progress in plum curculio management: a review
TRANSCRIPT
Progress in plum curculio management: a review
Charles Vincenta,*, GeÂrald Chouinardb, Stuart. B. Hillc
aHorticultural research and development centre, Agriculture and Agri-food Canada, 430 Gouin BLVD.
Saint-Jean-sur-Richelieu, QC, Canada J3B 3E6bCentre recherche en productions veÂgeÂtales, MinisteÁre de l'Agriculture, des PeÃcheries et de l'Alimentation du QueÂbec,
3300 Sicotte, C.P. 480, Saint-Hyacinthe, QC, Canada J2S 7B8cFaculty of Social Ecology UWS ± Hawkesbury, Richmond NSW 2753, Australia
Abstract
The plum curculio, Conotrachelus nenuphar Herbst (Coleoptera: Curculionidae), is a key pest of apple orchards of North
America. In absence of chemical treatments, up to 85% damage is caused to the fruit at harvest. In spite of its importance,
plum curculio is one of the least known pest of apple orchards. This paper reviews research advances that were made since
1992 to implement sound IPM programmes against this pest. These advances include behavioral studies, trapping methods,
attractants, entomogenous nematodes and treatments of peripheral zones, with special reference to apple orchards. Plum
curculio management achieved through peripheral-zone treatments have been successful in commercial situations. # 1999
Elsevier Science B.V. All rights reserved.
Keywords: Plum curculio; Conotrachelus nenuphar; Curculionidae; Apple orchard; IPM; Peripheral-zone treatments; Behavioral studies;
Monitoring
1. Introduction
Commercial apple orchards typically harbour many
insect species (Croft and Hull, 1983; Vincent and
Bostanian, 1988; Prokopy and Croft, 1995). In Eastern
North America, three species limit the implementation
of effective biological control programs: tarnished
plant bug, (Lygus lineolaris P. de B.: Miridae), plum
curculio (Conotrachelus nenuphar Herbst: Curculio-
nidae) and apple maggot (Rhagoletis pomonella
Walsh: Tephritidae). All three share the following
seven characteristics:
1. they are indigenous to North America;
2. their present distribution is restricted to North
America;
3. their natural enemies provide inefficient controls in
both untreated and treated orchards;
4. they are direct pests;
5. their life cycle is such that adults are the most
convenient target for effective control;
6. they are relatively mobile; and
7. in Quebec orchards, they complete only one gen-
eration per year on apple trees (Vincent and Roy,
1992).
So far, no resistance to synthetic insecticides has been
reported for these pests.
Agriculture, Ecosystems and Environment 73 (1999) 167±175
*Corresponding author. Tel.: +450-346-4494 ext. 202; fax: +450-
346-7740; e-mail: [email protected]
0167-8809/99/$ ± see front matter # 1999 Elsevier Science B.V. All rights reserved.
PII: S 0 1 6 7 - 8 8 0 9 ( 9 9 ) 0 0 0 2 5 - 0
As no biocontrol agents have yet been identi®ed that
can effectively control these pests in commercial
orchards (van Driesche et al., 1987), broad-spectrum
insecticides remain as the primary controls. Conse-
quently, the implementation of second-stage inte-
grated pest management (IPM) programs (Prokopy
et al., 1990) is a priority in order to reduce the
dependence of broad-spectrum insecticides. With
the apple maggot, for example, signi®cant reductions
in pesticide use have been achieved by placing sticky-
coated, red spheres on trees along the perimeter of
orchards to intercept immigrating ¯ies (Prokopy and
Croft, 1995).
The ecology and management of the plum curculio
has been reviewed most recently by Racette et al.
(1992). The present paper reviews subsequent
research on plum curculio management, based pri-
marily on the author's work on the univoltine (i.e.,
Northern) strain in southwestern Quebec. Most of the
work has focused on the behaviour and monitoring of
adults in commercial orchards, and on visual and
olfactory cues that would allow the design of reliable
monitoring tools to time more accurately, and there-
fore reduce dependence on, insecticide treatments.
