Invasive Fruit Fly Pests in Africa

A diagnostic tool and information reference for the four Asian species of fruit fly (Diptera, Tephritidae) that have become accidentally established as pests in Africa, including the Indian Ocean Islands.

Marc De Meyer¹, Salah Mohamed² & Ian M. White³

¹ Entomology Section, Royal Museum for Central Africa, Tervuren, Belgium

² Horticultural Insects Research Department, Plant Protection Research Institute, Dokki, Egypt

³ Scientific Associate, Entomology Department, Natural History Museum, London, UK

NOTE: Click “thumbnail” image to enlarge; use back button to return to this page.

Sections

1. Introduction

2. How to Recognise an Introduced Asian Bactrocera sp.

3. How to Separate the Introduced Asian Bactrocera spp.

4. Invasive History of Asian Introductions

5. African Distribution of the Introduced Asian Bactrocera spp.

6. Host Plants of the Introduced Asian Bactrocera spp.

7. Displacement of Native and Introduced Fruit Flies by Asian Bactrocera spp.

8. Further Reading

9. Acknowledgements and Contacts

10. Literature References

1. Introduction

1.1. Aims

1.2. What fruit flies do

1.3. What fruit flies are found in Africa

1.4. How fruit flies are found

1.1. Aims

Fruit flies (Diptera, Tephritidae) are considered an insect group of major economic significance. Several representatives are known to attack different types of commercial and wild fruits and vegetables, causing considerable damage to agricultural crops. The African fauna comprises almost 1000 described species. More than 50 of them are of economic significance. Although most of these are species native to the African mainland or to any of the Indian Ocean islands, some were accidentally introduced from other regions, in particular from Asia. So far, four Asian species belonging to the genus Bactrocera invaded Africa. Two of these were introduced in recent years and the risk for other introductions is great. There is, therefore, an urgent need for considerable strengthening of the human and physical quarantine and monitoring infrastructures in Africa, in order to avoid any further unwanted introductions.

This website aims to provide a diagnostic tool for the identification of the Asian introductions already established in Africa. It also gives information on their host plant range and distribution, and references to studies on their biology either in their native range or outside.

The conventional approach of a diagnostic key has been avoided, partly because we are only interested in four species, and partly because non-specialists often find them difficult to use. Instead the site is arranged as a series of questions, starting by testing to see if we have a fruit fly, then could it be a Bactrocera sp., and if so could it be one of the introduced species, and if so, which one. Features are listed to allow elimination of non-target flies and these are listed in increasing order of difficulty rather than according to more conventional scientific criteria, e.g. a great many non-target flies should be eliminated by size or colour pattern, before you have to worry about what acrostichal setae are.

The fact that these four Asian Bactrocera species could establish themselves in Africa, does not exclude other additional introductions. Therefore, caution should be taken when using this website, since one has to take into account that yet another hitherto undetected taxon could have been introduced.

1.2. What fruit flies do

The term “fruit fly” is used for two distantly related groups of flies, namely the families Drosophilidae and Tephritidae. The former include the “fruit flies” of geneticists, which are in reality micro-fungi feeders that have acquired this name because of their habit of feeding on decaying fruit. The Tephritidae are sometimes called the “true fruit flies” because most species attack living plant material, and an estimated 40% of the over 5000 described species attack intact and growing fruit. Female fruit flies have an ovipositor, similar to the “sting” of a wasp, with which they puncture the skin of a healthy fruit into which the eggs are laid. Larval development is completed within the fruit (which may become rotten as a result) and the fully grown larvae then drop into the soil where they form a puparium.

1.3. What fruit flies are found in Africa?

There are about 950 species and 150 genera of fruit fly (Tephritidae) known in Africa, most of which form a natural component of Africa’s rich and varied biodiversity, in many cases attacking wild fruits and flowers. Most species which attack commercially grown fruit crops belong to just two genera, Ceratitis (95 species) and Dacus (195 species) (White & Goodger, 2009). A few species belong to other genera such as the coffee fruit flies (Trirhithrum spp.) which are close relatives of Ceratitis, or to the genus Bactrocera, which are close relatives of Dacus.

In Africa there are only 11 native species of Bactrocera, one of which is a widespread pest in southern Europe, namely the olive fruit fly (B. oleae). However, the genus Bactrocera forms a very large group in Asia, Australia and the Pacific with about 500 described species in those regions, including numerous pests. Four of these have found there way to Africa as a result of humans moving fruit and vegetables between continents, for trade or personal use.

1.4. How fruit flies are found

Aside from general collecting methods, typically the use of a sweep net, most fruit fly collecting involves either rearing or trapping.

Rearing techniques have been explained in detail elsewhere, e.g. in the Host Fruit Survey section of the Pacific Fruit Fly Web. Traps can be baited with either a protein bait, which attracts both sexes and many non-target insects, or with a chemical lure that is specific to male fruit flies. For more information on protein baits see the FAO/IAEA’s Trapping Guidelines for Area-Wide Fruit Fly Programmes.

Male lures can only be used for sampling old world tropical (Africa, Asia, Pacific, Australia) fruit flies belonging to the genera Bactrocera, Dacus and Ceratitis. No species is known to respond to more than a single lure (discounting chemical analogues), and not all species respond to the known lures. These lures are therefore to some extent taxon specific, although if they are to be used as part of the identification process care must be taken to avoid mixing the lures; the flies can detect even a trace of cross contamination, e.g. even due to the same person servicing both types of trap (photo showing careful handling of trap using rubber gloves to reduce the chance of cross contamination). The two main lures for Bactrocera and Dacus are as follows:

Cue lure attracts numerous species of Dacus and many Bactrocera spp.

Methyl eugenol attracts a wide range of Asian, Pacific and Australasian Bactrocera spp.; it does not attract any native African species of Bactrocera; it does not attract species associated with Cucurbitaceae (bar one rare exception in Asia); it does not attract any species of Dacus.

For some species, like B. latifrons, species specific lures are developed:

B. latifrons is known to be attracted by alpha-ionol, and its effectiveness can be increased by the use of synergists from cade oil (McQuate et al., 2004). Both components are placed separately in a trap (photo).

Lures are usually used in traps (photo of Steiner trap in Brisbane, Australia; photo of McPhail trap in Tanzania) but in survey work they can be applied to a leaf (photo; B. caryeae Drew & Hancock, a very close relative of B. invadens, attracted to a methyl eugenol coated leaf; southern India).

2. How to Recognise an Introduced Asian Bactrocera sp.

2.1. How to recognise fruit flies

2.2. How to recognise fruit flies of the genera Bactrocera and Dacus

2.3. How to separate Asian from native African Bactrocera and Dacus spp.

2.1. How to recognise fruit flies (Tephritidae)

All Tephritidae have a distinctive form of the wing vein called the “subcosta” or “vein Sc”; it curves forward at almost a right angle before fading out towards the anterior edge of the wing (photo). Only the family Psilidae have a similar feature but they will not be reared from fruit or attracted to the fruit fly chemical lures.
Most fruit flies are colourful with at least some patterning on the wings.
The species we are interested in are small to medium sized flies, with each wing between 4mm and 7mm long.
Females have a long telescopic ovipositor (drawing, showing the three sections of the ovipositor: ovsc = oviscape; ev ovp sh = eversible ovipositor sheath; acul = aculeus). Because the ovipositor is telescopic it may not always be extended to show the aculeus apex. If it is extended then it can be of assistance in identification and the apex shape is illustrated for each species.

2.2. How to recognise fruit flies of the genera Bactrocera and Dacus

Most fruit flies requiring identification as potential pests have either been reared or trapped. Information on the trapping method, the type of attractant or the host plant, can sometimes provide an approximate identification before proceeding to morphological study.

Generalisations:

Male lure:

Any fly responding to cue lure is almost certain to be one of the native African Dacus sp. or perhaps Bactrocera cucurbitae.
Any fly responding to methyl eugenol is either a non-native species of Bactrocera (likely to be either B. invadens or B. zonata) or a Ceratitis sp. (a few Ceratitis spp. are attracted to methyl eugenol).

