Bulletin of Entomological Pest status, bio-ecology and management of
Research the false codling moth, Thaumatotibia
cambridge.org/ber leucotreta (Meyrick) (Lepidoptera: Tortricidae)
and its implication for international trade
Review M. Adom1 , K. O. Fening1,2, M. K. Billah1,3, D. D. Wilson1,3, W. Hevi4, V. A. Clottey4,
Cite this article: Adom M, Fening KO, Billah F. Ansah-Amprofi5 and A. Y. Bruce6
MK, Wilson DD, Hevi W, Clottey VA, Ansah-
Amprofi F, Bruce AY (2021). Pest status, bio- 1African Regional Postgraduate Programme in Insect Science (ARPPIS), College of Basic and Applied Sciences,
ecology and management of the false codling University of Ghana, P. O. Box LG. 68, Legon, Accra, Ghana; 2Soil and Irrigation Research Centre, School of
moth, Thaumatotibia leucotreta (Meyrick) Agriculture, College of Basic and Applied Sciences, University of Ghana, P. O. Box LG. 68, Accra, Ghana; 3The
(Lepidoptera: Tortricidae) and its implication Department of Animal Biology and Conservation Science, College of Basic and Applied Sciences, University of
for international trade. Bulletin of 4
Entomological Research 111, 17–30. https:// Ghana, P. O. Box LG. 68, Legon, Accra, Ghana; CAB International (CABI), CSIR Campus, No.6 Agostino Road,
doi.org/10.1017/S0007485320000358 Airport Residential Area P. O. Box CT. 8630, Cantonments, Accra, Ghana;
5Plant Protection and Regulatory Services
Directorate (PPRSD) of the Ministry of Food and Agriculture, P. O. Box M. 37, Accra, Ghana and and 6International
Received: 22 July 2019 Wheat and Maize Improvement Centre (CIMMYT), ICRAF house, UN Avenue, Gigiri, P.O. Box 1041-00621, Village
Revised: 26 February 2020 Market, Nairobi, Kenya
Accepted: 26 May 2020
First published online: 17 June 2020
Abstract
Keywords: The false codling moth (FCM), Thaumatotibia leucotreta (Lepidoptera: Tortricidae) is an
False codling moth; integrated pest insect pest which represents an important threat to the production and marketing of a
management; quarantine pest; Tortricid moth
wide range of agricultural crops in the African-Caribbean-Pacific (ACP) countries. The
Author for correspondence: FCM reduces not only the yield and quality of the crop but also as a quarantine insect
M. Adom, pest, restricts the trade of susceptible agricultural produce on the international market. In add-
Email: adomsons1@gmail.com ition, little research has been conducted in the ACP countries on the bio-ecology and sustain-
able management of this pest, especially on vegetables for export. Thus, action-oriented
research aimed at understanding the bio-ecology of this important pest is essential to achieve
effective management. Various management interventions against this pest have been used in
some parts of the world, especially in South Africa on citrus. Currently, farm sanitation is
regarded as the key management strategy. Exploring and improving on other interventions
such as Sterile Insect Technique, monitoring and mass trapping of male moths, augmentative
biological control, use of bio-pesticides, protected cultivation and cold treatment may help to
mitigate the expansion of FCM into other countries, especially in the European and
Mediterranean Plant Protection Organization region where it has become a regulated insect
pest since 2014. This review discussed the bio-ecology of FCM and highlighted some of the
challenges and opportunities for its effective management and its implication for international
trade, especially the export of chillies from the ACP countries into the European Union mar-
ket which requires strict phytosanitary regulations.
Introduction
The false codling moth (FCM), Thaumatotibia leucotreta (Meyrick) (Lepidoptera:
Tortricidae), was previously known as T. roerigii Zacher (EPPO, 2019) or Cryptophlebia
(= Argyroploce) leucotreta (Meyrick) (Komai, 1999). It is one of the most economically import-
ant Tortricid insect pests causing damage to a wide range of agricultural crops (Brown, 2005;
Timm, 2005). It is highly polyphagous and attacks various fruits, vegetables and cereal crops
(EPPO, 2013; Fening et al., 2016). The larva is the most destructive stage, feeding voraciously
on the pulp of the fleshy fruits, pods, seeds, bolls or cobs of the host plants which drop pre-
maturely reducing significantly the yield of the crop (Schulthess et al., 1991; Vaissayre, 1995;
Djieto-lordon et al., 2014). The yield loss caused by FCM infestation varies among crop spe-
cies, varieties and locations. An infestation of citrus in South Africa by FCM led to a yield loss
of up to 80% within 5 months (Hofmeyr, 2003). In both South Africa and Israel, more than
30% of yield loss was reported in infested macadamia farms (La Croix and Thindwa, 1986).
© The Author(s) 2020. Published by Cambridge Yield loss as much as 90% have been reported in Kenya in Capsicum spp. (Muchemi,
University Press 2015). In Cameroon, FCM was reported to attack different varieties of Capsicum spp. and
causes an average yield loss of 43.80% (Djieto-lordon et al., 2014). Up to 20% yield loss is
caused by FCM in Uganda on cotton, as well as citrus, peach and macadamia crops
(USDA, 2010). Moreover, feeding damage can lead to the development of secondary infections
by fungi or bacteria further reducing the quality of the infested produce (Newton, 1989). An
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https://doi.org/10.1017/S0007485320000358
18 M. Adom et al.
Figure 1. Distribution map of the false codling moth from CABI (https://www.cabi.org/isc/datasheet/6904#todistribution), accessed 15 February 2020.
