A Method for Laboratory Studies on the Polyphagous Predator

Macrolophus pygmaeus (Hemiptera: Miridae)

Author(s): Dionyssios Ch Perdikis
Source: Journal of Economic Entomology, 95(1):44-49. 2002.
Published By: Entomological Society of America
DOI: http://dx.doi.org/10.1603/0022-0493-95.1.44
URL: http://www.bioone.org/doi/full/10.1603/0022-0493-95.1.44

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 BIOLOGICAL AND MICROBIAL CONTROL

A Method for Laboratory Studies on the Polyphagous Predator

Macrolophus pygmaeus (Hemiptera: Miridae)
DIONYSSIOS CH. PERDIKIS
Laboratory of Agricultural Zoology and Entomology, Agricultural University of Athens, 75 Iera Odos,
118 55 Athens, Greece

J. Econ. Entomol. 95(1): 44Ð49 (2002)

ABSTRACT A method for studying the life history parameters of Macrolophus pygmaeus Rambur was
developed and described. Fecundity of the predator was studied by placing one female and one male
in a plastic cylinder in which a stem piece was used as an ovipositional substrate. Smaller cylinders
were used to study the percentage of egg hatch of M. pygmaeus. Results demonstrated that this method
is suitable for studying the fecundity and egg eclosion of M. pygmaeus. Allowing for easy modiÞcation,
the method offers suitable conditions for maintenance of both insect and plant material and can be
inexpensively modiÞed for different experiments and the parts are easily constructed. The proposed
method could also be employed for similar studies on other predatory hemipterans such as mirids and
anthocorids and very likely on other phytophagous insect species that lay their eggs in or on plant
stems.

KEY WORDS Macrolophus pygmaeus, method, oviposition, egg eclosion, Hemiptera

ALTHOUGH SOME SPECIES OF Miridae and Anthocoridae
are polyphagous predators, many are also phytopha-
gous. Several studies have been conducted on life
history parameters of some of those species. Proper or
at least acceptable methods for such studies are es-
sential and have to be developed. Frescata et al. (1994)
studied fecundity of Orius laevigatus (Fieber)
(Hemiptera: Anthocoridae) by placing one female
and one male in 2.1-liters plastic boxes, the top of
which was covered with a lid with two ventilation
holes. A potted cabbage plant infested with aphids as
prey was placed in each box and used as an oviposition
substrate. Females laid their eggs in plant tissue and
were counted on a daily basis. In that study, adult O.
laevigatus were removed using an aspirator or brush
when counting the eggs on the plant; however, high
adult mortality due to handling was observed. To avoid
the use of an aspirator or brush, the potted plant was
replaced by a leaf to allow the box to be opened for
removal of the leaf and to count the eggs. In the study
by Alauzet et al. (1994), the fecundity of O. laevigatus
was studied by placing one pair of adults in a plastic
box with plastic netting covering the top and bottom.
Humidity was maintained at high levels by placing the
box over a water tank. A geranium leaf placed in each
box was used as an oviposition substrate. Richards and
Schmidt (1996) conducted a study on the longevity
and fecundity of Orius insidiosus (Say) on several diets
by placing a pair of adults in a petri dish (5 cm in
diameter). Cocuzza et al. (1997) studied the fecundity
of Orius albidipennis (Reuter) and O. leavigatus by
placing a pair of adults in a Plexiglas container (9 cm
in diameter, 2.5 cm high) that had two holes at the
top covered with muslin for ventilation. Inside each
container a small pepper plant was placed as an ovi-
position substrate and a source of moisture. Percent-
age of egg eclosion of Nabis americoferus Carayon
(Hemiptera: Nabidae) and fecundity of Geocoris
punctipes (Say) (Hemiptera: Lygaeidae) also were
studied in petri dishes (Naranjo and Stimac 1985,
Guppy 1986).
Methods for rearing mirid species have been de-
scribed for Lygus hesperus Knight. Beards and Leigh
(1960) developed a method for rearing on green
beans, which served as both food supply and ovipo-
sition substrate, in jars with wide openings that al-
lowed the changing of plant material. Using this
method, Wilson (1973) observed that mold developed
in the boxes and that many adults escaped from the
wide opening when new bean were added. Wilson
(1973) therefore constructed two types of oviposition
cages with oviposition substrates (green beans) of-
fered through slits on the walls of the cages, which
prevented escape.
Species of the genus Macrolophus (Hemiptera: Miri-
dae) are polyphagous predators feeding on serious
pests like whiteßies and aphids on several vegetable
crops (Malausa et al. 1987, Goula and Alomar 1994,
Sampson and King 1996). Studies of life history pa-
rameters of Macrolophus spp. have been conducted
only with Macrolophus caliginosus Wagner. Fecundity
of M. caliginosus has been studied previously by Fauvel
et al. (1987). In that study one female and one male
predator were put in ventilated wood-framed boxes in

