Integrating Cultural Management Practices and Biological Control to

Suppress Citrus Mealybug

P.M. Shrewsbury and K. Bejleri
Department of Entomology
University of Maryland
4112 Plant Science
College Park, MD 20742
USA

J.D. Lea-Cox
Department of Natural Resource Science
and Landscape Architecture
University of Maryland
2120 Plant Science
College Park, MD 20742
USA

Keywords: cultural control, augmentation, water stress, fertilization, Cryptolaemus

montrouzieri, ornamentals
Abstract
The citrus mealybug, Planococcus citri (Risso) (Hemiptera: Pseudococcidae),
is one of the most common and damaging insect pests in greenhouses and protected
cultures. Pesticides are frequently used, often unsuccessfully, to control this pest. Our
objective was to determine the influence of water and nutrient management practices
on mealybug populations and their impact on the efficacy of augmentative releases of
Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae), a mealybug
predator. Studies were conducted on Heuchera micrantha Palace Purple growing in
protected culture. Heuchera plants received 3 rates of nitrogen: low (25 ppm/wk),
medium (75 ppm/wk), and high (150 ppm/wk). Within each fertilizer treatment
plants were either water stressed or not water stressed. In the absence of predators,
citrus mealybug populations increased dramatically on water stressed plants
receiving high and low levels of fertilizer. Cryptolaemus montrouzieri reduced
mealybug populations and consistently had the greatest impact on mealybugs where
mealybugs were the most abundant, regardless of fertilizer or water treatment. In the
presence of predators, mealybug populations are regulated by predation rather than
mealybug interactions with fertilizer or water. This data suggests plant management
practices that avoid high or low rates of fertilization of water stressed plants should
reduce the outbreak potential of citrus mealybug. If mealybugs do outbreak,
implementing augmentative biological control with C. montrouzieri should be
effective regardless of fertilization and irrigation practices.
INTRODUCTION
Insects and mites frequently outbreak in intensive plant production systems
causing economic and aesthetic damage to plants. Plant production practices such as
fertilization and irrigation, can influence herbivores directly or indirectly leading to
changes in pest population levels. In their comprehensive review of more than 450
studies, Waring and Cobb (1992) presented strong evidence that sucking insects and mites
overwhelmingly responded to elevated levels of nitrogen fertilization by increasing in
abundance through higher survival, increased growth, or greater fecundity. Responses of
sucking insects and mites to experimental water stress on herbaceous plants were less
clear (Waring and Cobb, 1992). Sucking insects were equally favored or disfavored by
plant water stress while mites generally benefited by water stress (Waring and Cobb,
1992). Kyto et al. (1996) reviewed 79 studies of insects and mites feeding on trees and
concluded that sucking insects and mites generally benefit at both the individual and
population levels to increased fertilization. A recent review of 70 studies involving insect
responses to plant stress found that sucking insects were generally favored by water stress
(Koricheva and Larsson, 1998). Fertilization has also been shown to influence
colonization of plants by insects. Azalea lace bug, Stephanitis pyrioides Scott, adults more
readily colonized azaleas fertilized with high rates of nitrogen compared to low rates
(Casey and Raupp, 1999). Greenhouse whitefly, Trialeurodes vaporariorum (Westwood),
Proc. XXVI IHC Protected Cultivation 2002
Ed. A.P. Papadopoulos
Acta Hort. 633, ISHS 2004
Publication supported by Can. Int. Dev. Agency (CIDA)

