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)

  1. 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.
  2. Most fruit flies are colourful with at least some patterning on the wings.
  3. The species we are interested in are small to medium sized flies, with each wing between 4mm and 7mm long.
  4. 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:

  1. Male lure:
    1. Any fly responding to cue lure is almost certain to be one of the native African Dacus sp. or perhaps Bactrocera cucurbitae.
    2. 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).
  2. Reared from commercial fruit or certain vegetables:
    1. 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).
    2. 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.).
    3. 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.).
  3. 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.

  1. 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).
  2. 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.
  3. Most Bactrocera and Dacus spp., including the invasive Bactrocera spp., have a pair of black spots on the face, one under each antenna (photo).
  4. 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.
  5. 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.

  1. 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.
  2. 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.:

  1. The terga (dorsal sclerites of the abdomen) are all fused into a single piece in Dacus spp.
  2. 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:

  1. Males attracted to cue lure, and (rest applies to both sexes):
  2. Scutum with a medial postsutural vitta (photo), and
  3. 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:

  1. Males attracted to methyl eugenol, and (rest applies to both sexes)
  2. Wing without a complete costal band (photo; reduced to a coloured mark in cell sc and an apical spot), and
  3. Wing without a well marked anal streak (photo), and
  4. 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:

  1. Yellow strip on side of thorax (the “anepisternal xanthine”) very broad, often almost contacting the shoulder (“postpronotal lobe”) (photo), and
  2. Abdomen mostly red-brown (no lateral marking on tergite IV; usually without a dark medial strip), and
  3. 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:

  1. Males attracted to methyl eugenol, and (rest applies to both sexes, except as indicated)
  2. Yellow strip on side of thorax (the “anepisternal xanthine”) narrow broad, always well separated from the shoulder (“postpronotal lobe”) (photo), and
  3. Abdomen, with extensive black markings (photo):
    1. Terga III-V with a distinct black mid-longitudinal line (photo), and
    2. 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
    3. tergite IV with broad antero-lateral markings which is usually almost rectangular (photo), or more extensively black, and
  4. Thorax (dorsal surface) variable from red-brown to black, often red-brown with black streaks, and
  5. 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.

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).

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 (Mozambique, Zambia, Namibia) have been reported (Correia et al., 2008; USDA-APHIS Pretoria office, unpublished reports).

 

 

Map showing countries in Africa from which B. invadens is reported (based upon records in White, 2006 and references therein; 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. Presently, it is not reported from the Afrotropical mainland below the Sahara.

 

 

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

Senegal 2005, IITA/CIRAD data

Mango

Mangifera indica

Benin 2004, IITA/CIRAD data

Tanzania 2003, MRAC collection data ; Kenya, Rwomushana et al., 2008a

Marula plum

Sclerocarya birrea

Ekesi et al., 2006

Western Africa, IITA data

 

Sorindeia madagascariensis

Kenya, Rwomushana et al., 2008a

Jew plum

Spondias cytherea

Tanzania, Mwatawala et al., 2006

Tropical plum

Spondias mombin

Western Africa, IITA data

 

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

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

Western Africa, IITA data

 

Family : Caricaceae

Common name

Scientific name

Country, Reference

Papaya

Carica papaya

Benin 2004, IITA/CIRAD data

 

Family : Clusiaceae

Common name

Scientific name

Country, Reference

Chewing stick

Garcinia mannii

Western Africa, IITA data

 

Family : Combretaceae

Common name

Scientific name

Country, Reference

Tropical almond

Terminalia catappa

Ekesi et al., 2006; Benin, Vayssières et al., 2008b

Tanzania 2006, MRAC/SUA data

 

Family : Cucurbitaceae

Common name

Scientific name

Country, Reference

Watermelon

Citrullus lanatus

Tanzania, Mwatawala et al., 2006

Western Africa, IITA data

 