2. Pest status and control of plum curculio
The plum curculio commonly causes severe
damage to fruit in North American apple orchards
(Croft and Hull, 1983). In Quebec, up to 85% of
harvested apples may be damaged by this pest in
unsprayed orchards (Vincent and Bostanian, 1988;
Vincent and Roy, 1992). In commercial orchards,
plum curculio populations return to levels of economic
importance within one to three years after cessation of
pesticide spraying (Glass and Lienk, 1971; Hall, 1974;
Hagley et al., 1977). Plum curculio causes two types of
damage. In spring, females oviposit in young fruit,
marking them with their half-moon shaped scars; and
in summer, adults puncture the fruit with their round
(2±3 mm diameter) feeding scars. The appearance of
plum curculio damage is highly variable and, among
all fruit damage rated by IPM specialists, damage of
plum curculio had the lowest average agreement level,
i.e. 71.8% (Vincent and Hanley, 1997).
Quaintance and Jenne (1912) is a valuable source of
information on the plum curculio's synonymy, distri-
bution, developmental stages, life history, habits, host
plants, natural enemies, and control measures. Other
key papers include Chapman (1938) and Smith and
Flessel (1968). Holloway (1977) provides a list of
papers published between 1911 and 1972, and Le
Blanc (1982) reviewed the literature up to 1982. A
review on the ecology and management of the plum
curculio has been published by Racette et al. (1992).
The present review examines advances on plum cur-
culio management, with special reference to the uni-
voltine, i.e. Northern strain. Since the review of
Racette et al. (1992), most of the work has focused
on the behaviour and monitoring of adults in orchards,
along with research on visual and olfactory cues that
would allow the design of reliable monitoring tools
that would improve the timing of insecticide treat-
ments.
3. Management of the plum curculio
The small number of predator and parasite species
of the plum curculio are unable to provide an effective
alternative to chemical insecticides in commercial
orchards (van Driesche et al., 1987; Racette et al.,
1992). Consequently, insecticide treatments are
usually recommended against the adults at petal fall
and once or twice thereafter at 10±14 day intervals
(Chouinard et al., 1998). No resistance to synthetic
insecticides has yet been recorded for this pest.
4. Behavioural studies
Observations on the behaviour of adults, both in the
®eld and in cages, has led to the design of better IPM
programs for plum curculio. Using Zn65 as a marker, it
has been shown that, in autumn, most adults move
from orchards to surrounding woodlots (La¯eur et al.,
1987). After overwintering, preferably under maple
leaves (La¯eur et al., 1987), the returning plum cur-
culios gradually invade adjacent apple orchards
between pink and petal fall (La¯eur and Hill,
1987), after spending several days on the ground under
the perimeter rows of trees, which most of them then
climb (Racette et al., 1990; Chouinard et al., 1993;
Chouinard et al., 1994). From full bloom to nine days
after fruit set, plum curculio adults were found to be
168 C. Vincent et al. / Agriculture, Ecosystems and Environment 73 (1999) 167±175
active mainly during the night (Racette et al., 1991). In
®eld cages, adults labelled with Zn65 also showed a
similar diel periodicity while foraging on dwarf apple
trees (Chouinard et al., 1992a). Because adults are
most active in the trees during the night, Chouinard et
al. (1992a) have recommended that insecticide treat-
ments are likely to be most effective if applied during
the ®rst part of the night.
There have been several attempts to relate adult
activity to ambient temperature to optimize the timing
of insecticide treatments (reviewed by Racette et al.,
1992). Two approaches have been investigated: the
development of a trap to evaluate adult populations
and relate population levels to risks (Prokopy and
Wright, 1998), and the development of day-degree
models to predict the appearance of damage in the
orchards (Reissig and Nyrop, 1995). So far, both
methods have not been used as the sole method to
manage plum curculio populations.
5. Trapping methods
The best available monitoring method for plum
curculio in commercial orchards still remains careful
examination of thousands of small fruit to detect fresh
egg-laying scars (Hoyt et al., 1983; Le Blanc et al.,
1984). In Quebec, Vincent et al. (1997) successfully
used a threshold of 1% damaged fruit, based on careful
monitoring of fruit three times a week, for managing
localized peripheral treatments following full-block
treatment at petal fall. A lower threshold and daily
monitoring would be required, however, in areas
where the pest pressure is very high. Limb tapping
as a monitoring technique is not popular with growers
because beating sticks damage the trees, and accuracy
varies with cultivar, tree shape, time of the day and
scout experience (Racette et al., 1990).