425

adults preferentially selected and oviposited more on tomato plants receiving high doses
of nitrogen (308 ppm) compared to plants receiving lower doses (140 and 84 ppm) (Jauset
et al., 1998). These studies suggest it may be possible to manage pest populations by
altering fertilization and water management practices to reduce herbivore performance
and fitness.
Altering fertilization and water management practices may not only influence
plant/herbivore interactions but also predator/prey dynamics. Some predators respond
numerically to increasing prey densities (Price, 1984; Murdoch, 1990) by increasing their
reproduction and/or by aggregating in areas of high prey density. This suggests that
predators may have a greater impact on herbivores that occur at higher densities than
lower. Altering cultural management practices may influence both bottom up (plant
quality) and top down (natural enemies) forces that regulate herbivore populations.
We evaluated the impact of two management strategies, cultural and biological
control, used singularly and in combination, to determine the influence of various
fertilization and water regimes on herbivore abundance and the efficacy of augmentative
releases of predators. We used the citrus mealybug, Planococcus citri (Risso), infesting
Heuchera micrantha Palace Purple a herbaceous perennial, and the mealybug predator,
Cryptolaemus montrouzieri Mulsant, as our herbivore/host plant/natural enemy study
system.
The citrus mealybug is a key pest of ornamental plants grown in interiorscapes and
protected cultures. It has a wide host range feeding on plants in over 25 families (Copland
et al., 1985; Dreistadt, 2001). Mealybugs have sucking mouthparts, feed on phloem, and
their feeding reduces plant vigor, causes yellowing and distortion of foliage, and
defoliation (Dreistadt, 2001). Citrus mealybug produces copius amounts of honeydew
(Copland et al., 1985; Dreistadt, 2001). Mealybugs have multiple, overlapping generations a year. C. montrouzieri is readily available from suppliers of biological control
agents (Hunter, 1997) and there are many anecdotal accounts and some empirical data of
C. montrouzieri suppressing citrus mealy bug populations (Copland et al., 1985).
To our knowledge no studies have evaluated the influence of fertilizer and water
stress on citrus mealybug and C. montrouzieri population dynamics. We hypothesized
that: 1) increased rates of nitrogen fertilizer will increase mealybug abundance, especially
when plants are water stressed; and 2) predators will have the greatest impact where
mealybug densities are highest.
MATERIALS AND METHODS
Experimental Design
This study was conducted in the nursery yard at the University of Maryland
Greenhouse Facility, College Park MD, USA, during the late summer and fall of 2001.
Heuchera plants were purchased as plugs and potted into #1 containers using Metro Mix
510 (Scotts Sierra Hort. Products Company, Marysville, OH). Two hundred and fifty
containerized Heuchera plants were lined out in a nursery bed. A clear plastic covering,
open on all sides, was erected approximately 1.6 m above the plants to prevent rainfall
from reaching the plants and interfering with water stress treatments. A 3 x 2 factorial
design, where treatments included three levels of nitrogen fertilizer (low, medium, and
high) and two levels of irrigation (water stressed and not water stressed), was used. Each
level of fertilizer received each irrigation treatment (= 6 treatment combinations total).
Heuchera plants received fertilizer and irrigation treatments beginning 10 May 2001 (3
months prior to infestation by mealybug) through 10 October 2001 at which time the
study was terminated.
Heuchera plants received weekly applications of nitrogen at rates of 25 ppm
(low), 75 ppm (medium), or 150 ppm (high). A balanced ratio of nitrogen to other
nutrients was maintained (see Handreck and Black, 1994; Nelson, 1998). Nutrient
solutions were composed of ammonium nitrate, potassium sulfate, potassium phosphate,
and potassium nitrate (Fisher Company, Burr Ridge, IL). Solutions for each treatment
426

