Experimental and Applied Acarology 25: 2536, 2001.

2001 Kluwer Academic Publishers. Printed in the Netherlands.

Biological control of strawberry tarsonemid mite

Phytonemus pallidus and two-spotted spider mite
Tetranychus urticae on strawberry in the UK using
species of Neoseiulus (Amblyseius) (Acari:
Phytoseiidae)
M.A. EASTERBROOK, J.D. FITZGERALD and M.G. SOLOMON
Horticulture Research International, East Malling, West Malling, Kent, ME19 6BJ, UK
(Received 12 September 2000; accepted 2 February 2001)

Abstract. Two species of Neoseiulus, N. californicus and N. cucumeris, showed potential

for biocontrol of phytophagous mites on strawberry. N. californicus controlled Tetranychus
urticae on potted strawberry plants in a gauze-sided glasshouse at temperatures comparable
to early summer in the UK (820 C). Both species of phytoseiid reduced numbers of the
tarsonemid Phytonemus pallidus on potted strawberry plants under glasshouse conditions (15
23 C). In several experiments reductions in the range of 7181% in numbers of tarsonemid
active stages and eggs, compared to non-release plants, were obtained. The importance of
establishing a suitable predator: prey ratio at an early stage was demonstrated in an experiment
where an initial ratio of 1 N. cucumeris: 10 P. pallidus gave a greater degree of control than
1:20 or 1:40.
Key words: Neoseiulus californicus, Neoseiulus cucumeris, Tetranychus urticae, Phytonemus
pallidus, strawberry, biological control

Introduction
The strawberry tarsonemid mite, sometimes known as strawberry mite,
Phytonemus (Steneotarsonemus) pallidus ssp. fragariae (Zimmerman), and
two-spotted spider mite, Tetranychus urticae Koch, are important pests of
strawberry in the UK. Both species can cause reductions in yield when their
population densities are high, and tarsonemid mite can damage the fruit
(Alford 1972; Stenseth and Nordby, 1976). Problems with these two pests are
becoming increasingly serious as more crops of strawberries are grown under
protection, mainly under polyethylene tunnels, but also under glass. Chemical
Author for correspondence: Tel.: + 44 (0) 1732 843833; Fax: + 44 (0) 1732 849067;
E-mail: jean.fitzgerald@hri.ac.uk

26
control is difficult, due to the restricted number of approved pesticides and,
in the case of T. urticae, resistance to many acaricides. Additionally, there are
many pressures to reduce pesticide use.
It is desirable to control these phytophagous mites by non-chemical means.
Many growers already use the exotic predator Phytoseiulus persimilis AthiasHenriot, which was shown to control T. urticae successfully (Easterbrook,
1992). However, this predator has to be re-introduced each year, as it rarely
survives the winter in the UK, and it is susceptible to many of the insecticides
used to control other pests. Also, it does not feed on P. pallidus (Easterbrook, unpublished data), so alternative predators are needed. Neoseiulus
(Amblyseius) californicus (McGregor), although not considered a native of
the UK, has been found in large numbers in some strawberry fields in southeast England (Easterbrook, 1998 and unpublished; Jolly, 2000), possibly as a
result of earlier releases. In laboratory experiments N. californicus was found
to survive doses of pyrethroid insecticides at several times field rate (Easterbrook and Fitzgerald, unpublished). Neoseiulus (Amblyseius) cucumeris
(Oudemans) is native to the UK (Chant, 1959), and has been available from
biological control supply companies for some time; N. californicus has also
become available recently. A series of experiments was done to examine
the suitability of these predatory phytoseiids to control tarsonemid and twospotted spider mites.
Materials and Methods
In 1997 an experiment was conducted to assess the potential of N. californicus
to control T. urticae, in a compartment of a glasshouse with a glass roof and
sides of nylon mesh. Temperatures were recorded with a Tinytag electronic
datalogger (Gemini Datalogger, Chichester); daily mean temperatures ranged
from 8 to 20 C during the course of the experiment (AprilJune), and on
most days were 14 C above ambient, as measured in a Stevenson screen
100 metres distant.
Thirty strawberry plants cv. Calypso, growing in 2 l pots, were infested
with T. urticae by placing 10 adult females on each plant on 26 March; the
mites were collected from a strawberry field, where they had started to lay
eggs. On 3 April, the plants were divided at random into three groups; one
group received four adult female N. californicus per plant, the second group
two N. californicus per plant, and the third did not receive any predators.
The phytoseiids were collected from a strawberry field at HRI-East Malling
and were placed on mature leaves of the plants using a fine paintbrush; at
this time the plants had an average of 5.5 leaves (range 39). Samples of
phytoseiids were mounted on microscope slides and examined to confirm

