Review Article

An Overview of Indian Dacine Fruit Flies (Diptera: Tephritidae: Dacinae: Dacini)

A. Vasudha, Md. Abbas Ahmad and M. L. Agarwal

  • Page No:  491 - 506
  • Published online: 23 Oct 2019
  • DOI : HTTPS://DOI.ORG/10.23910/IJBSM/2019.10.5.2016

  • Abstract
  •  mlagento@yahoo.com

Fruit flies belonging to tribe Dacini are pertinent pests of horticultural crops in Southeast Asia, Sub-Saharan Africa and Australasian Oceanian region. They pose enormous threats to fruit and vegetable production throughout the world, causing both quantitative and qualitative losses. Ninety two species of dacine fruit flies (28 endemic, 30.43% endemism) belonging to three genera, viz. Bactrocera Macquart (51 species, 17 endemic, 33.33% endemism), Dacus Fabricius (10 species, 3 endemic, 30.0% endemism), and Zeugodacus Hendel (31 species, 8 endemic, 25.80% endemism), and 19 subgenera have been listed from India. Of these, nine endemic species, viz. Bactrocera (Bactrocera) andamanensis, B. (B.) blairiae, B. (B.) curtivitta, B. (B.) patula, B. (B.) ranganathi, B. (Calodacus) harrietensis, Dacus (Mellesis) insulosus, Zeugodacus (Zeugodacus) fuscoalatus, Z. (Z.) havelockiae are known only from Andaman & Nicobar Islands. The current understanding of the morphology of the tribe, together with data on their biology, habitat associations, and distribution combine presents a picture of the phylogenetic relationships within the group. Besides, the knowledge of biology and behaviour of dacine species suggests that there is a close relationship between the fly species and their host plants which involves adult’s feeding, host recognition, courtship and mating, oviposition, larval feeding and bacteria associations. In the present paper notes on subgeneric classification of Dacini, their distribution, endemism, relative occurrence of different taxa, phylogeny, host associations, pest status, male lures and bacterial associations have been added.

Keywords :   Dacini, phylogeny, distribution, host associations, male lures, bacterial associations

  • Introduction

    Fruit flies are distributed in all biogeographic regions except in extreme desert and polar areas, where their hosts are scarce or absent (Foote et al., 1993). They have highly diverse biology and larvae of many species develop in fruits, vegetables, flower heads, buds, seeds, stems, etc. Fruit flies belonging to tribe Dacini are of greatest horticultural relevance and its most species are phytophagous and many pertinent pest species have been studied extensively, due to damage they cause in plants of economic interest (Norrbom et al., 1999). Many invasive dacine flies are pests of horticultural crops worldwide, due to their wide climatic tolerance, polyphagous nature, high reproduction potential, multivoltine nature and high capacity for dispersal (Prokopy, 1977). Tribe Dacini contains around a fifth of all known species in the family Tephritidae (Schutze et al., 2017). These flies are frugivorous or florivorous and about 10 per cent of the recognized species are pests of commercial fruits and vegetables (Vargas et al., 2015). They use host’s tissues for larval development causing severe economic impact and liable to quarantine restrictions imposed by many countries (Foote, 1967; Aluja and Mangan, 2008). In India, fruit flies have been identified as one of the ten most serious problems of horticulture because of their polyphagous nature and cause a huge economic loss which varies from 2.5-100% depending upon the crop and season (Verghese et al., 2004; Dhillon et al., 2005). With respect to the larval host plants, most species are polyphagous; a few are oligophagous, while the remaining <1% are monophagous (Drew, 1989).  Besides this, global invasion of fruit flies attracts a great deal of attention in the field of plant quarantine and invasion biology (Qin et al., 2015). India is a major producer of fruits and vegetables and with increasing globalization; it has become a challenge for this country not only to feed its own population but also to export fruits and vegetables to developed countries.


  • Tribe Dacini

    The genus Dacus was first described by Fabricius in 1805 by its type-species D. armatus. Flies of tribe Dacini are often wasp mimics, usually black to brown with yellow vittae and predominantly hyaline wing with well developed costal band and anal streak (David and Ramani, 2019). Presently four genera, viz. Bactrocera Macquart, Dacus Fabricius, Monacrostichus Bezzi and Zeugodacus Hendel and 932 species are recognized in this tribe (Virgilio et al., 2015; San Jose et al., 2018; Doorenweerd et al., 2018). Drew and Romig (2013, 2016) considered Ichneumonopsis to be a member of Dacini, whereas Kovac et al. (2006, 2013) and Freidberg et al. (2017) included this genus in tribe Gastrozonini. Earlier 85 species of Dacini were documented from India in 3 genera except for Monacrostichus (Agarwal and Sueyoshi, 2005; David and Ramani, 2011; Drew and Romig, 2013; David et al., 2017). The dacine flies are characterized by scutum black/ brown/ reddish-brown with or without yellow medial and lateral postsutural vittae; significantly reduced chaetotaxy of head and thorax. Face fulvous to black with a pair of dark spots or a band; wing with cell bm deeper/broader than bcu; extension of cell bcu longer than bcu; costal band vary in width and may expand at apex in small or large spot; males usually with pecten; ceromata present, female with 2 spermathecae.


