Phytoparasitica (2010) 38:313–325
DOI 10.1007/s12600-010-0105-9
Influence of weedy field margins on abundance patterns
of the predatory bugs Orius spp. and their prey, the western
flower thrips (Frankliniella occidentalis), on faba bean
Ekrem Atakan
Received: 7 January 2010 / Accepted: 28 June 2010 / Published online: 22 July 2010
# Springer Science+Business Media B.V. 2010
Abstract The influence of weedy field strips on the
abundance patterns of western flower thrips,
Frankliniella occidentalis (Pergande) (Thysanoptera:
Thripidae), and predatory bugs of Orius spp. (Hemi-
ptera: Anthocoridae), on faba bean and on weeds was
investigated in Adana province, Turkey, during 2005–
2006. There were two treatments: in one treatment,
weeds at the margins and inside the plots were
regularly controlled by tillage; in the other treatment
no weed control was done. Thrips and Orius were
sampled by beating faba bean plants and weeds. Orius
niger (Wolff) was the most abundant predatory insect
species in faba bean and flowering weeds, with
numbers of adults and nymphs significantly greater in
plots with weedy margins than in weed-free plots.
Flowering weeds did not contribute to the abundance
of F. occidentalis on faba bean. Abundance of adults of
Orius spp. did not coincide with the abundance of
F. occidentalis on faba bean or weeds. There were
significant negative associations for numbers of Orius
spp. among faba bean and the weed species Lamium
amplexicaule L. or Sinapis arvensis L. (P<0.05),
indicating movement of Orius individuals from the
weeds to faba bean during March–April. Finally, faba
bean and weeds may provide some benefits to
E. Atakan (*)
Plant Protection Department, Faculty of Agriculture,
Çukurova University,
Adana 01030, Turkey
e-mail: eatakan@mail.cu.edu.tr
predators, such as nectar, pollen, shelter and egg-
laying sites rather than as sources of insect prey.
Cultivation of faba bean could be useful for conserva-
tion and augmentation of beneficial insects, including
Orius spp. Furthermore, field margins bearing flower-
ing weeds such as S. arvensis and L amplexicaule
should be protected against destructive management
practices, because they host considerable numbers of
the Orius species.
Keywords Lamium amplexicaule . Prey-predator
interactions . Polyphagous predators . Sinapis
arvensis . Vicia faba . Weeds
Introduction
Faba bean (Vicia faba L.) is an important nutritious
human food crop containing high protein levels (Anon.
2001). It is widely cultivated, together with other winter
vegetable crops, in the eastern Mediterranean region of
Turkey. Faba bean is also a good rotational crop for
fixing nitrogen and thus enhancing soil fertility. The
production of this crop ranks fourth among leguminous
crops in Turkey. Faba bean is consumed as either green
pods or dried beans in the Aegean and Mediterranean
regions of Turkey (Anon. 2001).
The polyphagous predators Orius spp. (Hemi-
ptera: Anthocoridae) are often considered important
natural enemies of thrips (Riudavets et al. 1995). In
Turkey, Orius niger (Wolff) effectively suppresses
314
the Frankliniella flower thrips in cotton fields in the
Çukurova region that are not treated with insecti-
cides (Atakan 2006; Atakan and Gençer 2008).
Orius spp. were also recorded on various vegetable
crops grown in different parts of Turkey (Bulut and
Göçmen 2000; Yaşarakıncı and Hıncal 2000). Orius
spp. are important biological control agents that are
able to regulate populations of the serious pestifer-
ous thrips—the western flower thrips Frankliniella
occidentalis (Pergande) (Thysanoptera: Thripidae)—
especially in protected crops (Tavella et al. 2000,
2003; Tommasini 2004; van de Veire and Degheele
1992). It has been well established that O. niger can
manage F. occidentalis in glasshouse cultivation of
sweet pepper (van de Veire and Degheele 1992). In
contrast, the role of Orius spp. as predators of thrips
on winter crops, such as faba bean, or on weeds, has
not been well studied in the Mediterranean region.
It is well known that weeds affect the popula-
tions of herbivorous insects and their natural
enemies by providing alternative food as well as
sites for mating, hibernation and shelter (Frank and
Reichhart 2004; Gurr et al. 2005; Landis et al. 2000;
Marshall and Moonen 2002). Vegetative buffers such
as weedy field margins are often recognized as
having an important place in sustainable agriculture
because of their role in an increasing functional
biodiversity and in encouraging cyclic movements of
predators between crops and the natural environment
(Landis and Wratten 2002; K.Winkler, Ph.D. thesis,
Wageningen Univ., 2005). The effect of weedy field
margins on thrips and their generalist predators in
relation to crop plants has not been studied. Also, the
relationships between winter–spring crops and
winter-flowering weeds on prey–predator dynamics
are not known. Faba bean flowers from early winter
to spring, and during this period various ecological
interactions between F. occidentalis and Orius
populations are expected. This crop system may also
allow us to understand the predation abilities of
Orius spp. on pestiferous insects of other arable
crops in the region.
The main objectives of this study were to assess the (a)
species composition of Orius spp. and other beneficial
insects on faba bean plots with and without weedy field
margins, (b) abundance patterns of F. occidentalis on
faba bean plots with and without weedy field margins,
and (c) preference of F. occidentalis and O. niger for
certain faba bean plant parts.
Phytoparasitica (2010) 38:313–325
Materials and methods
Study site This work was carried out on the Research
and Application Farm (RAF) of the Faculty of
Agriculture, University of Çukurova, during two
growing seasons, 2004/05 and 2005/06. The experi-
mental area (37° 01.809´ N; 35° 21.694´ E) has high
plant biodiversity, i.e., it is a polyculture area which
includes winter vegetables (onion, broccoli, lettuce
and red cabbage), wheat, and annual summer crops
(cotton and maize) grown in small plots. Various
citrus trees (lemon, orange, mandarin and grapefruit)
and olive trees are also grown there commercially.
Winter or annual summer crops are usually grown in
plots of 1.5 ha; citrus and olives are grown in plots of