Reared from commercial fruit or certain vegetables:

Any colourful fly with a wing length of between 4mm and 7mm reared from a recently harvested commercial fruit, is probably a Ceratitis sp. or a Bactrocera sp. (possibly B. invadens or B. zonata).
Any colourful fly with a wing length of between 4mm and 7mm reared from a recently harvested vegetable of the cucumber family (Cucurbitaceae), is probably a Dacus sp., or perhaps Bactrocera cucurbitae (it is also remotely possible that it could be B. invadens, B. latifrons or B. zonata, or a Ceratitis sp.).
Any colourful fly with a wing length of between 4mm and 6mm reared from a recently harvested vegetable of the tomato family (Solanaceae) is likely to be Bactrocera latifrons (or possibly B. invadens or B. zonata, or a Ceratitis sp.).

Any colourful fly with a wing length of between 4mm and 7mm collected in a protein trap or by any general insect collecting method such as sweep netting cannot be assumed to be a fruit fly without careful study of its morphology.

Morphological features:

Any fly not having all of these features can be eliminated.

Most Bactrocera and Dacus spp., including the invasive Bactrocera spp., have a wing length of between 4mm and 7mm (some other fruit flies are larger and some are smaller).
Most Bactrocera and Dacus spp., including the invasive Bactrocera spp., have a distinctive wing pattern that includes either a dark coloured (typically red-brown to black) marking along the anterior edge of the wing (photo) known as the “costal band”, or the “costal band” may be reduced to just an apical dark coloured spot (photo); some species may have additional dark coloured markings such as a crossband (photo). Note that numerous native African species of Bactrocera and Dacus have similar patterns, and that some other genera and families may also have similar features.
Most Bactrocera and Dacus spp., including the invasive Bactrocera spp., have a pair of black spots on the face, one under each antenna (photo).
All Bactrocera and Dacus spp. have a greatly elongated extension of the basal wing cell called “cell bcu” (also known as “cell cup”) (photo; disregard the markings as they vary between species). Note that most Tephritidae have an extension of cell bm but it is always shorter than in Bactrocera and Dacus spp.
All Bactrocera spp., and most Dacus spp., have another distinctive feature of the basal wing cells; the cell called “bm” is considerably deeper (higher) than “cell bcu” (also known as “cell cup”) (photo; disregard the markings as they vary between species).

2.3. How to separate Asian from native African Bactrocera and Dacus spp.

As already noted, no native African Bactrocera or Dacus sp., is known to have males attracted to methyl eugenol. This means that any specimen caught at methyl eugenol (subject to the lure being uncontaminated by cue lure and that it matches the morphological features listed above) is probably an invasive Asian species of Bactrocera. However, two of the species we are interested in are not attracted to methyl eugenol so further morphological study is required.

The following two features can be used to eliminate native species from introduced Asian species.

There are two somewhat raised areas just behind and below the wing base. Collectively these are called “lateroterga”; the upper is the “anatergite” and the lower the “katatergite” (photo). They should both be yellow; if the upper one, or both, are the same colour as the main body colour (red-brown to black) then you do not have one of the introduced Asian Bactrocera spp.
Flies have long hairs called “setae” or “bristles” which are in recognizable positions to which special names have been given. Most Bactrocera spp., including the four introduced Asian species, have “acrostichal setae”, i.e. a pair of setae near the mid-line of the thorax and just in front of the scutellum (photo). No species of Dacus has these setae.

The following can be used to further confirm the identification to this stage:

1. The introduced Asian Bactrocera spp. all have “dorsocentral setae” (photo), as do many other Dacus and Bactrocera spp.

2. The introduced Asian Bactrocera spp. all have a lateral pair of yellow stripes on the thorax and one has a medial stripe; these are called the “lateral vittae” and “medial vitta” (photo).

3. The males of the introduced Asian Bactrocera spp. all have a “pecten” (photo), as do many other Dacus and Bactrocera spp.

Some of you may be interested to know how Dacus are separated from Bactrocera spp.:

The terga (dorsal sclerites of the abdomen) are all fused into a single piece in Dacus spp.
The first two terga of all higher Diptera are fused; the remaining terga are separate giving a flexible abdomen in Bactrocera spp.

That may sound like a simple feature but in practice it is not always easy to see if the terga are fused or not, and hence the feature has not been used in the above lists.

3. How to Separate the Introduced Asian Bactrocera spp.

3.1. The names of the four introduced species

3.2. Recognition of B. cucurbitae

3.3. Recognition of B. zonata

3.4. Recognition of B. latifrons

3.5. Recognition of B. invadens

3.1. The names of the four introduced species

The formal manner in which to cite any species name is to include the name of the specialist(s) who first described it. If the species was originally described in a genus other than Bactrocera, the author name appears in parentheses, as follows:

Bactrocera cucurbitae (Coquillett)

Bactrocera invadens Drew, Tsuruta and White

Bactrocera latifrons (Hendel)

Bactrocera zonata (Saunders)

The genus Bactrocera is a very large one (over 500 species) and it is divided into groups of species called subgenera. Optionally, the subgeneric names may be cited in parentheses after the generic name, as follows:

Bactrocera (Zeugodacus) cucurbitae (Coquillett)

Bactrocera (Bactrocera) invadens Drew, Tsuruta and White

Bactrocera (Bactrocera) latifrons (Hendel)

Bactrocera (Bactrocera) zonata (Saunders)

With the exception of B. invadens, all of these species were assigned to the genus Dacus in earlier classifications, i.e. Dacus cucurbitae, Dacus latifrons and Dacus zonatus.

No synonyms are listed here as none have been used in the economic literature or key works, and none have ever been in common use.

3.2. Recognition of B. cucurbitae

Features of B. cucurbitae:

Males attracted to cue lure, and (rest applies to both sexes):
Scutum with a medial postsutural vitta (photo), and
Wing with a crossband (photo; the crossband is along a crossvein called “DM-Cu”; other species may have crossbands elsewhere).

OR ELSE, try next species.

3.3. Recognition of B. zonata

Features of B. zonata:

Males attracted to methyl eugenol, and (rest applies to both sexes)
Wing without a complete costal band (photo; reduced to a coloured mark in cell sc and an apical spot), and
Wing without a well marked anal streak (photo), and
Narrow subbasal section of cell br without microtrichia (photo).

OR ELSE, as follows:

i. Wing with a complete costal band (photo), and

ii. Wing with a well marked anal streak (photo), and

iii. Wing with microtrichia in narrow subbasal section of cell br (photo).

3.4. Recognition of B. latifrons

Features of B. latifrons:

Yellow strip on side of thorax (the “anepisternal xanthine”) very broad, often almost contacting the shoulder (“postpronotal lobe”) (photo), and
Abdomen mostly red-brown (no lateral marking on tergite IV; usually without a dark medial strip), and
Female, aculeus (piercing apical segment of ovipositor) with a pair of preapical "shoulders" [if the aculeus is not extended disregard this feature].

OR ELSE, try next species.

3.5. Recognition of B. invadens

This is a very variable species but in any large sample specimens with the following features should be found:

Males attracted to methyl eugenol, and (rest applies to both sexes, except as indicated)
Yellow strip on side of thorax (the “anepisternal xanthine”) narrow broad, always well separated from the shoulder (“postpronotal lobe”) (photo), and
Abdomen, with extensive black markings (photo):

Terga III-V with a distinct black mid-longitudinal line (photo), and
tergite III with extensive black markings extending on most of the tergite, leaving only a small area each side of the mid-line (photo), and
tergite IV with broad antero-lateral markings which is usually almost rectangular (photo), or more extensively black, and

Thorax (dorsal surface) variable from red-brown to black, often red-brown with black streaks, and
Female, aculeus (piercing apical segment of ovipositor) pointed [if the aculeus is not extended disregard this feature].

OR ELSE, refer your specimens to an appropriate specialist for further study.

Important notice: Currently, there is increasing evidence that B. invadens may be synonymous to B. dorsalis (see for example Tan et al., 2011; San Jose et al., 2013). A formal decision is expected to be taken in 2014.