infestation of fruits shortly before harvest constitutes a source of Though FCM was reported in many African countries, in most
post-harvest decay (Moore, 2017). of these countries, except South Africa where extensive studies
In addition to the physical damage caused, FCM has been have been conducted, the distribution of FCM on the different
categorized as an A2 quarantine pest in European and hosts and geographical areas has not been established. The knowl-
Mediterranean Plant Protection Organization (EPPO) by the edge of in-country pest distribution may help to identify pest-free
European Commission for Plant health regulation (A2 pests are areas if any, areas with low and high prevalence. This is important
locally present in the EPPO region but not widely distributed), for informed decision making and the phytosanitary measures to
thus qualifying for inclusion as a harmful organism and restrict- put in place and enforcement of regulations at the national,
ing the international trade of its host crops such as citrus, maca- regional and international level. In South Africa, the Western
damia and a number of Solanaceae crops, including capsicum Cape region was initially free of FCM until it was introduced in
into the EU (EPPO, 2013; Muchemi, 2015). Follett and Neven 1974 from stone fruit production areas due to the lack of phyto-
(2006) defined a quarantine pest as a plant pest of potential eco- sanitary measures (Honiball, 2004; Stibick, 2006). Further studies
nomic importance to an area where it is not yet present or that is and surveying should then be conducted to determine the distri-
present but not widely distributed and is officially controlled. bution and other aspects of the bio-ecology of FCM in different
There are currently regulations on the export of FCM susceptible- regions per country. Timm (2005) reported a genetic variation
crops and phytosanitary measures are being implemented in some among FCM populations in South Africa. Analysis of population
ACP countries in order to enhance the trade of these produce into genetic variation may offer an insight into dispersal and gene flow
the EU market. of FCM which are important factors to consider when designing
This review discussed some important aspects of the bio-ecology pest management strategies (Han and Caprio, 2002, Salvato et al.,
and management of the FCM to ensure produce from the already 2002; Timm, 2005).
endemic areas such as the ACP countries, are free from the pest to
meet the demands of the new EU Council Directives on FCM
(European Commission’s Implementing Directives 2017/1279 and Host plants of FCM
2019/532) and to promote international trade. The FCM is highly polyphagous with more than 70 host plant spe-
cies, within 40 plant families, including cultivated and wild species.
A list of FCM host plants and associated countries is given in tables
Distribution of the FCM
1 and 2 of appendix 1 of the “Pest Risk Analysis for T. leucotreta”
The FCM is a native insect pest in Israel and Africa. It may be done by EPPO (2013). However, not all of the listed host plants
most associated with ecological zones that are generally classified such as apple, avocado, lime and lemon have been confirmed to
as desert and xeric shrubland, tropical and subtropical grasslands, support the feeding and development of the larvae (Newton,
savannas and moist broadleaf forest (Venette et al., 2003). In 1998; Grové et al., 1999; De Jager, 2013; Moore et al., 2015).
Israel, the pest was found for the first time in 1984 on macadamia Recently reported host plants from various African countries not
nut and has a restricted distribution (EPPO, 2002). However, in mentioned in EPPO (2013) review are listed in table 1.
Africa, FCM is thought to originate in the Ethiopian region and Though the FCM is polyphagous, host susceptibility varies
is currently widely distributed and has been reported in approxi- among plant species, varieties of the same species and may
mately 40 African countries and islands (CABI, 2000; Moore, depend on host plant characteristics (Djieto-lordon et al., 2014;
2002; EPPO, 2013) (fig. 1). Moore et al., 2017; Mkiga et al., 2019). For instance, in South
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Bulletin of Entomological Research 19
Table 1. Recently reported host plants from various African countries not represents a big concern for the citrus industry in the country
mentioned in EPPO’s (2013) review list (Schwartz, 1981, Newton, 1998). In tropical Africa, the FCM is
Country Host plant species Referencesa a serious pest of cotton and maize (Reed, 1974; Schulthess
et al., 1991; Vaissayre, 1995).
Angola Capsicum sp. EUROPHYT (2019) It was hypothesized that FCM biotypes having different host
Psydium gujava species may exist (Ford, 1934; Omer-Cooper, 1939), however, stud-
ies based on molecular analysis of specimens collected in South
Burundi Capsicum sp. EUROPHYT (2015) Africa, did not find enough evidence of the existence of these bio-
Cameroon Capsicum sp. Djieton-Lordon et al. (2014) types. This suggests an extensive gene flow (genes exchange)
Annona muricata EUROPHYT (2016) among populations from different hosts within the same location
(Timm, 2005). The polyphagous status of FCM and gene exchange-
Cote d’Ivoire Solanum EUROPHYT (2018) ability among host populations are important ecological factors to
aethiopicum consider in designing management strategies for it.
Capsicum sp. EUROPHYT (2015)
Ghana Annona muricata EUROPHYT (2017) Bio-ecology of FCM
Solanum GhanaVeg (2017)
aethiopicum The FCM is adapted to a wide ecological range and undergoes a
complete metamorphosis with four distinct stages: egg, larva,
Kenya Capsicum sp. EUROPHYT (2014); Muchemi pupa and adult (Daiber, 1979a, 1979b, 1979c, 1980; De Jager,
(2015) 2013). The different life stages are described below:
Citrus sp. Mkiga et al. (2019)
Abelmoschus EUROPHYT (2015) Egg stage
esculentus
Annona sp. Eggs are randomly laid singly or in batches by the female during
the night flight (between 5 and 11pm) in the depression of the
Dianthus sp. rind of the fruit, on foliage, on smooth surfaces, on fallen fruits
Mozambique Capsicum sp. EUROPHYT (2015) and occasionally on branches (USDA, 1984; Stibick et al.,
2010). At the optimum temperature of 25°C, the female lays
Zea mays EUROPHYT (2018)
between 3 and 8 eggs per fruit and can lay up to 800 eggs during
Nigeria Capsicum sp. EUROPHYT (2015) its lifetime with the highest daily record after 3–4 days of ovipos-
Citrus sp. Onah et al. (2016) ition period depending on the host species or the diet (Newton,
1998, De Jager, 2013; Mkiga et al., 2019). Eggs take 2–22 days
Rwanda Capsicum sp. EUROPHYT (2014)
to hatch, depending on the temperature and humidity. At 25°C,
Rosa sp. EUROPHYT (2016) De Jager (2013) reported 12.42, 9.46 and 6.47 days on orange,
Sierra Leone Capsicum sp. EUROPHYT (2018) grape and pear, respectively.