2.2 ha and 1.5 ha, respectively.
Experimental procedure The experimental area of faba
bean (cv. ‘Lara’) was located near citrus (mandarin and
orange) and olive trees. Wheat and lettuce (plots of 50 or
100 m2) and other winter vegetable crops (plots of
200 m2) were also grown nearby. There were two
treatments: in one treatment, weeds in the margins and
inside the plots were regularly controlled by tillage;
and in the other treatment, no weed control was
conducted in plot margins but weeds inside the plots
were regularly managed. In the latter, weeds such as
Calendula arvensis L., Fumaria officinalis L., Lamium
amplexicaule L., Melilotus officinalis (L.), Ochtodium
aegyptiacum (L.), Senecio vernalis Waldst. & Kit and
Sinapis arvensis L. grew naturally in the margins of the
plots. The total size of the experimental area was
1,375 m2 (55 m×25 m). A split-plot design was used
with four replicates for each treatment. The size of
each plot was 100 m2 (10 m×10 m). Each plot with
weedy margins (PWWM) was bounded on all four
sides by an untreated strip of 20 m2 (10 m long×2 m
wide). Experimental plots were separated by a
5-m-wide strip of bare ground.
Faba bean was planted on 5 October 2004 and 15
October 2005. No pesticides were applied during the
course of the experiment.
Sampling of insects on faba bean and weeds
Sampling of insects started when faba bean and weed
species began to flower, because populations of both
Frankliniella spp. and Orius spp. occurred mostly on
the flowers of field crops such as cotton (Atakan et al.
1998; Atakan and Özgür 2004) and weeds (Atakan
Phytoparasitica (2010) 38:313–325
and Uygur 2004) in the region. Sampling of flowers
was reported to show the population trends of both
thrips and Orius spp. accurately (Shipp and Zariffa
1991; Shipp et al. 1992). Faba bean and weeds were
sampled from January to April of both years. Sample
size was five randomly selected plants per plot of faba
bean and of each of the four prevalent weed species,
C. arvensis, L. amplexicaule, O. aegyptiacum and
S. arvensis in the plots' margins. Insect species were
sampled by beating the plants vigorously over a white
plastic container (37×28×7 cm) for about 5 s.
Collected F. occidentalis and Orius species were
counted in the field using a hand lens. Most Orius
specimens were returned to the plots and a few were
taken to the laboratory for identification. Insects that
could not be identified in the field were collected and
kept in 60% ethanol.
In the laboratory, the thrips samples were trans-
ferred to vials containing AGA solution (10 parts
60% ethanol, one part glacial acetic acid, and one
part glycerin) and kept for 1 day. Thrips species were
slide-mounted and identified under a binocular
microscope.
Insect identification Thrips species were identified
by the author. Orius spp. were identified following
Önder (1982) and Tommasini (2004). Other preda-
tors or pest insect species collected were identified
by using reference material deposited at the Ento-
mology Laboratory of the Plant Protection Depart-
ment, Faculty of Agriculture, University of
Çukurova, Adana, Turkey. Representative adult
thrips were slide-mounted for identification. Imma-
ture thrips (thrips larvae) and nymphs of the Orius
specimens were pooled into a single category
because no comprehensive keys are available for
their identification.
Orius niger was the most easily distinguishable
species, being almost entirely black except for the
antennae and front tibiae or the knee and distal part of
the front femur, which are light yellow-brown. Body
color of Orius laevigatus (Fieber) was dark or light
brown. In O. laevigatus the legs were completely dark
or light yellow-brown. In some specimens, a yellow-
ish prothoracic femur and dark brown mesothoracic
and metathoracic femurs with yellowish distal parts
were observed.
Identified insect samples were counted separately
under a stereomicroscope with ×45 magnification.
315
Distribution of thrips and Orius on plant parts of faba
bean To determine the abundance of thrips and
Orius on two plant parts, plants were sampled
weekly on eight different dates in February (four
sampling dates) and March (four sampling dates) of
both years. Five plants were randomly selected from
each plot. Six leaflets or flowers (two leaflets or
flowers from each of three plant sections, upper,
middle and lower) from each plant were collected
from each section and kept frozen. For extraction of
insects, plant parts were submerged in 2% detergent
solution and agitated for 25 s. The solutions were
then poured through a standard testing sieve No. 80
(180 μm). Thrips and predators extracted from
solution were transferred into small vials containing
60% ethanol for subsequent processing. Each empty
bag was rinsed two times in the same way. The thrips
and predators were identified according to the
procedures detailed by Atakan (2006). Identified
insect specimens were counted separately under a
stereomicroscope with ×45 magnification.
Data analysis The abundance of other insect
species, including thrips and other predators, was
not evaluated because their numbers were too few
on most sampling dates. Data collected on thrips
larvae were not analyzed because their numbers
were very low on faba bean or weeds.
Effects of treatment and sampling dates on the
abundance of Orius spp. adults (pooled) and Orius
spp. nymphs (pooled) and F. occidentalis adults
were analyzed by General Linear Model (GLM)-
repeated measure statistical analysis. Weekly com-
parisons of the numbers of thrips and Orius on faba
beans throughout the sampling period between the
two treatments were done using Student’s t-test at
P < 0.05.
Numbers of F. occidentalis and Orius spp. on each
plant part (leaves and flowers) were pooled over
sampling dates and sampling years. Their mean
numbers were grouped by Tukey’s honestly signifi-
cant difference (HSD) test at P<0.05.
Seasonal comparisons of the numbers of thrips and
Orius on the weed species were quantified by one-
way analysis of variance (ANOVA) at P<0.05. Their
mean numbers were grouped by the Tukey’s HSD test
at P<0.05.
Relationships between the numbers of Orius and
F. occidentalis in four weed species and in faba bean
316 Phytoparasitica (2010) 38:313–325