4. Invasive History of Asian Introductions

4.1. Bactrocera cucurbitae

4.2. Bactrocera invadens

4.3. Bactrocera latifrons

4.4. Bactrocera zonata

4.1. Bactrocera cucurbitae

Bactrocera cucurbitae is principally an Asian species and its main hosts are species of Cucurbitaceae. In Africa it is almost certainly a very long established introduction. The first specimens in collections date from the early 1930s from the African mainland, as well as from Mauritius and Reunion islands (White, 2006; White et al, 2001), but it might have been established much earlier. It is considered an introduced species from Asia since it has no immediate relatives outside of Asia. It was also discovered in Hawaii in the 1890s (Coquillett, 1899). Although it was restricted to eastern Africa for several decades, it has recently been reported from western Africa and the Seychelles (White, 2006).

4.2. Bactrocera invadens

Bactrocera invadens was first found in Kenya in 2003 (Lux et al., 2003) and somewhat later reported from Tanzania (Mwatawala et al., 2004). Although initially thought to be a variant form of Bactrocera dorsalis, it was recognized as a distinct species (Drew et al., 2005). In that description, its native range was indicated as Sri Lanka. Since its first discovery on the African mainland, it has rapidly spread and now it is now found in several countries of eastern, central and western Africa. Recently, observations from southern Africa are also reported (e.g. Correia et al, 2008). A brief outbreak of a methyl eugenol responding species in Mauritius in 1996, attributed to B. dorsalis (White et al, 2001), may have been this species. Available samples were re-examined but the results were inconclusive (White, 2006). A review of its invasion history and biology is given by Malavasi et al. (2013).

4.3. Bactrocera latifrons

Bactrocera latifrons was only very recently detected in Africa. The first specimens were trapped early in 2006 in Morogoro, Tanzania (Mwatawala et al., 2010). Surveys have shown that this species is widely distributed in Tanzania although not present in large numbers because of its limited host range (Mwatawala et al., 2010). In 2007, it was also found in Kenya near the border with Tanzania (Ekesi, unpublished records). So far, the species has not been reported from any other African country. B. latifrons is Asian in origin, and was found in the Hawaiian Islands around 1983 (Vargas & Nishida, 1985).

4.4. Bactrocera zonata

The Peach fruit fly is native to tropical Asia, and widely distributed in SE Asia (India, Thailand, Philippines, Pakistan, Vietnam, Sri Lanka, Bangladesh, and Nepal). It also occurs on the Indian Ocean islands of Mauritius (first record in 1942, and again established since 1987) and Reunion (regularly found since 1991) (White et al., 2001). By the early 1980s, it had become established in the Arabian Peninsula including Saudi Arabia, Oman, and Yemen (White, 2006).

It invaded the Egyptian ecosystem from which it was first erroneously identified as Bactrocera pallidus (Perkins & May) (Abuel-Ela et al., 1998). B. zonata was detected in Egypt in 1914 (Efflatoun, 1924) in Port Said (Red Sea coast) but there were no further records to suggest that quarantine interception was the start of establishment (label information for the two specimens, in the collection of the Ministry of Agriculture: “Port-said customs from India, Sample No. 1036.14 V 1914”).

The first record of B. zonata being established in Egypt was in Kalubia governorate (East Cairo) in 1993 from guava samples, and later the same year in Faiuom governorate (West Cairo). Both governorates are fruit producing areas and there is a continuous presence of the plant hosts during the year. The next year, the pest was found in Alexandria governorate (Agami), where the fig is widely distributed, and in Giza governorate (West Cairo) where different horticultural trees are cultivated in home gardens. By 1995, the insect was found in further fruit producing governorates. By 1997, it was distributed in throughout Egypt, including the Dakhla and Kharga oases (west Egypt) and in Sinai (East north Egypt), both of which are desert areas.

5. African Distribution of the Introduced Asian Bactrocera spp.

The current distribution of the different introduced species is briefly discussed here. Maps shown mark the countries (not individual localities) from which each of the introduced species is reported from the Africa, including the mainland as well as the islands of the Indian Ocean and the Arabian Peninsula. Specimen records can be found in White (2006).

Subsections:

5.1. Bactrocera cucurbitae

5.2. Bactrocera invadens

5.3. Bactrocera latifrons

5.4. Bactrocera zonata

5.1. Bactrocera cucurbitae

Bactrocera cucurbitae is currently reported from several countries in East and West Africa: Benin, Burkina Faso, Cameroon, Gambia, Guinea, Ivory Coast, Mali, Niger, Nigeria, Senegal and Togo in western Africa, and Kenya, Sudan, Tanzania and Uganda in eastern Africa. It has also been reported from Kinshasa in the Democratic Republic of Congo and from the islands of Mauritius and Réunion and the Seychelles. There is a specimen in the BMNH collection from “Arabia” for which the exact location is unknown (not included in the map below). An old record of B. cucurbitae from “Lower Nile Valley, Egypt” could not be confirmed or traced as to its origin. According to the second author, who made extensive surveys in Egypt, B. cucurbitae does not occur in that country, and hence this record should be removed.

Map showing countries in Africa from which B. cucurbitae is reported (based upon records in White, 2006 and references therein, and Vayssières et al., 2008a. Record from Sudan based upon material from Senga)

5.2. Bactrocera invadens

As mentioned in section 4.2, Bactrocera invadens was first recorded from eastern Africa in 2003. In a period of two years it was found in most western, Central and eastern African countries, as well as on the islands of the Comoro archipelago (including the French Overseas Territory of Mayotte). In recent years, several findings from southern African countries (Botswana, Mozambique, Zambia, Namibia) have been reported (Correia et al., 2008; USDA-APHIS Pretoria office, unpublished reports). The species has been detected in the northern part of South Africa in recent years but has been eradicated (Manrakhan et al., 2011). Since 2010, it is also found in Madagascar (Raoilijaona et al., 2012)

Map showing countries in Africa from which B. invadens is reported (based upon records in White, 2006 and references therein; references mentioned above, and unpublished records provided by CIRAD/PRPV, RID/CNEARC, IITA, ICIPE, RMCA and USDA-APHIS)

5.3. Bactrocera latifrons

Bactrocera latifrons was only recently discovered in Africa (since 2006). So far specific surveys have been only been conducted in Tanzania and Kenya. In Tanzania the species is found in several areas of the country. Researchers at ICIPE have also found the species in southern Kenya, close to the border with Tanzania. Its occurrence in other parts of Africa is currently unknown.

Map showing countries in Africa from which B. latifrons is reported (based upon records in Mwatawala et al., 2006 and unpublished records of specimens in collection of ICIPE)

5.4. Bactrocera zonata

Bactrocera zonata occurs in two regions in Africa. Firstly, it is reported from some of the islands in the Indian Ocean (Mauritius and Réunion). Secondly it is found in northern Africa (Egypt and Libya) and from a number of countries on the Arabian Peninsula: Oman, Saudi Arabia; United Arab Emirates and Yemen. Recently it has been reported from Gezira region in Sudan (Salah et al., 2012) which indicates a southward spread and potential risk for Sub-Saharan region.

Map showing countries in Africa and Arabian Peninsula from which B. zonata is reported (based upon records in White, 2006 and references therein).

6. Host Plants of the Introduced Asian Bactrocera spp.

Most species of Tephritidae are very host specific, i.e. they can only complete their larval development in a few closely related species of plants. However, it is a feature of many of the pest species that they are generalists, able to complete their larval development in a wide range of unrelated plants, although even these can exhibit preferences.

The host data presented here is mostly based on large scale rearing studies carried out in various areas of Asia and Africa. As far as possible spurious records (based on observation of an adult resting on a plant, or the name of a tree in which a trap had been placed) have been excluded; a great many records of that type were reviewed by White & Elson-Harris (1992) and need not be mentioned again. Common names also follow the list provided in White & Elson-Harris (1992).