27
that they were N. californicus. Each of the 10 replicate blocks consisted
of three potted plants (one of each treatment) standing on inverted saucers
surrounded by water in a 100 40 cm plastic tray, to minimise movement
of predators between plants. The plants were sampled non-destructively at 7
days after predator release (DAR), then at 1618 day intervals until 30 May,
by examining all leaves on each plant under a stereomicroscope mounted on
a stand and counting all the mites present. For the final sample on 26 June all
leaves were removed from the plants for examination.
In 1997 an initial experiment was done to determine whether N. californicus could reduce numbers of tarsonemid mite. Twelve young, partially
unfurled leaves of strawberry, cv. Tango, were cut off potted plants and kept
individually with the petiole inserted through a hole in the plastic lid of a
small pot (4.5 cm diameter) containing water. Twenty adult female P. pallidus
collected from the field were placed on each leaf, using a fine paintbrush, on
12 September. The leaves were kept in large plastic boxes (28 16 9.5 cm)
at 15 C. On 18 September, each leaf had a ring of OecoTAK (Oecos, UK)
glue put around the petiole and was kept individually in a small plastic box
(13.5 7.5 6 cm) with gauze inserts in the side to provide ventilation. One
gravid female N. californicus (ex Novartis BCM) was added to half of the
leaves and the boxes were placed in a random arrangement in a 15 C controlled environment cabinet, with 16 h light: 8 h dark period. Mites were counted on the leaves under a stereomicroscope on 24 September (6 DAR) and 1
October (13 DAR).
In 1998 three experiments were done to investigate biological control of
tarsonemid mite by species of Neoseiulus. In experiment 1998A 30 strawberry plants cv. Evita, growing in 2 l pots and showing symptoms of severe
attack by tarsonemid mite, were divided at random into three groups of 10
plants on 20 February. The three groups were placed into adjacent compartments of a glasshouse with heating and with supplementary lighting to
provide 16 h light: 8 h dark. Temperatures in the three compartments were
monitored using Tinytag dataloggers. During the course of the experiment,
temperatures ranged from 17 to 23 C, and were similar in all compartments.
On 20 February each plant in one group received 50 adult N. californicus;
another group had 50 N. cucumeris per plant, while the third group did not
receive any predators. The predatory mites, obtained from Novartis BCM,
were distributed on several mature leaves on each plant, using a fine paintbrush. On 18 March (26 DAR), five plants in each treatment were sampled by
cutting off all leaves and flower clusters, dissecting this material and counting
all active mites and eggs under a stereomicroscope. At this time, most plants
had 1721 leaves and several clusters of flowers/young fruits. The remaining

28
five plants in each treatment were sampled by the same method on 14 April
(53 DAR). Plant material was stored at 4 C until examined.
Experiment 1998B used strawberry plants cv. Elsanta growing in 2 l pots.
Twenty-three plants were inoculated with 20 field-collected adult female tarsonemid mites per plant between 1 and 7 July. On 7 August, the plants were
split at random into three groups, then eight plants received 30 adult N. californicus (ex Novartis BCM) per plant, eight had 30 N. cucumeris (ex Biological Crop Protection Ltd.) per plant, while the remaining seven did not
receive any predators. On this date the plants had 814 leaves (mean 11 per
plant), and there was evidence of tarsonemid mite damage on young leaves.
The plants were sampled on 4 September (26 DAR) by removing all the
leaves from each plant and counting mites under a stereomicroscope.
In experiment 1998C the plants used were cv. Tango, potted as runners
into 4 inch disposable pots a few weeks previously, and at the start of the
experiment they had 1014 leaves. On 5 March the plants were divided into
three groups of 12 and inoculated with either 20, 40 or 80 adult female
tarsonemid mites, placed on the youngest newly expanded leaf with a fine
paintbrush. Each group of plants was placed in a separate compartment of
a glasshouse with heating and with supplementary lighting to provide 16 h
light: 8 h dark. Temperatures, monitored by Tinytag dataloggers, were in the
range 1722 C during the course of the experiment, and were similar in the
three compartments. On 12 March two gravid adult female N. cucumeris (ex
Novartis BCM) were added to each plant, so providing nominal predator: prey
ratios of 1:10, 1:20 and 1:40 (not including any tarsonemid eggs laid in the
intervening period). Half of the plants from each group were sampled at 14
DAR by removing all leaves and flower clusters and counting mites on all the
plant material under a stereomicroscope; the remaining plants were sampled
at 28 DAR.
A glasshouse experiment in 1999 compared two release rates of N. cucumeris to control tarsonemids and T. urticae. Twenty-four strawberry plants,
cv. Evita, potted as runners two months earlier into 1 l pots, that had become naturally infested with T. urticae, were infested with 20 adult female
tarsonemids per plant from 7 to 12 September. On 1 October, the plants
were put at random into three groups of eight, and received either 20 gravid
adult female N. cucumeris (ex Biological Crop Protection Ltd) per plant, four
N. cucumeris per plant, or no predators. Each group of plants was then placed
in a separate compartment of a glasshouse with heating and supplementary
lighting (16 h light: 8 h dark). Temperatures during this experiment ranged
from 15 to 20 C. On 1 October the plants had a mean of 10.2 leaves per plant
(range 814). On 8 November all aerial growth was removed from each plant
and stored in plastic bags at 4 C until dissection under a stereomicroscope,

