A Safe Haven or a Temporary Alternative Host? - The Displaced Mango Fruit 
Fly, Ceratitis cosyra in the African Peach Plant
M. K. Billah1*, G. M. Oyinkah2, B. K. Badii3 and M. A. Cobblah1 
 
1  Department of Animal Biology and Conservation Science, University of Ghana, P.O. Box 
   LG 67, Legon-Accra, Ghana.
2  African Regional Post Graduate Programme in Insect Science (ARPPIS), University Of 
   Ghana, Legon.
3  Department of Agriculture, University for Development Studies, Nyankpala Campus, Ghana
*Corresponding Author: mxbillah@gmail.com 
Abstract
One of the difficulties in controlling fruit flies in cultivated crops is the use of alternative host plants as refugia 
when the preferred hosts are not in season. This study was aimed at collecting fruits and vegetables in localities 
across the five northern regions of Ghana (Northern, North-East, Savannah, Upper-East, and Upper-West 
regions) to catalogue the diversity of fruit flies and their host plants. A total of 1,722 fruits from all localities 
across the five regions were incubated, with 29.13% turning out to be fly-positive, yielding 1,141 individuals 
in four genera (Bactrocera, Ceratitis, Dacus, and Zeugodacus) and four species (Bactrocera dorsalis 
(Hendel), Ceratitis cosyra (Walker), Dacus bivittatus (Bigot), and Zeugodacus cucurbitae (Coquillet)). The 
African Peach plant, Nauclea latifolia, showed the highest incidence level of infestation, with the Mango 
fruit fly, Ceratitis cosyra as the dominant species, accounting for 97.19% (974) of the flies. The Oriental 
fruit fly, Bactrocera dorsalis and the Melon fly, Zeugodacus curcurbitae accounted for 1.23% (14 each), 
and Dacus bivittatus 0.35% (4). With evidence of displacement of C. cosyra from mango by the invasive 
Bactrocera dorsalis in most African countries, our results point to a plant that has hitherto not been known to 
be associated with fruit flies in Ghana for the displaced Mango fruit fly. Since information of previous fruit fly 
records is scanty, especially in the northern parts of the country, it is not known whether the African Peach has 
always been a host plant to C. cosyra, and served as a suitable alternative host during the long dry season, or 
is pointing to the new home after its displacement by Bactrocera dorsalis. There is therefore the need for an 
extended all-year-round collection to ascertain the host status and pattern of utilization of the African peach, 
as well as confirm the suspected host shift and displacement status of C. cosyra.
Keywords: Ceratitis cosyra; Bactrocera dorsalis; displacement; host shift; alternative host; African peach, 
Nauclea latifolia
Introduction of lucrative market opportunities results from 
imposition of strict quarantine regulations 
Fruits flies (Diptera: Tephritidae) cause by importing countries to prevent entry and 
enormous losses through direct damage establishment of fruit flies. This situation 
to fruits and vegetables. Larvae that feed is further aggravated by the introduction of 
and develop within the fruit cause the most the invasive species Bactrocera invadens of 
damage. They also introduce bacteria and Asian origin into mainland Africa. Despite the 
fungi which facilitates rotting in infested fruits economic significance associated with these 
causing them to fall to the ground prematurely insects, knowledge of their host spectrum 
(Christenson and Foote, 1960; Fletcher, 1987). remains scanty especially for northern Ghana.