were prepared in the laboratory. We determined that the water holding capacity of the
study pots was 0.9 liters. Plants were fertilized every Monday with the fertilizer solution
that provided them with the appropriate treatment dose of nitrogen in 0.9 liters water
using a Dosatron irrigation system (Dosatron International, Inc).
Irrigation treatments included water stressed and non-water stressed plants. Trials
were conducted to determine the soil water content at which Heuchera plants reached a
water stressed state and at what level they were non-stressed as indicated by stomatal
conductance. In a controlled environment, we determined the relationship between soil
water content using a Theta Probe (Type ML 2X Soil Moisture Sensor by Dynamax) and
stomatal conductance using a Porometer (LI-1600, LI-COR, Inc., Lincoln, NE) for the
plant/potting mix in our study. To maintain plants in a non-water stressed state, plants
were irrigated when water volumetric (Wv) content reached between 80-95%. This
corresponded to stomatal conductance of 80 mmol CO2/m2/sec. Plants in the water stress
treatment were watered when water volumetric content reached between 40-50%. This
corresponded to stomatal conductance of 50-60 mmol CO2/m2/sec. Every morning the
average stomatal conductance of Heuchera, in each water treatment, was measured using
a Theta Probe. Plants were irrigated as needed to maintain their assigned water treatment.
Citrus Mealybug Infestation Prior to Predator Release
To determine if mealybugs differentially infest Heuchera under the varying
fertilizer and water regimes, citrus mealybugs were allowed to naturally infest plants.
Mealybug counts were taken approximately one week after mealybugs were first noted on
the plants (20 August 2001). Sampling of mealybugs was conducted by removing 1
mature leaf (approx. the 8th leaf out from whorl) and 1 new leaf (approx. the 3rd leaf out
from whorl) from 3 plants (replicates) in each of the 6 treatment combinations (3 nitrogen
x 2 water). Leaves were brought back to the lab where mealybugs (all active stages) were
counted under the dissecting microscope on the upper and lower surface and the petiole of
the leaves.
Efficacy of Predator
To evaluate the efficacy of augmentative release of C. montrouzieri, three plants
from each of the six treatment combinations (3 replicates x 6 treatments = 18 plants) were
placed in a lightweight mesh cage to exclude predators (no predator treatment) and three
plants from each of the six treatment combinations (3 replicates x 6 treatments = 18
plants) were used to monitor citrus mealybug densities exposed to predators (predator
treatment). C. montrouzieri adults were purchased from a commercial supplier of beneficials (The Green Spot, Nottingham NH, USA) and releases were made onto the
remaining uncaged Heuchera in the nursery bed on 29 August 2001. Following guidelines
provided by the producer, two releases of C. montrouzieri were made at a rate of 9
predators/plant and 4 predators/plant, respectively with 8 days between releases. When
predators arrived, groups of ten were placed into small 25 mm diameter petri dishes.
Releases were made late in the day after sunset, plants were misted lightly with water,
petri dishes were evenly distributed throughout the nursery bed, and placed on the soil
surface of a pot. The petri cover was then removed and C. montrouzieri were allowed to
disperse from the petri dishes onto the plants. Two post counts were taken of citrus
mealybug. A post count was conducted 21 days after the first predator release. Mealybugs
were counted as described above on the 18 caged plants (no predator treatment) and 18
plants (3 from each of the 6 fertilizer/water treatments) exposed to predators. A second
post count was conducted 35 days after the first predator release. Mealybug counts were
taken only on 18 plants (3 from each of the 6 fertilizer/water treatments) exposed to
predators. No second post count was taken from caged plants (no predator treatments)
because all plants died prior to 35 days after treatment. Comparisons were made of citrus
mealybug densities in the presence and absence of predators across all fertilizer and water
treatments to determine the overall effect of predators on mealybugs, regardless of
cultural practices.
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Influence of Fertilizer and Water Regimes on Mealybugs and Predators