were evaluated by quadratic regression analysis at species (O. niger and O. laevigatus) were collected
P<0.05. from S. arvensis and L. amplexicaule, whereas the
All analyses were performed using the Microsoft numbers of Aeolothrips collaris Priesner on four weeds
statistics program SPSS 15.0. (SPSS, 2006). were usually similar in both years (Table 2).

Species compositon of Orius on weeds and faba

Results
Predatory insect species on faba bean and weeds
Seasonal total numbers of predators identified in faba
bean plots and weeds naturally grown in the plot
margins are summarized in Tables 1 and 2. A total of
ten predaceous insect species were identified in
PWWM and plots with no weedy margins (PNWM)
(Table 1). Seasonal numbers of both Orius species in
the plots were clearly different while numbers of
coccinellids, chrysopids, mirids, staphylinids and pred-
atory thrips in abundance in both plots were usually
similar. Faba bean plots had rich insect fauna (Table 1)
as compared with the insect species composition found
in weed species, where only three predator species
were detected (Table 2). The majority of both Orius
bean. Orius niger was the main predatory bug on all
four weed species sampled on most sampling dates.
O. laevigatus was the most common on S. arvensis
and L. amplexicaule on some sampling dates in
March in both years. O. niger was the most prevalent
insect predator on faba bean on most sampling dates,
in all plots in both years. However, in March,
O. laevigatus was relatively more abundant.
Abundance of Frankliniella occidentalis and Orius on
weeds Trends in population dynamics of either total
adults of Orius or F. occidentalis in abundance in all
sampled weed species growing in plot margins were
similar in both years (Fig. 1a, b, c, d). Mean numbers
of F. occidentalis were very low on C. arvensis and
L. amplexicaule throughout the sampling dates in
both years (Fig. 1a, b). F. occidentalis were collected

Table 1 Numbers of predators identified and sampled in faba bean plots with or without weeds during the winter–spring period of
2005–2006 in Adana province, Turkey

