JEN 128(9/10) doi: 10.1111/j.1439-0418.2004.00887.

569–575

Ó 2004 Blackwell Verlag, Berlin

Colonization of growing fruit by the armored scales Lepidosaphes

beckii, Parlatoria pergandii and Aonidiella aurantii (Hom.,
Diaspididae)
Ma E. Rodrigo, F. Garcia-Marı́, J. M. Rodrı́guez-Reina and T. Olmeda
Institut Agroforestal Mediterrani, Entomologı́a–ETSEA, Universitat Politècnica de Valencia, València, Spain

Ms. received: February 27, 2004; accepted: June 11, 2004

Abstract: The process of invasion of the fruit surface by three species of armored scale insects [Lepidosaphes beckii
(Newman), Parlatoria pergandii Comstock and Aonidiella aurantii (Maskell)], was studied in citrus orchards in
València, Spain. Three periods of intense crawler migration were detected annually for L. beckii and P. pergandii.
Abundance of crawlers increased throughout the year and they moved outwards in the tree canopy. The fruit was
continuously invaded during the period of fruit growth, starting at a very early stage of fruit development, by crawlers
of the first generation of all the three species. Differences in fruit invasion and settling behaviour were found between
the three species. First-generation crawlers tended to settle under the fruit calyx in L. beckii and P. pergandii, and on the
exposed fruit surface in A. aurantii. During the second generation, most A. aurantii scales originated from reproducing
females already settled on the fruit, whereas in L. beckii and P. pergandii new scales came preferentially from other tree
substrates or from females settled under the fruit calyx. These findings suggest changes to optimize the timing of
chemical sprays for controlling the scales.

