Bactrocera+dorsalis

=1. Introduction= toc The Oriental fruit fly, //Bactrocera dorsalis// (Hendel, 1912), is a member of the Tephritidae (fruit flies) family. Although its' name does not illicit much response here in Singapore, in countries such as the United States and Kenya, the mention of the Oriental fruit fly will send agricultural farmers, fruit vendors, immigration authorities fuming mad. Native to tropical Asian countries like Singapore, this species has since established populations in over 90 countries around the world, causing massive damage to over 400 types of fruit and other crops wherever they are found. Read on to find out more about this notorious species!
 * Oriental fruit fly ** //Bactrocera dorsalis// (Hendel, 1912)



The Tephritidae family comprises non-fruit feeding tephritids, which are rarely pestiferous [2], and frugivorous tephritids, which contain several genera which are of major economic concern globally [3]. Some notorious fruigivorous tephritid species include: the Mediterranean fruit fly or Medfly ( //__Ceratitis capitata __// (Wiedemann))//,// the Cherry fruit fly ( Rhagoletis cingulata (Loew)), the Mexican fruit fly, (//__ Anastrepha ludens __// (Loew)) and the Oriental fruit fly ( //Bactrocera dorsalis // (Hendel)).

The Oriental fruit fly belongs to the //Bactrocera dorsalis// species complex, a group of almost 100 morphologically similar tephritid taxa [4, 5]. Amongst the generally inconspicuous species in this species complex, the Oriental fruit fly and the Carambola fruit fly ( //__B. carambolae __//) stand out in their economic impact. These species have been recognised as possibly the world’s most important pests of horticulture [6, 7]. This page focuses solely on the Oriental fruit fly.

Due to fairly recent updates to the scientific nomenclature of the Oriental fruit fly, information with regards to the Oriental fruit fly can also be found online under the following names: the Asian Papaya fruit fly (//Bactrocera papayae//), the Philippine fruit fly (//Bactrocera philippinensis//) and the Invasive fruit fly //(Bactrocera invadens//). For more information with regards to the many names of this species, head over to the Scientific Name & Synonyms section.

1.1. Disclaimer: Fruit fly ≠ Drosophila melanogaster
Whenever the phrase "fruit fly" is mentioned, the image of the model organism, //Drosophila melanogaster//, comes to mind. However, albeit their similar common names, //Drosophila melanogaster//, the Common fruit fly, does not belong to the same family as //Bactrocera dorsalis//, the Oriental fruit fly; //Drosophila melanogaste//r belongs to the family Drosophilidae whilst //Bactrocera dorsalis// belongs to the family Tephritidae (See Figure 3).

Significance: Important model organism in genetics and developmental biology research
 * Common fruit fly || Oriental fruit fly || Size comparison ||
 * [[image:drosophila melanogaster square.jpg width="292" height="292" link="@http://www.discoverlife.org/mp/20q"]] || [[image:bactrocera dorsalis squared.jpg width="289" height="292" align="center" link="@http://nathist.science.nus.edu.sg/#A/Arth-Hexa-Dipt-Acalyptratae-0041"]] || [[image:B. dorsalis size comparison.jpg width="289" height="293" align="center"]] ||
 * Scientific name: //Drosophila melanogaster// Family: Drosophilidae

Adult's appearance: Red eyes; Yellow or brown (tan) colour; Black stripes on dorsal surface of abdomen Larvae's appearance: Minute white maggots lacking legs and a defined head || Scientific name: //Bactrocera dorsalis// Family: Tephritidae Significance: Notorious pest species known for causing immense economic losses

Adult's appearance: Variable colour & pattern; prominant yellow and dark brown to black markings on dorsal surface of thorax; Generally with two horizontal black stripes and a longitudinal mark on dorsal surface of abdomen Larvae's appearance: 10 mm long; creamy white colour || //Drosophila melanogaster// size: 1-3 mm body length //Bactrocera dorsalis// size: 6-8 mm body length Credits: //Drosophila melanogaster// Information: Animal Diversity Web [8] Image: Malcolm Storey © Copyright //Bactrocera dorsalis// Information: Plant Health Australia [9] Image: Arina Adom || Figure 2: Comparison table showing differences between the Common fruit fly and Oriental fruit fly.



