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Current status and potential of conservation biological control for agriculture

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DOI: 10.1016/j.biocontrol.2012.11.010

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 Biological Control 65 (2013) 152–167

Contents lists available at SciVerse ScienceDirect

Biological Control
journal homepage: www.elsevier.com/locate/ybcon

Review

Current status and potential of conservation biological control for agriculture

in the developing world
Kris A.G. Wyckhuys a,⇑, Yanhui Lu b, Helda Morales c, Luis L. Vazquez d, Jesusa C. Legaspi e,
Panagiotis A. Eliopoulos f, Luis M. Hernandez g
a
CIAT, Hanoi, Viet Nam
b
State Key Laboratory for Biology of Plant Diseases and Insect Pests, Chinese Academy of Agricultural Sciences, Beijing, China
c
El Colegio de la Frontera Sur ECOSUR, San Cristóbal de Las Casas, Mexico
d
Instituto de Investigaciones de Sanidad Vegetal INISAV, La Habana, Cuba
e
United States Department of Agriculture, Agricultural Research Service, CMAVE/Florida A&M University-Center for Biological Control, Tallahassee, FL, USA
f
Technological Educational Institute of Larissa, Larissa, Greece
g
Universidad Nacional de Colombia, Palmira, Colombia

h i g h l i g h t s g r a p h i c a l a b s t r a c t

" A total of 390 literature records from

53 different crops and 53 nations
were found.
" Most research focused on habitat
management and changes in
disturbance regimes.
" No CBC records were found for
several key staple crops and cash
crops.
" 70% of pests with high incidence of
insecticide resistance have been
overlooked.
" Many nations have high insecticide
use and import, but little CBC
research attention.

a r t i c l e i n f o a b s t r a c t

Article history: Conservation biological control (CBC), often described as the field of biological control with the greatest
Received 9 July 2012 potential for use in developing world agriculture, has received only marginal, scattered research attention
Accepted 28 November 2012 outside Western Europe or North America. As a consequence, pesticide overuse remains rampant in many
Available online 10 December 2012
cropping systems, while in others, a complete lack of safe, affordable and effective pest control options
leaves farmers vulnerable in face of herbivore attack. In this study, we describe the current status of
Keywords: CBC research in a wide variety of agro-production systems outside North America, Australia, New
Conservation biological control
Zealand, Japan and Western Europe. We summarize information on (1) a variety of CBC themes related
Natural enemies
Africa
to natural enemy biology and ecology, (2) factors that either disrupt or enhance natural enemy efficacy,
Developing world and (3) field evaluation of CBC schemes. A total of 390 CBC-related literature records from 53 different
Poverty crops were considered. Most records were from China, Brazil, or Cuba, while no CBC references were
Food security found from several developing countries. CBC research primarily focused on habitat management, with
71 records on general habitat manipulation and 80 records on the effects of inter-or cover-crops on nat-
ural enemy abundance or efficacy. The effects of deliberate modification of disturbance regimes, through
alterations in pesticide use or tillage, on natural enemies were well-characterized in many cropping sys-
tems. For each of the CBC themes, research progress was assessed and opportunities were identified to
translate current findings into practical solutions. On a crop level, most research was targeted at rice,

⇑ Corresponding author. Address: Km 17, Recta Cali-Palmira, Apartado Aéreo 6713, Cali, Colombia. Fax: +57 2 4450073.
E-mail address: kwyckhuys@hotmail.com (K.A.G. Wyckhuys).

1049-9644/$ - see front matter Ó 2012 Elsevier Inc. All rights reserved.
http://dx.doi.org/10.1016/j.biocontrol.2012.11.010
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167 153

maize and cotton. No CBC records were found for key staple crops such as yams, taro, sago or breadfruit;
fruits such as papaya, pineapple and avocado; or forage crops. Also, millet, lentils, barley and plantain, all
crops grown mainly in the developing world, received limited CBC research attention. CBC research has
been done on myriad arthropod pests, including species with high levels of insecticide resistance such as
Chilo suppressalis (Lepidoptera: Crambidae) and Helicoverpa armigera (Lepidoptera: Noctuidae). However,
almost 70% of pests with high incidence of insecticide resistance have been overlooked. Lastly, we con-
trast country-specific CBC research advances with the national level of insecticide use and importation,
and identify lucrative opportunities for countries to save funds through targeted research investment.
Based upon our delineation of the current status of CBC, we indicate potential for well-orchestrated regio-
nal research projects to pursue higher levels of CBC integration into current pest management schemes.
This work constitutes a first step in drawing a roadmap for developing-world research that provides local
farmers with safe, low-cost means to control damaging insect pests, safeguard harvests and secure their
livelihoods.
Ó 2012 Elsevier Inc. All rights reserved.

1. Introduction management through enhancement of biological control services

Arthropods provide many valuable ecosystem services, includ-
ing the natural control of agricultural pests (Daily, 1997). In the
United States alone, the annual value of these biological control
services is estimated at $4.5 up to $17 billion (Pimentel et al.,
1997; Losey and Vaughan, 2006), and this value may still be much
higher for small-scale agriculture in the developing world. Such
natural pest control services are delivered through a diverse com-
munity of arthropod predators, parasitoids and entomo-pathogens
that are present in the vegetation in or around farm fields. The
deliberate manipulation of agro-ecosystems to enhance the sur-
vival, fitness, and behavioral performance of these natural enemies,
and to improve their resulting pest control action, is termed con-
servation biological control (CBC) (Barbosa, 1998; Landis et al.,
2000). Through vegetation manipulation, CBC can provide control
of both primary and secondary pests, while reducing the likelihood
of pest outbreaks and resurgences (e.g., Naranjo and Ellsworth,
2009). Also, CBC practices do not lead to the development of insec-
ticide resistance, a process that prevents closure of yield gaps in
many developing world crops (Godfray et al., 2010). Lastly, CBC
does not bring about human health risks, as compared to many
chemically-defined insecticides.
Despite being one of the oldest forms of pest control, with re-
cords dating back to 300 BC (see Huang and Yang, 1987), CBC
has only received limited attention (Ehler, 1998). With the onset
of the pesticide era, chemical insecticides came to replace the ac-
tion of naturally occurring natural enemies. By displacing natural
biological control agents from farm fields, pesticides thus quickly
divorced agricultural production from ecology (Robertson and
Swinton, 2005). Similarly, the spectacular achievements with
importation biological control projects in the 1980s may have re-
duced interest in CBC. However, the volume of CBC research has
steadily increased and some related tactics are being adopted in
cropping systems in Europe and North America (e.g., McLeod
et al., 2004; Jonsson et al., 2008; Naranjo and Ellsworth, 2009;
Nilsson, 2011). Despite these evolutions, the potential for the
management of pest insects using CBC has yet to be fully exploited,
especially for control of indigenous crop pests that have large
natural enemy complexes.
In many parts of the developing world, CBC is believed to be
the area of biological control with the greatest scope for pest con-
trol in multiple crops (e.g., Waage and Schulthess, 1989; Yaninek
and Cock, 1989; Tamo et al., 1997; Gurr et al., 2011). Lower con-
sumer expectations regarding cosmetic standards for harvested
produce, relatively diverse agro-ecosystems, and the low resource
base supporting local production systems all make CBC an attrac-
tive pest management solution for local farmers (Neuenschwander,
2010). In systems where pest losses are high, better pest
can raise crop yields more than most other types of crop research
(Pretty et al., 2011). However, a range of problems impede the
development and implementation of CBC in developing countries.
Often, adequate ecological information describing the action of
indigenous natural enemies and an assessment of ways to exploit
them is missing (Yaninek and Cock, 1989; Legg et al., 2003). Also,
instead of being seen as an opportunity to advance CBC, the pre-
dominance of generalist predators is frequently thought an imped-
iment to effective biological control (Cherry et al., 2003).
Consequently, pesticide use in many parts of the developing world
remains high and interferes with natural biological control ser-
vices. In the Latin American horticultural sector, for example,
dependence on pesticides is at worrying levels, with the bulk of
growers naming insecticides as their sole pest management tool
(Nunes et al., 2005; Wyckhuys et al., 2011). In other systems,
farmers seem to be falling back into the pesticide treadmill. The
classic Peruvian Canete Valley success, in which insecticide appli-
cations in cotton fields were reduced from 16 to 2–3 per crop
(Doutt and Smith, 1971; Barducci Boza, 1972), had completely
degenerated within 30 years, with a return to pesticide-dominated
pest control (Way and van Emden, 2000). Even in Southeast Asia,
where increased understanding of agro-ecosystem functioning led
to far-reaching management and policy changes in the 1990s, pes-
ticide use is increasing and associated pest problems are mounting
once again (Bottrell and Schoenly, 2012). For those systems, there
is a need to seriously assess the potential of CBC as a complemen-
tary or alternative pest management tactic. In other parts of the
developing world, such as Sub-Saharan Africa, farmers do not have
the financial means to use insecticides and insects continue to
cause vast yield losses and imperil food security (Neuenschwan-
der, personal communication). In those systems, CBC could consti-
tute an equally attractive pest management option, representing a
low-cost means to promote resident natural enemies and contrib-
ute to pest control.
In some countries, such as Cuba, CBC has received a fair amount
of research attention and occupies a prominent place in pest man-
agement schemes (Vázquez et al., 2008). Also, in a small set of sub-
sistence crops, farmers have historically learned to manage their
pests with limited use of insecticides (Jago, 1991; Way and van
Emden, 2000; Wyckhuys and O’Neil, 2007a). These cases should
be documented to help guide the development of CBC practices
for other crops in the tropics and in the developed world.
Agricultural production systems tend to lower the abundance
and efficacy of natural enemies through increased levels of distur-
bance (i.e., pesticide use, tillage), simplification of landscapes, or
use of monocultures (Letourneau, 1998). CBC tactics, as tools to
correct these problems, have been categorized into those that re-
duce natural enemy mortality, provide supplementary resources
154 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
used by natural enemies, control secondary enemies, or manipu-
late host plant (or habitat) attributes to the benefit of natural ene-
mies (Rabb et al., 1976; van den Bosch and Telford, 1964). Initial
literature reviews show that CBC research has been carried out in
all of the above areas in the developing world, but to date this
information has not been compiled.
Here we review CBC advances in regions outside of North Amer-
ica, Australia, New Zealand, Japan and Western Europe. We de-
scribe its current status in a wide variety of agro-production
systems, by drawing on less accessible literature and work pub-
lished in languages other than English, which is less widely known.
The resulting information is organized along the following themes
(see Fig. 1): (1) elucidation of the role of non-prey foods in sustain-
ing natural enemies; (2) use of artificial food sprays; (3) on-farm
habitat manipulation; (4) deliberate modification of disturbance
regimes, with emphasis on tillage and pesticide use; (5) identifica-
tion of alternative host or prey items; (6) characterization of the ef-
fect of plant varietal traits on natural enemy performance and
survival; and (7) examples of actual use of CBC at the farm or land-
scape level.
In this exercise, we limited ourselves to records related to
arthropod natural enemies, and to their effect in controlling field
populations of agricultural pests. No records were included on nat-
ural enemy efficacy and life history (when fed prey items), as these
do not exclusively support CBC. Although this review is intended to
be as complete as possible, we acknowledge that some valuable re-
search may have escaped our attention. Despite its potential limi-
tations, this review should help identify opportunities to promote
pest management in the developing world, while providing a basis
for a more intensive cross pollination of CBC research globally.
Fig. 1. Focal themes of the manuscript, as identified within different sections of an existing conceptual framework for conservation biological control research (see Naranjo,
2001).
2. Assessment of the effect of non-prey foods on natural enemy
fitness
The importance of consumption of plant-based foods by natural
enemies was first reported by Thorpe and Caudle (1938), who re-
ported nectar consumption by parasitoids of the pine shoot moth
Rhyacionia buoliana. Gradually, researchers have come to acknowl-
edge that non-prey foods such as nectar, honeydew or pollen influ-
ence many life processes critical to the success of parasitoids and
predators alike (Wackers et al., 2005; Lundgren, 2009). These plant
compounds may supplement prey diets or act as alternative foods
that sustain natural enemies under sub-optimal conditions. For
many natural enemies, a diversity of plant-based foods can in-
crease many fitness parameters and consequent biological control
efficacy.
A total of 39 records were found in which a comparative assess-
ment was made of natural enemy performance on different plant-
based food resources, with the oldest record being work on winter
moth Operophtera brumata parasitoids in Uzbekistan (Appendix 1;
Eremenko, 1971). The majority of studies (n = 16) did not specify
the cropping system, while systems that received relatively more
attention were cassava (5), maize (3), and crucifer vegetables (2).
Most records were found for China (9), but Colombia (4) and Egypt
(4) were also well represented. A total of 9 studies dealt with brac-
onid wasps, nine with phytoseiid mites and four with earwigs (For-
ficulidae). The following food sources were tested: mixed flower
honey (16), pollen (14), synthetic mono-or polysaccharides (8), flo-
ral nectar (7), honeydew (5), and fungal spores (2). One record de-
scribed the role of other plant-based foods such as maize grains,
fruit juice or wetted raisins. Parameters that were computed for
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
the different food sources were longevity/survival (24), fecundity
or reproductive output (11), overall development (8), body nutri-
ent levels (4), parasitism (3), and egg load or maturation (2). Only
single records were found for resource consumption level and
duration of the oviposition period.
With exception of the work on cassava phytoseiids in Benin,
Brazil and Colombia (e.g., Gnanvossou et al., 2005; Onzo et al.,
2005), no evidence was found of sustained research on natural en-
emy nutritional ecology in a particular cropping system. However,
the consumption of pollen by earwigs in cereal crops has been re-
searched on separate occasions in Latin America and Africa (e.g.,
Boukary et al., 1997; Sueldo de Escano and Virla, 2009). Possibly,
this work was guided by the high abundance and conspicuousness
of these predators in several crops and initial US-based research on
pollen-dependence in the earwig Doru taeniatum (Jones et al.,
1988). Lastly, a multi-country initiative is currently assessing the
role of plant-based food sources for natural enemies of several
key rice pests in Southeast Asia (Gurr et al., 2011, 2012).
Pollen and carbohydrates are steadily gaining recognition as key
food sources for natural enemies. Aside from maize pollen, the rel-
ative nutritional value of pollen from several other crop and non-
crop plants has been determined (e.g., Pu et al., 1991). While honey
has been widely researched as a high-quality food source, isolated
initiatives from Chile, China, Turkey, Philippines, and Uzbekistan
have also assessed the role of floral nectar. A broad set of flowering
plants, such as soybean, Daucus carota, Brassica parachinensis, Vicia
angustifolia or wildflowers (e.g., Solidago altissima, Erigeron annuus)
were found to enhance key life history parameters of various
parasitoids. Nevertheless, aside from exploratory work in South
America on the importance of nectar composition and floral
architecture (Chalcoff et al., 2006), much remains to be done to
properly select plant species for habitat diversification schemes
that benefit natural enemies (e.g., Patt et al., 1997; Wackers,
2004; Fiedler and Landis, 2007). As is well documented in Europe,
New Zealand or North America, floral resources not only improve
fitness of natural enemies but may also benefit the pest (Baggen
et al., 1999; Lavandero et al., 2006; Wackers et al., 2007).
This has also been recognized in the developing world. In
Brazil, buckwheat Fagopyrum esculentum nectar was consumed
by the parasitoid Mirax sp. (Hymenoptera: Braconidae) but was
equally exploited by the coffee leaf miner, Leucoptera coffeella
(Lepidoptera: Lyonetiidae) (Rosado, 2007). Along the same lines,
floral nectar supplied for the benefit of natural enemies favored
the wheat armyworm, Pseudaletia sequax (Lepidoptera: Noctuidae)
(Marchioro and Foerster, 2012).
The effect of extra-floral nectar, an omnipresent food in several
cropping systems (Lundgren, 2009; Nicolson et al., 2010), has only
recently been assessed for phytoseiids, lacewings, and whitefly
parasitoids in Benin, Brazil, and Colombia, respectively. Although
some baseline work has been done on cassava extra-floral nectar
(e.g., Toko et al., 1994), the nutritional value of this resource from
other crops such as pulses has not been determined. A sound
appreciation of its role may be essential to manipulate resident
natural enemy populations, or help design pest-suppressive poly-
cultures. Overall, insights into the natural enemy’s food require-
ments appear to have been primarily used to fine-tune their
(laboratory) mass rearing instead of improving their efficacy under
field conditions (Chen et al., 2010). A rare exception is the exem-
plary way in which insights in earwig nutritional ecology have
helped define habitat manipulation schemes in East Africa
(Boukary et al., 1997; Koji et al., 2007). In the vast majority of
cropping systems, however; laboratory assays wait to be trans-
ferred to semi-field trials and management recommendations.
Natural enemy groups such Araneae, Carabidae, Formicidae,
Syrphidae or predaceous heteropterans have not received any
attention, while other groups such as Coccinellidae, Tachinidae or
155
aphid parasitoids have been frequently investigated. A large
literature exists on the interactions between ants and extra-floral
nectaries in natural habitats of the tropics (e.g., O’Dowd, 1979;
Cuautle and Rico-Gray, 2003). However, this wealth of information,
poorly known outside ecology circles, waits to be used for the de-
sign of crop diversification schemes that enhance natural control.
As a promising development, research in China and Turkey has
started to assess the effect of wildflower nectar on the life history
of Trichogramma spp., undoubtedly one of the most widely used
groups of wasps in pest control (Tuncbilek et al., 2010; Guo
et al., 2011).
3. Elucidation of associations with alternative hosts, food and
prey items
Earliest records on the importance of alternative hosts for nat-
ural enemies were by Hardy (1938). Since then, research has been
conducted on identifying alternative hosts, prey and food items in
many cropping systems (e.g., Agusti et al., 2003; Isaacs et al., 2009).
In our literature revision, we encountered a total of 51 references
identifying such resources (Appendix 2). A total of 20 studies iden-
tified alternative hosts or host plants, while other resources such as
non-prey food (10), alternative prey (4) or refuges (3) received less
attention. In a total of 12 studies, a broad community of parasitoids
was studied, while other natural enemy guilds that received major
attention were Coccinellidae (8), Braconidae (5) or pirate bugs (4).
In most cases, researchers studied natural enemy associations with
plants present in natural habitats, such as common weeds, shrubs
or (fruit) trees.
In many studies, high natural enemy abundance was considered
indicative of trophic linkages to a given plant species, but relative
importance of such linkages was rarely assessed (but see
Soroushmehr et al., 2008). No studies were encountered in which
nectar consumption on a given plant species was proven, e.g.,
through use of biochemical assays (e.g., Wyckhuys et al., 2008).
Only one record was found where pollen grain analysis was used
to identify plant associations for a given natural enemy (Medeiros
et al., 2010). A more in-depth elucidation of the nature of those
trophic linkages or refuge provision with certain plants could help
define appropriate conservation practices.
In a similar way, advances have been made in the identification
of alternative host items, with sometimes unexpected results such
as the identification of native trees as alternative hosts for parasit-
oids of the cowpea pod borer Maruca vitrata (Tamo et al., 2002).
These studies not only pinpoint plants necessary to sustain certain
natural enemies, but also plants that provide the pest with refuges
from natural enemy action (e.g., Morrill and Almazan, 1990). One
critical drawback in many of the studies is the lack of identification
of temporal patterns in natural enemy abundance on such alterna-
tive host (or food source). Long-term studies from Israel, for exam-
ple, show that it is essential to understand when alternative host
plants such as Rhamnus sp. shrubs are important to predatory
anthocorids (Shaltiel and Coll, 2004). Similar limitations have been
identified for studies on alternative host items. Although the iden-
tification of alternative food or host resources constitutes a neces-
sary first step in the research process, long-term focused studies
are needed to help guide the definition of CBC schemes and their
proper incorporation in cropping systems.
With exception of some Cuban and African studies (e.g.,
Schulthess et al., 2001; Cherry et al., 2003; Kahuthia-Gathu et al.,
2008), many initiatives remain isolated and miss follow-up re-
search. For example, Beingolea (1959) elegantly explored plant
associations of several key predators and parasitoids in Peruvian
cotton agro-ecosystems, and stressed the importance of taking into
account those ecological relationships to define sustainable pest
management programs. Fifty-three years after this publication,
156 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
however, no evidence was found of changes in management prac-
tices that derived from this study. In contrast, an identification of
alternative host plants for the African gall midge Orseolia oryzae
led to an evaluation of habitat manipulation schemes, and serves
as an example that different approaches are possible (Nwilene
et al., 2008). Also, separate research initiatives could easily guide
studies in other parts of the developing world. For example, find-
ings from Mexico or Brazil that native fruit trees help conserve
fruit fly parasitoids (Figueroa de la Rosa et al., 1998; Carvalho
et al., 2010) should constitute a basis for biological control of
native fruit flies in Thailand and Malaysia, amongst others (see
Chinajariyawong et al., 2000).
4. Determination of the role of artificial food supplements
The potential of artificial food sprays to increase abundance and
impact of natural enemies has been recognized for over 40 years
(Wade et al., 2008). In our literature revision, a total of 19 refer-
ences evaluated the effect of artificial food supplements on natural
enemy abundance and efficacy, with the oldest record being the
mention of yeast sprays to enhance aphid predation by Chrysopa
sp. (Appendix 3; Trujillo and Altieri, 1990). Even earlier, sugar
sprays were briefly mentioned for use in Indian pulse crops (Iswer-
an & Sen, 1972). Records were found for a range of crops, with
maize (3) and citrus (2) having received the most attention. Crops
included principally vegetables and fruits, such as tomato, almond,
peppermint or cocoa. In seven cases, target pests were not speci-
fied, while homopterans such as aphids, whitefly or membracids
were the focal pests of six studies. Ants were studied in a total of
12 studies, while other groups included Chrysopidae (3), Coccinel-
lidae (2), Vespidae (1), Tachinidae (1), Syrphidae (1), Trichogram-
matidae (1), Phytoseiid mites (1) or Anthocoridae (1). A diverse
range of food supplements were described, covering both carbohy-
drate (i.e., sucrose solution, molasses, honey) and protein sources
(i.e., pollen, protein hydrolysate, milk, powdered fish, intestines).
All studies with lacewings evaluated the role of yeast or pro-
tein + sugar mixtures. The main effects of the artificial food provi-
sion were increased abundance or sustained ant colony presence
(9) and reduced tending of homopterans (3). When describing
the effect of sugar sprays on ant-hemipteran interactions, no men-
tion was made of increased predation (but see Carabali-Banguero
et al., in press).
Although sugar sprays have been shown to boost efficacy of
Trichogramma spp. under field conditions (Yu and Byers, 1994),
their use was only recently reported from Pakistan (Ahmad et al.,
2011). Similarly, even though food sprays greatly benefit Coleop-
tera (Wade et al., 2008), no research has specifically focused on this
group. One possible reason for this lack of attention could be con-
cern over the viability of costly broadcast sprays of sucrose solu-
tions, especially for small-scale farmers. Nevertheless, sugar-rich
industry waste products such as molasses could be a valuable
alternative to refined sugar (see Perfecto and Castiñeiras, 1998;
Choate and Drummond, 2011) and make this more affordable. That
sugar sprays have potential even for small-scale farmers is exem-
plified by Colombian research, showing that sugar sprays can be
combined with selective insecticides to increase whitefly biological
control in cash crops such as tomato (Hernandez et al.,
unpublished).
In addition to external applications of sugar, two records were
found of farmers’ intentional infestation of their crop with (non-
pestiferous) hemipterans, as sources of sugar-rich honeydew for
the predatory ants Dolichoderus bituberculatus Mayr and D. thoraci-
cus Smith (Graham, 1991; Ho and Khoo, 1997). Through such pest
introductions, farmers secure a continuous availability of carbohy-
drates and a resulting high activity and intensive foraging by pred-
atory ants in cocoa orchards. Lastly, broadcasting of maize pollen
as supplemental food in Israeli avocado orchards increased natural
enemy abundance and pest control (Maoz et al., 2009). Other
promising low-cost approaches to boost predator abundance and
action include the deployment of powdered fish bones, as de-
scribed from work in Uganda (Sekamatte et al., 2001b, 2002).
Based upon this experience, an evaluation of similar low-cost
protein baits to increase abundance and foraging of other
ground-foraging ant predators such as Solenopsis geminata,
Wasmannia auropunctata or Pheidole spp. may carry particular
promise. Often, literature references mention the use of artificial
food supplementation as farmer innovations (e.g., Van Mele and
Cuc, 2000; Bentley, 2006), but do not document its effects as result
of scientific experimentation. The only exception may be the
validation of sugar-sprays and the burying of dead animals, as a
farmer innovation, in Honduran and Ugandan small-scale maize
fields (Canas and O’Neil, 1998; Sekamatte et al., 2001b). A more
formal evaluation of some of these practices could validate artifi-
cial food supplementation in various other cropping systems
(e.g., Mensah, 1996; Cook et al., 2007; Hazarika et al., 2009).
5. Effects of structural habitat manipulation on resident natural
enemies
The agricultural crop itself undoubtedly has the largest direct
effect on natural enemy abundance and efficacy. However, it was
not until the 1950s that European researchers recognized that
the ecological infrastructure of agricultural habitats could be
manipulated for the benefit of these natural enemies and biological
control (Gyorfi, 1951). Since then, research has come to show that
habitat management and plant diversification schemes, such as
intercropping or the establishment of flowering plants, can greatly
enhance natural enemy abundance (Letourneau et al., 2011). How-
ever, recent research shows that the efficacy of those schemes is
still tied to landscape complexity (Woltz et al., 2012). While the
science of habitat manipulation only became popular in the
1990s (Landis et al., 2000), it has advanced considerably during
the past decade, with ecological engineering taking shape as a sep-
arate discipline dedicated to its promotion (Gurr et al., 2004). This
has resulted in myriad promising technologies such as the use of
shelter habitats, beetle banks (McLeod et al., 2004) and wildflower
strips (Lavandero et al., 2005; Pfiffner et al., 2009), with some of
those approaches increasing pest control in specific cropping
systems.
In the developing world, habitat manipulation could easily be
incorporated in local agro-production systems, already being an
integral feature of Chinese agriculture (Olkowski and Zhang,
1998) and having been widely adopted under the form of polycul-
tures in African and Central American traditional agriculture alike
(Morales and Perfecto, 2000; Kfir et al., 2002; Greathead, 2003).
Early on, possibilities were discussed to manipulate agricultural
habitats to increase natural enemy abundance in Ugandan coffee
plantations (Taylor, 1945). Our literature revision found a total of
71 records of habitat manipulation (HM), and an additional 80
studies specifically on the effect of inter- and cover-cropping
(ICC) on natural enemy abundance or efficacy (Appendix 4). Crop-
ping systems that received most attention were coffee (9), rice (7),
maize and citrus (7), apple, cocoa and wheat (5) and beans (4). For
inter- and cover-cropping, great attention was paid to maize (11),
cotton (11), apple (5), sweetpotato, bean, tomato and sorghum (4)
as focal crops. In most cases, target pests were not specified for HM
(44), and ICC (30). In cases where the target pest was mentioned,
aphids (5), thrips (3) and corn earworm (3) were common in HM
studies, while aphids (14), sweet potato weevil (5) and corn ear-
worm (5) received great attention for ICC. The effect of habitat
manipulation was assessed on several natural enemies, such as
ants (17), coccinellids (10), spiders (10), hoverflies (6), lacewings
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167 157

Table 1
Plant species and associated natural enemy guilds, commonly studied as companion plants in inter- and cover-cropping schemes in the developing world.