0022-0493/02/0044Ð0049$02.00/0 䉷 2002 Entomological Society of America

February 2002 PERDIKIS: A METHOD FOR LABORATORY STUDIES ON Macrolophus
which a stem piece of Pelargonium peltatum (L.) LÕ
Hér. in Aiton (Geraniaceae) was placed as an ovipo-
sition substrate. Van Schelt et al. (1995) and Hansen
et al. (1999) studied the fecundity of M. caliginosus in
petri dishes that each contained one female and one
male.
Percentage of egg eclosion of M. caliginosus was
studied by Fauvel et al. (1987) by using stem pieces of
tomato and P. peltatum as oviposition substrate. In the
studies by Riudavets and Castañé (1998) and Hansen
et al. (1999) a number of females and males were held
on tomato plants until oviposition occurred, after
which the percentage of egg eclosion was determined.
Field studies on the occurrence and abundance of
natural enemies of aphids and whiteßies in central
Greece have demonstrated that the predator Macrolo-
phus pygmaeus Rambur is the most important among
the natural enemies recorded (Perdikis and Lykoures-
sis 1996, Lykouressis et al. 1999 Ð2000). However, fur-
ther laboratory studies on its life history parameters
were needed to obtain information on its food and
climatic preferences. Knowledge of these aspects is
essential for more effective use of this species in bi-
ological control programs.
Consequently, the development of a suitable
method that could provide appropriate plant and cag-
ing conditions was necessary for studying its biological
parameters. In the development of such a method it
had to be taken into consideration that M. pygmaeus
feeds not only on insects but also on plant sap (Per-
dikis and Lykouressis 2000) and that the females ovi-
posit mainly in the stems of host plants. Thus, for the
study of its fecundity it was necessary to provide the
female not only with prey on leaves of host plants but
also with a piece of plant stem as an oviposition sub-
strate. In addition, the extreme mobility of the adults
necessitated the development of a method that would
prevent them from escaping during handling but
would provide them with adequate space to move
normally. Previous studies on nymphal development
of this predator have been successfully conducted in
petri dishes because the nymphs are much less mobile
(Perdikis and Lykouressis 1999).
The aim of the current work was to develop a
method for the study of the egg eclosion, fecundity,
and longevity of M. pygmaeus, which are characteris-
tics necessary for the estimation of its life history
parameters. Such a method had to meet the require-
ments of allowing easy daily handling and observation
while providing optimal conditions for the mainte-
nance and proper conservation of plant material. It
also had to be easily adapted to the demands of dif-
ferent experiments like the study of the percentage of
egg hatch and to enable us to use it, if necessary, after
appropriate modiÞcations, for studies on other insect
species with similar biological characteristics and be-
havior, such as members of Miridae, Anthocoridae,
Nabidae, and Lygaeidae. Finally, the method had to be
developed using simple devices made of inexpensive
and easily available materials.
45
Fig. 1. Plastic cylinder used for laboratory studies of M.
pygmaeus in which a stem piece and a leaf have been placed.
Materials and Methods
A method was developed to study fecundity of M.
pygmaeus and is described as follows. Newly emerged
adults are separated into pairs (one female and one
male), and each pair is released in a cylinder made of
transparent 0.4-mm PVC (6.5 cm diameter and 30 cm
high) (Fig. 1). A rectangular window (6 by 13 cm) is
opened 7 cm from the cylinder base. The window was
not made exactly in the middle of the cylinder, but at
a shorter distance from the cylinder base than the top,
thus allowing for easier manipulations. The window, as
well as the top of the cylinder, is covered with Þne