=**2. Biology **=

**2.1. Life History **
In tropical environments, Oriental fruit flies breed throughout the year. Throughout her lifetime (typically 1-3 months), a female fly is able to typically lay between 1,200 to 1,500 eggs (under field conditions) and over 3,000 eggs (under optimum conditions) [11].

media type="custom" key="28869074" width="306" height="306" Figure 4: Gif showing adult Oriental fruit flies on an orchid. Video credits: YouTube [25]

Eggs are laid in batches of 1-20 eggs per fruit, just under the fruit's skin, where fruit-decaying bacteria is subsequently deposited. Although ripe fruit are preferred for oviposition, oviposition on immature fruits has also been observed. Development from egg to adult is usually completed within about 16 days, however cooler weather conditions are known to delay development. Larvae (commonly known as maggots) develop within the fruit [11, 12].

. When mature, larvae emerge from the fruit and jump or drop onto the ground (watch the video below to see how the larvae jumps!).

media type="youtube" key="z1_zBuUAcPk" width="560" height="315" Video 1: Video showing jumping tephritid larvae found inside a squash. Video credits: YouTube [26]

In the soil, they form tan to dark brown puparium. Once mature, the flies emerge from their puparia as adult flies. Young males often disperse over several kilometres before reaching sexual maturity and finding its mate! For this reason, Oriental fruit flies are known to spread far and fast, causing much damage to crop fruits [11, 12]. The Oriental fruit fly life <span style="font-family: Arial,sans-serif; font-size: 13.3333px;">cycle is summarised in the following figure.

2.2. Feeding Habits
The Oriental fruit fly is highly polyphagous and is able to feed on and infest a large variety of produce from fruits, vegetables and other plants. The USDA has recorded its' occurence on 478 kinds of fruit and vegetables as of 2016 [17]. Figure 7: Table showing some of the common host plants of //B. dorsalis//.
 * * Apple
 * Apricot
 * Avocado
 * Banana
 * Bell pepper
 * Cactus
 * Cashew
 * Cherry
 * Chili
 * Citrus
 * Coffee
 * Cucumber || * Date palm
 * Fig
 * Grape
 * Grapefruit
 * Guava
 * Lemon
 * Lime
 * Loquat
 * Mandarin
 * Mango
 * Nectarine
 * Orange || * Papaya
 * Passion fruit
 * Peach
 * Pear
 * Persimmon
 * Pineapple
 * Plum
 * Pomegranate
 * Roseapple
 * Tangerine
 * Tomato
 * Walnut ||

media type="custom" key="28869036" width="298" height="298" Figure 8: Gif showing a damaged fruit infected with a //B. dorsalis// larva [28].

=3. Economic Importance= <span style="font-family: Arial,sans-serif; font-size: 10pt;">The Oriental fruit fly has been known to cause up to billions of dollars in financial losses due to __<span style="color: #0070c0; font-family: Arial,sans-serif; font-size: 10pt;">(1) __<span style="font-family: Arial,sans-serif; font-size: 10pt;"> the direct damage caused to fruit crops, __<span style="color: #0070c0; font-family: Arial,sans-serif; font-size: 10pt;">(2) __<span style="font-family: Arial,sans-serif; font-size: 10pt;"> exportation and quarantine restrictions from areas with a history of fruit fly infestations and __<span style="color: #0070c0; font-family: Arial,sans-serif; font-size: 10pt;">(3) __<span style="font-family: Arial,sans-serif; font-size: 10pt;"> management costs to eradicate existing infestations [15].

3.1. Damage to Crops
Regarded as the "most damaging pest of tropical horticulture in the world ", Oriental fruit flies are feared for their indiscriminate infestations on a wide range of produce, whether ripe or unripe. Damage caused by the fruit fly result from 1) oviposition in fruit and soft tissues of vegetative parts of certain plants, 2) feeding by the larvae, and 3) decomposition of plant tissue by invading secondary microorganisms [19]. Often, much damage can occur before any obvious signs of infestation within a fruit are even observed [12].

The first signs of an infestation are small discoloured patches on the skin fruit skin, which develop from the "sting" of the female fly during ovipositon. Once hatched, fruit fly larvae able to tunnel deeply into the fruit flesh for feeding, spreading widely within infested fruit and making the whole fruit unpalatable. In certain fruit types, maggot infestation results in tissue breakdown and internal rotting associated with maggot infestation, but this varies with the type of fruit attacked. Affected young fruit tend to become distorted, callused and usually drop prematurely whereas affected mature fruits develop a water soaked appearance. The larval tunnels subsequently provide entry points for bacteria and fungi that cause the fruit to rot further. When only a few larvae develop, damage consists of an unsightly appearance and reduced marketability because of the egg laying punctures or tissue break down due to the decay [20]. Damage to crops has been estimated at between 50-80% in West Pakistan and even up to 100% in Malaysia [21].