Several traps designed for monitoring of other
curculionids (Mitchell and Hardee, 1974; Oehlschla-
ger et al., 1993; Tedders and Wood, 1994; Hardee
et al., 1996) have been tested for monitoring plum
curculio (Table 1). The pyramid trap (Tedders and
Wood, 1994), which has the silhouette of a small tree,
is based on the tendency of plum curculio to move in
spring towards tree silhouettes (La¯eur and Hill,
1987).
In Massachusetts, Prokopy and Wright (1997a, b)
found that pyramid traps, without attractants or baits,
were most effective when positioned near tree trunks,
and when temperatures were below 188C. This latter
restriction limits their value in this State as most
damage is done at temperatures above 188C. These
authors suggest that the addition of an attractant to this
trap should be tested as the next step in its develop-
ment.
5.1. Attractants
Early laboratory work with a McIndoo olfactometer
found that adult plum curculios are attracted to sali-
cycl-aldehyde in the early part of the season, and that
gallic acid was the best attractant tested in the late part
of the season (Snapp and Swingle, 1929a). Further
work showed that extracts of peach distillates exerted
different effects at different concentrations (Snapp and
Swingle, 1929b).
Over the past 20 years, attractants have been tested
for several curculionid species (Hardee et al., 1971;
Hedin et al., 1979; Booth et al., 1983). Hardee et al.
(1971) found that plant extracts signi®cantly increased
the attractive power of the boll weevil pheromone; and
Table 1
Traps tested for monitoring C. nenuphar and closely related species
Species Trap type Reference
C. nenuphar plastic funnel under trees Le Blanc et al., 1981
C. nenuphar sticky apple Yonce et al., 1995
C. nenuphar sticky plastic apple Yonce et al., 1995
C. nenuphar sticky band trunk trap Yonce et al., 1995
C. nenuphar pitfall trap under trees Yonce et al., 1995
C. nenuphar elevated funnel under trees Yonce et al., 1995
C. nenuphar funnel within tree canopy Eller and Bartelt, 1996
Curculio caryae pyramidal trap under trees Tedders and Wood, 1994
C. Vincent et al. / Agriculture, Ecosystems and Environment 73 (1999) 167±175 169
Dickens (1989) found that certain green leaf volatiles
(Visser and Ave, 1978) are also attractive to the boll
weevil.
So far attempts to identify potential attractants for
adult plum curculios in stored apples and aluminium
carbonate (Prokopy and Leskey, 1997), fresh apple
juice and synthetic apple blossom fragrance (Le
Blanc, 1982) have been unsuccessful. Leskey et al.
(1996) reported the attractiveness of a chemically
uncharacterized host odour produced by apples at
petal fall, and of water extracts of small apples and
apple twigs. Hexane extracts of various host parts were
only slightly attractive. Butkewich and Prokopy
(1993) found that because odours of host fruit were
signi®cantly less attractive at 4 and 8 cm than at 2 cm
from plum curculios, fruit odor-based traps are unli-
kely to be useful in commercial orchards.
Alm and Hall (1986) have described antennal sen-
sory structures on plum curculio that are similar to
pheromone receptors found on related curculionids.
Eller and Bartelt (1996) isolated and subsequently
synthesized an aggregation pheromone from male
plum curculios: (�)-(1R,2S)-Methyl-2-(1-methyl-
ethenyl) cyclobutaneacetic acid. This pheromone,
which they named grandisoõÈc acid, is attractive to
both sexes. Attempts to use this by placing live adults
in Tedder's pyramid traps were unsuccessful, baited
traps being no more attractive than unbaited traps,
possibly because of repulsive distress signals emitted
by the curculios (Prokopy and Leskey, 1997). How-
ever, lures impregnated with a racemic mixture of
grandisoõÈc acid have been reported by Johnson et al.
(1997) and Coombs et al. (1997) to increase signi®-
cantly the number of plum curculios trapped in Ted-
der's pyramid traps. Chouinard et al. (unpublished
data) demonstrated a two-fold increase in attractive-
ness when the lure was used in conjunction with small
amounts of green leaf volatiles; high amounts showed
a repulsive effect. Butkewich and Prokopy (1997)
found that both visual and olfactory lures are involved.