Cucumis figarei

Tanzania, Mwatawala et al., 2006 based on single specimen reared

Cucumber

Cucumis sativus

Tanzania, Mwatawala et al., 2006

Western Africa, IITA data

Pumpkin

Cucurbita maxima

Western Africa, IITA data

Gourd

Cucumis pepo

Western Africa, IITA data

 

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 2004, IITA/CIRAD data

 

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

Ekesi et al., 2006

Tanzania, Mwatawala et al., 2006

 

Family: Moraceae

Common name

Scientific name

Country, Reference

 

Ficus ? sycomorus

Western Africa, IITA data

 

Family : Musaceae

Common name

Scientific name

Reference

Plantain

Musa x  paradisiaca

Western Africa, IITA data

 

Musa sp.

Ekesi et al., 2006

 

Family: Myrtaceae

Common name

Scientific name

Country, Reference

Jambolan

Syzygium cumini

Tanzania, Mwatawala et al. , 2009

Rose apple

Syzygium jambos

Tanzania, SUA unpublished data

Malay apple

Syzygium malaccense

Western Africa, IITA data

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 2004, IITA/CIRAD data

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: Rosaceae

Common name

Scientific name

Country, Reference

Loquat

Eriobotrya japonica

Tanzania, Mwatawala et al., 2006

Western Africa, IITA data

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

Western Africa, IITA data

Grapefruit

Citrus  paradisi

Tanzania, Mwatawala et al., 2006

Western Africa, Vayssières et al., 2009a

Tangerine

Citrus reticulata

Benin 2004, IITA/CIRAD data

Tanzania, Mwatawala et al., 2006; Kenya, Rwomushana et al., 2008a

Sweet orange

Citrus sinensis

Benin 2004, IITA/CIRAD data; Kenya, Rwomushana et al., 2008a; Tanzania, Mwatawala et al., 2009

Tangelo

Citrus tangelo

TWestern Africa, Vayssièreset al., 2009a

Kumquat

Fortunella margarita

Tanzania, Mwatawala et al., 2010

 

Family: Sapindaceae

Common name

Scientific name

Country, Reference

 

Blighia sp.

Western Africa, IITA data

 

Family: Sapotaceae

Common name

Scientific name

Country, Reference


Achra sapota

Ivory Coast, N’Depo et al., 2010

White star-apple

Chrysophyllum albidum

Benin 2004, IITA/CIRAD data


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, IITA data

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

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

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

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

Population dynamics and distribution in Hawaii: Vargas et al., 1990

Population genetic structure: Hu et al., 2008

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

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

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

Invasion history in Africa: Khamis et al., 2009

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

Population dynamics in Ivory Coast: N’Dépoet al., 2009

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

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

Volatiles male rectal pheromone gland: Tan et al., 2010

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,

Entomology Section,

Royal Museum for Central Africa,

Leuvensesteenweg 13,

B-3080 Tervuren,

Belgium.

 

Email: demeyer<at>africamuseum<dot>be

10. Literature References

Abuel-Ela, R.G., Hashem, A.G & Mohamed, S.M.A. (998) Bactrocera pallidus (Perkin and May) (Diptera: Tephritidae), a new record in Egypt. Journal of the Egyptian German Society of Zoology (Entomology), 27,  221 - 229.

 

Agarwal, M.L., Kumar, P. & Kumar, V. (1999) Population suppression of Bactrocera dorsalis (Hendel) by Bactrocera zonata (Saunders) (Diptera: Tephritidae) in north Bihar. Shashpa, 6, 189-191.

 

Allwood, A.L., Chinajariyawong, A., Drew, R.A.I., Hamacek, E.L., Hancock, D.L., Hengsawad, C.,  Jipanin, J.C., Jirasurat, M., Kong Krong, C., Kritsaneepaiboon, S., Leong, C.T.S. &  Vijaysegaran, S.  (1999). Host plant records for fruit flies (Diptera: Tephritidae) in Southeast Asia.  Raffles Bulletin of Zoology, Supplement, 7, 1-92.