29
when all the mites present were counted. At this time most plants had 1316
leaves and 35 clusters with flowers and developing fruit.
In 1999 an unreplicated experiment was done on strawberry cvs EM639
and EM726, growing in peat-filled bags (five plants per bag) on top of straw
bales in a plastic tunnel. These plants had become naturally infested with
P. pallidus. On 30 June, N. cucumeris (ex Biological Crop Protection Ltd)
were released onto most of the plants in the tunnel by sprinkling the inert
carrier containing the mites onto the plants. Each plant received about 200
N. cucumeris. Two bags (10 plants) at the end of each row were left untreated;
there was a gap of 0.3 m between these plants and the nearest predator-release
plants. On 30 June (pre-release), 14 July (14 DAR) and 30 July (30 DAR)
samples of 20 young trifoliate leaves (with the upper surfaces of each leaflet
still adpressed) were taken from the non-release area (10 leaves per cv.), and
also from the 10 plants in each row nearest to the non-release area. Mites on
these leaves were counted under a stereomicroscope.
In all the experiments, counts of mites were log transformed and analysed
by ANOVA.
Results
Both release rates of N. californicus used in the 1997 experiment reduced
numbers of T. urticae, but control was better from the higher rate, particularly
during May, when there was a rapid increase in spider mite numbers on the
plants without predators (Figure 1A, B). Numbers of N. californicus were
highest on the plants that received the highest release rate until mid May
(Figure 1C). By late May N. californicus had dispersed to the non-release
plants, and reduced T. urticae numbers on those too in June.
In the detached leaf experiment in 1997 there were no significant differences in numbers of tarsonemids between leaves with or without a N. californicus female at 6 DAR. However, at 13 DAR there were significantly
fewer tarsonemid active stages on leaves with a predatory mite (mean SE of
19.4 10.3 per leaf) than on those without a predator (44.4 10.3 per leaf)
(on transformed data F1,8 = 5.23, P < 0.05). Differences between numbers of
eggs (14.0 per leaf with predator, 24.6 without) were not significant.
In experiment 1998A both species of Neoseiulus reduced the very high
number of tarsonemid mites present (Table 1). At the conclusion of the experiment, the maximum reduction compared to plants with no predators was
76% for tarsonemid active stages and 82% for eggs. At 26 DAR there was no
significant difference between numbers of tarsonemids on plants with N. cucumeris or N. californicus, but numbers of tarsonemid eggs were significantly
lower on plants with N. cucumeris at 53 DAR.

30

Figure 1. Control of Tetranychus urticae on potted strawberry plants in a gauzehouse by

release of Neoseiulus californicus, 1997. Vertical bars indicate least significant difference;
P = 0.05. Arrows indicate date of predator release; 0 N. californicus/plant; 2
N. californicus/plant; H 4 N. californicus/plant.

31
Table 1. Comparison of Neoseiulus cucumeris and N. californicus for control of tarsonemid
mite, Phytonemus pallidus, on potted strawberry plants
Treatment

Mean numbers of tarsonemid mites per plant (N)

26 DAR
53 DAR
Active stages
Eggs
Active stages
Eggs
N
loge N N
loge N N
loge N N
loge N

Experiment 1998A
No predators
1078.2
6.94 1029.6
N. cucumeris
436.4
5.62
556.2
N. californicus
280.4
5.56
348.2
SED (12df)
0.445
P -value
<0.05
Experiment 1998B
No predators
N. cucumeris
N. californicus
SED (15df)
P -value

229.8
49.3
30.2

5.24
3.59
2.92
0.628
<0.01

479.2
138.3
104.2

6.75 706.0
6.51 755.6
6.50
5.83 166.4
4.98 136.2
4.77
5.75 354.4
5.67 504.6
5.94
0.553
0.365
0.435
ns
<0.01
<0.01

6.08
4.60
3.94
0.613
<0.01

DAR = days after predator release.