Subsections:

6.1. Host plants of B. cucurbitae

6.2. Host plants of B. invadens

6.3. Host plants of B. latifrons

6.4. Host plants of B. zonata

6.1. Host plants of B. cucurbitae

B. cucurbitae, in common with most other species of the Asian subgenus B. (Zeugodacus), is primarily associated with the “fruits” of the cucumber family, Cucurbitaceae, but other plant families are sometimes attacked. These include melon (Cucumis melo), cucumber (Cucumis sativus), squash (Cucurbita spp.) and a variety of gourds (e.g. Coccinia and Momordica spp.). The part of the plant attacked is usually the “fruit”; however, in common with many of its Asian relatives, it can also attack flowers sometimes even stem and root tissue. In Hawaii pumpkin and squash fields (varieties of Cucurbita pepo) have been known to be heavily attacked before fruit has even set, with eggs being laid into unopened male and female flowers, and larvae even developing successfully in the taproots, stems and leaf stalks (Back & Pemberton, 1914). Sometimes it attacks plants belonging to other families, e.g. tomato (Lycopersicon esculentum).Observations on non-cucurbit hosts in Tanzania, however, indicate that infestations in these hosts are very minor (Mwatawala et al.,2010).

Hosts in Africa:

Family: Anacardiaceae

Common name	Scientific name	Country, Reference
Cashew	Anacardium occidentale	Benin, Burkina Faso, Vayssières et al., 2007
Mango	Mangifera indica	Benin, Mali, Vayssières et al., 2008a; Tanzania, Mwatawala et al., 2010 Mauritius, Quilici & Jeuffrault, 2001

Family: Annonaceae

Common name	Scientific name	Country, Reference
Wild custard apple	Annona senegalensis	Western Africa, Vayssières et al., 2007

Family: Cucurbitaceae

Common name	Scientific name	Country, Reference
Bitter apple	Citrullus colocynthis	Mauritius & Réunion, Quilici & Jeuffrault, 2001
Water melon	Citrullus lanatus	Western Africa, Vayssières et al., 2007; Tanzania, Mwatawala et al., 2010 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Ivy gourd	Coccinia grandis	Kenya 2000, R.S. Copeland data Réunion, Quilici & Jeuffrault, 2001
Bitter cucumber	Cucumeropsis mannii	Benin, Vayssières et al., 2008
	Cucumis anguria	Réunion, Quilici & Jeuffrault, 2001
	Cucumis dipsaceus	Kenya 2000, R.S. Copeland data ; Tanzania, Mwatawala et al., 2010
	Cucumis figarei	Kenya 2000, R.S. Copeland data
Melon	Cucumis melo	Western Africa, Vayssières et al., 2007; Tanzania, Mwatawala et al., 2010 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Cucumber	Cucumis sativus	Kenya 2002; Tanzania Mwatawala et al., 2010; Western Africa, Vayssières et al., 2007 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Pumpkin	Cucurbita maxima	Western Africa, Vayssières et al., 2007 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Pumpkin	Cucurbita moschata	Mwatawala et al., 2010
Spaghetti squash	Cucurbita pepo	Western Africa, Vayssières et al., 2007 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Pumpkin	Cucurbita sp.	Kenya 1937, SANC collection data ; Tanzania, Mwatawala et al., 2010
	Cylanthera pedata	Réunion, Quilici & Jeuffrault, 2001
	Diplocyclos palmatus	Kenya 2001, R.S. Copeland data
	Kedrostis leloja	Kenya 2004, R.S. Copeland data
Bottle gourd	Lagenaria leucaritha	Mauritius & Réunion, Quilici & Jeuffrault, 2001
	Lagenaria sphaerica	Mauritius & Réunion, Quilici & Jeuffrault, 2001
Water-bottle	Lagenaria siceraria	Western Africa, Vayssières et al., 2007; Tanzania, Mwatawala et al., 2010
Smooth luffa	Luffa acutangula	Tanzania, Mwatawala et al., 2010 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Smooth luffa	Luffa cylindrica	Western Africa, Vayssières et al., 2007 Mauritius & Réunion, Quilici & Jeuffrault, 2001
Bitter gourd	Momordica charantia	Kenya 1997, MRAC collection data; Western Africa, Vayssières et al., 2007; Tanzania, Mwatawala et al., 2010 Mauritius & Réunion, Quilici & Jeuffrault, 2001
	Momordica foetida	Kenya 2001, R.S. Copeland data; Tanzania, Mwatawala et al., 2010
	Momordica trifoliata	Kenya 2001, R.S. Copeland data; Tanzania, Mwatawala et al., 2010
	Secchium edule	Mauritius & Réunion, Quilici & Jeuffrault, 2001
Snakegourd	Trichosanthes cucumerina	Mauritius & Réunion, Quilici & Jeuffrault, 2001
	Telfairia occidentalis	Ivory Coast, Vayssières et al., 2007

Family: Canellaceae

Common name	Scientific name	Country, Reference
	Warburgia ugandensis	Kenya 1949, BMNH collection data

Family: Caricaceae

Common name	Scientific name	Country, Reference
Papaya	Carica papaya	Tanzania, Mwatawala et al., 2010

Family: Oxalidaceae

Common name	Scientific name	Country, Reference
Carambola, starfruit	Averrhoa carambola	Benin, Ivory Coast, Vayssières et al., 2007

Family: Passifloraceae

Common name	Scientific name	Country, Reference
Passion fruit	Passiflora edulis	Réunion, Quilici & Jeuffrault, 2001

Family: Rutaceae

Common name	Scientific name	Country, Reference
Tangerine	Citrus reticulata	Benin, Vayssières et al., 2008a
Sweet orange	Citrus sinensis	Benin, Burkina Faso, Vayssières et al., 2007

Family: Solanaceae

Common name	Scientific name	Country, Reference
Paprika	Capsicum annuum cov. longum	Tanzania, Mwatawala et al., 2010
Chili pepper	Capsicum frutescens	Western Africa, Vayssières et al., 2007
Tomato	Lycopersicon esculentum	Tanzania, Mwatawala et al., 2010
African eggplant	Solanum aethiopicum	Tanzania, Mwatawala et al., 2010
	Solanum anguivi	Tanzania, Mwatawala et al., 2010
African eggplant	Solanum macrocarpon	Tanzania, Mwatawala et al., 2010
Black nightshade	Solanum nigrum	Tanzania, Mwatawala et al., 2010

Major hosts in Asia:

Family: Cucurbitaceae

Common name	Scientific name	Reference
Melon	Cucumis melo	Allwood et al., 1999; Drew, 1989
Pumpkin	Cucurbita maxima	Allwood et al., 1999; Tsuruta et al., 1997
Spaghetti squash	Cucurbita pepo	Allwood et al., 1999; Drew, 1989
Snakegourd	Trichosanthes cucumerina	Allwood et al., 1999; Tsuruta et al., 1997

Other hosts in Asia:

Family: Cucurbitaceae

Common name	Scientific name	Reference
	Benincasa hispida	Allwood et al., 1999 (fruit and flowers)
Colocynth	Citrullus colocynthis	White & Elson-Harris, 1992
Watermelon	Citrullus lanatus	Allwood et al., 1999
Ivy gourd	Coccinia grandis	Allwood et al., 1999; Tsuruta et al., 1997 (fruit and flowers)
Cucumber	Cucumis sativus	Allwood et al., 1999; Drew, 1989; Tsuruta et al., 1997
	Cucumis trigonus	White & Elson-Harris, 1992
Pumpkin	Cucurbita moschata	Allwood et al., 1999 (fruit and flowers)
	Diplocyclos palmatus	Tsuruta et al., 1997
	Gymnopetalum integrifolium	Allwood et al., 1999
White-flowered gourd	Lagenaria siceraria	Allwood et al., 1999; Tsuruta et al., 1997
Angled luffa	Luffa acutangula	Allwood et al., 1999; Tsuruta et al., 1997
Luffa	Luffa aegyptiaca	Allwood et al., 1999 (fruit and flowers)
	Melothria wallichii	Allwood et al., 1999
Balsam-apple	Momordica balsamina	White & Elson-Harris, 1992
Bitter gourd	Momordica charantia	Allwood et al., 1999; Drew, 1989; Tsuruta et al., 1997
Teruah	Momordica cochinchinensis	White & Elson-Harris, 1992
	Momordica dioica	Ranganath & Veenakumari, 1995
	Mukia maderaspatana	Ranganath & Veenakumari, 1995
	Trichosanthes ovigera	Allwood et al., 1999
	Trichosanthes tricuspidata	Allwood et al., 1999
	Trichosanthes wallichiana	Allwood et al., 1999
	Trichosanthes wawraei	Allwood et al., 1999