Tanzania Capsicum sp. EUROPHYT (2016), Mkiga
et al. (2019) Larval stage
Citrus sp. Mkiga et al. (2019) Upon egg hatching, the neonate larvae cut through the fruit rind
Solanum and enter the fruit where it feeds on the pulp and seed of fruits.
aethiopicum The entry point of the larvae into the fruit varies with host species
Togo Capsicum sp. EUROPHYT (2017) (De Jager, 2013). The larvae are also opportunistic, entering
through wounds and cracks on the fruit as secondary infestation
Uganda Annona muricata EUROPHYT (2016)
(Stotter, 2009, De Jager, 2013). The entry point is recognizable
Solanum EUROPHYT (2019) through the presence of the frass that is left on the fruit surface,
macrocarpon the discolouration of the rind and from the clear entry hole
Zambia Zea mays EUROPHYT (2018) (Stibick, 2006; De Jager, 2013).
Zimbabwe Capsicum sp. EUROPHYT (2014) Different larval developmental periods varying from 12 to 67
days were reported by various authors. This wide variation is
aThe host plants with EUROPHYT as reference were intercepted produce infested by FCM in
EU Member States and Switzerland. mainly due to weather conditions (especially temperature) and
host fruit quality. Generally, the larval developmental period is
longer in cool conditions and in poor quality fruits (Daiber,
Africa, FCM infestation level varies among citrus varieties. 1979b; Stibick et al., 2010, De Jager, 2013). On artificial medium,
Lemons are not known to be infested at all, Valencia cultivars Daiber (1979b) reported larval development time of 46.6, 18.8
have low susceptibility, mid-season cultivars have moderate sus- and 11.6 days at 15, 20 and 25°C, respectively, whereas on a nat-
ceptibility while Navel oranges are very susceptible (Grout and ural diet, the larval development time of 31, 29 and 23 days on
Moore, 2015; Moore et al., 2015). Sweet pepper is more preferred orange, grape and pear, respectively at 25°C was reported (De
for oviposition than chilli pepper and African eggplant (Mkiga Jager, 2013). The larvae are cannibalistic and consequently rarely
et al., 2019). The preference of the FCM for host species may more than one larva develop in the fruit (Catling and
also vary with climatic conditions (Newton, 1990; Love et al., Aschenborn, 1974; Newton, 1998; Stibick, 2006).
2014). In South Africa, of the 21 cultivated host plants reported, The larval stage has five instars and by the time that the larva
Citrus spp. appears to be the most preferred by FCM which reaches maturity, the fruit might have fallen to the ground. If the
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20 M. Adom et al.
Figure 2. The life cycle of FCM, (a) (adult), (b) (eggs), (c) (larva), (d) (pupae). Photo credits: (a): Todd M. Gilligan and Marc E. Epstein, USDA APHIS PPQ; (b and d):
photo: J.H. Hofmeyr, Citrus Research Int.; (c): Marja van der Straten, NVWA Plant Protection Service, Bugwood.org.
fruit is still intact, on the branch, the last instar larva uses a silken Adult stage
thread to drop to the ground where it pupates (Schwartz, 1981;
st nd rd th th FCM adults are noctuid moths and are sexually dimorphic. TheNewton, 1998; Stibick, et al., 2010). The 1 , 2 , 3 , 4 and 5
male and female differ in overall size, wing shape and secondary
larval stage has an average head capsule width of 0.21, 0.37,
sexual characters (Gilligan et al., 2011; EPPO, 2019). Females are
0.61, 0.94, and 1.37 mm, respectively (Daiber, 1979b).
larger than males, with the hind wings lacking the semi-circular
pockets present in males. Females have a feature like a question
mark symbol and a white dot on the forewings, when in a spread
Pupal stage position. Males possess large genital tuft, pale grey colour; hind
The pupa undergoes firstly a prepupal stage which lasts for 2–27 legs with the dense brush of greyish-white hairs; inner side of
days depending on prevailing conditions and then moults into a the hind tibia with tufts of modified scales (Timm et al., 2007,
pupa (Stibick, 2006). Generally, male pupae are smaller than 2008; Muchemi, 2015; EPPO, 2019).
females with two knobs side by side in the centre of the ventral The complete life cycle of FCM (fig. 2) ranges from 30 days
side of the ninth abdominal segment (USDA–APHIS–PPQ, (under optimal conditions) to 174 days (under unfavourable condi-
1983; Komai, 1999). The pupal stage is the most sensitive of tions) and requires approximately 800-degree days. Two to ten gen-
FCM developmental cycle. The development can be influenced erations can succeed annually (Daiber, 1980; Venette et al., 2003).
by the relative humidity, temperature, soil cover and rainfall With an uninterrupted supply of host plants, FCM remains active
(Daiber, 1979c). The pupae sex ratio, 1:2 between males and throughout the year. Diapause is not found to occur in FCM during
females was recorded for wild moth population (Myburgh and field observations and laboratory experiment (Reed, 1974; De Jager,
Bass, 1969; Daiber, 1979c). 2013). The latter author attempted inducing diapause in FCM by
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Bulletin of Entomological Research 21
simulating the start of winter in the laboratory and measured four other commodities (Rosa) exported from the ACP countries into
different traits characterizing the diapause in insects, namely the EU (table 2).
