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Biological Control of Eotetranychus lewisi and Tetranychus urticae
(Acari: Tetranychidae) on Strawberry by four Phytoseiids (Acari:
Phytoseiidae)
Author(s): Anna D. Howell and Oleg Daugovish
Source: Journal of Economic Entomology, 106(1):80-85.
Published By: Entomological Society of America
URL: http://www.bioone.org/doi/full/10.1603/EC12304
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2. BIOLOGICAL AND MICROBIAL CONTROL
Biological Control of Eotetranychus lewisi and Tetranychus urticae
(Acari: Tetranychidae) on strawberry by four phytoseiids
(Acari: Phytoseiidae)
ANNA D. HOWELL1
AND OLEG DAUGOVISH
University of California Cooperative Extension, 669 County Square Drive, Suite 100, Ventura, CA 93003
J. Econ. Entomol. 106(1): 80Ð85 (2013); DOI: http://dx.doi.org/10.1603/EC12304
ABSTRACT The spider mite, Eotetranychus lewisi (McGregor) (Acari: Tetranychidae), is a new
emerging pest in California commercial strawberries. The predatory mite Phytoseiulus persimilis
(AthiasÐHenriot) (Acari: Phytoseiidae), typically used for biocontrol of Tetranychus urticae (Koch)
(Acari: Tetranychidae), provided growers little to no control of E. lewisi. Four commonly used
phytoseiidpredatorymites:P.persimilis,Neoseiuluscalifornicus(McGregor),N.fallacis(Garman),and
Amblyseius andersoni (Chant), were used in lab studies to investigate which is best at managing E.
lewisi populations. We also investigated the interactions between T. urticae and E. lewisi and in relation
to phytoseiid efÞciency given the potential for indirect effects of biocontrol. When E. lewisi and T.
urticae are present on the same leaf, T. urticae populations increase and begin displacing E. lewisi. P.
persimilis did not feed on E. lewisi, but the other three predatory mites consumed the spider mites
and lowered their populations. When both E. lewisi and T. urticae are present on the same leaf, N.
fallacis and A. andersoni fed on both types of mites equally and were capable of decreasing both
populations. N. californicus fed on E. lewisi Þrst and decreased its population, but allowed T. urticae
populations to increase. P. persimilis may be insufÞcient at controlling E. lewisi and its use may instead
enhance E. lewisi populations.
KEY WORDS Eotetranychus lewisi, T. urticae, biological control, strawberry, phytoseiids
The spider mite, Eotetranychus lewisi (McGregor)
(Acari: Tetranychidae), has traditionally been con-
sidered a major pest of greenhouse poinsettias and a
minor pest of citrus (McMurtry 1985, Jeppson et al.
1975, Doucette 1962). Pest Control Advisors (PCAs)
along CaliforniaÕs central coast had occasionally ob-
served E. lewisi on strawberries (Fragaria x ananassa
Duchesne), but E. lewisi populations has remained
latent compared with Tetranychus urticae (Koch), the
dominant mite pest of strawberries (Strand 2008; Oat-
man 1971; Oatman 1982, 1981). However, in recent
years E. lewisi populations have increased relative to
T. urticae. A hypothesis to the increase in E. lewisi may
be that the biological and chemical control tools used
to suppress T. urticae may have indirectly released E.
lewisi from competition, thereby allowing their pop-
ulations to increase. Damage to strawberry by E. lewisi
is similar to that of T. urticae: chlorosis and bronzing
of the leaves where feeding occurs, light to high web-
bing and reduction in fruit production at high mite
densities. In coastal California, the recent E. lewisi
outbreaks have caused signiÞcant damage to produc-
tion Þelds, particularly organic Þelds, thereby becom-
ing a problematic pest in commercially cultivated
strawberries, and raspberries.
