New Zealand Journal of Crop and Horticultural Science

ISSN: 0114-0671 (Print) 1175-8783 (Online) Journal homepage: www.tandfonline.com/journals/tnzc20

Distribution of greenhouse whitefly Trialeurodes

vaporariorum (Homoptera, Aleyrodidae) and
Encarsia formosa (Hymenoptera, Aphelinidae)
in a greenhouse tomato crop: implications for
sampling

N.A. Martin, R.D. Ball, L.P.J.J. Noldus & J.C. van Lenteren

To cite this article: N.A. Martin, R.D. Ball, L.P.J.J. Noldus & J.C. van Lenteren (1991) Distribution
of greenhouse whitefly Trialeurodes vaporariorum (Homoptera, Aleyrodidae) and Encarsia
formosa (Hymenoptera, Aphelinidae) in a greenhouse tomato crop: implications for
sampling, New Zealand Journal of Crop and Horticultural Science, 19:3, 283-290, DOI:
10.1080/01140671.1991.10421812

To link to this article: https://doi.org/10.1080/01140671.1991.10421812

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New Zealand Journal ofCrop and Horticultural Science, 1991, Vol. 19: 283-290
0114--0671/91/1903--0283 $2.50/0 © Crown copyright 1991
Distribution of greenhouse whitefly Trialeurodes vaporariorum
(Homoptera, Aleyrodidae) and Encarsia formosa
(Hymenoptera, Aphelinidae) in a greenhouse tomato crop:
implications for sampling
N.A.MARTIN
DSIR Crop Protection
Private Bag, Auckland, New Zealand
R.D.BALL
DSIR Physical Science
Private Bag, Auckland, New Zealand
L. P. J. J. NOLDUS*
J. C. VAN LENTEREN
Department of Entomology
Agricultural University
Wageningen, The Netherlands
*Present address: Noldus Information Technology
bv, University Business & Technology Centre,
Vadaring 51, 6702 EA Wageningen, The
Netherlands.
Abstract Vertical and horizontal distributions of
whitefly puparia in a Dutch commercial greenhouse
tomato crop were studied over 16 weeks in order to
test sampling techniques. In the period Week 7-
8,2270 puparia infested 137 out of the 18144 plants,
whereas in Week 15-16,2005 puparia infested 1024
plants. Calculation of spatial autocorrelations showed
that at both times infested plants were grouped along
rows with a much lower probability of them being
alongside one another in adjacent rows. The seventh
leaf layer (seven leaves up from the base of the plant)
was the most heavily infested in Week 7-8 and the
distribution of the infestation was the same as for the
whole plant. It is concluded that presence-absence
sampling for whitefly puparia and for estimates of
H91053
Received 5 June 1991; accepted 16 October 1991
283
parasitism should be restricted to the zone of three-
five leaf layers with most puparia and involve the
examination of a single leaf taken from widely spaced
plants in each row of a small greenhouse or in areas
of infestation in a large greenhouse.
Keywords spatial distribution; greenhouse whitefly;
Trialeurodes vaporariorum; Homoptera;
Aleyrodidae; Encarsia formosa; Hymenoptera;
Aphelinidae; greenhouse tomatoes; parasitism;
puparia; monitoring
INTRODUCTION
A study of greenhouse whitefly Trialeurodes
vaporariorum (Westwood) (Homoptera: Aleyrodidae)
populations in a large greenhouse in the Netherlands
revealed that the degree of aggregation of puparia
amongst the plants decreased with time and that
clumping occurred at two spatial levels (Eggenkamp
et al. 1982; Noldus et al. 1986; Lenteren & Noldus
1990; Noldus & Lenteren 1990). The basic component
of aggregation was a compact clump occupying one
plant. Groups of clumps, comprising several or many
plants, formed spatially separate infested zones. The
authors concluded that sampling methods derived
from small greenhouses are not applicable to large
greenhouses, though they noted that the groups of
clumps were spread over enough plants to fill a small
greenhouse.
A study of the distribution of greenhouse white-
fly and its parasite Encarsia formosa Gahan
(Hymenoptera: Aphelinidae) on tomato crops in small
greenhouses in New Zealand provided a sampling
method for estimating if parasitism was above or
below preset thresholds (Martin & Dale 1989). The
method may be applicable to part of a large greenhouse
that encompasses one group of whitefly clumps.
Sampling systems may be designed to study
population dynamics or to determine whether pest
control is required. Different sampling objectives
require different levels of precision (Ekbom & Xu
284 New Zealand Journal of Crop and Horticultural Science, 1991, Vol. 19