Plant family Genus Species Common # Focal crop Focal natural enemies
name Records
Aliaceae Allium ampeloprasum Leek 1 Vegetables NS
sativum Garlic 1 Chinese cabbage Th, Ly
cepa Onion 1 Vegetables NS
Apiaceae Coriandrum sativum Coriander 4 Tomato, chickpea, safflower Ar, Ch, Co, Fr, Ic
Galinsoga parviflora Gallant 1 Tomato Ar, Co, Fr
soldier
Pimpinella anisum Anise 1 Vegetables NS
Arecaceae Cocos nucifera Coconut 1 Cocoa Fr
Asteraceae Ageratum conyzoides Billy-goat 1 Citrus Ph
weed
houstonianum Bluemink 1 Pear Co, Ph, Ch
Carthamus tinctorius Safflower 1 Cotton Ch
Lactuca sativa Lettuce 1 Chinese cabbage Th, Ly
Tagetes patula French 1 Pear Co, Ph, Ch
marigold
Brassicaceae Brassica campestris Rape 3 Apple, cotton, wheat NS
oleraceae Green 1 Chinese cabbage Th, Ly
cabbage
Commelinaceae Commelina diffusa Climbing 1 Coffee Fr, Br, Ic
dayflower
Zebrina pendula Wandering 1 Coffee Fr, Br, Ic
jew
Cucurbitaceae Cucurbita pepo Squash 2 Maize Ch, Fr
Trichosanthes cucumerina Snake 1 Tea Ar
gourd
Euphorbiaceae Manihot esculenta Cassava 1 Maize Sc
Fabaceae Arachis hypogaea Peanut 3 Sorghum, maize Ar, Fr, Tr
pintoi Pinto 1 Tea Ar, Ca
peanut
Cyamopsis tetragonoloba Cluster 1 Cotton Ar, Co
bean
Desmodium sp. Tick clover 1 Maize Ar
Glycine max Soybean 4 Maize, sweetpotato Ar, Fr, Sc
Medicago sativa Alfalfa 5 Apple, cotton An, Ar, Ca, Ch, Co, Na, Ge, Ch, Br,
St, Sy
Melilotus albus Honey 1 Apple Ch, Ph
clover
Neonotonia wightii Perennial 1 Citrus Ph
soybean
Phaseolus vulgaris Kidney 6 Cereals, maize, sorghum, sugarcane Br, Ch, Eu, Fr, Ic, Ph, Ta, Tr
bean
Pisum sativum Pea 2 Grape, wheat Co, Ve, Fr, Br
Trifolium fragiferum Strawberry 1 Apple, Ca, St, Co, Na, Ge, An, Ch, Ar, Br
clover
repens White 1 Apple Ch, Co, Sy
clover
Vigna radiata Mungbean 3 Cowpea, maize, sorghum An, Ar, Fo, Tr
unguiculata Cowpea 3 Cotton, maize, sorghum Ar, Br, Co, Ic, Sc
Lamiaceae Lagopsis supina NA 1 Apple NS
Melissa officinalis Lemon 1 Vegetables NS
balm
Mentha haplocalyx Mint 1 Pear Ch, Co, Ph
x piperita Peppermint 1 Vegetables NS
Ocimum basilicum Basil 2 Pear Co, Ph, Ch
Salvia officinale Sage 1 Vegetables NS
Satureja hortensis Summer 1 Pear Co, Ph, Ch
savory
Malvaceae Gossypium hirsutum Cotton 1 Pigeonpea Sc
Hibiscus sabdariffa Roselle 1 Tomato Br, Ic
Myricaceae Myrica rubra Waxberry 1 Tea Ar
Poaceae Avena strigosa Black oat 1 Grape Ve, Fr, Co
Brachiaria decumbens Suriname 1 Banana, plantain Ar, Ci, Fr, Fo
grass
Festuca arundinaceae Fescue 1 Apple Ca, St, Co, Na, Ge, An, Ch, Ar, Br
Lolium perenne Perennial 1 Apple Ch, Co, Sy
ryegrass
Melinis minutiflora Molasses 1 Maize Br
grass
Pennisetum glaucum Pearl millet 3 Cereals, cowpea, pigeonpea Br, Eu, Ic, Ta, Tr
purpureum Napier 1 Maize Ar

(continued on next page)

158 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
Table 1 (continued)
Plant family Genus Species Common
name
grass
Sorghum bicolor Sorghum
Triticum aestivum Wheat
Zea mays Maize
Rutaceae Citrus sinensis Orange
Theaceae Camellia sinensis Tea
NS : non-specified.
An: Anthocoridae; Ar: Araneae (spiders, unspecified); Br: Braconidae ; Ca : Carabidae ; Ch : Chrysopidae ; Ci : Chilopoda (centipedes, unspecified) ; Co : Coccinellidae ; Eu :
Eulophidae ; Fo : Forficulidae ; Fr : Formicidae ; Ge: Geocordiae; Ic : Ichneumonidae ; Ly : Lycosidae ; Ma : Mantidae ; Na: Nabidae; Ph : Phytoseiidae ; Sc : Scelionidae ; Sy :
Syrphidae ; St : Staphylinidae ; Ta : Tachinidae ; Th : Theridiidae ; Tr : Trichogrammatidae ; Ve : Vespidae.
and phytoseiids (3), for HM. For ICC, natural enemies that received
most attention were spiders (20), coccinellids (17), ants (12),
anthocorids (8) and lacewings (7).
Several HM tactics were tested, with the most common being
the use of herbaceous strips in the field plot (21), tolerance of
weeds (13), judicious selection of border plants (11), and the estab-
lishment of shade trees (7). Flower strips or weed borders were
evaluated, using a total of 21 different plant species belonging to
6 families. Members of Poaceae (6), Brassicaceae (5) and Fabaceae
(4) were widely used. In four studies, existing weeds were toler-
ated in some sections of the field plot, instead of intentionally
establishing new plants in separate strips. In two studies from In-
dia, Sri Lanka and Malaysia, researchers assessed the effect of prac-
tices such as partial weeding on the beneficial arthropod fauna
(Rickson and Rickson, 1998; Rekha et al., 2009). Tolerance of some
weed cover could be a labor-saving approach to help conserve nat-
ural enemies (Norris and Kogan, 2000), and merits more intensive
research as a tailor-made HM tactic for small-scale farmers. Lastly,
six studies on the provision of artificial shelter and nesting re-
sources introduce (possibly) effective, simple and low-cost prac-
tices to conserve predatory ants or mites. However, if those
tactics actually contribute to pest control waits to be evaluated
in many of above cases. In ICC schemes, an equally broad diversity
of plant species was evaluated either as cover crops or intercrops
(Table 1). A total of 54 different plant species were tested, with var-
ious species belonging to Fabaceae (14) or Poaceae (10). A great
number of records were found for maize (16), sorghum (6) and al-
falfa (5) as companion plans. For Poaceae, the effect of different
plant species was primarily quantified on Araneae (5), Coccinelli-
dae (5), Forficulidae, Formicidae or Chrysopidae (3). For Fabaceae,
the effect of several species was assessed on Araneae (8), Coccinel-
lidae (6) and Formicidae (4). In the bulk of ICC studies, researchers
evaluated the effect of a given practice on natural enemy abun-
dance (43) and to lesser extent on pest pressure (21). No records
were found in which the underlying drivers for increased natural
enemy abundance were determined. Although limited research
has been conducted on proper management of cover- or intercrops,
findings from Chinese apple (Du and Yan, 1994), Chinese cotton
(Zhang et al., 2000), Chinese chestnut (Lu et al., 2008) and Guade-
loupian citrus orchards (Mailloux et al., 2010) indicate this to be a
fertile field for future research. However, in many of those systems,
with exception of Chinese apple orchards, little evidence exists of
widespread adoption of those cover crop management schemes.
In certain systems, clear potential exists for follow-up research
on HM or ICC tactics. For example, the effect of grass borders on
natural enemies waits to be determined in Latin American bean
production (Altieri, 1981). In Central America, in-field abundance
of earwigs is directly related to grass cover in neighboring fields,
but little attention has been paid to its related potential as HM
tactic (Wyckhuys and O’Neil, 2007b). Lastly, Vietnamese citrus
# Focal crop Focal natural enemies
Records
6 Blackgram, cereals, cowpea, cotton, pigeonpea An, Ar, Br, Eu, Ic, Fo, Ta, Tr
1 Cotton Co, Ar
16 Beans, cabbage, cereals, cowpea, cotton, groundnut, An, Ar, Br, Ca, Co, Ch, Eu, Fo, Fr, Ic,
pigeonpea, potato, soybean, sweetpotato, tomato Ly, Ma, Sc, St, Sy, Ta, Tr, Ge
1 Tea Ar
1 Citrus Br, Hy
growers recognize that pest problems are lower in weedy fields
(Van Mele and van Lenteren, 2002), which could point towards
effective HM schemes. In other cases, early research has been
translated into HM practices and widely adopted, as is the case
for the use of Ageratum conyzoides cover crops in 135,000 ha of
Chinese citrus orchards or the broad adoption of sunflower strips
in Cuban small-scale agriculture (Liang and Huang, 1994; Vázquez
et al., 2007). Similarly, an exhaustive evaluation of the role of flow-
ering plants in attracting natural enemies has been translated in
HM tactics for Tajik cotton and vegetable production (Saidov and
Landis, 2008; Saidov et al., 2011).
Although considerable effort has gone into the evaluation of
habitat manipulation schemes, little or no attention has been paid
to consolidating a mechanistic basis for this research. Instead of
simply adding diversity to agro-production systems or evaluating
the effect of (a small set of) common crops or N-suppliers on the
resident natural enemy community, it may be preferable to a priori
identify the ‘right kind’ of diversity (Landis et al., 2000; Gurr, 2009),
or pursue the promotion of healthy ecosystems (J. Lundgren, per-
sonal communication). With applied ecological research identified
as a necessary step to develop pest-suppressive crop diversification
schemes in the Western hemisphere (Barbosa et al., 2009; Letour-
neau et al., 2011), such work is urgently needed in the developing
world. By linking new mechanistic modeling work (e.g., Tixier
et al., 2011; Vinatier et al., 2012) to recent advances in natural
enemy ecology (e.g., Wackers, 2004; Fiedler and Landis, 2007;
Tompkins et al., 2010), such habitat manipulation schemes could
be within close reach for certain (perennial) systems in the tropics.
6. Deliberate manipulation of disturbance regimes
Several types of disturbance, such as tillage or weed manage-
ment, can have far-reaching effects on the detritus food web and
consequently on the resident natural enemy community (Wardle,
1995). Also, while insecticide use directly impacts natural enemies,
relatively little research has been conducted in the Western hemi-
sphere to make chemical control compatible with (natural) biolog-
ical control (Naranjo and Ellsworth, 2009). In our literature
revision, a total of 77 references were found that describe the effect
of disturbances (e.g., pesticide use, tillage, fertilizer application) on
arthropod natural enemies (Appendix 5). Research was commonly
conducted in major crops such as rice (13), cotton (12), coffee (10)
or soybean-corn rotations (5). A total of 25 accounts were from
(semi-)perennial systems, while all other references were on
short-term crops. Many references evaluated the effect on ants
(12), spiders (8), coccinellids (4) and ground beetles (5). A range
of disturbance factors were taken into account, with most research
focusing on insecticide use frequency or timing (22), use of organic
fertilizer or soil amendments (8) and alteration in tillage regimes
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
(8). Other accounts dealt with practices such as pruning (3), collec-
tion of fallen fruits (2), hand weeding (1), modification of fallow re-
gimes (1), overhead irrigation (1), and the use of insectivorous
farm animals such as ducks (1). Insecticide use alterations were
mostly evaluated on natural enemy abundance (15), but rarely
on pest pressure (2) or natural enemy efficacy (3). Organic fertilizer
use was commonly evaluated on natural enemy abundance (8), but
only once on predation levels and pest pressure. Similarly, altera-
tions in tillage regimes were mostly evaluated on abundance (5)
and not on pest pressure. This is surprising, as tillage is frequently
mentioned in folk knowledge as an efficient way to reduce pests.
For example, among the Tzeltal Maya in present-day Mexico, land
preparation is widely seen as a way to expose Phyllophaga sp.
white grubs to predation by a variety of natural enemies (Gomez
et al., 2000). This shows that much research waits to be conducted,
and that an exclusive focus on natural enemy abundance may ob-
scure (possibly) stronger effects of certain disturbance factors on
biological control efficacy.
Fertilizer use and the addition of organic matter generate bot-
tom-up forces that conserve resident natural enemies in many
cropping systems, with tangible increases of foliage-foraging pre-
dators such as spiders in Asian rice (Settle et al., 1996), aphid pre-
dators in African bean (Karungi et al., 2006) or earwigs in Brazilian
maize (Galvao et al., 2001). Similarly, crop residue retention led to
a 7- and 15-fold increase in the respective abundance of predatory
ants and springtails in Brazilian common bean (Pereira et al.,
2010). In several systems, pest outbreaks proved less likely upon
addition of organic matter, but its effect on natural enemies was
not assessed (Neuenschwander et al., 1990). Given the (relatively)
steep and increasing costs of chemical fertilizers for small-scale
farmers, there is a need to assess effects of organic fertilizers, man-
ure or crop residue retention schemes on biological control. This
work is particularly relevant in light of the rising use of unsustain-
able practices in various developing world nations, such as straw
burning in Indian rice–wheat systems (Gupta et al., 2004).
Along the same lines, research has shown that rational insecti-
cide use can bring about dramatic increases in natural enemy lev-
els. Such practices led to natural enemy increases of 300% in
Chinese cotton fields (Xing et al., 1991), or parasitism levels up
to 97% in Indian tea plots (Hazarika et al., 2001). In South Africa,
native parasitoids exerted sufficient control of the invasive potato
tuber moth, even when farmers refrained from insecticides (Kfir,
2003). In the meantime, several commonly-used herbicides and
fungicides greatly affect natural enemies, either by causing direct
mortality (Chen et al., 1999) or through changes in habitat struc-
ture, composition or health (Pereira et al., 2007). Consequently, a
reduction of pesticide use in certain crops can boost the contribu-
tion of biological control and thus become a valuable pest manage-
ment approach for resource-poor farmers (e.g., Greathead and
Girling, 1981). Classic experiments in Asian rice systems (Kenmore
et al., 1984; Settle et al., 1996) or Peruvian cotton (Beingolea, 1959)
helped create broad awareness on the side-effects of pesticide
over-use and highlighted the potential of biological control. Recent
renewed pest outbreaks in several of those systems signal that the
time may be ripe to revisit those historic works and use them to re-
orient deficient pest management schemes.
7. Germplasm effects on natural enemy performance
Agricultural crops, as highly suitable host plants of arthropod
pests, may equally affect natural enemies through multi-trophic
interactions (Bottrell et al., 1998). In the Western hemisphere,
researchers have come to acknowledge that a sound appreciation
of plant–pest–natural enemy evolution can help develop CBC
schemes. In our literature search, we found a total of 27 studies
that describe crop-mediated influences on natural enemies
159
(Appendix 6). Crops that received most attention were pigeon
pea (6), cotton (4) and cassava (4), with a third of studies from In-
dia. The effect of crop germplasm was assessed on a diverse range
of natural enemy groups, with Phytoseiidae (4), Coccinellidae (3)
and Trichogrammatidae (3) commonly studied. Crop attributes
that received most importance were leaf pubescence or trichome
density (8), pest resistance (4) and varietal-dependent production
of volatiles (3).
In some crops, such as cassava, leaf trichomes have been shown
to facilitate establishment and presence of certain predators while
trapping alternate foods such as pollen (Zundel et al., 2009). As
multiple natural enemies readily feed upon cassava extra-floral
nectar (Bakker and Klein, 1992; Bruce-Oliver et al., 1996), the po-
tential for CBC by selecting cassava varieties with enhanced nectar
secretion or pubescence has been repeatedly identified (Salick,
1983). This characteristic however, in addition to leaf pubescence,
has yet to be taken into account in breeding programs (A. Bellotti,
personal communication). Along the same lines, several research-
ers have identified the considerable potential of breeding crops
that resist specific pests and simultaneously encourage their
antagonists (e.g., Sharma and Yadav, 1993; Lou and Cheng, 2003).
Host plant resistance does not necessarily interfere with natural
enemy fitness, mobility or host searching ability (Tandon, 2001)
and moderate levels of resistance could easily boost natural enemy
abundance and efficacy (Kartohardjono and Heinrichs, 1984;
Shannag and Obeidat, 2008).
With insecticide- and herbicide-resistant crops steadily
adopted in many developing countries, the assessment of their
impact on biological control could prove a fertile field of research
(Romeis et al., 2006; Lundgren et al., 2009). Herbicide-tolerant
crops can lead to substantial losses of in-field vegetational diver-
sity and thus (possibly) interfere with biological control. On the
other hand, Bt-transgenic varieties can reduce insecticide use and
give natural enemies a chance (e.g., Lu et al., 2012). While our
study focused on CBC studies related to conventional germplasm,
we also found work that dealt with the effect of transgenic crops
on resident natural enemies in Indian Bt chickpea (Romeis et al.,
2004), Brazilian Bt cotton (Faria et al., 2006), amongst others. The
bulk of these studies provide guidelines for pre-release risk
assessment of transgenics. However, in most developing countries,
laboratory and field trials wait to be conducted that assess compat-
ibility of transgenic crops and natural enemies (Lovei et al., 2009).
Although transgenic crops can help promote natural biological
control in conventional agriculture in certain countries, they are
not suitable for organic production systems.
Above findings show that germplasm research could pave the
road towards more pest-suppressive cropping systems. Neverthe-
less, few breeding programs in the tropics have carried out sus-
tained research on CBC promotion through judicious selection of
crop germplasm. In the meantime, CBC researchers in the develop-
ing world may need to consider including transgenic crops in their
agendas.
8. CBC at the landscape level
In the broader agricultural landscape, the presence of more sta-
ble and heterogeneous (non-crop) habitats such as forests, hedge-
rows, fallows or meadows is thought essential to CBC services
(Tscharntke et al., 2008). They provide important resources for nat-
ural enemies such as refuge sites, alternative prey, pollen and nec-
tar (e.g., Landis et al., 2000; Cronin and Reeve, 2005; Bianchi et al.,
2006). In the Western hemisphere, the importance of landscape-
level contributions to biological control was first reviewed half a
century ago (van Emden, 1965). In our literature revision, a total
of 29 records were found that describe landscape-level effects
on biological control (Appendix 7). Amongst those, ten studies
160 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167

describe rice agro-ecosystems, while less attention has gone to cot-
ton (3) and another eight crops. Natural enemy guilds that received
most attention were spiders (5) and Trichogrammatidae (3). In the
broader agricultural landscape, the effect of surrounding crop
fields (8), landscape complexity and diversity (7) and the presence
or (relative) cover of natural habitats (9) was assessed.
Despite comparatively little research attention, several promis-
ing findings have been reported. In SE Asia, asynchronous rice
planting over large areas allowed natural enemies to build up
(Way and Heong, 1994; Ives and Settle, 1997). In China, high abun-
dance and efficacy of Trichogramma spp. on certain vegetables or
(early-season) sweet potato crops has been used to diversify
grain-cropping landscapes (Wang et al., 1988; Zhou et al., 1997).
Also, landscape mosaics composed of rice and horticultural crops
benefited from high natural enemy abundance and increased pest
suppression (Zhang et al., 2011). In those systems, biological con-
trol can be enhanced either through overall increase of landscape
diversity (e.g., Gardiner et al., 2009), or through the provision of
habitats that benefit one or more key natural enemies, such as
Trichogramma spp. Along the same lines, a mosaic of natural habi-
tats at differing stages of succession supports a diverse natural en-
emy community in Central American small-scale maize production
(Wyckhuys and O’Neil, 2007b). It is suspected that shifting agricul-
ture inadvertently preserves the diversity of habitats and biological
control services within those systems. Despite these promising
findings, much remains to be investigated in this field. More so,
in certain regions, such as Africa, virtually nothing is known about
the effect of landscape structure on biological control (Neuensch-
wander et al., 2003).
9. General tendencies in developing world CBC research
Our literature review reported a wide range of CBC-related
studies from the developing world, on a total of 53 different
focal crops. Research was reported from 16 different fruits, ten
vegetables, eight pulses, six cereals, three tuber/root crops and a
small set of cash or forage crops. Research effort greatly differed
between crop species and crop types. The greatest number of stud-
ies was recorded from rice (36), cotton (35), maize (30), coffee (21),
citrus (14) and beans (13). The number of records per crop species
was highest for cereals (14.33 ± 6.31; mean ± SE) and lowest for
fruits (4.75 ± 1.48) and vegetables (2.80 ± 0.94). Although most
fruit crops were only covered in one or two publications, citrus, ap-
ple and coffee were the focus of more than ten studies each. Similar
patterns were found for vegetables, but research effort was more
equally distributed between crop species for pulses. CBC research
in some important staple crops proved very limited or entirely
missing (Fig. 2). For example, only one record was found for bana-
na/plantain production systems, lentils, barley or millet. This is
particularly worrisome, as crops such as millet or banana/plantain
are grown on 35.1 and 10 million hectares, respectively, and are
key staples in many parts of the tropics. Also, comparatively little
research effort has been dedicated to CBC in potato (3), groundnut
(2) or fava bean (2). For other important staple crops such as yams,
taro, sago or breadfruit, we found no references at all. Possibly, the
limited research effort to some root or tuber crops could be due to
the cryptic nature of (many of) its pests, and associated natural
enemies. CBC research could take forward pest management in
several of those crops, while crop species that already received cer-
tain coverage could benefit from sustained research built upon a
meticulous review of actual advances.
Perennial fruit orchards are optimum habitats to promote con-
servation biological control (e.g., Brown, 1999; Landis et al., 2000;
Simon et al., 2010). Nevertheless, it is rather unfortunate to note an
absence of sustained research in perennial fruits, with the number
of CBC references for major fruits such as mango (2), avocado (1),
papaya (0) or pineapple (0) being strikingly low. For minor fruits,
the situation is even more critical, with no records found for most
crops. In many fruit cropping systems, the current lack of research
attention is reflected in an ever more acute proliferation of

Fig. 2. Number of CBC records per crop, in relation to its 2010 global acreage (log10) and the share of its production outside the European Union and North America. Crop
production data were obtained from FAO Stat (http://faostat.fao.org). Selected crops are identified in the figure, while we did not include crops for which no CBC records were
found or that did not have reliable global production data.Size of the circles represents the number of CBC records for a given crop.
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167 161

Table 2
Number of CBC records for a given pest species, as contrasted with its respective number of worldwide records of insecticide resistance. Insecticide resistance records were
obtained through the Arthropod Pesticide Resistance Database (www.irac-online.org), and are solely reported for agricultural arthropod pests that are important in the
developing world. We included the 30 pest species with highest incidence of insecticide resistance.

Species name Family Number of pesticide Number of

resistance records CBC records
Helicoverpa armigera Noctuidae 640 10
Plutella xylostella Plutellidae 455 5
Bemisia tabaci Aleyrodidae 436 9
Myzus persicae Aphididae 392 1
Tetranychus urticae Tetranychidae 389 2
Spodoptera exigua Noctuidae 322 0
Spodoptera litura Noctuidae 238 2
Panonychus ulmi Tetranychidae 187 1
Aphis gossypii Aphididae 160 12
Frankliniella occidentalis Thripidae 153 0
Nilaparvata lugens Delphacidae 123 7
Heliothis virescens Noctuidae 121 0
Delia antiqua Anthomyiidae 90 0
Thrips tabaci Thripidae 72 3
Trichoplusia ni Noctuidae 68 0
Bactrocera dorsalis Tephritidae 65 0
Earias vittella Noctuidae 64 0
Pectinophora gossypiella Gelechiidae 56 2
Chilo suppressalis Crambidae 51 15
Heliothis assulta Noctuidae 50 0
Spodoptera littoralis Noctuidae 50 0
Panonychus citri Tetranychidae 49 0
Anthonomus grandis Curculionidae 41 0
Sogatella furcifera Delhacidae 39 1
Liriomyza trifolii Agromyzidae 38 0
Spodoptera frugiperda Noctuidae 29 9
Tetranychus cinnabarinus Tetranychidae 26 0
Nephotettix cincticeps Cicadellidae 25 0
Tuta absoluta Gelechiidae 25 0
Pseudoplusia includens Noctuidae 23 0

ripe for a serious appreciation of CBC opportunities in many of

Fig. 3. Number of historical CBC literature records for a subset of the 15 developing
countries with highest research output. One single publication can be considered as
multiple records, depending upon the number of CBC aspects (as described in this
manuscript) that are covered. Publications from the former Soviet Union are not
grouped, but reported per member state.
unsustainable practices. In small-scale Colombian passionfruit pro-
duction for example, >90% of farmers rely on calendar-based insec-
ticide sprays, despite the presence of an abundant and effective
natural enemy community (Wyckhuys et al., 2011; Carrero and
Wyckhuys, unpublished). Similar patterns can be found in vegeta-
ble cropping systems (e.g., Grzywacz et al., 2010), where only to-
mato (6) and certain crucifers (10) appear to have received
sustained CBC research attention. Last but not least, with the sole
exception of one study on alfalfa, forage crops have largely been
deprived from CBC research. As many of the above crops are grown
by smallholders and sustain rural economies, the time might be
those systems.
CBC research covered a wide range of pest insects, with Busseola
fusca or Chilo suppresalis stemborers (15), cotton aphid (12), corn
earworm (11) or cotton bollworm (10) receiving most attention.
Some of these pests, such as Chilo suppressalis and Helicoverpa
armigera, are suitable targets for CBC research given their high sus-
ceptibility to insecticide resistance development. In contrast, other
pest insects with high pesticide resistance incidence, such as Spo-
doptera exigua, Frankliniella occidentalis or Heliothis virescens, have
received little or no CBC research attention (Table 2). Strikingly,
no CBC records were found for more than half of the 30 agricultural
arthropod pests with highest incidence of insecticide resistance.
Increased CBC research attention to some of those pests could help
slow insecticide resistance development, and contribute to a clo-
sure of yield gaps (Godfray et al., 2010). Also, lack of CBC research
for certain large-scale cropping systems (e.g., West African cotton)
in concert with irrational insecticide use seems to trigger
resistance development in other noxious organisms, such as the
malaria vector Anopheles gambiae (Yadouleton et al., 2011).
Many developing countries invest millions of dollars annually in
insecticide imports to fight agricultural pests (Fig. 4). Countries
such as Algeria, Thailand or Morocco have comparatively high
insecticide imports, but seem not to fully recognize the potential
of natural biological control. Similarly, while interest in CBC re-
search steadily grows in countries with high insecticide use such
as Colombia, nations such as Bangladesh seem not to explore other
cost-saving alternatives such as CBC (Fig. 4). Certain mega-diverse
countries, such as Peru, Ecuador or the Philippines possibly have
ample resident natural enemies effective against pests that are
presently controlled with insecticides. For many of these countries,
CBC research could help identify cost-saving pest control tactics
and save scarce funds for more pressing development issues.
162 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167

Fig. 4. Number of CBC records from developing countries, in relation to country-specific annual insecticide imports (A) or annual insecticide consumption levels (B).
Insecticide use data were obtained for the year 2005, from FAO Stat (http://faostat.fao.org). For many developing countries, no reliable information was available on
insecticide consumption or import.