water loss, fat content, metabolic rate and super-cooling point. It Although the EU has lifted the ban since 1 January 2018 for
showed that these measurements are not consistent with the direc- Ghana, some challenges are still needed to be addressed. For
tion of a priori predictions for diapause induction. instance, currently, the export of eggplant and chillies from
Most of the studies on FCM biology were done in the labora- ACP countries to the EU countries could only be possible if mea-
tory using citrus fruit as host. Future research should attempt to sures are put in place to produce and export these commodities in
determine the development of FCM on other economically such a way that the quality will meet not only the expectations of
important host crops under laboratory and field conditions. consumers but also the new EU Plant Health Directives in force
Determining life table parameters of FCM on different host spe- (Fening and Billah, 2017b). According to the EU Implementing
cies will be a valuable tool in predicting the population dynamics directive (EU 2017/1279) (see Annex IV, Part A, Point 16.6) of
of this pest and will contribute significantly in designing manage- Article 13a of Council Directive 2000/29/EC, FCM host produce
ment interventions (De Jager, 2013). could be exported only from a country, area or place recognized
as being free of FCM in accordance with relevant international
standards for phytosanitary measures. Since FCM is widely dis-
Quarantine pest status of FCM
tributed in ACP countries, this requirement may be difficult to
Pest risk analyses for FCM conducted by the EPPO demonstrated be fulfilled unless chillies are grown in a screen house (place of
clearly that this indigenous pest from ACP countries could be intro- production free from FCM and under supervision of the
duced and established in EPPO countries and the USA as a direct National Plant Protection Organization, NPPO) (Option 3). The
result of increased international trade of host produce (Bloem other alternative according to the directive is that the produce is
et al., 2007; Gilligan et al., 2011, EPPO, 2013). This is mainly due subjected to an effective cold treatment (or other effective treat-
to the fact that the host plants of the pest in its native environment ment) (Option 4), that ensures it is FCM free. As stated, other
are also grown in EPPO countries, for example, Capsicum crops in effective treatment may refer to all other insect control methods
northern Europe, whilst in southern Europe Citrus crops would be that will ultimately lead to the effective and sustainable manage-
mostly at risk (Ostojá-Starzewski et al., 2017). Moreover, though ment of FCM in order to produce FCM-free produce as well as
FCM is a tropical pest, it can survive under cold temperatures post-harvest treatments to kill immature stages of FCM (Fening
and establish under favourable conditions especially in protected and Billah, 2017b). Another provision of the new directive is
cultivation (EPPO, 2013). The most concerned EPPO countries that the NPPO of each ACP country must send advance data
with this potential invasion of FCM are UK, Netherlands, on the effectiveness of the proposed treatment method to the
Belgium, Spain, Italy, France, Sweden and Ireland. An outbreak of EU, and this data is likely to be regularly updated, for the EU
FCM was reported in a greenhouse pepper in the Netherlands in to decide on the fate of that country (Fening and Billah,
2009 (Potting and van der Straten, 2011). However, the possibility 2017b). Thus, the EU has already tasked ACP countries that
of establishment outdoors in EPPO region has not yet been proven. chose option 4 and whose dossier was approved in 2017 to sub-
In the USA, Venette et al. (2003) estimated approximately 20% of mit a revised and updated version based on the requirements of
the continental land to be suitable for FCM. a new EU Implementing Directive in March 2019 (2019/523)
Therefore, in order to mitigate its dispersion, FCM has been which takes effect from 1st September 2019 in order to continue
declared as a quarantine insect pest by the European Commission to export chillies to the EU (E-OJ, 2019; Fening and Billah,
for plant health regulation in 2014. It has been categorized as A2 2019).
quarantine pest (i.e. it is present in EPPO region but not widely dis- The guidelines for option 3 or cultivation of chillies in a screen
tributed), thus qualifying for inclusion as a harmful organism house to eliminate the FCM remains unchanged under the new
(EPPO, 2013; Muchemi, 2015). The FCM is regarded as a high pri- Directive. Thus, for Ghana and other ACP countries subscribing
ority quarantine organism in the USA and is listed as a regulated to option 4 of the guidelines, continuous research in the area of
pest in other countries or recommended to be regulated by other the surveillance of the FCM levels and thresholds on the targeted
Regional Plant Protection Organizations (RPPOs). It is also a quar- commodities, to ascertain how effective control interventions have
antine pest in Argentina, Brazil, Chile, Paraguay, Uruguay, Israel, been, is crucial to inform pest management decisions and interven-
Jordan, Japan and New Zealand (SA-DAFF, 2015). tions to adopt in order to satisfy the guidelines of the new EU
The presence of FCM in one country might have immediate implementing directives (Fening and Billah, 2017b; E-OJ, 2019).
effects on export markets and the detection of a single larva in
fruits marked for export could result in the entire consignment
Management strategies of FCM
being rejected (Moore, 2002; Hattingh, 2006). Effects on export
markets may be in the form of a ban on the export of host fruits The aim of quarantine pest management is to eliminate or sterilize
from the country where the pest becomes established. For the target pests in exported commodities to prevent their introduc-
instance, the FCM was the key pest intercepted in chillies exported tion and establishment to new areas (Follett and Neven, 2006). The
to the UK in 2014 and 2015, thereby contributing to the ban on most commonly used FCM management strategies can be cate-
chillies and other affected commodities from Ghana into the EU gized into pre-harvest and post-harvest methods (fig. 3).
in October 2015 (table 2) (EUROPHYT, 2014, 2015; Fening et al.,
2016; GhanaVeg Sector Reports, 2017). (1) Pre-harvest methods
Pest Monitoring
Phytosanitary regulations on FCM
Monitoring FCM population is important to assist in the accurate
Since the declaration of the FCM as a quarantine insect pest, it has timing of treatment application and to compare FCM activity
been one of the major pests intercepted in vegetables (chillies) and levels between seasons which enable one to gauge probable post-
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22 M. Adom et al.