13 unparasitised Parasitised Fig. 1 Number of unparasitised

12 (white) and parasitised (black)
11 whitefly puparia in each leaf layer
during Week 7-8.
10
...Gl 9

-
>. 8
.!!!
7
co 6
Gl
..J 5

4
3
2
1
-
0 100 0 100 200 300 400
Number of whitefly puparia

1990)-decisions on control measures can be made MATERIALS AND METHODS

using low levels of precision. Ekbom & Xu (1990)
also pointed out that the within plant distribution of The basic methodology has been described in detail
each pest stage must be understood. by Eggenkamp et al. (1978, 1982) and is summarised
The present work analyses the application of the here. The greenhouse was 86.~ X 75.0 m with a
method for estimating parasitism (Martin & Dale central path. The young tomato plants were kept for 2
1989) to data from a large Dutch greenhouse weeks in one part of the greenhouse. During 2-5
(Eggenkamp et al. 1982). The data are also analysed January 1974 the plants, 30 cm high and with four
to establish the vertical and horizontal distribution of leaves, were planted out in 216 rows each of 84
whitefly puparia in the leaf layers during one of the plants. There were four rows ofplants between pillars
eight sampling periods. A further comparison is made (roof supports) with a path, 1.2 m wide, between
between some aspects of the horizontal distribution every second row and 0.4 m between each row in a
of puparia on the plants during two time periods pair. Within the row the plants were 0.43 m apart.
when the populations were similar but the degree of The experiment lasted 16 weeks during which time
aggregation was very different. the plants were regularly layered (lowered) after they
frrst reached 2 m high. Whitefly adults (121) were
found at planting out and E.fonrwsa was introduced
Table 1 Vertical distribution of whitefly puparia, four times with 0.7, 2.1, 4.5, and 2.5 black puparia
percentage of puparia parasitised and infested leaves in a per tomato plant distributed in Weeks 4, 6, 8, and 10,
greenhouse tomato crop in a large greenhouse with 18 144 respectively. At regular intervals, total counts were
plants and 7-8 weeks from planting. regularly made of white (unparasitised) and black
~~ Th~ (parasitised) puparia on each leaf of every plant. The
(lowest leaf To~ % infested numbers of larvae and adults on each plant were also
= no. 1) puparia parasitism leaves recorded. The complete counts were initially every
13 3 0 1 week (fIrst 6 weeks). From Week 7 to 16 each
U ~ 44 5 complete count of all puparia took 2 weeks to make.
11 77 35 12
10 56 36 18 The present analysis concerns only the spatial
9 303 65 35 distribution of parasitised and unparasitised whitefly
8 286 41 51 puparia recorded during Week 7-8 and Week 15-16.
7 319 52 63 These two periods were chosen because high numbers
6 550 69 53
5 339 52 37 of puparia were found on both occasions but with
4 23~ 76 13 different degrees of aggregation (Noldus et al.
3 ~ n 7 1986). In Week 7-8 the position ofeach infested leaf
2 8 75 2 on the plant was recorded but in Week 15-16 the
1 0 0 positions on a plant of very few infested leaves were
To~s 2270 297 determined.
Martin et al.-Distribution of greenhouse whitefly and Encarsia
The data from Week 7-8 were segregated into the
separate leaf layers and the vertical distribution of
whitefly puparia was examined. Grid sampling
(Martin & Dale 1989) was tried on the data from Leaf
layer 7 (seven leaves up from the base of the plant) in
one group of whitefly clumps (See Table 1). A
rectangular sampling grid consisted ofplants at regular
intervals along the chosen rows which were spaced
an odd number of rows apart. Sampling grids can
have different between-plant and between-row
spacings, but they always have a random starting
plant. Leaf layer 7 was chosen because it had the
highest number of infested leaves and the total
numbers of black and white puparia were repre-
sentative of the population when parasitised whitefly
puparia tum black and before adult whitefly emerge
from unparasitised puparia (Table 1, Fig. 1). The data
on the number of whitefly puparia on each leaf in
Leaf layer 7 were also used to analyse the spatial
distribution ofplants for the purpose of sample design.
This was achieved by calculating autocorrelations.
The square root transformation was applied to all
counts before taking autocorrelations were attempted.
Distance (or lags) was defined as numbers of plants
along and across rows.
The data from both Week 7-8 and Week 15-16
were used to calculate autocorrelations for the number
of puparia on the whole plant irrespective of which
leaf was infested. The frequency distributions of the
numbers of infested leaves per plant and puparia per
plant from the two sampling occasions were
compared.
Definition of autocorrelation
Leti,j, m, n,l], 12 be integers.
Given data Xij for k=i<=m, k=j<=n, the auto-
correlation at lag (1], 12 ) is the ordinary correlation
cor(x,x') where
x'·',J·=X·1+1/, J+1
. 2
and where we restrict x and x' to indices for which
they are both defined. Our defmition is a generalisation
of the autocorrelation of time series (Fuller 1976).
Note that in the greenhouse numbers i and j index
rows and columns and m and n are the numbers of
rows and columns respectively. The data value x.+11,
j+12 is said to be at lag (1], 12) from Xi,j' We will refer
to a lag of (k, 0) as a lag of k along rows and a lag of
(0, k) as a lag along co~umns, or if the direction is
clear from the context, simply a lag of k. We looked
at autocorrelations at lags (/1, 0) (between plants in
different rows but in the same column or at lags (0, 12)
(between plants in the same row) (cf. Table 4).
285
RESULTS
Vertical distribution of whitefly puparia in
Week 7-8
During Week 7-8, whitefly puparia were found on
Leaf layers 2-13 with highest numbers of puparia on
Leaf layers 4-9 (Table 1). In Leaf layer 4 (the older
leaves) the puparia are on only a few leaves and a
high proportion were parasitised. Only 137 (0.76%)
of the 18 144 plants had any leaves with any
infestation, but those leaves held up to 18 puparia per
leaf. This non-random distribution indicates that the
initial infestation in the greenhouse was concentrated
on a few plants. The maximum number of infested
leaves in a layer (Leaf level 7) was 63 or 0.35% of
leaves in the layer.
Horizontal distribution of whitefly puparia in
Leaf layer 7 in Week 7-8
Grid sampling was used in an area of high whitefly
density, in an attempt to estimate whitefly populations
and to determine if the proportion of parasitised
whitefly puparia was above or below the action
threshold of 80%, the threshold currently used in
New Zealand. The sample area of 1032 plants
consisted of 24 rows (180-203, or Bays 46-51) with
42 plants/row starting from the aisle. Only seven
(1.62%) of the 432 leaves sampled in Leaf layer 7
contained whitefly puparia. This was too few to
estimate parasitism.
The spatial autocorrelations along rows in Leaf
layer 7 were high, up to 30% at lags (distances) of up
to 12 plants. In contrast the autocorrelations between
rows were low, below 10% at Lag 1, i.e., adjacent
plants in adjacent rows. This means that plants with
infested leaves in Leaf layer 7 and probably other
layers, tend to be aggregated in groups of 12 plants
arranged in a single row. (For those unfamiliar with
autocorrelations, full figures are given for the
autocorrelations for whole plants in Table 4 and these
are discussed in the next section).
Broadly speaking, in operating a sampling
procedure it is optimal to choose sampling points so
that autocorrelations are equal between neighbouring
points along a row of plants and at right angles to the
row. If a sampling procedure for this crop and pest is
to have a high probability offmding a zone ofinfested
plants and only to sample it once (Martin & Dale
1989), the procedure should include every row and
examine a leaf from plants more than 12 plants apart
along the rows. Because the infested plants are
aggregated along single rows, the grid sampling
system proposed by Martin & Dale (1989) needs to
286 New Zealand Journal of Crop and Horticultural Science, 1991, VoL 19