Our literature search encountered a total of 390 CBC-related lit-

Fig. 5. Temporal patterns in the number of literature references on different aspects
of conservation biological control from developing world nations.
erature records from 53 different nations (Fig. 3), with the greatest
number of records from China, Brazil and Cuba. We acknowledge
that records from certain countries such as Russia and Middle East-
ern nations may be far from complete, as some studies are only re-
ported in the grey literature or local languages. On the other hand,
our focused search in the Chinese and Cuban literature could have
disproportionately increased the number of records from those
countries.
With the earliest records on developing-world CBC research
dating from the 1940s, the number of references has steadily in-
creased since the 1980s (Fig. 5). During 2005–2010, a total of 50
studies were found on habitat manipulation and up to 25 on the
modification of disturbance regimes. Research attention to crop
germplasm or the assessment of alternative host or prey items ap-
pears to be diminishing, while food sprays have continued to re-
ceive low but constant levels of research over the past 20 years.
A promising trend is the mounting number of references on the
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
identification of non-prey food sources for (omnivorous) natural
enemies. During 1985–90, only two of 19 studies assessed aspects
related to natural enemy nutrition. Since 2010 onwards, a total of
12 (out of 49) studies described plant-based food sources of certain
natural enemies. Possibly, this latter tendency can generate much-
needed baseline data to take CBC forward in many cropping
systems.
Time-held and novel research approaches alike wait to be em-
ployed to elucidate ecological particularities of natural enemies
in many cropping systems. Ecological experiments, such as addi-
tion/exclusion trials, combined with molecular gut content analy-
sis or biochemical assays could help assess the real potential for
conservation biological control. Only in a few cases, researchers
have moved beyond initial natural enemy surveys, and used exclu-
sion trials, palynology, gut content analysis or feeding studies to
assess the role of certain natural enemies or identify plant-natural
enemy associations (see Santos-Neto et al., 2010; Jaramillo et al.,
2010; Medeiros et al., 2010; Duyck et al., 2011; Narvaez et al.,
2012). Such approaches carry considerable potential to help iden-
tify resident natural enemies of a variety of (cryptic) pests, deter-
mine their nutritional requirements and generate much-needed
information to guide habitat modification schemes or artificial
food sprays.
Looking at historic patterns in CBC research, we note a particu-
lar emphasis on two domains: survey and identification of (poten-
tially) important natural enemies and on-farm evaluation of
possible CBC schemes (see Naranjo, 2001). In many developing
countries, a wealth of information has been generated through di-
rect observation or (descriptive) population censuses of natural
enemies. Observations were primarily carried out in agricultural
crops, and a very small share in natural or non-crop areas. In cer-
tain systems, natural enemy surveys have spanned several years
(Sharma and Agarwal, 2007). In others, natural enemies have only
recently been identified (Rao, 2005), or still wait to be described
(e.g., Baskaran et al., 1999). Surprisingly, there is a critical lack of
natural enemy surveys in certain major staple crops or for key
pests, such as whitefly or Phyllophaga white grubs (e.g., Legg
et al., 2003; Bellotti et al., 2012). Only recently, the predator com-
plex of cassava whitefly has been characterized (Lundgren et al.,
unpublished) and promising research venues have been identified
for cassava whitefly CBC (Ewesi, 2011). For many other crops or
pests, sensible CBC practices can only be defined once insights have
been gained on the identity, abundance and action of resident nat-
ural enemy community.
On the other hand, researchers have gone to great lengths
studying effects of disturbance schemes (20% historic records)
and evaluating habitat manipulation approaches (38%). For habitat
manipulation, companion plants have traditionally been selected
based upon their N-fixing capacity or status as food or cash crop,
mostly with little insights into their benefits for resident natural
enemies. Through time, only a handful of studies have conducted
the necessary, sequential set of experiments to successfully devel-
op CBC or habitat manipulation schemes. Poster-child examples of
such long term studies to boost natural enemy efficacy or abun-
dance are the work on African stemborers (Ndemah et al., 2001;
Kfir et al., 2002), West African cassava mites and mealybugs (e.g.,
Onzo et al., 2005), Mexican fruit fly parasitoids (Aluja, 1994), SE
Asian rice pests (Matteson, 2000), and the recent revival of weaver
ant research (van Mele, 2008). Amongst in-field evaluations of CBC
tactics, we found few records where the extent of target pest sup-
pression was assessed. By missing this pest angle, these studies
also have little relevance to pest management (see Jonsson et al.,
2008; Furlong and Zalucki, 2010).
Although we note numerous promising tendencies in CBC re-
search in the developing world, its future may be threatened by
agricultural development in several regions. First and foremost
163
may be the rising level of mechanization, intensification and sim-
plification of present-day cropping systems. Although pest severity
and related insecticide use in Chinese cotton-wheat intercrops is
far lower than in monoculture crops (Lu, personal communication),
these systems are gradually disappearing. Amounting to >50% of
Chinese cotton acreage at the turn of the century, its proportion
has recently dropped to under 15% (Mao, 2010). In Indian rice–
wheat cropping systems, field borders have long been valued as a
refuge habitat for a broad diversity of natural enemies (Jaipal
et al., 2002). With increasing mechanization in major wheat-grow-
ing regions (e.g., Punjab), straw burning has risen dramatically in
the past decade and its impact on biological control is thought to
be devastating (e.g., Gupta et al., 2004). Lastly, global cassava pro-
duction is rapidly shifting towards large-scale monocultures with
little place for on-farm biodiversity. Dropping natural enemy num-
bers, the appearance of novel pests and more severe outbreaks of
existing pests all point to considerable deficiencies in those pro-
duction systems (Bellotti et al., 2012).
Under these scenarios, CBC may be promoted by capitalizing
on human ingenuity, deployed for centuries to solve agricultural
challenges (Kiers et al., 2008). In many parts of the developing
world, small-scale farmers have acquired an intimate knowledge
of (part of) the local pest and natural enemy community (Bentley
and Rodriguez, 2001; Wyckhuys and O’Neil, 2007a but see Abate
et al., 2000). Based upon this knowledge, many farmers use sim-
ple, on-farm experiments to gradually adapt management prac-
tices to local farming systems (Scoones and Thompson, 1994;
Sumberg and Okali, 1997). Farmer inventions for pest manage-
ment have been documented from all over the globe, with records
as old as written history (Kiritani and Nakasuji, 1977; Peng,
1983). Even though farmer inventions have been derided as a re-
sult of inaccurate folk science, a multitude of valuable pest man-
agement practices can be documented. For example, Chinese
citrus farmers relied on observation to understand key aspects
of weaver ant ecology and devise tactics to improve their efficacy
in pest control. Many of these inventions have survived the test of
time, and have been adopted by generations of farmers world-
wide. A systematic documentation of such local CBC innovations
and their possible insertion into modern-day farming could carry
tremendous potential (see Morales and Perfecto, 2000; Wyckhuys
and O’Neil, 2007a; van Mele, 2008). Such approach fits seamlessly
within emerging interests for traditional knowledge and practices
to help guide ecological engineering processes (Martin et al.,
2010).
10. Conclusions
As a practice that is neither self-perpetuating (i.e., as classical
biological control) nor benefits from political support or financial
backing through vested industry interests, the future of CBC has of-
ten been termed as bleak (Ehler, 1998). However, our documenta-
tion of 390 CBC-related literature records from 53 different nations
and >50 crops shows exactly the opposite. Active research in this
field is being conducted, and critical insights in natural enemy
ecology and CBC effectiveness are slowly but gradually being gen-
erated. Given that past work has mainly been conducted in isola-
tion and long-term, sustained research on a given cropping
system is still very much a rarity, we identify vast potential for
well-orchestrated regional CBC projects that make strategic use
of historic data, carry out additional research and wisely employ
(local) knowledge to pursue higher levels of CBC integration into
current pest management schemes. Now more than ever, CBC in
the developing world is at a crossroads. In times of far-reaching
budget cuts in international agricultural research, scientists have
become increasingly concerned of focusing on ‘‘trendy’’ issues,
164 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
such as biotechnology or computational science. However, with
pest management improvements causing five times higher multi-
plicative increases than plant breeding in West African food pro-
duction (Pretty et al., 2011), a closer look should be taken at
present-day research priorities. Our work shows that the time is
ripe to broadly recognize the benefits of biological control, and
consider it as an integral part of sustainable crop intensification
strategies in the developing world (Neuenschwander, 2010). By
doing so, one could reduce greenhouse gas emissions, improve hu-
man and environmental health, circumvent insecticide resistance
development and promote stability in yield gaps (Godfray et al.,
2010). To achieve this, we identify an urgent need of funding to
conduct fundamental agricultural ecology research, and ensure
continuity of emerging initiatives in several cropping systems.
While this work may be unfashionable, it could provide vital guid-
ance for the design of cost-effective, environmentally-sound pest
management solutions that are tailor-made to small-scale farming
systems worldwide.
Acknowledgments
We are grateful to Jonathan Lundgren, Douglas Landis and Roy
Van Driesche for their revision of earlier drafts of the manuscript.
Nurali Saidov (ICARDA) and Dora Carmona (INTA) provided valu-
able information on CBC practices in Russia and Argentina,
respectively.
Appendix A. Supplementary data
Supplementary data associated with this article can be found, in
the online version, at http://dx.doi.org/10.1016/j.biocontrol.2012.
11.010.
References
Abate, T., van Huis, A., Ampofo, J.K.O., 2000. Pest management strategies in
traditional agriculture: an African perspective. Annual Review of Entomology
45, 631–659.
Agusti, N., Shayler, S.P., Harwood, J.D., Vaughan, I.P., Sunderland, K.D., Symondson,
W.O.C., 2003. Collembola as alternative prey sustaining spiders in arable
ecosystems: prey detection within predators using molecular markers.
Molecular Ecology 12, 3467–3475.
Ahmad, N., Sarwar, M., Wagan, M.S., Muhammad, R., Tofique, M.T., 2011.
Conservation of biocontrol agents in cotton, Gossypium hirsutum L. field by
food supplements for insect pest management. The Nucleus 48, 255–260.
Altieri, M.A., 1981. Weeds may augment biological control of insects. Calif. Agric.
May 1981, 22–24.
Aluja, M., 1994. Bionomics and management of Anastrepha. Annual Review of
Entomology 39, 155–178.
Baggen, L.R., Gurr, G.M., Meats, A., 1999. Flowers in tritrophic systems: mechanisms
allowing selective exploitation by insect natural enemies for conservation
biological control. Entomologia Experimentalis et Applicata 91, 155–161.
Bakker, F.M., Klein, M.E., 1992. Transtrophic interactions in cassava. Experimental
and Applied Acarology 14, 293–311.
Barbosa, P., 1998. Conservation Biological Control. Academic Press, New York, NY.
Barbosa, P., Hines, J., Kaplan, I., Martinson, H., Szczepaniec, A., Szendrei, Z., 2009.
Associational resistance and associational susceptibility: having right or wrong
neighbors. Annual Review of Ecology Evolution and Systematics 40, 1–20.
Barducci Boza, T., 1972. Ecological consequences of pesticides used for the control
of cotton insects in Canete Valley, Peru. In: Taghi Farvar, M., Milton, J.P. (Eds.),
The Careless Technology: Ecology and International Development. Natural
History Press, New York, pp. 423–438.
Baskaran, R.K.M., Mahadevan, N.R., Sathiyanandam, V.K.R., Thangavelu, S., 1999.
Approaches for prevention and control of sesame pests. In: Sinha, M.P. (Ed.),
Recent Advances in Ecobiological Research, vol. 2. A.P.H. Publishing
Corporation, New Delhi.
Beingolea, O.D., 1959. Notas sobre la bionómica de arañas e insectos benéficos que
ocurren en el cultivo de algodón. Revista Peruana de Entomología Agrícola 2,
36–44.
Bellotti, A., Herrera, B.V., Hyman, G., 2012. Cassava production and pest
management: present and potential threats in a changing environment.
Tropical Plant Biology 5, 39–72.
Bentley, J.W., Rodriguez, G., 2001. Honduran folk entomology. Current
Anthropology 42, 285–301.
Bentley, J.W., 2006. Folk experiments. Agriculture and Human Values 23, 421–462.
Bianchi, F.J.J.A., Booij, C.J.H., Tscharntke, T., 2006. Sustainable pest regulation in
agricultural landscapes: a review on landscape composition, biodiversity and
natural pest control. Proceedings of the Royal Society of London. Series B 273,
1715–1727.
Bottrell, D.G., Barbosa, P., Gould, F., 1998. Manipulating natural enemies by plant
variety selection and modification: a realistic strategy? Annual Review of
Entomology 43, 347–367.
Bottrell, D.G., Schoenly, K.G., 2012. Resurrecting the ghost of green revolutions past:
The brown planthopper as a recurring threat to high-yielding rice production in
tropical Asia. Journal of Asia-Pacific Entomology 15, 122–140.
Boukary, I.B., Gingras, J., Tourneur, J.C., 1997. Feeding habits of Forficula senegalensis
(Dermaptera: Forficulidae) on millet in the Sudanese-Sahelian zone of Niger:
crop content analysis. Entomophaga 42, 537–542.
Brown, M.W., 1999. Applying principles of community ecology to pest management
in orchards. Agriculture, Ecosystems & Environment 7, 103–106.
Bruce-Oliver, S.J., Hoy, M.A., Yaninek, J.S., 1996. Effect of some food sources
associated with cassava in Africa on the development, fecundity and longevity
of Euseius fustis (Acari: Phytoseiidae). Experimental and Applied Acarology 20,
73–85.
Canas, L.A., O’Neil, R.J., 1998. Applications of sugar solutions to maize, and the
impact of natural enemies on fall Armyworm. International Journal of Pest
Management 44, 59–64.
Carabali-Banguero, D.J., Wyckhuys, K.A.G., Montoya-Lerma, J., Kondo, T., Lundgren,
J.G., in press. Do additional sugar sources affect the degree of attendance of
Dysmicoccus brevipes by the fire ant Solenopsis geminata? Entomologia
Experimentalis et Applicata.
Carvalho, R., Soares, W., Ritzinger, R., 2010. Umbu-cajá como repositório natural de
parasitoide nativo de moscas-das-frutas. Pesquisa Agropecuária Brasileira 45,
1222–1225.
Chalcoff, V.R., Aizen, M.A., Galetto, L., 2006. Nectar concentration and composition
of 26 species from the temperate forest of South America. Annals of Botany 97,
413–421.
Chen, J.M., Yu, X.P., Lu, Z.X., Zheng, X.S., Cheng, J.A., 1999. Effects of herbicides and
fungicides on the brown planthopper, Nilaparvata lugens and its predatory
enemies. Acta Phytophylacica Sinica 26, 162–166.
Chen, Q., Peng, Z.Q., Xu, C., Tang, C., Lu, B.Q., Wen, H., Wan, F.H., 2010. Biological
assessment of Tetrastichus brontispae, a pupal parasitoid of coconut leaf beetle
Brontispa longissima. Biocontrol Science and Technology 3, 283–295.
Cherry, A., Cock, M., van den Berg, H., Kfir, R., 2003. Biological control of Helicoverpa
armigera in Africa. In: Neuenschwander, P., Borgemeister, C., Langewald, J.
(Eds.), Biological Control in IPM Systems in Africa. CABI Publishing, Wallingford,
UK, pp. 329–346.
Chinajariyawong, A., Clarke, A.R., Jirasurat, M., Kristaneepiboon, S., Lahey, H.A.,
Vijaysegaran, S., Walter, G.H., 2000. Survey of Opiine parasitoids of fruit flies
(Diptera: Tephritidae) in Thailand and Malaysia. Raffles Bulletin of Zoology 48,
71–101.
Choate, B., Drummond, F., 2011. Ants as biological control agents in agricultural
cropping systems. Terresterial Arthropod Reviews 4, 157–180.
Cook, S.M., Khan, Z.R., Pickett, J.A., 2007. The use of push-pull strategies in
Integrated Pest Management. Annual Review of Entomology 52, 375–400.
Cronin, J.T., Reeve, J.D., 2005. Host-parasitoid spatial ecology. a plea for a landscape-
level synthesis. Proceedings of the Royal Society of London. Series B 272, 2225–
2235.
Cuautle, M., Rico-Gray, V., 2003. The effect of wasps and ants on the reproductive
success of the extrafloral nectaried plant Turnera ulmifolia (Turneraceae).
Functional Ecology 17, 417–423.
Daily, G.C., 1997. Nature’s services: societal dependence on natural ecosystems.
Island, Washington, DC.
Doutt, R.L., Smith, R.E., 1971. The pesticide syndrome – diagnosis and suggested
prophylaxis. In: Huffaker, C.B. (Ed.), Biological Control. Plenum Press, New York,
USA, p. 511.
Du, X.G., Yan, Y.H., 1994. Effect of the cover crops on the pest mite Panonychus ulmi
and its predator Orius sauteri. Chinese Journal of Biological Control 10, 114–117.
Duyck, P.F., Lavigne, A., Vinatier, F., Achard, R., Okolle, J.N., Tixier, P., 2011. Addition
of a new resource in agroecosystems: do cover crops alter the trophic positions
of generalist predators? Basic and Applied Ecology 12, 47–55.
Ehler, L.E., 1998. Conservation Biological Control: Past, Present and Future. In:
Barbosa, P. (Ed.), Conservation Biological Control. Academic Press, San Diego,
pp. 1–8.
Eremenko, T.S., 1971. Importance of the adult parasite feeding of winter moth in
increasing their efficiency. In: Biological methods of protection fruit and
vegetable crops from insect pest, diseases and weed as bases of the integrated
systems. Thesis’ report, October 1971, Kishinev, pp. 30–31.
Ewesi, E., 2011. Potentials and prospects of strip cropping in the management of
cassava whitefly (Bemisia tabaci) in peri - urban agroecosystems. Texas A&M
University, PhD thesis.
Faria, M.R., Lundgren, J.G., Fontes, E.M.G., Fernandes, O.A., Schmidt, F., Nguyen, V.T.,
Andow, D.A., 2006. Assessing the effects of Bt cotton on generalist arthropod
predators. In: Hilbeck, A., Andow, D.A., Fontes, E.M.G. (Eds.), Environmental risk
assessment of genetically modified organisms, Methodologies for Assessing Bt
Cotton in Brazil, vol. 2. CABI, Wallingford, UK, pp. 175–199.
Fiedler, A.K., Landis, D.A., 2007. Plant characteristics associated with natural enemy
abundance at Michigan native plants. Environmental Entomology 36, 878–
886.
Figueroa de la Rosa, J.I., Ovruski Alderete, S., Liedo Fernandez, J.P., Cancino Díaz, J.L.,
1998. Parasitoides nativos y exóticos asociados a Anastrepha spp. en frutales
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
silvestres y cultivados de la región del Soconusco, Chiapas, México. XXI
Congreso Nacional de Control Biológico. México.
Furlong, M.J., Zalucki, M.P., 2010. Exploiting predators for pest management: the
need for sound ecological assessment. Entomologia Experimentalis et Applicata
135, 225–236.
Galvao, J.C., Silva, E.C., Miranda, G.V., Bastos, C.S., Picanco, M.C., Silva, R.G., 2001.
Densidade populacional de alguns insetos em milho exclusivo e consorciado
com feijao, em dois sistemas de adubacao. Revista Ceres 48, 25–35.
Gardiner, M.M., Landis, D.A., Gratton, C., DiFonzo, C.D., O’Neal, M., Chacon, J.M.,
Wayo, M.T., Schmidt, N.P., Mueller, E.E., Heimpel, G.E., 2009. Landscape
diversity enhances biological control of an introduced crop pest in the north-
central USA. Ecological Applications 19, 143–154.
Gnanvossou, D., Hanna, R., Yaninek, S.J., Toko, M., 2005. Comparative life history
traits of three neotropical phytoseiid mites maintained on plant-based diets.
Biological Control 35, 32–39.
Godfray, H.C.J., Beddington, J.R., Crute, I.R., Haddad, L., Lawrence, D., Muir, J.F.,
Pretty, J., Robinson, S., Thomas, S.M., Toulmin, C., 2010. Food security: the
challenge of feeding 9 billion people. Science 327, 812–817.
Gomez, B., Castro, A., Junghans, C., Montoya, L., Villalobos, F.J., 2000. Ethnoecology of
white grubs (Coleoptera: Melolonthidae) among the Tzeltal Maya of Chiapas.
Journal of Ethnobiology 20, 43–59.
Graham, M.K., 1991. Biological control of Helopeltis spp. in mature cocoa by the
black ant (Dolichoderus bituberculatus) and the cocoa mealybug (Planococcus
lilacinus). Planter 67, 516–520.
Greathead, D.J., 2003. Historical overview of biological control in Africa. In:
Neuenschwander, P., Borgemeister, C., Langewald, J. (Eds.), Biological Control
in IPM Systems in Africa. CABI Publishing, Wallingford, UK, pp. 1–26.
Greathead, D.J., Girling, D.J., 1981. Possibilities for natural enemies in Heliothis
management and the contribution of the Commonwealth Institute of Biological
Control. In: Proceedings of the International Workshop on Heliothis
Management, 15–20 November 1981. ICRISAT Center, Patancheru, India.
Grzywacz, D., Rossbach, A., Rauf, A., Russell, D.A., Srinivasan, R., Shelton, A.M., 2010.
Current control methods for diamondback moth and other brassica insect pests
and the prospects for improved management with lepidopteran-resistant Bt
vegetable brassicas in Asia and Africa. Crop Protection 29, 68–79.
Guo, X.L., Wang, D.S., He, Y.R., Luo, Y.L., 2011. Effects of different nutritional
resources on longevity and fecundity of Trichogrammatoidea bactrae Nagaraja.
Chinese Journal of Biological Control 27, 448–452.
Gupta, P.K., Sahai, S., Singh, N., Dixit, C.K., Singh, D.P., Sharma, C., Tiwari, M.K., Gupta,
R.K., Garg, S.C., 2004. Residue burning in rice–wheat cropping system: Causes
and implications. Current Science 87, 1713–1717.
Gurr, G.M., 2009. Prospects for ecological engineering for planthoppers and other
arthropod pests in rice. In: Heong, K.L., Hardy, B. (Eds.), Planthoppers: New
Threats to the Sustainability of Intensive Rice Production Systems in Asia. Los
Banos, Philippines, International Rice Research Institute, pp. 371–388.
Gurr, G.M., Liu, J., Read, D.M.Y., Catindig, J.L.A., Cheng, J.A., Lan, L.P., Heong, K.L.,
2011. Parasitoids of Asian rice planthopper (Hemiptera: Delphacidae) pests and
prospects for enhancing biological control by ecological engineering. Annals of
Applied Biology 158, 149–176.
Gurr, G.M., Read, D.M.Y., Catindig, J.L.A., Cheng, J., Lium, J., Lan, L.P., Heong, K.L.,
2012. Parasitoids of the rice leaffolder Cnaphalocrocis medinalis and prospects
for enhancing biological control with nectar plants. Agricultural and Forest
Entomology 14, 1–12.
Gurr, G.M., Wratten, S.D., Altieri, M.A., 2004. Ecological Engineering for Pest
Management: Advances in Habitat Manipulation for Arthropods. CSIRO
Publishing, Collingwood, Australia.
Gyorfi, J., 1951. Die Schlupfwespen und der Unterwhuchs des Waldes. Zeitschrift für
Angewandte Entomologie 33, 32–47.
Hardy, J.E., 1938. Plutella maculipennis Curt., its natural and biological control in
England. Bulletin of Entomological Research 29, 343–372.
Hazarika, L.K., Bhuyan, M., Hazarika, B.N., 2009. Insect pests of tea and their
management. Annual Review of Entomology 54, 267–284.
Hazarika, L.K., Puzari, K.C., Wahab, S., 2001. Biological control of tea pests. In:
Upadhyay, R.K., Mukerji, K.G., Chamola, B.P. (Eds.), Biocontrol Potential and Its
Exploitation in Sustainable Agriculture, vol. 2: Insect Pests, ed. Kluwer
Academic, New York, pp. 159–180.
Ho, C.T., Khoo, K.C., 1997. Partners in biological control of cocoa pests: mutualism
between Dolichoderus thoracicus (Hymenoptera: Formicidae) and Cataenococcus
hispidus (Hemiptera: Pseudococcidae). Bulletin of Entomological Research 87,
461–470.
Huang, H.T., Yang, P., 1987. The ancient cultured citrus ant. BioScience 37, 665–671.
Isaacs, R., Tuell, J., Fiedler, A., Gardiner, M., Landis, D., 2009. Maximizing arthropod-
mediated ecosystem services in agricultural landscapes: the role of native
plants. Frontiers in Ecology and the Environment 7, 196–203.
Ives, A.R., Settle, W.H., 1997. Metapopulation dynamics and pest control in
agricultural systems. Am. Nat. 149, 220–246.
Jago, N.D., 1991. Integrated pest management for rainfed millet in northwest Mali.
In: Kiss, A., Meerman, F. (Eds.), Integrated Pest Management and African
Agriculture. World Bank Technical Paper 142, World Bank, Washington, DC, pp.
25–33.
Jaipal, S., Singh, S., Yadav, A., Malik, R.K., Hobbs, P., 2002. Species diversity and
population density of macrofauna of rice-wheat cropping habitat in semi-arid
subtropical north-west India in relation to modified tillage practices of wheat
sowing. International workshop on herbicide resistance management & zero
tillage in rice-wheat cropping system, March 4–6, 2002, Department of
Agronomy, CCS Haryana Agricultural University, Hisar-125 004, India.
165
Jaramillo, J., Chapman, E.G., Vega, F.E., Harwood, J.D., 2010. Molecular diagnosis of a
previously unreported predator–prey association in coffee: Karnyothrips flavipes
Jones (Thysanoptera: Phlaeothripidae) predation on the coffee berry borer.
Naturwissenschaften 97, 291–298.
Jones, R.W., Gilstrap, F.E., Andrews, K.L., 1988. Biology and life tables for the
predaceous earwig, Doru taeniatum (Dermaptera: Forficulidae). Entomophaga
33, 43–54.
Jonsson, M., Wratten, S.D., Landis, D.A., Gurr, G.M., 2008. Recent advances in
conservation biological control of arthropods by arthropods. Biological Control
45, 172–175.
Kahuthia-Gathu, R., Lohr, B., Poehling, H.M., 2008. Effect of common wild crucifer
species of Kenya on fitness of two exotic diamondback moth parasitoids, Cotesia
plutellae and Diadegma semiclausum. Crop Protection 27, 1477–1484.
Kartohardjono, A., Heinrichs, E.A., 1984. Populations of the brown planthopper,
Nilaparvata lugens (Stal) (Homoptera: Delphacidae), and its predators on rice
varieties with different levels of resistance. Environmental Entomology 13,
359–365.
Karungi, J., Ekbom, B., Kyamanywa, S., 2006. Effects of organic versus conventional
fertilizers on insect pests, natural enemies and yield of Phaseolus vulgaris.
Agriculture, Ecosystems & Environment 115, 51–55.
Kenmore, P.E., Carino, F.O., Perez, C.A., Dyck, V.A., Guttierez, A.P., 1984. Population
regulation of the rice brown planthopper (Nilaparvata lugens Stal) within rice
fields in the Philippines. Journal of Plant Protection in the Tropics 1, 19–37.
Kfir, R., 2003. Biological control of the potato tuber moth Phthorimaea operculella in
Africa. In: Neuenschwander, P., Borgemeister, C., Langewald, J. (Eds.), Biological
Control in IPM Systems in Africa. CABI International, Wallingford, UK, pp. 77–
85.
Kfir, R., Overholt, W.A., Khan, Z.R., Polaszek, A., 2002. Biology and management of
economically important Lepidopteran cereal stem borers in Africa. Annual
Review of Entomology 47, 701–731.
Kiers, E.T., Leakey, R.R.B., Izac, A.M., Heinemann, J.A., Rosenthal, E., Nathan, D.,
Jiggins, J., 2008. Agriculture at a crossroads. Science 320, 320–321.
Kiritani, K., Nakasuji, F., 1977. Battle with Insect Pests: From Control Towards
Management. NHK Publishing Company, Tokyo.
Koji, S., Khan, Z.R., Midega, C.A.O., 2007. Field boundaries of Panicum maximum as a
reservoir for predators and a sink for Chilo partellus. Journal of Applied
Entomology 131, 186–196.
Landis, D.A., Wratten, S.D., Gurr, G.M., 2000. Habitat management to conserve
natural enemies of arthropod pests in agriculture. Annual Review of
Entomology 45, 175–201.
Lavandero, B.I., Wratten, S.D., Didham, R.K., Gurr, G.M., 2006. Increasing floral
diversity for selective enhancement of biological control agents: a double-
edged sward? Basic and Applied Ecology 7, 236–243.
Lavandero, B., Wratten, S., Shishehbor, P., Worner, S., 2005. Enhancing the
effectiveness of the parasitoid Diadegma semiclausum (Helen): movement
after use of nectar in the field. Biological Control 34, 152–158.
Legg, J., Gerling, D., Neuenschwander, P., 2003. Biological control of whiteflies in
sub-Saharan Africa. In: Neuneschwander, P., Borgemeister, C., Langewald, J.
(Eds.), Biological Control in IPM Systems in Africa. CABI International,
Wallingford, UK, pp. 87–100.
Letourneau, D.K., 1998. Conservation biology: lessons for conserving natural
enemies. In: Barbosa, P. (Ed.), Conservation Biological Control. Academic
Press, San Diego, pp. 9–38.
Letourneau, D.K., Armbrecht, I., Salguero, B., Montoya, J., Jimenez, E., Daza, M.C.,
Escobar, S., Galindo, V., Gutierrez, C., Duque, S., Lopez, J., Acosta, A.M., Herrera, J.,
Rivera, L., Saavedra, C.A., Torres, A.M., Reyes, A., 2011. Does plant diversity
benefit agroecosystems? A synthetic review. Ecological Applications 21, 9–21.
Liang, W., Huang, M., 1994. Influence of citrus orchard ground cover plants on
arthropod communities in China: a review. Agriculture, Ecosystems &
Environment 50, 29–37.
Losey, J.H., Vaughan, M., 2006. The economic value of ecological services provided
by insects. BioScience 56, 311–323.
Lou, Y.G., Cheng, J.A., 2003. Role of rice volatiles in the foraging behaviour of the
predator Cyrtorhinus lividipennis for the rice brown planthopper Nilaparvata
lugens. BioControl 48, 73–86.
Lovei, G.L., Andow, D.A., Arpaia, S., 2009. Transgenic insecticidal crops and natural
enemies: a detailed review of laboratory studies. Environmental Entomology
38, 293–306.
Lu, X.Y., Xu, J., Li, Q.Y., 2008. Impact of Lolium perenne planted in chestnut orchards
and its mowing methods on population densities of Oligonychus ununguis and
its natural enemy Amblyseius castaneae. Chinese Journal of Biological Control 24,
108–111.
Lu, Y.H., Wu, K.M., Jiang, Y.Y., Guo, Y.Y., Desneux, N., 2012. Widespread adoption of
Bt cotton and insecticide decrease promotes biocontrol services. Nature 487,
362–365.
Lundgren, J.G., 2009. Relationships of Natural Enemies and Non-prey Foods.
Springer Verlag, Dordrecht, The Netherlands.
Lundgren, J.G., Gassmann, A.J., Bernal, J., Duan, J.J., Ruberson, J., 2009. Ecological
compatibility of GM crops and biological control. Crop Protection 28, 1017–
1030.
Mailloux, J., Le Bellec, F., Kreiter, S., Tixier, M.S., Dubois, P., 2010. Influence of ground
cover management on diversity and density of phytoseiid mites (Acari:
Phytoseiidae) in Guadeloupean citrus orchards. Experimental and Applied
Acarology 52, 275–290.
Mao, S.C., 2010. Report on China‘s Cotton Production Prosperity 2011. Chinese
Agriculture Press.
166 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
Marchioro, C.A., Foerster, L.A., 2012. Importance of carbohydrate sources to the
reproductive output of the wheat armyworm Pseudaletia sequax. Agricultural
and Forest Entomology 14, 29–35.
Martin, J.F., Roy, E.D., Diemont, S.A.W., Ferguson, B.G., 2010. Traditional Ecological
Knowledge (TEK): Ideas, inspiration, and designs for ecological engineering.
Ecological Engineering 36, 839–849.
Maoz, Y., Palevsky, E., Gal, S., Zilberstein, M., Noy, M., Izhar, Y., Ganmor, J.A.S., Coll,
M., 2009. Integrated pest management of Oligonychus perseae: developing
action thresholds and the identification and conservation of natural enemies.
In: Proceedings of the IOBC/WPRS working group ‘‘Integrated control of plant-
feeding mites’’, Florence, Italy, 9–12 March 2009. IOBC/WPRS Bulletin 50, 57–
60.
Matteson, P.C., 2000. Insect pest management in tropical Asian irrigated rice.
Annual Review of Entomology 45, 549–574.
McLeod, A., Wratten, S.D., Sotherton, N.W., Thomas, M.B., 2004. ‘Beetle banks’ as
refuges for beneficial arthropods in farmland: long-term changes in predator
communities and habitat. Agricultural and Forest Entomology 6, 147–154.
Medeiros, M.A., Ribeiro, P.A., Morais, H.C., Castelo Branco, M., Suji, E.R., Salgado-
Laboriau, M.L., 2010. Identification of plant families associated with the
predators Chrysoperla externa (Hagen) (Neuroptera: Chrysopidae) and
Hippodamia convergens Guérin-Menéville (Coleoptera: Coccinelidae) using
pollen grain as a natural marker. Brazalian Journal of Biology 70, 293–300.
Mensah, R.K., 1996. Suppression of Helicoverpa spp. (Lepidoptera: Noctuidae)
oviposition by use of the natural enemy food supplement Envirofeast.
Australian Journal of Entomology 35, 323–329.
Morales, H., Perfecto, I., 2000. Traditional knowledge and pest management in the
Guatemalan highlands. Agriculture and Human Values 17, 49–63.
Morrill, W.L., Almazan, L.P., 1990. Effect of host plant species and age of rice bug
(Hemiptera: Alydidae) eggs on parasitism by Gryon nixoni (Hymenoptera:
Scelionidae). Journal of Entomological Science 25, 450–452.
Naranjo, S.E., 2001. Conservation and evaluation of natural enemies in IPM systems
for Bemisia tabaci. Crop Protection 20, 835–852.
Naranjo, S.E., Ellsworth, P.C., 2009. Fifty years of the integrated control concept:
moving the model and implementation forward in Arizona. Pest Management
Science 65, 1267–1286.
Narvaez, A., Cancino, J., Canal Daza, N., Wyckhuys, K.A.G., 2012. Effect of different
dietary resources on longevity, carbohydrate metabolism, and ovarian
dynamics in two fruit fly parasitoids. Arthropod-Plant Interactions 6,
361–374.
Ndemah, R., Schulthess, F., Phoeling, H.M., Borgemeister, C., 2001. Natural enemies
of lepidopterous borers on maize and elephant grass in the forest zone of
Cameroon with special reference to Busseola fusca (Fuller) (Lepidoptera:
Noctuidae). Bulletin of Entomological Research 91, 205–212.
Neuenschwander, P., 2010. Importance of biological control for tropical Africa. J.
Kulturpflanzen 62, 97–101.
Neuenschwander, P., Borgemeister, C., Langewald, J., 2003. Biological Control in IPM
Systems in Africa. CAB International, Wallingford, UK.
Neuenschwander, P., Hammond, W.N.O., Ajuonu, O., Gado, A., Echendu, N.,
Bokonon-Ganta, A.H., Allomasso, R., Okon, I., 1990. Biological control of the
cassava mealybug, Phenacoccus manihoti (Hom., Pseudococcidae) by
Epidinocarsis lopezi (Hym., Encyrtidae) in West Africa, as influenced by
climate and soil. Agriculture, Ecosystems & Environment 32, 39–55.
Nicolson, S.W., Nepi, M., Pacini, E., 2010. Nectaries and Nectar. Springer.
Nilsson, U., 2011. Conservation biological control of insect pests in two horticultural
crops. PhD thesis, Swedish University of Agricultural Sciences, Alnarp, Sweden.
Norris, R.F., Kogan, M., 2000. Interactions between weeds, arthropod pests, and their
natural enemies in managed ecosystems. Weed Science 48, 94–158.
Nunes, C., Lucas, E., Coderre, D., 2005. Diagnóstico sobre el conocimiento y manejo
de Bemisia tabaci por los productores del norte nicaragüense. Manejo Integrado
de Plagas y Agroecologia (Costa Rica) 76, 75–79.
Nwilene, F.E., Togola, A., Agunbiade, T.A., Ogah, E.O., Ukwungwu, M.N., Hamadoun,
A., Kamara, S.I., Dakouo, D., 2008. Parasitoid biodiversity conservation for
sustainable management of the African rice gall midge, Orseolia oryzivora
(Diptera: Cecidomyiidae) in lowland rice. Biocontrol Science and Technology
18, 1075–1081.
O’Dowd, D.J., 1979. Foliar nectar production and ant activity on a neotropical tree,
Ochroma pyramidale. Oecologia 43, 233–248.
Olkowski, W., Zhang, A., 1998. Habitat management for biological control, examples
from China. In: Pickett, C.H., Bugg, R.L. (Eds.), Enhancing Biological Control:
Habitat Management to Promote Natural Enemies of Agricultural Pests.
University of California Press, Berkeley, pp. 255–270.
Onzo, A., Hanna, R., Negloh, K., Toko, M., Sabelis, M.W., 2005. Biological control of
cassava green mite with exotic and indigenous phytoseiid predators—Effects
of intraguild predation and supplementary food. Biological Control 33, 143–
152.
Patt, J.M., Hamilton, G.C., Lashomb, J.H., 1997. Foraging success of parasitoid wasps
on flowers: interplay of insect morphology, floral architecture and searching
behavior. Entomologia Experimentalis et Applicata 83, 21–30.
Peng, S.J., 1983. Biological control: one of the fine traditions of ancient Chinese
agricultural techniques. Scientia Agricultura Sinica 1, 92–98.
Pereira, J.L., Picanco, M.C., Pereira, E.J., Silva, A.A., Jakelaitis, A., Pereira, R.R., Xavier,
V.M., 2010. Influence of crop management practices on bean foliage arthropods.
Bulletin of Entomological Research 100, 679–688.
Pereira, J.L., Picanco, M.C., Silva, A.A., Barros, E.C., Xavier, V.M., Gontijo, P.C., 2007.
Effect of herbicides on soil arthropod community of bean cultivated under no-
tillage and conventional systems. Planta Daninha 25, 61–69.
Perfecto, I., Castiñeiras, A., 1998. Deployment of the predaceous ants and their
conservation in agroecosystems. In: Barbosa, P. (Ed.), Conservation Biological
Control. Academic Press, San Diego, California, pp. 269–289.
Pfiffner, L., Luka, H., Schlatter, C., Juen, A., Truagott, M., 2009. Impact of wildflower
strips on biological control of cabbage lepidopterans. Agriculture, Ecosystems &
Environment 129, 310–314.
Pimentel, D., Wilson, C., McCullum, C., Huang, R., Dwen, P., Flack, J., et al., 1997.
Economic and environmental benefits of biodiversity. BioScience 47, 747–757.
Pretty, J., Toulmin, C., Williams, S., 2011. Sustainable intensification in African
agriculture. International Journal of Agricultural Sustainability 9, 5–24.
Pu, T.S., Zeng, T., Wei, D.W., 1991. An integrated evaluation on 20 plant pollen as
diet for mass rearing Amblyseius makuwa (Acari: Phytoseiidae). Chinese Journal
of Biological Control 7, 111–114.
Rabb, R.L., Stinner, R.E., van den Bosch, R., 1976. Conservation and augmentation of
natural enemies. In: Huffaker, C.B., Messenger, P.S. (Eds.), Theory and Practice of
Biological Control. Academic Press, New York, pp. 23–254.
Rao, G.V.R., 2005. Integrated pest management in legumes: present status and
future needs. IPM research at ICRISAT: present status and future priorities.
Proceedings of the in-house review, 5th April 2005.
Rekha, B.S., Ramkumar, J., Kandibane, M., Raguraman, S., Swamiappan, M., 2009.
Diversity of Coccinellids in cereals, pulses, vegetables and in weeded and
partially-weeded rice-cowpea ecosystems in Madurai District of Tamil Nadu.
Madras Agricultural Journal 96, 251–264.
Rickson, F.R., Rickson, M.M., 1998. The cashew nut, Anacardium occidentale
(Anacardiaceae) and its perennial association with ants: extrafloral nectary
location and the potential for ant defense. American Journal of Botany 85, 835–
849.
Robertson, G.P., Swinton, S.M., 2005. Reconciling agricultural productivity and
environmental integrity: a grand challenge for agriculture. Frontiers in Ecology
and the Environment 3, 38–46.
Romeis, J., Meissle, M., Bigler, F., 2006. Transgenic crops expressing Bacillus
thuringiensis toxins and biological control. Nature Biotechnology 24, 63–71.
Romeis, J., Sharma, H.C., Sharma, K.K., Das, S., Sarmah, B.K., 2004. The potential of
transgenic chickpeas for pest control and possible effects on non-target
arthropods. Crop Protection 23, 923–938.
Rosado, M.C., 2007. Plantas favoraveis a agentes de controle biologico. MSc thesis,
Universidade Federal de Vicosa.
Saidov, N.S., Jalilov, A.U., Landis, D.A., Fiedler, A., Nazirov, V.K., Mirzoev, T.,
Karimjanov, C., 2011. Enrichment of agro landscapes by flowering plants for
increasing of efficiency of natural enemies in integrated pest management of
cotton. In: Proceedings of scientific work of Research Institute of Plant
Production of the Academy of Agricultural Sciences of the Republic of
Tajikistan. Dushanbe 2011, vol. 6, pp. 106–115.
Saidov, N.S., Landis, A.D., 2008. Evaluation of flowering plants to attract natural
enemies in Tajikistan. In: Proceeding of the Conference devoted to 50 years
Kazakh Research Institute of Plant Protection and Quarantine, Almata,
Kazakhstan, November 2008, vol. I, pp. 127–132.
Salick, J., 1983. Natural history of crop-related wild species: uses in pest habitat
management. Environ. Manag. 7, 85–90.
Santos-Neto, J.R., Mezencio, J.M.S., Chagas, A.T.A., Michereff-Filho, M., Serrao, J.E.,
2010. Use of serological techniques for determination of Spodoptera frugiperda (J
E Smith) predators (Lepidoptera: Noctuidae). Neotropical Entomology 39, 420–
423.
Schulthess, F., Chabi-Olaye, A., Goergen, G., 2001. Seasonal flucturations of noctuid
stemborer egg parasitism in southern Benin with special reference to Sesamia
calamistis Hampson (Lepidoptera: Noctuidae) and Telenomus spp. (Hymnoptera:
Scelionidae) on maize. Biocontrol Science and Technology 11, 765–777.
Scoones, I., Thompson, J., 1994. Beyond Farmer First - Rural People’s Knowledge,
Agricultural Research and Extension Practice. Intermediate Technology
Publication, London.
Sekamatte, M.B., Kyamanywa, S., Willson, H.R., Russell-Smith, A., 2002. Effect of
placement method and rate of application of crushed fish bones on the activity
of predatory ants and impact on termite damage to maize. International Journal
of Tropical Insect Science 22, 199–204.
Sekamatte, M.B., Ogenga-Latigo, M., Russel-Smith, A., 2001. The potential of
protein- and sugar-based baits to enhance predatory ant activity and reduce
termite damage to maize in Uganda. Crop Protection 20, 653–662.
Settle, W.H., Ariawan, H., Astuti, E.T., Cahyana, W., Hakim, A.L., Hinayana, D., Lestari,
A.S.P., 1996. Managing tropical rice pests through conservation of generalist
natural enemies and alternative prey. Ecology 77, 1975–1988.
Shaltiel, L., Coll, M., 2004. Reduction of pear psylla damage by the predatory bug
Anthocoris nemoralis (Heteroptera: Anthocoridae): the importance of orchard
colonization time and neighboring vegetation. Biocontrol Science and
Technology 14, 811–821.
Shannag, H.K., Obeidat, W.M., 2008. Interaction between plant resistance and
predation of Aphis fabae (Homoptera: Aphididae) by Coccinella septempunctata
(Coleoptera: Coccinellidae). Annals of Applied Biology 152, 331–337.
Sharma, R.P., Yadav, R.P., 1993. Response of lentil varieties to the incidence of bean
aphid (Aphis craccivora Koch.) and its predatory coccinellids. LENS Newsletter
20, 60–62.
Sharma, A., Agarwal, V.K., 2007. Role of predatory mites in IPM of phytophagous
mites. In: Jain, P.C., Bhargava, M.C. (Eds.), Entomology: Novel Approaches. New
India Publishing Agency, New Delhi, pp. 453–459.
Simon, S., Bouvier, J.C., Debras, J.F., Sauphanor, B., 2010. Biodiversity and pest
management in orchard systems: a review. Agronomy for Sustainable
Development 30, 139–152.
 K.A.G. Wyckhuys et al. / Biological Control 65 (2013) 152–167
Soroushmehr, Z., Sahragard, A., Salehi, L., 2008. Comparative life table statistics for
the ladybeetle Scymnus syriacus reared on the green citrus aphid, Aphis
spiraecola, fed on two host plants. Entomological Science 11, 281–288.
Sueldo de Escano, M.R., Virla, E.G., 2009. Doru lineare (Dermaptera: Fortifuclidae),
insecto benéfico en cultivos de maíz del norte argentino: preferencias
alimenticias y tasas de consumo. Boletin de Sanidad Vegetal 35, 39–47.
Sumberg, J.E., Okali, C., 1997. Farmer’s Experiments: Creating Local Knowledge.
Lynne Rienner, Boulder, Colorado.
Tamo, M., Arodokoun, D.Y., Zenz, N., Tindo, M., Agboton, C., Adeoti, R., 2002. The
importance of alternative host plants for the biological control of two key
cowpea insect pests, the pod borer Maruca vitrata (Fabricius) and the flower
thrips Megalurothrips sjostedti (Trybom). In: Fatokun, C.A., Tarawali, S.A., Singh,
B.B., Kormawa, P.M. Tamò, M. (Eds.), Challenges and opportunities for
enhancing sustainable cowpea production. Proceedings of the World Cowpea
Conference, 4–8 September, 2000. IITA. Ibadan, Nigeria, pp. 81–93.
Tamo, M., Bottenberg, H., Arodokoun, D.Y., Adeoti, R., 1997. The feasibility of
biological control of two major cowpea insect pests. In: Singh, B.N., Mohan Raj,
D.R., Dashiell, K.E., Jackai, L.E.N. (Eds.), Advances in Cowpea Research. IITA,
Ibadan, Nigeria.
Tandon, P.L., 2001. Negative aspects of interaction between host plant resistance
and biological control and its implication in integrated pest management of
crops. In: Upadhyay, R.K., Mukerji, K.G., Chamola, B.P. (Eds.), Biocontrol
Potential and its Exploitation in Sustainable Agriculture: Insect Pests. Kluwer
Academic/Plenum Publishers, New York.
Taylor, T.H.C., 1945. Recent investigations of Antestia spp. in Uganda. Part I. East
African Agricultural Journal 10, 223–233.
Thorpe, W.H., Caudle, H.B., 1938. A study of the olfactory responses of insect
parasites to the food plant of their host. Parasitology 30, 523–528.
Tixier, P., Lavigne, C., Alvarez, S., Gauquier, A., Blanchard, M., Ripoche, A., Achard, R.,
2011. Model evaluation of cover crops, application to 11 species for banana
cropping systems. European Journal of Agronomy 34, 53–61.
Tscharntke, T., Bommarco, R., Clough, Y., Crist, T.O., Kleijn, D., Rand, T.A., Tylianakis,
J.M., van Nouhuys, S., Vidal, S., 2008. Reprint of ‘‘Conservation biological control
and enemy diversity on a landscape scale’’. Biological Control 45, 238–253.
Toko, M., O’Neil, R.J., Yaninek, J.S., 1994. Effect of cassava exudate and prey densities
on the survival and reproduction of Typhlodromalus limonicus (Garman &
McGregor) s.l. (Acari: Phytoseiidae), a predator of the cassava green mite,
Mononychellus tanajoa (Bondar) (Acari:Tetranychidae). Experimental and
Applied Acarology 18, 221–231.
Tompkins, J.M.L., Wratten, S.D., Wackers, F.L., 2010. Nectar to improve parasitoid
fitness in biological control: does the sucrose:hexose ratio matter? Basic and
Applied Ecology 11, 264–271.
Trujillo, J., Altieri, M.A., 1990. A comparison of aphidophagous arthropods on maize
polycultures and monocultures, in Central Mexico. Agriculture, Ecosystems &
Environment 31, 337–349.
Tuncbilek, A.S., Cinar, N., Campolat, U., 2010. Comparing flower nectar and artificial
diet on the longevity and progeny production of Trichogramma turkestanica. In:
Proceedings of the 10th International Working Conference on Stored Product
Protection. Julius-Kühn-Archiv, pp. 425, 754.
van Emden, H.F., 1965. The role of uncultivated land in the biology of crop pests and
beneficial insects. Scientia Horticulturae 17, 121–136.
van den Bosch, R., Telford, A.D., 1964. Environmental modification and biological
control. In: DeBach, P. (Ed.), Biological Control of Pests and Weeds. Reinhold,
New York, pp. 459–488.
Van Mele, P., 2008. A historical review of research on the weaver ant Oecophylla in
biological control. Agricultural and Forest Entomology 10, 13–22.
Van Mele, P., Cuc, N.T.T., 2000. Evolution and status of Oecophylla smaragdina
(Fabricius) as a pest control agent in citrus in the Mekong Delta Vietnam.
International Journal of Pest Management 46, 295–301.
Van Mele, P., van Lenteren, J.C., 2002. Survey of current crop management practices
in a mixed-ricefield landscape, Mekong Delta, Vietnam—potential of habitat
manipulation for improved control of citrus leafminer and citrus red mite.
Agriculture, Ecosystems & Environment 88, 35–48.
Vázquez, L.L., Fernández, E., Alfonso, E., 2007. Desarrollo de prácticas de manejo de
la diversidad de plantas en sistemas de producción de la agricultura urbana.
Agric. Orgánica (La Habana) 3, 39–41.
Vázquez, L.L., Matienzo, Y., Veitia, M., Alfonso, J., 2008. Manejo y conservación de
enemigos naturales de insectos fitófagos en los sistemas agrícolas de Cuba.
CIDISAV, Habana.
Vinatier, F., Lescourret, F., Duyck, P.F., Tixier, P., 2012. From IBM to IPM: Using
individual-based models to design the spatial arrangement of traps and crops in
integrated pest management strategies. Agriculture, Ecosystems & Environment
146, 52–59.
Waage, J.K., Schulthess, F., 1989. Relevant research activities in support of
sustainable pest management. In: Yaninek, J.S., Herren, H.R. (Eds.), Biological
control: a sustainable solution to crop pest problems in Africa. Proceedings of
167
the inaugural conference and workshop of the IITA Biological Control Program
Center for Africa, 5–9 December 1998, Cotonou, People’s Republic of Benin, pp.
133–140.
Wackers, F.L., 2004. Assessing the suitability of flowering herbs as parasitoid food
sources: flower attractiveness and nectar accessibility. Biological Control 29,
307–314.
Wackers, F.L., Romeis, J., van Rijn, P., 2007. Nectar and pollen feeding by insect
herbivores and implications for multitrophic interactions. Annual Review of
Entomology 52, 301–323.
Wackers, F.L., van Rijn, P.C.J., Bruin, J., 2005. Plant-provided Food for Carnivorous
Insects: A Protective Mutualism and its Applications. Cambridge University
Press, Cambridge, UK.
Wade, M.R., Zalucki, M.P., Wratten, S.D., Robinson, K.A., 2008. Conservation
biological control of arthropods using artificial food sprays: Current status
and future challenges. Biological Control 45, 185–199.
Wang, F., Zhang, S., Hou, S., 1988. Inoculative release of Trichogramma dendrolimi
in vegetable gardens to regulate populations of cotton pests. In: Voegele, J.,
Waage, J., van Lenteren, J.C. (Eds.), Trichogramma and other egg parasites.
Second International Symposiom on Trichogramma, Guangzhou, China. Les
Colloques de l’INRA 43, pp. 613–621.
Wardle, D.A., 1995. Impacts of disturbance on detritus food webs in agro-
ecosystems of contrasting tillage and weed management practices. Advances
in Ecological Research 26, 105–185.
Way, M.J., Heong, K.L., 1994. The role of biodiversity in the dynamics and
management of insect pests of tropical irrigated rice – a review. Bulletin of
Entomological Research 84, 567–587.
Way, M.J., van Emden, H.F., 2000. Integrated pest management in practice –
pathways towards successful application. Crop Protection 19, 81–103.
Woltz, J.M., Isaacs, R., Landis, D.A., 2012. Landscape structure and habitat
management differentially influence insect natural enemies in an agricultural
landscape. Agriculture, Ecosystems & Environment 152, 40–49.
Wyckhuys, K.A.G., Lopez, F., Rojas, M., Ocampo, J., 2011. The relationship of farm
surroundings and local infestation pressure to pest management in cultivated
Passiflora species in Colombia. International Journal of Pest Management 57, 1–10.
Wyckhuys, K.A.G., O’Neil, R.J., 2007a. Agro-ecological knowledge and its
relationship to farmers’ pest management decision making in rural Honduras.
Agriculture and Human Values 24, 307–321.
Wyckhuys, K.A.G., O’Neil, R.J., 2007b. Influence of extra-field characteristics to
abundance of key natural enemies of Spodoptera frugiperda Smith (Lepidoptera:
Noctuidae) in subsistence maize production. International Journal of Pest
Management 53, 89–99.
Wyckhuys, K.A.G., Strange-George, J.E., Kulhanek, C.A., Wackers, F.L., Heimpel, G.E.,
2008. Sugar feeding by the aphid parasitoid Binodoxys communis: How does
honeydew compare with other sugar sources? Journal of Insect Physiology 54,
481–491.
Xing, J.M., Feng, S.L., Fan, X.H., Li, B.L., 1991. The application of Bt emulsion against
cotton bollworm and its preserving effect for natural enemies to suppress
summer cotton aphid. Acta Phytophylacica Sinica 18, 206–209.
Yadouleton, A., Martin, T., Padonou, G., Chandre, F., Asidi, A., Djogbenou, L., Dabiré,
R., Aikpon, R., Boko, M., Glitho, I., Akogbeto, M., 2011. Cotton pest management
practices and the selection of pyrethroid resistance in Anopheles gambiae
population in Northern Benin. Parasites & Vectors 4, 1–11.
Yaninek, J.S., Cock, M.J.W., 1989. Identifying pest problems in relation to
implementing biological control in Africa. In: Yaninek, J.S., Herren, H.R. (Eds.),
Biological control: a sustainable solution to crop pest problems in Africa.
Proceedings of the inaugural conference and workshop of the IITA Biological
Control Program Center for Africa, 5–9 December 1998, Cotonou, People’s
Republic of Benin, pp. 116–126.
Yu, D.S., Byers, J.R., 1994. Inundative release of Trichogramma brassicae (Hym.:
Trichogrammatidae) for control of European corn borer in sweet corn. Canadian
Entomolology 12, 291–301.
Zhang, R.Z., Liang, H.B., Tian, C.Y., Zhang, G.X., 2000. Biological mechanism of
controlling cotton aphid (Homoptera: aphididae) by the marginal alfalfa zone
surrounding cotton field. Chinese Science Bulletin 45, 355–358.
Zhang, Y.B., Liu, W.X., Wang, W., Wan, F.H., Li, Q., 2011. Lifetime gains and patterns
of accumulation and mobilization of nutrients in females of the synovigenic
parasitoid, Diglyphus isaea Walker (Hymenoptera: Eulophidae), as a function of
diet. Journal of Insect Physiology 57, 1045–1052.
Zhou, D.R., Song, Y.Y., He, K.L., Wang, Z.Y., Zheng, L., Zhang, G.Y., 1997. Study on the
preference habitat of Trichogramma ostriniae: I. Natural population distribution
and dynamics of the parasitoid in different habitats. Chinese Journal of
Biological Control 13, 1–5.
Zundel, C., Nagel, O., Hanna, R., Korner, F., Scheidegger, U., 2009. Environment and
host-plant genotype effects on the seasonal dynamics of a predatory mite on
cassava in sub-humid tropical Africa. Agricultural and Forest Entomology 11,
321–331.
Appendix 1. Documentation of non-prey food sources for natural enemies
Country Target crop / pest Natural enemy Type of food source Effect Reference
Argentina Maize / fall Doru lineare (Derm.: Maize pollen, milky maize grains, armyworm Feeding preference for eggs, Sueldo de Escano
armyworm Forficulidae) eggs development on pollen & Virla, 2009
Benin Cassava / cassava Amblyseius idaeus, Pollen of Zea mays, Leucaena leucocephala, Maize pollen and cassava nectar Gnanvossou et al.,
green mite Typhlodromalus manihoti, Ricinus communis, cassava extra-floral nectar enhance mite development 2005
Typhlodromalus aripo (Acari:
Phytoseiidae)
Cassava / cassava Euseius fustis (Acari: Pollen of maize, castor bean and cassava, Maize plus castor bean pollen Bruce-Oliver et al.,
green mite, cotton red Phytoseiidae) cassava extra-floral nectar, mealybug increased survival and reproduction 1996; Onzo et al.,
mite honeydew 2005
Brazil Coffee / various Chrysoperla externa (Neur.: Mixed flower honey, pollen of pigeon pea, Access to pollen and honey increased Venzon et al., 2006
Chrysopidae) sunhemp or castor bean reproductive success and
development
Various Chrysoperla externa (Neur.: Buckwheat floral nectar, castor bean Nectar increased survival, pollen Rosado, 2007
Chrysopidae) extrafloral nectar and sunn hemp pollen increased survival but not fecundity
Various / Fall Telenomus remus (Hym.: Mixed flower honey, glucose, fructose and Honey and single sugar solutions Meirelles et al.,
armyworm Scelionidae) sucrose improved parasitism 2009
Cameroon Maize, sorghum / Diaperasticus erythrocephala Maize pollen Permits rapid development Aroga, 2003
maize stalk borer (Derm. : Forficulidae)
Chile Pine trees / Pine bud Orgilus obscurator (Hym.: Mixed flower honey, flowers of Conium Improved longevity Ide & Lanfranco,
moth Braconidae) maculatum and Daucus carota 2001
China Coconut / coconut Tetrastichus brontispae (Hym.: Mixed flower honey, sucrose and glucose Increased longevity, parasitism and Chen et al., 2010
hispid Eulophidae) solutions fecundity
Maize / Asian corn Macrocentris linearis (Hym.: Corn pollen dissolved in water Increased longevity Lu & Yang, 1995
borer Braconidae)
Unspecified Amblyseius makuwa (Acari: Pollen of 14 plant species Improved development and Pu et al., 1991
Phytoseiidae) reproduction for pollen from
Cucumis sativus, Jasminum sumbad,
Lantana camara
Various Meteorus pulchricornis (Hym.: Varying concentration of sucrose-glucose- Greatest longevity on 30% solution Wu et al., 2008
Braconidae) fructose mixture
Various / leafminer Diglyphus isaea (Hym.: Host feeding, mixed honey solution Greater levels of body nutrients on Zhang et al., 2011
Eulophidae) honey
Rice / rice brown Anagyrus nilaparvatae (Hym.: Mixed flower honey, corn pollen, soybean Honey, pollen and nectar increased Zheng et al., 2003
planthopper Mymaridae) floral nectar, planthopper honeydew longevity
Rice / rice leaffolder Apanteles cypris (Hym.: Mixed flower honey Increased longevity Cheng, 1984
Braconidae)
Cotton / cotton Microplitis mediator (Hym.: Sucrose, glucose, fructose, mannose, Differing effects on longevity, Luo et al., 2010
bollworm Braconidae) galactose fecundity and nutrient reserves
Various / Trichogrammatoidea Mixed flower honey, flowers of Brassica Honey and floral nectar increased Guo et al., 2011
 Lepidopteran pests bactrae(Hym.: parachinensis longevity
Trichogrammatidae)
Colombia Guava / Tephritid Diachasmimorpha longicaudata Fruit juice, mixed flower honey, pollen Greater longevity, egg load and body Narvaez et al., 2012
fruit flies (Hym.: Braconidae); Aganaspis nutrient levels on honey and mixed
pelleranoi (Hym.: Figitidae) diets
Bean / whitefly Amitus fuscipennis (Hym.: Extra-floral nectar of several weed species Increased longevity Hernandez et al.,
Platygasteridae) 2009
Cassava / cassava Typhlodromalus limonicus Extrafloral nectar Nectar sustains predator populations, Bakker & Klein,
green mite (Acari: Phytoseiidae) and increases reproduction 1992; Toko et al.,
1994
Egypt Mango / spider mite Typhlodromus mangiferus, Mango powdery mildew, Oidium mangiferae Sustains reproduction in absence of Abou-Awad et al.,
Typhlodromips swirskii (Acari: prey 2011
Phytoseiidae)
Strawberry, Neosieulus barkeri (Acari: Fungi associated with the host plants: Increased survival and development, Momen &
cucumber, eggplant / Phytoseiidae) Aspergillus niger, Alternaria solani, some sustained oviposition Abdelkhader, 2010
spider mite Penicillium digitatum, P. italicum
Various / whitefly Encarsia bimaculata (Hym.: Glucose, fructose, trehalose, trehalulose Increased longevity Mandour et al.,
Aphelinidae) compared to melizitose and honeydew 2007
Various Amblyseius zaheri, Euseius Ricinus communis and Helianthus annuus Depending upon mite species and Momen, 2004
yousefi, Amblyseius lindquisti, pollen grains pollen type, different effects on
Typhlodromus balanites, longevity, fecundity and
Typhlodromus sennarensis, development
Amblyseius cabonus (Acari:
Phytoseiidae)
Honduras Maize / various Doru taeniatum (Derm.: Plant pollen, lepidopteran eggs Most rapid development under Jones et al., 1988
Forficulidae) combined or animal diets
India Pigeonpea / coreid Gryon clavigrallae (Hym.: Mixed flower honey With access to honey, adults live 28- Shanower et al.,
bugs Scelionidae) 96 days, versus 6 days for water-fed 1999
individuals
Iran Various / Black bean Lysiphlebus fabarum (Hym.: Honey, aphid honeydew Greatest longevity on honey Matin et al., 2009
aphid Aphidiidae)
Malaysia Cabbage / Tetrastichus sokolowskii (Hym. Mixed flower honey, 10% sucrose solution Higher longevity, compared to water Ooi, 1988b
Diamondback moth Eulophidae)
Niger Millet / various Forficula senegalensis (Derm. : Pollen from different plant species Heavy reliance on pollen from millet Boukary et al.,
Forficulidae) anthers 1997
Pakistan Various / lepidopteran Trichogramma chilonis (Hym.: Honey, glucose, fructose, sucrose Greatest longevity and parasitism on Saljoqi & Khajjak,
pests Trichogrammatidae) glucose, fructose and sucrose 2007
Philippines Cabbage / Diadegma semiclausum (Hym.: Floral nectar of Zantedeschia albomaculata, Increased longevity Amend &
diamondback moth Ichneumonidae) Galinsoga parvjflora Basedow, 1997
Turkey Citrus / woolly Cales noacki (Hym.: Sugarwater, honey, whitefly honeydew Greatest longevity with sugarwater Sakin & Ulusoy,
whitefly Aphelinidae) 2009
Unspecified Apanteles galleriae (Hym.: Comb without honey, honey solution at 5°%; Greatest longevity with 30% solution Uckan & Ergin,
 Braconidae)
Unspecified Bracon hebetor (Hym.:
Braconidae)
Various / lepidopteran Trichogramma turkestanica
pests (Hym.: Trichogrammatidae)
Uzbekistan Various / winter moth Apanteles congestus,
Microplitis spectabilis (Hym.:
Braconidae), Amblyteles
panzeri (Hym.:
Ichneumonidae), Tachinidae
Venezuela Various / Fall Eiphosoma vitticolle (Hym.:
armyworm Ichneumonidae)
30%, 10% 2003
Mixed flower honey Increased body nutrient levels Gunduz et al., 2010
Nectar of different wildflowers, honey, grape Improved longevity and fecundity Tuncbilek et al.,
molasses, wetted raisins, beet molasses 2010
Sugar solution, nectar of winter cress, carrot Improved longevity and egg Eremenko, 1971
or onion maturation
Mixed flower honey, maize pollen Increase in oviposition period, Giraldo-Vanegas &
augmented fecundity with access to Garcia, 1995
foods
 