 

Back, E.A. & Pemberton, C.E. (1914) Life history of the melon fly. Journal of Agricultural Research, 3, 269-274.

 

Bokonon-Ganta, A.H., McQuate, G.T. & Messing, R.H. (2007) Natural establishment of a parasitoid complex on Bacrocera latifrons (Diptera: Tephritidae) in Hawaii. Biological Control, 42, 365-373.

 

Chang, C.L., Caceres, C. & Jang, E.B. (2004) A novel liquid larval diet and its rearing system for melon fly, Bactrocera cucurbitae (Diptear: Tephritidae). Annals of the Entomological Society of America, 97, 524-528.

 

Chaudhary, F.K. & Patel, G.M. (2007) Biology of melon fly, Bactrocera cucurbitae Conquillett [sic] on pumpkin. Indian Journal of Entomology, 69, 168-171.

 

Chen, Z.L. (1997) Description of the larval morphology of Bactrocera latifrons (Hendel). Entomological Knowledge, 34, 302-303.

 

Correia, A.R., Rego, J.M. & Olmi, M. (2008) A pest of significant economic importance detected for the first time in Mozambique: Bactrocera invadens Drew, Tsuruta & White (Diptera: Tephritidae: Dacinae). Bollettino di Zoologia Agraria e di Bachicoltura, ser II, 40 , 9-13.

 

De Meyer, M., Robertson, M.P., Mansell, M.W., Ekesi, S., Tsuruta, K., Mwaiko, W., Vayssières J.-F. & Peterson A.T. 2010. Ecological niche and potential geographic distribution of the Invasive Fruit Fly Bactrocera invadens (Diptera, Tephritidae). Bulletin of Entomological Research , 100, 35-48.

 

Dhillon, M.K., Singh, R., Naresh, J.S. & Sharma, H.C. (2005) the melon fruit fly, Bactrocera cucurbitae: a review of its biology and management. Journal of Insect Science, 5, unpaginated.

 

Drew, R.A.I. (1989a) The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Memoirs of the Queensland Museum, 26, 1-521.

 

Drew, R.A.I., Tsuruta, K. & White, I.M. (2005) A new species of pest fruit fly (Diptera: Tephritidae: Dacinae) from Sri Lanka and Africa. African Entomology, 13, 149-154.

 

Drew, R.A.I., Romig, M.C. & Dorji, C. (2007) Records of Dacine fruit flies and new species of Dacus (Diptera: Tephritidae) in Bhutan. The Rafles Bulletin of Zoology, 55, 1-21.

 

Duyck, P.F., David, P. & Quilici, S. (2004a) A review of relationships between interspecific competition and invasions in fruit flies (Diptera: Tephritidae). Ecological Entomology, 29, 511-520.

 

Duyck, P.F., Sterlin, J.F. & Quilici, S. (2004b) Survival and development of different life stages of Bactrocera zonata (Diptera: Tephritidae) reared at five constant temperatures compared to other fruit fly species. Bulletin of Entomological Research, 94, 89-93.

 

Duyck, P.F., David, P., Junod, G., Brunel, C., Dupont, R. & Quilici, S. (2006a) Importance of competition mechanisms in successive invasions by polyphagous tephritids in La Réunion. Ecology, 87, 1770-1780.

 

Duyck, P.F., David, P. & Quilici, S. (2006b) Climatic niche partitioning following successive invasions by fruit flies in La Réunion. Journal of Animal Ecology, 75, 518-526.

 

Duyck, P.F., David, P. & Quilici, S. (2007) Can more K-selected species be better invaders ? A case study of fruit flies in La Réunion. Diversity and Distributions, 13, 535-543.

 

Efflatoun, H.C. (1924) A monograph of Egyptian Diptera, part II., Fam. Trypaneidae. Mémoires de la société entomologique de l’Egypte 2, 1-132.