be slightly modified so that the grid includes every showed that there was strong correlation between
row. It is probably more sound not to sample plants infested plants within rows and much lower correlation
the same distance from the row ends in the adjacent between infested plants in adjacent rows (Table 4).
rows but better to have the equivalent of two These autocorrelations describe mathematically the
interlocking grids. For example if the samples are observations of Eggenkamp et al. (1982). A
taken from every 15th plant, the starting plant in the
odd numbered rows could be Plant 4 and the starting
Table 3 Frequency distribution of the numbers of
plant in the even numbered rows could be Plant 10. whitefly puparia per tomato plant in a large greenhouse on
two occasions.
Comparison between Week 7-S and
Week 15-16 No. of plants
The distribution of whitefly puparia bo$ within and No. of puparia Weeks after planting
between the plants was different in Week 7-8 to that per plant 7-8 15-16
in Week 15-16 (Tables 2 and 3), even though the 1 21 618
same plants were examined and the total numbers of 2 19 200
puparia found were similar (Table 3). During Week 3 10 96
7-8,137 plants were infested with whitefly puparia. 4 13 52
One plant had eight leaves infested whereas 83% of 5 6 18
the infested plants had less than four infested leaves, 6 5 13
and nearly half had only one infested leaf (Table 2). 7 7 5
8 6 5
This contrasts with Week 15-16 when 1024 plants 9 5 3
were infested but >90% had only one leaf infested. 10 5 4
Over half the infested plants in Week 15-16 were 11 3 0
infested with only one puparium and the highest 12 3 0
number found on a plant was 26 (Table 3). In Week 13 2 1
7-8, half the infested plants had five or more puparia 14 1 1
15 0 1
present. Four plants were infested with more than 16 1 3
100 puparia, and one with 497 puparia. Two of these 17 1 1
heavily infested plants had> 100 puparia on some 18 2 0
leaves. The changing distribution of the whitefly 19 2 0
puparia in the greenhouse has been described 20 1 0
mathematically by Noldus et al. (1986). -
23 2 1
Spatial autocorrelations of whitefly puparia on 24 1 0
whole plants for both Week 7-8 and Week 15-16 25 1 1
26 2 1
-
Table 2 Frequency distribution of the number of whitefly 29 2 0
infested leaves per tomato plant in a large greenhouse on 30 2 0
two occasions. 31 2 0
32 1 0
No.of plants 33 3 0
No. of infested Weeks after planting 34 2 0
leaves per plant 7-8 15-16 35 0 0
36 1 0
1 58 992 -
2 38 13 43 1 0
3 18 7 50 1 0
4 12 4 102 1 0
5 7 3 171 1 0
6 3 3 242 1 0
7 o 1 497 1 0
8 1 0
Total number of
9 o 0
137 1024
10 o 1 infested plants
Total number of plants Total number of
with infested leaves 137 1024 whitefly puparia 2296 2005
Martin et aI.-Distribution of greenhouse whitefly and Encarsia 287