Appendix 2. Identification of alternative host plants, naturally occurring food items or alternative prey items for natural enemies
 
Country Natural enemy Alternative host plant, prey item Effect Reference
Benin Parasitoids of cowpea pod borer Native tree Pterocarpus santalinoides Increased parasitism Tamo et al., 2002
Predators and parasitoids of Helicoverpa armigera Sorghum, maize Increased abundance Cherry et al., 2003
Scelionid egg parasitoids Wild grasses Increased parasitism Schulthess et al.,
2001 ; Ndemah et
al., 2001, 2003 ;
Overholt et al., 2003
Brazil Orius sp. (Heteroptera : Anthocoridae) Broad list of weeds and cultivated plants Observed foraging, benefit from refuge, Silveira et al., 2003
pollen or prey
Hippodamia convergens (Col.: Coccinelidae), Asteraceae (H. convergens) and Poaceae (C. Pollen grains attached to insect body Medeiros et al.,
Chrysoperla externa (Neuroptera : Chrysopidae) externa) 2010
Coccinellid beetles Dill, in comparison with coriander and sweet High abundance Lixa et al., 2010
fennel
Lynx spiders Wild plants with glandular trichomes Observed association Vasconcellos-Neto
et al., 2007
Doru luteipes (Derm.: Forficulidae), Nephila clavipes Crotalaria juncea Observed association Tavares et al., 2011
(Aran.: Nephilidae), Orius insidiosus (Het.:
Anthocoridae), Pheidole sp., Solenopsis sp. (Hym.:
Formicidae)
Ants, parasitoids and predators of the coffee Inga sp. trees Association with extra-floral nectaries Rezende, 2010
leafminer
Parasitoids of Anastrepha obliqua (Dipt.: Hog plum (Spondias sp.) High rate of attack of fruitfly Carvalho et al.,
Tephritidae) 2010
China Trichogramma ostriniae (Hym.: Trichogrammatidae) Sweet potato High levels of parasitism Zhou et al., 1997a
Cuba Cycloneda sanguinea (Col. : Coccinellidae), social Wildflowers and flowers of Morinda citrifolia Observed foraging Matienzo et al.,
wasps 2007; 2011
Coffee leafminer parasitoids Flowers of Zebrina pendula, Commelina diffusa Provision of refuge and pollen Simon, 1999
Orius insidiosus (Het.: Anthocoridae) Wildflower Bidens pilosa Observed foraging Veitia, 2008
Lacewings, coccinellids, earwigs, parasitoids, Sunflower Higher population levels Rijo et al., 1999
predatory stinkbugs
Aphid predators such as Cycloneda sanguinea (Col.: Parthenium hysterophorus Population sustained on aphids Veitia, 2008
Coccinellidae)
Parasitoid complex of fall armyworm Several weeds Population reservoir Vazquez, 2009
Several predators Leguminous tree Gliricidia sepium Population reservoir, pollen feeding Vazquez, 2011
Several guilds and species, including Vespidae Weeds, e.g. Parthenium hysterophorus Source of pollen and nectar Fernandez et al.,
2001
El Salvador Paratheresia claripalpis (Dipt.: Tachinidae) Broad leaf weeds Serve as reservoir Argueta &
Hernandez, 2009
Greece Macrolophus caliginosus (Het.: Miridae) Wildflower Dittrichia viscosa Higher population levels Perdikis et al., 2007
Aphidius colemanii, Ephedrus persicae (Hym.: Wild shrubs Vitex agnus castus, Euphoribia Serve as reservoirs Kavallieratos et al.,
Braconidae) characais 2008
Aphidius matricariae, Lysephlebus fabarum (Hym: Wild shrubs Ditricchia viscosa, Rubus Maintain local populations Kavallieratos et al.,
Braconidae) ulmifolius 2002
India Parasitoids of Spodoptera litura Castor bean, as compared to tobacco Higher parasitism Chari, 1985; Rao et
al., 1990
Coccinellid predators of green peach aphid Weed Gynandropsis pentaphylla High abundance Sitaramaiah et al.,
2001
Indonesia Dolichoderus bituberculatus (Hym. : Formicidae) Colonies of the mealybug Planococcus Rapid establishment of ant colonies Graham, 1991
lilacinus
Iran Scymnus syriacus (Col. : Coccinellidae) Spirea sp., compared to Citrus sinensis Shorter development, higher fecundity, Soroushmehr et al.,
(infested with Aphis spiraecola) higher survival 2008
Israel Anthocoris nemoralis (Heteroptera : Anthocoridae) Rhamnus, Laurus, Pistacia trees Greater reproduction, early population Shaltiel & Coll,
build-up 2004
Kenya, West Stem borer parasitoids Wild grasses, Sudan grass Greater efficacy, differing development Khan et al ., 1997 ;
Africa time Ndemah et al.,
2003 ; Setamou et
al., 2005
Cotesia plutellae (Hym.: Braconidae), Diadegma Wild crucifers (Erucastrum arabicum, Differing development time, weight Kahuthia-Gatu et
semiclausum (Hym.: Ichneumonidae) Raphanus raphanistrum, Rorippa micrantha, al., 2008
Rorippa nudiuscula, Brassica juncea)
Mexico Thirty ant species, some of which key predators 102 plant species, Leguminosae being the most Temporal fluctuations in dependence Rico-Gray, 1993
visited
Parasitoids of Anastrepha sp. fruit flies Mesoamerican native fruit trees, belonging to Serve as reservoir Aluja et al., 2003 ;
Melastomataceae, Annonaceae, Apocynaceae, Figueroa de la Rosa
etc. et al., 1998
Insect predators birds in coffee plantation Epiphytes Serve as reservoir Cruz-Angon &
Greenberg 2005
Trichogrammatidae sp. parasitoids of pink bollworm Onobrychis vicrifolia (Fabacea) Serve as reservoir Loya Ramírez et al.,
2003
Whitefly parasitoids: Eretmocerus spp., Encarsia Several wild associated plants Serve as reservoir Bravo Luna et al.,
spp. 2003.
Pakistan Coccinella septempunctata (Col.: Coccinellidae) Cotton mealybug host plants, such as cotton, Greater fitness Sana-Ullah et al.,
lantana, itsit and shoeflower 2011
Philippines Parasitoids of rice bug Alternative hosts such as goosegrass, jungle Lower parasitism than on rice Morrill & Almazan,
rice 1990
Russia Tachinid flies Flowering weeds, belonging to Apiaceae, Observed foraging Rogochaya, 1971
Asteraceae and Cruciferae
South Africa Euseius rubicolus, Neosieulus californicus, Weeds on orchard floor Predator and herbivore records on De Villiers &
Typhlodromus sp. (Acari: Phytoseiidae) different plants Pringle, 2010
Tajikistan, Braconid wasps Wildflowers Ferula assa-foetida, F. Observed foraging Saidov et al., 2007
Turkmenistan badrakema, Senecio subdentatus, Lepidium
draba, Anethum graveolens
Broad range of natural enemies Native flowering plants Observed foraging Saidov & Landis,
2008
Tanzania Parasitoids of Helicoverpa armigera, predatory ants Sorghum, cleome, sunflower Great attractivity to parasitoid, Nyambo, 1986; Van
increased ant foraging on sunflower den Berg et al.,
1990
Turkey Parasitoids of Helicoverpa armigera Several crops, such as tomato Up to 10 times higher parasitism levels Koclu &
Karsavauran, 1998
 
 