 

Ekesi, S., Nderitu, P.W. & Rwomushana, I. (2006) Field infestation, life history and demographic parameters of the fruit fly Bactrocera invadens (Diptera: Tephritidae) in Africa. Bulletin of Entomological Research, 96, 379-386.

 

Ekesi, S., Nderitu, P.W. & Chang, C.L. (2007) Adaptation to and small-scale rearing of invasive fruit fly Bactrocera invadens (Diptera: Tephritidae) on artificial diet. Annals of the Entomological Society of America, 100, 562-567.

 

Ekesi, S., Billah, M.K., Nderitu, P.W., Lux, S.A. & Rwomushana, I. (2009) Evidence for competitive displacement of Ceratitis cosyraby the invasive fruit fly Bactrocera invadens (Diptera: Tephritidae) on mango and mechanisms contributing to the displacement. Journal of Economic Entomology , 102, 981-991.

 

El-Aw, M.A., Draz, K.A.A., Hashem, A.G. & El-Gendy, I.R. (2003) Biology and life table parameters of the peach fruit fly, Bacterocera zonata, (Saunders) (Diptea: Tephritidae) reared on different host fruits. Alexandria Journal of Agricultural Research, 48, 31-37.

 

Hu, J., Zhang, J.L., Nardi, F. & Zhang, R.J. (2008) Population structure of the melon fly, Bactrocea cucurbitae (Diptera: Tephritidae), from China and Southeast Asia. Genetica, 134, 319-324.

 

Fetoh, B.A. and Afia Y.I. (2004) Preliminary biological aspects of the new recorded parasitoids, Dirhinus luzonesis, (Hymenoptera: Chalcididae) of Cucurbit fruit fly, Dacus ciliatus (Loew.) and Peach fruit fly, Bactrocera (Dacus) zonata (Saunders) (Diptera: Tephritidae) in Egypt. Egyptian Journal on Biological Pest Control, 14,

 

Hardy, D.E. (1973) The fruit flies (Tephritidae Diptera) of Thailand and bordering countries. Pacific Insects Monograph, 31, 1-353.

 

Harris, E.J., Liquido, N.J. & Spencer, J.P. (2002) Distribution and host utilization of Bactrocera latifrons (Diptera: Tephritidae) on the island of Kauai, Hawaii. Proceedings of the Hawaiian Entomological Society, 35, 55-6.

 

Hashem, A.G., Mohamed, S.M.A. & El-Wakkad, M.F. (2001) Diversity and abundance of Mediterranean and Peach fruit flies (Diptera: Tephritidae) in different horticultural orchards. Egyptian Journal of Applied Sciences, 16, 303-314.

 

Hu, J., Zhang, J.L., Nardi, F. & Zhang, R.J. (2008) Population structure of the melon fly, Bactrocea cucurbitae (Diptera: Tephritidae), from China and Southeast Asia. Genetica, 134, 319-324.

 

Hussain, T., Siddiqui, Q.H. & Qureshi, Z.A. (1995) Visual responses of Bactrocera zonata (Saunders) to traps of different colours. Proceedings of Pakistan Congress of Zoology, 15, 313-318.

 

Ishida, T., Nakahara, S., Minoura, K. & Dohino, T. (2005) Development and reproductive ability of Bactrocera latifrons (Hendel) (Diptera: Tephritidae) on Yonaguni Island, Okinawa. Research Bulletin of the Plant Protection Service Japan, 41, 39-42.

 

Ishida, T., Enomoto, H. & Nishida, R. (2008) New attractants for males of the Solanaceous fruit fly Bactrocera latifrons. Journal of Chemical Ecology, 34, 1532-1535.

 

Iwahashi, O. & Routhier, W. (2001) Aedeagal length and its variation of the peach fruit fly, Bactrocera zonata (Saunders) (Diptera: Tephritidae), which recently invaded Egypt. Applied Entomology and Zoology, 36, 13-17.