Key words: Citrus, crawlers, diaspidids, fruit, invasion

1 Introduction armored scales like L. beckii and P. pergandii may

settle beneath the calyx when the infestation of fruit
Armored scales (Hom., Diaspididae) are important
begins. The calyx seems to protect scales from adverse
pests of citrus all over the world, causing reduction of
climatic conditions and natural enemies. The fruit
tree health and production, and severe downgrading of
surface may be infested by insects coming out or
fresh marketed fruit at the packinghouse (Quayle,
emerging from under the calyx (Bodenheimer, 1951;
1941; Ebeling, 1959; Talhouk, 1975; University of
Carnegie, 1957; Gerson, 1967).
California, 1991a). In Spain, the main world exporter
It is important to know the manner and the moment
of fresh citrus fruit, three diaspidid species require
when diaspidids infest growing fruit in order to apply
regular applications of pesticides, Lepidosaphes beckii
measures of control, especially the succesful applica-
(Newman), Parlatoria pergandii Comstock and Aonid-
tion of chemical treatments (Walker et al., 1990). The
iella aurantii (Maskell) (Moner, 1992; Francés et al.,
objectives of this study were to investigate the move-
1998; Costa-Comelles et al., 1999). The three species
ment of crawlers in the tree, analyse the process of
develop two to four generations per year in the
infestation of fruit by successive crawler generations
Mediterranean area and in California (Benassy and
and study the importance of the calyx in the infestation
Soria, 1964; Abbasi, 1975; Moreno and Kennett, 1985;
process in the three species of armored scales, L. beckii,
Hafez et al., 1987; Walker et al., 1990; Rodrigo and
P. pergandii and A. aurantii.
Garcı́a-Marı́, 1992). Fruit of oranges [Citrus sinensis
(L.) Osbeck] and clementines [Citrus reticulata
(Blanco)] start to grow in spring and reach their final
size in autumn, being invaded by successive broods of 2 Materials and Methods
scale crawlers that develop during this period. The
The present study was conducted in València, Spain in four
second annual crawler generation which takes place in groves (Xest1, Xest2, Sueca and València) during 1995 and
the summer is generally considered to be mainly 1996, and at six groves (Catarroja1, Catarroja2, Catarroja3,
responsible for the infestation of fruit (Smirnoff, Picassent, Sueca and Riola) in 2001. All 10 groves consisted
1960; Benassy et al., 1980; Orphanides, 1982; Alexan- of 12–15-year-old navel orange commercial plantations. No
drakis, 1983; Atkinson, 1983; Moreno and Kennett, insecticides were applied to the groves during the sampling
1985). It has been observed that certain species of period and for at least 2 years prior to the present study. All
570
groves supported medium to heavy infestations of L. beckii,
P. pergandii and/or A. aurantii. In the four groves sampled in
1995 and 1996, the seasonal trend in the proportion of
different stages was monitored by sampling infested leaves
every 2 weeks and recording the number of different stages of
100 live scales found in a minimum of 10 leaves.
2.1 Movement of L. beckii and P. pergandii crawlers
in the tree
The movement of L. beckii and P. pergandii crawlers in trees
was studied in 1995 at the València grove. The study started
in April, once the first flush of growth had been completed.
Sticky tape traps were wrapped around infested twigs,
following the method used in California to detect crawler
generations of A. aurantii (University of California, 1991a).
Fifteen traps were placed on each of two study trees. The
traps were wrapped around green twigs, branches and wood,
five on every one. Green twigs (0–1 year old) corresponded to
the last flush of the previous year. Branches (1–2 years old)
corresponded to a more interior region of the tree, were
partly lignified (grey and green wood) and measured between
0.5 and 1 cm in diameter. Wood (2–3 years old) was the most
interior branches of the tree, completely lignified and
measuring 2–5 cm in diameter. Traps were replaced every
10 days from April to October.
Traps consisted of a base of black friction tape (tesaÒ, tesa
AG, Hamburg, Germany) tightly wrapped around branches
upon which transparent tape sticky on both sides (tesaÒ,
Beiersdorf AG, Hamburg, Germany) was placed to capture
crawlers. Transparent tapes were collected and mounted on
glass slides for counting crawlers under a stereomicroscope.
The identification of the two scale species was based on the
form and colour of crawlers, oval–elongated, pale yellow in