Appendix 3. Use of (artificial) food supplements to increase natural enemy efficacy or abundance
Country Target crop / pest
Brazil Unspecified
Colombia Tomato / whitefly
Various / pineapple
mealybug
Cuba Sweet potato / sweet potato
weevil
El Salvador Maize / various
Honduras Maize / Fall armyworm
India Pulses / unspecified
Israel Almond & peppermint /
cotton aphid
Avocado / persea mite
Mexico Various / aphids
Citrus / brown citrus aphid
Wolf Apple / membracids
Malaysia Cocoa / unspecified
Mahogany / shoot borer
Pakistan Cotton / unspecified
Uganda Maize / unspecified
Vietnam Citrus / unspecified
Sapodilla / unspecified
Natural enemy Type of food source Effect Reference
Polybia occidentalis (Hym.: Vespidae) Sugar solution in artificial Increased foraging Hrncir et al., 2007
carbohydrate feeders
Various Sucrose sprays combined with Increased abundance of natural enemies Hernandez et al.,
IGR unpublished
Solenopsis geminata (Hym.: Formicidae) Access to vials with sucrose Reduced homopteran tending Carabali et al.,
solution unpublished
Pheidole megacephala (Hym.: Formicidae) Sugarcane molasses, table Conservation and augmentation of Perfecto &
leftovers predatory ants Castineiras, 1998
Solenopsis geminata (Hym.: Formicidae) Table leftovers, maize bread Conservation of predatory ants Bentley, 2006
Solenopsis geminata (Hym. : Formicidae), Sugarwater Increased abundance, reduced Canas & O’Neil,
Lespesia archippivora (Dipt. : Tachinidae), armyworm infestation 1998
coccinellid beetles
Unspecified Sugar sprays Unspecified Iswaran & Sen,
1972
Tapinoma erraticum (Hym.: Formicidae) Artificial sugar source Reduced homopteran tending Martinez et al.,
2011
Euseius scutalis (Acari: Phytoseiidae) Maize pollen Increased predator populations, Maoz et al., 2009
enhanced mite control
Chrysopa sp. (Neur.: Chrysopidae) Yeast application Increased aphid predation Trujillo &Altieri,
1990
Chrysopidae, Coccinellidae, Syrphidae Powder milk and sugar Attracts and retains predators López Arroyo,
2009a
Camponotus crassus (Hym.: Formicidae) Mixed flower honey Reduced homopteran tending Rico-Gray &
Morais, 2006
Dolichoderus thoracicus (Hym.: Blood clams, fish Increased colony establishment Ho & Khoo, 1997
Formicidae)
Oecophylla smaragdina (Hym.: Live mealworms, fish, honey, Sustain existing and relocated colonies Lim, 2007
Formicidae) sucrose-solution
Chrysoperla carnea (Neur.: Chrysopidae), Protein hydrolysate and sugar Higher abundance of C. carnea and T. Ahmad et al., 2011
Trichogramma chilonis (Hym. : chilonis under combined sprays, higher
Trichogrammatidae), Orius sp. (Het. : Orius sp. under sugar treatments
Anthocoridae)
Pheidolesp.,Lepisiota sp. (Hym.: Protein baits, such as powdered Increased ant nesting, reduced termite Sekamatte et al.,
Formicidae) fish damage and increased maize yield 2001b, 2002
Oecophylla smaragdina (Hym.: Fish, chicken intestines Maintenance of ant colonies Van Mele & Cuc,
Formicidae) 2000
Dolichoderus thoracicus (Hym.: Sugar solution applied to plant Increased distribution, colony Van Mele & Cuc,
Formicidae) residues maintenance during dry season 2001
Appendix 4. Effects of (structural) habitat manipulation, including inter- and cover-cropping practices, on natural enemy abundance and efficacy
Country Target crop / pest Natural enemy Habitat manipulation Effect Reference
Argentina Potato / unspecified Harmonia axyridis (Col.: Weed strips Increased abundance Martinoia &
Coccinellidae) Carmona, 2011
Wheat / unspecified Syrrphidae, Coccinellidae Strips of flowering Brassica Increased abundance Bertolaccini et
campestris, weed strips al., 2008
Soybean / unspecified Generalist predators Weed strips Increased abundance Weyland &
Zaccagnini, 2008
Brazil Onion / onion thrips Parasitoids and predators Strips of marigold Increased abundance, possible pest Silveira et al.,
control 2009
Onion / onion thrips Toxomerus spp. (Dipt. : Syrphidae) Border plants of buckwheat, fodder No difference Goncalves &
radish, carrot, maize, rocket, weeds Sousa, 2003
Cacao / various Parasitoid wasps Richness and density of (shade) tree Increased diversity Sperber et al.,
species 2004
Central Asia – Apple / various Parasitoids in genera Microdus, Areas with nectariferous plants Increased parasitism Vorotynseva,
unspecified Ascogaster, Pimpla 1975
China Apple / various Metaseiulus occidentalis (Acari: Leaf and grass debris on orchard floor Increased overwinter survival Deng et al., 1988
Phytoseiidae)
Apple / wooly apple aphid Aphelinus mali (Hym.: Aphelinidae) Artificial shelter: dry grass attached to Increased overwinter survival Zhou et al., 2010
tree trunk
Apple / various Orius sauteri (Het.: Anthocoridae) Tolerance of weeds, esp. Lamiaceae Increased abundance, pest control Yu et al., 1998
Citrus / various Oecophylla smaragdina (Hym.: Strips of bamboo between trees Increased foraging Huang & Yang,
Formicidae) 1987
Various / grasshoppers Rosy starling (Sturnus roseus) Rock piles, shrubs Increased abundance, lower Olkowski &
grasshopper numbers Zhang, 1998
Wheat, maize / unspecified Ground-dwelling spiders Strips of Amorpha fructose, Vetiveria Increased abundance and diversity Wu et al., 2009
zizanoides, Eulaliopsis binate and within strips
Medicago sativa
Colombia Coffee / unspecified Predatory ants Shaded coffee Greater species diversity, possible Gallego-Ropero
effect on pest pressure et al., 2009
Coffee / unspecified Predatory ants Addition of leaf litter, bamboo twigs Increased nesting opportunities, Armbrecht et al.,
higher abundance 2006
Costa Rica Oil palm / various Various ‘Islands’ and strips of selected weed (Expected) conservation of natural Mexzon &
species enemies Chinchilla, 1998
Coffee / various Predatory ants Shade and conservation of leaf litter Increased species diversity Perfecto &
Vandermeer,
1996
Cuba Various Cycloneda sanuinea, Chilocorus cacti Coconut or guava in field borders Increased abundance Matienzo, 2007
(Col.: Coccinellidae), Chrysopa poeyi
(Neur.: Chrysopidae)
Various Various Tolerance of flowering weeds, e.g., Increased abundance Vazquez et al.,

Bean / unspecified
Eggplant / unspecified
Cucumber / unspecified
Cabbage / mustard aphid
Coffee / coffee berry borer
Sweet potato / sweet potato
weevil
Ecuador Coffee / unspecified
Egypt Sugar beet, cotton /
unspecified
El Salvador Sugar cane/sugarcane borer
Ethiopia Bean / African bollworm
Ghana Cocoa / unspecified
Guatemala Maize / various
India Rice-wheat / various
Chickpea / corn earworm
Rice, Cowpea / unspecified
Ivory Coast Rice / various
Kenya Maize / stemborer
Kenya Maize, bean / various
Malaysia Maize / Asiatic corn borer
Rice / tungro virus vectors
Amaranthus sp., Parthenium 2008
hysterophorus, Argemone mexicana,
Bidens pilosa
Various predators and parasitoids Establishment of maize strips Increased abundance Matienzo, 2005
Various predators and parasitoids Establishment of maize strips Increased abundance Matienzo et al.,
2007
Cycloneda sanguinea (Col.: Tolerance of weeds, esp. Portulaca Increased abundance Matienzo, 2005
Coccinellidae), Lysiphlebus oleraceae
testaceipes (Hym.: Aphidiidae),
Polistes cubensis (Hym.: Vespidae),
syrphids
Diaretiella rapae (Hym.: Braconidae) Mustard strips Increased parasitism Mujica et al.,
2009
Tetramorium bicarinatum (Hym.: Shading with Gliricida sepium, Musa Increased diversity and activity Vazquez et al.,
Formicidae) spp., Theobroma cacao 2009
Pheidole megacephala (Hym.: Temporary nests with banana stems and Increased abundance Perfecto &
Formicidae) rolled leaves Castineiras, 1994
Spiders Orchard abandonment Increased abundance Richter et al.,
2007
Coccinellid and staphylinid beetles Bermuda grass strips Increased abundance Msebah & El-
Husseini, 2009
Paratheresia claripalpis (Dipt. : Broad leaf weed rows Increased abundance Argueta &
Tachinidae) Hernandez 2009
Tachinidae, predatory wasps Tolerance of weeds Increased abundance Abate, 1991
Predatory ants Shaded plots Increased diversity Bigger, 1981
Various Conservation trees Increased abundance Morales &
Perfecto 2000
Spiders, ants, earwigs, coccinellid Grasses and weeds on bunds Increased abundance, colonization Jaipal et al., 2002
beetles
Campoletis chloridae (Hym.: Coriander strips at field border Increased parasitism Pimbert &
Ichneumonidae) Srivastava, 1989
Coccinellid beetles Partial weeding, compared to complete Increased abundance Rekha et al.,
weeding 2009
Predatory ants Tolerance of weeds Increased abundance, no effect on Afun et al., 2000
pest pressure
Earwigs, spiders Strips of Guinea grass Increased abundance Koji et al., 2007
Predatory wasps, spiders, coccinellid Nine different types of hedgerows Increased abundance for certain Girma et al.,
beetles guilds 2000
Trichogramma papilionis (Hym.: Tolerance of weeds in field plots Increase in parasitism of egg Sulaiman et al.,
Trichogrammatidae) masses 2004
Various Weedy bunds without tungro hosts Increased abundance Bottenberg et al.,
1990
 Cocoa / various
Mexico Citrus / brown citrus aphid
Maize / fall armyworm
Beans / bean pod weevil,
Mexican bean beetle
Cotton / pink bollworm
Coffee/ lepidopteran larvaes
Coffee / unspecified
Coffee/ unspecified
Niger Okra / fruit worm
Nigeria Cowpea / cowpea aphid
Rice / African gall midge
Pakistan Wheat / unspecified
Peru Cotton / various
Philippines Rice / various
Rice / various
Russia Cabbage / various
Onion / unspecified
Unspecified / fruit moth,
black pine leaf scale
Cabbage / cabbage moth
Unspecified
South Africa Citrus / phytophagous mites
Dolichoderus thoracicus, D. Deployment polyethylene bags with Colony maintenance Chong & Fee,
bituberculatus (Hym.: Formicidae) leaf litter, leaf bundles, rolls of 1989; Graham,
cardboard 1991; Hosang et
al., 2010
Chrysopidae, Coccinellidae, Establishment of wild sunflower, Increased predator population López Arroyo,
Syrphidae grasses, white flowers 2009b
Ground beetles, parasitoids Tolerance of weeds Increased beetle abundance, lower Penagos et al.,
parasitism, lower pest pressure 2003
Various Increased plant diversity Increased abundance López Arroyo et
al., 1992
Trichogrammatidae Strips of Onobrychis viciifolia Increased abundance Loya Ramírez et
(Fabacea) al 2003.
Insectivorous birds Shade trees Increased predation Perfecto et al,
2004
Predatory ants Shaded plots Increased diversity Philpott et al.,
2006
Predatory ants Artificial bamboo nests Increased diversity Philpott & Foster
2005
Spiders Pigeonpea borders Increased abundance, possibly Ratnadass, 2010
greater predation
Predatory coccinellids and syrphid Tolerance of weeds Increased abundance, less aphid Ofuya, 1989
larvae colonization
Platygaster diplosisae (Hym.: Paspalum scrobiculatum strips at field Increased parasitoid abundance Nwilene et al.,
Platygastridae), Aprostocetus edge 2008
procerae (Hym.: Eulophidae)
Orb web spiders Brassica napus in field borders Increased in-field abundance Butt & Sherawat,
2011
Various Mowing regimes of vegetation in field (Expected) conservation of natural Beingolea, 1959
borders and drainage canals, enemies
maintenance of habitat diversity
Spiders Weeds on rice bunds Source of colonization Sigsgaard, 2000
Web-building and hunting spiders Grazing of weedy rice bunds Lower abundance Barrion, 1999
Different natural enemies Flower strips every 50-60 m Increased fecundity Mikhalsev, 1994
Predatory thrips Early-blooming plants such as hyssop, (Expected) increased abundance Saidov et al.,
celery and spring rape 2007
Ageniaspis sp. (Hym.: Encyrtidae), Strips of dill, buckwheat and Phacelia Increased parasitoid efficacy Saidov et al.,
Aphytis sp. (Hym.: Aphelinidae) sp. 2007
Ernestia sp. (Dipt.: Tachinidae) Strips of flowering carrot, dill and Attraction of parasitoids, increased Saidov et al.,
onion moth parasitism 2007
Trichogramma spp. (Hym.: (Unspecified) nectarious plants Increase in biological control Voronin, 1982
Trichogrammatidae) efficacy
Amblyseius tutsi (Acari: Phytoseiidae) Eucalyptus trees in crop border Increased abundance Grout & Stephen,
 and thrips
South Korea Apple / unspecified
Sri Lanka, India, Cashew / unspecified
Malaysia
Tajikistan Wheat, barley / Sunn pest
Turkey Tomato / unspecified
Ukraine Pea / various
Vietnam Sapodilla / various
Citrus / various
Citrus / various
Fruits / various
Argentina Apple / unspecified
Benin Cowpea / legume pod borer
Brazil Coffee / coffee leaf-miner
Tomato / silverleaf whitefly
Tomato / tomato pinworm
Soybean / unspecified
Vineyards / unspecified
Soybean / various
Cameroon Maize / maize stalk borer
China Apple / unspecified
1995
Neoseiulus makuwa, N. womersleyi Artificial ground shelters, e.g., urethane Increased overwinter survival Kawashima &
(Acari: Phytoseiidae) foam, nets Jung, 2010
Predatory ants Partial cleaning of groundcover and Increased abundance and diversity Rickson &
understory vegetation Rickson, 1998
Microphanurus vassilievi, M. Apiaceae in field borders Increased parasitoid abundance Rubtsov, 1944
semistriatus (Hym.: Scelionidae)
Ground beetles, spiders, ants Tolerance of weeds Increased abundance Yardim &
Edwards, 2002
Different species of natural enemies Phacelia sp. within pea plots Increased natural enemy abundance Galunko &
Dyadechko, 1971
Dolichoderus thoracicus (Hym.: Banana plants in crop border Increased nesting Van Mele & Cuc,
Formicidae) 2001
Natural enemies Tolerance of weeds Increased abundance, superior pest Van Mele, 1998
control
Oecophylla smaragdina (Hym.: Connecting or disconnecting trees Increased foraging range, lower Van Mele & Cuc,
Formicidae) competition 2000
Oecophylla smaragdina (Hym.: Conservation of non-crop trees such as Superior nesting Van Mele & van
Formicidae) Spondias dulcis, Mangifera indica, Lenteren, 2002
Eucalyptus tereticornis, Ceiba
pentandra
Intercropping or strategic use of cover crops
Predaceous beetles, parasitoids, Alfalfa-fescue, strawberry clover as Increased abundance and diversity Fernandez et al.,
predator bugs, lacewings cover crops 2008
Trichogrammatiodea eldaneae Intercropping with sorghum, millet, Increased persistence, eventually Tamo et al., 2002
(Hym.: Trichogrammatidae) maize efficacy
Predatorsand parasitoids of Intercropping with perennial peanut, Increased predation in unshaded, Amaral et al.,
Leucoptera coffeella sunn hemp and lucerne (in shaded and and lower predation in shaded 2010
unshaded systems) system
Various Intercropping with cilantro Increased abundance and diversity Togni, 2009
Spiders, coccinellids, ants Intercropping with coriander and Increased abundance, lower pest Medeiros et al.,
Gallant soldier pressure 2009
Dermaptera Intercropping with corn Increased abundance Cividanes, 2002
Various Black oat and pea as cover crops Increased abundance and diversity Fadini et al.,
2001
Geocoris sp. (Het.: Geocoridae), Intercropping with maize Increased abundance, some pests Cividanes &
Orius sp. (Het.: Anthocoridae), more severe Barbosa, 2001
Braconid and Scelionid wasps
Telenomus sp. (Hym.: Scelionidae) Intercropping with legumes or cassava Increased parasitism, up to two-fold Chabi-Olaye et
al., 2005
Chrysopa sinica (Neur.: Melilotus albus as cover crop Increased abundance Yan & Duan,
Chrysopidae), Amblyseius 1988
 pseudolongispinosus (Acari :
Phytoseiidae)
Apple / unspecified Unspecified Lagopsis supina and Medicago sativa, Stabilized natural enemy Yan et al., 1997
compared to Brassica campestris as communities
cover crops
Apple / European red mite Orius sauteri (Het.: Anthocoridae) Medicago sativa as cover crop Increased abundance Du & Yan, 1994
Apple / Various Chrysopa sinica (Neur.: Lolium perenne, Trifolium repens and Earlier appearance, greater overall Zhao et al., 2011
Chrysopidae), Coccinella Medicago sativa as cover crop species richness
septempunctata (Col.: Coccinellidae),
Episyrphus balteatus (Dipt.:
Syrphidae)
Chinese cabbage / Arthropod predators Intercropping with green cabbage, Increased abundance and diversity Cai et al., 2007,
unspecified garlic, lettuce 2010
Citrus / various Predatory mites Intercropping with Ageratum Increased abundance Zhou et al., 1994;
conyzoides Olkowski &
Zhang, 1998
Pear / various Coccinella septempunctata, Aromatic plants as cover crops: Increased abundance, improved Song et al., 2010,
Phytoseiulus persimilis, Chrysoperla Satureja hortensis, Ageratum niche overlap with pests, pest 2011
sinica houstonianum, Tagetes patula, Mentha suppression
haplocalyx and basil
Sorghum, maize / Trichogramma ostriniae (Hym. : Intercropping with mungbean, kidney Higher parasitism Zhou et al.,
lepidopteran pests Trichogrammatidae) bean, spring peanut 1997a, b
Tea / green leafhopper Spiders Intercropping with citrus, waxberry and Increased abundance, reduction in Ye et al., 2010
snake gourd pest pressure
Cotton / cotton aphid Several natural enemies, including Intercropping with winter wheat Increased abundance by 70%, Zhao et al., 1987:
coccinellids and spiders earlier population build-up, lower Wang et al.,
pest pressure 1993; Xia, 1997 ;
Men et al., 2004 ;
Ma et al., 2006,
2007
Cotton / various Spiders, coccinellids, anthocorids and Intercropping with corn Increased abundance by 61-115%, Wu et al., 1991
lacewings superior pest control
Cotton / cotton aphid Spiders, ladybirds, lacewings Intercropping with alfalfa Increased species abundance, Chen et al., 2007
diversity, effective aphid control
Cotton / unspecified Various Intercropping with rape and sorghum Increased abundance by 25-50% Zhao et al., 1991
Cotton / cotton aphid Lacewings Intercropping with safflower Increased abundance Li, 1987
Wheat / cereal aphid Coccinellid beetles, aphid parasitoids Intercropping with peas Increased abundance, reduced pest Zhou et al., 2009
pressure
Wheat / cereal aphid Unspecified Intercropping with oilseed rape Increased diversity, possible Wang et al., 2010
superior pest control
Cuba Cabbage / mustard aphid Diaeretiella rapae (Hym.: Intercropping with maize Increased parasitism Mujica et al.,
Braconidae) 2009
 Cabbage / various Parasitoids Intercropping with maize Increased diversity and parasitism Vazquez, 1999;
Vazquez et al.,
2008
Coffee / coffee leafminer Predatory ants, parasitoids Commelina diffusa (evt. with Zebrina Increased abundance Vazquez, 2001;
pendula) as cover crops Simon, 1999
Sweetpotato / sweetpotato Predatory ants Intercropping with maize Increased abundance Pérez &
weevil Vazquez, 2001;
Vazquez et al.,
2005, 2008
Vegetables / various Various predators and parasitoids Polycultures Reservoirs and increase diversity Vazquez et al.,
and abundance 2010 ; Alfonso et
al., 2010
Potato and beans / thrips Predators Intercropping with maize Reservoir and increased abundance Vazquez, 1999
Tomato and beans / Various Intercropping with maize Reservoir and increased abundance Vazquez &
whitefly Lopez, 2000
Ethiopia Bean / corn earworm Tachinidae Intercropping with maize Increased abundance Abate, 1991
Georgia Citrus / citrus aphid Aphidophagous parasitoids Intercropping with tea Increased abundance, pest Hazarika et al.,
suppression 2001
Guadeloupe Citrus / various Phytoseiid mites Neonotonia wightii as a cover crop High abundance and sustained Mailloux et al.,
diversity year-round 2010
Guatemala Tomato / whitefly Parasitoids Intercropping with roselle and corn Higher diversity Smith et al., 2001
India Blackgram / unspecified Predatory spiders Intercropping with sorghum Increased abundance Rao, 2008
Chickpea / corn earworm Campoletis chlorideae (Hym.: Intercropping with coriander Increased abundance Pawar et al.,
Ichneumonidae) 1999
Cotton / unspecified Arthropod predators, spiders and Intercropping with clusterbean or Increased abundance Rao, 2008
coccinellids cowpea
Sorghum / unspecified Parasitoids Intercropping with cowpea Increased parasitism Rao, 2008
Tea / unspecified Carabid beetles, spiders Arachis pintoi as cover crop Increased abundance, reduced pest Hazarika et al.,
pressure 2009
Maize / unspecified Predatory spiders and spiderlings Intercropping with groundnut Increased abundance Rao, 2008
Pigeonpea / corn earworm Trichogramma spp. (Hym.: Intercropping with cereals such as Increased activity Pawar et al.,
Trichogrammatidae), coccinellids, (short cycle) sorghum, maize or pearl 1999; Rao, 2008
anthocorids, spiders millet
Pigeonpea / coreid pod bug Gryon sp. (Hym. : Scelionidae) Intercropping with cotton Lower parasitism Rao, 2008
Safflower / corn earworm, Chrysoperla carnea (Neuroptera: Intercropping with coriander Increased abundance Hanumantharaya
safflower aphid Chrysopidae), coccinellids et al., 2008
Indonesia Sweetpotato / sweetpotato Spiders Intercropping with corn, soybean, corn Increased abundance, reduced Yaku, 1992
weevil + soybean infestation levels
Kenya Cereals / stemborer Parasitoids Intercropping of maize, sorghum, millet Increased diversity Songa et al.,
and beans 2007
Maize / stemborer Spiders Intercropping with Desmodium plus Increased in-field abundance Midega et al.,
Napier grasses 2008
 Maize / stemborer Cotesia sesamiae (Hym.: Braconidae) Intercropping with Melinis minutiflora Increased parasitism Khan et al., 1997
Malaysia Cocoa / cocoa mirid Dolichoderus thoracicus, Oeophylla Intercropping with coconut palm Increased nesting Way & Khoo,
smaragdina (Hym. : Formicidae) 1991
Martinique Banana, plantain / banana Spiders, ants, centipedes, earwigs Bracchiaria decumbens as cover crop (Possible) increase in abundance, Duyck et al.,
weevil improved pest suppression 2011
Mexico Maize / various Various Intercropped trees Increased abundance Carmona, 2006
Maize / various Various Polyculture (corn, beans, squash) Increased abundance Altieri & Trujillo
1987
Maize / aphids Chrysopa sp. (Neur.: Chrysopidae) Polyculture (corn, beans, squash) Increased abundance Trujillo & Altieri
1990
Nicaragua Maize / Fall armyworm, Predatory ants Polyculture (corn-beans) Lower herbivory Perfecto &
corn leafhopper Sediles 1992
Nigeria Cowpea / various Orius sp. (Het.: Anthocoridae) Intercropping with maize Reduced abundance, depending Matteson, 1982
upon location
Peru Bean / various Paratriphleps sp. (Het.: Intercropping with maize (and weeds) No effect Gianoli et al.,
Anthocordiae), coccinellids, spiders 2006
Philippines Coconut / slug caterpillar Various Cover crops or leguminous intercrops Increased abundance PCA, 2003
Russia Vegetable crops / various Different natural enemies Intercropping with peppermint, sage, Increased abundance Nagirnyaka &
lemon balm, basil, anise, onion and leek Krasavina, 2005
Uganda, DRC Cowpea / various Orius sp. (Het.: Anthocoridae), Intercropping with greengram or Lowest abundance in cowpea - Nampala et al.,
spiders and earwigs sorghum sorghum 1999
Cowpea / various Arthropod predators Intercropping with greengram Higher predator abundance, higher Munyuli et al.,
yield, lower pest pressure 2007
Groundnut / various Arthropod predators Intercropping with maize Higher predator abundance, higher Munyuli et al.,
yields, lower pest pressure 2008
Maize / Termite Predatory ants (e.g., Myrmicaria, Intercropping with soybean, groundnut, Increased nesting Sekamatte et al.,
Lepisiota spp.) bean 2003
South Africa Sugarcane / African stalk Ants, predatory mites Intercropping with beans Stable population levels Beje, 2011
borer
Zanzibar Coconut / various Oecophylla longinoda (Hym.: Interplanting, increased plant diversity Increased abundance Way, 1954
Formicidae)
 

Appendix 5. Manipulation of disturbance regimes and their benefits for the resident natural enemy fauna
Country Target crop / pest
Argentina Wheat / unspecified
Various
Brazil Soybean-corn / unspecified
Common bean / various
Maize
Hog plum, guava /
Tephritid fruitfly
Benin Cassava / cassava mealybug
China Apple / herbivorous mites
Cotton / various
Cotton-Barley / unspecified
Chestnut / spruce spider
mite
Rice / whitebacked
planthopper, rice brown
planthopper
Rice / various
Natural enemy
Coccinellids
Carabidae
Carabidae, Staphylinidae, ants,
spiders
Cycloneda sanguinea (Col.:
Coccinellidae), Doru sp..(Derm.:
Forficulidae), Eulophid wasps,
carabids
Predatory ants
Solenopsis sp. (Hym.: Formicidae),
syrphids, Staphylinid beetles
Doru taeniatum (Derm.:
Forficulidae)
Parasitoids
Parasitoids
Orius sauteri (Het.: Anthocoridae)
Coccinellids, chrysopids, syrphids
Predators
Parasitoids and predators
Amblyseius castaneae (Acari:
Phytoseiidae)
Predators and parasitoids
Various
Predaceous spiders
Generalist predators
Disturbance scheme Effect
No till Unspecified
No till Unspecified
No till Increased diversity, increased
abundance of ants, carabids and
spiders
No till Increased abundance
No till and crop residue retention Increased abundance
Use of herbicides Sharp reduction abundance
Use of organic fertilizer instead of 2-5 times higher abundance
mineral
Routine removal of fallen fruits Reduced abundance
Frequent insecticide use Lower abundance, increased
pest pressure
Altered mowing regime of Medicago Increased abundance
sativa cover crop
Timely mowing of alfalfa intercrop or Increased abundance on target
cover crop crop
Use of Bt sprays vs chemically defined Up to 300% increased
insecticides abundance
Early removal of harvest residues Reduction in abundance
Altered mowing of Lolium perenne Increased abundance
cover crops
Addition of organic matter Increased abundance esp. of
spiders and parasitoids
Avoidance of calendar-based Increased abundance
insecticide sprays, use of ducks for pest
control
Appropriate timing of insecticide use Sustained presence in cropping
systems
Application nitrogen fertilizer Decreased abundance, lowered
(sentinel) prey removal
Reference
Lopez et al., 2008
Carmona et al., 2004
Martins et al., 2009;
Cividanes, 2002
Cividanes & Yamamoto,
2002; Cividanes &
Barbosa, 2001; Fernandes
et al., 2009
Pereira et al., 2010
Pereira et al., 2007
Galvao et al., 2001
Aguiar-Menezes &
Menezes, 2002
Neuenschwander et al.,
1987
Du & Yan, 1995
Zhang et al., 2000
Xing et al., 1991
Zhang et al., 1990
Lu et al., 2008
Jiang & Cheng, 2004;
Zhong et al., 2005b
Zhong et al., 2005a;
Zhang et al., 2009
Jiang et al., 2002
Ly et al., 2006a,b
 Tea / unspecified Carabid beetles Nitrogen application Increased abundance and Chen et al., 2009
richness of detritivores and
associated predators
Colombia Snap bean / whitefly Parasitoids Reduction insecticide use Increased abundance Manzano et al., 2003
Cuba Coffee / Unspecified Tetramorium bicarinatum, Addition of organic matter Sustained abundance Vazquez et al., 2009
Solenopsis geminata (Hym.:
Formicidae)
Sweet potato / sweet potato Pheidole megacephala (Hym.: Reduction insecticide use Conservation of ant colonies Perfecto & Castineiras,
weevil Formicidae) 1994
El Salvador Sugar cane / sugarcane Paratheresia claripalpis (Dipt. : Reduction insecticide use Increased abundance Argueta & Hernandez,
borer Tachinidae) 2009
Guatemala Corn / Various Various Organic fertilizer Increased abundance Morales et al., 2001
India Eggplant / fruit and shoot Trathala flavoorbitalis (Hym.: Reduction or suspension of insecticide Increased abundance Alam et al., 2003
borer Ichneumonidae) use
Groundnut / lepidopteran Spiders, carabids and coccinellids Reduction insecticide application Increased abundance, reduced Rao, 2005
pests frequency pest pressure
Wheat / unspecified Natural enemies Shaving or burning of rice stubble Lower abundance Jaipal et al., 2002
Sorghum / stemborer Natural enemies Short cropping cycle, frequent Reduced abundance and action Nwanze & Mueller, 1989
disturbance
Tea / various Egg parasitoids Reduction insecticide use Egg parasitism up to 97% Hazarika et al., 2001
Indonesia Cacao / various Spiders Reduction insecticide use Increased abundance Stenchly, 2010
Rice / Unspecified Generalist predators Addition of organic matter Increased abundance Settle et al., 1996
Ivory Coast Rice / Various Predatory ground spiders Hand weeding, compared to herbicide Increased spider abundance Afun et al., 1995
use
Kenya Coffee / coffee leafminer Hymenopteran parasitoids Refraining from pruning Higher levels of parasitism Wanjala, 1978
Predatory ants Addition of organic matter Lowered predation Crowe, 1964
Coffee / coffee berry borer Prorops nasuta (Hym.: Bethylidae) Removal of fallen berries Lowered abundance and Baker, 1999; Oduor &
performance Simons, 2003; Jaramillo
et al., 2009
Coffee / giant coffee looper Various Use of pyrethrum at a knock-down, and Increased efficacy Wheatley, 1963
DDT painted on tree trunk
Malawi, Coffee berry / Antestia bug, Parasitoids Pruning Increased efficacy Taylor, 1945; Oduor &
Kenya, white stem borer Simons, 2003; Jaramillo
Uganda et al., 2009
Mexico Cotton / tarnished plant bug Eulophidae, Braconidae, Elimination of pesticide that most Increased abundance Pulido-Herrera et al.,
Ichneumonidae, Chalcididae affect the parasitoids 1998
Coffee / various Spiders Organic management Higher diversity Pinkus-Rendón et al.,
2006
Predatory ants Pruning of coffee shade-trees Decreased diversity Philpott 2005
Fruits / Tephritid fruit fly Solenopsis geminata (Hym.: Overall disturbance Increased abundance Aluja et al., 2005
Formicidae)
Mango / Trips Various Reduced pesticide use Increased abundance Rocha & Infante 2009
 Oranges / citrus snow scale
Various / white grubs
Nicaragua Maize / various
Pakistan Wheat / unspecified
Cotton / various
Peru Cotton / various
Potato / Andean potato
weevil
Philippines Cotton / Asian corn borer
Rice / various
Rice / rice leaf folder
South Africa Potato / potato tuber moth
Cotton / cotton bollworm
Sri Lanka Cashew nut / various
Sudan, Israel, Cotton / cotton aphid,
Egypt whitefly
Thailand Rice / various
Uganda Common bean / black bean
aphid, legume pod borer
Maize / termites
Vietnam Sapodilla / various
Citrus / various
Arrhenophagus chionaspidis Appropriate timing of insecticide use Increased abundance Coronado-Blanco et al
(Hym.: Encyrtidae) 1998
Various predators Land preparation Exposure to natural enemies Gomez et al., 2000
Various Non till Increased abundance Amador et al., 1994
Spiders, staphylinid and coccinellid No till Increased abundance of hunting Butt & Sherawat, 2011
beetles spiders and predatory beetles,
high species richness
Arthropod predators Reduction insecticide spray frequency Increased abundance Hafeez et al., 2006;
Solangi et al., 2008
Various Careful selection of timing and type of Conserve natural enemies Beingolea, 1959
insecticide use
Carabid beetles Modification of tillage regimes (Possibly) increased abundance Yabar et al., 2006
Generalist predators and egg Modification of planting time No differences in abundance Litsinger et al., 2007
parasitoids
Various Adoption of short fallow period More stable populations Way & Heong, 1994
Various Refraining from early-season Increased abundance Kenmore et al., 1984;
insecticide use Way & Heong, 1994;
Settle et al., 1996
Various Nitrogen application Increased abundance De Kraker et al., 2000
Indigenous parasitoids Increasing use of insecticides Reduced efficacy Kfir, 2003
Mirids, anthocorids, spiders Minimization insecticide use Increased abundance, superior Van Hamburg & Guest,
pest attack 1997
Predatory ants Reduction of insecticide use Increased abundance Rickson & Rickson, 1998
Various Rational use of insecticides Increased abundance, superior Abou-Elhagag, 1998;
pest control Abdelrahman & Munir,
1989; Devine et al., 1998
Various Calendar-based insecticide sprays Reduced abundance Jahn, 1992
Various parasitoids and predators Use of organic fertilizer Increased abundance Karungi et al., 2005
Predatory ants, Lepisiota sp., Use of maize stover as organic Increased abundance, lower pest Sekamatte et al, 2001a
Myrmicaria sp. fertilizer pressure
Dolichoderus thoracicus (Hym.: Use of water bailing instead of Stable population levels Van Mele & Cuc, 2001
Formicidae) overhead irrigation
Oecophylla smaragdina (Hym.: Reduction insecticide use Conservation ant populations Van Mele & Cuc, 2000
Formicidae)

