Dacus ciliatus (lesser pumpkin fly)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Dacus ciliatus Loew
Preferred Common Name
- lesser pumpkin fly
Other Scientific Names
- Dacus appoxanthus var. decolor Bezzi
- Dacus brevistylus Bezzi
- Dacus cocciniae Premlata & Singh
- Dacus insistens Curran
- Dacus sigmoides Coquillett
- Didacus ciliatus (Loew)
- Leptoxyda ciliata (Loew)
- Tridacus mallyi Munro [nomen nudum]
International Common Names
- English: cucurbit fly; Ethiopian fruit fly; lesser melon fly
- DACUCI (Dacus ciliatus)
Summary of InvasivenessTop of page
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Diptera
- Family: Tephritidae
- Genus: Dacus
- Species: Dacus ciliatus
Notes on Taxonomy and NomenclatureTop of page
DescriptionTop of page
The genus Bactrocera belongs to the family Tephritidae, which is part of the superfamily Tephritoidea. In common with most species of Tephritoidea, this species has patterned wings, and the female has a long telescopic and pointed ovipositor; these features are hardly known outside the Tephritoidea. The family Tephritidae may also be separated from all other Diptera by the shape of the subcostal vein, which bends abruptly through a right-angle and fades to a fold before reaching the wing edge, combined with the presence of setulae along the dorsal side of vein R1. At the wing base, species of Bactrocera and Dacus have a very deep cell bm and a very long pointed extension of cell bcu (= cup). The genus Dacus, is separated from Bactrocera, by the terga (dorsal sclerites of the abdomen) being fused into a single sclerotized plate.
D. ciliatus is a member of subgenus Didacus, which is separated from other African subgenera as follows: anterior supra-alar setae absent; male with a pecten (comb of setae along each postero-lateral edge of abdominal tergum 3). It can be separated from other pest species within the subgenus by its lack of yellow vittae (stripes) on the scutum, and by the yellow spot in each haltere base being small and separated from the scutellum by at least its own diameter. Non-reared specimens from Africa should be identified using the keys provided by Munro (1984), but it should be noted that those keys are very difficult to use. Non-reared specimens from Asia should be identified using Drew et al. (1998).
White and Elson-Harris (1994) gave some details, but they were insufficient to permit separation from other pest species of subgenus Didacus.
DistributionTop of page
D. ciliatus is a major pest of cucurbits throughout Africa, except in the Maghreb, being distributed from Egypt to South Africa (Hancock, 1989). The fly is also a pest in the Indian Ocean (Reunion and Mauritius) and Oriental Asia (Pakistan and India). It has also been reported in the Middle East (Iran, Saudi-Arabia, Yemen, amongst others). D. ciliatus has spread to Israel (Norrbom et al., 1999) and to Oman (Azam et al., 2004). It is found in a small area of the Arava desert and is still a quarantine insect in Israel (Maklakov et al., 2001). As the common name, Ethiopian fruit fly, suggests, D. ciliatus is native to East Africa (Vayssières et al., 2008).
Due to confusion with other species, some country records have been based on misidentifications. Records of D. ciliatus from Cape Verde have been found to be based on a misidentification of Dacus frontalis (White and Elson-Harris, 1994). Similarly, Sri Lankan specimens examined by Drew et al. (1998) were Dacus keiseri. A record for Botswana, cited in the 1995 edition of the Distribution Maps of Plant Pests (UK International Institute of Entomology, 1995), was based on an NHM specimen that has not been traced, and the record for Madagascar is likely to be based on a misidentification. The following records, also cited in this map, are not confirmed by voucher specimens available to either NHM staff or to Munro (1984) and are listed as requiring further confirmation: Bangladesh; Myanmar; Benin; Chad; and Sierra Leone. Refer to CABI/EPPO (2002) for further information.