Watch the video below to see the impacts of Oriental fruit fly infestations on the community in Mpumalanga, South Africa.

media type="youtube" key="FEy_SN7JWbo" width="560" height="315" align="center"

Video 2: News report of the impacts of Oriental fruit fly infestations in Mpumalanga, South Africa. Video credits [27]

**3.2. Quarantine**
Due to the potential damage caused by these flies, coupled with their ability to be active dispersers and breed agressively as adults, the Oriental fruit fly is regarded as an invasive species in many countries around the world. These countries guard heavily against the introduction of the Oriental fruit fly through immigration in hopes of avoiding massive economic loss to agricultural farmers. Fresh agricultural produce being transported into each country is meticulously monitored and quarantined in order to isolate and eliminate the accidental introduction of these flies. As a result, much economic loss is caused by trade restrictions. Malaysian fruits, for example, cannot be freely exported to lucrative markets like USA, Japan and Australia, because of strict quarantine regulations in these countries, which prohibit entry of fresh fruits from the fruit flies infested countries unless proper disinfestation treatments are carried out [23]. This situation is also faced in the Philippines, Thailand, Indonesia, India, Taiwan and China.

3.3. Management
Fortunately, populations of the Oriental fruit fly are manageable and can be eradicated, albeit a huge financial cost incurred. In 1985, all Japanese territories were declared free of the Oriental fruit fly after an 18-year program of eradication combining insecticide-impregnated fiberblocks or cotton containing the powerful male attractant methyl-eugenol, and the sterile insect (sterile male) technique. Steiner traps baited with a lure and toxicant are also used to monitor the presence and control of the flies. In 1995, Oriental fruit fly established near Cairns and cost $33.5 million and took 4 years to eradicate [11].

=4. Distribution=

4.1. Worldwide distribution
Native to the tropical Asian region, the fruit flies have spread throughout the world and become a prominent pest species across multiple continents. The fly is believed to be introduced through the import of infected fruits from native regions in Asia.

4.2. Asian distribution
//Bactrocera dorsalis// is native to tropical Asian countries such as Singapore, India, Malaysia and Bangladesh. In countries such as Cambodia and China, //B. dorsalis// is regarded as an invasive species [30].

4.3. African distribution
First introduced to Africa through Kenya in 2003. Since then, most countries within sub-Saharan Africa have faced //B. dorsalis// infestations [29].

4.4. Pacific Islands distribution
//Bactrocera dorsalis// was accidentally introduced in the United States between 1944 to 1945 and is now present and a major threat on all major Hawaiian islands [19]

Not shown on map: //Bactrocera dorsalis// has also been reported in Queensland, Australia [30].

=5. Description=

5.1. Adult
The adult, which is noticeably larger than a house fly, has a body length of about 7.0-8.0 mm; the wing is about 7.3 mm in length. The color of the fly is very variable, but there are prominant yellow and dark brown to black markings on the thorax (a). Generally, the abdomen has two horizontal black stripes (b) and a longitudinal median stripe extending from the base of the third segment to the apex of the abdomen (c). These markings may form a T-shaped pattern, but the pattern varies considerably. The wings are clear (d). In females, the ovipositor is very slender and sharply pointed (e) [11].

**5.2. Larva**
The third-instar, which has a typical maggot appearance, is about 7.5-10.0 mm long and 1.5-2.0 mm wide and creamy white coloured [33]. The only band of spinules encircling the body is found on the first segment. The external part of the anterior respiratory organs, the spiracles, located one on each side of the pointed or head end of the larva, has an exaggerated and deflexed lobe at each side and bears many small tubercles. The caudal segment is very smooth. The posterior spiracles are located in the dorsal third of the segment as viewed from the rear of the larva [11].

The mature larva emerges from the fruit, drops to the ground, and forms a tan to dark brown puparium about 4.9 mm in length.

** 5.3. Puparium **
Barrel-shaped with most larval features unrecognisable. White to yellow-brown. Usually approximately 60-80% the length of the larva [33].