Boundary of greenhouse

<>

o
(j): ~. ' ..
o 0 .

Fig. 2 The position of infested plants and the density of whitefly puparia on each plant in the greenhouse during
Week 7--S. The greenhouse was 86.4 X 75 m. The size of the circle is proportional to the number of puparia per plant, see
Table 3.

comparison of the autoeorre1ations for Week 7-8 and
Week 15-16 (Table 4), shows the more aggregated
distribution of whitefly in Week 7-8 compared with
Week 15-16 (Eggenkamp et al. 1982; No1dus et al.
1986) and is illustrated in Figs 2 and 3. The distribution
of whitefly along rows can be interpreted as evidence
that whitefly adults tend to move along rows rather
than from row to row. This was also mentioned by
Eggenkamp etal. (1982).
DISCUSSION
The most important result of this analysis of data
from a Dutch commercial greenhouse tomato crop
was the mathematical. confirmation using auto-
correlation analysis that'plants infested with whitefly
puparia tend to be grouped along the rows (Table 4),
(cf. Figs 2 and 3; Eggenkamp et al. 1982; No1dus et
al. 1986). The advantage of the autocorrelation
analysis is that it provides information on the size of
the groups of infested plants which has implications
for sampling procedures. This is discussed below.
The distribution of whitefly puparia reflects where
females laid eggs because all the non-adult whitefly
stages stay on the same leaf on which the eggs are
laid. However, the position of the puparia is the
starting point for subsequent dispersal of the next
generation of adults.
Whitefly dispersal could be influenced by two
distinctive features of commercial crops. First, the
plants are arranged in rows with a shorter distance
between plants in the same row than plants in the row
on the other side of the passage. It is possible that
when newly emerged whitefly adults move up to the
young leaves at the top ofthe plants, and are dispersing
from the plant of emergence (Vianen et al. 1988;
Noldus & Lenteren 1990), that the adults tend to stay
amongst the leaves of closely spaced plants rather
288 New Zealand Journal of Crop and Horticultural Science, 1991, Vol. 19

Boundary of greenhouse
: :

:: ...
Q :;
•• 0

,
0
:~ ;"
".!"

o
,
...
~. z

..
'. '
'0

"
0 ••••
.. ,: ! I

! I ,·0.
:~~iJ~;'
"'U":::;..:.
.
0' •

o 0. ."~:i' : .
.,. ..
• • 0 I .::' •

"t:i~08i
. I'
Oi: .• I •
I.
. !·.~I 0A- 0 ,.

