spectrum organophosphates and synthetic pyrethroids. Diflubenzuron is successfully used in IPM programs (Gruys, 1982).
 
 

2.15.2.-Natural enemies.

Parasitoids:

Different hymenopterous and dipterous insect mostly generalist parasitoids have been shown to attack A. podana in the different regions of Europe where it is mostly distributed.

*Hymenoptera:

-Ichneumonidae:

Campoplex mutabilis Holmgren, in Britain (Hammond & Smith, 1960).

Glypta cicatricosa Ratzeburg, in Germany (Hedwig, 1958).

Itoplectis maculator Fabr., in Britain (Hammond & Smith, 1960).

Lissonota mutanda Schmiedeknecht,in Switzerland (C.I.L.F. List No.1).

Tranosema arenicola Thomson, in Germany (Hedwig, 1958).

-Braconidae:

Apanteles ater Ratzeburg, in Britain and Germany (Wilkinson, 1945).

Apanteles xanthostigmus Haliday, in Poland (Wiackowski, 1957).

Macrocentrus abdominalis Fabr., in the Czech Republic (Capek, 1961).

-Eulophidae:

Colpoclypeus florus Walker, in Germany (Janssen, 1958).
 
 

*Diptera:

-Tachinidae:

Actia crassicornis Meigen, in Britain (Audcent, 1942).

Bessa selecta Mg.,in different ex-sovietic republics (Kudel, 1959).

Eumea mitis Mg., in different ex-sovietic republics (Kudel, 1959).

Nemorilla floralis Fallen, in Britain (Audcent, 1942).

Platymya fimbriata Mg., in different ex-sovietic republics (Kudel, 1959).

Pseudoperichaeta insidiosa R.D., in Britain (Emden, 1954; Hammond & Smith, 1955).
 
 
 
 
 
 
 
 

2.16.-Ctenpseustis obliquana sensu (Green and Dugdale): brownheaded leafroller.
 
 

2.16.1.-Principal characteristics of the pest.

This polyphagous species is found in New Zealand and Australia, attacking principally apple trees (Thomas, 1965; Green, 1984,; Tomkins, 1984), but also in kiwifruits (Steven, 1987).

It has four generations four year and it overwinters as nondiapausing larvae. A complete description of its morphology, biological cycle and feeding habits can be found in Wearing et al. (1991).

As a control practices, the use of organophosphate (such as azinphosmethyl and, more recently, chlorpyrifos targeting eggs and young larvae) and carbamate (e.g. carbaryl, methomyl) chemicals is the most common. Synthetic pyretroids, although providing effective control of this leafroller in trials, are not registered for use in pome and stone fruits (Wearing et al., 1991). Different cultural techniques to reduce leafroller populations and improve the efficacy of insecticidal control can also be used. Since throughout the year reservoirs of leafrollers occur in the ground cover of orchards, a combination of regular mowing, strip spraying with herbicides, and winter grazing by sheep greatly reduces leafroller populations. Furthermore, removal of mummified fruits during winter pruning also reduces overwintering leafroller population (Thomas, 1975).
 
 

2.16.2.-Natural enemies.

While chemicals are the main defense against leafrollers in pome and stone fruits, biological control is also important. Later larval instars of the brownheaded leafroller are attacked by many of endemic predators and parasitoids from Australia and New Zealand.

Parasitoids:

The most important parasitoids species of the brownheaded leafroller are hymenopterous and dipterous belonging to the Braconidae, Bethylidae, Trichogrammatidae, and Tachinidae families. All of them are not very specific, and they attack most of the others leafrollers as other tortricid pests in Australia and New Zealand, like the lightbrown apple moth (Epiphyas postvittana (Walker)). Apanteles spp and Goniozus sp are the parasitoids that have a better control of the brownheaded leafroller in pome and stone fruit orchards, but not the only ones.

*Hymenoptera:

-Braconidae:

Different species of the genus Apanteles spp. attack the brownheaded leafroller: Apanteles tasmanicus (Cameron) and Apanteles sicarius Marshall are the most frequent and effective (Dumbleton, 1935; Thomas, 1965). Both species attack the brownheaded leafroller in the first and second larval instars and kill it before it molts to pupa.

Cotesia (=Glyptapanteles) demeter (Wilkinson), species that belongs to the genus Adelius sp. (Braconidae), attacks larvae and kills the host before it molts to pupa (Green, 1984; Valentine, 1967; Thomas, 1965).

-Bethylidae:

Different species of the genus Goniozus sp. attack the third and fourth larval instar of the brownheaded leafroller. The death of the host occurs during the same instar it is attacked (Dumbleton, 1932; Green, 1984).

-Trichogrammatidae:

Different species of the genus Trichogramma spp. (basically Trichogramma (Trichogrammanza) funiculatum Carver (Valentine, 1967; Green, 1984) and Trichogrammatoidea bactrae fumata Nagaraja (Valentine, 1975; Thomas, 1989), have been shown to control very frequently the brownheaded leafroller. They attack and kill the host in the stage of egg (Valentine, 1967).
 
 

*Diptera:

-Tachinidae:

Pales funesta (Hutton) attacks later larval instars of the brownheaded leafroller and provokes the death of its host in the pupal stage (Thomas 1965, Green, 1984), but it is not as frequent as Apanteles spp and Goniozus sp.
 
 
 
 

Note: The lightbrown apple moth (Epyphias postvittana (Walker)), a tortricid with leafroller behavior sometimes, and the greenheaded leafroller complex (Planotortrix excessana sensu Dugdale) are occasionally important pests. Their natural enemies complex is basically the same than the brownheaded leafroller.
 
 
 
 

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