  • Generic and Subgeneric Classification

    Tribe Daciniis one of the most species rich clades within the family Tephritidae and includes 4 genera, viz. Bactrocera, Dacus, Zeugodacus and Monacrostichus. These genera are well distributed in and around Indian subcontinent and extending their range to Pacific Australasian countries, while greatest diversity of genus Dacus occurs in Afrotropical region. Shiraki (1933) discussed the generic classification of the Dacini and provided a key to 6 genera, viz. Chaetodacus Bezzi, Zeugodacus Hendel, Parazeugodacus Shiraki, Paratridacus Shiraki, Tetradacus Miyake and Mellesis Bezzi based on characters like presence or absence of supernumerary lobe and dense aggregation of microtrichia around vein CuA + 1A in male wing; presence or absence of bristles and a brown anal stripe in wing; presence or absence of a pecten on 3rd abdominal tergite of male and shape of basal segment of ovipositor. Munro (1947) recognized only one genus Dacusand divided it into a number of subgenera. Hardy (1955) arranged Dacini species in 4 genera and 24 subgenera by considering groups of characters. Drew (1972) recognized two valuable characters in showing intra-tribe relationships, i.e. (1) Abdominal sternite V of male– posterior margin with a deep concavity or a very slight concavity. (2) size of male surstylus - either short-at most only twice as long as anterior lobeor long-at least 6times as long as anterior lobe. He divided genus Dacus into Dacus group and Strumeta group of subgenera on the basis of these characters.

    Munro (1984) elevated subfamily Dacinae to full family status namely Dacidae for African taxa and described many new subcategories in the family; however, his classification was not accepted by most tephritid taxonomists. Drew (1989) divided Dacini into four groups, viz.Bactrocera group, Zeugodacus group, Melanodacus group Queenslandacus group,and 28 Asian and Pacific subgenera. Drew and Hancock (1999) recognized four genera in the tribe: Bactrocera, Dacus, Ichneumonopsis, and Monacrostichus and placed majority of species of the first two genera in different subgenera. Agarwal (2006) placed Indian Dacini species in two groups Bactrocera group and Zeugodacus group; and 10 subgenera, namely - Bactrocera, Bulladacus, Daculus, Gymnodacus, Tetradacus, Hemigymnodacus, Javadacus, Paradacuss, Paratridacus and Zeugodacus. Hancock and Drew (2015) listed subgenera in genus Bactrocera in four groups namely Bactrocera Group, Melanodacus Group, Tetradacus – ancestral subgenus?, and Zeugodacus Groups. Hancock and Drew (2018) listed subgenera in the Zeugodacus group as: Subgroup (1) - Aglaodacus, Heminotodacus, Nesodacus, Paradacus, Parasinodacus, Perkinsidacus, Subgroup (2) - Asiadacus, Austrodacus, Diplodacus, Javadacus, Niuginidacus, Papuodacus, Sinodacus, Zeugodacus.


  • Distribution and Relative Occurrence of Different Taxa

    Most Dacini are found in Afrotropical region, Southeast Asia to Australasian Oceanian region and only a few have invaded in other areas. The Asian-Pacific Dacini, primarily consisting of Bactrocera and Zeugodacus species mainly occur in South East Asia and Papua New Guinea (Drew, 2004). The distributions of known species within the tribe Dacini in India is given in Table 1.


    In India, Dacini is represented by 3 genera Bactrocera, Dacus and Zeugodacus and 92 species (28 endemic, i.e. 30.43% endemism). The genus Bactrocera includes (51 species, 17 endemic – 33.33% endemism), Dacus (10 species, 3 endemic - 30% endemism) and Zeugodacus (31 species, 8 endemic – 25.80% endemism) (Figure 1, Table 1). The relative occurrence of species in each genus and subgenus is depicted in Figure 2.


    Nine species of Indian dacine flies (9.78%) are described only from Andaman and Nicobar Islands (Table 1). Dacus and Zeugodacus species namely, D. longicornis, Z. caudatus, Z. diversus, Z. scutellaris and Z. tau andare well distributed in most countries of Southeast Asia. Indian dacine species also show relationship with the corresponding fauna of other regions. B. (Daculus) oleae, an African species spread to Mediterranean region and northwest India and Pakistan. The genus Dacus is known by 10 species in 5 subgenera, of which, D. craboniformis, D. icariiformis and D. insulosus are endemic. Two subgenera of African origin, Didacus and Leptoxyda, have radiated back to Asia with the occurrence of D. (Didacus) ciliatus (cucurbit feeders) and D. (Leptoxyda) persicus (in asclepiads) (Drew et al., 1998). B. carambolae has introduced into Neotropical region. Species namely, B. albistrigata, B. correcta, B. carambolae, B. dorsalis, B. latifrons, B. oleae, B. zonata, B. calophylli, D. ciliatus, D. persicus, and Z. cucurbitae have also been recorded/ intercepted/ eradicated (not established) from other zoogeographical areas (Agarwal, 2006).


  • Host Associations

    Agarwal (2019) listed 296 species of fruit flies belonging to 90 genera from India. Species belonging to the genera Bactrocera and Zeugodacus are most economically significant with at least 50 polyphagous species considered to be important pests in tropical and subtropical areas of the world (White and Elson-Harris, 1992). The adults of polyphagous species have high mobility, relatively long life span (often more than 3 months) and high fecundity (> 1000 eggs/female) (Vargas et al., 1984), scramble type competition in the larval stage, several generations per year and the ability to pass unfavourable periods in a facultative reproductive diapause (Fletcher, 1987). Adult dacines mostly feed on plant secretions, nectar, sap, honey dew, bird dropping and microorganisms (Christenson and Foote, 1960) and direct damage begins with female puncturing the host’s skin and ovipositing underneath it. During oviposition, fruit-rottening bacteria from the intestinal flora of the fly are introduced into the host which multiply and cause its tissues to rot (Vayssières et al., 2009). The larvae make feeding galleries resulting in conversion of host’s tissues in a spongy mess. The second and especially third instar stages are voracious feeders and generally, the fruit falls to the ground as, or just before the maggots pupate (Ekesi and Billah, 2006). The larvae feeding forms a “non-interactive grazing system” while other major class of food substrates constitute plant parts, e.g. shoots, flowers, roots and species utilizing such food operate in an “interactive grazing system” (Zwolfer, 1983).