Regional biological control of T. urticae and other
latent pest mites found on strawberries, such as the
carmine spider mite, Tetranychus cinnabarinus (Bois-
duval), has traditionally relied on the use of the pred-
atory mite Phytoseiulus persimilis (AthiasÐHenriot)
(Acari: Phytoseiidae; Strand 2008, McMurtry 1991).
PCAs and growers noted that while populations of T.
urticae decreased with effective control, E. lewisi pop-
ulations increased. The effectiveness of P. persimilis
on E. lewisi has never been evaluated, yet strawberry
growers released P. persimilis in hopes of controlling
E. lewisi. However, several growers and PCAs have
reported that P. persimilis failed to effectively control
E. lewisi in their Þelds. Additionally, several miticides
that were effective against T. urticae failed to control
E. lewisi (F. Zalom, personal communication).
The use of phytoseiid predators is increasing be-
cause of the quick development of resistance to acar-
icides (Wysoki 1985) and environmental concerns of
pesticide applications. Thus, to better assist growers in
the application of proper biocontrol it is crucial to
understand competitive and predatorÐprey interac-
tions. In the current study we investigated E. lewisi
and four commonly and commercially available
phytoseiid predatory mites used on strawberry: P.
persimilis (AthiasÐHenriot), Neoseiulus californicus
(McGregor), N. fallacis (Garman), and Amblyseius
1 Corresponding author, e-mail: adhowell@ucdavis.edu.
0022-0493/13/0080Ð0085$04.00/0 䉷 2013 Entomological Society of America

3. andersoni (Chant). Similarly, we investigated the in-
teractions between T. urticae and E. lewisi and in
relationtophytoseiidefÞciencygiventhepotentialfor
indirect effects of biocontrol.
Materials and Methods
Mites and Experimental Arena. E. lewisi were col-
lected in 1 March 2011 from organically grown straw-
berry plants at the University of California Hanson
Agricultural Center in Santa Paula, CA. The mites
were placed on clean, mature, fully expanded de-
tached strawberry trifoliates (variety San Andreas)
and reared for at least two generations in a laboratory
in Ventura, CA. Rearing conditions were: 24 ⫾ 1⬚C,
50Ð55% relative humidity (RH) with a photoperiod of
16:8 (L:D) h. T. urticae were provided by Syngenta
Bioline, Oxnard, CA, and reared for at least two gen-
erations on mature, fully expanded, detached straw-
berry trifoliates (variety San Andreas) in the same
laboratory and conditions as those for E. lewisi. The
predatory mites P. persimilis, A. andersoni, N. califor-
nicus, and N. fallacis were provided by several local
insectaries in Ventura County, CA. Predatory mites
were not reared in the laboratory. All strawberry
leaves used for rearing and during monitoring were
collected from organic Þelds and placed in water con-
taining a drop of dish detergent, rinsed several times
with water, and examined under a stereoscope to
assure that no other mites and arthropods were pres-
ent. The leaves were changed as often as needed to
provide adequate forage for each mite species.
The following experimental setup was used for all
monitoring experiments. A clean, whole, mature, fully
expanded detached with an intact petiole strawberry
leaf (variety San Andreas; area ⬇112.3 ⫾ 22.8 cm2
(area of ellipse ⫽ plant length/2 * plant width/2 * ␲))
was placed into a 150 ⫻ 15 mm petri dish (Fisher,
Pittsburg, PA) with double-sided poster tape (Scotch
poster tape; 3M, Scotch, Maplewood, MN) surround-
ing the inside of the dish to prevent escape and outside
immigration. The petiole was placed into a 0.6 ml
polymerase chain reaction (PCR) tube with a hole cut
into the cap and Þlled with water to keep the leaf
hydrated (Fig. 1). Leaves were replaced and tubes
reÞlled with water as needed. Experimental labora-
tory conditions were similar to rearing conditions.