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 12 May 2022
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Cabo Verde||Absent, Invalid presence record(s)|
|Chad||Absent, Unconfirmed presence record(s)|
|Congo, Democratic Republic of the||Present|
|Ethiopia||Present||Originally from East Africa|
|Madagascar||Absent, Unconfirmed presence record(s)|
|Réunion||Present||Recorded in Reunion after Mauritius|
|South Africa||Present, Widespread|
|Myanmar||Absent, Unconfirmed presence record(s)|
|Thailand||Absent, Unconfirmed presence record(s)|
|Turkey||Present, Localized||Diyarbakir, Mardin, Siirt and Sirnak.|
|United Arab Emirates||Present|
|New Zealand||Absent, Confirmed absent by survey|
History of Introduction and SpreadTop of page
Adult flight and the transport of infested fruit are the major means of movement and dispersal to previously uninfected areas.
Risk of IntroductionTop of page
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Principal habitat||Productive/non-natural|
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat||Productive/non-natural|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Secondary/tolerated habitat||Natural|
Hosts/Species AffectedTop of page
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
List of Symptoms/SignsTop of page
|Fruit / discoloration|
|Fruit / extensive mould|
|Fruit / gummosis|
|Fruit / internal feeding|
|Fruit / lesions: black or brown|
|Fruit / lesions: scab or pitting|
|Fruit / obvious exit hole|
|Fruit / odour|
|Fruit / ooze|
Biology and EcologyTop of page
D. ciliatus adults reach their sexual maturity at 14-15 days although there are data from Egypt stating that the adults reach their reproductive maturity after 5-6 days during the summer and after 20-30 days during the winter (El Nahal et al., 1971). As in most of the fruit flies belonging to the Dacinae, mating in D. ciliatus occurs at dusk under low light intensities (Fletcher, 1987; Smith, 1989). At light intensities lower than 50 lux, mating pairs start to copulate (Dehecq, 1995). They usually stay coupled during the entire night and are separated by the morning light (Dehecq, 1995). Although not yet confirmed, it seems that females mate only once in their life span (El Nahal et al., 1971).
Generally, mature females of the genus Bactrocera oviposit into fruit. Although in most species of Bactrocera this is usually at the start of ripening, some cucurbit-associated species may attack at fruit set or even attack the ovaries. Vayssières and Dal (2002) observed that orange spheres mostly attract sexually mature females for oviposition whereas immature females prefer yellow colours.
Females oviposit an average of 210 eggs (El Nahal et al., 1971). Fetoh (2006) recorded a mean egg production of 322.6. The eggs are laid in groups of 5-15 (El Nahal et al., 1971). After the eggs hatch, the young larvae start to feed in the host, causing damage to the fruit. The final instar larvae of Bactrocera drop to the ground, find a crack to drop into, and then form a puparium within which pupation takes place.
Patel and Patel (1998) reared D. ciliatus in the laboratory (in Coccinia grandis) and noted that larval development took 4-7 days, pupariation 7-14 days, and adult longevity was 13-27 days, depending on the season. El Nahal et al. (1971) reported similar results from Cucurbita pepo, except that they reported a pupation period of up to 40 days. Fetoh (2006) reared D. ciliatus under laboratory conditions on marrow fruits and recorded the following mean life spans: egg, 3.0 ± 0.8 days; larva, 7.3 ± 2.7 days; pupa, 9.3 ± 1.9 days; adult, 14-45 days.
In laboratory conditions, the adults may live for up to 4 months with constant sources of protein and sucrose (El Nahal et al., 1971; Yarom et al., 1997). On the other hand, if deprived from food, the adults die in about 2-3 days (El Nahal et al., 1971). Fetoh (2006) recorded a sex ratio of 1:1.
In comparison to the related species B. cucurbitae,D. ciliatus is characterised by early reproduction, lower oviposition time, shorter life span, and lower fecundity (Vayssières et al., 2008). The adults are classified as K-strategists (Vayssières et al., 2002).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Fopius arisanus||Parasite||Eggs; Arthropods|Larvae; Arthropods|Pupae|
Notes on Natural EnemiesTop of page
Silvestri (1913) noted that in watermelon [Citrullus lanatus], melon [Cucumis melo] and cucumber [Cucumis sativus], the larvae were not parasitized because the larvae are too deep within such large fruits. However, he reported 60% parasitism in the fruits of Momordica.