    Host utilization by fruit fly species, whether mono-, oligo-, or polyphagous, must depend on adult’s choice in terms of attraction to the host for oviposition. Female uses visual cues such as plant colour and odours emitted by a host (mainly but not exclusively host’s specific).  Light and Jang (1996) noted that there are 3 types of volatiles that can draw females (or males) to a particular habitat, “green leaf volatiles” such as aliphatic aldehydes and alcohols emitted by leaves and unripe fruit; (b) volatiles, mainly esters, emitted by ripening fruit; and (c) volatiles emanating from rotting fruit, bacteria, and other food sources, and species-specific volatiles emitted by the hosts of specialized fruit fly species. After alighting on a host, the female assesses its surface texture and chemical properties with tarsi and decide to bore or not. Finally, sensors at the tip of the aculeus send the last series of signals, allowing the female to reject the fruit or to accept it and lay eggs (Rice, 1989).


  • Phylogeny

    Dacini is a tropical and subtropical evolutionary radiation of flies with centers of diversity in Southeast Asia and Sub-Saharan Africa. Early molecular phylogenetic studies focussed on pest species, often of a particular region, leading to biased results on the relationships between species that may not precisely reflect monophyletic origins or sister-group assignments (Smith et al., 2003; Nakahara and Muraji, 2008). Further phylogenetic studies molecular data resulted into splitting of large genus Bactrocera into Bactrocera and Zeugodacus (Krosch et al., 2012; Virgilio et al., 2015; San Jose et al., 2018). Besides, the Dacini taxa have been variably assigned to species complexes, species groups, subgenera and species-complex groups (Clarke et al., 2005; White, 2006; Drew and Romig, 2013). The largest and most intensively studied is the Bactrocera dorsalis complex with 88 species includes the largest number of pest species (Doorenweerd et al., 2018). This complex, like most others, is not monophyletic (Leblanc et al., 2015; Virgilio et al., 2015; San Jose et al., 2018). For Southeast Asian Dacini recent two-part work including a revision (Drew and Romig, 2013) and the accompanying keys (Drew and Romig, 2016) are pertinent. For other regions, all treatments are older and with perplexity.

    Phylogenetic studies on Dacini using morphological characters are few except for the work by White (1999), wherein 51 pest species were analysed using 38 morphological characters. His studies revealed Monacrostichus as a sister group to Bactrocera and Dacus as that of Drew and Hancock (1999). A subgeneric phylogeny of genus Dacus using morphological characters was provided by Hancock and Drew (2006). Phylogenetic studies on Dacini using molecular markers were explored by several researchers: Smith et al. (2002) confirmed the monophyly of Dacus and Bactrocera. Smith et al. (2003) also reported monophyly of Bactrocera and noted the paraphyletic status of Zeugodacus in their analysis. White (2006) also indicated a sister group relationship between Zeugodacus group and genus Dacus. Han and Ro (2009) reconfirmed monophyly of subfamily Dacinae using 12s, 16s and COII gene fragments. Asokan et al. (2011) and Yong et al. (2015) revealed monophyly of Bactrocera using COI and 13 protein coding genes, respectively.

    Krosch et al. (2012) concluded that genus Bactrocera consists of 2 clades, Bactrocera s.s. and Zeugodacus group of subgenus. They considered Zeugodacus clade, a sister group of Dacus and recommended to raise subgenus Zeugodacus to genus level. Virgilio et al. (2015) raised the subgenus Bactrocera (Zeugodacus) to generic rank (Zeugodacus Hendel, stat. nov.) and placed all species of subgenus (Zeugodacus) in the genus Zeugodacus; however, they concluded that exact relationship between Zeugodacus, Dacus and Bactrocera still needs to be resolved. San Jose et al. (2018) also confirmed the monophyly of genera Bactrocera, Dacus and Zeugodacus based on 7 genes in 167 species of the tribe Dacini. David and Ramani (2019) analysed phylogenetic relationships between Bactrocera, Dacus and Zeugodacus from India based on morphological characters. Cladistic analysis revealed the monophyly of Dacini, Bactrocera and Dacus with supporting non-homplasious synapomorphies. Zeugodacus was retrieved as a monophyletic sister-group to Dacus.


  • Male Lures

    Chemical cues and signals influence the behavior, physiology, and ecology of fruit flies and lures are used for surveillance, suppression, and ecological studies. Males of many dacine fruit flies are attracted to plant-derived secondary compounds termed male lures (Sivinski and Calkins, 1986). Howlett (1912) reported existence of fruit fly male lures in citronella (Cymbopogon nardus, Fam. Poaceae) oil. Howlett (1915) reported that the attractive component was phenyl proponoid methyl eugenol (ME) or 3-4 dimethoxy-1 allylbenzene and its effectiveness was rediscovered by Steiner (1952). Barthel et al. (1957) observed anisyl acetone or 4(p-methoxyphenyl)-2-butanone as an effective attractant for the melon fly. A derivative cue-lure (CL) or 4(p-acetoxyphenyl)-2-butanone was found to be more attractive to some dacine species (Beroza et al., 1960). Drew and Hooper (1981) reported that each dacine species responded only to one of these attractants and some species did not respond to either. However, several studies (Tan and Nishida, 1996; Tan et al., 2011) have demonstrated that certain male lures (e.g., methyl eugenol, raspberry ketone, and zingerone) are used in synthesizing male sex pheromones. ME is a widely distributed natural plant product that occurs in >450 plant species in 80 families found mainly in the tropics (Tan and Nishida, 2012). CL has not been isolated as a natural product but is rapidly hydrolyzed to form 4-(p-hydroxyphenyl)-2-butanone, rheosimin, or raspberry ketone (RK), a constituent of raspberries (Rubus idaeus and R. strigosus, Fam. Rosaceae) with a raspberry-like odour (Metcalf and Metcalf, 1992). RK was originally isolated from an orchid, Dendrobium superbum (Nishida et al., 1993).