Losses of up to 40 % have been recorded in One of the difficulties of controlling these 
mango in East Africa and 12-50 % in Benin fruit flies is the fact that when major crops 
(Lux et al., 2003a; Vayssières et al. 2005). Loss of economic importance such as mango, 
West African Journal of Applied Ecology, vol. 31(1), 2023: 56 - 63
57                       West African Journal of Applied Ecology, vol. 31(1), 2023
cashew and shea nuts are not in season, they baobab, and neem trees. Major cultivated crops 
find refuge in alternative host plants till their include mango and cashew. There are two 
preferred host plants are in season. This study main seasons, the dry and the wet seasons. The 
was aimed at cataloguing the host ranges of wet season commences from early April and 
fruit fly species, their preferred host plants and ends in October. The dry season, characterized 
any natural enemies associated with them from by the cold and hazy harmattan weather, starts 
both cultivated and wild fruits and vegetables from early November and ends in the latter 
from the five northern regions of Ghana. The part of March when the hot weather begins, 
information will help to plan effective fruit with intensity and ends only with the onset 
fly management strategies. Knowledge of of the early rainfall in April. Temperatures 
associated biological control agents will also fall to as low as 15 °C in the night during the 
be useful in understanding where they live harmattan season and as high as 40 °C in the 
during the crop off-season. day during the host season.
Fruit collection and incubation
Materials and methods Fruits were sampled at the beginning of the dry 
season in November when major cultivated 
Study area fruits such as mango, cashew and shea nuts 
Northern Ghana is located within latitudes were out of season. Fruits were sampled 
8°- 11°N and longitudes 0.5°-3°S. This randomly throughout the study period. Due 
area consists of the Northern, North-East, to the fact that this program had no definite 
Savannah, Upper-East, and Upper-West sampling interval as a result of variation in 
regions. It is bordered to the north by Burkina fruit availability during the sampling period, 
Faso, East by Togo, West by the Ivory Coast, number of fruit samples collected varied 
and in the south by Ghana’s Brong-Ahafo considerably, and only from areas where trees 
region (to the west), Bono East region (in were fruiting. The number of fruits in each 
the middle), and the Oti region (to the east) sample and the number of samples incubated 
(Figure 1). The northern regions are located depended mainly on fruit availability and 
in the guinea savannah vegetation belt. The abundance. Fruits were sampled from backyard 
vegetation consists of grass with scattered gardens, roadside, forest areas, orchards, and 
drought resistant trees such as the shea, the irrigated farmlands. Fruits from each sample 
Figure 1 Map of the five northern regions of Ghana and their boundary regions.
                      (Modified from source: https://www.ghanamissionun.org/wp-content/uploads/2020/09/Ghana_Regional_Map.png)
Billah M. K. et al:  A Safe Haven or a Temporary Alternative Host? - The Displaced Mango Fruit Fly                      58
were kept in plastic bags with labels indicating water as and when needed to prevent fruits 
sampling number, locality, sampling date and from drying up. Incubation chambers were 
fruit name, if known. In the laboratory, fruits in observed at 3-day intervals for puparia, and 
each sample were counted, weighed and each any puparia collected were counted, kept 
set of the same fruits kept in the same chamber in petri dishes lined with moistened filter 
for bulk incubation. The incubation chambers papers, and held in holding cages till fly and/
were made from plastic containers measuring or parasitoid emergence. Holding cages were 
18 cm x10 cm x 13.5 cm. Holes were made made from 3 mm thickness perspex sheets 
at the base of one of the plastic containers. A with measurements 14 cm x 15 cm x 10 cm. 
section of the cover was cut open and fitted Circular holes (8-10 cm diameter) were made 
with an organza material for ventilation on one side of each cage and fitted with a fine 
purposes. This container was then nested into netting sleeve to provide easy access to flies in 
another container provided with moistened the cage by hand. One side of each cage was 
sand (Figure 2). Prior to usage, the sand was also covered with a netting material to enhance 
sieved to remove debris and gravels, washed to ventilation. Petri dishes in cages were checked 
remove dust and heat-sterilized at 100 °C for daily for fly and/or parasitoid emergences, 
2 hours. Duplicate labels of the ones made in which were then counted and released into 
the field were affixed to incubation chambers. appropriately designated cages (Figure 3). 