5.4. Egg
The white, elongate and elliptical egg measures about 1.17 x 0.21 mm and has a chorion without sculpturing.

**5.5.** Official Diagnosis
First flagellomere shorter than ptilinal fissure, face with a black spot in each antennal furrow. Tomentum pattern without longitudinal gap in the middle of prescutum. Scutum colour (other than vittae) red-brown to black. Lateral vittae of scutum present, parallel-sided, yellow, ending at or just behind intra-alar seta. Medial vitta of scutum absent. Scutellum largely yellow. Setae: anterior supra-alar present, prescutellar present, scutellar one pair. Anepisternal stripe not extended forward. Wing costal band width from vein subcostal to slightly below vein R4+5 at wing apex; confluent with vein R2+3 in depth. Cell basal costal without microtrichia. Cell costal with microtrichia in the anterodistal corner only. Cell basal radial with microtrichae at the base. Wing length: 5.4–6.9mm. Foretibia pale fuscous to dark fuscous, mid-tibia pale fuscous to fuscous, hind tibia fuscous to dark fuscous. Femora largely fulvous. Abdominal terga free, except I and II. Terga markings: terga III–V withmedial longitudinal black band, tergum III with a basal narrow transverse black band, terga IV and V with black triangular lateral markings, sometimes longitudinal black bands on lateral margins and sometimes without any mark. Tergum III (males) with pecten. Sternum V (males) with V-shaped notch. Posterior surstylus lobe short. Males attracted to methyl eugenol [34].

=<span style="font-family: Arial,sans-serif; font-size: 10pt;">**6. Taxonomy & Systematics** =

<span style="font-family: Arial,sans-serif; font-size: 10pt;">6.1. Original Description
//Bactrocera dorsalis// was first described by Friedrich Hendel as //Dacus dorsalis// in the "//Entomologischen Mitteilungen//" or "Entomological Messages" in 1912.

<span style="font-family: Arial,sans-serif; font-size: 10pt;">6.2. Type Information
<span style="font-family: Arial,sans-serif; font-size: 10pt;">The holotype of //<span style="font-family: Arial,sans-serif; font-size: 10pt;">Dacus ferrugineus //<span style="font-family: Arial,sans-serif; font-size: 10pt;"> Fabricius is currently located in the Natural History Museum of Denmark. although the specimen is nearly entirely destroyed, the taxonomically informative ‘red-brown’ colour of the thorax is still evident (See Fig.19) [31].

<span style="font-family: Arial,sans-serif; font-size: 10pt;">6.3. Common Names
<span style="font-family: Arial,sans-serif; font-size: 10pt;">English: Oriental fruit fly <span style="font-family: Arial,sans-serif; font-size: 10pt;">Malay: //lalat buah// <span style="font-family: Arial,sans-serif; font-size: 10pt;">Spanish//: mosca oriental de la fruta// <span style="font-family: Arial,sans-serif; font-size: 10pt;">French: //mouche de fruits asiatique; mouche orientale des arbres fruitiers// <span style="font-family: Arial,sans-serif; font-size: 10pt;">Portuguese: //la mosca oriental das frutas// <span style="font-family: Arial,sans-serif; font-size: 10pt;">Germany: //Orientalische Fruchtfliege// <span style="font-family: Arial,sans-serif; font-size: 10pt;">Japan: //mikan-ko-mibae// <span style="font-family: Arial,sans-serif; font-size: 10pt;">Netherlands: //mangga-vlieg//

<span style="font-family: Arial,sans-serif; font-size: 10pt;">**6.4. Scientific Name & Synonyms**
Scientific Name: //Bactrocera dorsalis// (Hendel, 1912)

Originally described by Hendel in 1912 as //Dacus dorsalis//, the Oriental fruit fly has since been redescribed, reclassified and renamed multiple times by numerous authors internationally. One major cause was the large variability in colouration and patterning, between the otherwise morphologically and genetically similar pest species within the //Bactrocera dorsalis// species complex.This resulted in the hindrance of the development of reliable diagnostic characters to distinguish between intra- and inter-specific variation. The prolonged difficulties in differentiating putative species, coupled with the highly disjunct geographic distribution of closely-related species within the //Bactrocera dorsalis// species complex, has resulted in grouping errors in the form of separate populations of //B. dorsalis// worldwide being described as new separate Bactroceran species [34].