. ..
,-' &. i·.: a 'f._fr O -
:
:0.: : ... i': ~: 10 0
I . .. .'.. I

I;o~ : ' ~ Ii. ;. .o•

:!·i.~ ~.~ .. !;~ ...
•0

&

• • .. •
,1.1
j'"
6°:"
.' i 0
;01 ,"
.. v!_.. . :. 0.

.. :1..: :,'.17: '!,.'f'Oi I:':.·

:. '.'o;fb~'"
Y'?<
or •• 1'
.2~1". lAo
,. Atll.~·
.tQ_ .. ~Q·t
"of!'
••

Fig. 3 The position of infested plants and the density of whitefly puparia on each plant in the greenhouse during Week
15-16. The greenhouse was 86.4 X 75 m. The size of the circle is proportional to the number of puparia per plant, see
Table 3.

than cross the more open space of the passage. Where
plants are arranged in pairs of rows as in the
commercial glasshouse it might be expected that
there could be a correlation of infestation between
adjacent plants in a pair ofrows. This was not checked
in the statistical analysis, but examination of Figs 2
and 3 suggest that there is no correlation. A possible
reason for the movement of adult whitefly along the
plant row rather than to plants in the adjacent rows
may be a preference to fly along the interface between
vegetation and free flying space. Second, in
commercial greenhouse tomato crops the plants are
subject to regular handling (removal of side shoots,
twisting of stems around support strings) which
disturbs the adult Whitefly and cause some to fly.
These adults may then settle on another plant The
distribution of puparia suggests that some whitefly
may fly or be taken (e.g., on clothes) a long way to
uninfested parts of the greenhouse but that most tend
to stay in the same row of plants. One of the other
features that needs to be accounted for in any model
of whitefly and E.formosa distribution in greenhouse
tomatoes is the high proportion of plants in Week
15-16 that had only one puparium (fable 3).
Monitoring and sampling systems for whitefly
and its parasite will be most soundly based if there is
a full understanding of the population interactions of
whitefly, E.formosa, and the crop. Aspects of whitefly
behaviour that influence dispersal in commercial
greenhouse tomato crops are directly relevant to
sampling systems. Recent studies have shown that
the whitefly adults disperse during the first few days
after emergence (Lenteren & Noldus 1990; Noldus
& Lenteren 1990) and then dispersal ceases.
Experiments using evenly spaced plants showed that
young adult whitefly dispersed to plants in all
Martin et al.-Distribution of greenhouse whitefly and Encarsia 289

directions from the point of release (Vianen et al.
1988; Noldus & Lenteren 1990). However,
observations on the commercial crop used in this
study showed that there was local dispersal ofwhitefly
(mainly along rows) and on the evidence of new
areas of infected plants, longer distance dispersal
(Figs 2 and 3; Eggenkamp et al. 1982). Both features
need to be accounted for in any model of whitefly
crop interaction.
Advisors and growers want to know if whitefly
populations are going to stay below the economic
threshold (i.e., the population level that causes honey
dew and sooty mould on fruit), if there is sufficient
parasites in the crop and do they need to do anything.
The description of whitefly distribution in this paper
is in a form that growers and advisors can more
readily appreciate than the mathematical analysis of
Noldus et al. (1986). For example the number of
puparia on each plant on the two sampling occasions
(Table 3) can be related to the risk: ofgetting honeydew
and sooty mould on fruit. In Week 7-8 the two most
heavily infested plants each had leaves with > 100
puparia. Honeydew from these high levels of
infestation caused no problems.
The main practical benefit from this study has
been to improve the efficiency of the sampling
procedure proposed by Martin & Dale (1989). The
autocorrelation analysis of both the whole plants and
a single leaf layer confirmed that infested plants tend
to be in rows and gave information on the length of
groups of infested plants (Table 4). Before discussing
this, it should be pointed out that only part of a large
greenhouse needs to be sampled and that the grower
should identify the area where the whitefly infestation
is of concern. The monitoring system already takes
into account the distribution of puparia along plant
rows (Eggenkamp et al. 1982; Martin & Dale 1989),
but the grid sampling plan needs slight modification
so that the grid includes every row and leaves are
examined from more widely spaced plants, e.g., every