Table 2. Interception of produce from African countries imported into the EU and Switzerland with FCM (2014–2019)
Number of interceptions
Country of export Host plant species 2014 2015 2016 2017 2018 2019 Total/country
Angola Capsicum sp. – – – – – 1 2
Psydium gujava – – – – – 1
Burundi Capsicum sp. – 1 – – – – 2
Capsicum chinense – 1 – – – –
Cameroon Capsicum frutescens – 6 1 1 – – 10
Solanum aethiopicum – 1 – – – –
Annona muricata – – 1 – –
Cote d’Ivoire Solanum aethiopicum – – – – 1 – 5
Capsicum frutescens – 1 1 2 – –
Ethiopia Rosa sp. – – – – – 1 1
Ghana Capsicum sp. 68 65 – 10 – 150
Capsicum annuum – – – – 1 –
Capsicum frutescens 2 – – 1 –
Capsicum chinense – 1 – – – –
Annona muricata – – – 1 – 1
Kenya Capsicum sp. 10 68 14 15 4 11 229
Capsicum annuum – – 10 1 6 1
Capsicum frutescens – 1 – 1 – –
Capsicum chinense – – – – 1 –
Rosa sp. – – – – 37 39
Annona sp. – – 1 – – –
Abelmoschus esculentus – 1 – – – –
Gypsophila sp. – – – – 5 1
Zea mays – – – – 1 –
Dianthus sp. – – – – 1 –
Mozambique Capsicum sp. – 4 4 7 3 3 27
Capsicum frutescens – – 2 – 1 –
Capsicum annuum – – 1 1 – –
Zea mays – – – 1 –
Nigeria Capsicum sp. – 1 – 1 1 9
Capsicum annuum – – – 2 – –
Zea mays – – – – 1 3
Rwanda Capsicum sp. 1 3 – 2 3 7 34
Capsicum annuum – 1 – 3 3 6
Capsicum chinense – – – – – 3
Annona muricata – – – 1 – –
Rosa sp. – – – – – 1
Sierra Leone Capsicum frutescens – – 1 – 1 – 2
South Africa Capsicum sp. – – 1 1 – – 89
Capsicum annuum – – 1 1 –
Capsicum frutescens – – – 1 2 –
(Continued )
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Bulletin of Entomological Research 23
Table 2. (Continued.)
Number of interceptions
Country of export Host plant species 2014 2015 2016 2017 2018 2019 Total/country
Citrus reticulata 1 1 2 2
Citrus paradisi 3 3 – 7 – 2
Citrus sinensis 16 14 1 2 7 15
Fortunella sp. – – – 1 1 2
Punica granatum – – – – 1 –
Othera 1 – – – – –
Swaziland Citrus paradisi 1 – – – – 2 4
Citrus sinensis – – – 1 – –
Tanzania Capsicum sp. 1 – 2 – 1 1 54
Capsicum annuum 1 – – – 1 1
Capsicum frutescens – – – – – 1
Rosa sp. – – – – 33 12
Togo Capsicum sp. – – – 1 – 3 4
Uganda Capsicum sp. 40 64 40 30 17 7 385
Capsicum annuum 2 11 22 21 16 13
Capsicum chinense – 1 4 9 2 6
Capsicum frutescens 18 3 7 5 5 1
Rosa sp. 1 – 1 – 9 19
Annona muricata – – 1 – 4 5
Solanum macrocarpon – – – – – 1
Zambia Capsicum sp. – 1 4 1 – – 18
Capsicum annuum – – 1 2 – –
Rosa sp. – – – – 4 4
Zea mays – – – – 1 *
Zimbabwe Capsicum sp. 1 7 12 7 1 3 67
Capsicum annuum – – 1 3 – 1
Capsicum frutescens 1 – – – – –
Rosa sp. – – – – 13 –
Citrus sinensis 2 1 1 9 – 4
Total 170 260 136 138 203 185 1092
– No report.
aUnidentified host plant.
The bold values represent total interceptions per country.
Source: EUROPYT (2014–2019).
harvest risk (Moore, 2017). Monitoring is usually done by setting The monitoring can also be done by determining the presence
traps baited with female synthetic sex pheromone to trap males on of larvae in fallen fruits (Fening and Billah, 2019).
the farm. The pheromone is a blend of (E) and (Z)-8-dodecenyl
acetate (Persoons et al., 1976, 1977, Newton et al., 1993; Venette Farm sanitation
et al., 2003). The efficacy of the pheromone to attract FCM male It is known that poor farm sanitation provides breeding sites and
depends on the content of each component, (E) and harbouring places and promotes the development of FCM infest-
(Z)-8-dodecenyl acetate. It is most effective at a ratio between ation in chillies and eggplant in the field (GhanaVeg Sector
70:30 and 30:70 (E: Z) (Persoons et al., 1976, 1977; Angelini, Report, 2017; Fening and Billah, 2018). Therefore, regular removal
1979; Angelini et al., 1981; Bourdouxhe, 1982). Currently, different and destruction of all fallen and hanging fruits which are infested,
synthetic pheromone such as FCM PheroLure®, Chempac FCM damaged or is decaying and regular weeding will contribute signifi-
Lure®, P250-lure® and CRYPTACK® are commercially available. cantly to reducing FCM populations. Farm sanitation represents
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24 M. Adom et al.
Figure 3. Summary of the management strategies for the false codling moth.