colour in L. beckii and oval, short, burgundy in colour in
P. pergandii (Avidov and Harpaz, 1969).
2.2 Infestation of fruit
The process of infestation of fruit by L. beckii, P. pergandii
and A. aurantii was monitored in three different groves
(Xest1, Xest2 and Sueca) in 1995 and 1996. Ten trees were
selected in each grove, 20 fruit were marked in each tree and
the fruit were examined visually every week. We recorded the
number of scales of the three species found on each fruit
between May and October, except for A. aurantii in Sueca,
for which sampling was started in June.
2.3 Role of the calyx
To determine the role of the calyx in the process of
infestation, samples of 100 fruit, with 10 fruit per tree from
10 trees, were collected. The València and Sueca groves were
sampled every 2 weeks between June and October 1995. A
single sample was taken in June from the six groves in 2001.
The fruit were taken to the laboratory, each fruit was
inspected beneath a stereomicroscope and the number of
scales (L. beckii, P. pergandii and A. aurantii) found under
the calyx and in the rest of the fruit was noted for each fruit
separately.
2.4 Exclusion technique
In order to establish where the scales infesting the fruit in the
second generation originate from, an exclusion technique was
used at the Sueca grove in July 1996. Before the second
annual flush of crawlers, 27 trees were selected and 12 fruit
Ó 2004 Blackwell Verlag, Berlin, JEN 128(9/10) doi: 10.1111/j.1439-0418.2004.00887.569–575
Ma E. Rodrigo et al.
from each tree, in four groups of three, were marked. The
three fruit in each group were situated in the same branch.
The four groups of each tree were selected from the four
quadrants of the tree canopy. If necessary, leaves or branches
situated near the fruit were eliminated so that the marked
fruit were not in contact with them. We placed a 5-mm-wide
ring of insect-trapping adhesive (TanglefootÒ, The Tangle-
foot Company, Grand Rapids, MI) around the peduncle of
one of the fruit in each group, a second fruit had adhesive
over the calyx to seal it completely, and the third one with no
adhesive acted as control. A total of 324 fruit were marked.
In December 1996, fruit were picked, carried to the labor-
atory and inspected under a stereomicroscope, recording
numbers of scales found on each fruit. For subsequent
analyses, only groups with the complete set of three fruit
were considered, rejecting those groups in which any fruit
had dropped.
2.5 Data analysis
SAS/STAT software (SAS Institute, 1989) was used for the
statistical procedures. One-way analysis of variance (anova)
and Fisher’s protected least significant difference (LSD) tests
were applied to the comparison in number of crawlers
between types of branches and between generations. Paired
t-tests were performed for comparing settlement preferences
in first-generation crawlers. Percentages of infested fruit were
arcsine square root-transformed before the analysis. The
effect of exclusion with adhesive barriers was analysed by
one-way anova followed by the LSD procedure.
3 Results
3.1 Movement of L. beckii and P. pergandii crawlers in
the tree
Approximately 2000 crawlers of each species (L. beckii
and P. pergandii) were caught on the sticky tape traps
during the sampling period (April to October).
Seventy-two and 67% of L. beckii and P. pergandii
crawlers, respectively, were caught in the inner edge of
the trap, showing that crawlers moved preferentially
outwards in the tree canopy. Sticky tape traps detected
three peak periods of crawler movement in the tree
branches (figs 1a and 2a), which coincided with the
three annual generations found when sampling popu-
lations on leaves (figs 1b and 2b). The trend was
similar for the three plant substrates studied, green
twigs, branches and wood. The three peaks in crawler
movement occurred in the first half of May for the first
generation, in mid-July for the second and from mid-
September to the beginning of October for the third
generation. These dates correspond to the year of
sampling and variation in timing of peak crawler
emergence is likely to occur, as it depends on degree
day units that will vary from year to year. Immature
(first and second instar larvae) peaks (figs 1b and 2b)
occured within 15–30 days following crawler settling
(figs 1a and 2a).
For L. beckii (fig. 1a), the average number of crawlers
per trap found during any sampling period was greater
on branches (32.13 ± 16.52) (mean ± SEM) than on
green twigs (18.07 ± 6.65) and wood (22.83 ± 12.85),
Colonization of citrus fruit by armored scales 571