To determine the effects of fertilizer and water on mealybug populations in the
absence of predators, we examined the change in mealybug populations over time.
Population change was expressed as the number of mealybugs at day 21 minus the
number of mealybugs at day 0 divided by the number of mealybugs at day 0 times 100 (=
proportional change in mealybug population).
The impact of C. montrouzieri on mealybug populations with respect to fertilizer
and water was evaluated in two ways. First, we compared the abundance of mealybugs on
plants exposed to predators among fertilizer and water treatments, 21 and 35 days after
the release of predators. Next, we calculated the change in mealybug population growth
attributable to the predator. Population change was estimated by subtracting the number
of mealybugs remaining at day 21 in the presence of predators from the number of
mealybugs remaining at day 21 in their absence (cage controls) divided by the number of
mealybugs in the cage controls times 100 (= % reduction in mealybug population).
Statistical Analysis
All data were tested to determine if assumptions of normality and homogeneity
were met. Log10 or arc sin transformations were performed where data did not meet the
assumptions. Non-transformed means are presented in figures. ANOVA for a factorial
design (Proc Mixed, SAS Institute, 1999) were used to test for interactions and main
effects of water, nitrogen, and time unless stated otherwise. Means were separated by a ttest, Bonferoni test, or pairwise comparisons for differences in least squares means. Data
that could not be normalized by a transformation were analyzed by a Kruskal Wallis nonparametric test (SAS Institute, 1999) for differences among nitrogen/water treatments.
RESULTS
Citrus Mealybug Infestation Prior to Predator Release
When mealybugs were allowed to naturally infest Heuchera growing under
different fertilizer and water stress regimes there was no significant water by nitrogen
interaction (F = 1.34, df = 2, 12, P = 0.2978) or main effects of water (F = 3.77, df = 1,
12, P = 0.0762) or nitrogen (F = 0.76, df = 2, 12, P = 0.4882) on mealybug densities.
Although, water stress was barely not significant, mean ( SEM) mealybug densities were
as follows on the 6 different nitrogen/water treatment combinations: high nitrogen, water
stress = 34.7 (16.9); high nitrogen, no water stress = 138.2 (24.6); medium nitrogen, water
stress = 114.7 (28.4); medium nitrogen, no water stress = 128.1 (15.1); low nitrogen,
water stress = 81.1 (21.7); low nitrogen, no water stress = 106.2 (54.4). Mean
comparisons found mealybug densities on plants receiving high nitrogen and water stress
were significantly lower than on plants receiving high nitrogen and no water stress (P =
0.037) and plants receiving medium nitrogen and no water stress (P = 0.0474). There were
no significant differences in mealybug densities between all other treatments.
Efficacy of Predators
To evaluate the efficacy of C. montrouzieri in suppressing citrus mealybug we first
examined the population growth of mealybugs over time in the absence of predators.
Mealybug density (mean SEM) increased dramatically from 106 25.5 per leaf on day
0 to 1,869 384 per leaf on day 21 (t = -7.6, df = 68, P = 0.0001). To assess the impact of
C. montrouzieri on citrus mealybug populations, we compared mealybug densities on
Heuchera plants that were exposed to predators (predator treatment) to mealybug
densities with predators excluded by cages (no predator treatment) 21 days after predators
were released. There was a significant decrease, 16 fold, in mealybug densities when they
were exposed to C. montrouzieri (F = 124.4, df = 2, 101, P = 0.0001) (Fig. 1). A second
post count of mealybugs on plants exposed to predators 35 days after predators were
released found a further reduction in mealybug densities (12.36 2.98). All control plants
(predators excluded) died prior to day 35, apparently from high mealybug populations,
and therefore could not be compared with predator treatment plants.
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Influence of Fertilizer and Water Regimes on Mealybugs and Predators