Efficiency of Four Phytoseiid Mites in Controlling
E. lewisi. Forty mobile stages (larvae ⫹ nymphs ⫹
adults combined) of E. lewisi, male and female (un-
known ratio), were carefully transferred using a soft
paintbrush (size 0), and haphazardly placed onto a
wholestrawberryleafandallowedtoestablishfor24h.
Ten phytoseiid mites (adults only; male and female,
unknown ratio) were then introduced onto the leaf. E.
lewisi were counted before the introduction of phyto-
seiids. Phytoseiid mites were haphazardly placed onto
the leaf. All phytoseiid mites were starved for 24 h
before introduction into the experimental arena. Four
days after phytoseiid introduction, all mobile stages
(larvae ⫹ nymphs ⫹ adults combined) and eggs of E.
lewisi and the phytoseiid were counted using a ste-
reomicroscope and clicker counters. Monitoring con-
tinued for 2 wk, with counts every 4 d. The experiment
was replicated four times for each phytoseiid, P. per-
similis, A. andersoni, N. californicus, N. fallacis, and a
control (no predators).
Population Development of E. lewisi and T. urticae
on the Same Leaf. Twenty mobile stages (male and
female, unknown ratio) of E. lewisi and 20 mobile
stages (male and female, unknown ratio) of T. urticae
were haphazardly placed onto a whole clean detached
strawberry leaf. One day after placement, all mobile
stages of each mite species were counted. Counting
continued every fourth day for 2 wk. The experiment
was replicated four times.
Efficacy of Preference of Three Phytoseiid Mites in
Controlling E. lewisi and T. urticae on the Same Leaf.
Twenty mobile stages of E. lewisi and 20 mobile stages
of T. urticae (male and female, unknown ratio) were
Fig. 1. Experimental arena set up with detached strawberry trifoliate (including the intact petiole), surrounded by double
sided poster tape, inside a petri dish. (Online Þgure in color.)
February 2013 HOWELL AND DAUGOVISH: BIOLOGICAL CONTROL OF Eotetranychus lewisi 81

4. haphazardly placed onto a whole clean detached
strawberry leaf and allowed to establish for 24 h. After
24 h of starvation, 10 adult phytoseiid mites (male and
female, unknown ratio) were introduced and haphaz-
ardly placed onto the leaf. E. lewisi and T. urticae were
counted before phytoseiids were introduced into the
experimental arena. Four days after predator intro-
duction,allmobilestagesofE.lewisi,T.urticae,andthe
phytoseiid were counted. The experiment was repli-
cated four times for each predatory mite: A. andersoni,
N. californicus, N. fallacis, and a control (no preda-
tors).
Statistical Analysis. All experiments were designed
as randomized complete block design with four rep-
licates. Data were analyzed using repeated measure
analysis of variance (ANOVA) with time as a repeated
measure (SPSS Inc. 2008) to compare values between
treatments over the sampling period. We used the
GreenhouseÐGeisser correction to adjust degrees of
freedom in any case where data violated the assump-
tion of sphericity or homogeneity of covariance
(Greenhouse and Geisser 1959). Means were sepa-
rated using Tukey honestly signiÞcant difference
(HSD) (P ⬍ 0.05).
Results
P. persimilis would not feed on E. lewisi and either
died of starvation or by running off the leaf onto the
sticky tape. This behavior was observed for two trials
(eight replicates total). Therefore, P. persimilis was
excluded from all analyses.
Efficiency of Three Phytoseiid Mites in Controlling
E. lewisi. The number of E. lewisi mobiles (larvae ⫹
nymphs ⫹ adults) varied throughout the experimental
period (repeated measures ANOVA with Green-
houseÐGeisser correction, F ⫽ 5.28; df ⫽ 1.47, 17.6; P ⫽
0.023; Fig. 2A), with all three phytoseiids, N. califor-
nicus, N. fallacis, and A. andersoni, decreasing E. lewisi
populations compared with controls (repeated mea-
sures ANOVA; F ⫽ 13.14; df ⫽ 3, 12; P ⬍ 0.0001). E.
lewisi eggs also varied throughout the experiment (re-
peated measures ANOVA with GreenhouseÐGeisser
correction, F ⫽ 2.98; df ⫽ 5.28, 21.12; P ⫽ 0.033; Fig.