Source: Myburgh (1963, 1965); Moore and Kirkman (2008); EPPO (2013); Moore and Hattingh (2016); Fening and Billah, (2019).
the cornerstone of integrated management of FCM without which susceptible to attack by FCM (Erichsen and Schoeman, 1992). In
the efficacy of all treatments directed against FCM will be severely Cameroon, Djieto-lordon et al. (2014) found that yellow pepper is
compromised (EPPO, 2013; GhanaVeg Sector Report, 2017). Farm more susceptible to FCM infestation than the red pepper and
sanitation can improve pesticide efficacy in chillies and eggplant sweet pepper. Moreover, breeding for new resistant varieties to
farms and reduced infestation levels up to 75% in the citrus FCM will contribute to mitigating the dispersal of the pest.
orchard (Moore and Kirkman, 2008; GhanaVeg Sector Report,
2017). However, for farm sanitation to be effective, it is imperative Protected cultivation
to be initiated early before the larvae emerge in infested fruit to Protected cultivation is among the strategies used in pest manage-
complete their life cycle and will contribute towards a sizeable sub- ment (Fening and Billah, 2017b). It consists of protecting culti-
sequent generation (Moore and Kirkman, 2008). In South Africa, vated crops using a physical barrier to exclude the FCM,
weekly orchard sanitation is recommended for effective control preventing, therefore, the moth from reaching the crop. This
of the FCM in orchards. In the warmer regions and periods, larvae can be done by using a mesh, exclusion net, greenhouse or net
would develop more quickly, and sanitation can be conducted house. However, this strategy is not widely adopted in the control
more regularly than once a week (Moore and Kirkman, 2008). of the FCM in Africa (Fening and Billah, 2017b). Some countries
The collected fruits should be disposed of properly. Different such as Israel and Kenya are using this method with additional
ways of disposing them of were reviewed by Moore and Kirkman measures to protect Capsicum spp., against FCM (EPPO, 2013).
(2008). In Ghana, it is recommended to destroy infested fruit by The NPPO of Ghana known as the Plant Protection and
putting them in thick black polythene or plastic bags and exposing Regulatory Services Directorate (PPRSD) of the Ministry of
them to the sun for 10–14 days or burying them to a depth of 60– Food and Agriculture (MoFA), stated in its roadmap to the EU
90 cm (Fening et al., 2016; GhanaVeg Sector Report, 2017). the intention of exploring the potential for using protected culti-
Although farm sanitation is effective for the control of FCM, it vation soon as part of the management strategy for FCM in vege-
may have an adverse effect on the parasitoids which are their nat- table production (Fening and Billah, 2017b). So, preliminary
ural enemies. Therefore, a reduction in farm sanitation when research and data are needed to serve as a baseline for further
parasitoid numbers are low, in order to aid the build-up of the studies to actualize this goal by Ghana’s NPPO.
parasitoid population is necessary. However, this might be coun-
terproductive and parasitoid survival should be considered of sec- Biological control
ondary importance to proper farm sanitation (Keeton, 2008). There are a number of natural enemies which contribute to redu-
cing FCM population in the field (Grout and Moore, 2015). The
Use of resistant host plants egg parasitoid Trichogrammatoidea cryptophlebiae (Hymenoptera:
Where available, resistant varieties of crops can be used for plant- Trichogrammatidae) occurs naturally in the field and can be used
ing to reduce the build-up of FCM population in the field. In for augmentative control (Moore and Hattingh, 2012). When
South Africa, the level of susceptibility to FCM infestation varies undisrupted, T. cryptophlebiae naturally causes egg parasitism
significantly among citrus types and within varieties of the same ranging from 80 to 100%, leading to at least 67% reduction in
citrus type (Newton, 1990; Love et al., 2014). Among avocado cul- FCM infestation in Navel oranges in South Africa (Moore and
tivars in South Africa, Edranol, Hass and Pinkerton are the most Hattingh, 2012). The use of T. cryptophlebiae as an augmentative
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Bulletin of Entomological Research 25
biological agent has not been used extensively in Africa probably (Hofmeyr and Hofmeyr, 2002; Moore and Kirkman, 2011;
due to the cost involved in mass rearing of the parasitoids as a Moore and Hattingh, 2012).
high number of parasitoids are required. Four to five monthly ‘Last Call FCM’ (Insect Science, South Africa), is an attract and
releases of 25,000 parasitoids per hectare in citrus are necessary kill product registered for the control of FCM in citrus in South
for effective control of FCM (Moore and Richards, 2001; Africa. (Hofmeyr, 2003; Kirkman, 2007). Mating disruption,
Hofmeyr, 2003). The augmentative release of T. cryptophlebiae attract and kill are negatively density-dependent approaches.
needs also to be evaluated in other climatic conditions and Therefore, for the control to be effective, they should be initiated
crops especially in the countries where FCM represents a threat. early in the season before a peak in moth activity occurs (Moore,
Some larval parasitoids including the braconids Apanteles 2017). In Ghana, the lures P-250® and CRYPTACK® have been
sp. (Hymenoptera: Braconidae) and Agathis bishopi (Nixon) used by farmers to mass trap the FCM male moths in chilli
(Hymenoptera: Braconidae), and the Ichneumonid Apophua farms (Fening and Billah, 2019).
leucotreta (Wilkinson) (Hymenoptera: Ichneumonidae) were In Ghana, the Delta trap embedded with a card with a sticky
reported to parasitize FCM (Prinsloo, 1984). Orius bugs, assassin surface and the FCM lure were used to mass trap and kill the
bugs and ants prey on FCM eggs, larvae and pupae, respectively male moths as a population suppression tool (Fening et al.,
(Moore, 2017). Agathis bishopi can cause larval parasitism to 2016; GhanaVeg Sector Report, 2017; Fening and Billah, 2017a,
rate up to 38% in Citrus in South Africa (Sishuba, 2003). 2019). This method is relatively safe as it does not require a toxi-
But the efficacy of most of these parasitoids and predators cant or an insecticide to kill the pest.
has not yet been proven and are not commercially available. In Ghana’s NPPO roadmap for the management of the FCM,
Investigations need to be done to determine the effectiveness of it was recommended that the farmer should set up FCM phero-
those parasitoids in suppressing FCM populations and assess mone traps at the beginning of the season (within the crop and
the option of their release through augmentative biological on the borders), just before fruit set, to establish the presence
control programmes. and build-up of adults (males) in traps (Fening and Billah,
Cryptophlebia leucotreta granulovirus (CrleGV) has been used 2017b).