To evaluate the influence of fertilizer and water regimes on suppression of citrus
mealybug, we examined mealybug densities at each of the six fertilizer/water stress
treatment combinations at 21 days post predator release in the presence and absence of
predators and at 35 days post release in the presence of predators. When we compared the
proportional change in population of mealybugs, on day 21 in the absence of predators,
there was no significant water by nitrogen interaction (F = 1.56; df = 2, 11; P = 0.2534) or
main effect of water (F = 1.83; df = 1, 11; P = 0.2028). However, there was a significant
main effect of nitrogen (F = 4.35; df = 2, 11; P = 0.0405). There were significant
differences between water/nitrogen treatment combinations (Fig. 2). Plants receiving the
highest level of nitrogen and water stress supported a greater increase in mealybug
populations than plants receiving high nitrogen, no water stress and medium levels of
nitrogen (water stressed and not water stressed) but not plants receiving low levels of
nitrogen (water stressed and not water stressed).
When we compared the abundance of mealybugs on plants exposed to predators
among fertilizer and water treatments, 21 and 35 days after the release of predators there
was a significant interaction between nitrogen, water, and day (F = 13.6; df = 2, 23; P =
0.001) (Fig. 3). Therefore, all lower level interactions (nitrogen*day: F = 2.23; df = 2, 23;
P = 0.1300; water*day: F = 2.93; df = 1, 23; P = 0.1007; water*nitrogen: F = 0.29; df = 2,
23; P = 0.7501) and main effects (day: F = 16.7; df = 1, 23; P = 0.0001; nitrogen: F =
1.46; df = 2, 23; P = 0.2535; water: F = 1.00; df = 1, 23; P = 0.3271) can not be addressed.
Plants/treatments with the highest level of mealybugs on day 21 had declining mealybug
populations on day 35 (low nitrogen, water stress; medium nitrogen, water stress; and
high nitrogen, no water stress). Whereas plants/treatments with lower level of mealybugs
on day 21 had similar (low nitrogen, no water stress) or increasing mealybug populations
(medium nitrogen, no water stress; (high nitrogen, no water stress) on day 35 (Fig. 3).
When examining the ability of C. montrouzieri to suppress citrus mealybug
populations (% reduction) on Heuchera receiving the various nitrogen and water regimes,
variances in the percent change in mealybug population were non-homogeneous and
could not be corrected by transformations. Therefore, we compared population change
across the six treatment regimes with a non-parametric Kruskal Wallis test. The percent
reduction in mealybug densities when C. montrouzieri was present did not differ among
water and fertilizer treatment combinations (X2 = 1.18, df = 5, P = 0.9471). Mealybug
reduction from C. montrouzieri was remarkably high in all nitrogen/water treatment
combinations ranging from 78 to 98%.
DISCUSSION
Citrus mealybug populations increased dramatically on Heuchera micrantha in the
absence of natural enemies. When exposed to the coccinellid predator, C. montrouzieri,
mealybug populations were reduced to very low densities. The impact of C. montrouzieri
on mealybug populations in this study is similar to other studies of C. montrouzieri
releases. C. montrouzieri provided successful control of citrus mealybug on various
ornamentals (Copland et al., 1985), coffee mealybug on coffee in India (Chacko, 1979),
and striped mealybug, Ferrisia virgta (Ckll.) on guava in India (Mani et al., 1990).
The greatest increase in citrus mealybug populations, in the absence of natural
enemies, occurred on Heuchera plants that were water stressed and receiving high or low
levels of nitrogen fertilizer. These results support our hypothesis and are consistent with
the findings of Koricheva and Larsson (1998) in that sucking insects are generally favored
by water stress. Additionally, sucking insects are generally more abundant on plants with
elevated levels of fertilization (reviewed by Waring and Cobb, 1992).
In a study similar to ours using melon aphid, Aphis gosypii Glover, on
chrysanthemum, Bethke et al. (1998) found that varying nitrogen (3 rates) and irrigation
(high and low levels) did not affect aphid fecundity, longevity, or survival. Our results
differ from the results of this study. However, in a related study by the same group,
Schuch et al. (1998) found thrips on chrysanthemum responded similarly to the
429