6.4.1. Case study: //Bactrocera invadens//
One example of a grouping error was a group of //B. dorsalis// specimens detected in Kenya in 2003. After comparison with //B. dorsalis// specimens from the purported native range of Sri Lanka, the Kenyan material was described as a new species, //Bactrocera invadens// [36]. This was in spite of the considerably similar morphology of Kenyan specimens to the //B. dorsalis// specimens obtained from Sri Lanka [37]. What led to this decision was in particular, the scutum colour of the Kenyan specimens which ranged "from pale red-brown to black with the existence of variable lanceolate-patterned intermediates". This was in contrast with the //B. dorsalis// description in which the scutum had been defined as "strictly black" [4]. As a result of the confusion over the unreliable discrimination between //B. dorsalis// and //B. invadens,// significant problems with African horticulture and food security emerged, especially with regards to quarantine and pest management policies [7].

6.4.2. Final resolution
<span style="font-family: Arial,sans-serif; font-size: 10pt;">Fortunately, in 2009, a major international collaboration was established by the [|United Nations Food and Agriculture Organization] (UN-FAO) and the [|International Atomic Energy Agency] (IAEA) with the aim of resolving biological species limits among several highly morphologically and genetically similar species: //Bactrocera papayae// Drew & Hancock, //Bactrocera philippinensis// Drew & Hancock, //Bactrocera carambolae// Drew & Hancock, //Bactrocera invadens// Drew, Tsuruta & White and //Bactrocera dorsalis// (Hendel, 1912). The study, published in 2014, r<span style="font-family: Arial,sans-serif; font-size: 10pt;">esolved the problem by using multidisciplinary evidence (mo rphological, molecular, cytogenetic, behavioural and chemoecological data) <span style="font-family: Arial,sans-serif; font-size: 10pt;">to synonymise //B. dorsalis, B. invadens// and //B. papayae. Bactrocera dorsalis// (Hendel) was subsequently declared a senior synonym and a more comprehensive re-description was published to include intra-specific colour variations. The following are the synonyms of //Bactrocera dorsalis// (Hendel, 1912) listed in the paper [34].

6.4.3. Other synonyms
Figure 20: Table of //Bactrocera dorsalis// synonyms [34].
 * **Synonyms** || **Author, Year** || **Remarks** ||
 * //Musca ferruginea// || Fabricius, 1794 || Preoccupied by Musca ferruginea Scopoli, 17634 ||
 * //Dacus ferrugineus// || Fabricius, 1805 ||  ||
 * //Dacus dorsalis// || Hendel, 1912 || Lectotype ♀ in The Natural History Museum, London, U.K. ||
 * //Bactrocera ferruginea// || Bezzi, 1913 ||
 * //Chaetodacus ferrugineus// var. //dorsalis// || Hendel, 1915 ||  ||
 * //Chaetodacus ferrugineus// || Bezzi, 1916 ||  ||
 * //Chaetodacus ferrugineus dorsalis// || Bezzi, 1916 ||  ||
 * //Chaetodacus ferrugineus// var. //okinawanus// || Shiraki, 1933; Hardy & Adachi, 1956; Hardy, 1969 ||  ||
 * //Dacus (Strumeta) dorsalis// || Hardy & Adachi, 1956; Hardy, 1969; 1973; 1974 ||  ||
 * //Strumeta dorsalis// || Hering, 1956 ||  ||
 * //Strumeta ferruginea// || Hering, 1956 ||  ||
 * //Strumeta dorsalis okinawa// || Shiraki, 1968 ||  ||
 * //Dacus (Bactrocera) dorsalis// || Hardy, 1977; Drew, 1982; 1989 ||  ||
 * //Dacus (Bactrocera) semifemoralis// || Tseng et al., 1992 || synonymized by Drew & Romig, 2013 ||
 * //Bactrocera (Bactrocera) dorsalis// || Drew & Hancock, 1994; Norrbom et al., 1998; Mahmood & Hasan, 2005; White, 2006; Drew et al., 2007 || Lectotype designation by Drew & Hancock, 1994 ||
 * //Bactrocera (Bactrocera) variabilis// || Lin & Wang, 2011 in Lin et al., 2011 || Holotype in HEIQ; synonymized by Drew & Romig, 2013: 76 ||
 * //Bactrocera (Bactrocera) philippinensis// || Drew & Hancock, 1994 || synonymized with //B. papayae// by Drew & Romig, 2013 ||
 * //Bactrocera (Bactrocera) papayae// || Drew & Hancock, 1994 || syn.n. ||
 * //Bactrocera (Bactrocera) invadens// || Drew et al., 2005 || syn.n. ||