6. Entomogenous nematodes
Entomogenous nematodes are promising biocontrol
agents, especially for soil inhabiting pests. They can
enter their hosts by any ori®ces (i.e. spiracles, mouth
or anus) or through the cuticle (Tanada and Kaya,
1993). Tedders et al. (1982) tested several species of
nematodes for the control of plum curculio larvae and
concluded that they were ineffective under the con-
ditions tested. In laboratory studies the highest larval
mortality, i.e. 95.1%, was caused at a concentration of
400 Steinernema carpocapsae All nematodes per
larvae (Olthof and Hagley, 1993). There was no sig-
ni®cant increase in larval mortality from 200 to 400
nematodes per larvae. At these concentrations, 73.4%
larval mortality was achieved in natural sods. Nema-
tode treatments would also effect larvae of the apple
saw¯y, Hoplocampa testudinea Klug (Hymenoptera:
Tenthredinidae) (Vincent and BeÂlair, 1992). Nema-
tode treatments applied to the soil would not prevent
damage to apples. However, it could lower popula-
tions of both pests for the subsequent growing season.
To prevent damage to fruit, the effectiveness of
repeated applications of Steinernema carpocapsae
to the foliage or areal parts of apple trees was tested
(Chouinard et al., 1996). Although nematodes
remained infective on foliage from 24 to 48 h after
application, persistence was lower on ¯owers and
twigs (10±70 h). Some ®eld ef®cacy of canopy and
ground applications was detected in two years of ®eld
trials (5±55% plum curculio damage vs. 80±85% in
the control), but a replicated test performed in the third
year showed no reduction of fruit damage under
climatic conditions favourable to plum curculio activ-
ity, even after four weekly applications of S. carpo-
capsae, beginning at petal fall (28% damage vs. 31%
in the control). In caged trees, localized applications of
nematodes at the base of tree trunks signi®cantly
reduced adult populations maintained there (82±
100% mortality vs. 0±18% in the control).
7. Treatments of peripheral zones of appleorchards
The strategy to treat 20-m wide peripheral zones of
apple orchards (when needed) in spring was based on
the ®nding that plum curculio damage is frequently
more abundant at this time in peripheral zones (Le
Blanc et al., 1984), and that during the tight cluster
stage most plum curculio adults move only 1 to 4 m
per day when returning to the orchards from their
overwintering sites in adjacent woodlots (La¯eur and
Hill, 1987). During this 5±20 day re-invasion period in
170 C. Vincent et al. / Agriculture, Ecosystems and Environment 73 (1999) 167±175
southern Quebec, petal fall was selected as the most
appropriate time for this peripheral-zone treatment.
Using this approach in a 1.7 ha experimental orch-
ard (with standard-size trees) in Frelighsburg, Quebec,
fruit damage at harvest was reduced from 57% to 2.4%
(Chouinard et al., 1992b), while reducing the amount
of insecticide used by 70%, and the plum curculio
adult population by 83%. These results were consis-
tent over a two-year period in one locality. The
mortality data were based on recaptures of plum
Fig. 1. Field layout of an experiment on peripheral-zone versus full plot treatment (i.e. reference plot) in a commercial apple orchard.
C. Vincent et al. / Agriculture, Ecosystems and Environment 73 (1999) 167±175 171
curculio adults, radio-labelled with Zn65, that had
been released in a woodlot adjacent to an orchard
in which the peripheral zones had been treated with
insecticides.
In 1991 and 1992, peripheral-zone spraying was
compared for effectiveness with full-block spraying in
four commercial apple orchards in southern Quebec
(Fig. 1). All plots were subdivided into ®ve zones, i.e.,
peripheral zones (North, South, East and West) and a
central zone. To avoid economic losses, additional
sprays were applied if more than 1% of the fruit
examined in a given zone showed fresh curculio
oviposition scars. This was estimated by recording
oviposition scars on 20 apples, randomly selected,
from one side only of each apple tree in the outermost
row of the peripheral zones; and by examining 20
apples per tree on 25, randomly chosen, early cultivar
trees in the central zones.
Plum curculio damage at harvest varied from 0.0%
to 0.7% and from 0.0% to 0.8% fruit in plots receiving
peripheral sprays (experimental) and full-plot sprays
(reference) respectively; and most damaged apples
(95%) were found in peripheral zones. Total insect
damage on fruit at harvest varied from 1.3% to 3.8% in
experimental plots, and from 0.4% to 5.0% in refer-
ence plots.