Appendix 6. Influence of crop germplasm on mobility, host searching behavior and fitness of natural enemies
Country Target crop
Benin Cassava / cassava green mite
Cassava / various
Pigeonpea / coreid pod bug
Brazil Cotton / cotton aphid
China Wheat / Grain aphid
Rice / brown planthopper
Rice / brown planthopper
Congo Cassava / cassava mealybug
India Cotton / cotton aphid
Cotton / cotton bollworm
Lentil / bean aphid
Chickpea / corn earworm
Pigeonpea / corn earworm
Pigeonpea / bean aphid
Jordan Faba bean / bean aphid
Natural enemy Germplasm attribute Effect Reference
Typhlodromalus aripo (Acari: Pubescence Facilitate establishment, increase Zundel et al., 2009
Phytoseiidae) abundance
Predatory mites Extra-floral nectar Sustains populations and increases Bakker & Klein,
reproduction 1992; Bruce-Oliver et
al., 1996
Ooencyrtus utetheisae (Hym.: Unspecified No parasitism on pigeonpea plants Dryer, 1994
Encyrtidae) within cowpea field
Chrysoperla externa (Neur.: Pubescence Reduced longevity Santos et al., 2003
Chrysopidae)
Aphidius sp. (Hym.: Braconidae) Unspecified Parasitism higher on resistant Cai et al., 2009
varieties
Anagrus nilaparvatae (Hym.: Texture and siliceous cell density of Host size and parasitoid fitness Lou & Cheng, 1996
Mymaridae) leaves
Cytorhinus lividipennis (Het. : Plant volatiles, other attributes Differing attraction, variable egg Lou & Cheng, 2003
Miridae) predation
Exochomus flaviventris (Col.: Mealybug resistance Reduced pre-imaginal mortality, Le Ru & Mitsipa,
Coccinellidae) oviposition time and fecundity 2002
Various Phloem nutritional composition Sugar composition of honeydew Lawo et al., 2009
Chrysopa scelestes (Neur.: Trichome density Lower egg and larval predation rate Ramnath &
Chrysopidae) Uthamasamy, 1992
Trichogramma chilonis (Hym.: H. armigera resistance Increased efficacy Annadurai et al., 1992
Trichogrammatidae)
Coccinellids Unspecified Great variability in suitability for Sharma & Yadav,
natural enemies 1993
Trichogramma spp. (Hym.: Glandular hairs, exudates Interference with parasitoid activity Bhatnagar, 1981
Trichogrammatidae)
Trichogramma chilonis (Hym.: Glandular trichomes, exudates Reduce searching efficacy Sithanantham et al.,
Trichogrammatidae) 1982; Pawar et al.,
1986
Orius tantillus (Het.: Anthocoridae) Plant chemistry and architecture Lower efficacy on pigeonpea vs Sigsgaard & Esbjerg,
sorghum 1997
Paratrechina longicornis (Hym.: Long trichomes, sticky exudates Lower egg predation on buds and Romeis et al., 1995
Formicidae) pods vs leaves
Trioxys indicus (Hym.: Aphidiidae) Foliar pubescence Lower searching efficacy Kumar et al., 1983
Coccinella septempunctata (Col.: Aphid resistance Partial resistance benefited Shannag & Obeidat,
Coccinellidae) biological control, greater fertility 2006, 2008
on susceptible lines
Mexico Maize / fall armyworm Cotesia marginiventris (Hym.: Herbivore-induced volatiles Differing degree of attraction Hoballah et al., 2002
Braconidae)
Nigeria Cowpea / cowpea aphid Coccinellids Unspecified Higher abundance on aphid Ofuya, 1995
susceptible varieties
Pakistan Pumpkin, cucumber / Predatory mites Hairiness, hair length, leaf area, Abundance Afzal & Hamid, 2007
unspecified surface waxes
Philippines Rice / brown planthopper Cyrtorhinus lividipennis (Het.: Volatiles from different rice Differential attraction Rapusas et al., 1995
Miridae) genotypes
Rice / brown planthopper Various predators, Lycosid spiders Planthopper resistance Moderate resistance benefited Karothardjono &
biological control Heinrichs, 1984
Tanzania Pigeonpea / coreid pod bug Gryon fulviventre (Hym.: Unspecified Lower egg parasitism on Phaseolus Materu, 1971
Scelionidae) beans vs pingeonpea or lablab bean
 
 
 
 

Appendix 7. Landscape level effects on natural enemy abundance and efficacy

Country Target crop / pest Natural enemy Landscape level component Effect Reference
Argentina Wheat Spiders Herbaceous field margins Increased colonization Armendano &
Gonzalez, 2011
Brazil Eucalyptus / Microhymenopterans Native cerrado vegetation Higher wasp abundance near native Silva et al., 2010
unspecified vegetation
Trichogramma maxacalii (Hym.: Forest fragments Increased abundance, superior Murta et al., 2008
Trichogrammatidae) parasitism
Maize / fall Social wasps Forest fragments Higher abundance Sousa et al., 2011
armyworm
Chile Alfalfa / unspecified Coccinellid beetles Surrounding habitats, tall vegetation near Earlier arrival in crop Grez et al., 2010
crop field
China Various Trichogramma spp. (Hym.: Landscape diversity, natural forest edges, Conservation of parasitic wasps Olkowski & Zhang,
Trichogrammatidae) tree and shelterbelt systems 1998
Rice / planthopper, Anagyrus nilaparvatae (Hym.: Non-crop habitats Provide alternative hosts, boost Zheng et al., 2003
rice leafroller Encyrtidae) abundance
Arthropod predators Diverse landscapes, with white melon, Increased abundance Liu et al., 1999;
cabbage, tea orchards, soybean and grassy Zhang et al., 2011a
borders
Apple / various Orius sauteri (Het.: Anthocoridae) Presence of alfalfa fields, corn and Increased abundance Yu et al., 1998
vegetable plots
Cotton / Trichogramma spp. (Hym.: Presence of vegetable gardens (with Stable populations Wang et al., 1988
lepidopteran pests Trichogrammatidae) inoculative releases)
Cotton / cotton Coccinellid beetles, lacewings, Regional adoption of Bt cotton Increased abundance, improved pest Lu et al.,
aphid spiders suppression unpublished
Cuba Various Various Reduction share monocultures, increased Increased abundance and diversity Vazquez, 2007
number of farms
Ecuador Unspecified Arthropod predators Increasing agricultural transformation and Lower diversity Tscharntke et al.,
intensification 2008
Honduras Maize / fall Doru taeniatum (Dermaptera: Grassy habitats (earwigs), late successional Increased abundance Wyckhuys &
armyworm Forficulidae), spiders, carabid beetles habitats O’Neil, 2007b
Indonesia Unspecified Telenomus remus (Hym.: Landscape complexity Increased parasitism Buchori et al., 2008
Scelionidae)
Rice / various Arthropod predators Larger fields, landscape simplification Delayed colonization Settle et al., 1996
Iran Cotton / various Dipteran parasitoids, predators Surrounding grasslands Differing abundance and activity Ghahari et al., 2008
Israel Wheat / unspecified Several spider families Semi-desert habitats in the agro-landscape Increased activity-density Pluess et al., 2008
Kenya Crucifers / Plutella Diadegma spp., Apanteles spp. Presence of wild crucifers Increased parasitoid abundance and Kahuthia-Gatu et
xylostella (Hym.: Braconidae),, Brachymeria diversity, recolonization sites for al., 2009
 sp. (Hym.: Chalcidae), Oomyzus parasitoids
sokolowski (Hym. Eulophidae)
Mexico Various Ichneumonoidea Higher diversity in surrounding vegetation Increased abundance Chay-Hernandez et
al., 2006
Philippines Rice / various Various Asynchronous rice planting over large areas Conservation of natural enemies Way & Heong,
1994; Ives & Settle,
1997; Matteson,
2000
Rice / various Wolf- and web-building spiders Establishment of vegetable fields, fruit or Increased abundance Barrion, 1999;
timber trees Drechsler & Settele,
2001; Gurr, 2009
Ukraine Grape / grape leaf Unspecified Proximity to forest plantations Increased parasitism levels Saidov et al., 2007
roller
Various Olive / olive fly Pnigalio agraules (Hym. : Habitat connectivity Increased abundance Boccaccio &
Eulophidae), Eupelmus urozonus Petacchi, 2009
(Hym.: Chalcidae), Eurytoma martelli
(Hym.: Eurytomidae)
 