6.5. Taxonomic Hierarchy
<span style="font-family: Arial,sans-serif; font-size: 10pt;">The classification below reflects above species-level rankings (from Kingdom to Species) for //B. dorsalis//: <span style="font-family: Arial,sans-serif; font-size: 10pt;">

=7. Phylogeny= The position of the Tephritidae family within Diptera: Schizophora can be observed in Fig. 3, a subsection of the combined molecular phylogenetic tree for Diptera. The combined phylogenetic tree was produced through partitioned Maximum Likelihood (ML) analysis of data calculated in RAxML. M olecular and morphological data from 149 of 157 dipteran families, including 30 kb from 14 nuclear loci and complete mitochondrial genomes as well as 371 morphological characters were used in this analysis[10].

Within the family Tephritidae, the following phylogenetic tree has been obtained (Fig. 21) using Bayesian and ML analysis of mitochondrial DNA sequences obtained through next-generation sequencing (NGS) [40]. The tree is fairly stable with 8 out of 16 nodes within the Tephritidae tree being well supported (Bayesian posterior probabilities = 1.00 or ML bootstrap = 100 and 4 out of the remaining nodes having majority relatively high support (posterior probabilities ≥ 0.90 or ML bootstrap ≥ 70). Results show that both the family Tephritidae and the tribe Dacini (represented in the figure by genera //Dacus// and //Bactrocera//) are monophyletic. The genus //Bactrocera,// on the other hand is not monophyletic.



Within the //Bactrocera dorsalis// species complex, a separate phylogenetic study was conducted in 2013 (before //B. dorsalis// was synonymised with //B. invadens, B. papayae// and //B. philippinensis// [34]) to study the phylogeny of several //Bactrocera// species. A molecular phylogenetic tree was build with molecular data obtained from six loci (cox1, nad4-3′, CAD, period, ITS1, ITS2) for approximately 20 individuals from each of 16 sample sites. Specimens used were morphoogically sorted into the "ingroups" //B. dorsalis, B. papayae, B. philippinensis// and //B. carambolae//. Several "outgroup species" were also used in this study, some being species within the //B. dorsalis// species complex (namely the Australian species //B. cacuminata// and //B. opiliae//, and the Philippine species //B. occipitalis// (which occurs sympatrically with //B. philippinensis//) and some not being within the species complex (namely //B. musae// and //B. tryoni//). The resultant tree obtained using Bayesian and ML analysis showed the non-monophyly of //B. dorsalis, B. papayae// and //B. philippinensis// and supports the more recent synonymisation of the three species as a single species under //B. dorsalis// (Box 6 in Fig. 22). The tree also strongly supported the monophyly of //B. carambolae a//nd its' distinction from the remaining 3 ingroup species (Box 5 in Fig 22).



Thank you for reading! :)