Means of Movement and DispersalTop of page
Adult flight and the transport of infested fruits are the major means of movement and dispersal to previously uninfested areas. The flight capability of D. ciliatus has not been measured. Transport of plants of host species that are transported with roots from countries where these pests occur can also be a cause of dispersal of puparia.
[Erratum: In previous versions of this datasheet, it was stated that “many Bactrocera spp. can fly 50-100 km (Fletcher, 1989)” but a review of Fletcher (1989a) and Fletcher (1989b) by Hicks et al. (2019) found no evidence to support this statement and it has been removed. Fletcher (1989b) provides dispersal data for only 11 of 651 species of Bactrocera, many of the case studies lack the necessary numerical data, and the study did not discern between active flight and passive wind-assisted dispersal. There are differences among fruit fly species and further studies are required to determine dispersal distances for individual species. For further information on trapping Bactrocera species to monitor movement, see Weldon et al. (2014).]
Pathway CausesTop of page
Pathway VectorsTop of page
|Bulk freight or cargo||infested fruit||Yes|
|Clothing, footwear and possessions||fruit in cases or bags||Yes|
|Containers and packaging - wood||Of fruit cargo||Yes|
|Land vehicles||infested fruit||Yes|
|fruit in post||Yes|
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Fruits (inc. pods)||arthropods/eggs; arthropods/larvae||Yes||Pest or symptoms usually visible to the naked eye|
|Growing medium accompanying plants||arthropods/pupae||Yes||Pest or symptoms usually visible to the naked eye|
Impact SummaryTop of page
ImpactTop of page
Economic ImpactTop of page
D. ciliatus is a serious pest of cucurbit crops (Hancock, 1989). It is reported to cause serious economic damage in Egypt (El Nahal et al., 1971) and South Africa (Hancock, 1989). In the Reunion Island, D. ciliatus, together with Dacus cucurbitae (and out of nine species of Tephritidae), represents the primary pests of Cucurbitaceae, having been reported in nine genera of this plant family (Dehecq, 1995; Vayssières et al., 2008).
It is an EPPO A1 quarantine pest within the category 'non-European Trypetidae' and is also of quarantine significance to CPPC (Caribbean Plant Protection Commission). However, the risk of establishment in most of the EPPO area is minimal, although in some areas, populations might enter and multiply during the summer months and, in southern areas may survive several winters (EPPO, 2009). The direct losses from these introductions are not believed to be high. However, its presence may cause indirect economic impact by restricting exports.
In southern areas, some populations might survive one or several winters, although the direct losses from such introductions would probably not be high. The major risk for EPPO countries arises from the probable imposition of much stricter phytosanitary restrictions on exported fruits (particularly to America) if Bactrocera spp. or tropical Dacus spp., such as D. ciliatus, enter and multiply, even temporarily.
Risk and Impact FactorsTop of page
- Has a broad native range
- Abundant in its native range
- Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
- Highly mobile locally
- Benefits from human association (i.e. it is a human commensal)
- Fast growing
- Has high reproductive potential
- Has propagules that can remain viable for more than one year
- Host damage
- Monoculture formation
- Negatively impacts agriculture
- Negatively impacts cultural/traditional practices
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
Uses ListTop of page
- Laboratory use
- Research model
Detection and InspectionTop of page
However, both sexes may be monitored using protein bait traps (either protein hydrolysate or protein autolysate); see Drew (1982) for further details. More recent experimental essays led to the conclusion that the fly seems to be attracted to low concentrations of some “food lures” and to volatiles emanating from mature melons (D Nestel, Agriculture Research Center, Israel, personal communication, 2008). Sookar et al. (2001), during a field cage trial in order to determine the preference of fruit flies to two types of bait (commercial and local), found no significant differences among the trap catches for D. ciliatus.
Similarities to Other Species/ConditionsTop of page
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
When detected, it is important to gather all infested host fruits and destroy them. Protein baited traps should be used to monitor population size and spread continuously.
