Biological Control
journal homepage: www.elsevier.com/locate/ybcon
Review
Laboratory of Agricultural Zoology and Entomology, Agricultural University of Athens, Iera Odos 75, 118 55 Athens, Greece
Laboratory of Ecology and Environmental Sciences, Agricultural University of Athens, Iera Odos 75, 118 55 Athens, Greece
a r t i c l e
i n f o
Article history:
Available online 23 April 2011
Keywords:
Conservation biological control
Predator
Heteroptera
Zoophytophagous
Omnivorous
Vegetable
a b s t r a c t
Conservation biological control (CBC) combines a diverse set of actions aiming to enhance pest regulation
by the provision of pest natural enemies preservation in the crop environment. This is accomplished
through appropriate implementation practices and the creation of infrastructures of non-crop vegetation.
On most annual crops, omnivorous heteropteran predators comprise the main assemblage of native natural enemies. Their populations, when protected from pesticides, have shown a high potential in biological control. Aiming to promote their functioning in CBC, efforts targeted to modify the crop environment
are focusing on the determination of their non-crop host plants that can support the predators persistence as well as the emigration of these predators to the nearby crops. Nonetheless, relatively few studies
seek to collect essential and adequately veried data for the practical implementation of CBC approaches.
In this review, the advances already developed are illustrated, the shortcomings are discussed and the
future research needs are highlighted in an effort to intensify efforts for CBC developments on annual
crops.
2011 Elsevier Inc. All rights reserved.
1. Introduction
Biological control (BC) is the regulation of pest populations as a
result of the activity of their living antagonists natural enemies
(predators, parasitoids and pathogens). BC offers a more sustainable and less expensive alternative to chemicals (DeBach and
Rosen, 1991; Bale et al., 2008). In fact, despite the high number
of releases of BC agents that have been performed globally, very
seldom have adverse effects been recorded (van Lenteren et al.,
2006).
Effectiveness in biological control necessitates conservative
measures in the crop environment targeting to the protection of
the natural enemies for a shorter or longer period. In classical BC,
a natural enemy is imported and released in a new area aiming
to provide permanent control of an invasive pest, whereas in augmentative BC periodical releases are performed. In both types, conservation refers to the protection of the released natural enemies
from adverse effects of pesticides, incompatible cultivation practices, but also favoring their establishment and impact by the
adjustment of the environmental conditions or providing food supplements, when possible.
Conservation biological control (CBC) is realized by the activity
of the natural enemies assemblages that already exist in the area
(Barbosa, 2003). CBC consists of premeditated actions for protecting and maintaining natural enemies (Rabb et al., 1976). In the CBC
Corresponding author. Fax: +30 210 529 4577.
E-mail address: dperdikis@aua.gr (D. Perdikis).
1049-9644/$ - see front matter 2011 Elsevier Inc. All rights reserved.
doi:10.1016/j.biocontrol.2011.03.014
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high disturbance in the established populations of the natural enemies (Altieri, 1991). The reduction of the populations of natural
enemies deteriorates BC potential and necessitates the colonization of the new crop from natural sources which may result in
weak or late establishment and nally in failures in pest control.
Additionally, intensication and high input investment on these
crops are not favorable for the natural enemies activity. Annual
crops do have some intrinsic attributes that may favor BC as a
desirable alternative to chemical control: (a) the pests in annual
crops are multivoltine and this increases their inherent potential
to develop resistance to chemicals, particularly in greenhouse
environments and (b) the risk for residual contamination of the
products of annual crops by chemical pesticides is increased since
these crops are harvested frequently at close intervals and their
products are mostly consumed fresh. In comparison to augmentative BC, CBC can be less costly because native natural enemies are
used and also safer because releases of exotic agents used in augmentative BC might be associated with environmental risks (van
Lenteren et al., 2006).
Due to their zoophytophagous feeding characteristics, the predatory Heteroptera may offer control of a wide range of pests and for
a prolonged period. Therefore, these predators have been considered as well equipped to cope with ephemeral crops on which prey
availability is variable in space and time (Pimm and Lawton, 1978;
Coll and Guershon, 2002). These characteristics align also with the
statement of Wiedenmann and Smith (1997) that most suitable
natural enemies on annual crops should keep the numbers of pests
at low levels and prevent their increase, since attempting to reduce
high pest populations would be much less efcient.