muslin to keep temperature and humidity inside the
cylinder at similar levels to that inside the growth
cabinet. A disc of foam (ßower foam, Smithers Oasis,
Ohio) 2 cm high was placed inside the base of the
cylinder (Fig. 1). This material is used widely for
preservation of ßowers and characteristically absorbs
water and retains it for an extended period after being
soaked (⬇3 d at 25⬚C). The upper surface of the foam
was covered with aluminum foil to prevent an increase
in relative humidity inside the cylinder.
A section of a plant stem and two to three leaves
bearing insect prey for the predator were placed in
each cylinder (Fig. 1). Each stem was 6 Ð7 cm long and
5Ð7 mm in diameter. Stems of this length provide
adequate substrate for oviposition for 24 h. This length
allows us to obtain numerous stem pieces from a plant
stem and conserves plant material.
The base of each stem piece and the petioles of the
leaves were inserted (1 cm) into moistened foam,
which prevents dehydration. The foam is sufÞciently
hard to keep the stem piece and leaves in vertical
positions, which enhances the detection and obser-
vation of the predators. This also facilitates proper
manipulation of leaves and stems when being changed
and reduces escape of insects.
46 JOURNAL OF ECONOMIC ENTOMOLOGY
Fig. 2. Parts of the model (cylinder, base, and plastic tray) used for laboratory studies of M. pygmaeus.
When the stem piece and the leaves have been
inserted into the foam, each cylinder is placed upright
in a round hole (6.5 cm diameter by 3 cm deep) in a
base of expanded polystyrene foam (Fig. 2). In this
way, cylinders are held vertically and easily trans-
ported. This base is 45 by 26 cm and has eight holes.
The base is placed on a plastic tray, with the same
dimensions, 2 cm deep. Water added to the plastic tray
can maintain moisture in the foam. The base is Þrmly
attached to the plastic tray with transparent sticky tape.
Leaves and stem pieces are changed through the
lower part of the window in the cylinder. Muslin on
the lower edges of the window is held in place with
transparent sticky tape to allow reopening. Long for-
ceps are used to remove stems and leaves through the
window.
We used this method to study the percentage of egg
hatch of M. pygmaeus, and placed stems bearing eggs
in small cylinders (PVC 0.21 mm thick, 4 cm in diam-
eter, 11 cm high).
We studied the fecundity of the predator M. pyg-
maeus following the method described earlier (in a
growth cabinet at 20⬚C, 65 ⫾ 5% RH, and a photope-
riod of 16:8 [L:D] h, on eggplant, pepper, and tomato,
in the presence of prey). The aphid Myzus persicae
(Sulzer) (Homoptera: Aphididae) was used as prey
Vol. 95, no. 1
on the eggplant and pepper plant and the whiteßy
Trialeurodes vaporariorum (Westwood) (Homoptera:
Aleyrodidae) was used on tomato. Each combination
was replicated 20 times (one pair in each cylinder) on
eggplant and tomato and 17 times on pepper plants.
The numbers of eggs were counted at 24-h intervals.
The success of egg hatch was examined in growth
cabinets with controlled conditions at 15, 20, 25, and
30⬚C; 65 ⫾ 5% RH; and a photoperiod of 16:8 (L:D) h.
Stem pieces of each of eggplant, pepper plant, or
tomato, bearing 20 eggs of M. pygmaeus in total, were
placed in small cylinders and held at each of the above
temperatures. The stems and the inside of cylinders
were inspected every 24 h and emerged nymphs were
collected. Stems were held in the cylinders for a pe-
riod much longer than the maximum egg incubation
period to make sure that all hatched eggs were de-
tected.
To test for signiÞcance, a chi-square test (P ⬍ 0.05)
was used to compare the data on egg hatch. Data were
arcsine-transformed before statistical analysis.
Results and Discussion
Leaves in the cylinders were in very good condition
for two or more days, depending on the temperature
February 2002 PERDIKIS: A METHOD FOR LABORATORY STUDIES ON Macrolophus 47