 

Jackson, C.G. & Long, J.P. (1997) Mating behaviour of Bactrocera latifrons (Diptera: Tephritidae) in field cages. Annals of the Entomological Society of America, 90, 856-860.

 

Jang, E.B., Casana-Giner, V. & Oliver, J.E. (2007) Field captures of wild Melon fly (Diptera: Tephritidae) with an improved male attractant, raspberry ketone formate. Journal of Economic Entomology, 100, 1124-1128.

 

Jilani, G., Khattak, M.K. & Shahzad, F. (2006) Toxic and growth regulating effect on ethanol extract and petroleum ether extract of Valariana officianlis L. against Bactrocera zonata Saunders. Pak. Entomology 28, 11-14.

 

Kandybina, M.N. (1977) The larvae of fruit flies (Diptera, Tephritidae). Opredeliteli po Fauna SSR, Izdavaemye Zoolologischeskim Muzeem Akademii Nauk 144, 1 – 212.

 

Keum, Y.-S., McQuate, G.T. & Qing, X.L. (2007) Synergists isolated from cade oil for the parapheromone alpha-ionol for male Bactrocera latifrons (Diptera: Tephritidae). Biochemical Systematics and Ecology, 35, 188-195.

 

Khamis, F.M., Karam, N., Ekesi, S., De Meyer, M., Bonomi, A., Gomulski, L.M., Scolari, F., Gabrieli, P., Siciliano, P., Masiga, D., Kenya, E.U., Gasperi, G., Malacrida, A.R. & Guglielmino, C.R. (2009) Uncovering the tracks of a recent and rapid invasion: the case of the fruit fly pest Bactrocera invadens (Diptera: Tephritidae) in Africa. Molecular Ecology (online publication).

 

Khan,R.J. &Khan, M.A.J. (1987) A comparative morphological study on third instar larvae of some Dacus Species (Tephritidae: Diptera) in Pakistan. Pakistan Journal for Scientific and Industrial Research 30, 534 - 538.

 

Koyama, J., Kakinohana, H. & Miyatake, T. (2004) Eradication of the melon fly, Bactrocera cucurbitae, in Japan: importance of behaviour, ecology, genetics, and evolution. Annual Review of Entomology, 49, 331-349.

 

Liquido, N.J., Harris, E.J., Dekker, L.A. (1994) Ecology of Bactrocera latifrons (Diptera: Tephritidae) populations: host plants, natural enemies, distribution, and abundance. Annals of the Entomological Society of America, 87, 71-84.

 

Lux, S.A., Copeland, R.S., White, I.M., Manrakhan, A. & Billah, M.K. (2003) A new invasive fruit fly species from the Bactrocera dorsalis (Hendel) group detected in East Africa. Insect Science and its Application, 23, 355-360.

 

McQuate, G.T., Keum, Y.S., Sylva, C.D., Li Q.X. & Jang, E.B. (2004) Active ingredients in cade oil that synergize attractiveness of alpha-ionol to male Bactrocera latifrons (Diptera: Tephritidae). Journal of Economic Entomology, 97, 862-870.

 

Mohamed, S.M.A. (2001) Efficiency of different trap combinations for detecting and monitoring the Peach fruit fly Bactrocera zonata (Saunders) in Egypt. Egyptian journal of Applied Sciences, 16, 315-334.

 

Mohamed, S.M.A. (2004) Competition between Mediterranean fruit fly and Peach fruit fly in fruit infestation. Journal of the Egyptian German Society of Zoology (Entomology), 43, 17-23.

 

Mohamed, S.M.A. & El-Wakkad, M.F. (2003) the role of killing bags on the reduction of Peach fruit fly in horticultural orchards. Journal of the Egyptian German Society of Zoology (Entomology), 42, 11-19.

 

Mohamed, S.A., Ekesi, S. & Hanna, R. (2008) Evaluation of the impact of Dachasmimorpha longicaudata on Bactrocera invadens and five African fruit fly species. Journal of Applied Entomology, 132, 789-797.