Occasionally samples were sprinkled with Fly cages were provided with artificial diet of 
Figure 2 Improvised plastic containers used as incubation chambers. A = Plastic containers with holes made in the 
bottom of one (to hold fruits) and the other left intact (to hold sand). Lid of one cut and fitted with netting material 
for ventilation, B = Nested incubation chamber with fruits, C = Arrangement of labeled incubated chambers in a 
rearing room
Figure 3 Cages for holding puparia and flies from incubated fruits. A = Cage for holding petri dishes with puparia, 
B = Freshly-collected puparia in a petri dish lined with filter paper, C = Emergence of flies in a Holding cage
59                       West African Journal of Applied Ecology, vol. 31(1), 2023
hydrolysate yeast and sugar in a ratio of 1:3, standards require (IAEA, 2003, 2009).
while parasitoid cages were provided with a 
streak of pure honey on the inside of the top 
side of each cage. Two balls of cotton wool Results 
were also provided - one soaked in a 20% 
honey solution and the other in water (Ekesi From 103 samples, a total of 1,722 fruits and 
and Billah, 2007). Emerged insects were held vegetables were collected, weighing 50.98 kg, 
for at least four days (to ensure development and representing 15 species in 11 plant families 
of full adult features) before they were freeze- (Table 1). Fruit flies emergence was detected 
killed and preserved in 70% ethanol for in 30 field samples (29.13%), producing a 
subsequent identification. total of 1,141 individuals from four fruit 
fly genera (Bactrocera, Ceratitis, Dacus 
Identification of materials and Zeugodacus) and four species (Table 
Plants were identified by staff of the 1) - Bactrocera dorsalis (Hendel), Ceratitis 
Herbarium of the Plant and Environmental cosyra (Walker), Dacus bivittatus (Bigot), 
Health Department of the University of and Zeugodacus cucurbitae (Coquillet). Out 
Ghana, Legon, while fruit flies and their of the identified flies, 1,109 were C. cosyra 
natural enemies were identified using keys by representing 97.19%, 14 individuals each of 
Billah et al. (2007), with the help of a digital B. dorsalis and Z. cucurbitae - representing 
Leica EZ 4HD stereomicroscope. 1.23%, while 4 individuals of D .bivittatus 
were recorded, representing 0.35% (Table 2). 
The highest level of infestation was recorded 
Data analysis by C. cosyra reared from Nauclea latifolia. It 
Infestation indices were calculated in two was the highest in both puparia per fruit and 
different ways (i) by dividing the total number puparia per gram (Table 2). Ceratitis cosyra 
of puparia obtained in a given sample by the was recorded in four of the five northern 
number of fruits in the sample (puparia per regions (Northern, Savannah, Upper-East, 
fruit); and (ii) by dividing the total number and Upper-West) where fruits were collected, 
of puparia by the total mass (g) of fruits in with the highest average level of infestation in 
the sample (puparia per kg) as international terms of puparia per fruit recorded from the 
TABLE 1
List of sampled hosts, indicating number of samples and total weight of each sample
Host Plant Common and/or Vernacular Name Family No. of samples Weight (kg)
Mangifera indica Mango Anacardiaceae 1 0.85
Carica papaya Pawpaw Caricaceae 1 0.65
Cucumeropsis mannii White seed melon or “Egushi” Cucurbitaceae 1 0.50
Luffa aegyptiaca Vietnamese gourd or Sponge gourd Cucurbitaceae 7 6.70
Diospyros mespiliformis Jackalberry or African Ebony Ebenaceae 10 5.75
Strychnos spinosa The Spiny or Green Monkey Orange Loganiaceae 5 4.50
Ficus sur Cape fig or Broom cluster fig “Kankana” Moraceae 34 10.00
Ziziphus abyssinica - Rhamnaceae 1 0.20
Ziziphus mauritiana Jujube or Indian plum Rhamnaceae 1 0.35
Gardenia ternifolia “Kundozugo” Rubiaceae 2 0.75
Nauclea latifolia African Peach Rubiaceae 31 17.55
Blighia sapida Akee apple Sapindaceae 4 1.90
Pachystela brevipes - Sapotaceae 1 0.10
Capsicum annum Pepper Solanaceae 2 0.18
Lycoperiscum esculentum Tomato Solanaceae 2 1.00
Total 103 50.98
Billah M. K. et al:  A Safe Haven or a Temporary Alternative Host? - The Displaced Mango Fruit Fly                     60
TABLE 2
Host plants that were positive for fruit flies and levels of infestation by species
Fruit Fly Species
Family Host Plant C. B. D. Z. 