(a) Xest295 Xest195 Xest196

2.5 Green twigs Branches Wood 80

Percentage of infested fruits

mean no. crawlers / trap day

2
60 L. beckii

1.5
40
1

0.5 20

0 0
A M J J A S O M J J A S O

(b) L1 + L2
100 80 Xest295 Xest195 Xest196
% immature stages

Percentage of infested fruits

80
P. pergandii
60 60

40
40
20

0 20
A M J J A S O

Fig. 1. Phenology of L. beckii. (a) Crawlers on three 0

M J J A S O
plant substrates; (b) immature stages on leaves
SUECA 95
120
Percentage of infested fruits

(a) 100 A. aurantii

2.5 Green twigs Branches Wood
80
mean no. crawlers / trap day

2 60
40
1.5
20
1
0
M J J A S O
0.5
Fig. 3. Progress of infestation of fruit during the period
0
A M J J A S O of fruit growing by L. beckii, P. pergandii and
A. aurantii, in different groves from Vale`ncia, Spain,
(b) in 1995 and 1996
L + L2
100
% immature stages

80 of crawlers caught was higher on wood (49.37 ± 9.85),

60
40
20
0 significantly greater than that for the first generation
A M
Fig. 2. Phenology of P. pergandii. (a) Crawlers on
three plant substrates; (b) immature stages on leaves
although no significant differences were detected
between the three types of branches (F ¼ 1.35;
d.f. ¼ 2, 2; P > 0.05). The number of crawlers during
each emergence increased significantly during the
course of the year with each new generation
(F ¼ 10.68; d.f. ¼ 2, 2; P < 0.05). The average number
of crawlers per trap was 5.1 ± 0.46 for the first
generation, 22.63 ± 3.31 for the second and
45.3 ± 10.77 for the third. For P. pergandii (fig. 2a),
significant differences were detected between branches
(F ¼ 28.68; d.f. ¼ 2, 2; P < 0.01). The average number
intermediate on branches (25.8 ± 8.22) and lower on
green twigs (8.39 ± 3.41). The average number of
crawlers per trap for the second and third generations
(37.2 ± 15.72 and 32.19 ± 11.19, respectively) was
J J A S O
(14.17 ± 8.95) (F ¼ 9.95; d.f. ¼ 2, 2; P < 0.01).
3.2 Infestation of fruit
In Valencia, fruit start to grow in April and navel
oranges attain their final colour and size in November,
when the harvest starts. We observed an increase in
the percentage of infested fruit throughout the period
of fruit development, from April to November (fig. 3).
We detected fruit infested with the three species of
scales in June, and this suggests that the fruit became
infested early, by the first crawler generation in May.
For L. beckii the process of infestation of fruit started
earlier at the Xest1 95 and Xest2 95 groves than at
Xest1 96, probably because of the high levels of
infestation at those groves or because of differences in
temperature between 1995 and 1996. For P. pergandii

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572 Ma E. Rodrigo et al.

infestation of fruit started at the same time at the
three groves. For A. aurantii, the percentage of
infested fruit was above 40% already in June,
probably because of high levels of infestation at the
grove.
P. pergandii for settling under the calyx, more definite
in the first species, whereas A. aurantii tended to settle
preferably on the fruit rather than under the calyx. In
100

Percentage of infested fruits

In July, just before the second annual generation, the L. beckii
80
percentage of infested fruit reached at most 10% for
P. pergandii, ranged from 5 to 30% for L. beckii and 60
was approximately 50% for A. aurantii. After the
40
second crawler generation, fruit infestation increased
sharply, reaching by the end of September 40–60% of 20 Calyx
fruit infested for P. pergandii, 40–80% for L. beckii Free-fruit surface
and 100% for A. aurantii. 0
J J A S O

3.3 Role of the calyx 100

Percentage of infested fruits

P. pergandii Calyx
During June, the percentage of fruit with live scales of 80 Free-fruit surface
P. pergandii and L. beckii under the calyx was nearly
60
double the percentage of fruit with live scales settled on
the free fruit surface (figs 4 and 5). A sharp increase 40
in the number of infested fruit, that equalled or 20
exceeded the calyx infestation, was detected in mid-
July at the València grove, and during August at the 0
J J A S O
Sueca grove. In A. aurantii, fruit surface infestation
was nearly greater than calyx infestation at the Sueca
Percentage of infested fruits

100
grove (fig. 5), suggesting that the trend followed by
A. aurantii
A. aurantii is different from that followed by 80
P. pergandii and L. beckii. In some cases the percentage
60
of infested fruit decreased, presumably because of
mortality of the scales on fruit caused by abiotic 40 S
factors or because of the action of natural enemies. 20 Calyx
In order to confirm that first-generation crawlers Free-fruit surface
tend to settle under the calyx, we sampled six groves 0
J J A S O
once in June 2001. The results are shown in table 1 and
corroborate the results obtained from previous experi- Fig. 5. Percentage of calyx and free fruit surface
ences. There was a clear preference in L. beckii and infested with live scales of L. beckii, P. pergandii and
A. aurantii at the Sueca grove
100
Calyx
Free-fruit surface
Percentage of infested fruits

L. beckii Table 1. Comparison between calyx and free fruit sur-

80
face in percentage infestation and mean number of
60 diaspidid scales, at six orange groves sampled in June
2001 in Vale`ncia (Spain)
40