    Of the 86 Dacini species that are horticultural pests, 41 respond to CL/RK and 18 to ME (Dacine Fruit Flies of the Asia-Pacific website, 2012). Based on this attraction, detection and monitoring traps and the suppression/eradication technique called male annihilation technique (MAT) was developed using these chemicals. Vargas et al. (2014) summarized future trends for use of male lures (Figure 3), such as the use of reduced risk insecticides, new lures, lure mixtures, and new dispenser formulations.


    Among Indian Dacini males of 47 species respond to cue-lure, 20 species to methyl eugenol, 6 species to zingerone while B. latifrons males to latilure (Table 1). These attractants are also used in the surveillance system targeting more than one species at a time and are a powerful monitoring tool for the early detection of a species and population monitoring. Stringer et al. (2019) reported that in fruit fly management using various lure combinations reduces the cost of operation. Recently in Oceania and Asia, more attractive male lures (isoeugenol, methylisoeugenol, dihydroeugenol, and zingerone) were identified for several weakly CL- and methyl eugenol (ME) responsive species (Royer et al., 2019).


  • Bacterial Associations

    Tephritid flies harbour communities of bacteria dominated by species of Enterobacteriaceae. These microbes are involved in nitrogen fixation, reproductive success, temporal host range expansion, protection from pathogens and detoxification (Akami et al., 2019). Petri (1909) first observed symbiotic association with bacteria in olive fly, B. oleae. The attractancy of protein solutions containing bacteria to fruit flies was first reported by Gow (1954). Cultures of fruit fly type bacteria growing on peptone yeast extract agar (Drew et al., 1983) and hydrolyzed protein solutions inoculated with these bacteria are strong attractants for dacine species (Drew and Fay 1988). These bacteria provide nutrients for adult females and possibly larvae as a food substrate, olfactory cue to attract flies to the host plant, lure flies to the plant to ensure courtship and mating, and may play a role in the fly defense mechanisms against bacterial pathogens such as Serratia species. Krischik and Jones (1991) defined the bacteria associated with dacine fruit flies as insect mutualists, not symbionts and stated that the bacteria beneficially affect the capacity of the fly to explore the plant, and in turn the microorganism is affected by the insect-plant interaction.

    In dacine flies bacterial mediation is hypothesized as being integral to the larval host plant being the ‘centre of activity’ of the fly (Raghu et al., 2002). The role of bacteria as a food source for adult fruit flies and how they affect their behaviour and fitness have been studied extensively by Drew et al., 1983; Drew and Lloyd, 1987, 1989, 1990; Drew, 1987 and by Fitt and O’Brien (1985), who reported that some bacteria found on ripening fruits also exist in the digestive tract of flies and that females transmit these bacteria to their offspring during oviposition. Gujjar et al. (2017) attempted to decipher the gender specificity of gut bacterial communities of two major fruit fly species of India and based on molecular identification, B. dorsalis females were found to predominantly harbor the bacterial species Enterobacter cloacae, E. asburiae and Citrobacter freundii, while B. dorsalis males were found to harbor Providencia rettgerii, Klebsiella oxytoca, Enterococcus faecalis and Pseudomonas aeruginosa. The cultivable diversity from females of Z. cucurbitae comprised mainly of Morganella morganii and Bacillus pumilis while Z. cucurbitae males were predominantly colonized by aerobic endospore formers, viz. Bacillus cereus, B. licheniformis and B. subtilis.


  • Conclusion

    The Indian Dacini fauna comprises of 92 species belonging to 3 genera and 19 subgenera. Efforts are in vogue to manage dacine fruit flies by using male lures which have been found of profound significance in monitoring, suppression and population eradication programmes. However, there is ample scope for further researches to study the molecular characterization of Indian dacine flies and to discover new attractants, particularly for females. Efforts should also be made by quarantine authorities to remain aware so that invasive pest species may not enter in India.