Predatory insects [Order/family] Total no. of predatory insects on faba bean

Year 2005 Year 2006

Plot with Plot without Total Plot with Plot without Total
weeds weeds weeds weeds

Coccinella semptempunctata L. [Coleoptera/ 7 3 10 3 1 4

Coccinellidae]
Scymnus levaillanti Mulsant [Coleoptera/ 7 2 9 3 1 4
Coccinellidae]
Scymnus pallipediformis Goeze [Coleoptera/ 2 1 3 0 0 0
Coccinellidae]
Paederus litoralis Gravenhorst [Coleoptera/ 2 1 3 3 1 4
Staphylinidae]
Tachyporus sp. [Coleoptera/Staphylinidae] 6 2 8 25 14 39
Orius niger (Wolff) [Hemiptera/Anthocoridae] 74 33 107 174 83 257
Orius laevigatus (Fieber) [Hemiptera/Anthocoridae] 18 9 27 34 4 38
Orius nymphs 148 85 233 103 28 131
Campylomma sp. [Hemiptera/Miridae] 4 1 5 3 2 5
Chrysoperla carnea (Stephens) [Neuroptera/ 6 1 7 8 3 11
Chrysopidae]
Aeolothrips collaris Priesner [Thysanoptera/ 16 12 28 12 10 22
Aeolothripidae]
Total 290 150 440 368 147 515
Phytoparasitica (2010) 38:313–325 317

Table 2 Numbers of
predators sampled on weed Weed species Total no. of predators on weeds
species during the
winter–spring period of Aeolothrips collaris Orius niger Orius laevigatus
2005–2006 in Adana
province, Turkey 2005 2006 Total 2005 2006 Total 2005 2006 Total

Calendula arvensis 2 1 3 9 33 42 4 2 6
Lamium amplexicaule 1 3 4 48 55 103 13 19 32
Ochtodium aegyptiacum 8 12 20 12 18 30 1 6 7
Sinapis arvensis 21 19 40 67 62 129 16 25 41
Total 32 35 67 136 168 304 34 52 86

mainly from O. aegyptiacum and S. arvensis in both
years (Fig. 1c, d). Mean numbers of F. occidentalis
increased clearly on both weed species after the first
week of April. Thrips were very few or nonexistent
until the end of March in the presence of high
densities of total Orius spp. Population densities of
F. occidentalis on O. eagyptiacum and S. arvensis
increased steadily to considerable levels after late
March, when Orius individuals were absent or their
numbers were very low in both years (Fig. 1c, d).
Most adults of Orius spp. were detected on
L. amplexicaule and S. arvensis during the sampling

Fig. 1 Infestation levels of

adult Frankliniella occiden-
talis and Orius spp. on
weeds grown on the mar-
gins of faba bean plots:
Calendula arvensis a, Lam-
ium amplexicaule b, Ochto-
dium aegyptiacum c and
Sinapis arvensis d, in 2005–
2006
318 Phytoparasitica (2010) 38:313–325

Table 3 Regression analysis among mean numbers of Orius spp. and Frankliniella occidentalis in four weed species growing in the
margins of faba bean plots in Adana province, Turkey, in 2005–2006

Weed species Associations/years df R2 F P Equations

Calendula arvensis Orius adult and thrips adult/2005 2, 15 0.06 0.397 0.681 Y ¼ 18:715x2 þ 5:9127x þ 0:3558
Orius adult and thrips adult/2006 2, 16 0.25 2.269 0.143 Y ¼ 7:8802x2  2:9955x þ 0:3099
Lamium amplexicaule Orius adult and thrips adult/2005 2, 15 0.15 1.785 0.371 Y ¼ 1:2436x2  1:198x þ 0:1819
Orius adult and thrips adult/2006 2, 16 0.25 2.266 0.143 Y ¼ 1:092x2  1:5723x þ 0:4128
Ochtodium aegyptiacum Orius adult and thrips adult/2005 2, 15 0.30 2.636 0.113 Y ¼ 303:89x2  97:032x þ 0:5931
Orius adult and thrips adult/2006 2, 16 0.08 0.169 0.559 Y ¼ 49:705x2 þ 25:167x þ 1:6804
Sinapis arvensis Orius adult and thrips adult/2005 2, 15 0.76 23.854 0.0001 Y ¼ 54:632x2  66:693x þ 17:771
Orius adult and thrips adult/2006 2, 16 0.40 4.326 0.036 Y ¼ 15:369x2  21:728x þ 7:8689