Family: Agavaceae

Common name	Scientific name	Reference
	Dracaena curtissi	Allwood et al., 1999

Family: Capparidaceae

Common name	Scientific name	Reference
	Capparis sepiaria	Allwood et al., 1999
	Capparis thorellii	Allwood et al., 1999
	Maerua siamensis	Allwood et al., 1999

Family: Caricaceae

Common name	Scientific name	Reference
Papaya	Carica papaya	Tsuruta et al., 1997

Family : Fabaceae

Common name	Scientific name	Reference
Garden bean	Phaseolus vulgaris	Allwood et al., 1999
	Vigna sinensis	Allwood et al., 1999
Southern pea	Vigna unguiculata	Allwood et al., 1999

Family: Loganiaceae

Common name	Scientific name	Reference
Strychnine	Strychnos nux-vomica	Tsuruta et al., 1997

Family : Malvaceae

Common name	Scientific name	Reference
	Abelmoschus moschatus	Allwood et al., 1999

Family: Moraceae

Common name	Scientific name	Reference
	Ficus chartacea	Allwood et al., 1999

Family: Myrtaceae

Common name	Scientific name	Reference
Guava	Psidium guajava	Allwood et al., 1999

Family: Pandanaceae

Common name	Scientific name	Reference
	Pandanus odoratissimus	Tsuruta et al., 1997

Family: Passifloraceae

Common name	Scientific name	Reference
Purple granadilla	Passiflora edulis	Tsuruta et al., 1997

Family : Rhamnaceae

Common name	Scientific name	Reference
Zizyphus	Ziziphus jujuba	Allwood et al., 1999

Family: Rutaceae

Common name	Scientific name	Reference
	Citrus hystrix	Allwood et al., 1999

Family: Sapotaceae

Common name	Scientific name	Reference
Sapodilla, chicle	Manilkara zapota	Allwood et al., 1999

Family: Solanaceae

Common name	Scientific name	Reference
Tomato	Lycopersicon esculentum	Allwood et al., 1999
	Solanum trilobatum	Allwood et al., 1999

Family: Vitaceae

Common name	Scientific name	Reference
	Tetrastigma lanceolarium	Allwood et al., 1999

6.2. Host plants of B. invadens

B. invadens has been recorded attacking plants belonging to several unrelated families and it may be expected to have as broad a host range as the Oriental fruit fly, B. dorsalis (Hendel). It is particularly injurious to mango (Mangifera indica) and guava (Psidium guajava). Several studies on the host plant range of B. invadens are ongoing and it is to be expected that the host list will be increase in future.

Hosts in Africa:

Family: Anacardiaceae

Common name	Scientific name	Country, Reference
Cashew	Anacardium occidentale	Benin, Goergen et al., 2011
Mango	Mangifera indica	Western Africa, Goergen et al., 2011
Mango	Mangifera indica	Tanzania 2003, MRAC collection data ; Kenya, Rwomushana et al., 2008a
Marula plum	Sclerocarya birrea	Ekesi et al., 2006 Benin, Goergen et al., 2011
	Sorindeia madagascariensis	Kenya, Rwomushana et al., 2008a
Jew plum	Spondias cytherea	Tanzania, Mwatawala et al., 2006 Benin, Goergen et al., 2011
Tropical plum	Spondias mombin	Benin, Goergen et al., 2011

Family: Annonaceae

Common name	Scientific name	Country, Reference
Cherimoya	Annona cherimola		Kenya, Rwomushana et al., 2008a; Tanzania, Mwatawala et al., 2009
	Annona diversifolia		Ivory Coast, N’Depo et al., 2010
	Annona montana		Ivory Coast, N’Depo et al., 2010
Wild custard apple	Annona senegalensis		Western Africa, Vayssières et al., 2009a,
	Annona squamosa		Kenya, Rwomushana et al., 2008a
Soursop	Annona muricata		Ekesi et al., 2006; Vayssières et al., 2009a
			Tanzania, Mwatawala et al., 2006

Family : Apocynaceae

Common name	Scientific name	Country, Reference
	Landolphia sp.	Western Africa, IITA data
Saba nut	Saba senegalensis	Cameroon, Goergen et al., 2011
Lucky nut	Thevetia peruviana	Tanzania, Mwatawala et al.,2009

Family : Boraginaceae

Common name	Scientific name	Country, Reference
	Cordia sp. cf myxa	Kenya, Rwomushana et al., 2008a

Family : Caesalpiniaceae

Common name	Scientific name	Country, Reference
Cayor pear tree	Cordyla pinnata	Western Africa, Vayssières et al., 2009a

Family : Capparaceae

Common name	Scientific name	Country, Reference
	Maerua duchesnei	Benin, Goergen et al., 2011

Family : Caricaceae

Common name	Scientific name	Country, Reference
Papaya	Carica papaya	Western Africa, Goergen et al., 2011; Tanzania, SUA data

Family : Clusiaceae

Common name	Scientific name	Country, Reference
Chewing stick	Garcinia mannii	Cameroon, Goergen et al., 2011

Family : Combretaceae

Common name	Scientific name	Country, Reference
Tropical almond	Terminalia catappa	Ekesi et al., 2006; Benin, Vayssières et al., 2008b
Tropical almond	Terminalia catappa	Tanzania 2006, MRAC/SUA data

Family : Cucurbitaceae

Common name	Scientific name	Country, Reference
Egusi	Citrullus colocynthis	Benin, Goergen et al., 2011
Watermelon	Citrullus lanatus	Tanzania, Mwatawala et al., 2006 Western Africa, Goergen et al., 2011
	Cucumis figarei	Tanzania, Mwatawala et al., 2006 based on single specimen reared
Cucumber	Cucumis sativus	Tanzania, Mwatawala et al., 2006 Western Africa, Goergen et al., 2011
Pumpkin	Cucurbita maxima	Western Africa, Goergen et al., 2011
Gourd	Cucumis pepo	Western Africa, Goergen et al., 2011
Bottle gourd	Lagenaria siceraria	Benin, Goergen et al., 2011
Bitter melon	Momordica charantia	Western Africa, Goergen et al., 2011

Family : Dracaenaceae

Common name	Scientific name	Country, Reference
	Dracaena steudneri	Kenya 2004, BMNH collection data

Family : Ebenaceae

Common name	Scientific name	Country, Reference
Japanese persimmon	Diospyros kaki	Western Africa, IITA data
Mountain persimmon	Diospyros montana	Benin, Goergen et al., 2011

Family: Flacourtiaceae

Common name	Scientific name	Country, Reference
Governor’s plum	Flacourtia indica	Tanzania, Mwatawala et al., 2006

Family: Irvingiaceae

Common name	Scientific name	Country, Reference
African wild mango	Irvingia gabonensis	Western Africa, Vayssières et al., 2009a

Family : Lauraceae

Common name	Scientific name	Country, Reference
Avocado	Persea americana	Kenya, Ekesi et al., 2006; Tanzania, Mwatawala et al., 2006
Avocado	Persea americana	Wetsern Africa, Goergen et al., 2011

Family: Moraceae

Common name	Scientific name	Country, Reference
	Ficus cf. ottoniifolia	Benin, Goergen et al., 2011

Family : Musaceae

Common name	Scientific name	Reference
Plantain	Musa x paradisiaca	Western Africa, Goergen et al., 2011
Cavendish banana	Musa acuminata	Benin, Goergen et al., 2011 East Africa, Cugala et al., 2013 (dwarf variety)
	Musa sp.	Ekesi et al., 2006