2.2. CBC in greenhouses: is it feasible?
Annual crops cultivation also includes the greenhouse environments. The application possibilities of CBC in greenhouses have
been considered as a debated issue because they are closed systems
that may provide barriers against the entry of the native natural
enemy fauna. However, there are some types of greenhouses that
may permit the entrance of natural enemies. For example, the plastic tunnels, as well as the open greenhouses of the Mediterranean
area, allow the indigenous natural enemies to move between the
crops, with positive impact in pest control (Albajes and Alomar,
1999; Bosco et al., 2008). In greenhouses with screened sides, the
entrance of natural enemies can be facilitated when kept open,
i.e., in hot conditions. The nethouses might be more suitable as
their sides can be opened. The greenhouses may also act as source
for predators to move out to colonize other close by crops or to nd
refuges in the surrounding area (Gabarra et al., 2004).
In modern greenhouses where the natural entry is prohibited
due to covering of the openings by insect-excluding nets, compatible measures with CBC could be the introduction of non-crop
plants infested with predators to act as sources or refuges of the
predators in the greenhouse (Sanchez et al., 2003). An alternative
approach could be the intended release of heteropteran predators
collected from the native ora of the neighboring area since they
can survive and reproduce on wild, naturally grown, non-crop
plants (i.e., Yano, 2008). Furthermore, at the end of the crop cycle,
the populations that developed on the greenhouse crop may be
collected and transferred to a newly established (greenhouse) crop.
This management or recycling of the natural enemies reservoir are within the context of CBC where sustainable management
of the natural resources in the wider scale and cooperation among
practitioners should be promoted.
Ultimately, in the greenhouse environments, IPM or BC programs are often disrupted or even collapsed by the introduction
of new, sometimes destructive pests (Desneux et al., 2010). The
heteropteran predators prey on a wide range of pests and have
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example, Orius insidiosus (Say) was the dominant predator on cotton crops and the most important mortality factor of Helicoverpa
zea (Boddie) eggs in Texas (Sansone and Smith, 2001). Also, Orius
spp. were the most abundant predators consisting 69.5% of the
predator numbers on non-chemically treated cotton (Greenberg
et al., 2009). In another example, Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) is common on cotton in South and Central
America and effective against noctuids in Brazil (Medeiros et al.,
2000; Lemos et al., 2001).
Deraecoris nebulosus (Uhler) (Heteroptera: Miridae) is commonly recorded in high numbers on cotton elds [14 nymphs
and adults/m/row (>137,000/ha)]. In laboratory tests, it readily
consumed cotton aphids and eggs of Heliothis virescens (F.) and rst
instar larvae of Spodoptera exigua (Hbner). Therefore, it appears to
be a naturally occurring predator that can have a signicant impact
on the pest populations on cotton crops (Snodgrass, 1991). In
northern China, the native Orius similis Zheng and Deraeocoris
punctulatus Fall largely supported the suppression of Bemisia tabaci
(Gennadius) (Homoptera: Aleyrodidae) (Lin et al., 2008).
Naranjo et al. (2002) showed that the application of conventional insecticides on cotton highly reduced the populations of major natural enemies, including G. punctipes, G. pallens (Stl), Orius
tristicolor (White), Nabis alternatus Parshley, Zelus renardii Kolenati,
in California and Arizona. The authors recognized that enhancing
the establishment of the predators early in the season is the key
issue in CBC and proposed raising the initial damage threshold to
delay the rst insecticide application, or alternatively, to use more
selective insecticides. Further study proved the compatibility of
selective insecticides with the conservation of natural enemies
on cotton (Naranjo et al., 2004).
G. punctipes acting together with O. insidiosus managed to keep
the populations of the sucking insect pests below the economic injury level at all the phenological stages of the cotton plants. The
major management practice that made this outcome possible
was the non-application of pesticides (Solangi et al., 2008).
The widely recognized importance of heteropteran predators on
cotton crops led to efforts to evaluate their potential in CBC using
other crop or non-crop host plants. Loya-Ramrez et al. (2003)
showed that the interplanted plants: canola (Brassica napus L.), alfalfa (Medicago sativa L.), hairy vetch (Vicia villosa Roth) and sanfoin
(Onobrychis viciifolia Scop.) supported high populations of heteropteran predators accompanied by a signicant migration toward
the intercropped cotton. In particular, Geocoris spp. attained highest densities on hairy vetch that increased towards the end of the
crop season. On the same plant, Nabis spp. were initially favored,
but densities were higher in alfalfa and sanfoin. On alfalfa, Orius
spp. had the highest density during the whole season. In comparison, treated cotton attained the lowest number of predators.
Plantings of Sorghum bicolor (L.) Moench (grain sorghum) beside cotton crops effectively favored the abundance of heteropteran predators but also their timely emigration to the cotton crops
(Fye, 1971). This approach was further explored by Prasifka et al.