% Infestation1 Scales per fruit2

20
(mean ± SEM) (mean ± SEM)
0 L. beckii
J J A S
Calyx 27.67 ± 8.35* 1.07 ± 0.51*
Free fruit surface 10.17 ± 3.96* 0.19 ± 0.09*
100
Calyx P. pergandii
Calyx 42.67 ± 10.48** 1.76 ± 0.96*
Percentage of infested fruits

P. pergandii Free-fruit surface

80 Free fruit surface 24.33 ± 7.27** 0.66 ± 0.37*
A. aurantii3
60 Calyx 15.50 ± 7.75* 0.34 ± 0.17*
Free fruit surface 39.75 ± 10.97* 1.41 ± 0.68*
40 Significant differences (* P < 0.05; ** P < 0.01) were deter-
mined using paired t-test.
20 1
Percentage of infestation, L. beckii (t ¼ )3.31; d.f. ¼ 5; P <
0.05), P. pergandii (t ¼ )4.49; d.f. ¼ 5; P < 0.01) and
0 A. aurantii (t ¼ 4.01; d.f. ¼ 3; P < 0.05).
J J A S 2
Number of scales per fruit, L. beckii (t ¼ )3.06; d.f. ¼ 5; P <
0.05), P. pergandii (t ¼ )3.67; d.f. ¼ 5; P < 0.05) and
Fig. 4. Percentage of calyx and free fruit surface A. aurantii (t ¼ 3.48; d.f. ¼ 3; P < 0.05).
3
In A. aurantii, only four groves were considered because of the
infested with live scales of L. beckii and P. pergandii
lower level of fruit infestation in the other two groves.
at the Vale`ncia grove