 
References for Appendices

Abate, T., 1991. Intercropping and weeding: effects on some natural enemies of African
bollworm, Heliothis armigera (Hbn.) (Lep., Noctuidae), in bean fields. Journal of Applied
Entomology 112, 38-42.
Abdelrahman, A.A., Munir, B., 1989. Sudanese experience in integrated pest management of
cotton. Insect Science and Application 10, 787–794.
Abou-Awad, B.A., Afia, S.I., Al-Azzazy, M.M., 2011. Mango powdery mildew Oidium
mangiferae an alternative food for the predatory mites Typhlodromus mangiferus and
Typhlodromips swirskii (Phytoseiidae) in absence or presence increasing prey density of
Oligonichus mangiferus (Tetranychidae) in Egypt. Archives of Phytopathology and Plant
Protection 44, 1703-1710.
Abou-Elhagag, G.H., 1998.Effect of spraying cotton plants during the early-season against
cotton aphid on cotton pests, natural enemies and some crop characters in southern Egypt.
Assiut Journal of Agricultural Science 29, 91–100.
Afun, J.K.V., Heinrichs, E.A., Johnson, D.E., Russell-Smith, A., 1995.The influence of weeds in
the natural control of upland rice insect pests in Cote d'Ivoire. Proceedings (Vol 1) of the
Brighton Crop Protection Conference, 20-23 Nov 1995. British Crop Protection Council,
Farnham, UK. pp 181-186.
Afun, J.K.V., Johnson, D.E., Russell-Smith, A., 2000.Weeds and natural enemy regulation of
insect pests in upland rice; a case study from West Africa. Bulletin of Entomological Research
89, 391-402.
Afzal, M., Hamid, M., 2007. Influence of certain leaf characters of some summer vegetables
with incidence of predatory mites of the family Cunaxidae. Pakistan Journal of Botany 39, 205-
209.
Aguiar-Menezes, E.L., Menezes, E.B. 2002. Effect of time of permanence of host fruits in the
field on natural parasitism of Anastrepha spp. (Diptera: Tephritidae). Neotropical Entomology
31, 589-595.
Ahmad, N., Sarwar, M., Wagan, M.S., Muhammad, R., Tofique, M.T., 2011. Conservation of
biocontrol agents in cotton, Gossypium hirsutum L. field by food supplements for insect pest
management. The Nucleus 48, 255-260.
Alam, S.N., Rashid, M.A., Rouf, F.M.A., Jhala, R.C., Patel, J.R., Satpathy, S., Shivalingaswamy,
T.M., Rai, S., Wahundeniya, I., Cork, A., Ammaranan, C., Talekar, N.S., 2003. Development
of an integrated pest management strategy for eggplant fruit and shoot borer in South Asia,
Technical Bulletin TB28, AVRDC – The World Vegetable Center, Shanhua, Taiwan, 66 pp .
Alfonso, J., Matienzo, Y., Vazquez, L., 2010. Fauna de hormigas (Hymenoptera: Formicidae)
asociadas a un sistema de producción agrícola urbano. Fitosanidad 14, 153-158.
Altieri, M., Trujillo, J., 1987.The agroecology of maize production in Tlaxcala, Mexico. Human
Ecology 15, 189-220.
Aluja, M., Rull, J., Sivinski, J., Norrbom, A.L., Wharton, RA., Macias-Ordonez, R., Diaz-
Fleischer, F., Lopez, M., 2003. Fruit flies of the genus Anastrepha (Diptera : Tephritidae) and
associated native parasitoids (Hymenoptera) in the tropical rainforest biosphere reserve of
Montes Azules, Chiapas, Mexico. Environmental Entomology 32, 1377-1385.
Aluja, M., Sivinski, J., Rull, J., Hodgson, P.J., 2005. Behavior and predation of fruit fly larvae
(Anastrepha spp.) (Diptera: Tephritidae) after exiting fruit in four types of habitats in tropical
Veracruz, Mexico.Environmental Entomology 34, 1507-1516.
Amador, T.A., Escobar, J.E., Gladstone, S., 1994. Efecto de labores culturales en la conservación
de enemigos naturales en cultivo de maíz. Anales del Curso y Foro Sub Regional
Centroamericano y del Caribe de Control Biológico de Plagas.León (Nicaragua). Universidad
Nacional Autónoma de Nicaragua, León (Nicaragua).
Amaral, D.S., Venzon, M., Pallini, A., Lima, P.C., DeSouza, O.G. 2010. Does vegetational
diversification reduce coffee leaf miner Leucoptera coffeella (Guérin-Mèneville) (Lepidoptera:
Lyonetiidae) attack? Neotropical Entomology 39, 543-548.
Amend, J., Basedow, T., 1997. Combining release/establishment of Diadegma semiclausum
(Hellen) (Hym., Ichneumonidae) and Bacillus thuringiensis Berl. for control of Plutella
xylostella (L.) (Lep., Yponomeutidae ) and other lepidopteran pests in the Cordillera Region of
Luzon (Philippines). Journal of Applied Entomology 121, 337–342.
Annadurai, R.S., Murugesan, S., Senrayan, R., Gurusubramanian, G., Ananthakrishnan, T. 1992.,
Tritrophic interactions in Helitohis armigera Hubner and its natural enemy systems : a
chemical ecology approach (Noctuidae:Lepidoptera), in: Ananthakrishnan, T.N. (Ed.),
Emerging trends in biological control of phytophagous insects. Oxofrd & IBH Publishing Co.
Ltd., New Delhi, pp. 83-101.
Argueta, A., Hernandez, W., 2009. Parasitoidismo y control microbiano del barrenador
(Diatraea saccharalis F.) de la caña de azúcar (Saccharum officinarum L.), en el departamento
de Sonsonate, El Salvador. Tesis Ingeniero Agronomía. Facultad de Ciencias Agronómicas,
Universidad de El Salvador, San Salvador.
Armbrecht, I., Perfecto, I., Silverman, E., 2006. Limitation of nesting resources for ants in
Colombian forests and coffee plantations. Ecological Entomology 31, 403-410.
Armendano, A., Gonzalez, A., 2011. Spider fauna associated with wheat crops and adjacent
habitats in Buenos Aires, Argentina. Revista Mexicana de Biodiversidad 82, 1176-1182.
Aroga, R., 2003. Le pollen de mais (Zea mays L.), un aliment adéquat pour un élevage massif de
Diaperasticus erythrocephala Olivier (Dermaptera:Forficulidae), in: Badu-Apraku, B. (Ed.),
Maize revolution in West and Central Africa. IITA, Ibadan, Nigeria.
Baker, P.S., 1999. The coffee berry borer in Colombia.Final report of the DFID-Cenicafe-CABI
Bioscience IPM of Coffee project, Chinchina, Colombia.
Bakker, F.M., Klein, M.E., 1992. Transtrophic interactions in cassava. Experimental and Applied
Acarology 14, 293-311.
Barrion, A., 1999. Ecology of spiders in selected non-rice habitats and irrigated rice fields in two
southern Tagalog provinces in the Philippines. PhD Thesis, University of the Philippines at Los
Banos.
Beingolea, O.D., 1959. Notas sobre la bionómica de arañas e insectos benéficos que ocurren en
el cultivo de algodón. Revista Peruana de Entomologia Agricola 2, 36-44.
Beje, S., 2011. The effect of intercropping beans on Eldana saccharina Walker (Lepidoptera:
Pyralidae) arthropod predator populations in sugarcane. MSc Thesis. University of Durban,
Department of Zoology.
Bentley, J.W., 2006. Folk experiments. Agriculture and Human Values 23, 421-462.
Bertolaccini, I., Andrada, P., Quaino, O., 2008. Efecto de franjas marginales en la atracción de
coccinellidae y syrphidae, depredadores de áfidos en trigo, en la zona central de la provincia de
Santa fe, Argentina. Agronomía Tropical 58, 267-276.
Bhatnagar, V.S., 1981. Are effective parasites of Heliothis eggs found on pigeonpea and
chickpea? International Pigeonpea Newsletter 1, 32.
Bigger, M., 1981.Observations on the insect fauna of shaded and unshaded Amelonado cocoa.
Bulletin of Entomological Research 71, 107–119.
Boccaccio, L., Petacchi, R., 2009. Landscape effects on the complex of Bactrocera oleae
parasitoids and implications for conservation biological control. BioControl 54, 607-616.
Bottenberg, H., Litsinger, J.A., Barion, A.T., Kenmore, P.E., 1990. Presence of Tungro vectors
and their natural enemies in different rice habitats in Malaysia. Agriculture, Ecosystems and
Environment 31, 1–15.
Boukary, I.B., Gingras, J., Tourneur, J.C., 1997. Feeding habits of Forficula senegalensis
(Dermaptera: Forficulidae) on millet in the Sudanese-Sahelian zone of Niger: crop content
analysis. Entomophaga 42, 537-542.
Bravo Luna, L., Flores Moctezuma, E., Villa Ayala, P. Corona Rangel, M.L., Bermúdez Torres,
K., 2003. Parasitismo natural de mosquita blanca en Morelos, México. XXVI Congreso
Nacional de Control Biológico, México.
Bruce-Oliver, S.J., Hoy, M.A., Yaninek, J.S., 1996. Effect of some food sources associated with
cassava in Africa on the development, fecundity and longevity of Euseius fustis (Acari:
Phytoseiidae). Experimental and Applied Acarology 20, 73-85.
Buchori, D., Sahari, B., Nurindah, 2008. Conservation of agro-ecosystem through utilization of
parasitoid diversity: lesson for promoting sustainable agriculture and ecosystem health. Hayati
Journal of Biosciences 15, 165-172.
Butt, A., Sherawat, S.M., 2011.Effect of different agricultural practices on spiders and their prey
populations in small wheat fields. Acta Agriculturae Scandinavica, Section B.
Cai, H.J., Li, Z.S., You, M.S., 2007. Impact of habitat diversification on arthropod communities:
a study in the fields of Chinese cabbage, Brassica chinensis. Insect Science 14, 241-249.
Cai, H.J., You, M.S., Lin, C., 2010. Effects of intercropping systems on community composition
and diversity of predatory arthropods in vegetable fields. Acta Ecologica Sinica 30, 190-195.
Cai, Q.N., Ma, X.M., Zhao, X., Cao, Y.Z., Yang, X.Q., 2009. Effects of host plant resistance on
insect pests and its parasitoid: a case study of wheat-aphid-parasitoid system. Biological
Control 49, 134-138.
Canas, L.A., O’Neil, R.J., 1998. Applications of sugar solutions to maize, and the impact of
natural enemies on fall armyworm.International Journal of Pest Management 44, 59-64.
Carmona, M., 2006. Impacto de arboles sobre abundancia y diversidad de artrópodos en maíz, en
huertos familiares de los Altos de Chiapas. MSc thesis, El Colegio de la Frontera Sur.
Carvalho, R., Soares, W., Ritzinger, R., 2010.Umbu-cajá como repositório natural de parasitoide
nativo de moscas-das-frutas. Pesquisa Agropecuaria Brasileira 45, 1222-1225.
Chari, M.S., Bharpoda, T.M., Patel, N.M., 1985.Studies on integrated management of
Spodoptera litura F. in tobacco nursery. Tobacco Research 11, 93-97.
Chabi-Olaye, A., Nolte, C., Schulthess, F., Borgemeister, C., 2005. Abundance, dispersion and
parasitism of the stem borer Busseola fusca (Lepidoptera: Noctuidae) in maize in the humid
forest zone of southern Cameroon. Bulletin of Entomological Research 95, 169-177.
Chay-Hernandez, D.A., Delfin-Gonzalez, H., Parra-Tabla, V., 2006. Ichneumonoidea
(Hymenoptera) community diversity in an agricultural environment in the state of Yucatan,
Mexico. Environmental Entomology 35, 1286-1297.
Chen, M., Li, G.Q., Luo, J.C., 2007. The structure and dynamics of natural enemy community
and controlling effect against Aphis gossypii in cotton alfalfa intercrop fields. Acta
Phytophylacica Sinica 34, 631-636.
Chen, Q., Peng, Z.Q., Xu, C., Tang, C., Lu, B.Q., Wen, H., Wan, F.H., 2010. Biological
assessment of Tetrastichus brontispae, a pupal parasitoid of coconut leaf beetle Brontispa
longissima. Biocontrol Science and Technology 3, 283-295.
Chen, S.B., Wei, Z.J., Zeng, Z.H., Chen, L.L., Chen, H.T., You, M.S., 2009. Effects of nitrogen
application on beetle communities in tea plantations. Insect Science 16, 185-192
Cheng, Z.F., 1984. Biology study on Apanteles cypris Nixon. Natural Enemies of Insects 6, 71-
80.
Cherry, A., Cock, M., van den Berg, H., Kfir, R., 2003. Biological control of Helicoverpa
armigera in Africa, in: Neuenschwander, P., Borgemeister, C., Langewald, J. (Eds.), Biological
control in IPM systems in Africa. CABI Publishing, Wallingford, pp. 329-346.
Chong, P.P., Fee, C.G., 1989. Use of black cocoa ant to control mirid damage in cocoa.Planter,
Kuala Lumpur 65, 370-383.
Cividanes, F.J., 2002. Effects of tillage systems and intercropping of soybean and corn on soil-
inhabiting arthropods.Pesquisa Agropecuaria Brasileira 37, 15-23.
Cividanes, F.J., Barbosa, J.C., 2001. Efeitos do plantio direto e da consorciação soja-milho sobre
inimigos naturais e pragas. Pesquisa Agropecuaria Brasileira 36, 235-241.
Cividanes, F.J., Yamamoto, F.T., 2002. Pragas e inimigos naturais na soja e no milho cultivados
em sistemas diversificados. Scientia Agricola (Piracicaba, Braz.) 59, 683-687.
Coronado Blanco, J.M., Ruiz Cancino, E., Evans, G., Myortseva, S., Trjapitzin, V., 1998. Fauna
parasítica de Unaspis citri (Comstock) (Homoptera: Diaspididae) en Llera, Tamaulipas. XXI
Congreso Nacional de Control Biológico. México.
Crowe, T.J., 1964. Coffee leafminer in Kenya. II. Causes of outbreaks. Kenya Coffee 29, 223-
231.
Cruz-Angon, A., Greenberg, R., 2005 Are epiphytes important for birds in coffee plantations?
An experimental assessment. Journal of Applied Ecology 42, 150-159.
De Kraker, J., Rabbinge, R., Van Huis, A., Van Lenteren, J.C., Heong, K.L., 2000. Impact of
nitrogenous-fertilization on the population dynamics and natural control of rice leaffolders
(Lep.: Pyralidae). International Journal of Pest Management 46, 225-235.
Deng, X., Zheng, Z.Q., Zhang, N.X., Jia, X.F., 1988. Methods of increasing the winter-survival
of Metaseiulus occidentalis (Acari: Phytoseiidae) in northwestern China. Chinese Journal of
Biological Control 4, 97-101.
De Villiers, M., Pringle, K.L., 2010. The presence of Tetranychus urticae (Acari: Tetranychidae)
and its predators on plants in the ground cover in commercially treated vineyards. Experimental
and Applied Acarology 53, 121-137.
Devine, G.J., Ishaaya, I., Horowitz, A.R., Denholm, I., 1998. Effects of piperonyl butoxide on
Bemisia tabaci Genn (Homoptera: Aleyrodidae): mortality, development, parasitism and
predation in Israeli cotton fields. Crop Protection 17, 717–726.
Drechsler, M., Settele, J., 2001. Predator-prey interactions in rice ecosystems: effects of guild
composition, trophic relationships and land use changes: a model study exemplified for
Philippine rice terraces. Ecological Modeling 137, 135-159.
Dryer, H., 1994. Seed damaging field pests of cowpea (Vigna unguiculata) in southern Benin,
with special reference to Clavigralla tomentosicollis Stal (Het., Coreidae). PhD thesis. Swiss
Fed. Inst.Tech., Zurich.
Du, X.G., Yan, Y.H., 1994.Effect of the cover crops on the pest mite Panonychus ulmi and its
predator Orius sauteri. Chinese Journal of Biological Control 10, 114-117
Du, X.G., Yan, Y.H., 1995.Manipulating the ground cover of apple orchard to increase the
number of Orius sauteri in apple trees. Chinese Journal of Biological Control 11, 1-4.
Duyck, P.F., Lavigne, A., Vinatier, F., Achard, R., Okolle, J.N., Tixier, P., 2011. Addition of a
new resource in agroecosystems: do cover crops alter the trophic positions of generalist
predators? Basic and Applied Ecology 12, 47-55.
Eremenko, T.S., 1971. Importance of the adult parasite feeding of winter moth in increasing their
efficiency.Biological methods of protection fruit and vegetable crops from insect pest, diseases
and weed as bases of the integrated systems.Thesis’ report, October 1971, Kishinev, pp. 30-31.
Fadini, M.A.M., Regina, M.A., Fraguas, J.C., Louzada, J.N.C., 2001. Efeito da cobertura vegetal
do solo sobre a abundância e diversidade de inimigos naturais de pragas em vinhedos. Revista
Brasileira de Fruticultura 23, 573-576.
Fernandes, I.C., Cividanes, F.J., Barbosa, J.C., de Souza, E., Queiroz, G., 2009. Faunal analysis
and population fluctuation of Carabidae and Staphylinidae (Coleoptera) in no-tillage and
conventional tillage systems.Revista Brasileira de Entomologia 53, 432-443.
Fernandez, D.E., Cichon, L.I., Sanchez, E.E., Garrido, S.A., Gittins, C., 2008.Effect of different
cover crops on the presence of arthropods in an organic apple (Malus domestica Borkh)
orchard.Journal of Sustainable Agriculture 2, 197-211.
Fernández, J. L, Garcés, G., Portuondo, E., Valdéz, P., Expósito, I., 2001. Insectos asociados con
flores de malezas del Jardín Botánico de Santiago de Cuba, con énfasis en Hymenoptera.
Revista de Biologia Tropical 49, 3-4.
Figueroa de la Rosa, J.I., Ovruski Alderete, S., Liedo Fernandez, J.P., Cancino Díaz, J.L., 1998.
Parasitoides nativos y exóticos asociados a Anastrepha spp. en frutales silvestres y cultivados
de la región del Soconusco, Chiapas, México. XXI Congreso Nacional de Control Biológico.
México.
Gallego-Ropero, M.C., Montoya-Lerma, J., Armbrecht, I., 2009. Es la sombra benéfica para la
diversidad de hormigas y peso del café: una experiencia en Pescador, Cauca, Colombia. Boletin
Cientifico del Museo de Historia Natural Universidad de Caldas 13, 106-116.
Galunko, V.I., Dyadechko, N.P., 1971. About attraction entomophages to pea fields.Biological
methods of protection fruit and vegetable crops from insect pest, diseases and weed as bases of
the integrated systems.Thesis’s report, October 1971, Kishinev, pp. 24-25.
Galvao, J.C., Silva, E.C., Miranda, G.V., Bastos, C.S., Picanco, M.C., Silva, R.G., 2001.
Densidade populacional de alguns insetos em milho exclusivo e consorciado com feijao, em
dois sistemas de adubacao. Revista Ceres 48, 25-35.
Ghahari, H., Hayat, R., Chao, C., Ostovan, H., 2008. A contribution to the dipteran parasitoids
and predators in Iranian cotton fields and surrounding grasslands. Munis Entomology &
Zoology 3, 699-706.
Gianoli, E., Ramos, I., Alfaro-Tapia, A., Valdez, Y., Echegaray, E.R., Yabar, E., 2006. Benefits
of a maize–bean–weeds mixed cropping system in Urubamba Valley, Peruvian Andes.
International Journal of Pest Management 52, 283-289.
Giraldo-Vanegas, H., Garcia, J.L., 1995. Influencia de la alimentacion sobre la capacidad
reproductiva de Eiphosoma vitticolle Cresson (Hymenoptera: Ichneumonidae), parasito de
Spodoptera frugiperda (J.E. Smith). Agronomía Tropical 45, 159-170.
Girma, H., Rao, M.R., Sithanantham, S., 2000. Insect pests and beneficial arthropods population
under different hedgerow intercropping systems in semiarid Kenya. Agroforestry Systems 50,
279-292.
Gnanvossou, D., Hanna, R., Yaninek, S.J., Toko, M., 2005. Comparative life history traits of
three neotropical phytoseiid mites maintained on plant-based diets. Biological Control 35, 32-
39.
Gomez, B., Castro, A., Junghans, C., Montoya, L., Villalobos, F.J., 2000. Ethnoecology of white
grubs (Coleoptera: Melolonthidae) among the Tzeltal Maya of Chiapas. Journal of
Ethnobiology 20, 43-59.
Goncalves, P.A., Sousa, C.R., 2003. Efeito de espécies vegetais em bordadura em cebola sobre a
densidade populacional de tripes e sirfídeos predadores. Horticultura Brasileira 21, 731-733.
Graham, M.K., 1991. Biological control of Helopeltis spp. in mature cocoa by the black ant
(Dolichoderus bituberculatus) and the cocoa mealybug (Planococcus lilacinus).Planter 67,
516-520.
Grez, A.A., Torres, C., Zaviezo, T., Lavandero, B., Ramirez, M., 2010. Migration of coccinellids
to alfalfa fields with varying adjacent vegetation in Central Chile. Ciencia e Investigacion
Agraria 37(2).
Grout, T.G., Stephen, P.R., 1995. New windbreak tree contributes to integrated pest management
of citrus. Citrus Journal 5, 26–27.
Gunduz, E.M., Gulel, A., Isitan, O., Boz, A., Cesur, O., 2010. Effects of sugar feeding on lipid,
glycogen, and total sugar levels of a female parasitoid, Bracon hebetor (Say) (Hymenoptera:
Braconidae). Turkish Journal of Agriculture and Forestry 34, 343-347.
Guo X.L., Wang, D.S., He, Y.R., Luo, Y.L., 2011.Effects of different nutritional resources on
longevity and fecundity of Trichogrammatoidea bactrae Nagaraja. Chinese Journal of
Biological Control 27, 448-452.
Gurr, G.M. 2009. Prospects for ecological engineering for planthoppers and other arthropod
pests in rice, in: Heong, K.L., Hardy, B. (Eds.) Planthoppers: New Threats to the Sustainability
of Intensive Rice Production Systems in Asia. Los Banos, Philippines: International Rice
Research Institute, pp. 371–388.
Hafeez, F., Arif, M.J., Gogi, M.D., Zia, K., Karar, H., Arshad, M., 2006.Survey of
entomophagous insects in cotton belt of Punjab, Pakistan. Pakistan Journal of Biological
Sciences 9, 1357-1380.
Hanumantharaya, L., Venkateshalu, K.V.S., Raju, S.G., 2008. Role of cropping pattern for the
management of insect pests of Safflower, Carthamus tinctorius L. Proceedings of the 7th
International Safflower Conference, Waga Waga, Australia.
Hazarika, L.K., Bhuyan, M., Hazarika, B.N., 2009.Insect pests of tea and their management.
Annual Review of Entomology 54, 267-284.
Hazarika, L.K., Puzari, K.C., Wahab, S., 2001. Biological control of tea pests, in: Upadhyay,
R.K., Mukerji, K.G., Chamola, B.P. (Eds.), Biocontrol Potential and Its Exploitation in
Sustainable Agriculture. Vol. 2: Insect Pests. New York: Kluwer Academic, pp. 159–80.
Hernandez, L.M., Manzano, M., Otero, T., 2009.Control biológio de Trialeurodes vaporariorum
Westwood (hemiptera; Aleyrodidae) por Amitus fuscipennis MacGown (Hymenoptera:
Platygasteridae) en frijol y habichuela: bases para planear la diversidad de plantas
acompañantes. Resúmenes. XXXVI Congreso de la Sociedad Colombiana de Entomologia.
Medellin. Colombia.
Ho, T.T.G., Ueno, T., 2007. Improving parasitoid performance by improving adult food quality:
a case study for the leafminer parasitoid Hemiptarsenus varicornis (Hymenoptera : Eulophidae).
Journal of the Faculty of Agriculture, Kyushu University 52, 57-61.
Hoballah, M.E.F., Tamo, C., Turling, T.C.J., 2002. Differential attractiveness of induced odors
emitted by eight maize varieties for the parasitoid Cotesia marginiventris: is quality or quantity
important? Journal of Chemical Ecology 28, 951-968.
Hosang, M.L.A., Schulze, C.H., Tscharntke, T., Buchori, D., 2010. The potential of artificial
nesting sites for increasing the population density of the black cacao ants. Indonesian Journal of
Agriculture 3, 45-50.
Hrncir, M., Mateus, S., Nascimento, F.S., 2007. Exploitation of carbohydrate food sources in
Polybia occidentalis: social cues influence foraging decisions in swarm-founding wasps.
Behavioral Ecology and Sociobiology 61, 975-983.
Huang , H.T., Yang, P., 1987. The ancient cultured citrus ant.BioScience 37, 665 -671.
Ide, S., Lanfranco, D., 2001. Longevidad de Orgilus obscurator Ness. (Hymenoptera:
Braconidae) en presencia de diferentes fuentes de alimento. Revista Chilena de Historia Natural
74, 469-475.
Iswaran, V., Sen, A., 1972. Sugar spray boosts up pulse yields. AGRI Digest 24, 40–41.
Ives, A.R., Settle, W.H., 1997. Metapopulation dynamics and pest control in agricultural systems.
American Naturalist 149, 220-246.
Jahn, G.C., 1992. Rice pest control and effects on predators in Thailand. Insecticide & Acaricide
Tests 17, 252-253.
Jaipal, S., Singh, S., Yadav, A., Malik, R.K., Hobbs, P., 2002. Species diversity and population
density of macrofauna of rice-wheat cropping habitat in semi-arid subtropical north-west India
in relation to modified tillage practices of wheat sowing. International workshop on herbicide
resistance management & zero tillage in rice-wheat cropping system, March 4-6, 2002,
Department of Agronomy, CCS Haryana Agricultural University, Hisar-125 004, India.
Jaramillo, J., Chabi-Olaye, A., Borgemeister, C., Kamonjo, C., Poehling, H.m., Vega, F.E., 2009.
Where to sample? Ecological implications of sampling strata in determining abundance and
impact of natural enemies of the coffee berry borer, Hypothenemus hampei. Biological Control
49, 245-253.
Jiang, Y.H., Wu, J.C., Xu, J.X., Liu, J.L., Qiu, H.M., Yang, G.Q., Wang, H.Q., 2002. Influence
of seasonal and daily changes of spatial niche of spiders in paddy field and two insecticides to
spatial niche and predatory function. Acta Ecologica Sinica 22, 1286-1292.
Jiang, M.X., Cheng, J.A., 2004. Effects of manure use on seasonal patterns of arthropods in rice
with special reference to modified biological control of whitebacked planthopper, Sogatella
furcifera Horvath (Homoptera: Delphacidae). Journal of Pest Science 77, 185-189.
Jones, R.W., Gilstrap, F.E., Andrews, K.L., 1988. Biology and life tables for the predaceous
earwig, Doru taeniatum (Dermaptera: Forficulidae). Entomophaga 33, 43-54.
Kahuthia-Gathu, R., Lohr, B., Poehling, H.M., 2008.Effect of common wild crucifer species of
Kenya on fitness of two exotic diamondback moth parasitoids, Cotesia plutellae and Diadegma
semiclausum. Crop Protection 27, 1477-1484.
Kahuthia-Gathu, R., Lohr, B., Poehling, H.M., Mbugua, P.K., 2009.Diversity, distribution and
role of wild crucifers in major cabbage and kale growing areas of Kenya. Bulletin of
Entomological Research 99, 287-297.
Kartohardjono, A., Heinrichs, E.A., 1984. Populations of the brown planthopper, Nilaparvata
lugens (Stal) (Homoptera: Delphacidae), and its predators on rice varieties with different levels
of resistance. Environmental Entomology 13, 359–365.
Karungi, J., Ekbom, B., Kyamanywa, S., 2006. Effects of organic versus conventional fertilizers
on insect pests, natural enemies and yield of Phaseolus vulgaris. Agriculture, Ecosystems and
Environment 115, 51-55.
Kavallieratos, N.G., Stathas, G.J., Athanassiou, C.G., Papadoulis, G.T., 2002. Dittrichia viscosa
and Rubus ulmifolius as reservoirs of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae)
and the role of certain coccinellid species. Phytoparasitica 30, 1-12.
Kavallieratos, N.G., Tomanovic, Z., Stary, P., Emmanouel, N.E., 2008. Vitex agnus castus and
Euphorbia characias spp. wulfenii as reservoirs of aphid parasitoids (Hymenoptera: Braconidae:
Aphidiinae). Florida Entomologist 91, 179-191.
Kawashima, M., Jung, C., 2010. Artificial ground shelters for overwintering phytoseiid mites in
orchards. Experimental and Applied Acarology 52, 35-47.
Kenmore, P.E., Carino, F.O., Perez, C.A., Dyck, V.A. Guttierez, A.P., 1984. Population
regulation of the rice brown planthopper (Nilaparvata lugens Stal) within rice fields in the
Philippines. Journal of Plant Protection in the Tropics 1, 19-37.
Khan, Z.R., Chiliswa, P., Ampong-Nyarko, K., Smart, L.E., Polaszek, A., et al. 1997.Utilisation
of wild gramineous plants for management of cereal stemborers in Africa.Insect Science and its
Application 17, 143–50.
Koclu, T., Karsavuran, Y., 1998. Natural effectiveness of the egg parasitoids of Helicoverpa
armigera (Huebner) (Lepidoptera: Noctuidae). Turk.Entomol.Derg. 22, 269-283.
Koji, S., Khan, Z.R., Midega, C.A.O., 2007.Field boundaries of Panicum maximum as a reservoir
for predators and a sink for Chilo partellus. Journal of Applied Entomology 131, 186-196.
Kumar, A., Tripathi, C.PM., Singh, R., Pandey, R.K., 1983. Biomass of Trioxys indicus, an aphid
parasitoid of Aphis craccivora, effect of host plant on the activities of the parasitoids.
Zeitschrift fur Angewandte Entomologie 96, 304-307.
Lawo, N.C., Wackers, F.L., Romeis, J., 2009. Indian Bt cotton varieties do not affect the
performance of cotton aphids. Plos One 4, e4804.
Le Ru, B.R., Mitsipa, A., 2002. Comparative life table statistics of Exochomus flaviventris reared
on the cassava mealybug, Phenacoccus manihoti, fed on four host plants. International Journal
of Tropical Insect Science 22, 175-182.
Li, H.C., 1987. Augmentation of Chrysopa species to control cotton aphids by intercropping
cotton and safflower. Chinese Journal of Biological Control 8, 109-111.
Lim, G.T., 2007. Enhancing the weaver ant, Oecophylla smaragdina (Hymenoptera: Formicidae),
for biological control of a shoot borer, Hypsipyla robusta (Lepidoptera: Pyralidae), in
Malaysian mahogany plantations. PhD dissertation, Virginia Tech.
Litsinger, J.A., Dela Cruz, C.G., Canapi, B.L., Barrion, A.T., 2007. Maize planting time and
arthropod abundance in southern Mindanao, Philippines. II. Population dynamics of natural
enemies. International Journal of Pest Management 53, 161-173.
Liu, Z.L., Wang, L.S., Du, Y.X., Yu, X.P., Lu, Z.X., Zheng, X.S., Mei, X.Q., Pan, H.P., 1999.
The dispersal patterns of predators between single-season rice fields and the neighboring non-
rice habitats, and its application in conserving natural enemies of rice pests. Acta Agriculturae
Zhejiangensis 11, 344-348.
Lixa, A.T., Campos, J.M., Resende, A.L.S., Silva, J.C., Almeida, M.M.T.B., Aguiar-Menezes,
E.L., 2010.Diversidade de Coccinellidae (Coleoptera) em plantas aromáticas (Apiaceae) como
sítios de sobrevivência e reprodução em sistema agroecológico. Neotropical Entomology 39,
354-359.
López Arroyo, I., 2009a. Alternativa para atraer y retener depredadores del pulgón café de los
cítricos. Ficha Tecnológica por Sistema Producto. Campo Experimental INIFAP-Gral. Terán,
N.L., México.
López Arroyo, I., 2009b. Aprovechamiento de la maleza para incrementar y conservar agentes de
control biológico del pulgón café de los cítricos. Ficha Tecnológica por Sistema Producto.
Campo Experimental INIFAP-Gral. Nuevo León, México.
López Arroyo, J.I., Trujillo Arriaga, J., Carrillo Sánchez, J.L., 1992. Hipótesis de los enemigos
naturales en el control de insectos en cultivos diversificados: una prueba en plagas de frijol. XV
Congreso Nacional de Control Biológico. México.
Lou, Y.G., Cheng, J.A., 1996. Influence of rice varieties on development, survival and fecundity
of Anagrus nilaparvatae. Acta Entomologica Sinica 39, 28-36.
Loya Ramírez, J.G., Ellington, J., Lucero Pulido M. del C., Carrillo, T., 2003. Impacto de
tratamientos sobre especies parasitoides de huevecillos de gusano rosado (Pectinophara
gossypyella) en algodón (Gossypium hirsutum). XXVI Congreso Nacional de Control
Biológico. México.
Lou, Y.G., Cheng, J.A., 2003. Role of rice volatiles in the foraging behaviour of the predator
Cyrtorhinus lividipennis for the rice brown planthopper Nilaparvata lugens. Biocontrol 48, 73-
86.
Luo, S.P., Li, J.C., Liu, X.X., Lu, Z.Y., Pan, W.L., Zhang, Q.W., Zhao, Z.W., 2010. Effects of
six sugars on the longevity, fecundity and nutrient reserves of Microplitis mediator. Biological
Control 52, 51–57.
Lu, X.Y., Xu, J., Li, Q.Y., 2008. Impact of Lolium perenne planted in chestnut orchards and its
mowing methods on population densities of Oligonychus ununguis and its natural enemy
Amblyseius castaneae. Chinese Journal of Biological Control 24, 108-111.
Lu, Z.X., Yang, Z.F., 1995. The effect of corn pollen on the adult longevity of Macrocentrus
linearis. Entomological Journal of East China 4, 61-64.
Ly, Z.X., Yu, X.P., Heong, K.L., Hu, C., 2006a. Dynamics of predators in rice canopy and
capacity of natural control on insect pests in paddy fields with different nitrogen regimes.Acta
Phytophylacica Sinica 33, 225-229
Ly, Z.X., Yu, X.P., Heong, K.L., Hu, C., 2006b. Impact of nitrogen fertilizer on natural control
capacities of invertebrate predators and parasitoids and its demonstration in rice-based
ecosystem Acta Agriculturae Zhejiangensis 18, 128-132.
Ma, X.M., Liu, X.X., Zhang, Q.W., Zhao, J.Z., Cai, Q.N., Ma, Y.A., Chen, D.M., 2006.
Assessment of cotton aphids, Aphis gossypii, and their natural enemies on aphid-resistant and
aphid-susceptible wheat varieties in a wheat–cotton relay intercropping system. Entomologia
Experimentalis et Applicata 121, 235–241.
Ma, X.M., Zhang, H.Z., Liu X.X., Ding, J., Zhang, Q.W., 2007. Economic profit of wheat
transgenic Bt cotton intercropping culture and its effect on cotton aphids and natural enemies.
Acta Phytophylacica Sinica 4, 167-172.
Mailloux, J., Le Bellec, F., Kreiter, S., Tixier, M.S., Dubois, P., 2010. Influence of ground cover
management on diversity and density of phytoseiid mites (Acari: Phytoseiidae) in
Guadeloupean citrus orchards. Experimental and Applied Acarology 52, 275-290.
Mandour, N.S., Ren, S.X., Qiu, B.L., 2007.Effect of Bemisia tabaci honeydew and its
carbohydrates on search time and parasitization of Encarsia bimaculata. Journal of Applied
Entomology 131, 645-651.
Manzano, M.R., van Lenteren, J.C., Cardona, C., 2003. Influence of pesticide treatments on the
dynamics of whiteflies and associated parasitoids in snap bean fields. Biocontrol 48, 685-693.
Martinez, J.J.I., Cohen, M., Mgocheki, N., 2011.The response of an aphid tending ant to artificial
extra-floral nectaries on different host plants.Arthropod Plant Interactions 5, 185-192.
Martinoia, G.I., Carmona, D.M., 2011. Presencia de Harmonia axyridis (Coleoptera:
Coccinellidae) en los refugios vegetales del cultivo de papa y su papel como controladora
biológica. Geociencies 18, 5-11.
Martins, I.C., Cividanes, F.J., Barbosa, J.C., Araujo, E.S., Haddad, G.Q., 2009.Faunal analysis
and population fluctuation of Carabidae and Staphylinidae (Coleoptera) in no-tillage and
conventional tillage systems.Revista Brasileira de Entomologia 53, 432-443.
Materu, M.E.A., 1971. Population dynamics of Acanthomia spp. (Hemiptera: Coreidae) on beans
and pigeonpeas in the Arusha area of Tanzania. East African Agriculture and Forestry Journal
36, 361–383.
Matienzo, Y., 2005. Conservación de artrópodos benéficos en un sistema de producción agrícola
urbano. MSc thesis, Universidad Agraria de La Habana, Junio.
Matienzo, Y., 2007. Prácticas agroecológicas para la conservación de la entomofauna benéfica
en sistemasagrícolasurbanos. 2007. Memorias del curso taller ‘Producción local de
entomófagos’.
Matienzo, Y., Rijo, E., Milán, O., Torres, N., Larrinaga, J., Massó, E., 2007. Contribución al
conocimiento de especies botánicas con potencialidad para el fomento de reservorios de
insectos benéficos en la agricultura urbana. Taller internacional Producción y manejo
agroecológico de artrópodos benéficos, Ciudad de la Havana, Cuba.
Matienzo, Y., Veitia, M.M., Alayon, G., 2011. Composición y riqueza de insectos y arañas
asociados a plantas florecidas en sistemas agrícolas urbanos. Fitosanidad 15, 25-30.
Matin, S.B., Sahragard, A., Rasollian, G., 2009. Some biological parameters of Lysiphlebus
fabarum (Hymenoptera : Aphidiidae), a parasitoid of Aphis fabae (Homoptera : Aphidiidae)
under laboratory conditions. Munis Entomology & Zoology 4, 193-200.
Matteson, P.C., 1982. The effects of intercropping with cereals and minimal permethrin
applications on insect pests of cowpea and their natural enemies in Nigeria. Tropical Pest
Management 28, 372-380.
Matteson, P.C., 2000. Insect pest management in tropical Asian irrigated rice. Annual Review of
Entomology 45, 549-574.
Medeiros, M.A., Ribeiro, P.A., Morais, H.C., Castelo Branco, M., Suji, E.R., Salgado-Laboriau,
M.L., 2010. Identification of plant families associated with the predators Chrysoperla externa
(Hagen) (Neuroptera: Chrysopidae) and Hippodamia convergens Guérin-Menéville (Coleoptera:
Coccinelidae) using pollen grain as a natural marker. Brazilian Journal of Biology 70, 293-300.
Medeiros, M.A., Suji, E.R., Morais, H.C., 2009. Effect of plant diversification on abundance of
South American tomato pinworm and predators in two cropping systems. Horticultura
Brasileira 27, 300-306.
Men, X.Y., Ge, F., Yardim, E.N., Parajulee, M.N., 2004.Evaluation of winter wheat as a
potential relay crop for enhancing biological control of cotton aphids in seedling
cotton.BioControl 49, 701–714.
Meirelles, A.P., Carneiro, T.R., Fernandes, O.A., 2009. Efeito de diferentes fontes de carboidrato
e da privacao de alimento sobre aspectos biologicos de Telenomus remus Nixon (Hymenoptera :
Scelionidae). Revista Brasileira de Entomologia 53, 457–460.
Mexzon, R.G., Chinchilla, C.M., 1998. Especies vegetales atrayentes de la entomofauna benéfica
en plantaciones de palma aceitera (Elaeis guineensis Jacq.) en Costa Rica. ASD Oil Palm
Papers 19, 1-22.
Midega, C.A.O., Khan, Z.R., Van Den Berg, J., Ogol, C.K.P.O., Dippenaar-Schoeman, A.S.,
Pickett, J.A., Wadhams, L.J., 2008. Response of ground-dwelling arthropods to a ‘push–pull’
habitat management system: spiders as an indicator group. Journal of Applied Entomology 132,
248-254.
Mikhalsev, V.P., 1994. The ecologically safe and pesticide-free technologies of plant-growing
production.Proceedings of the All-Russian Scientific-production Conference. Krasnodar,
August 24-26, 1994. Part 1, Pushchino 1994, pp. 97-99.
Momen, F.M., 2004. Suitability of the pollen grains, Ricinus communis and Helianthus annuus
as food for six species of Phytoseiid mites (Acari: Phytoseiidae). Acta Phytopathologica et
Entomologica Hungarica 39, 415-422.
Momen, F.M., Abdelkhader, M.M., 2010. Fungi as food source for the generalist predator
Neoseiulus barkeri (Hughes) (Acari: Phytoseiidae). Acta Phytopathologica et Entomologica
Hungarica 45, 401-409.
Morales, H., Perfecto, I., 2000.Traditional knowledge and pest management in the Guatemalan
highlands. Agriculture and Human Values 17, 49-63.
Morales, H., Perfecto I., Ferguson, B., 2001. Traditional Cakchiquel soil fertilization and its
impact on insect pest populations in corn. Agriculture, Ecosystems and Environment 84, 145-
155.
Morrill, W.L., Almazan, L.P., 1990. Effect of host plant species and age of rice bug (Hemiptera:
Alydidae) eggs on parasitism by Gryon nixoni (Hymenoptera: Scelionidae). Journal of
Entomological Science 25, 450-452.
Msebah, A.H., El-Husseini, M.M., 2009. Effect of weed borders on insect pests and their
associated predators on sugar beet and cotton plant fields. Egyptian Academic Journal of
Biological Sciences 2, 73-80.
Mujica, Y., Martinez, M., Aleman, J., Ravelo, J., 2009. Fluctuacion poblacional de plagas de la
col (Brassica oleracea) y otros enemigos naturales en dos agroecosistemas. Cultivos Tropicales
30(4).
Munyuli, T.M.B., Kyamanywa, S., Luther, G.C., 2008. Effects of groundnut genotypes, cropping