Family: Myrtaceae

Common name	Scientific name	Country, Reference
Pitanga cherry	Eugenia uniflora	Benin, Goergen et al., 2011
Jambolan	Syzygium cumini	Tanzania, Mwatawala et al. , 2009
Rose apple	Syzygium jambos	Tanzania, SUA unpublished data Benin, Goergen et al., 2011
Malay apple	Syzygium malaccense	Benin, Goergen et al., 2011
Java apple	Syzygium samarangense	Western Africa, IITA data
Strawberry guava	Psidium littorale	Tanzania, Mwatawala et al., 2009
Guava	Psidium guajava	Ekesi et al., 2006
		Benin, Goergen et al., 2011
		Cameroon 2004, MRAC collection data
		Tanzania 2004, Mwatawala et al., 2006; Kenya, Rwomushana et al., 2008a

Family: Oxalidaceae

Common name	Scientific name	Country, Reference
Carambola, starfruit	Averrhoa carambola	Western Africa, Vayssières et al., 2009a

Family Punicaceae

Common name	Scientific name	Country, Reference
Pomegranate	Punica granatum	José et al., 2013

Family: Rhamnaceae

Common name	Scientific name	Country, Reference
Jujube	Ziziphus mauritiana	Benin, Goergen et al., 2011

Family: Rosaceae

Common name	Scientific name	Country, Reference
Loquat	Eriobotrya japonica	Tanzania, Mwatawala et al., 2006 Cameroon, Goergen et al., 2011
Apple	Malus domestica	Tanzania, Mwatawala et al., 2009
Peach	Prunus persica	Tanzania, Mwatawala et al.,2006

Family: Rubiaceae

Common name	Scientific name	Country, Reference
Arabica coffee	Coffea arabica	Tanzania, SUA unpublished data
Robusta coffee	Coffea canephora	Tanzania, Mwatawala et al., 2009
African peach	Sarcocephalus latifolius	Westen Africa, Vayssièreset al., 2009a

Family: Rutaceae

Common name	Scientific name	Country, Reference
Sour orange	Citrus aurantiaum	Western Africa, IITA data
Pomelo	Citrus grandis	Tanzania, Mwatawala et al., 2009
Lemon	Citrus limon	Ekesi et al., 2006 Tanzania, Mwatawala et al., 2009 Benin, Goergen et al., 2011
Grapefruit	Citrus paradisi	Tanzania, Mwatawala et al., 2006 Western Africa, Vayssières et al., 2009a
Tangerine	Citrus reticulata	Western Africa, Goergen et al., 2011
Tangerine	Citrus reticulata	Tanzania, Mwatawala et al., 2006; Kenya, Rwomushana et al., 2008a
Sweet orange	Citrus sinensis	Western Africa, Goergen et al., 2011; Kenya, Rwomushana et al., 2008a; Tanzania, Mwatawala et al., 2009
Tangelo	Citrus tangelo	Western Africa, Vayssières et al., 2009a
Kumquat	Fortunella margarita	Tanzania, Mwatawala et al., 2010

Family: Sapindaceae

Common name	Scientific name	Country, Reference
Ackee	Blighia sapida	Benin, Goergen et al., 2011

Family: Sapotaceae

Common name	Scientific name	Country, Reference
	Achra sapota	Ivory Coast, N’Depo et al., 2010
White star-apple	Chrysophyllum albidum	Benin, Goergen et al., 2011
	Chrysophyllum Cainito	Ivory Coast, N’Depo et al., 2010
Bully tree	Manilkara zapota	Western Africa, Vayssières et al., 2009a
	Richardella campechiana	Ivory Coast, N’Depo et al., 2010
Sheanut	Vitellaria paradoxa	Western Africa, Vayssières et al., 2009a

Family: Solanaceae

Common name	Scientific name	Reference
Bell pepper	Capsicum annuum	Western Africa, Vayssières et al., 2005
Chili pepper	Capsicum frutescens	Western Africa, Vayssières et al., 2009a
Tomato	Lycopersicon esculentum	Ekesi et al., 2006; Mziray et al., 2010 Western Africa, Goergen et al., 2011
African eggplant	Solanum aethiopicum	Tanzania, Mwatawala et al., 2009
African eggplant	Solanum anguivi	Tanzania, Mziray et al., 2010
	Solanum incanum	Tanzania, Mziray et al., 2010
Black nightshade	Solanum nigrum	Tanzania, Mziray et al., 2010
Sodom apple	Solanum sodomeum	Tanzania, Mziray et al., 2010

Family: Strychnaceae

Common name	Scientific name	Country, Reference
	Strychnos mellodora	Kenya 2003, R.S. Copeland data

Hosts in Asia:

No available data.

6.3. Host plants of B. latifrons

B. latifrons is a pest of tomato family (solanaceous) crops. However, data from Hawaii, where B. latifrons is also an introduction, has confimed that it can also develop in a few species of Cucurbitaceae, but it was out-numbered by B. cucurbitae in those hosts. In Asia it has been shown that other families can also be attacked but only rarely (Allwood et al., 1999), e.g. guava (Psidium guajava), pomegranate (Punica granatum) and citrus (Citrus aurantifolia). Recent studies in Tanzania (Mziray et al., 2010) have confirmed the predominance in Solanaceae with occasional infestation in Cucurbitaceae.

Hosts in Africa:

Family: Solanaceae

Common name	Scientific name	Country, Reference
Paprika	Capsicum annuum var. longum	Tanzania, Mziray et al., 2010
Habanero pepper	Capsicum chinense	Tanzania, Mziray et al., 2010
Tomato	Lycopersicon esculentum	Tanzania, Mziray et al., 2010
Cherry tomato	Lycopersicon pimpinellifolium	Tanzania, Mziray et al., 2010
African eggplant	Solanum aethiopicum	Tanzania, Mwatawala et al., 2007
African eggplant	Solanum anguivi	Tanzania, Mziray et al., 2010
	Solanum incanum	Tanzania, Mziray et al., 2010
African eggplant	Solanum macrocarpon	Tanzania, Mwatawala et al., 2007
Eggplant, aubergine	Solanum melongena	Tanzania, Mziray et al., 2010
Black nightshade	Solanum nigrum	Tanzania, Mziray et al., 2010
	Solanum scabrum	Tanzania, Mziray et al., 2010
Sodom apple	Solanum sodomeum	Tanzania, Mziray et al., 2010

Family: Cucurbitaceae

Common name	Scientific name	Country, Reference
	Citrullus lanatus	Tanzania, Mziray et al., 2010
Teasel gourd	Cucumis dipsaceus	Tanzania, Mziray et al., 2010
	Momordica cf trifoliata	Tanzania, Mziray et al., 2010

Major hosts in Asia:

Family: Solanaceae

Common name	Scientific name	Reference
	Capsicum anuum	Allwood et al., 1999
Eggplant, aubergine	Solanum melongena	Allwood et al., 1999
Tomato	Lycopersicon esculentum	Allwood et al., 1999

Other hosts in Asia and Hawaii:

Family: Solanaceae

Common name	Scientific name	Reference
	Lycianthes macrodon	Allwood et al., 1999
	Solanum aculeatissimum	Allwood et al., 1999
	Solanum anguivi	Allwoodet al., 1999
	Solanum donianum	Hardy, 1973
	Solanum granuloso-leprosum	Allwood et al., 1999
	Solanum incanum	Allwood et al., 1999
	Solanum mammosum	Allwood et al., 1999
Black nightshade	Solanum nigrum	Allwood et al., 1999
	Solanum santiwongsei	Allwood et al., 1999
	Solanum sarmentosum	Hardy, 1973
	Solanum sisymbriifolium	White & Elson-Harris, 1992
	Solanum stramoniifolium	Allwood et al., 1999
Terongan	Solanum torvum	Allwood et al., 1999
	Solanum trilobatum	Allwood et al., 1999
	Solanum xanthocarpum	Tsuruta et al., 1997

Family: Cucurbitaceae

Common name	Scientific name	Country, Reference
	Benincasa hispida	Hawaii, Liquido et al., 1994
Ivy gourd	Coccinea grandis	Hawaii, Liquido et al., 1994
Cucumber	Cucumis sativus	Hawaii, Liquido et al., 1994
White-flowered gourd	Lagenaria siceraria	Hawaii, Liquido et al., 1994

Family: Euphorbiaceae

Common name	Scientific name	Reference
	Baccaurea motleyana	White & Elson-Harris, 1994