(1999). They used marked and recapture methods to record the
movements of the predators including Orius in adjacent plantings
of cotton and grain sorghum. The outcomes revealed that cotton
gained approximately twice as many dispersing predators from
sorghum than those it lost, suggesting that the association is benecial. A mean speed of movement of about 2232 m/day was
estimated, indicating fast moving, showing that relatively large
elds may be colonized by predators ying from the edges, reducing the density of strip of sorghum needed inside the cotton crops
in CBC implementation. Therefore, the sorghum identities were
considered to make this scenario possible since: (a) both plants
share the same predator species, (b) sorghum is tolerant to insect
damage and seldom requires application of pesticides and thus
can act as a source of predators, and (c) cotton received inuxes
3.4. Legumes
On leguminous crops, the conservation of heteropteran omnivorous predators may largely contribute to the control of whiteies
or other pests but their contribution can be signicant against the
highly polyphagous pest, the southern green stinkbug Nezara viridula L. (Heteroptera: Pentatomidae). This pest causes economic
damage in several crops like cotton, tomato or eggplant. The control of N. viridula relies heavily upon insecticides and this is a major
constraint in the IPM programs on these crops (Knight and Gurr,
2007).
The oriental asopine, Amyotea malabarica (F.) (Heteroptera:
Pentatomidae), has demonstrated a strong preference for N. viridula over lepidopterous prey (Singh et al., 1973). A number of other
heteropterans, mainly reduviids, have been noted to act as predators of N. viridula with biocontrol potential (Ambrose, 1999;
Grundy and Maelzer, 2000a).
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L., had been introduced, providing also a refuge for the predator
when prey on tomato was scarce. Establishment of the predators
populations on these non-crop plants would largely contribute to
the early colonization of the following greenhouse crop and the
effective control of whitey pests (Sanchez et al., 2003). In tomato
unheated greenhouses, Arn et al. (2000) showed that tobacco
plants can serve as a refuge for the mirid predators during winter
that may enhance the timely and efcient colonization of the next
spring tomato crop.
As a measure within the frame of CBC, it has been also suggested that the farmers could collect predators on crops plants at
the end of the crop cycle or on major non-crop plants for subsequent release in greenhouses (Lykouressis et al., 19992000;
Roditakis et al., 2003; Cano et al., 2009).
3.6. Other crops
Strawberries are infested with several species of thrips worldwide. Coll et al. (2007) demonstrated that releases of O. laevigatus
were not justied in insecticide-free strawberry crops grown in
low tunnels in Israel, since the thrips populations were successfully controlled by native O. niger and O. albidipennis. Bosco et al.
(2009) evaluated the thrips control in plots where articial releases
of Orius bugs were performed in comparison to greenhouses where
chemical control was applied. The results showed that both treatments were equally effective. Furthermore, in the biological control plots, the native O. niger prevailed over the released O.
majusculus. Therefore, native predators play a signicant role in
thrips control on strawberries and should be conserved. In greenhouse roses, introduction of Tagetes plants during their owering
period may increase the effectiveness of the released O. insidiosus
and this technique worth further evaluation (Bueno et al., 2009).
4. Conclusions and suggestions
The omnivorous predators comprise distinct functional groups,
commonly present on annual crops with a signicant contribution
in natural pest control. This evidence lies in support of the view
that effectiveness in pest regulation may not be positively associated with increased complexity but simplied natural enemy communities may be highly effective (Bianchi et al., 2010).
The high potential of heteropteran predators in CBC can be largely attributed to their (a) wide prey range covering almost the entire spectrum of pests and thus, when conserved, they can offer
effective protection of the crop, (b) zoophytophagous feeding habit
so that they can survive for a longer or shorter period on crop
plants without prey and already existing populations can suppress
pest numbers when the pests appear and start to increase, keeping
them below economic threshold, and (c) populations can persist
and increase on certain non-crop plants that act as refuges and natural reservoirs enabling their persistence and the colonization of
the new crop.
Thus far, the high proportion of this encouraging evidence is derived from studies where the only means to enhance the effectiveness of the naturally occurring populations of these predators was
the drastic reduction of pesticide use. Although it reects the high
potential of CBC if these predators are conserved, this is only the
rst step towards the development of CBC implementation
strategies.
Recent research efforts have been devoted to systematically
explore and evaluate non-crop host plants of omnivorous
heteropteran predators as elements of CBC agro-environment
schemes in annual crops. In this regard, evidence from related research highlights the areas where future efforts should be directed
to (a) correctly identify the predator species on each candidate
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