dates in both years (Fig. 1b, d). In general, the
numbers of Orius individuals peaked on the weed
species in early March in 2005 and 2006.
Overall, the numbers of F. occidentalis and Orius spp.
(adults + nymphs) were significantly higher on S.
arvensis than on other weed species sampled in 2005
(F. occidentalis: F=655.562, d.f.=3, 76, P=0.0001;
Orius spp.: F=30.206, d.f.=3, 76, P=0.0001, respec-
tively) and in 2006 (F. occidentalis: F=469.236, d.f.=3,
76, P=0.0001; Orius spp.: F=12.882, d.f.=3, 76,
P=0.0001, respectively).
Seasonal population patterns of Orius spp. were
significantly compatible with the population fluctua-
tions of F. occidentalis only on S. arvensis in both
years (Table 3 and Fig. 2d; P<0.05). No relationship
was detected among the mean numbers of Orius spp.
and F. occidentalis on the other weed species sampled
(Table 3 and Fig. 2a, b, c).
Distribution of Frankliniella occidentalis and Orius
on two plant parts of faba bean. Frankliniella
occidentalis (adults) and Orius spp. (adults +
nymphs) were collected mainly from the flowers of
faba bean in PWWM (Fig. 3a) and in PNWM
(Fig. 3b) in both years. Orius nymphs were found
only on flowers, but their numbers were very low on
most sampling dates in all plots.
Abundance of Frankliniella occidentalis and Orius
populations on faba bean Plot type, sampling dates,
sampling years and various interactions had significant
effects on the abundance of F. occidentalis (Table 4).

Fig. 2 Relationships among

mean numbers of Orius spp.
and numbers of Frankli-
niella occidentalis in Ca-
lendula arvensis a, Lamium
amplexicaule b, Ochtodium
aegyptiacum c and Sinapis
arvensis d, in 2005–2006
Phytoparasitica (2010) 38:313–325 319

Fig. 3 Distributions of
adults of Frankliniella occi-
dentalis and Orius spp.
(mean±SE) on faba bean in
plots with a and without b
weedy margins, in 2005–
2006. Asterisk denotes the
level of significance; t-test
at *P<0.05, **P<0.001

Early colonization of F. occidentalis on faba bean was
detected in PWWM in both years (Fig. 4a). The
number of adult thrips peaked (1.21±0.22 per plant) on
14 March 2005 and 11 March 2006 (2.52±0.61 per
plant) in PWWM. In PNWM, the number of adult
thrips peaked (0.22±0.12 per plant) on 14 March 2005
and 23 March 2006 (0.61±0.30 per plant) (Fig. 4a). In
2005, the mean numbers of thrips in PWWM were
significantly higher than the numbers in PNWM on 21
February, 7 March and 14 March (t 38 = 3.230,

Table 4 Effects of plot

type, year and sampling Sources df MS F
date on population densities
of adult Frankliniella occi- Frankliniella occidentalis
dentalis and Orius spp. Plot type (with weeds and without weeds) 1 36.053 76.524 ***
(nymphs + adults) Year 1 8.670 18.402 ***
Plot type x year 1 4.083 8.667 **
Error 76 0.741
Date 14 6.907 14.089 ***
Date x plot type 14 2.050 4.181 *
Date x year 14 1.513 3.086 *
Date x plot type x year 14 2.883 5.882 *
Error (date) 1064 0.490
Orius spp.
Plot type 1 126.101 55.113 ***
Year 1 0.041 0.018 NS
Plot type x year 1 6.601 2.885 NS
Error 76 2.288
Date 14 14.388 7.413 ***
Date x plot type 14 13.403 6.605 ***
Date x year 14 17.253 8.889 ***
* P<0.05, ** P<0.01, *** Date x plot type x year 14 29.313 15.102 ***
P<0.001
Error (date) 1064 1.941
NS: not significant
320 Phytoparasitica (2010) 38:313–325

Fig. 4 The effects of weedy margins on the infestation levels of adult Frankliniella occidentalis a and Orius spp. b (mean±SE) on
faba bean in plots with and without weedy margins, in 2005–2006

P=0.0002; t38 =2.508, P=0.01; t38 =3.946, P=0.0003, P=0.001; t38 =4.328, P=0.0001, t38 =3.928, P=0.0003,
respectively; Fig. 4a). In 2006, mean numbers of thrips respectively; Fig. 4a).
in PWWM were significantly higher on 12 January, 11 Plot type, sampling dates, sampling years and
March and 30 March than in PNWM (t38 =3.321, various interactions had significant effects on the