=8. References= [1] “Oriental fruit fly (//Bactrocera dorsalis//) (Hendel, 1912)”, by Florida Division of Plant Industry, Florida Department of Agriculture and Consumer Services, Bugwood.org. URL: http://www.invasive.org (accessed on 15 Nov 2016). [2] Headrick, D. H. & R. D. Goeden, 1998. The biology of nonfrugivorous tephritid fruit flies. //Annual Review of Entomology//, 43: 217-241. [3] White, I. M. & M. M. Elson-Harris, 1992. Fruit flies of economic significance: their identification and bionomics. C.A.B International in association with ACIAR, Wallingford, Oxon. [4] Drew R. A. I. & D. L. Hancock, 1994. The //Bactrocera dorsalis// complex of fruit flies in Asia. //Bulletin of Entomological Research (Suppl. 2)//, CAB International, Wallingford, UK. [5] Drew, R.A.I.& M.C. Romig, 2013. Tropical fruit flies of South-east Asia. CAB International, Wallingford, UK. [6] Clarke, A.R., Armstrong, K.F., Carmichael, A.E. et al., 2005. Invasive phytophagous pests arising through a recent tropical evolutionary radiation: the //Bactrocera dorsalis// complex of fruit flies. //Annual Review of Entomology//, 50: 293–319. [7] Khamis, F.M., D.K. Masiga, S.A. Mohamed et al., 2012. Taxonomic identity of the invasive fruit fly pest, //Bactrocera invadens//: concordance in morphometry and DNA barcoding//. PLoS One//, 7, e44862. [8] “//Drosophila melanogaster//,” by Conrad Miller. Animal Diversity Web. URL: http://animaldiversity.org/accounts/Drosophila_melanogaster/ (accessed on 14 Nov 2016). [9] “Oriental fruit fly complex – Fact sheet,” by Plant Health Australia. URL: http://www.planthealthaustralia.com.au/wp-content/uploads/2013/09/Oriental-fruit-fly-complex-FS-Papaya.pdf (accessed on 14 Nov 2016). [10] Wiegmann, B. N., M. D. Trautwein, I. S. Winkler et al., 2011. Episodic radiations in the fly tree of life. //Proceedings of the National Academy of Sciences//. Available from: https://www.researchgate.net/50395098_fig1_Fig-1-Combined-molecular-phylogenetic-tree-for-Diptera-Partitioned-ML-analysis-of (accessed on 22 Nov 2016). [11] “oriental fruit fly,” by H.V. Weems, J.B. Heppner, James L. Nation & Gary Steck. University of Florida, Florida Department of Agriculture and Consumer Services, Aug 2016. URL: [] (accessed on 14 Nov 2016). [12] “Oriental fruit fly,” byDepartment of Agriculture and Fisheries, Queensland Government, 19 Feb 2015. URL: [] (accessed on 14 Nov 2016). [13] Daniel, C. & J.Grunder, 2012. Integrated Management of European Cherry Fruit Fly //Rhagoletis cerasi// (L.): Situation in Switzerland and Europe, “//Pupae of R. cerasi © 2012//”. //Insects//, 3(4): 956-988 URL: [](accessed on 22 Nov 2016). [14] “A female Oriental fruit fly (//Bactrocera dorsalis//) lays eggs by inserting her ovipositor in the skin of a papaya on Dec. 27, 2011,” U.S. Department of Agriculture (USDA) photo by Scott Bauer. Flickr.com, 27 Dec 2011. URL: @https://flic.kr/p/dQtRto (accessed on 14 Nov 2016). (permission pending) [15] “Oriental fruit fly larvae” U.S. Department of Agriculture (USDA) photo by Scott Bauer. Flickr.com, 05 March 2012. URL: [] (accessed on 14 Nov 2016). (permission pending) [16] “//Bactrocera zonata// eggs” by Russell IPM. URL: http://russellipm-agriculture.com/en/insect/bactrocera-zonata (accessed on 22 Nov 2016). [17] USDA, 22 July 2016. A Review of Recorded Host Plants of Oriental Fruit Fly, // Bactrocera dorsalis // (Hendel) (Diptera: Tephritidae). Version 2.1 // A Product of the USDA Compendium of Fruit Fly Host Information (CoFFHI), a Farm Bill Project //. [18] "Oriental Fruit Fly FACT SHEET," by California Department of Food & Agriculture. CDFA, 25 Aug 2008. URL: https://www.cdfa.ca.gov/plant/factsheets/OFF_FactSheet.pdf (accessed on 15 Nov 2016). [19] “Bactrocera dorsalis (Hendel),“ by Ronald F.L. Mau, Jayma L. Matin & J. M. Diez. //Crop Knowledge Master,// Apr 2007, URL: [] (accessed on 15 Nov 2016). [20] Steiner, L. F., 1957. Field Evaluation of Oriental Fruit Fly Insecticides in Hawaii. // Journal of Economic Entomology //, 50: 16-24. [21] Mahmood, K., 2004. Identification of Pest Species in Oriental Fruit Fly, // Bactrocera dorsalis // (Hendel) (Diptera: Tephritidae) Species Complex. // Pakistan Journal of Zoology, // 36(3): 219-230. [22] Bhagat, D., Samanta, S. K. & Bhattacharya, S., 2013. Efficient Management of Fruit Pests by Pheromone Nanogels. //Scientific