In experiment 1998B, where numbers of tarsonemid mites were much

lower than in experiment 1998A, both Neoseiulus species reduced numbers
of tarsonemids by 7887% for actives and 7178% for eggs at 26 DAR,
compared to non-release plants (Table 1). At the conclusion of the experiment (26DAR) there were no significant differences between the numbers of
N. cucumeris and N. californicus, mean number of active stages per plant
being 14.7 and 13.3 on the two treatments, respectively.
When different initial predator: prey ratios of N. cucumeris: P. pallidus
were compared in experiment 1998C, control of the tarsonemid was significantly better at 1:10 and 1:20 than at 1:40 at 28 DAR (Table 2). Numbers
of tarsonemid eggs were significantly lower on the plants with 1:10 predator:
prey ratio than at the higher ratios on that date. Numbers of N. cucumeris
were not significantly different between treatments on either sampling date,
and ranged from 2.1 to 4.9 active stages per plant.
In the 1999 glasshouse experiment the higher rate of release of N. cucumeris gave significantly greater reductions of numbers of both active stages
and eggs of P. pallidus than did the lower rate (Table 3). Numbers of active
stages were reduced by 81% and eggs by 80% at the higher release rate.
Numbers of T. urticae eggs were reduced significantly at the higher release

32
Table 2. Experiment 1998C: Control of Phytonemus pallidus by Neoseiulus cucumeris
at different predator: prey ratios
Initial
predator:
prey ratio

Mean numbers of tarsonemids per plant (N)

14 DAR
28 DAR
Active stages
Eggs
Active stages
Eggs
N
loge N
N
loge N N
loge N
N
loge N

1:10
1:20
1:40
SED(15df)
P -value

2.3
1.7
9.3

1.07
0.74
1.98
0.456
< 0.05

2.2
2.3
2.8

0.54
0.68
0.94
0.586
ns

7.3
10.5
41.3

1.64
2.37
3.51
0.473
< 0.01

5.8
18.0
95.7

1.50
2.77
4.22
0.514
< 0.001

DAR = days after predator release.

Table 3. Control of Phytonemus pallidus and Tetranychus urticae on potted strawberry

plants at different release rates of Neoseiulus cucumeris (1999)
N.
cucumeris
release
rate/plant

Numbers of mites per plant (N) at 38 DAR

Phytonemus pallidus
Tetranychus urticae
Active stages
Eggs
Active stages
Eggs
N
loge
N
loge
N
loge
N
loge
(N + 1)
(N + 1)
(N + 1)
(N + 1)

0
4
20
SED
(21df)
P -value

169.0
61.8
31.4

4.99
4.09
3.23
0.333
< 0.001

64.9
39.0
12.8

3.85
3.36
1.91
0.558
0.006

617.1
1089.1
251.2

5.72
6.64
4.63
0.645
0.018

1761.6
1604.3
548.3

7.11
6.97
5.68
0.520
0.022

rate, though the difference in numbers of active stages was not significant.
The lower release rate of N. cucumeris had no significant effect on T. urticae
numbers. At the end of the experiment (38 DAR) there was no significant
difference between the number of N. cucumeris on the low and high release
rates (6.7 and 5.5 active stages per plant, respectively). Numbers on the high
release rate plants may have started to decline as numbers of prey diminished.
Mean numbers of N. cucumeris on non-release plants at this time were 0.8
active stages per plant, though five out of eight plants were free of predatory
mites.