Table 5 Regression analysis between mean numbers of Orius spp. and Frankliniella occidentalis in faba bean plots with weedy
margins or plots without weedy margins in Adana province, Turkey, in 2005–2006

Plot type Associations/years df R2 F P Equations

Plot with weeds Orius adult and thrips adult/2005 2, 15 0.22 1.767 0.212 Y ¼ 1:0533x2 þ 2:0675x þ 1:5536
Orius adult and thrips adult/2006 2, 16 0.12 0.944 0.414 Y ¼ 0:3618x2 þ 1:1617x þ 1:7071
Orius nymph and thrips adult/2005 2, 15 0.02 0.153 0.860 Y ¼ 0:1477x2  0:5624x þ 0:7421
Orius nymph and thrips adult/2006 2, 16 0.05 0.400 0.679 Y ¼ 0:1477x2 þ 22x þ 0:9667
Plot without weeds Orius adult and thrips adult/2005 2, 15 0.19 1.489 0.264 Y ¼ 96:667x2  14x þ 0:1333
Orius adult and thrips adult/2006 2, 16 0.29 2.758 0.100 Y ¼ 2:6138x2 þ 3:6013x þ 0:7868
Orius nymph and thrips adult/2005 2, 15 0.42 4.515 0.035 Y ¼ 68:330x2  14x þ 0:1333
Orius nymph and thrips adult/2006 2, 16 0.13 0.995 0.396 Y ¼ 0:4786x2  0:5754x þ 0:1415
Phytoparasitica (2010) 38:313–325
Fig. 5 Relationships be-
tween mean numbers of
Orius spp. and numbers of
Frankliniella occidentalis
on faba bean in plots with a
and without b weedy mar-
gins, in 2005–2006
abundance of Orius spp. (Table 4). Populations of
Orius spp. adults fluctuated throughout the sampling
period in all plots during 2005 (Fig. 4b). In 2005, the
numbers of adult Orius spp. peaked on 28 March
(3.21±0.21 per plant) in PWWM and on 11 April
(3.22±0.51 per plant) in PNWM. The numbers of
Orius adults in PWWM were significantly greater
than in PNWM on most sampling dates of January
and February in 2005 (P<0.001). The numbers of
Orius adults in PNWM were significantly higher
than in PWWM only on 4 April and 11 April in 2005
(t38 =2.367, P=0.023; t38 =4.681, P=0.0001, respec-
tively). In 2006, the numbers of adult Orius spp.
peaked on 30 March (3.24 ± 0.51 per plant) in
PWWM and on 19 January (2.42±0.42 per plant)
in PNWM (Fig. 4b). Mean numbers of Orius adults
in PWWM were greater on most sampling dates,
after excluding 2 March, 11 March and 30 March,
compared with numbers in PNWM. Mean numbers
of Orius adults in PNWM were significantly higher
on 9 February and 16 February (t 38 = 4.368,
P=0.0001; t38 =2.997, P=0.004, respectively) than
in PWWM (Fig. 4b).
Nymphs of Orius species could not be identified
to the species level in this study. However, most
nymphs probably belonged to O. niger because this
bug was the most prevalent predator on the weeds
and faba bean on most sampling dates in all plots
and in both years (Fig. 4b). The first nymphs of
Orius were detected on faba bean in the first or
321
second week of March in all plots. Mean numbers of
Orius nymphs peaked on 28 March in both PWWM
and PNWM in 2005 (2.93±0.33 and 1.22±0.31 per
plant, respectively), while in 2006, the peak was on
30 March (2.50±0.20 per plant) in PWWM and on
6 April (0.52±0.21 per plant) in PNWM. In 2005,
the numbers of Orius nymphs in PWWM were
significantly higher than in PNWM on 28 March
(t38 = 4.917, P= 0.0001) and 4 April (t38 = 3.596,
P=0.0009) (Fig. 4b). In 2006, numbers of nymphs
in PWWM were significantly higher on most
sampling dates (Fig. 4b; P<0.001).
Populations of Orius spp. (adults + nymphs) were
greater than those of thrips on all sampling dates in
both years, and numbers of thrips were very low or
negligible in the presence of high numbers of Orius
populations in both plots. Population densities of
thrips were very low when the numbers of Orius
nymphs or adults attained the highest levels in both
plots. Population trends of adults of Orius spp. in
abundance did not follow the population patterns of
F. occidentalis in both treatments in both years
(Table 5 and Fig. 5a, b). However, there was a
significant relationship between the population fluc-
tuation of adult F. occidentalis and the Orius
nymphal population only in PNWM in 2005 (Table 5
and Fig. 5b; P<0.05).
Other predators on faba bean Other predators that
were identified occurred sporadically and their
322
mean numbers in both plots of faba bean were very
low throughout the sampling periods. Coccinellid
and chrysopid predators were detected together
with aphids on the leaves in April. Most adults of
the staphylinid predators (mainly Tachyphorus sp.)
were observed on the leaves in both plots in
February 2006. Their numbers ranged between
0.05 and 0.90 individuals per plant in both plots in
February. The first females of predatory thrips A.
collaris occurred in both plots in the first week of
March. The population sizes of Aeolothrips in both
plots were relatively large in March when the
numbers of thrips were also relatively high. Mean