cosyra dorsalis bivittatus cucurbitae
Cucurbitaceae Luffa aegyptiaca 26 6,700 36 1.38 0.010 18 4 14
Ebenaceae Diospyros mespiliformis 579 5,250 10 0.02 0.002 8 8
Moraceae Ficus sur 512 9,650 32 0.06 0.003 31 31
Rhamnaceae Ziziphus mauritiana 121 350 24 0.20 0.070 6 6
Rubiaceae Nauclea latifolia 290 13,670 2,505 8.64 0.180 1,078
Total number of flies 1,141 1,109 14 4 14
Percentage Emergence 97.19 1.23 0.35 1.23
TABLE 3
Level of infestation of African Peach by C. cosyra from the four northern regions
District Locality No. fruits Weight (g) No. puparia Puparia per Puparia fruit per g
Northern Region
Gushiegu Gushiegu 17 750 119 7.00 0.16
Gushiegu Gushiegu 12 600 197 16.42 0.33
Yendi Yendi 9 1500 242 26.89 0.16
Yendi Puriya 8 750 54 6.75 0.07
Tolon-Kumbungu Gbrimani 4 550 125 31.25 0.23
Karaga Karaga 11 450 91 8.27 0.20
Karaga Gaa 13 200 28 2.15 0.14
Karaga Digblah 9 950 115 12.78 0.12
Total 83 5,750 971 11.70 0.17
Savannah Region
East Gonja Kpabulsi 17 1950 266 15.65 0.14
West Gonja Mole 3 100 89 29.67 0.89
West Gonja Achuburnyo 10 450 53 5.30 0.12
Central Gonja Yapei 14 650 218 15.57 0.34
Central Gonja Buipe 16 450 65 4.06 0.14
Central Gonja Buipe 7 200 7 1.00 0.04
Central Gonja Buipe 12 120 19 1.58 0.16
Central Gonja Buipe 5 350 30 6.00 0.09
Bole-Bamboi Bamboi 10 400 218 21.80 0.55
Bole-Bamboi Banda-Nkwanta 3 300 27 9.00 0.09
Bole-Bamboi Malawe 4 350 132 33.00 0.38
Bole-Bamboi Sakpa 6 150 15 2.50 0.10
Bole-Bamboi Kiape 12 250 23 1.92 0.09
Bole-Bamboi Gboddae 7 250 22 3.14 0.09
Bole-Bamboi Mankuma 21 550 148 7.05 0.27
Total 147 6,520 1,332 9.06 0.20
Upper-East Region
Bawku East Tilli 7 300 36 5.14 0.12
Bawku West Zebillah 14 250 8 0.57 0.03
Builsa Kpabense 12 250 23 1.92 0.09
Total 33 800 67 2.03 0.08
Upper-West region
Sissala East Sakai 17 250 47 2.76 0.19
Sissala West Lilixsi 10 350 88 8.80 0.25
Total 27 600 135 5.00 0.23
No. fruits
Weight (g)
No. puparia
Puparia/fruit
Puparia/g
No. flies
61                       West African Journal of Applied Ecology, vol. 31(1), 2023
Northern region (11.7 puparia/fruit), followed found in the Sudano-Guinean agro-ecological 
by the Savannah region (9.06 puparia/fruit), zone, which is abundantly available in all 
Upper-West region (5.0 puparia/fruit), and the inter-tropical Africa, and its distribution has 
least from the Upper-East region (2.03 puparia/ been confirmed in the four regions in northern 
fruit) (Table 3). Both B. dorsalis and C. cosyra Ghana. Discussions about pest invasions, new 
were recorded from two host plants belonging host records, and host ranges have always 
to two different families. Luffa aegyptiaca involved consideration of agro-ecological 
was the only host plant that recorded two fruit zones, proper sampling methodologies, and 
fly species, D. bivittatus and Z. cucurbitae. careful taxonomic considerations (Lux et al., 
2003b; Clark et al., 2005; Ekesi and Billah, 
2007; Nboyine et al., 2012; Mwatawala et al., 
Discussion and Conclusion 2006, 2009a, 2009b, 2015).