2B). All three phytoseiids consumed E. lewisi eggs and
decreased their numbers compared with the controls
(repeated measures ANOVA; F ⫽ 6.82; df ⫽ 3, 12; P ⬍
0.006). E. lewisi mobiles and eggs slightly increased for
theÞrst4daftertheintroductionofN.californicus,but
this phytoseiid was able to decrease both eggs and
mobile stages to nearly zero by the 16th day (Fig. 2).
N. fallacis showed the same predatory pattern as N.
californicus (Fig. 2). A. andersoni Þrst began feeding
on E. lewisi mobiles decreasing their populations and
then shifted to eggs (Fig. 2). It took A. andersoni
longer, compared with the other phytoseiids, but it
eventually lowered the number of E. lewisi eggs, but
did not eliminate E. lewisi mobiles. When E. lewisi was
free from predation the population rapidly increased
in the control. By the end of the sampling period, the
Fig. 2. Population trends of E. lewisi. (A) Mobiles (larvae ⫹ nymphs ⫹ adults) and (B) eggs developing on a detached
strawberry leaf with the addition of the following predatory mites: N. californicus (Nc), N. fallacis (Nf) A. andersoni (Aa),
and no predator (control). Data are presented as means of four replicates ⫾ SE.
82 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 106, no. 1

5. population had grown to ⬇1,000 mobiles and 830 eggs
and caused signiÞcant leaf deterioration (Fig. 2).
Population Dynamics of E. lewisi and T. urticae on
the Same Leaf. The populations of both mites did not
begin to increase until the fourth day from the begin-
ning of the experiment (Fig. 3). However, by the
eighth day T. urticae populations were 1.5 times
greater than E. lewisi, and by the 12th day T. uticae
populations were 3.5 times greater than E. lewisi pop-
ulations. After the 12th day, T. urticae populations
continued to increase while E. lewisi struggled to grow
and began to decrease (Fig. 3). By the end of the study
T. uticae populations had surpassed 400 mobiles, while
E. lewisi had dropped down to ⬇60 mobiles (mean
total).
Three Phytoseiids Controlling E. lewisi and T. ur-
ticae on the Same Leaf. Populations of E. lewisi varied
across the experimental period (repeated measures
ANOVA with GreenhouseÐGeisser correction, F ⫽
4.17; df ⫽ 8.04, 32.14; P ⫽ 0.002; Fig. 4). All three
phytoseiids decreased E. lewisi populations substan-
tially compared with the controls (repeated measures
ANOVA; F ⫽ 15.34; df ⫽ 3, 12; P ⬍ 0.0001). T. urticae
populations also varied with each treatment across the
experimental period (repeated measures ANOVA
with GreenhouseÐGeisser correction, F ⫽ 10.72; df ⫽
4.18, 16.71; P ⬍ 0.0001; Fig. 4). All three phytoseiids
decreased T. urticae populations compared with the
control (repeated measures ANOVA; F ⫽ 83.99; df ⫽
3, 12; P ⬍ 0.0001). However, N. californicus’ predation
efÞciency was signiÞcantly different from N. fallacis
and A. andersoni (Tukey HSD; P ⬍ 0.001). N. fallacis
and A. andersoni decreased T. urticae populations to
almostzerobythe12thday(Fig.4A,B).N.californicus,
however, Þrst fed on E. lewisi allowing T. urticae to
rapidly increase their populations (Fig. 3C). By the
12th day, T. urticae populations had increased to over
300 mobiles in that treatment. By the end of the ex-
periment N. californicus had decreased T. urticae pop-
ulations to just below 180 mobiles, but E. lewisi pop-
ulations began to increase.