  • Reference
  • Agarwal, M.L., 2006. Indian Dacini (Diptera: Tephritidae) and their management. In: Dwivedi, S.C., Dwivedi N. (Eds.), Integrated Pest Management and Biocontrol. Pointers Publ., Jaipur, 353–373.
    Agarwal, M.L., 2019. A checklist of Indian fruit flies (Diptera: Tephritidae) with notes on some species. Journal of the Bombay Natural History Society, in press.
    Agarwal, M.L., Sueyoshi, M., 2005. Catalogue of Indian fruit flies (Diptera: Tephritidae). Oriental Insects 39, 371-433. Available from: https://www.tandfonline.com/doi/pdf/10.1080/00305316.2005.10417450
    Akami, M., Andongma, A.A., Zhengzhong, C., Nan, J., Khaeso, K., Jurkevitch, E., Niu, C.-Y., Yuval, B., 2019. Intestinal bacteria modulate the foraging behavior of the oriental fruit fly Bactrocera dorsalis (Diptera: Tephritidae). PLoS ONE 14(1), e0210109. Available from: https://doi.org/10.1371/journal.pone.0210109
    Aluja, M., Mangan, R.L., 2008. Fruit fly (Diptera: Tephritidae) host status determination: critical conceptual, methodological, and regulatory considerations. Annual Review of Entomology 53, 473-502. Available from: http://dx.doi.org/10.1146/annurev.ento.53.103106.093350
    Asokan, R., Rebijith, K.B., Singh, S.K., Sidhu, A.S., Siddharthan, S., Karanth, P.K., Ellango, R., Ramamurthy, V.V., 2011. Molecular identification and phylogeny of Bactrocera species (Diptera: Tephritidae). Florida Entomologist 94(4), 1026–1035.
    Barthel, W.F., Green, N., Keiser, I., Steiner, L.F., 1957. Anisyl acetone, synthetic attractant for male melon fly. Science 126, 654.
    Beroza, M., Alexander, B.H., Steiner, L.F., Mitchell, W.C., Miyashita, D.H., 1960. New synthetic lures for the male melon flies. Science 131, 1044–1045.
    Christenson, L.D., Foote, R.H., 1960. Biology of fruit flies. Annual Review of Entomology 5, 171–192.
    Clarke, A.R., Armstrong, K.F., Carmichael, A.E., Milne, J.R., Raghu, S., Roderick, G.K., Yeates, D.K., 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. Available from: https://doi.org/10.1146/annurev.ento.50.071803.130428
    Dacine Fruit Flies of the Asia-Pacific Website, 2012. University of Hawaii at Manoa. Available from:  http://www. herbarium.hawaii.edu/fruitfly/index.php.
    David, K.J., Ramani, S., 2011. An illustrated key to fruit flies (Diptera: Tephritidae) from Peninsular India and the Andaman and Nicobar Islands. Zootaxa 3021, 1–31.
    David, K.J., Ramani, S., 2019. New species, redescriptions and phylogenetic revision of tribe Dacini (Diptera: Tephritidae: Dacinae) from India based on morphological characters. Zootaxa 4551(2), 101–146.
    David, K.J., Hancock, D.L., Singh, S.K., Ramani, S., Behere, G.T., Salini, S., 2017. New species, new records and updated subgeneric key of Bactrocera Macquart (Diptera: Tephritidae: Dacinae: Dacini) from India. Zootaxa 4272(3), 386-400. Available from: https://doi.org/10.11646/zootaxa.4272.3.4
    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. 16. Journal of Insect Science 5, 40. Available from: insectscience.org/5.40.
    Doorenweerd, C., Leblanc, L., Norrbom, A.L., San Jose, M., Rubinoff, D., 2018. A global checklist of the 932 fruit fly species in the tribe Dacini (Diptera, Tephritidae). ZooKeys 730, 17-54. Available from: https://zookeys.pensoft.net/article/21786/
    Drew, R.A.I., 1972. The generic and subgeneric classification of Dacini (Diptera: Tephritidae) from the South Pacific area. Journal of the Australian Entomological Society 11, 1-22.
    Drew, R.A.I., 2004. Biogeography and Speciation in the Dacini (Diptera: Tephritidae: Dacinae). In: Evenhuis, N.L., Kaneshiro, K.Y. (Eds.), D. Elmo Hardy Memorial Volume. Contributions to the Systematics and Evolution of Diptera. Bishop Museum Bulletin in Entomology 12, 165–178.
    Drew, R.A.I., Fay, H.A., 1988. Elucidation of the role of ammonia and bacteria in the attraction of Dacus tryoni (Froggatt) (Queensland fruit fly) to proteinaceous suspensions. Journal of Plant Protection in the Tropics 5(1), 127–130.
    Drew, R.A.I., Hancock, D.L., 1999. Phylogeny of the Tribe Dacini (Dacinae) based on morphological, distributional, and biological data. In: Aluja, M., Norrbom, A.L. (Eds.), Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, 491–504.
    Drew, R.A.I., Hooper, G.H.S., 1981. The responses of fruit fly species (Diptera: Tephritidae) in Australia to various attractants. Journal of the Australian Entomological Society 20, 201–205.
    Drew, R.A.I., Lloyd, A.C., 1987. Relationship of fruit flies (Diptera: Tephritidae) and their bacteria to host plants. Annals of the Entomological Society of America 80, 629–636.
    Drew, R.A.I., Lloyd, A.C., 1989. Bacteria associated with fruit flies and their host plants. In: Robinson, A.S., Hooper, G. (Eds.), Fruit Flies: Their Biology, Natural Enemies and Control. World Crop Pests Ser. Vol. 3A, Elsevier, Amsterdam, 131–140.