This observation should be the subject of 
Even though a comprehensive catalogue of critical investigation to identify the factors 
host plants of fruit flies does not exist in the contributing to the variability in dominance 
country, especially for the fauna of northern of C. cosyra in the different agro-ecological 
Ghana (Billah, unpublished data), studies by zones (Papadopoulos, 2014). One possible 
Oyinkah (2012), Nboyine (2012), and Badii reason for this could be the ability of C. 
et al. (2014, 2015a,b) are good enough to cosyra to use wild hosts such as the African 
form the basis of building a comprehensive peach, Nauclea latifolia Smith, False yam, 
database for the region. These findings may Icacina senegalensis Juss. and the Broom 
serve as new records of fruit fly host plants cluster figs, Ficus sur Forsk. in the northern 
in the northern regions, and add up to the regions. These important wild hosts should 
numerous discrete host record data scattered be considered in future fruit fly management 
in Ghana. Aside from Luffa aegyptiaca, from strategies in that region, considering the 
which D. bivittatus and Z. cucurbitae have fact that they were sampled around mango 
reportedly been reared, all the other host plants plantations. These plants may thus be serving 
may be new records as White and Elson- as alternative hosts for C. cosyra, particularly 
Harris (1992) did not list them as host plants. during the dry seasons (Billah & Wilson, 
This is particularly important especially for 2016). The three host plants, which are shrubs/
the recording of B. dorsalis from the African small trees of about 4 m high, are abundant in 
Ebony plant, Diospyros mespiliformis and the dry savanna zones, and could be important 
the Jujube or Indian plum plant, Ziziphus refugia for C. cosyra, should the displacement 
mauritiana. These plants belong to two trend in mango continue. It would, therefore, 
different families supporting the polyphagous be very important to sample fruit flies across 
nature of this recently introduced invasive different agro-ecological zones and compare 
species to the African continent and elsewhere infestation levels all-year-round to confirm the 
(Clark et al., 2005; Rwomushana et al., 2008; potential alternative host status of these wild 
Oyinkah, 2012; Badii et al., 2014, 2015a, plants for C. cosyra. Until these studies have 
2015b; Billah & Wilson, 2016; Rwomushana been conducted, it still remains as a preferred 
& Tanga, 2016; Tanga & Rwomushana, 2016). host plant of C. cosyra. The African peach 
With evidence of a displacement of C. cosyra therefore may be an important host plant to be 
by B. dorsalis in mango (Ekesi et al., 2009), the considered in the formulation or development 
detection and recording of the high numbers of of strategies for the management of fruit flies, 
C. cosyra in the African peach plant, Nauclea since it produced over 97 % of the displaced 
latifolia may be an indication of a new home pest. This may also indicate the crucial need 
for C .cosyra or suitable alternative host plant, for all-year-round surveys to determine the 
especially during the long dry season in Ghana. population dynamics of fruit flies associated 
The African peach plant, Nauclea latifolia is with those plants.
Billah M. K. et al:  A Safe Haven or a Temporary Alternative Host? - The Displaced Mango Fruit Fly                     62
Acknowledgement Tephritid Fruit Flies in Africa. Second 
Edition. ICIPE Science Press, Nairobi, 
The authors are grateful to the staff of the Ghana Kenya. 160 pp.
Herbarium at the Plant and Environmental Christenson, L. D and Foote, L. D. 1960. 