Family: Lythraceae

Common name	Scientific name	Reference
	Lagerstroemia indica	Allwood et al., 1999

Family: Myrtaceae

Common name	Scientific name	Reference
Guava	Psidium guajava	Allwood et al., 1999

Family: Oleaceae

Common name	Scientific name	Reference
	Linociera parkinsoni	Allwood et al., 1999
	Linociera xanthocarpum	Allwood et al., 1999

Family: Passifloraceae

Common name	Scientific name	Reference
Wild watermelon	Passiflora foetida	Allwood et al., 1999

Family: Punicaceae

Common name	Scientific name	Reference
Pomegranate	Punica granatum	Allwood et al., 1999

Family: Rhamnaceae

Common name	Scientific name	Reference
	Ziziphus rotundifolia	Allwood et al., 1999

Family: Rutaceae

Common name	Scientific name	Reference
Lime	Citrus aurantifolia	Allwood et al., 1999
Orange jessamine	Murraya paniculata	Allwood et al., 1999

Family: Sapindaceae

Common name	Scientific name	Reference
	Sapindus rarak	Allwood et al., 1999

Family: Verbenaceae

Common name	Scientific name	Reference
	Gmelina philippensis	Allwood et al., 1999

6.4. Host plants of B. zonata

Bactrocera zonata is highly polyphagous, i.e. it attacks a wide range of plants belong to lots of different families. Its major hosts include peach (Prunus persica), guava (Psidium guajava) and mango (Mangifera indica).

Hosts in African region:

Family: Anacardiaceae

Common name	Scientific name	Country, Reference
Mango	Mangifera indica	Egypt S. Mohamed own observations
Mango	Mangifera indica	Mauritius & Réunion, Quilici & Jeuffrault, 2001

Family: Annonaceae

Common name	Scientific name	Country, Reference
Custard apple	Annona reticulata	Mauritius, Quilici & Jeuffrault, 2001
Annona	Annona squamosa	Egypt, S. Mohamed own observations

Family: Arecaceae

Common name	Scientific name	Country, Reference
Date	Phoenix dactylifera	Egypt, S. Mohamed own observations

Family: Caricaceae

Common name

Scientific name

Country, Reference

Papaya

Carica papaya

Egypt, S. Mohamed own observations

Mauritius, Quilici & Jeuffrault, 2001

Family: Combretaceae

Common name	Scientific name	Country, Reference
Topical almond	Terminalia catappa	Mauritius & Réunion, Quilici & Jeuffrault, 2001

Family: Cucurbitaceae

Common name	Scientific name	Country, Reference
Watermelon	Citrullus lanatus	Mauritius, Quilici & Jeuffrault, 2001
Cucurbit	Cucurbita sp.	Egypt, S. Mohamed own observations
Bottle gourd	Lagenaria leucaritha	Mauritius, Quilici & Jeuffrault, 2001

Family: Lauraceau

Common name	Scientific name	Country, Reference
Avocado	Persea americana	Mauritius, Quilici & Jeuffrault, 2001

Family: Lythraceae

Common name	Scientific name	Country, Reference
Pomegranate	Punica granatum	Egypt, S. Mohamed own observations

Family: Moraceae

Common name

Scientific name

Country, Reference

Fig

Ficus carica

Egypt, S. Mohamed own observations

Mauritius, Quilici & Jeuffrault, 2001

Family: Myrtaceae

Common name	Scientific name	Country, Reference
Rose apple	Syzygium jambos	Mauritius, Quilici & Jeuffrault, 2001
Java apple	Syzygium samarangense	Mauritius, Quilici & Jeuffrault, 2001
Guava	Psidium guajava	Egypt, S. Mohamed own observations
Guava	Psidium guajava	Mauritius, Quilici & Jeuffrault, 2001

Family: Oleaceae

Common name	Scientific name	Country, Reference
Olive	Olea europaea	Egypt, S. Mohamed own observations

Family Rhamnaceae

Common name	Scientific name	Country, Reference
Indian jujube	Ziziphus mauritiana	Mauritius, Quilici & Jeuffrault, 2001

Family: Rosaceae

Common name	Scientific name	Country, Reference
Apple	Malus domestica	Mauritius, Quilici & Jeuffrault, 2001
Apricot	Prunus armeniaca	Egypt, S. Mohamed own observations
Peach	Prunus persica	Egypt, S. Mohamed own observations Mauritius, Quilici & Jeuffrault, 2001
Pear	Pyrus sp.	Egypt, S. Mohamed own observations
Plum	Prunus sp.	Egypt, S. Mohamed own observations

Family: Rutaceae

Common name	Scientific name	Country, Reference
Grape fruit	Citrus paradisi	Egypt, S. Mohamed own observations
Orange	Citrus sp.	Egypt, S. Mohamed own observations
Orange	Citrus sp.	Saudi Arabia 1982, BMNH collection data

Family: Solanaceae

Common name	Scientific name	Country, Reference
Bugtree	Solanum mauritianum	Mauritius, Quilici & Jeuffrault, 2001

Note: All records from Egypt were confirmed by the second author through rearing from infested samples collected in the field. In addition B. zonata is recorded in Egypt from the following hosts but these records were not verified and need confirmation: Cucumis melo; Cucumis sativus (Cucurbitaceae); Abelmoschus esculentus (Malvaceae); Ziziphus sp. (Rhamnaceae); Capsicum annuum; Lycopersicon esculentum, Solanum melongena (Solanaceae); Vitis sp. (Vitaceae).

Hosts in Asia:

Family: Anacardiaceae

Common name	Scientific name	Reference
Mango	Mangifera indica	White & Elson-Harris, 1992

Family: Annonaceae

Common name	Scientific name	Reference
Annona	Annona squamosa	White & Elson-Harris, 1992

Family: Arecaceae

Common name	Scientific name	Reference
Date	Phoenix dactylifera	White & Elson-Harris, 1992

Family: Caricaceae

Common name	Scientific name	Reference
Papaya	Carica papaya	White & Elson-Harris, 1992

Family: Combretaceae

Common name	Scientific name	Reference
Tropical almond	Terminalia catappa	Allwood et al., 1999; Tsuruta et al., 1997

Family: Cucurbitaceae

Common name	Scientific name	Reference
Bitter gourd	Momordica charantia	White & Elson-Harris, 1992

Family: Fabaceae

Common name	Scientific name	Reference
	Afzelia xylocarpa	Allwood et al., 1999

Family: Lecythidaceae

Common name	Scientific name	Reference
	Careya arborea	Allwood et al., 1999; Tsuruta et al., 1997

Family: Malpighiaceae

Common name	Scientific name	Reference
Huesito	Malpighia glabra	Allwood et al., 1999

Family: Malvaceae

Common name	Scientific name	Reference
Okra	Abelmoschus esculentus	White & Elson-Harris, 1992

Family: Myrtaceae

Common name	Scientific name	Reference
Guava	Psidium guajava	White & Elson-Harris, 1992
Rose-apple	Syzygium jambos	Allwood et al., 1999

Family: Punicaceae

Common name	Scientific name	Reference
Pomegranate	Punica granatum	White & Elson-Harris, 1992

Family: Rosaceae

Common name	Scientific name	Reference
Quince	Cydonia oblonga	White & Elson-Harris, 1992
Apple	Malus domestica	White & Elson-Harris, 1992
Paradise apple	Malus pumila	White & Elson-Harris, 1992
Peach	Prunus persica	White & Elson-Harris, 1992

Family: Rutaceae

Common name	Scientific name	Reference
Indian bael	Aegle marmelos	Allwood et al., 1999
Sweet orange	Citrus sinensis	White & Elson-Harris, 1992

Family: Tiliacea

Common name	Scientific name	Reference
Phalsa	Grewia asiatica	White & Elson-Harris, 1992

7. Displacement of Native and Introduced Fruit Flies by Asian Bactrocera spp.

The impact of accidental introductions of Asian Bactrocera pests on the fruit and vegetable production in Africa cannot be underestimated. Some of the introductions, e.g. B. latifrons have a host range that is not fully exploited by the native fruit fly species. This could aggravate the problems already encountered by local farmers, by attacking crops that were relatively fruit fly pest free.