33
In the non-replicated experiment on tunnel-grown plants in 1999, numbers
of tarsonemid mites in the N. cucumeris release area declined from 188 active
stages per 20 leaves pre-release to 55 and 10 per 20 leaves at 14 DAR and 30
DAR, respectively. In the non-release area there were 313 per 20 leaves pretreatment, 240 at 14 DAR and 186 at 30 DAR, so the reduction in numbers
was much greater in the predator-release zone. The probable reasons for the
drop in tarsonemid numbers in the non-release area are that plant quality
declined during this period and this area was gradually colonised by predatory mites. At 30 DAR there were 17 tarsonemid eggs per 20 leaves in the
predator-release area, compared with 489 per 20 leaves in the no-predator
area.
In all the experiments most tarsonemid mites were found on young leaves
where the lamina had not expanded and the adaxial surfaces of each leaflet
were still pressed together, under bracts and bracteoles on flower stalks, at the
base of young fruits where the sepals joined the receptacle, and at the base of
leaf and flower stalks in the crown of the plant.
Discussion
The experiment in 1997 on potted strawberry plants, at temperatures representative of early summer in southern England, showed that N. californicus
was capable of exerting control of T. urticae. However, it is important that
suitable predator: prey ratios are established if rapid control is to be achieved.
In this experiment T. urticae numbers were reduced by N. californicus when
predator: prey ratios were in the range 1: 8 to 1: 11. These are similar ratios
to those reported for successful control of T. urticae by N. californicus in the
field in Spain by Garcia-Mari and Gonzalez-Zamora (1999). They found that
at spider mite levels of above 1 per leaflet, a ratio of 1 N. californicus per
510 T. urticae resulted in a decline in the prey population in the following
sample, taken 12 weeks later.
Greco et al. (1999) found that on strawberries grown in glasshouses in
Argentina T. urticae usually declined greatly after one peak in population
density when N. californicus was present. The high spatial coincidence of
N. californicus with T. urticae found in their study suggested a significant
dispersal capacity of the predator, as well as a high ability to detect leaves
with prey, even at low spider mite density. Similar results were found in
individual leaf counts in our experiment.
Neoseiulus californicus reduced numbers of T. urticae in field releases in
southern California (Oatman et al., 1977) and can delay rapid increase of
this species there (Zalom et al., 1990), and an Italian strain of N. californicus demonstrated a good capacity to control T. urticae in laboratory studies

34
(Castagnoli and Amato, 1991). In Belgium, N. californicus gave better control of T. urticae on strawberry than did Phytoseiulus persimilis, both under
plastic tunnels and outdoors (Anonymous, 1998). Other species of Neoseiulus
(Amblyseius) have also given successful biological control of T. urticae. Neoseiulus fallacis (Garman) has been used successfully in north-east USA and
eastern Canada (Raworth, 1990; Coop and Croft, 1995; Cooley et al., 1996)
and N. womersleyi (Schicha) in Australia (Waite, 1988).
Control of P. pallidus was similar for the two species of Neoseiulus. Neither species of predator eliminated the tarsonemid. Part of the reason for
this is probably that the tarsonemids occupy extremely cryptic microhabitats,
which the larger life stages of the phytoseiids find difficulty in accessing.
Also, neither Neoseiulus species consumed tarsonemid eggs when presented
with them in laboratory feeding tests, although they developed successfully
to adult on other tarsonemid life stages (Easterbrook, unpublished data).
The experiment with different initial N. cucumeris: P. pallidus ratios showed that it is important to release the predatory mites before tarsonemid
populations get too large, if successful regulation is to be achieved. Otherwise, it may be necessary to release very large numbers of predators per
plant to achieve control, as in the unreplicated experiment. This will often be
uneconomic, unless only small numbers of plants are affected.
McMurtry and Croft (1997) categorised phytoseiid mites into four broad
divisions, according to the level of specialization in their prey preferences.
When Croft et al. (1998) compared the generalist predator N. cucumeris
with a more specialised predator of spider mites, N. fallacis, for control of
P. pallidus, they obtained better control with N. fallacis over the time period
of their experiments (48 weeks). However, they speculated that, if the period
had been longer, regulation of tarsonemids might have been better from N. cucumeris. This is because generalist phytoseiids are better regulators of pest
mites, whereas specialists are better at numerically responding to, and controlling, high-density populations (McMurtry and Croft, 1997). Neoseiulus
californicus, like N. fallacis, is a specialised consumer of spider mites
(McMurtry and Croft, 1997). The 1999 glasshouse experiment suggested that
when P. pallidus and T. urticae are both present, N. cucumeris will give
reasonable control of the tarsonemid, but not of the spider mite. In these
circumstances, a mixture of N. cucumeris and N. californicus may provide
control of both phytophagous mites, but further research is required to study
the interactions between the two phytoseiid species.

35
Acknowledgements
We thank Catherine Pinch for technical assistance and Martin Ridout for
statistical analyses. The research was funded by the Ministry of Agriculture,
Fisheries and Food, Horticulture Division.

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