Table 1. Methods used in studies on fecundity and egg hatch of Macrolophus caliginosus

Life history
Method Temp,⬚C Results Reference
parameter studied
Fecundity One female and one male in a wood-framed 20 268 eggs/씸 Fauvel et al. (1987)
box with stem pieces of Pelargonium
peltatum
One female and one male in a petri dish- 23 2Ð7 eggs/씸/day van Schelt et al. (1995)
type plastic tray with a tomato leaf disc
One female and one male, in a petri dish 22 20.3 eggs/씸 Hansen et al. (1999)
with 3 leaves of tomato
Egg hatch Stem pieces of tomato with eggs were 20 23.1% Fauvel et al. (1987)
transferred to a cabinet and kept there
until egg hatch
25 55.6%
30 42.4%
Cohort of females oviposited on plants, eggs 25 80.0% Riudavets and Castañé (1998)
were marked and their eclosion was
recorded
Cohort of females oviposited on plants, the 22 97.4% Hansen et al. (1999)
nymphs emerged were counted and
Þnally the number of eggs oviposited was
measured

(⬇2 d at 25⬚C) and plant stems also were in good (1999) put one male and one female of the predator
condition at least until eggs hatched, i.e., ⬇25 d at 15⬚C in a petri dish (15 cm diameter, with three mesh-
and 15 d at 25⬚C. Therefore, this method allowed covered holes of 1 cm in the lid) along with three
maintenance of plant material in good condition for a leaves of tomato. Results of the Hansen et al. (1999)
long enough period to conduct insect studies. This is (Table 1) study were not comparable to ours because
of crucial importance for a high percentage of egg the prey species used (Tetranychus urticae Koch [Aca-
hatch (Constant et al. 1996). rina: Tetranychidae]) was less suitable for fecundity
This method can easily be adapted for various ex- of M. caliginosus. However, the study of M. caliginosus
periments concerning studies on life history parame- fecundity in petri dishes does not seem to be the best
ters of M. pygmaeus. The size of the cylinders can be approach because females can escape from the dishes
changed easily to accommodate the requirements of (Hansen et al. 1999). We also observed this in pre-
different experiments. liminary experiments on fecundity of M. pygmaeus and
In the experiment on the fecundity of M. pygmaeus, we discontinued using this method.
all females of M. pygmaeus laid eggs, with fecundity Egg hatch of M. pygmaeus reached 90% at 15 and
reaching an average of 214, 204, and 228 eggs on 20⬚C on eggplant at both temperatures, 85 and 90% on
eggplant, pepper plant, and tomato, respectively, at pepper plant, and 90 and 85% on tomato, respectively
20⬚C. Fauvel et al. (1987) studied fecundity of M. (Table 2). Egg hatch at 30⬚C was 60, 60, and 55% on
caliginosus fed on eggs of Ephestia kuehniella (Zeller) eggplant, pepper plant, and tomato, respectively. Egg
(Lepidoptera: Pyralidae) by placing one female and hatch at each temperature did not differ signiÞcantly
one male in a ventilated box (23 cm high by 10 cm among the three host plants (Table 2), indicating that
diameter), with a stem piece of P. peltatum as ovipo- this method was almost equally suitable for the main-
sitional substrate. In that study, M. caliginosus had a tenance of stem pieces of all the host plants tested.
relatively higher level of fecundity (average 268 eggs In a similar study by Fauvel et al. (1987), the per-
at the same temperature) (Table 1) than what we centage of egg eclosion of M. caliginosus was examined
observed from M. pygmaeus in the current study. Fau- by using stems of tomato (10 cm long and ⬇8 mm
vel et al. (1987) reported that E. kuehniella eggs were diameter) as an oviposition substrate of the predator.
a more suitable diet than T. vaporariorum for ovipo-
sition of M. caliginosus; however, fecundity on the
latter was not reported. Therefore, considering that Table 2. Hatch of eggs of M. pygmaeus (%) at different tem-
these two predators are close relatives and results of peratures and on different host plants and results of statistical
both studies on fecundity are similar it seems that the analysis of those percentages among the host plants at each
method developed here is suitable for studies on the temperature