    Drew, R.A.I., Lloyd, A.C., 1990. The role of bacteria in the life cycle of tephritid fruit flies. In: Barbosa, P., Krischik, V., Jones, C.L. (Eds.), Microbial Mediation of Plant-Herbivore Interactions. Wiley, NY, 441–465.
    Drew, R.A.I., Romig, M.C., 2013. Tropical Fruit Flies (Tephritidae: Dacinae) of South-East Asia. CAB International, Wallingford, 653.
    Drew, R.A.I., Romig, M.C., 2016. Keys to The Tropical Fruit Flies (Tephritidae: Dacinae) of South-East Asia. CAB International, Wallingford, 487.
    Drew, R.A.I., Courtice, A.C., Teakle, D.S., 1983. Bacteria as a natural source of food for fruit flies (Diptera: Tephritidae). Oecologia 60(3), 279–284.
    Drew, R.A.I., Hancock, D.L., White, I.M., 1998. Revision of the tropical fruit flies (Diptera: Tephritidae: Dacinae) of South-east Asia. II. Dacus Fabricius. Invertebrate Taxonomy 12, 567–654.
    Ekesi, S., Billah, M.K., 2006. Field Guide to the Management of Economically Important Tephritid Fruit Flies in Africa. ICIPE Science Press, Nairobi, Kenya, 160.
    Fitt, G.P., O’Brien, R.W., 1985. Bacteria associated with four species of Dacus (Diptera: Tephritidae) and their role in the nutrition of the larvae. Oecologia 67, 447–454.
    Fletcher, B.S., 1987. The biology of dacine fruit flies. Annual Review of Entomology 32, 115–144.
    Foote, R.H., 1967. 57. Family Tephritidae (Trypetidae, Trupaneidae). In: Papavero, N. (Ed.), A Catalogue of the Diptera of the Americas South of the United States. Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo, 91.
    Foote, R.H., Blanc, F.L., Norrbom, A.L., 1993. Handbook of the Fruit Flies (Diptera: Tephritidae) of America and North of Mexico. Comstock Publishing Associates, Ithaca, 571.
    Freidberg, A., Kovac, D., Shiao, S.F., 2017. A revision of Ichneumonopsis Hardy, 1973 (Diptera: Tephritidae: Dacinae: Gastrozonini), Oriental bamboo shoot flies. European Journal of Taxonomy 317, 1–23. Available from: https://doi.org/10.5852/ejt.2017.317
    Gow, P.L., 1954. Proteinaceous bait for the oriental fruit fly. Journal of Economic Entomology 47(1), 153–160.
    Gujjar, N.R., Govindan, S., Verghese, A., Subramaniam, S., More, R., 2017. Diversity of the cultivable gut bacterial communities associated with the fruit flies Bactrocera dorsalis and Bactrocera cucurbitae (Diptera: Tephritidae). Phytoparasitica 45, 453–460. Available from: https://doi.org/10.1007/s12600-017-0604-z
    Han, H.Y., Ro, K.E., 2009. Molecular phylogeny of the family Tephritidae (Insecta: Diptera): new insight from combined analysis of the mitochondrial 12S, 16S, and COII genes. Molecules and Cells 27(1), 55-66. Available from: https://doi.org/10.1007/s10059-009-0005-3
    Hancock, D.L., Drew, R.A.I., 2006. A revised classification of subgenera and species groups in Dacus Fabricius (Diptera, Tephritidae). Instrumenta Biodiversitatis VII, 167–205.
    Hancock, D.L., Drew, R.A.I., 2015. A review of the Indo-Australian subgenus Parazeugodacus Shiraki of Bactrocera Macquart (Diptera: Tephritidae: Dacinae). Australian Entomologist 42(2), 91–104.
    Hancock, D.L., Drew, R.A.I., 2017. A review of the Indo-Australian subgenera Heminotodacus Drew, Paradacus Perkins and Perkinsidacus subgen. n. (Diptera: Tephritidae: Dacinae). Australian Entomologist 44(3), 137–146.
    Hancock, D.L., Drew, R.A.I., 2018. A review of the subgenera Asiadacus Perkins, Diplodacus May, Hemigymnodacus Hardy, Niuginidacus Drew, Papuodacus Drew and Sinodacus Zia of Bactrocera Macquart (Diptera: Tephritidae: Dacinae). Australian Entomologist 45(2), 181–208.
    Hardy, D.E., 1955. A reclassification of the Dacini (Tephritidae-Diptera). Annals of the Entomological Society of America 48, 425–437.
    Howlett, F.M., 1912. The effect of oil of citronella on two species of Dacus. Transactions of the Entomological Society of London 60, 412–418.
    Howlett, F.M., 1915. Chemical reactions of fruit flies. Bulletin of Entomological Research 6, 297–305.
    Kovac, D., Dohm, P., Freidberg, A., Norrbom, A.L., 2006. Catalog and revised classification of the Gastrozonini (Diptera: Tephritidae: Dacinae). In: Freidberg, A. (Ed.), Biotaxonomy of Tephritoidea. Israel Journal of Entomology 35-36, 163–196.
    Kovac, D., Freidberg, A., Steck, G.J., 2013. Biology and description of the third instar larvae and puparium of Ichneumonopsis burmensis Hardy (Diptera: Tephritidae: Dacinae: Gastrozonini), A bamboo-breeding fruit fly from the Oriental region. Raffles Bulletin of Zoology 61(1), 117–132
    Krischik, V.A., Jones, C.G., 1991. Prologue – Microorganisms: the unseen mediators. In: Barbosa, P., Krischik, V.A., Jones, C.G. (Eds.), Microbial Mediation of Plant–Herbivore Interactions. John Wiley & Sons, NY, 1–6.
    Krosch, M.N., Schutze, M.K., Armstrong, K.F., Graham, G.C., Yeates, D.K., Clarke, A.C., 2012. A molecular phylogeny for the tribe Dacini (Diptera: Tephritidae): systematic and biogeographical implications. Molecular Phylogenetics and Evolution 64, 513-523. Available from: https://doi.org/10.1016/j.ympev.2012.05.006