Discussion
Our experiments showed that three commercially
reared phytoseiids, N. fallacis, N. californicus, and A.
andersoni were able to control E. lewisi populations,
but the commonly used phytoseiid P. persimilis would
not feed on E. lewisi. N. californicus and N. fallacis
suppressed E. lewisi populations rather quickly,
whereas A. andersoni initially faltered, but was able to
suppress mite populations compared with the con-
trols. We also found that control efÞciency can change
depending on whether E. lewisi is populating an area
alone or in a mixed population with T. urticae. N.
californicus’ predation efÞciency differed from the
other phytoseiids in mixed populations of E. lewisi and
T. urticae. This phytoseiid appeared to initially predate
onE.lewisi,allowingT.urticaepopulationstoincrease.
Previous studies have shown N. californicus to be an
effective suppressor of T. urticae (Oatman et al. 1977,
Rhodes et al. 2006, Fraulo and Liburd 2007, Ahn et al.
2010) and even showing preference for T. urticae over
other spider mite species (Moraes and McMurtry
1985, Easterbrook et al. 2001, Escudero and Ferragut
2005). This type of variation in response has been
observed and attributed to factors such as previous
exposure to prey (Carillo et al. 2012) and the ability to
develop adequately on speciÞc prey (Moraes and Mc-
Murtry 1985). Perhaps the population of N. californi-
cus we used was less adapted to feeding on T. urticae
or found E. lewisi to be a more suitable prey than T.
urticae. Similarly, Moraes and McMurtry (1985) found
that different populations of P. persimilis varied in
their survival and oviposition rates when consuming T.
urticae. The reasons as to why P. persimilis would not
feed on E. lewisi were not evaluated in the current
study, but may be because of the nutritional quality of
E. lewisi or perhaps the phytoseiids type I specialized
life-style (McMurtry and Croft 1997). Future exper-
iments would be required to address the nutritional
aspect.
The observed competitive interactions between T.
urticae and E. lewisi may explain why E. lewisi popu-
lations have been increasing in strawberry Þelds in
Ventura Co. When E. lewisi and T. urticae are occu-
pying the same resource, T. urticae outcompetes and
begins to displace E. lewisi. However, if P. persimilis is
insufÞcient at controlling E. lewisi, it can release E.
lewisi from competition allowing the mite to prolifer-
ate and dominate. A similar situation was observed in
greenhouse strawberries in western Japan with T. ur-
Fig.3. PopulationsofE.lewisiandT.urticaedevelopingonthesamestrawberryleafwithoutpredation.Dataarepresented
as means of four replicates ⫾ SE.
February 2013 HOWELL AND DAUGOVISH: BIOLOGICAL CONTROL OF Eotetranychus lewisi 83

6. ticae and Eotetranychus asiaticus (Ehara) (Acari: Tet-
ranychidae). Osakabe (2002) found that the release of
P. persimilis to control T. urticae enhanced the occur-
rence of E. asiaticus, allowing it to become a dominant
pest on greenhouse strawberry.
This study conÞrms that the release of the common
predatory mite P. persimilis may be insufÞcient at
controlling E. lewisi and may be enhancing its popu-
lation growth. Little research has been conducted on
the biocontrol of E. lewisi on strawberry, but here we
identiÞed three commercially available phytoseiids
capable of controlling E. lewisi and two phytoseiids
capable of controlling E. lewisi and T. urticae simul-
taneously. Additionally, the study demonstrates how
phytoseiid predatory behavior can change when pre-
sented with a mixed population of E. lewisi and T.
urticae. These Þndings, along with ongoing miticide
bioassays, are essential components of a successful
integrated pest management (IPM) program in Cali-
fornia strawberry production.
Acknowledgments
We thank R. Alarcón and two anonymous reviewers for
helpfulcommentsonanearlierversion,andSyngentaBioline
for T. urticae, P. persimilis, and A. andersoni donations.
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