8. Effects of peripheral treatments on non-targetarthropods
In 1991, there were no signi®cant differences
between population abundance estimates in peripheral
versus central zones of spotted tentiform leafminer
(Phyllonorycter blancardella Fab.: Gracillariidae),
apple aphids (Aphis pomi (DeGeer) and Aphis citri-
cola (Van der Goot): Aphididae), leafrollers (includ-
ing Choristoneura rosaceana Harris, Archips
argyrospilus Wlk., Argyrotaenia velutinana (Wlk.):
Tortricidae) and phytophagous mites (Tetranychus
urticae Koch: Tetranychidae, and Aculus schlechten-
dali (Nal.): Eriophyidae).
In 1991, in the experimental plots, populations of
woolly apple aphid, two-spotted spider mite and apple
rust mite were signi®cantly higher in peripheral zones
than in the central zones. In 1992, there were no
signi®cant differences between population estimates
of peripheral versus central zones for both experi-
mental and reference plots, for all species examined
and in all orchards. When the results from all localities
and years were pooled, peripheral zones of experi-
mental orchards received 1.22 insecticide treatments,
compared with 1.29 for reference orchards. However,
the central zones of experimental orchards received
0.38 insecticide treatments, versus 1.13 treatments for
reference plots.
The expected savings in pesticides are a function of
orchard size, the larger the area the larger the savings
(Fig. 2). Assuming a 20 m peripheral zone, depending
on plot size, the percentage saving of insecticide could
range from 20% to 85%.
Peripheral-zone treatments for plum curculio man-
agement is compatible with the agronomic complex-
ities of commercial orchards, e.g., multi-cultivar
plantations and odd-shaped orchards. It does require,
however, increased scouting efforts. From 1992 to
1996, implementation of peripheral-zone treatments
for plum curculio proved to be reliable under all
situations encountered by Quebec commercial apple
growers relying on scouting services: if plum curculio
was found in large numbers in all zones, these were all
treated (equivalent to a full-block treatment) (Y.
Morin, personal communication 1997).
Peripheral-zone treatments have also been used
successfully in Ontario against apple maggot, Rhago-
letis pomonella Walsh, and the codling moth, Cydia
pomonellla (L.) (Trimble and Solymar, 1997). As the
life cycle of each of these pests is quite different from
that of plum curculio (Table 2), they require different
Fig. 2. Expected savings of insecticides as a function of orchard
surface and geometry of the plot.
172 C. Vincent et al. / Agriculture, Ecosystems and Environment 73 (1999) 167±175
control strategies. The net impact of the plum curculio,
codling moth and apple maggot managed with per-
ipheral-zone treatments used sequentially is yet to be
evaluated on a short (within a season) and long (over
several seasons) term basis. On a long term basis,
concerns pertaining to the long term stability (sensu
Wildbolz, 1988) of partially treated ecosystems are to
be addressed.
Acknowledgements
The authors thank Y. Morin (Agrilus Inc., St-Alex-
andre, Quebec) for his participation in ®eld work.
Jean-Pierre R. Le Blanc, Nova Scotia Agricultural
College, Truro, N.S., N.J. Bostanian and Gaetan Rac-
ette (HRDC-AAAC/Saint-Jean-sur-Richelieu) kindly
commented an early version of the manuscript. C.
Brodeur (MAPAQ, St-Hyacinthe, Qc) assisted in writ-
ing the ®rst draft of the sections on trapping and
attractants.
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Differences among the tarnished plant bug, the plum curculio and the apple maggot, three insects that are entomological limits to biological
control programs in apple orchards of Eastern North America (Modified after Vincent and Roy, 1992)
Tarnished plant bug Plum curculio Apple maggot
Order Hemiptera Coleoptera Diptera
Family Miridae Curculionidae Tephritidae
Host plants ca. 100 Rosacae: apple, plum
blueberry, pear
apple, Crataegus spp.
Window of treatment pink petal fall summer
Larvae in fruit no yes yes
Target stage adult or Nymph adult adult
Activity day night and day day
Maximum damage to fruit in
unsprayed orchard*
15±20% 85% 50±60%
Monitoring white sticky trap Tedders traps (pyramidal) red spheres or AM1 traps
Known attractive compound none Grandisoic acid ammonium sulfate and
apple volatiles
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