as an entomopathogenic agent for the control of FCM. There are
three CrleGV-based products on the market: Cryptogran®, Sterile Insect Technique (SIT)
Cryptex® and Gratham® (Moore, 2017). The application of SIT is an area-wide pest management strategy. It is being devel-
CrleGV in citrus reduced the fruit infestation level up to 92% oped in some regions in South Africa since 2002 for the control
(Moore et al., 2015). In citrus orchards in South Africa, virus for- of FCM (Boersma et al., 2018). It is an effective strategy which
mulations are recommended to be applied shortly before FCM can reduce infestation by 95.2%, as a stand-alone treatment
egg hatching period which would occur 1–2 weeks at the peak (Hofmeyr et al., 2016a). The response of FCM to radiation for
in trap catches (Moore, 2017). adult sterilization is dose-dependent. Total sterility (100%) can
The potential of the use of some other entomopathogenic be achieved in irradiated females for both adults and pupae at
agents such as nematode (Heterorhabditis bacteriophora), and 200 Gy (Bloem et al., 2003). In-field flight ability of released
fungi (Beauveria bassiana and Metarhizium anisopliae) for con- FCM moth is sensitive to changes in temperature. Therefore,
trol of the pupal stage in the soil was also reported (Moore the release time of irradiated moths should target optimal
et al., 2013; Coombes et al., 2016; Steyn et al., 2017). Both nema- temperature to allow appropriate dispersal which is an important
todes and fungi have been shown to reduce T. leucotreta infest- factor for a successful release (Stotter and Terblanche, 2009; De
ation of fruit by 80% or more for the full duration of the Jager, 2013).
season, with a single application (Moore et al., 2013; Coombes Though the SIT technique is effectively used in South Africa,
et al., 2016). In Ghana, some preliminary studies on the efficacy further research is needed to improve on production, handling,
of biological insecticide Bacillus thuringiensis (Ecopel® and processing, transport, and release protocols to ensure the delivery
Bypel 1®) were effective against FCM larvae in chillies and egg- of high-quality sterile moths while considering the cost-
plant (GhanaVeg Sector Reports, 2017). effectiveness of the whole process (De Jager, 2013; Boersma and
The aqueous neem kernel extract (ANKE) (Azadirachtin) Carpenter, 2016).
applied at 50 g L−1 of water offered 100% protection of chilli
and eggplant fruits against FCM infestation in the field Chemical control
(GhanaVeg Sector Reports, 2017). Other Botanicals (plant The effectiveness of chemical control of the destructive larval
extracts and essential oils) may also be effective for the control stage of the FCM is reduced due to the protection the larva
of FCM, especially in vegetable production. However, there is a gains by living within the fruit or boll of the attacked host. This
need to determine the optimum application rates for these natural leads to relying on systemic insecticides for effective control.
insecticides against FCM. Using botanicals may also be a way to Different groups of insecticides are used to control FCM in citrus
reduce the application of synthetic insecticides and may aid in orchards in South Africa. The insect growth regulators and ovi-
the management of insecticide resistance development in FCM. cides Triflumuron (Alsystin®) and Teflubenzuron (Nomol®) are
Mating disruption, attract and kill technology chitin synthesis inhibitors of benzoylurea group. Novel insecticide
Pheromones have been much used for mating disruption and molecules chlorantraniliprole (Coragen®), an anthranilic diamide
mass trapping of this species and constitute one of the most tar- of ryanodine group and Methoxyfenozide (Runner® and
geted control methods (Fening et al., 2016; GhanaVeg Sector Walker®) of Diacylhydrazine group are effective against egg and
report, 2017; Fening and Billah, 2017a). Some mating disruption larvae of FCM (Newton, 1989; Moore, 2017). Cypermethrin is
products for FCM male mass trapping such as Isomate®, of the pyrethroid group and has a larvicidal effect on FCM.
Checkmate FCM®, Splat-FCM® and X-Mate FCM® are commer- Spinetoram (Delegate®) of spinosyn group is active across multiple
cially available. Isomate® was reported to reduce pest pressure in insect growth stages. Other insecticides have been recommended
the range of 55–75% during a period of 5 months in citrus for the control of FCM as well: Azinphos methyl (Guthion®
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26 M. Adom et al.
Solupak® 50%) Methomyl and Cypermethrin 200 EC are recom- demonstrated that at 100 Gy, moth emerged from irradiated fruits
mended in South Africa (Sweet et al., 1983). USDA–APHIS– were infertile and are not able to fly precluding the prospect of
PPQ (1983) recommended application of Azinphos methyl dispersal and mate location under field conditions. This finding
(Gusathion®, Guthion®), Diazinon and Fenvalerate to eradicate validates, therefore, the probit-9 level efficacy of 100 Gy of ioniz-
the false codling moth in the USA. ing radiation for phytosanitary treatment.
In Ghana, the binary insecticides Acetamiprid 16 g l−1 + Though ionising radiation is seen as an alternative treatment, it
Indoxacarb 30 g l−1 (Viper®) and Lambda cyhalothrin 15 g l−1 + also has a dose-dependent negative effect on fruit quality which
Acetamiprid 20 g L−1 EC (Protocol®) gave a 100% protection can, therefore, be avoided with enough reduction in dose. For
to the chilli fruits against FCM, while Dimethoate (400 g l−1) + each produce, there is a need to determine the appropriate treat-
Cypermethrin (36 g l−1) (Cydim Super®), Bacillus Thuringiensis ment dose to preserve its quality. Moreover, the potential of the
(Ecopel®) and Maltodextrin (Eradicoat T GH®) offered 71.2, 85.8 use of cold treatment associated with ionizing radiation should be
and 85.8% protection, respectively, in chilli. (Fening et al., 2016; explored. However, it is worth remembering ionizing radiation is
GhanaVeg Sector Reports, 2017; Fening and Billah, 2017b). not universally accepted and this is slowing its adoption (Follett
Recently FCM has developed resistance to some insecticides in and Neven, 2006).
