Professional Documents
Culture Documents
J.D. Lea-Cox
Department of Natural Resource Science
and Landscape Architecture
University of Maryland
2120 Plant Science
College Park, MD 20742
USA
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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
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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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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
2500
2000
1500
1000
500
b
0
No Predators
Predators
Treatment
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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.
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90
Low N
80
Med N
High N
70
60
50
40
30
NS
20
NS
NS
S
10
0
Day 21
Day 35
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.
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