mealybugs in our study. Thrips were more abundant on plants receiving higher levels of
nitrogen and on water stressed plants.
C. montrouzieri consistently had the greatest impact where mealybugs were the
most abundant. On plants where mealybugs were the most abundant 21 days post predator
release, populations declined by day 35. Whereas on plants where populations of
mealybugs were the lowest on day 21, populations increased or remained the same by day
35. These results support our hypothesis. The patterns we see suggest mealybug
populations are influenced by an aggregative or numerical response by predators to
mealybug density (Murdoch, 1990), rather than by mealybug interactions with fertilizer or
water. Further support for this response is suggested when we examine mealybug
densities on day 21 in the absence of predators. Without predators mealybug densities are
high on water stressed plants receiving high nitrogen. In contrast, mealybug densities are
the lowest on these plants when predators are present. We did not specifically test for
numerical response in our predator/prey system and we recognize other factors may also
be influencing mealybug populations.
To our knowledge the only other study that examined the influence of cultural
practices on augmentative releases of natural enemies evaluated the influence of water
stress on the performance of cassava mealybug and three associated parasitoids
(Calatayud et al., 2002). This study was conducted in the laboratory and found water
stress favored cassava mealybug development and decreased parasitism. Water stress
enhanced the ability of mealybugs to encapsulate its parasitoids (Calatayud et al., 2002).
These studies support our hypotheses that: higher rates of fertilizer increase
mealybug abundance, especially when plants are water stressed; and that predators have
the greatest impact where mealybug densities are highest. We also found that overall there
were very high reductions in mealybug populations by C. montrouzieri, irrespective of
fertilizer or water treatments.
From a pest management perspective, our data suggest that fertilizer and water
management practices can influence citrus mealybug populations on Heuchera.
Preventing plants from becoming water stressed, regardless of fertilization rate, should
reduce the outbreak potential of citrus mealybug. In addition, moderate rates of nitrogen
(75 ppm/wk) should not promote mealybug outbreaks whether plants are water stressed or
not. If plants do outbreak with mealybugs, implementing augmentative biological control
of citrus mealybug with C. montrouzieri should be effective at reducing mealybug
populations regardless of fertilization and irrigation practices.
ACKNOWLEDGEMENTS
We thank Stacey Bealmear and Andrew Ristvey for their technical assistance with
this project. We also thank Michael Raupp and an anonymous reviewer for comments on
an earlier draft of this manuscript and Andrea Huberty for statistical guidance. This
project was funded in part by the Maryland Agricultural Experiment Station.
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Figurese

Number of Mealybugs / Leaf

2500

2000

1500

1000

500

b
0

No Predators

Predators

Treatment

Fig. 1. Mean SEM number of citrus mealybugs on Heuchera micrantha following

augmentative release of Cryptolaemus montrouzieri. Mealybugs were counted 21
days after predator release from plants exposed to predators and plants with
predators excluded (control). Bars with the same letter are not significantly
different (Bonferroni test, P < 0.05).

432

Proportional Change in Mealybugs

550
No Water Stress

Water Stress

500
450
400

ab

350
300
250
200
150
100

a
a

50

Low

Medium

High

Fertilization Level

Fig. 2. Mean SEM proportional change in citrus mealybug populations in the absence of
natural enemies after 21 days on Heuchera micrantha receiving low (25 ppm),
medium (75 ppm), and high (150 ppm) levels of nitrogen. Within each fertilizer
treatment plants were either water stressed or not water stressed. There was no
significant nitrogen by water interaction (F = 1.56; df = 2, 11; P = 0.2534) or main
effect of water (F = 1.83; df = 1, 11; P = 0.2028). However, there was a main
effect of nitrogen (F = 4.35; df = 2, 11; P = 0.0405). Bars with the same letter
represent no significant difference between nitrogen/water treatment combinations. Proportional change = (abundance day 21 - abundance day 0) / abundance
day 0.

433

Number of Mealybugs / Leaf

90

Low N
80

Med N
High N

70
60

50
40

30

NS

20

NS
NS
S

10
0

Day 21

Day 35

Days After Predator Release

Fig. 3. Mean SEM number of citrus mealybug 21 and 35 days following augmentative
release of Cryptolaemus montrouzieri. Mealybugs were sampled from Heuchera
micrantha receiving low (25 ppm) (Low N), medium (75 ppm) (Med N), and high
(150 ppm) (High N) levels of nitrogen. Within each fertilizer treatment plants
were either water stressed (S) or not water stressed (NS). There was a significant
interaction between nitrogen*water*day (F = 13.6; df = 2, 23; P = 0.001). There
were no other significant interactions or main effects.

434