Health Department of the University of Biology of fruit flies. Annual Review of 
Ghana, Legon for the identification of all Entomology, 5: 171-192.
plant samples. We also thank University Clarke, A.R., Armstrong, K.F., Carmichael, 
of Ghana Graduate Fellowships through A.E., Milne, J.R., Raghu, S., Roderick, 
Carnegie Corporation of New York under the G.K. & Yeates, D.K. 2005. Invasive 
“Next Generation of Academics in Africa” for phytophagous pests arising through a 
providing funds to conduct this study. recent tropical evolutionary radiation: the 
Bactrocera dorsalis complex of fruit flies. 
Annual Review of Entomology, 50: 293–319.
References Ekesi, S and Billah, M.K. 2007. A Field 
Guide to the Management of Economically 
Badii, K.B., Billah, M.K., Afreh-Nuamah, Important Tephritid Fruit Flies In Africa. 
K. & Obeng-Ofori, D. 2014. Seasonal ICIPE Science Press, Nairobi, Kenya. 160 
phenology of Bactrocera invadens Drew, pp. 
Tsuruta, and White and Ceratitis cosyra Ekesi, S., Billah, M.K., Nderitu, P.W., Lux, 
(Walker) (Diptera: Tephritidae) in Northern S.A., Rwomushana, I. 2009. Evidence for 
Ghana. Bioscience Methods, 5(3): 1-11. competitive displacement of the mango fruit 
DOI: 10.5376/bm.2014.05.0003. fly, Ceratitis cosyra by the invasive fruit fly, 
Badii, K.B., Billah, M.K., Afreh-Nuamah, Bactrocera invadens (Diptera: Tephritidae) 
K. and Obeng-Ofori, D. 2015a. Species on mango and mechanisms contributing 
composition and host range of fruit-infesting to the displacement. Journal of Economic 
flies (Diptera: Tephritidae) in northern Entomology, 102: 981–991.
Ghana. International Journal of Tropical IAEA [International Atomic Energy 
Insect Science, 35(3): 137-151. Agency] 2003. Trapping guidelines for area-
Badii, K.B., Billah, M.K., Afreh-Nuamah, wide fruit fly programmes. IAEA, Vienna, 
K. Obeng-Ofori, D. & Nyarko, G. 2015b. 48 pp.
Review of the pest status, economic impact IAEA [International Atomic Energy Agency] 
and management of fruit-infesting fruit flies 2009. Development of bait stations for fruit 
(Diptera: Tephritidae) in Africa. African fly suppression in support of SIT. Report and 
Journal of Agricultural Research, 10:1488– recommendations of the consultants group 
1498. meeting organized by the Joint FAO/IAEA 
Billah M.K. & Wilson, D.D. 2016. Integrated Division of Nuclear Techniques in Food and 
management of fruit flies – case studies Agriculture, Mazatlán, Mexico, 30 October 
from Ghana, pp. 601-627. In: Ekesi, S., −1 November 2008.
Mohamed S.A. and De Meyer, M. (Editors). Lux, S.A., Ekesi, S., Dimbi, S., Mohamed, 
Fruit Fly Research and Development in S., Billah, M. 2003a. Mango-infesting fruit 
Africa - Towards a Sustainable Management flies in Africa: perspectives and limitations of 
Strategy to Improve Horticulture. Springer biological approaches to their management, 
International Publishing AG, Switzerland. pp 277–293. In: Neuenschwander P, 
778 pp. ISBN: 978-3-319-43224-3. Borgemeister C, Langewald J (eds) 
Billah, M.K., Mansell, M. W., De Meyer, M. Biological Control in IPM systems in Africa. 
& Goergen, G. 2007. Fruit fly Taxonomy CAB International, Wallingford, UK. ISBN: 
and Identification, pp H1-19. In: Ekesi, S & 0-85199-639-6.
Billah, M. K. (Eds.). A Field Guide to the Lux, S.A., Copeland, R.S., White, I.M., 
Management of Economically Important Manrakhan, A. & Billah, M.K. 2003b. 