South Africa, principally benzylureas (Hofmeyr and Pringle,
1998). Rational use of synthetic insecticides by alternating differ- Fumigation
ent classes will minimize the possibility of the pests becoming The potential of fumigation in FCM disinfestation is less
resistant (Fening et al., 2016; GhanaVeg Sector Reports, 2017). explored. Nevertheless, some have reported the efficacy of
methyl bromide and ethylene dibromide in fumigation of T. leu-
(2) Post-harvest methods cotreta in citrus fruit. Complete disinfestation was achieved
when ethylene dibromide fumigation was followed by exposure
Cold treatment to 4.4°C for 18 days or 11.1°C for 21 days (Myburgh, 1963;
Post-harvest cold treatment of infested fruits is one of the import- Schwartz and Milne, 1972; Schwartz and Kok, 1976).
ant measures recommended by the Food and Veterinary Office However, in 1992, methyl bromide was recognized as an ozone-
(FVO) of EU for phytosanitary risk mitigation of FCM because depleting substance under the Montreal protocol and recently
it can reduce the risk to an acceptable level on its own (EPPO, their use for quarantine purposes is restricted (Fields and
2013). Cold treatment can be compulsory for some export mar- White, 2002; EPPO, 2013).
kets irrespective of the efficacy of other control measures taken Integrated Pest Management (IPM) of FCM
(Hofmeyr and Hofmeyr, 2005; SA-DAFF, 2015). According to Moore (2017), no single control measure is cur-
The current cold treatment methods for FCM disinfestation rently available that will suppress FCM population satisfactorily.
were initially developed by Myburgh (1963, 1965) with citrus. For instance, Bloem et al. (2003) suggested the combination of
He recommended a post-harvest treatment protocol of 21-day the SIT and biological control by releasing the egg parasitoid
exposure to – 0.55°C which would provide at least a probit 9 T. cryptophlebiae can provide pest control that is more effective
level of control. This protocol has been adopted by many than when either tactic is employed separately. As a result, a
EPPO countries and others such as the USA, China and Korea multidisciplinary approach (system approach) with integrated
(Hofmeyr and Hofmeyr, 2005; USDA, 2005). Recently, Moore strategies is needed for effective and sustainable management of
et al. (2017) demonstrated that as the temperature increases, FCM. Rational integration of various strategies (fig. 3) including
the duration required for effective treatment increases. proper farm sanitation particularly may lead to effective control
Moore and Hattingh (2016) argued that cold treatment of of FCM and minimize the possibility of insecticide resistance
fruits would not be feasible since cold temperatures for a pro- development in the pest. The control of FCM using an integrated
tracted duration are known to be damaging to fruit quality, par- strategy in the field can be highly effective (Carpenter et al., 2007;
ticularly certain mandarin, orange, lemon and grapefruit Moore and Hattingh, 2012).
cultivars (Lafuente et al., 2003; Lafuente and Zacarias, 2006; In South Africa, the IPM of FCM using SIT as the main
Cronjé, 2007; Wettergreen, 2015). Therefore, the mandatory component has succeeded in reducing pest abundance by 97%
cold disinfestation required for some export markets could not in the Western Cape Province. In Ghana, the control of FCM
be applied to all fruits. Indeed, many fruits such as peppers do as outlined by NPPO’s Roadmap to address important amend-
not tolerate cold treatment as they are sensitive to chilling injury ment to EU plant health regulations employs a holistic and sus-
when stored below 7°C and symptoms can appear after a few tainable approach to crop health known as Integrated Crop
days at 0°C (EPPO, 2013). In regard to phytosanitary risk miti- Management (ICM) of which IPM is an integral part. This
gation of FCM, specific treatments involving a system approach approach involves the use of the best agronomic practices in
would need to be developed and implemented (EPPO, 2013). terms of pest and plant nutrient management and follows
Also, the possibility of cold disinfestation of harvested chilli three steps:
fruits and other produce needs to be deeply explored through
active research.
• Prevention: this avoids the introduction and build-up of a
Ionizing radiation pest population in the farm by selecting production sites in
The potential use of ionizing radiation treatment for post-harvest areas with low FCM prevalence, healthy seed, adoption of
phytosanitary disinfestation of FCM has risen as an alternative proper farm sanitation, crop rotation and best agronomic
and/or complementary method to cold treatment due to increas- practices.
ing chilling injury concern. It was established that treated larvae at • Monitoring: FCM lures Delta sticky trap is set at the beginning
doses of 200–400 Gy could not develop into moths (Barry et al., of the season (within the crop and on the borders) to establish
2007, Hofmeyr et al., 2016b). Hofmeyr et al. (2016b) the presence and build-up of adults (males) in traps.
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Bulletin of Entomological Research 27
• Acting: Use the most effective management methods and strat- Bourdouxhe L (1982) Résultats de deux années de piégeage sexuel de
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trapping of Cryptophlebia Leucotreta Meyr. in Senegal]. FAO Plant protec-
tion Bulletin 30, 125–129.
Conclusion Brown JW (2005) World Catalogue of Insects. Vol 5. Tortricidae (Lepidoptera).
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Successful IPM is based on a sound understanding of the pest’s
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Djieto-lordon C, Heumou CR, Elono PS, Alene CD, Ngueng A and
Acknowledgement. The authors acknowledge the financial and logistics sup-
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port from GhanaVeg (now Hortifresh), CABI, GIZ, USAID, Ghana Association
L.1753 (Solanaceae) in a cultivation cycle in Yaoundé. International
of Vegetable Exporters (GAVEX) and PPRSD in undertaking the initial studies
Journal of Biological and Chemical Sciences 8, 621–632.
which provided the baseline information for the development of the roadmap
E-0J: Official Journal of European Union (2019) Commission Implementing
for the management FCM on chillies for export in Ghana.
Directive (EU) 2019/523 of 21 March 2019. Amending Annexes I to V to
Council Directive 2000/29/EC on protective measures against the introduc-
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