Table 4 Autocorrelations of the numbers of whitefly puparia per plant in Week

7-8 and Week 15-16. Numbers of puparia transformed to square root for
analysis.
Along rows Along columnsa
Lagb Week 7-8 Week 15-16 Lagc Week 7-8 Week 15-16
0 1.00 1.00 0 1.00 1.00
1 0040 0.30 1 0.23 0.22
2 0.38 0.29 2 0.19 0.23
3 0.30 0.27 3 0.08 0.18
4 0.33 0.29 4 0.03 0.19
5 0.25 0.25 5 0.02 0.17
6 0.25 0.28 6 0.00 0.16
7 0.23 0.27 7 0.14
8 0.18 0.27 8 0.13
9 0.16 0.25 9 0.12
10 0.18 0.26 10 0.09
11 0.18 0.26 15 0.04
12 0.21 0.25 20 0.01
13 0.16 0.23 25 0.00
14 0.22 0.23
15 0.16 0.22
20 0.08 0.16
25 0.00 0.14
30 0.12
35 0.10
40 0.09
45 0.07
aLines of plants with the same position in row.
bDistarlce (units within row inter plant spacing) between corresponding plants in
autocorrelation calculation.
cDistance (units of inter row spacing) between corresponding plants in the
autocorrelation calculation.
290 New Zealand Journal of Crop and Horticultural Science, 1991, Vol. 19
IS-20th plant. Detailed instructions for using a grid
sampling plan in greenhouse tomato crops and
accompanying recording charts are now being
prepared.
REFERENCES
Eggenkamp-Rotteveel Mansveld, M. H.; Ellenbrook, F. 1
M.; Lenteren, 1 C. van; Woets, 1 1978: The
parasite-host relationship between Encarsia
formosa Gah. (Hym., Aph~linidae) and
Trialeurodesvaporariorum (Westw'.) (Homoptera,
Aleyrodidae) vrn. Comparison and. evaluations
of an absolute count and a stratified random
sampling programme. Zeitschrift fur angewandte
Entomologie 85: 133-140.
Eggenkamp-Rotteveel Mansveld, M. H.; Lenteren, 1 C.
van; M. H., Ellenbrook, F. 1 M.; Woots, J. 1982:
The parasite-host relationship between Encarsia
formosa (Hym., Aphelinidae) and Trialeurodes
vaporariorum (Hom., Aleyrodidae) XII.
Population dynamics of parasite and host in a
large, commercial glasshouse and test of the
parasite-introduction method used in the
Netherlands. Zeitschrift fur angewandte
Entomologie 85: 133-140 (first part), 258-279
(second part).
Ekbom, B. S.; Xu, R. M. 1990: Sampling and spatial
patterns of whiteflies. pp. 107-121 in: Whiteflies:
their bionomics, pest status and management,
Gerling, D. ed. Andover, England, Intercept Ltd.
Fuller, W. A. 1976: Introduction to statistical time series.
Wiley.
Lenteren, J. C. van; Noldus, L. P. J. 1 1990: Whitefly-
plant relationships: behaviourial and ecological
aspects. pp. 47-89 in: Whiteflies: their bionomics,
pest status and management. Gerling, D. ed.
Andover, England, Intercept Ltd.
Martin, N. A; Dale, 1 1989: Monitoring greenhouse
whitefly and parasitism: a decision approach. New
Zealand journal ofcrop and horticultural science
17: 115-122.
Noldus, L. P. J. 1; Lenteren, J. C. van 1990: Host
aggregation and parasitoid behaviour: biological
control in a closed system. Pp. 229-269 in: Critical
issues in biological control. Mackauer, D.; Ehler,
L. E.; Roland, 1 ed. Andover, England, Intercept
Ltd.
Noldus, L. P. 1 1; Xu, R. M.; Eggenkamp-Rotteveel
Mansveld, M. H.; Lenteren, 1 C. van 1986: The
parasite-host relationship between Encarsia
formosa Gahan (Hymenoptera, Aphelinidae) and
Trialeurodes vaporariorum (Westwood)
(Homoptera, Aleyrodidae) XX. Analysis of the
spatial distribution of greenhouse whiteflies in a
large glasshouse. Journal of applied entomology
102: 484-496.
Vianen, van A.; Xu R. M.; Lenteren, 1 C. van 1988: The
parasite-host relationship between Encarsia
formosa Gahan (Hymenoptera, Aphelinidae) and
Trialeurodes vaporariorum (Westwood)
(Homoptera, Aleyrodidae) xxvrn. The influence
of whitefly densities and temperature on the
horizontal dispersal of greenhouse whiteflies.
Journal ofapplied entomology 105: 436-449.