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Colonization of citrus fruit by armored scales
Table 2. Mean number of diaspidid scales on orange
fruit in December 1996, as influenced by a barrier of
adhesive located around the fruit peduncle, or around the
calyx, or without barrier (control), in July 1996
Adhesive L. beckii P. pergandii A. aurantii
No 18.26 ± 5.44 a 3.48 ± 0.95 a 11.22 ± 3.69 a
Peduncle 7.93 ± 1.70 b 1.38 ± 0.41 b 12.40 ± 2.92 a
Calyx 2.84 ± 0.63 c 0.25 ± 0.09 c 12.53 ± 2.54 a
Mean ± SEM within columns followed by same letters are not
significantly different according to LSD test (P < 0.05).
L. beckii the percentage of fruit with scales under the
calyx was 2.7 times higher than the percentage of fruit
with scales on the surface. In P. pergandii calyx
infestation was 1.7 as high as fruit infestation.
Conversely, in A. aurantii the percentage of fruit with
scales under the calyx was 2.5 times lower than the
percentage of fruit with scales on the free fruit surface.
Comparing the average number of scales, we found 5.6
times more scales under the calyx than on the fruit in
L. beckii, 2.7 times more in P. pergandii and 4.1 times
less in A. aurantii.
3.4 Exclusion technique
As a result of placing adhesive barriers on fruit in July,
we observed that the average number of scales per fruit
in December was significantly lower when compared
with control fruit in the case of L. beckii and
P. pergandii (F ¼ 12.85; d.f. ¼ 2, 240; P < 0.01; F ¼
9.74; d.f. ¼ 2, 240; P < 0.01, respectively). Popula-
tions were lower when the barrier of adhesive was
placed over the calyx than when it was placed around
the peduncle of the fruit. In A. aurantii, the barrier of
adhesive was irrelevant and no significant differences
were found when we compared the average number of
scales found in the control fruit with the fruit isolated
with adhesive barriers neither around the peduncle nor
over the calyx (F ¼ 0.47; d.f. ¼ 2, 240; P > 0.05)
(table 2). From these results, we estimate that most
A. aurantii crawlers originated from the reproduction
of females already settled on the fruit, whereas we
found that in L. beckii and P. pergandii, 57 and 60% of
new settlers originated from crawlers walking across
the peduncle and 28 and 34% of settlers proceeded
from the calyx area, respectively.
4 Discussion
Our results showed the existence of three defined
periods of crawler production for L. beckii and
P. pergandii during the season in the study area, in
May, July–August and September–October. In other
areas of similar climate, such as other Mediterranean
countries or California, two to four discrete genera-
tions are usually found during the year in these two
species of diaspidids (Benassy and Soria, 1964; Abbasi,
1975; Moreno and Kennett, 1985; Hafez et al., 1987;
Walker et al., 1990). The movement of crawlers along
the branches seemed to increase throughout the year,
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573
at least from first to second generation, and they
moved preferentially outwards in the tree canopy.
Crawler population density on branches was similar
for the two species of diaspidids, but in P. pergandii it
was more abundant in the inner part of the tree canopy
than in the exterior. This is possibly because of the fact
that the proportion of the whole tree population
settled on the bark of the trunk and main limbs is
higher in this species (Bodenheimer, 1951; Gerson,
1967). Our results demonstrated the utility of the
sticky tape traps, already used in other scales as
A. aurantii and Quadraspidiotus perniciosus (Com-
stock) (University of California, 1991a,b), to monitor
the population fluctuations of L. beckii and
P. pergandii.
Several authors have observed that fruit become
infested by A. aurantii at the beginning of their
development and fruit infestation by this species
continues as long as fruit remain on the trees. This
process is initiated at a very early stage of fruit
development by crawlers of the first generation (Geor-
gala, 1963; Orphanides, 1982; Alexandrakis, 1983;
Atkinson, 1983; Carroll and Luck, 1984). Previous
information on this behaviour in L. beckii and
P. pergandii is scarce (Bodenheimer, 1951; Gerson,
1967). It has been suggested that L. beckii crawlers
invade the developing fruit only in the second genera-
tion in August (Benassy et al., 1980). Our study
demonstrated that citrus fruit became invaded by
L. beckii and P. pergandii crawlers continuously during
the entire period of fruit growth, similar to what was
previously reported for A. aurantii.
According to our results, the three scale species
showed different behaviour during infestation of the
developing fruit in the first generation. Crawlers of
L. beckii and P. pergandii invading the fruit tended to
settle preferentially under the fruit calyx and not on the
exposed fruit surface. At the end of the first flush of
crawlers, the percentage of fruit infested was 2.7–3.4
times higher under the calyx compared with the
exposed fruit surface in L. beckii, and 2.7–3.3 times
higher in P. pergandii. In contrast, A. aurantii prefered
the exposed fruit surface and populations were always
higher on the exposed fruit surface compared with the
calyx. We have found no references to the calyx as a
settling place in the numerous papers published on
A. aurantii. Bodenheimer (1951) indicates that first
larvae of L. beckii infesting fruit at a very early stage,
from the end of April on, settle on the fruit in the
shelter of the sepal leaves and do not settle visibly on
the surface of the fruit before mid-July. In P. pergandii,