numbers of Aeolothrips in both plots were similiar,
ranging between 0.05 and 0.50 individuals per plant.
Relationships between numbers of Orius spp. or
Frankliniella occidentalis on weed species and faba
bean Mean numbers of Orius on S. arvensis were
significantly negatively associated with the numbers
on faba bean (2005: R2 = 0.42, F =4.403, d.f.=2,
14; P=0.041, Y=–4.4616x2 +5.79.34x+ 0.6607 and
2006: R2 = 0.42, F = 4.943, d.f. = 2, 15, P= 0.025;
Y= –7.3578x2–6.8758x + 3.224). There were also
significant negative relationships between the
numbers of Orius spp. on L. amplexicaule and the
numbers on faba bean in 2005 (R2 =0.64, F =10.913,
d.f. = 2, 14 P = 0.002; Y = –0.0073x 2 + 0.1173x–
0.1025) and in 2006 (R2 =0.51, F =6.802, d.f.=2,
15, P= 0.010; Y= –5.415x 2 + 7.01918x + 0.9879).
These significant and negative relationships mean
that the numbers of Orius spp. found on these two
weed species were decreasing while the numbers on
faba bean were increasing during March–April.
No relationship appeared among the mean numbers
of F. occidentalis on four weed species and the
numbers on faba bean in both years (P>0.05).
Discussion
The results show that Orius niger was the main Orius
species on both faba bean and weeds. Here, O. niger
appeared to be the more active predator in the hard
winter months in the region. This may be a result of
both year round availability of its thrips prey (Atakan
and Uygur 2005) and supporting weedy plants and
arable crops in the region, e.g. various vegetables and
fruits (Atakan 2008a, b) and field crops, including
Phytoparasitica (2010) 38:313–325
cotton (Atakan 2006). While O. niger was found to be
the most common anthocorid species in all parts of
Turkey, O. laevigatus was recorded mostly in the
southeastern part of Turkey (Önder 1982). O. niger,
O. laevigatus, Orius majusculus (Reuter) are the
commonly found anthocorid species in the Mediter-
ranean basin. However, the predominance of any of
these species may depend on location. For instance, in
Italy, O. niger predominated in the northwest,
whereas O. laevigatus predominated in the warmest
locations of the country (Bosco et al. 2008; Bosco and
Tavella 2008; Tommasini 2004).
In general, population trends of both thrips or
Orius spp. on the weeds were similiar but their
density levels were different in both years. S.
arvensis bore high numbers of both thrips and Orius
in both years. The data related to thrips and Orius
seasonal abundance on only S. arvensis allowed us
to establish statistical relationships in both years
(Table 3 and Fig. 2d). Previous studies reported that
weeds served as alternative sources of prey, and on
which Orius spp. and thrips populations interacted
very well, in Spain (Lacasa et al. 1995) and in Japan
(Kakimoto et al. 2006). In the present study, the
presence of Orius spp. on other major weed species,
C. arvensis and L. amplexicaule, which host no or
extremely low numbers of thrips, suggests that such
weeds may play some other role. For example, they
may provide nectar and pollen or shelter sites in
supporting Orius spp. regardless of the presence of
F. occidentalis.
Orius species were more abundant on flowering
weeds in the winter—early spring period (Fig. 1).
After mid-March the numbers of Orius species on
these weeds were very low. These findings are
similar to those of previous work done at the same
location (Atakan and Uygur 2004). The first Orius
nymphs were found only in the flowers of S.
arvensis in late April, whereas Orius nymphs were
present on faba bean in both plots in early March.
This may indicate that Orius spp. may prefer faba
bean plants as a site for egg-laying; therefore, they
may move from wild plants to faba bean in the early
spring despite the greater availability of flowers and
the presence of reasonable numbers of thrips prey
on most weed species (Atakan and Uygur 2005).
The relationship between abundance of Orius spp.
on the flowering weeds and faba bean that yielded a
significant negative correlation, may be evidence of
Phytoparasitica (2010) 38:313–325
movements of Orius from the weedy plants to the
faba bean. Many polyphagous predators often move
between arable crops and native habitats, thus
exhibiting cyclic colonization patterns (Bommarco
and Ekbom 2000; Murdoch et al. 1985; Thomas et
al. 1991). Such movements from native habitats to
arable crops include redistribution to early season
crop habitats where pests are beginning to be
established, enabling generalist predators to sup-
press agricultural pests (Ekbom et al. 1992; Ives and
Settle 1997; Murdoch et al. 1985).
Despite the large populations of Orius spp. in