systems and insecticides on the abundance of native arthropod predators from Uganda and
Democratic Republic of Congo.Bulletin of Insectology 61, 11-19.
Munyuli, M.B.T., Luther, G.C., Kyamanywa, S., 2007. Effects of cowpea cropping systems and
insecticides on arthropod predators in Uganda and Democratic Republic of the Congo.Crop
Protection 26, 114-126.
Murta, A.F., Ker, F.T.O., Costa, D.B., Espirito-Santo, M.M., Faria, M.L., 2008. Efeitos de
remanescentes de Mata Atlântica no controle biológico de Euselasia apisaon (Dahman)
(Lepidoptera: Riodinidae) por Trichogramma maxacalii (Voegelé e Pointel) (Hymenoptera:
Trichogrammatidae). Neotropical Entomology 37, 229-232.
Nagirnyak, A.F., Krasavina, L.P., 2005. Use the nectar plants in the system of biological method
protection of the plants. The second “All-Russian congress on plant protection,” St. Petersburg,
December 5-10 2005.Recovery of the plant-sanitary of eco-landscape, volume 2, pp. 99-100.
Nampala, P., Adipala, E., Ogenga-Latigo, M.W., Kyamanywa, S., Obugo, J.E., 1999. Effect of
cowpea monocultures and polycultures with sorghum and greengram on predatory arthropods.
Annals of Applied Biology 135, 457-461.
Narvaez, A., Cancino, J., Canal Daza, N., Wyckhuys, K.A.G., 2012. Effect of different dietary
resources on longevity, carbohydrate metabolism, and ovarian dynamics in two fruit fly
parasitoids. Arthropod Plant Interactions, in press.
Ndemah, R., Schulthess, F., Phoeling, H.M, Borgemeister, C., 2001. Natural enemies of
lepidopterous borers on maize and elephant grass in the forest zone of Cameroon with special
reference to Busseola fusca (Fuller) (Lepidoptera: Noctuidae). Bulletin of Entomological
Research 91, 205-212.
Ndemah, R., Schulthes, F., Poehling, H.M., Borgemeister, C., Cardwell, K.F., 2003. Factors
affecting infestations of the maize stalk borer Busseola fusca (Lepidoptera: Noctuidae) on
maize in the forest zone of Cameroon, with special reference to Scelionid egg parasitoids.
Environmental Entomology 32, 51-60.
Neuenschwander, P., Hammond, W.N.O., Herren, H.R., 1987. Biological control of the cassava
mealybug (Phenacoccus manihoti) by the exotic parasitoid, Epidinocarsis lopezi, in: Terry,
E.R., Akoroda, M.O., Arene, O.B. (Eds.). Proceeding of the symposium on Tropical root crops:
Root crops and the African food crisis. Owerri, Nigeria.
Nwanze, K.F., Mueller, R.A.E., 1989. Management options for sorghum stem borers for farmers
in the semi-arid tropics. Proceedings of the International Workshop on Sorghum Stem Borers,
17-20 Nov 1987, lCRISAT Center, India. Patancheru, India.
Nwilene, F.E., Togola, A., Agunbiade, T.A., Ogah, E.O., Ukwungwu, M.N., Hamadoun, A.,
Kamara, S.I., Dakouo, D., 2008. Parasitoid biodiversity conservation for sustainable
management of the African rice gall midge, Orseolia oryzivora (Diptera: Cecidomyiidae) in
lowland rice. Biocontrol Science and Technology 18, 1075-1081.
Nyambo, B.T., 1986. Studies of the bollworm, Helitohis armigera Hubner, the key cotton pest in
Tanzania, as a basis for improved integrated pest management.PhD thesis, University of
London, UK.
Oduor, G.I., Simons, S.A., 2003. Biological control in IPM for coffee, in: Neuenschwander, P.,
Borgemeister, C., Langewald, J. (Eds.), Biological control in IPM systems in Africa, pp. 347-
362. CABI Publishing, Wallingford, UK.
Ofuya, T.I., 1989. Effect of weeds on colonization of cowpea by Aphis craccivora Koch
(Homoptera: Aphididae) and its major predators in Nigeria. Tropical Pest Management 35,
403-405.
Ofuya, T.I., 1995. Colonization and control of Aphis craccivora Koch (Homoptera: Aphididae)
by coccinellid predators in some resistant and susceptible cowpea varieties in Nigeria. Crop
Protection 14, 47-50.
Olkowski, W., Zhang, A., 1998. Habitat management for biological control, examples from
China, in: Pickett, C.H., Bugg, R.L. (Eds.), Enhancing biological control: habitat management
to promote natural enemies of agricultural pests. University of California Press, Berkeley, pp.
255–270.
Onzo, A., Hanna, R., Negloh, K., Toko, M., Sabelis, M.W., 2005. Biological control of cassava
green mite with exotic and indigenous phytoseiid predators—Effects of intraguild predation
and supplementary food. Biological Control 33, 143-152.
Ooi, P.A.C., 1988. Laboratory studies of Tetrastichus sokolowskii. Biocontrol 33, 145-152.
Overholt, W.A., Conlong, D.E., Kfir, R., Schulthess, F., Sétamou, M., 2003. Biological control
of gramineous Lepidopteran stem borers in sub-Saharan Africa, in: Neuenschwander, P.,
Borgemeister, C., Langewald, J. (Eds.), Biological control in IPM systems in Africa. CABI
Publishing, Wallingford, UK, pp. 131-144.
Pawar, C.S., Bhatnagar, V.S., Jadhav, D.R., 1986. Heliothis species and their natural enemies
with their potential for biological control. Proceedings of the Indian Academy of Sciences 95,
695-703.
Pawar, C.S., Reddy, M.V., Haware, M.P., Ramakrishna, A., 1999.Insect pests, diseases and
weeds of pigeonpea and their management. In Sinha, M.P. (Ed.), Recent advances in
ecobiological research, volume 2. A.P.H. Publishing Corporation, New Delhi.
PCA, Philippine Coconut Authority, 2003. Integrated pest management for slug caterpillar.CPD
Technoguide No. 10.Crop Protection Division, Davao Research Center, Davao City,
Philippines.
Penagos, D.I., Magallanes, R., Valle, J., Cisneros, J., Martinez, A.M., Goulson, D., Chapman,
J.W., Caballero, P., Cave, R.D., Williams, T., 2003.Effect of weeds on insect pests of maize
and their natural enemies in Southern Mexico. International Journal of Pest Management 49,
155-161.
Perdikis, D., Favas, C., Lykouressis, D., Fantinou, A., 2007. Ecological relationships between
non-cultivated plants and insect predators in agroecosystems: the case of Dittrichia viscosa
(Asteraceae) and Macrolophus melanotoma (Hemiptera: Miridae). Acta Oecologica 31, 299-
306.
Pereira, J.L., Picanco, M.C., Pereira, E.J., Silva, A.A., Jakelaitis, A., Pereira, R.R., Xavier, V.M.,
2010. Influence of crop management practices on bean foliage arthropods. Bulletin of
Entomological Research 100, 679-688.
Pereira, J.L., Picanco, M.C., Silva, A.A., Barros, E.C., Xavier, V.M., Gontijo, P.C., 2007. Effect
of herbicides on soil arthropod community of bean cultivated under no-tillage and conventional
systems.Planta Daninha 25, 61-69.
Pérez, N., Vázquez, L.L., 2001. Manejo ecológico de plagas, in: Transformando el campo
cubano. Avances de la Agricultura Sostenible. Ed. ACTAF. La Habana. Pp. 191-223.
Perfecto, I., Castiñeiras, A., 1998. Deployment of the predaceous ants and their conservation in
agroecosystems, in: Barbosa, P. (Ed.), Conservation Biological Control. Academic Press, San
Diego, California, pp. 269-289.
Perfecto, I., Sediles, A., 1992.Vegetational diversity, ant (Hymenoptera, Formicidae), and
herbivorous pests in a Neotropical agroecosystem. Environmental Entomology 21, 61-67.
Perfecto, I., Vandermeer, J., 1996.Microclimatic changes and the indirect loss of ant diversity in
a tropical system. Oecologia 108, 577–582.
Perfecto, I., Vandermeer, J.H., Bautista, G.L., Nunez, G.I., Greenberg, R., Bichier, P., Langridge,
S., 2004. Greater predation in shaded coffee farms: The role of resident neotropical birds.
Ecology 85, 2677-2681.
Philpott, S.M., 2005. Changes in arboreal ant populations following pruning of coffee shade-
trees in Chiapas, Mexico. Agroforestry Systems 64, 219-224.
Philpott, S.M., Foster, P.F., 2005. Nest-site limitation in coffee agroecosystems: Artificial nests
maintain diversity of arboreal ants. Ecological Applications 15, 1478-1485.
Philpott, S.M., Perfecto, I., Vandermeer, J., 2006. Effects of management intensity and season on
arboreal ant diversity and abundance in coffee agroecosystems. Arthropod Diversity and
Conservation 1, 125-141.
Pimbert, M.P., Srivastava, C.P., 1989. Vegetation management and the biological control of
Helicoverpa armigera in chickpea. International Chickpea Newsletter 21, 16-19.
Pinkus-Rendón, M., Ibarra-Nuñez. G., Parra, V., García-Ballinas, J., Henaut, Y., 2006. Spider
diversity in coffee plantations with different management in Southeast Mexico. Journal of
Arachnology 34, 104-112.
Pluess, T., Opatovsky, I., Gavish-Regev, E., Lubin, Y., Schmidt, M.H., 2008.Spiders in wheat
fields and semi-desert in the Negev (Israel). Journal of Arachnology 36, 368-373.
Pu, T.S., Zeng, T., Wei, D.W., 1991. An integrated evaluation on 20 plant pollen as diet for mass
rearing Amblyseius makuwa (Acari: Phytoseiidae). Chinese Journal of Biological Control 7,
111-114.
Pulido Herrera, A., Capiz Angeles, M.A., Quintero Meza, M.J., 1998. Evaluación del impacto de
seis insecticidas sobre fauna benéfica en el control de chinche Lygus spp. en el Valle de
Méxicali, Baja California. XXI Congreso Nacional de Control Biológico. México.
Ramnath, S., Uthamasamy, S., 1992. Interaction of host plant resistance and natural enemes for
the management of bollworm Heliothis armigera on cotton, in: Ananthakrishnan, T.N. (Ed.),
Emerging trends of biological control of phytophagous insects. Oxford and IBH Publishing Co.
Ltd., New Delhi, pp. 37-42.
Rao, G.V.R., 2005. Integrated pest management in legumes: present status and future needs. IPM
research at ICRISAT: present status and future priorities. Proceedings of the in-house review,
5th April 2005.
Rao, M.S., 2008. Pest management in organic farming using crop-crop diversity, in:
Venkateswarlu, B., Balloli, S.S., Ramakrishna, Y.S. (Eds.), Organic farming in rainfed
agriculture: opportunities and constraints. ICAR, Central Research Institute for Dryland
Agriculture, Hyderabad, India.
Rao, R.S.N., Chari, M.S., Gunneswara Rao, S., 1990. Further record of natural enemies on the
insect pests of tobacco in Andhra Pradesh. Journal of Biological Control 4, 65-66.
Rapusas, H.R., Bottrell, D.G., Coll, M., 1996.Intraspecific variation in chemical attraction of rice
to insect predators. Biological Control 6, 394-400.
Ratnadass, A., 2010. Using pigeon pea border to regulate fruit worm on okra. In: ICRISAT West
and Central Africa: towards greater impacts. Annual Report, ICRISAT, Bamako, Mali.
Rekha, B.S., Ramkumar, J., Kandibane, M., Raguraman, S., Swamiappan, M., 2009.Diversity of
Coccinellids in cereals, pulses, vegetables and in weeded and partially-weeded rice-cowpea
ecosystems in Madurai District of Tamil Nadu.Madras Agricultural Journal 96, 251-264.
Rezende, M.Q. 2010. Etnoecologia e controle biológico conservativo em cafeeiros sob sistemas
agroflorestais. MSc thesis, Universidad Federal de Vicosa, Brazil, 79 p.
Richter, A., Klein, A.M., Tscharntke, T., Tylianakis, J.M., 2007. Abandonment of coffee
agroforests increases insect abundance and diversity. Agroforestry Systems 69, 175–182.
Rickson, F.R., Rickson, M.M., 1998. The cashew nut, Anacardium occidentale (Anacardiaceae)
and its perennial association with ants: extrafloral nectary location and the potential for ant
defense. American Journal of Botany 85, 835-849.
Rico-Gray, V., 1993.Use of plant-derived food resources by ants in the dry tropical lowlands of
coastal Veracruz, Mexico.Biotropica 25, 301-315.
Rico-Gray, V., Morais, H.C., 2006. Efecto de una fuente de alimento experimental sobre una
asociacion hormiga-hemiptero. Acta Zoologica Mexicana 22, 23-28.
Rijo, E., Rodríguez, T., Acosta, N., Mollineda, M., Acosta, B., 1999. La presencia de fitófagos y
sus reguladores biológicos en organopónicos. Informe de proyecto. Instituto Investigaciones de
Sanidad Vegetal, Ciudad de La Habana.
Rocha, F., Infante, F., 2009. Diversidad de enemigos naturales asociados a inflorescencias de
mango ataulfo en el Soconusco, Chiapas, México. XXXII Congreso Nacional de Control
Biológico. México.
Rogochaya, L.G., 1971. To question about food relationship tachina flies with wild flowering
vegetation and attraction them in orchards. Biological methods of protection fruit and vegetable
crops from insect pest, diseases and weed as bases of the integrated systems.Thesis’s report,
October 1971, Kishinev, pp. 86-87.
Romeis, J., Romeis, O., Shanower, T.G., 1995. Paratrechina longicornis (Hymenoptera:
Formicidae), a predator of Helicoverpa armigera (Lepidoptera: Noctuidae) eggs. Journal of
Biological Control 9, 56–58.
Rosado, M.C., 2007. Plantas favoraveis a agentes de controle biologico. MSc thesis,
Universidade Federal de Vicosa.
Rubtsov, I.A., 1944. Sunn pest egg parasitoid in Tajikistan.Scientifically popular library, Tajik
Branch of the Academy of USSR, Stalinabad, 4, pp. 1-56.
Saidov, N.S., Landis, D.A., Bohssini, M., 2007. A history of habitat management in the former
USSR and the Commonweatlh of Independent States and current research in Central Asia, in:
Maredia, K.M., Baributsa, D.N. (Eds.), Integrated pest management in central Asia.
Proceedings of the central Asia Region integrated pest management stakholders forum,
Dushanbe, Tajikistan, May 27-29, 2007.
Sakin, G.V., Ulusoy, M.R., 2009. The effects of different temperatures and diets on the biology
of Cales noacki Howard (Hymenoptera: Aphelinidae), a parasitoid of the citrus woolly whitefly.
Turkish Journal of Agriculture and Forestry 33, 267-275.
Saljoqi, A.U.R., Khajjak, A.S.K., 2007. Effect of different artificial diets on the efficiency and
development of Trichogramma chilonis (Ishii) (Hymenoptera: Trichogrammatidae). Sarhad
Journal of Agriculture 23, 129-133.
Sana-Ullah, M., Arif, M.J., Gogi, M.D., Shabid, M.R., Adid, A.M., Raza, A., Alli, A., 2011.
Influence of different plant genotypes on some biological parameters of cotton mealybug,
Phenacoccus solenopsis and its predator, Coccinella septempunctata under laboratory
conditions. International Journal of Agricultural Biology 12, 125–129.
Santos, T.M., Boica, A.L., Soares, J.J., 2003. Influence of cotton trichomes on biological
trichomes on biological aspects and predatory ability of Chrysoperla externa (Hagen) fed with
Aphis gossypii Glover. Bragantia 62, 2.
Schulthess, F., Chabi-Olaye, A., Goergen, G., 2001. Seasonal flucturations of noctuid stemborer
egg parasitism in southern Benin with special reference to Sesamia calamistis Hampson
(Lepidoptera: Noctuidae) and Telenomus spp. (Hymnoptera: Scelionidae) on maize. Biocontrol
Science and Technology 11, 765-777.
Sekamatte, M.B., Kyamanywa, S., Willson, H.R., Russell-Smith, A., 2002. Effect of placement
method and rate of application of crushed fish bones on the activity of predatory ants and
impact on termite damage to maize. International Journal of Tropical Insect Science 22, 199-
204.
Sekamatte, M.B., Ogenga-Latigo, M.W., Russell-Smith, A., 2001a. The effect of maize stover
used as mulch on termite damage to maize and activity of predatory ants. African Crop Science
Journal 9, 411-419.
Sekamatte, M.B., Ogenga-Latigo, M., Russel-Smith, A., 2001b. The potential of protein- and
sugar-based baits to enhance predatory ant activity and reduce termite damage to maize in
Uganda. Crop Protection 20, 653-662.
Sekamatte, B.M., Ogenga-Latigo, M., Russell-Smith, A., 2003. Effects of maize-legume
intercrops on termite damage to maize, activity of predatory ants and maize yields in Uganda.
Crop Protection 22, 87-93.
Setamou, M., Jiang, N., Schulthess, F., 2005. Effect of the host plant on the survivorship of
parasitized Chilo partellus Swinhoe (Lepidoptera: Crambidae) larvae and performance of its
larval parasitoid Cotesia flavipes Cameron (Hymenoptera: Braconidae). Biological Control 32,
183-190.
Settle, W.H., Ariawan, H., Astuti, E.T., Cahyana, W., Hakim, A.L., Hinayana, D., Lestari, A.S.P.,
1996. Managing tropical rice pests through conservation of generalist natural enemies and
alternative prey. Ecology 77, 1975-1988.
Shaltiel, L., Coll, M., 2004. Reduction of pear psylla damage by the predatory bug Anthocoris
nemoralis (Heteroptera: Anthocoridae): the importance of orchard colonization time and
neighboring vegetation. Biocontrol Science and Technology 14, 811-821.
Shannag, H.K., Obeidat, W.M., 2008. Interaction between plant resistance and predation of
Aphis fabae (Homoptera: Aphididae) by Coccinella septempunctata (Coleoptera:
Coccinellidae). Annals of Applied Biology 152, 331-337.
Shannag, H.K., Obeidat, W.M., 2006. Voracity and conversion efficiency by larvae of
Coccinella septempunctata L. (Coleoptera: Coccinellidae) on Aphis fabae Scop. (Homoptera:
Aphididae) reared on two faba bean cultivars with different levels of resistance. Applied
Entomology and Zoology 41, 521-527.
Shanower, T.G., Romeis, J., Minja, E.M., 1999. Insect pests of pigeonpea and their management.
Annual Review of Entomology 44, 77-96.
Sharma, R.P., Yadav, R.P., 1993. Response of lentil varieties to the incidence of bean aphid
(Aphis craccivora Koch.) and its predatory coccinellids. LENS Newsletter 20, 60-62.
Sigsgaard, L., 2000. Early season natural biological control of insect pests in rice by spiders -
and some factors in the management of the cropping system that may affect this control.
Proceedings of the 19th European Colloquium of Arachnology, Århus 17-22 July 2000, pp. 57-
64.
Sigsgaard, L., Esbjerg, P., 1997. Cage experiments on Orius tantillus predation of Helicoverpa
armigera. Entomologia Experimentalis et Applicata 82, 311-318.
Silva, J.O., Oliveira, K.N., Santos, K.J., Espirito-Santo, M.M., Neves, F.S., Faria, M.L., 2010.
Effects of landscape structure and Eucalyptus genotype on the abundance and biological
control of Glycaspis brimblecombei Moore (Hemiptera: Psyllidae). Neotropical Entomology 39,
91-96.
Silveira, L.C.P., Bueno, V.H.P., Pierre, L.S.R., Mendes, S.M., 2003. Plantas cultivadas e
invasoras como habitat para predadores do gênero Orius (Wolff) (Heteroptera: Anthocoridae).
Bragantia 62, 261-265.
Silveira, L.C.P., Filho, E.B., Pierre, L.S.R., Cunha, F.S., Cassa, J.N., 2009. Marigold (Tagetes
erecta L.) as an attractive crop to natural enemies in onion fields.Scientia Agricola 66, 780-787.
Simón, F., 1999. Los plaguicidas en el manejo integrado de plagas del café y su impacto
ecotoxicológico sobre ecosistemas de montaña. PhD Thesis, Instituto de Investigaciones de
Sanidad Vegetal. La Habana, Cuba.
Simon, S., Bouvier, J.C., Debras, J.F., Sauphanor, B., 2010.Biodiversity and pest management in
orchard systems: a review. Agronomy for Sustainable Development 30, 139-152.
Sithanantham, S., Bhatnagar, V.S., Jadhav, D.R. & Reed, W., 1982. Some aspects of
Trichogramma spp. in eggs of Heliothis armigera (Hb.) (Lepidoptera: Noctuidae). International
symposium on Trichogramma and other egg parasitoids, Antibes, France.
Sitaramaiah, S., Rao, S.G., Ramaprasad, G., Sreedhar, U., 2001. Role of biological control
agents in tobacco insect pest management, in: Upadhyaya, K., Mukerji, K.G., Chamola, B.P.
(Eds.), Biocontrol potential and its exploitation in sustainable agriculture. New York, Kluwer
Academic/Plenum Publishers, pp. 389-400.
Smith, H.A., McSorley, R., Sierra, J.A., 2001. Effect of intercropping common bean with poor
hosts and nonhosts on numbers of immature whiteflies (Homoptera: Aleyrodidae) in the
Salamá Valley, Guatemala. Environmental Entomology 30, 89-100.
Solangi, G.S., Mahar, GM., Oad, F.C., 2008. Presence and abundance of different insect
predators against sucking insect pests of cotton. Journal of Entomology 5, 31-37.
Song, B.Z., Wang, M.C., Yun, K., Yao, Y.C., Wu, H.Y., Li, Z.R., 2010. Interaction of the
dominant pests and natural enemies in the experimental plots of the intercropping aromatic
plants in pear orchard.Scientia Agricultura Sinica 43, 3590-3601.
Song, B.Z., Zhang, J., Hu, J.H.., Wu, H.Y., Kong, Y., Yao, Y.C., 2011. Temporal dynamics of
the arthropod community in pear orchards intercropped with aromatic plants. Pest Management
Science 67, 1107–1114.
Songa, J.M., Jiang, N., Schulthess, F., Omwega, C., 2007. The role of intercropping different
cereal species in controlling lepidopteran stemborers on maize in Kenya. Journal of Applied
Entomology 131, 40-49.
Soroushmehr, Z., Sahragard, A., Salehi, L., 2008. Comparative life table statistics for the
ladybeetle Scymnus syriacus reared on the green citrus aphid, Aphis spiraecola, fed on two host
plants. Entomological Science 11, 281-288.
Sousa, E.H.S., Matos, M.C.B., Almeida, R.S., Teodoro, A.V., 2011. Forest fragments'
contribution to the natural biological control of Spodoptera frugiperda Smith (Lepidoptera:
Noctuidae) in maize. Brazilian Archives of Biology and Technology 54, 755-760.
Sperber, C.F., Nakayama, K., Valverde, M.J., Neves, F., 2004. Tree species richness and density
affect parasitoid diversity in cacao agroforestry. Basic and Applied Ecology 5, 241-251.
Stenchly, K., 2010. Spider communities in Indonesian cacao agroforestry: diversity, web density
and spatio-temporal turnover. Georg-August Universitat zu Gottingen, Germany. Dissertation.
Sueldo de Escano, M.R., Virla, E.G., 2009. Doru lineare (Dermaptera: Fortifuclidae), insecto
benéfico en cultivos de maíz del norte argentino: preferencias alimenticias y tasas de consumo.
Boletin Sanidad Vegetal 35, 39-47.
Sulaiman, G., Hussein, M.Y., Idris, A.B., 2004. The abundance and parasitism on the egg masses
of the Asiatic corn borer Ostrinia furnacalis Guenee in weedy and weed-free cornfields in
Malaysia. International Journal of Agriculture & Biology 6, 36-38.
Tamo, M., Arodokoun, D.Y., Zenz, N., Tindo, M., Agboton, C., Adeoti, R., 2002. The
importance of alternative host plants for the biological control of two key cowpea insect pests,
the pod borer Maruca vitrata (Fabricius) and the flower thrips Megalurothrips sjostedti
(Trybom), in: Fatokun, C.A., Tarawali, S.A., Singh, B.B., Kormawa, P.M. Tamò, M (Eds.),
Challenges and opportunities for enhancing sustainable cowpea production. Proceedings of the
World Cowpea Conference III held at the International Institute of Tropical Agriculture (IITA),
Ibadan, Nigeria, 4-8 September, 2000. IITA, Ibadan, Nigeria, pp 81-93.
Tavares, W.S., Cruz, I., Silva, R.B., Figueiredo, M.L.C., Ramalho, F.S., Serrao, J.E., Zanuncio,
J.C., 2011. Soil organisms associated to the weed suppressant Crotalaria juncea (Fabaceae)
and its importance as a refuge for natural enemies.Planta daninha29, 473-479.
Taylor, T.H.C., 1945. Recent investigations of Antestia spp. in Uganda. Part I. East African
Agricultural Journal 10, 223-233.
Tscharntke, T., Bommarco, R., Clough, Y., Crist, T.O., Kleijn, D., Rand, T.A., Tylianakis, J.M.,
van Nouhuys, S., Vidal, S., 2008. Reprint of ‘‘Conservation biological control and enemy
diversity on a landscape scale’’. Biological Control 45, 238-253.
Togni, P.H.B., 2009. Bases ecológicas para o manejo de Bemisia tabaci (Genn.) biótipo B
(Hemiptera: Aleyrodidae) em sistemas orgânicos de produção de tomate. MSc thesis,
Universidad de Brasilia.
Toko, M., O’Neil, R.J., Yaninek, J.S., 1994. Effect of cassava exudate and prey densities on the
survival and reproduction of Typhlodromalus limonicus (Garman & McGregor) s.l. (Acari:
Phytoseiidae), a predator of the cassava green mite, Mononychellus tanajoa (Bondar)
(Acari:Tetranychidae). Experimental & Applied Acarology 18, 221-231.
Trujillo, J., Altieri, M.A., 1990. A comparison of aphidophagous arthropods on maize
polycultures and monocultures, in Central Mexico. Agric. Ecosyst. Environ. 31, 337–49.
Tuncbilek, A.S., Cinar, N., Campolat, U., 2010. Comparing flower nectar and artificial diet on
the longevity and progeny production of Trichogramma turkestanica.Proceedings of the 10th
International Working Conference on Stored Product Protection.Julius-Kühn-Archiv, 425, 754.
Uckan, F., Ergin, E., 2003. Temperature and food source effects on adult longevity of Apanteles
galleriae Wilkinson (Hymenoptera: Braconidae). Environmental Entomology 32, 441-446.
Van den Berg, H., Nyambo, B.T., Waage, J.K., 1990. Parasitism of Helicoverpa armígera
(Lepidoptera: Noctuidae) in Tanzania: analysis of parasitoid-crop associations. Environmental
Entomology 19, 1141-1145.
van Hamburg, H., Guest, P.J., 1997. The impact of insecticides on beneficial arthropods in cotton
agro-ecosystems in South Africa. Archives of Environmental Contamination and Toxicology
32, 63-68.
Van Mele, P., 1998. Integrated fruit production of mango and durian in the eastern region of
Thailand. Vietnamese-Belgian IPM Project Technical Report No. 2, 22 pp.
Van Mele , P., Cuc, N.T.T., 2000 . Evolution and status of Oecophylla smaragdina (Fabricius) as
a pest control agent in citrus in the Mekong Delta, Vietnam. International Journal of Pest
Management 46, 295-301.
Van Mele, P., Cuc, N.T.T., 2001. Farmers' perceptions and practices in use of Dolichoderus
thoracicus (Smith) (Hymenoptera: Formicidae) for biological control of pests of sapodilla.
Biological Control 20, 23-29.
Van Mele, P., van Lenteren, J.C., 2002. Survey of current crop management practices in a
mixed-ricefield landscape, Mekong Delta, Vietnam—potential of habitat manipulation for
improved control of citrus leafminer and citrus red mite. Agriculture, Ecosystems and
Environment 88, 35-48.
Vasconcellos-Neto, J., Romero, G.Q., Santos, A.J., Dippenaar-Schoeman, A.S., 2007.
Associations of spiders of the genus Peucetia (Oxyopidae) with plants bearing glandular
hairs.Biotropica 39, 221-226.
Vázquez, L., 1999. La conservación de los enemigos naturales de plagas en el contexto de la
fitoprotección. BoletínTécnico del INISAV (Ciudad de La Habana) 5, 1-75.
Vázquez, L.L., 2001. Principales estrategias y componentes del programa de manejo integrado
de plagas del cafeto en Cuba. En: XVII Congreso Venezolano de Entomología. Conferencias.
pp. 55-63.
Vázquez, L.L., 2007. Desarrollo del manejo agroecológico de plagas en los sistemas agrarios de
Cuba. Fitosanidad 11, 39.
Vázquez, L.L., 2009. Una visión entomológica de la introducción del maíz transgénico FR Bt 1
en Cuba, in: Funes-Monzote, F., Freire, E. (Eds.), Transgénicos. ¿Qué se gana?, ¿qué se pierde?
Textos para un debate en Cuba. Ed. Acuario, Centro Félix Varela. La Habana, pp. 159-188.
Vazquez, L.L., 2011. La cerca viva perimetral de la finca como practica agroecológica en el
manejo de plagas, in: Manual para la adopción del manejo agroecológico de plagas en fincas de
la agricultura suburbana. La Habana, pp. 69-83.
Vázquez, L.L., Figueroa, S.C., Pérez, A.C., Michelena, J.G., García, J.A., Fernández, A.R.,
Barrios, M.B., Rodríguez, L.A., Sánchez, R.G., Santos, T.C., Mollineda, M.F., Rodríguez, M.P.,
Campbell, I.E., Cardona, L.L., Peña, E.C., Torres, T.R., Suárez, O.C., 2010. Diagnóstico de la
utilización de entomófagos y entomopatógenos para el control biológico de insectos por los
agricultores en Cuba. Fitosanidad 14, 159-169.
Vázquez, L.L., Fernández, E., Lauzardo, J., García, Alfonso, J., Ramírez, R., 2005. Manejo
agroecológico de plagas en fincas de la agricultura urbana (MAPFAU). Ed. CIDISAV, Ciudad
de La Habana.
Vázquez, L.L., López, D., 2000. Comportamiento de las poblaciones de la chinchita Cyrtopeltis
tenuis Reuter (Heteroptera: Miridae) en el cultivo del tomate infestado con la mosca blanca
Bemisia tabaci. (Homoptera:Aleyrodidae). Fitosanidad 4, 3-4.
Vázquez, L.L., Matienzo, Y., Alfonso, J., Moreno, D., Álvarez, A., 2009. Diversidad de especies
de hormigas (Hymenoptera: Formicidae) en cafetales (Coffea arabica) afectados por
Hypothenemus hampei (Coleoptera: Curculionidae: Scolytinae). Fitosanidad 13.
Vázquez, L.L., Matienzo, Y., Veitia, M., Alfonso, J., 2008. Manejo y conservación de enemigos
naturales de insectos fitófagos en los sistemas agrícolas de Cuba. Ed. CIDISAV, Ciudad de la
Habana.
Veitía, M., 2008. Estrategia de utilización de umbelíferas, poáceas y compositáceas para el
manejo de insectosplagas en cincohortalizas. Informe de avance de proyecto.
Venzon, M., Rosado, M.C., Euzebio, D.E., Souza, B., Schoereder, J.H., 2006. Suitability of
leguminous cover crop pollens as food source for the green lacewing Chrysoperla externa
(Hagen) (Neuroptera: Chrysopidae). Neotropical Entomology 35, 371-376.
Voronin, K.E., 1982. Biocenotic aspects of Trichogramma utilization in integrated plant
protection control. Les trichogrammes. First International symposium on Trichogramma,
Antibes, France. Les Colloques de l’INRA 9, 260-275.
Vorotynseva, A.F., 1975. The influence of nectariferous plants on contamination parasite to
codling moth and appleleaf moths. In: "Protection of the plants from pest and diseases, Cartiya
Moldovenyaske, pp. 43-46.
Wanjala, F.M.E., 1978. Relative abundance and within-canopy distribution of the parasites of the
coffee miner, Leucoptera meyricki (Lepidoptera: Lyonetiidae) in Kenya. Entomophaga 23, 57-
62.
Wang, G., Ji, X.L., Wang, W.L., Liu, Y., 2010. Community dynamics of wheat aphid natural
enemies in wheat-oilseed rape or wheat-garlic intercropping fields. Journal of Environmental
Entomology 32, 295-298.
Wang, F., Zhang, S., Hou, S., 1988. Inoculative release of Trichogramma dendrolimi in
vegetable gardens to regulate populations of cotton pests, in: Voegele, J., Waage, J., van
Lenteren, J.C. (Eds.), Trichogramma and other egg parasites. Second International Symposiom
on Trichogramma, Guangzhou, China. Les Colloques de l’INRA 43, 613-621.
Wang, H.Z., Zhao, H.L., Su, J.D., Zhang, D.X., Gao, Z.M., 1993. The ecology effects of cotton
intercropped in wheat field on cotton pests. Acta Phytophylacica Sinica 20, 163–167 (in
Chinese).
Way, M.J., 1954. Studies of the life history and ecology of the ant Oecophylla longinoda
Latreille. Bulletin of Entomological Research 45, 93-112.
Way, M.J., Heong, K.L., 1994. The role of biodiversity in the dynamics and management of
insect pests of tropical irrigated rice – a review. Bulletin of Entomological Research 84, 567-
587.
Way, M.J., Khoo, K.C., 1991. Colony dispersion and nesting habits of ants, Dolichoderus
thoracicus and Oecophylla smaragdina (Hymenoptera: Formicidae), in relation to their success
as biological control agents on cocoa. Bulletin of Entomological Research 81, 341–350.
Weyland, F., Zaccagnini, M.E., 2008. Efecto de las terrazas sobre la diversidad de artrópodos
caminadores en cultivos de soja. Ecologia Austral 18, 357-366.
Wheatley, P.E., 1963. The giant coffee looper Ascotis selenaria reciprocaría. East African
Agricultural and Forestry Journal 29, 143-146.
Wu, H.P., Meng, L., Li, B.P., 2008. Effects of feeding frequency and sugar concentrations on
lifetime reproductive success of Meteorus pulchricornis (Hymenoptera: Braconidae).
Biological Control 45, 353-359.
Wu, G.J., Chen, Z.J., Ji, M.Z., Dong, S.W., Li, H.Z., An, J.H., Shi, J.C., 1991. Influence of
interplanting corn in cotton fields on natural enemy population and its effect on pest control in
southern Shaanxi. Chinese Journal of Biological Control 7, 101-104.
Wu, Y.H., Cai, Q.N., Lin, C.W., Chen, Y.B., Li, Y.Y., Cheng, X., 2009. Responses of ground-
dwelling spiders to four hedgerow species on sloped agricultural fields in Southwest China.
Progress in Natural Science 19, 337-346.
Wyckhuys, K.A.G., O’Neil, R.J., 2007b. Influence of extra-field characteristics to abundance of
key natural enemies of Spodoptera frugiperda Smith (Lepidoptera: Noctuidae) in subsistence
maize production. International Journal of Pest Management 53, 89-99.
Xia, J.Y., 1997. Biological control of cotton aphid (Aphis gossypii Glover) in cotton (inter-
)cropping systems in China: a simulation study. Thesis Landbouwuniversiteit Wageningen, The
Netherlands.
Xing, J.M., Feng, S.L., Fan, X.H., Li, B.L., 1991. The application of Bt emulsion against cotton
bollworm and its preserving effect for natural enemies to suppress summer cotton aphid. Acta
Phytophylacica Sinica 18, 206-209.
Yabar, E., Castro, E., Melo, L., Gianoli, E., 2006. Predación de Bembidion sp., Notiobia
peruviana (Dejean) y Metius sp. (Coleóptera: Carabidae) sobre huevos de Premnotrypes
latithorax (Pierce) (Coleóptera: Curculionidae) en condiciones de laboratorio. Revista Peruana
de Entomologia 45, 91-94.
Yaku, A., 1992. Effects of intercropping sweet potato on the population density of sweet potato
weevil, Cylas formicarius (F.) (Coleoptera: Curculionidae). MSc Thesis, Department of
Entomology, McGill University, Canada.
Yan Y.H., Duan, J.J., 1988. The effect of cover cropping in apple orchards on the predator
community on the apple tree. Acta Phytophylacica Sinica 15, 23-27.
Yan, Y.H., Yu, Y., Du, X.G., Zhao, B.G., 1997. Conservation and augmentation of natural
enemies in pest management of Chinese apple orchards. Agriculture, Ecosystems and
Environment 62, 253-260.
Yardim, E.N., Edwards, C.A., 2002. Effects of weed control practices on surface-dwelling
arthropod predators in tomato agroecosystems. Phytoparasitica 30, 379-386.
Ye, H.X., Cui, L., He., X.M., Han, B.Y., 2010. Effect of intercropping tea with citrus, waxberry,
or snake gourd on population density and spatial distribution of the tea green leafhopper and
araneids.Acta Ecologica Sinica 30, 6019-6026.
Yu, Y., Zhang, A.S., Yan, Y.H., 1998. Studies on the occurrence and dispersion of Orius sauteri
in relation to ground cover vegetation in apple orchard and adjacent farms. Chinese Journal of
Biological Control 14, 148-151.
Zhang, J., Zhao, B.L., Chen, X., Luo, S.M., 2009. Insect damage reduction while maintaining
rice yield in duck-rice farming compared with mono rice farming. Journal of Sustainable
Agriculture 33, 801-809.
Zhang, R.Z., Liang, G.W., Zeng, L., 2011a. Effects of rice-planting neighboring with vegetable
crops on the population dynamics of Cnaphalocrocis medinalis, planthopper, and their
predatory enemies. Chinese Journal of Ecology 30, 281-289.
Zhang, R.Z., Liang, H.B., Tian, C.Y., Zhang, G.X., 2000. Biological mechanism of controlling
cotton aphid (Homoptera: aphididae) by the marginal alfalfa zone surrounding cotton field.
Chinese Science Bulletin 45, 355-358.
Zhang Y.J., Cang, H., Zhu, Y.Y., Wu, W.H., 1990. Influence of cultivation methods on the
natural enemy populations in the cotton-barley intercropping systems in Jiangsu. Chinese
Journal of Biological Control 6, 1-4.
Zhang, Y.B., Liu, W.X., Wang, W., Wan, F.H., Li, Q., 2011b. Lifetime gains and patterns of
accumulation and mobilization of nutrients in females of the synovigenic parasitoid, Diglyphus
isaea Walker (Hymenoptera: Eulophidae), as a function of diet. Journal of Insect Physiology 57,
1045-1052.
Zhao, J.Z., Yang, Q.H., Cheo, M.T., Ma, Z.S., Li, B.L., 1991. The effects of attractive crops
interplanted in cotton field on the production and propagation of natural enemies of cotton
insect pests. Acta Phytophylacica Sinica 18, 339-343.
Zhao, J.Z., Yang, Q.H., Xie, Y.Q 1987. Effect of cotton wheat–intercropping on cotton
pests.China Cotton 14, 41–43 (in Chinese).
Zhao, X.Q., Chen, A.D., Li, X.Y., Zhao, G.H., Gong, S.X., 2011. Effect of cover crops on
populations of insect pests and natural enemies in apple orchards. Chinese Journal of Biological
Control 27, 470-478.
Zheng X.S., Yu, X.P., Lu, Z.X., Chen, J.M., Xu, H.X., 2003. Effects of different nutritional
resources on the longevity and parasitic ability of egg parasitoid Anagrus nilaparvatae. Chinese
Journal of Applied Ecology 14, 1751-1755.
Zhong, P.S., Liang, G.W., Zeng, L., 2005a. Effect of different ecological control on population
of brown planthopper and its natural enemy community. Journal of Anhui Agricultural
University 32, 428-431.
Zhong, P.S., Liang, G.W., Zeng, L., 2005b. Effect of organic fertilizer treatment on rice
planthopper population and its natural enemy community. Entomological Journal of East China
14, 136-140.
 Zhou, C.A., Zou, J.J., Huang, S.D., 1994.The effects of Ageratum interplantation in hilly citrus
orchard in Hunan province on citrus mite and insect populations. Acta Phytophylacica Sinica
21, 57-61.
Zhou, D.R., Song, Y.Y., He, K.L., Wang, Z.Y., Zheng, L., Zhang, G.Y., 1997a. Study on the
preference habitat of Trichogramma ostriniae: I. Natural population distribution and dynamics
of the parasitoid in different habitats. Chinese Journal of Biological Control 13, 1-5.
Zhou, D.R., Song, Y.Y., Wang, Z.Y., He, K.L., Zheng, L., Zhang, G.Y., 1997b. Study on the
preference habitat of Trichogramma ostriniae: II. Effect of summer corn interplanted with
creeping type mungbean on the parasitic rate. Chinese Journal of Biological Control 13, 49-52.
Zhou, H.B., Chen, J.L., Cheng, D.F., Liu, Y., Sun, J.R., 2009. Effects of wheat-pea intercropping
on the population dynamics of Sitobion avenae (Homoptera: Aphididae) and its main natural
enemies. Acta Entomologica Sinica 52, 775-782.
Zhou, H.X., Guo, J.Y., Wan, F.H., Yu, Y., 2010. Natural control of Aphelinus mali on Eriosoma
lanigerum and the parasitoid’s overwintering protection measures. Acta Phytophylacica Sinica
37, 153-158.
Zundel, C., Nagel, O., Hanna, R., Korner, F., Scheidegger, U., 2009. Environment and host-plant
genotype effects on the seasonal dynamics of a predatory mite on cassava in sub-humid tropical
Africa.Agricultural and Forest Entomology 11, 321-331.

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