Gerson (1967) observed that the first infestation of fruit
occurred mainly in the calyx area and throughout the
period of fruit development. He reported a continuous
flow of scales from the calyx to other areas of the fruit
surface. Our experiences were the first quantification of
this process for the three diaspidid species.
Several reactions to external stimuli when deciding
on a place to settle down and start feeding have been
described in crawlers of these diaspidid species. In
general, diaspidids tend to show positive thigmotactism
and this causes them to settle near obstacles, crevices,
contact surfaces or irregularities of the substrate
574
(Beardsley and Gonzalez, 1975). This was experiment-
ally demonstrated for A. aurantii (De Bach and White,
1960) and L. beckii (Carnegie, 1957; Hulley, 1962).
Bodenheimer (1951) indicated that positive thigmotac-
tism is probably the main reason why crawlers of
L. beckii prefer to settle below the sepals of the fruit.
A strong positive response to the light (phototactism)
was also demonstrated in A. aurantii (De Bach and
White, 1960). L. beckii was considered positively
phototactic by Hulley (1962) but not by Carnegie
(1957). Differences in settling behaviour among dias-
pidid species suggest that they probably exhibit differ-
ences in the response to thigmotactism and/or
phototactism, although other types of stimuli, like
chemotactism as suggested for L. beckii by Benassy
et al. (1980), could also be involved.
There were also apparent differences among the
three especies in the fruit invasion behaviour during
the second generation. According to the results of our
exclusion experiment, in the previous generation, it
was found that aerial invasion could also be involved
as dispersal among armored scale crawlers is accom-
plished not only by active wandering but also by wind
(Willard, 1974; Beardsley and Gonzalez, 1975). How-
ever, crawlers of L. beckii and P. pergandii invading the
fruit were preferentially from other tree substrates or
from females settled under the fruit calyx. We found
the percentage of invaders from the calyx area to be
considerably lower than that reported by Bodenheimer
(1951), who pointed out that 90% of L. beckii scales on
fruit originate from the first settlers below the sepals
and not from immigrants.
Differences in settling and fruit invasion behaviour
between the three diaspidid scales observed in this
study could lead to the use of different management
strategies to reduce scale infestation of mature citrus
fruit at harvest in areas or orchards where the three
species coexist. Diaspidid scales are usually sprayed
with two types of pesticides, organophosphate com-
pounds (OP) and mineral oils. Two key aspects for the
success of this chemical spray are the developmental
stage that predominates in the population and the
moment of invasion of the citrus fruit by the scales. It
is known for A. aurantii that the proportion of
development stages changes gradually and the mortal-
ity produced by the pesticides is dependent on the
predominant stage, older stages being more tolerant to
chemical sprays, OPs (Abdelrahman, 1973; Schoonees
and Giliomee, 1982) and mineral oil (Riehl, 1988;
Davidson et al., 1991).
The efficacy of OP sprays is higher for scales living
on leaves and branches, whereas mineral oil shows
higher efficacy for scales settled on fruit (Cressman
et al., 1953; Carman, 1977). This, together with the
phytotoxicity produced by oils in the early stages of
fruit development, led to the application of oils (or oil
plus OP) in mid-summer to control diaspidid scales in
citrus when the scale species present were L. beckii and
P. pergandii, in Spain. This has been the most common
time period for spraying scales for many years in
Spain. Sprays with OPs at the end of the spring, in the
early stages of fruit development, were applied only in
cases of heavy infestations (Moner, 1994; Francés et al.,
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Ma E. Rodrigo et al.
1998; Costa-Comelles et al., 1999). Our finding of
differences in settling behaviour and origin of scales on
fruit between A. aurantii and the former two scale
species, together with the recent expansion and out-
break of A. aurantii in all the Spanish citrus area
suggest changes in the pest control strategy that could
improve scale management in Spain. For example, the
application of OPs should be reserved preferentially to
the spring, to the first period of crawler production,
whereas mineral oils could be applied early in the
summer, before the scales settled on the fruit start to
reproduce. Comparative trials should be carried out in
the field to test the validity of these suggestions. Other
aspects of the strategy for controlling diaspidids in
Spain, as the definition and use of treatment thresholds
or the role of biological control agents in regulating the
diaspidid populations, should be also addressed to
optimize the management of these pests in Spanish
citrus groves.
Acknowledgements
The authors thank M.C. Torralba and cooperating growers
(Cooperativa de Xest and Cooperativa de Catadau) for
logistical support. This research was funded by Ministerio
de Ciencia y Tecnologı́a (AGF98-0330). Special thanks to
M.C. Lahoz for her disinterested help in the grammatical
correction of this manuscript.
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Author’s address: Ma Eugènia Rodrigo Santamalia (corres-
ponding author), Entomologı́a- ETSEA, Universitat Polit-
ècnica de Valencia, 14, Camı́ de Vera. 46022 València, Spain.
E-mail: erodrigo@eaf.upv.es