Other introduced Asian Bactrocera spp. are polyphagous and encounter interspecific competition with other polyphagous tephritid flies that are already well established (either as native species or as African species that were introduced before). For example, casual observations in western and eastern Africa suggest that B. invadens is largely displacing the native mango fruit fly (Ceratitis cosyra) in infestation of mangoes. In Egypt, because of the spread of B. zonata, the Mediterranean fruit fly, Ceratitis capitata has become more restricted in the horticulture areas (Hashem et al. 2001; Safaan, 2005, 2006) and the mixed infestation in fruits of both fruit flies produced flies mostly of B. zonata irrespective of which insect infested the fruit first (Mohamed, 2004).

Displacement can lead to shifts to hosts that were considered of minor importance before, or to climatic niche partitioning between the different pest species. Extensive work on these aspects has been carried out on the island of La Réunion where one native fruit fly species is found and three exotic species have successively invaded the island (Duyck et al., 2006a, 2006b), including B. zonata. The trade-offs from their respective life histories (with the larger flies of the genus Bactrocera having a more K-selection orientated profile than the smaller Ceratitis flies) can have its consequences in Asian Bactrocera introductions being able to displace Ceratitis, as observed in recent invasions (Duyck et al., 2004a, 2007).

8. Further Reading

This section groups relevant literature on morphology, ecology, and other aspects for each of the introduced species. The aim is to provide guidance to the available literature. We have emphasized literature references based upon research conducted in Africa, although a selection of recent literature references originating from research elsewhere are included. If a series of papers dealing with the same topic are available, the most recent one is listed below if it includes the references to older literature. Papers dealing with aspects of fruit flies in general, or general review monographs, are excluded.

Subsections:

8.1. Bactrocera cucurbitae

8.2. Bactrocera invadens

8.3. Bactrocera latifrons

8.4. Bactrocera zonata

8.1. Bactrocera cucurbitae

Behaviour: Rahman et al., 2003

Biocontrol agents in Thailand: Ramadan & Messing, 2003

Biology: Chaudhary & Patel, 2007

Demography in Hawaii: Vargas et al., 1996

Development of immature stages: Vayssières et al., 2008a

Distribution and host plants in West and Central Africa: Vayssières et al., 2007

Distribution and host plants in Tanzania: Mwatawala et al., 2010

Eradication in Japan: Koyama et al., 2004

Improved male attractant: Jang et al., 2007

Interspecific competition with Dacus ciliatus: Qureshi et al., 1987

Larval diet for rearing: Chang et al., 2004

Life tables: Huang & Chi, 2012

Mating compatibility: Sookar et al., 2013

Population dynamics and distribution in Hawaii: Vargas et al., 1990 and Tanzania: Geurts et al., 2012; 2014

Population genetic structure: Hu et al., 2008; Virgilio et al., 2010; Wu et al., 2011; 2012; Jacquard et al., 2013

Review on biology and management: Dhillon et al., 2005

8.2. Bactrocera invadens

Biocontrol with weaver ants in Benin: Van Mele et al., 2007; 2009

Competitive success in Kenya: Ekesi et al., 2009; Rwomushana et al., 2009

Control with spinosad in Benin: Vayssières et al., 2009b

Detection in India: Sithanantham et al., 2006

Detection in Bhutan: Drew et al., 2007

Detection in Mozambique: Correia et al., 2008

Diversity in citrus crops: Vayssières et al., 2010a

Ecological modeling: De Meyer et al., 2010

Eradication in South Africa: Manrakhan et al., 2011

Food based attractants: Ekesi et al., 2014

Host preference in eastern Africa: Rwomushana et al., 2008a; Mwatawala et al., 2009; Cugala et al., 2013; José et al., 2013

Host preference in western Africa: Vayssières et al., 2009a; N'Dépo et al., 2010; Goergen et al., 2011

Infestation rates and correlation with biotic and abiotic factors in Benin: Vayssières et al., 2009c

Invasion history in Africa: Khamis et al., 2009; Malavasi et al., 2013

Parasitoids: Mohamed et al., 2010 ; Vayssières et al., 2010b

Population dynamics in Ivory Coast: N’Dépoet al., 2009 and Tanzania: Geurts et al., 2012, 2014

Post harvest treatment: Grout et al., 2011; Hallman et al., 2011

Rearing on artificial diet: Ekesi et al., 2007

Seasonality and host utilization in Tanzania: Mwatawala et al., 2006

Survival and development of immature life stages: Rwomushana et al., 2008b

Life history in Kenya: Ekesi et al., 2006

Life history: in East Africa: Ekesi et al., 2006; Salum et al., 2013; review Malavasi et al., 2013

Infestation in mango in Benin: Vayssières et al., 2005

Taxonomic status: San José et al., 2013

Volatiles male rectal pheromone gland: Tan et al., 2011

8.3. Bactrocera latifrons

Allozyme variation: Yong, 1993

Demography in Hawaii: Vargas et al., 1996

Development and reproductive ability on Yonaguni Island, Japan: Ishida et al., 2005.

Detection in Africa: Mwatawala et al., 2007

Distribution inTanzania: Mwatawala et al., 2010

Host plant range and distribution in Hawaii: Liquido et al., 1994, Harris et al., 2002, Peck & McQuate, 2004

Host plant range and temporal distribution inTanzania: Mziray et al., 2009, 2010

Identification with PCR: Yu et al., 2004

Invasion to Yonaguni Island, Japan: Shimizu et al., 2007

Larval morphology: Chen, 1997

Mating behaviour: Jackson & Long, 1997

Parapheromones: McQuate et al., 2004 ; Keum et al., 2007; Ishida et al., 2008

Parasitoids, Hawai: Bokonon-Ganta et al., 2007

Rearing: Vargas et al., 1993

8.4. Bactrocera zonata

Abundance in orchards in Egypt: Hashem et al., 2001

Biology and life table parameters: El-Aw et al., 2003

Control measures with killing bags in Egypt: Mohamed & El-Wakkad, 2003; with plant extracts: Jilani et al., 2006

Detection and monitoring in Egypt: Mohamed, 2001; Saafan, 2005

Interspecific competition in India: Agarwal et al., 1999; in Egypt: Mohamed, 2004; and climatic niche partitioning in Réunion: Duyck et al., 2006a, b.

Larval morphology and identification: Kandybina, 1977; Khan & Khan, 1987

Male terminalia: Singh & Premlata, 1985

Morphology and differentiation from other Bactrocera species: Iwahashi & Routhier, 2001

Ecology and population dynamics in Egypt: Afia, 2007 [unpublished PhD thesis]; Hashem et al., 2001; Safaan et al., 2006

Parasitoids in Egypt: Fetho & Afia, 2004; in Réunion: Rousse et al., 2006

Post harvest disinfestations: Hashem et al., 2001

SIT, effect of radiation: Shehata et al., 2006

Survival and development of different life stages: Duyck et al., 2004b

Visual responses to coloured traps: Hussain et al., 1995

9. Acknowledgements and Contacts

The authors gratefully acknowledge the financial support provided by the International Atomic Energy Agency through IAEA technical contract (Nr 14151/R0 to the Royal Museum for Central Africa, RMCA) that enabled the production of this website, as well as an IAEA fellowship (C6/EGY/06047) to the second author for his stay at RMCA.
Many thanks to M. Mwatawala (SUA, Morogoro Tanzania), S. Ekesi (ICIPE, Nairobi Kenya), G. Goergen (IITA, Benin), S. Quilici (CIRAD, La Réunion), and J.-F. Vayssières (CIRAD/IITA, Benin) for allowing us to cite some of their unpublished observations; to G. McQuate (USDA-ARS, Hawaii), R.S. Copeland (ICIPE) and S. Quilici (CIRAD) for allowing us to use photographs, their copyright.

Queries, comments or suggested additions to this website can be sent to:

Marc De Meyer,

Invertebrates Unit,

Royal Museum for Central Africa,

Leuvensesteenweg 13,

B-3080 Tervuren,

Belgium.

Email: demeyer<at>africamuseum<dot>be

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(Updated on February 26, 2014)