PWWM, the numbers of F. occidentalis were
relatively higher there than in PNWM in both years
(Fig. 4). This may be due to the flowering of some
weed species bearing F. occidentalis. The numbers
of thrips on faba bean were low, especially in
PNWM, where the populations of Orius spp. were
less abundant in both years. In fact, thrips in both
plots of faba bean were less abundant throughout
the growing season. This may be due to the
unsuitability of this crop as a host for thrips growth
and reproduction, rather than to predation by Orius
species or the other sampled and identified preda-
tors. Individuals of Orius spp., especially nymphs,
were more abundant on faba bean grown in the
PWWM. This may be due to the presence of
flowering weed species, some of which host
considerable numbers of Orius spp. and thrips prey
(Fig. 1). Several studies have shown that vegetative
buffers that provide ecological benefits such as
nectar and pollen can improve the reproductive
success of natural enemies and that this may lead to
lower abundance of pest arthropods in the crop
plants (Coll 2009; Gurr et al. 2005; Heimpel and
Jervis 2005).
Mean numbers of both Orius spp. (adults +
nymphs) were greater than the numbers of F.
occidentalis on faba bean in both plots throughout
the sampling periods in both treatments (Fig. 4).
Orius populations were not related to thrips pop-
ulations in both faba bean plots, even where there
was a weak relationship between populations of
Orius nymphs and thrips in PNWM in 2005
(Table 5). This may indicate that Orius spp.
benefited more from the faba bean plants for plant
meal, shelter, mating and ovipositon sites than from
thrips prey or other insect prey species such as
aphids, whose numbers were low and were detected
323
sporadically in both years. Colonization of the
flowers by greater numbers of Orius spp. compared
to colonization of the leaves may be evidence for
this (Fig. 3). Faba bean, rather than other arable
crops, may be a good host plant in the winter–spring
period for growth and reproduction of Orius spp.
because no or few Orius spp. (mostly O. niger) were
detected on the flowers of various winter vegetables
(e.g. broccoli, garden cress, lectuce, red and white
cabbage, radish and spinach) grown in the same area
(Atakan 2008a). Faba bean also had a rich fauna of
predatory insects (Table 1). Similarly, Nuessly et al.
(2004) found that numerous beneficial insect
species, including Orius, visited faba bean plants
growing in southern Florida (USA).
In conclusion, controlling weeds can create a
problem in many cases because destruction of
weeds surrounding agricultural crops could cause
rapid dispersal of pestiferous thrips such as F.
occidentalis to crops and disturbance of the natural
enemies of thrips on the weeds. Proper management
of mowing, including timing, is recommended
during the entire season in order to protect and
encourage the development of insect predators
(Burgio et al. 2006). With respect to the results of
the present study, removing flowering weedy strips
bearing high numbers of predators (mainly Orius
species) by different control methods before early
April in the region could be destructive to beneficial
insects, and management procedures of field borders
should take into account the phenology of the
beneficials (Hodek and Honek 1996; Honek 1982).
On the other hand, control measures should be taken
for the weeds which host only a few predators but
high numbers of thrips (e.g. O. aegyptiacum).
Control of such weeds should be done at an
appropriate phenological stage, that is, before they
are colonized by thrips (e.g. before flowering),
because their destruction after bearing high numbers
of thrips may lead to rapid and serious infestation of
crop plants.
In this study, faba bean appared to be the more
attractive plant species, hosting considerable numbers
of Orius spp. in winter. Cultivation of faba bean in
agricultural locations, especially in monoculture areas
in the Mediterranean region or other countries with
similar ecological conditions, could be useful for
conservation and augmentation of beneficial insects,
including Orius spp.
324
Acknowledgments I would like to thank Dr. David Ben-Yakir
(Department of Entomology, Agricultural Research Organization,
The Volcani Center, Bet Dagan, Israel) for reviewing an earlier
draft of the manuscript and Murat Ölçülü (Institute of Basic and
Applied Sciences, Çukurova University, Adana, Turkey) for field
assistance. I would like also to thank three anonymous reviewers
for their valuable comments and suggestions which improved the
manuscript.
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