Reports 3//, 1294. doi:10.1038/srep01294 [23] Singh, R.B., 1991. Significance of fruit flies in fruit and vegetable production in the Asia-Pacific region. Proceedings First International Symposium on Fruit flies in the Tropics (eds. S. Vijaysegaran and A.G. Ibrahim), pp. 11- 29. Kuala Lumpur, 1988. Malaysian Agricultural Research and Development Institute, Kuala Lumpur. [24] “Florida growers, Adam Putnam debate aerial pest spray for invasive Oriental fruit fly,” by Alex Harris. Miami Herald, 21 Sep 2015. Hoested on MiamiHerald.com: [](accessed on 21 Nov 2016). [25] "Fruit flies, Oriental Mindoro, Philippines," by [|warrenlaurde]. YouTube, 06 Jun 2011. Gif made using gifyoutube.com. [26] “Jumping Tephritidae fruit fly maggots,” by [|hillbournesian]. Youtube, 05 Oct 2011. [27] “Mpumalanga citrus farmers hit hard by Oriental Fruit Flies,” by SABC Digital News. SABC YouTube Channel, 08 Apr 2016. [28] “Alerta Verde: La Mosca Oriental de la Fruta,” by DaniSilva TV. Youtube, 21 Sep 2015. Gif made using gifyoutube.com. [29] Goergen, G., J. F. Vayssières, D. Gnanvossou & M. Tindo, 2011. // Bactrocera invadens // (Diptera: Tephritidae), a new invasive fruit fly pest for the Afrotropical Region: host plant range and distribution in West and Central Africa. // Environmental Entomology. // 40: 844-854 [30] EPPO (2015) PQR - EPPO database on quarantine pests (available online). http://www.eppo.int [31] “Holotype of //Dacus ferrugineus// Fabricius located in the Natural History Museum of Denmark,” by Verner Michelsen, Oct 2014. One and the same: Integrative taxonomic evidence that Bactrocera invadens (Diptera: Tephritidae) is the same species as the Oriental fruit fly Bactrocera dorsalis - Scientific Figure on ResearchGate. URL: https://www.researchgate.net/267761592_fig3_Figure-7-Holotype-of-Dacus-ferrugineus-Fabricius-located-in-the-Natural-History-Museum (accessed on 23 Nov 2016). [32] Bactrocera dorsalis (Oriental fruit fly) Datasheet,” by Centre for Agriculture and Biosciences International. CABI.org, 23 Sep 2016. URL: [] (accessed 14 Nov 2016). [33] White, I.M., Elson-Harris, M.M., 1994. Fruit Flies of Economic Significance. Their Identification and Bionomics. Wallingford, UK: CAB International. [34] Schutze, M. K., N. Aketarawong, W. Amornsak et al., 2014. Synonymization of key pest species within the //Bactrocera dorsalis// species complex (Diptera: Tephritidae): Taxonomic changes based on a review of 20 years of integrative morphological, molecular, cytogenetic, behavioural and chemoecological data. //Systematic Entomology//, DOI: 10.1111/syen.12113. [35] “Original description of //Dacus dorsalis// in //<span style="font-family: Arial,sans-serif; font-size: 10pt;">Entomologischen Mitteilungen,” ////<span style="font-family: Arial,sans-serif; font-size: 10pt;">by //<span style="font-family: Arial,sans-serif; font-size: 10pt;">Friedrich Hendel, 1912. Image obtained from Biodiversity Heritage Library (BHL). URL: [] (accessed 14 Nov 2016). [36] Lux, S.A., R.S. Copeland, I.M. White et al., 2003. A new invasive fruit fly species from the Bactrocera dorsalis (Hendel) group detected in East Africa. //Insect Science and its Application//, 23, 355–361. [37] Drew, R.A.I., K. Tsuruta & I.M. White, 2005 A new species of pest fruit fly (Diptera: Tephritidae: Dacinae) from Sri Lanka and Africa. // African Entomology //, 13, 149–154. [38] Retrieved [14 Nov 2016], from the UniProt Consortium, UniProt: a hub for protein information, Nucleic Acids Res, []. [39] Retrieved [14 Nov 2016], from the Integrated Taxonomic Information System on-line database, http://www.itis.gov. [40] Jiang F., X. Pan, X. Li et al., 2016. The first complete mitochondrial genome of // Dacus longicornis // (Diptera: Tephritidae) using next-generation sequencing and mitochondrial genome phylogeny of Dacini tribe. // Scientific Reports //, 6. [41] Boykin, L. M., M. K. Schutze, M. N. Krosch et al., 2014. Multi-gene phylogenetic analysis of south-east Asian pest members of the //Bactrocera dorsalis// species complex (Diptera: Tephritidae) does not support current taxonomy. //Journal of Applied Entomology//, 138: 235–253. doi:10.1111/jen.12047 [42] Hardy, D.E., 1969. Taxonomy and distribution of the Oriental fruit fly and related species (Tephritidae-Diptera). //Proceedings of the Hawaiian Entomological Society//, 20, 395–428.