    Leblanc, L., San Jose, M., Barr, N., Rubinoff, D., 2015. A phylogenetic assessment of the polyphyletic nature and intraspecific color polymorphism in the Bactrocera dorsalis complex (Diptera, Tephritidae). ZooKeys 540, 339-367. Available from: https://doi.org/10.3897/zookeys.540.9786
    Light, D.M., Jang, E.B., 1996. Plant volatiles evoke and modulate tephritid behavior. In: McPheron, B.A., Steck, G.J. (Eds.), Fruit Fly Pests: A World Assessment of Their Biology and Management. Delray Beach, FL: St. Lucie Press, 123–133.
    Metcalf, R.L., Metcalf, E.R., 1992. Plant Kairomones in Insect Ecology and Control. Chapman and Hall, London, 184.
    Munro, H.K., 1947. African Trypetidae (Diptera): A review of the transition genera between Tephritinae and Trypetinae with a preliminary study of the male terminalia. Memoir of the Entomological Society of Southern Africa 1, 284.
    Munro, H.K., 1984. A taxonomic treatise on the Dacidae (Tephritoidea, Diptera) of Africa. Entomology Memoir Department of Agriculture Republic South Africa No. 61, 313.
    Nakahara, S.M.M., Muraji, M., 2008. Phylogenetic analyses of Bactrocera fruit flies (Diptera: Tephritidae) based on nucleotide sequences of the mitochondrial COI and COII genes.      Research Bulletin of the Plant Protection Service Japan 44, 1–12.
    Nishida, R., Iwahashi, I., Tan, K.H., 1993. Accumulation of Dendrobium (Orchidaceae) flower fragrance in the rectal glands by males of the melon fly, Dacus cucurbitae (Tephritidae). Journal of Chemical Ecology 19, 713-722.
    Norrbom, A.L., Carroll, L.E., Freidberg, A., 1999. Status of Knowledge. In: Thompson, F.C. (Ed.), Fruit Fly Expert Identification System and Systematic Information Database. Myia 9, 9–48. [1998].
    Petri, L., 1909. Ricerche sopra i batteri intestinali della mosca olearia. Memorie della Regia Stazione di Patologia Vegetale di Roma, Rome, Italy, 130.
    Prokopy, R.J., 1977. Stimuli influencing trophic relations in Tephritidae. In: Labeyrie, V. (Ed.), Comporement des Insects et Millieu Trophique. Coll. Int. CNRS, 265, 305–336.
    Qin, Y., Paini, D.R., Wang, C., Fang, Y., Li, Z., 2015. Global establishment risk of economically important fruit fly species (Tephritidae). PLoS ONE 10(1), e0116424. Available from: doi:10.1371/journal.pone.0116424
    Raghu, S., Clarke, A.R., Bradley, J., 2002. Microbial mediation of fruit fly–host plant interactions: is the host plant the ‘‘centre of activity’’? OIKOS 97, 319–328.
    Rice, M.J., 1989. The sensory physiology of pest fruit flies: conspectus and prospectus. In: Robinson, A.S., Hooper, G. (Eds.), Fruit Flies: Their Biology, Natural Enemies and Control. World Crop Pests Ser. Vol. 3A, Elsevier, Amsterdam, 249–272.
    Royer, J.E., Mille, C., Cazeres, S., Brinon, J., Mayer, D.G., 2019. Isoeugenol, a more attractive male lure for the cue-lure-responsive pest fruit fly Bactrocera curvipennis (Diptera: Tephritidae: Dacinae), and new records of species responding to zingerone in New Caledonia. Journal of Economic Entomology XX(XX), 1–6.
    San Jose, M., Doorenweerd, C., Leblanc, L., Barr, N., Geib, S.M., Rubinoff, D., 2018. Incongruence between molecules and morphology: A seven-gene phylogeny of Dacini fruit flies paves the way for reclassification (Diptera: Tephritidae). Molecular Phylogenetics and Evolution 121: 139-149. Available from: https://doi.org/10.1016/j.ympev.2017.12.001
    Schutze, M.K., Virgilio, M., Norrbom, A., Clarke, A.R., 2017. Tephritid integrative taxonomy: Where we are now, with a focus on the resolution of three tropical fruit fly species complexes. Annual Review of Entomology 62, 147-164. Available from: https://doi.org/10.1146/annurev-ento-031616-035518
    Shiraki, T., 1933. A systematic study of Trypetidae in the Japanese Empire. Memoir of the Faculty of Science and Agriculture Taihoku Imperial University 8, 509, pls. 14.
    Sivinski, J., Calkins, C., 1986. Pheromones and parapheromones in the control of tephritids. Florida Entomologist 69, 157–168.
    Smith P.T., Kambhampati, S., Armstrong, K.A., 2003. Phylogenetic relationships among Bactrocera species (Diptera: Tephritidae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 26, 8-17. Available from: https://doi.org/10.1016/S1055-7903(02)00293-2
    Smith, P.T., McPheron, B.A., Kambhampati, S., 2002. Phylogenetic analysis of mitochondrial DNA supports the monophyly of Dacini fruit flies (Diptera: Tephritidae). Annals of the Entomological Society of America 95(6), 658-664. Available from: https://doi.org/10.1603/0013-8746(2002)095[0658:PAOMDS]2.0.CO;2
    Steiner, L.F., 1952. Methyl eugenol as an attractant for the oriental fruit fly. Journal of Economic Entomology 45, 241–248.
    Stringer, L.D., Soopaya, R., Butler, R.C., Vargas, R.I., Souder, S.K., Jessup, A.J. Woods. B., Cook, P.J., Suckling, D.M., 2019. Effect of lure combination on fruit fly surveillance sensitivity. Scientific Reports 9, 2653. Available from:  https://doi.org/10.1038/s41598-018-37487-6