Mohamed, S.A., Ekesi, S. & Hanna, R. (2010) Old and new host-parasitoid associations: parasitism of the invasive fruit fly Bactrocera invadens (Diptera: Tephritidae) and five African fruit fly species by Fopius arisanus, an Asian opiine parasitoid. Biocontrol Science and Technology, 20, 183-196.

 

Mwatawala, M.W., White, I.M., Maerere, A.P., Senkondo, F.J. & De Meyer, M. (2004) A new invasive Bactrocera species (Diptera: Tephritidae) in Tanzania. African Entomology, 12, 154-156.

 

Mwatawala, M.W., De Meyer, M., Makundi, R.H. & Maerere, A.P. (2006) Seasonality and host utilization of the invasive fruit fly, Bactrocera invadens (Dipt., Tephritidae) in central Tanzania. Journal of Applied Entomology, 130, 530-537.

 

Mwatawala, M.W., De Meyer, M., White, I.M., Maerere, A. & Makundi, R.H. (2007) Detection of the solanum fruit fly, Bactrocera latifrons (Hendel) in Tanzania (Dipt., Tephritidae). Journal of Applied Entomology, 131, 501-503.

 

Mwatawala, M., De Meyer, M., Makundi R. & Maerere, A. 2009 Host range and distribution of fruit-infesting pestiferous fruit flies (Diptera, Tephritidae) in selected areas of Central Tanzania. Bulletin of Entomological Research , 99, 629-641.

 

Mwatawala, M., Maerere, A., Makundi R. & De Meyer, M. 2010. Incidence and host range of Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae) in Central Tanzania. International Journal of Pest Management, 56, 265-273.

Mwatawala, M., Makundi, R., Maerere, A.P. & De Meyer, M. 2010. Occurrence of the invasive solanum fruit fly Bactrocera latifrons (Hendel) (Diptera: Tephritidae) in Tanzania. Journal of Afrotropical Zoology, 6, 83-89.

Mziray, H., Makundi, R., Mwatawala, M., Maerere, A. & De Meyer, M. 2009. Spatial and temporal abundance of the solanum fruit fly, Bactrocera latifrons (Hendel) in Morogoro, Tanzania. Crop Protection (online publication).

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N’depo, O.S., Hala, N., Gnago, A., Allou, K., Kouassi, K.P., Vayssières, J.-F. & De Meyer, M. 2010. Inventaire des mouches des fruits de trois regions agro-écologiques et des plantes-hôtes associées à l’espèce nouvelle, Bactrocera (Bactrocera) invadens Drew et al. (Diptera : Tephritidae) en Côte-d’Ivoire. European Journal of Scientific Research, 46, 62-72.

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Qureshi, Z.A., Hussain, T. & Siddiqui, Q.H. (1987). Interspecific competition of Dacus cucurbitae Coq. and Dacus ciliatus Loew in mixed infestation of cucurbits. Journal of Applied Entomology, 104, 429-432.

 

Rahman, A.K.M.Z., Alam, M.Z., Islam, M.N. & Hossain, M.M. (2003) Studies on the behavior of adult fruit fly (Bactrocera cucurbitae Coquillett) after emergence from pupae. Bulletin of the Institute of Tropical Agriculture Kyushu University, 26, 1-9.

 

Ramadan, M.M. & Messing, R.H. (2003) A survey for potential biocontrol agents of Bactrocera cucurbitae (Diptera: Tephritidae) in Thailand. Proceedings of the Hawaiian Entomological Society, 36, 115-122.

 

Ranganath, H.R. & Veenakumari, K. (1995) Notes on the dacine fruit flies (Diptera: Tephritidae) of Andaman and Nicobar Islands. The Raffles Bulletin of Zoology, 43, 235-238.

 

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(Last update : February 28, 2011)