oviposition of Macrolophus species. Fecundity of M.
Host plant
caliginosus was also studied by van Schelt et al. (1995), Temp,⬚C
Eggplant Pepper plant Tomato ␹2 P
who placed one female and one male in a petri dish (8
cm diameter by 3 cm high) with T. vaporariorum. In 15 90 85 90 0.19 0.90
each petri dish a tomato leaf disc was placed upside 20 90 90 85 0.19 0.90
25 75 80 75 0.10 0.95
down on a 1-cm agar layer; however, fecundity of the 30 60 60 55 0.09 0.96
predator was studied only for the Þrst 15 d of its
reproductive life. In a similar study, Hansen et al. Twenty replicates were used for each host plant and temperature.
48 JOURNAL OF ECONOMIC ENTOMOLOGY
The lower part of each stem was inserted in a tube
Þlled with a nutrient solution. The stem and the tube,
together with a female, were placed in a 3-cm-diam-
eter glass tube, both ends of which were tapered with
cork. The eggs deposited every 24 h were counted and
each stem was transferred to a cabinet with 60% RH
and kept there until egg hatch. In their study the
percentage of egg hatch was 23.1% at 20⬚C, 55.6% at 25,
and 42.4% at 30⬚C (Table 1), which are values much
lower than those in our study (Table 2).
Previous studies encountered problems when to-
mato stems were not kept at suitable conditions for egg
hatch (Fauvel et al. 1987). In many cases, humidity
was a problem and in high humidity, mold developed
on stems; whereas in low humidity, they became too
dry. Therefore, the higher percentages of egg hatch in
M. pygmaeus than in M. caliginosus, are mainly due to
the better condition of tomato stems used in our
method. However, the percentage of egg hatch of M.
caliginosus when oviposited in stems of P. peltatum was
similar to those found in M. pygmaeus (87, 88, and 76%
at 15, 20, and 25⬚C, respectively), mainly due to the
better tolerance of the stems of that plant than the
stems of tomato either in humid or dry conditions
(Fauvel et al. 1987). In the study by Riudavets and
Castañé (1998), egg eclosion of M. caliginosus was
examined by placing 50 Ð75 adults (males and females)
on Þve tomato plants. Eleven eggs per plant were
marked and their eclosion was recorded. In a similar
study by Hansen et al. (1999), females that were 6 Ð13
d old were kept for 1 wk in a ßower pot with two
tomato plants for oviposition. Emergent nymphs were
counted, and 1 mo after introduction of the females
the number of eggs oviposited was determined. The
results of both of these studies are similar to ours
(Table 1). In their studies, however, it is not possible
to determine either the number and/or the hatchibil-
ity of eggs oviposited per day by each female, or the
consumption of prey by each female. These results
may be important in studies, such as examination of
food preferences or determining female consumption
of prey.
In conclusion, the method developed and described
here provides appropriate conditions for the study of
life history parameters of M. pygmaeus based on ovi-
position and egg hatch studies, which were consistent
between all host plants tested. Moreover, this method
has some advantages compared with other methods
used for similar studies; because it is easily adapted to
the needs of various experiments, it offers suitable
conditions for maintenance of both insects and plant
material, it enables easy manipulation through the
windows, and inexpensive materials are used. Further-
more, it can be easily constructed and observations are
more easily made than in other methods used for
similar studies of insects within this genus. This
method could also be suitable for study on life history
parameters of other hemipteran species as well as on
other phytophagous insects that show similar ovipo-
sitional behavior laying their eggs within or on the
surface of plant stems.
Vol. 95, no. 1
Acknowledgments
The author thanks Dionyssios Lykouressis (Agricultural
University of Athens) for the useful suggestions and improve-
ments he made to the manuscript.
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Received for publication 16 November 2000; accepted 20
May 2001.