    Tan, K.H., Nishida, R., 1996. Sex pheromone and mating competition after methyl eugenol consumption in Bactrocera dorsalis complex. In: McPheron, B.A., Steck, G.J. (Eds.), Fruit fly Pests – A World Assessment of their Biology and Management. St. Lucie Press, Delray Beach, 147–153.
    Tan, K.H., Nishida, R., 2012. Methyl eugenol – its occurrence, distribution, and role in nature, especially in relation to insect behaviour and pollination. Journal of Insect Science 20(56), 1–74. Available from: http://www.insectscience.org/12.56/i1536-2442-12-56.pdf
    Tan, K.H., Tokushima, I., Ono, H., Nishida, R., 2011. Comparison of phenylpropanoid volatiles in male rectal pheromone gland after methyl eugenol consumption, and molecular phylogenetic relationship of four global pest fruit fly species – Bactrocera invadens, B. dorsalis, B. correcta and B. zonata. Chemoecology 21, 25–33.
    Vargas, R.I., Leblanc, L., Piñero, J.C., Hoffman, K.M., 2014. Male annihilation, past, present, and future. In: Shelly, T., Epsky, N., Jang, E.B., Reyes-Flores, J., Vargas, R. (Eds.), Trapping and the Detection, Control, and Regulation of Tephritid Fruit Flies-Lures, Area-Wide Programs, and Trade Implications. Springer Dordrecht Heidelberg, NY, 493–511.
    Vargas, R.I., Miyashita, D., Nishida, T., 1984. Life-history and demographic parameters of three laboratory reared tephritids (Diptera: Tephritidae). Annals of the Entomological Society of America 77, 651–656.
    Vargas, R.I., Leblanc, L., Piñero, J.C., Leblanc, L., 2015. An overview of pest species of Bactrocera fruit flies (Diptera: Tephritidae) and the integration of biopesticides with other biological approaches for their management with a focus on the Pacific region. Insects 6, 297–318. Available from: https://doi.org/10.3390/insects6020297.
    Vayssières, J.F., Sinzogan, A., Korie, S., Ouagoussounon I., Thomas-Odjo, A., 2009.  Effectiveness of spinosad bait sprays (GF-120) in controlling mango-infesting fruit flies (Diptera: Tephritidae) in Benin. Journal of Economic Entomology 102, 515–521.
    Verghese, A., Madhura, H.S., Kamala Jayanthi, P.D., Stonehouse, J.M., 2004. Fruit flies of economic significance in India, with special reference to Bactrocera dorsalis (Hendel). In: Barnes, B.N. (Ed.), Proceedings 6th International Fruit Fly Symposium, 6-10 May 2002, Stellenbosch, South Africa, ARC Irene:  Isteg Scientific Publ., S. Africa, 317–324. Available from: https://nucleus.iaea.org/sites/naipc/twd/Documents/6thISFFEI_Proceedings/VERGHESE.pdf
    Virgilio, M., Jordaens, K., Verwimp, C., White, I.M., De Meyer, M., 2015. Higher phylogeny of frugivorous flies (Diptera, Tephritidae, Dacini): Localised partition conflicts and a novel generic classification. Molecular Phylogenetics and Evolution 85, 171–179. Available from: https://www.sciencedirect.com/science/article/pii/S1055790315000111?via%3Dihub
    White, I.M., 1999. Morphological Features of the Tribe Dacini (Dacinae): Their Significance to Behavior and Classification. In: Aluja, M., Norrbom, A.L. (Eds.), Fruit flies (Tephritidae): Phylogeny and Evolution of Behavior. CRC Press, Boca Raton, 505–534.
    White, I.M., 2006. Taxonomy of the Dacina (Diptera: Tephritidae) of Africa and the Middle East. African Entomology Memoir, 2, 156.
    White, I.M., Elson-Harris, M.M., 1992. Fruit Flies of Economic Significance: Their Identification and Bionomics, CAB. Int. Publ., Wallingford, 512.
    Yong, H.S., Song, S.L., Lim, P.E., Chan, K.G., Chow, W.L., Eamsobhana, P., 2015. Complete mitochondrial genome of Bactrocera arecae (Insecta: Tephritidae) by next-generation sequencing and molecular phylogeny of Dacini tribe. Scientific Reports 5, 15155. Available from: https://doi.org/10.1038/srep15155
    Zwolfer, H., 1983. Life system and strategies of resource exploitation in tephritids. In: Cavalloro, R. (Ed.), Fruit Flies of Economic Importance. CEC/IOBC Symp. Athens, 1982. Balkema, Rotterdam, 16–30.


Cite

1.
Vasudha A, Ahmad MA, Agarwal ML. An Overview of Indian Dacine Fruit Flies (Diptera: Tephritidae: Dacinae: Dacini) IJBSM [Internet]. 23Oct.2019[cited 8Feb.2022];10(1):491-506. Available from: http://www.pphouse.org/ijbsm-article-details.php?article=1297

People also read

Full Research

Regeneration of Commercial Tree Species and Long Term Compositional and Structural Changes in a Logged and Silviculturally Treated Brazilian Rainforest, 1955-1993

T. N. S. Karfakis

Silviculture, commercial tree, Brazil, rainforest, biodiversity value, tree regeneration

Published Online : 07 Jun 2014

Review Article

An Overview of Indian Dacine Fruit Flies (Diptera: Tephritidae: Dacinae: Dacini)

A. Vasudha, Md. Abbas Ahmad and M. L. Agarwal

Dacini, phylogeny, distribution, host associations, male lures, bacterial associations

Published Online : 23 Oct 2019

Editorial

Editorial

Published Online : 07 Jun 2010