63                       West African Journal of Applied Ecology, vol. 31(1), 2023
A new invasive fruit fly species from R (eds) Trapping and the detection, control 
the Bactrocera dorsalis (Hendel) group and regulation of Tephritid fruit flies: lures, 
detected in East Africa. Insect Science and area-wide programs and trade implications. 
Its Application, 23: 355–361. Springer Science + Business Media, 
Mwatawala, M.W., De Meyer, M., Makundi, Dordrecht.
R.H. and Maerere, A.P. 2006. Biodiversity Rwomushana, I., Ekesi. S., Gordon. I. & 
of fruit flies (Diptera: Tephritidae) at Ogol C.K.P.O. 2008. Host plants and host 
orchards in different agro-ecological zones preference studies for Bactrocera dorsalis 
of the Morogoro region, Tanzania. Fruits, (Diptera: Tephritidae) in Kenya, a new 
61: 321–332. invasive fruit fly species in Africa. Annals of 
Mwatawala, M.W., De Meyer, M., Makundi, the Entomological Society of America, 101: 
R.H. and Maerere, A.P. 2009a. Host range 331–340.
and distribution of fruit-infesting pestiferous Rwomushana, I. & Tanga, C.M. 2016. Fruit 
fruit flies (Diptera: Tephritidae) in selected Fly Species Composition, Distribution and 
areas of Central Tanzania. Bulletin of Host Plants with Emphasis on Mango-
Entomological Research, 99: 629–641. Infesting Species, pp. 71-106. In: Ekesi, S., 
Mwatawala, M.W., De Meyer, M., Mohamed S.A. and De Meyer, M. (Editors). 
Makundi, R.H. and Maerere, A.P. 2009b. Fruit Fly Research and Development in 
An overview of Bactrocera (Diptera: Africa - Towards a Sustainable Management 
Tephritidae) invasions and their speculated Strategy to Improve Horticulture. Springer 
dominancy over native fruit fly species in International Publishing AG, Switzerland. 
Tanzania. Journal of Entomology, 6: 18–27. 778 pp. ISBN: 978-3-319-43224-3.
Mwatawala, M.W., Mziray, H., Malebo, H. Tanga, C.M. and Rwomushana, I. 2016. 
& De Meyer, M. 2015. Guiding farmers’ Fruit Fly Species Composition, Distribution 
choice for an integrated pest management and Host Plants with Emphasis on Vegetable-
program against the invasive Bactrocera Infesting Species, pp.107-126. In: Ekesi, S., 
dorsalis Hendel (Diptera: Tephritidae) Mohamed S.A. and De Meyer, M. (Editors). 
in mango orchards in Tanzania. Crop Fruit Fly Research and Development in 
Protection, 76: 103–107. Africa - Towards a Sustainable Management 
Nboyine J.A, Billah M.K. and Afreh- Strategy to Improve Horticulture. Springer 
Nuamah K. 2012. Species range of fruit International Publishing AG, Switzerland. 
flies associated with mango from three 778 pp. ISBN: 978-3-319-43224-3.
agro-ecological zones in Ghana. Journal of Vayssières, J.-F., Goergen, G., Lokossou, 
Applied Biosciences, 52: 3696–3703. O., Dossa, P. & Akponon, C. 2005. A new 
Oyinkah, G.M. 2012. Host range, field Bactrocera species in Benin among mango 
preference and size relationship studies fruit fly (Diptera: Tephritidae) species. 
of fruit flies from Northern Ghana. M.Phil Fruits, 60: 371–377.
Thesis, University of Ghana, Legon. 91 pp. White, I. M. and. Elson-Harris, M. M. 
Papadopoulos, N.T. 2014. Fruit fly invasion: 1992. Fruit Flies of Economic Significance: 
historical, biological, economic aspects Their Identification and Bionomics. C.A.B. 
and management, pp 219–252. In: Shelly T, International, Wallingford, UK. 602 pp.
Epsky N, Jang EB, Reyes-Flores J, Vargas