Insecticide protection is possible by using a cover spray (the bait sprays used against fruit flies in tree fruits are not generally used for cucurbit crops). Shaheen et al. (1973) found that sprays of pirimiphos-methyl or trichlorphon were effective against D. ciliatus on autumn plantings of Cucumis melo (snake cucumber and sweetmelon). Saddik and Rizkallah (1970) found that fenthion and trichlorphon all increased yield. Kazi (1976) found that adult D. ciliatus moved to non-host 'rest plants' between 1700h and 0800h, and suggested that the flies might be better controlled there, rather than on the larval host.
D.ciliatus adults are highly sensitive to low concentrations of spinosad baited with 1% yeast hydrolysate and 10% sucrose as phagostimulant; specifically 8.5 ppm had a 90% efficacy on females (Nestel et al., 2004). Chakravarthy et al. (2007) found that a garlic barrier (GB Ag; garlic juice 99.98% pure) lowered the infestation of D. ciliatus when used on gherkin cultivation. Moreover, Hussein et al. (2006) showed that the entomopathogenic nematodes, Steinernema feltiae (NC) and Heterorhabditis bacteriophora (BA1) efficiently controlled D. ciliatus larvae and pupae, whereas treatment with profenofos and pirimiphos-methyl caused even higher mortality. On the other hand, organophosphates were proved to be ineffective for the control of D. ciliatus, but pyrethroids have high potential for controlling it, showing satisfactory killing ability, massive knockdown effect, and prevention of oviposition (Maklakov et al., 2001).
Nestel found that pesticides such as Telstar (pyrethroid), Mospilan (neonicotinoid) and Avisect were effective against D. ciliatus. Organic insecticides such as Karlic@ and Hot-Pepper@ did not show any effectiveness in killing or deterring the fly (D Nestel, Agriculture Research Center, Israel, personal communication, 2008).
Physical protection provides a more environmentally acceptable approach. In Senegal, where D. ciliatus and Dacus vertebrates cause heavy losses to cantaloupe melon [Cucumis melo] production, it was found that growing under the protection of tunnels increased yield (Froissart et al., 1995; Vaissière and Froissart, 1996).
Consignments of fruits from countries where these pests occur should be inspected for symptoms of infestation and suspect fruit should be cut open in order to look for larvae. EPPO recommends that such fruits should come from an area where D. ciliatus does not occur and where routine intensive control measures are applied (OEPP/EPPO, 1990). The fruits may also be treated in transit by cold treatment (13 or 14 days at 0.0 or 0.6°C, respectively) or, for certain types of fruits, by vapour heat (keeping at 43-44°C for 6-9 h, according to commodity) (FAO, 1983), or hot water treatment. Treatment methods against fruit flies are currently under review within EPPO and as part of a common programme of the regional plant protection organizations.
Plants of host species transported with roots from countries where these pests occur should be free from soil, or the soil should be treated against puparia. The plants should not carry fruits. Importation of such plants may indeed be prohibited.
Gaps in Knowledge/Research NeedsTop of page
There are many gaps in knowledge concerning this species, including:
- Few studies have been carried out on the biology of this species
- There is only one study on the demographic parameters of this species
- There is little information about trapping methods or attractive substances - there is only one study about the attraction of D. ciliatus to several known lures for fruit flies
- There is little information about pesticide sensitivity.
ReferencesTop of page
Aldawood, A. S., 2013. Comparative study of cucurbit fly: Dacus ciliatus Loew (Diptera: Tephritidae) infestation on zucchini squash (Cucurbita pepo L.) at Huraimila and Diraab, Riyadh Region, Saudi Arabia. Egyptian Academic Journal of Biological Sciences: Entomology, 6(2), 91-96. http://entomology.eajbs.eg.net/pdf/vol6-num2/9.pdf
Armstrong JW, Couey HM, 1989. Control; fruit disinfestation; fumigation, heat and cold. In: Robinson AS, Hooper G, eds. Fruit Flies; their Biology, Natural Enemies and Control. World Crop Pests. Amsterdam, Netherlands: Elsevier, 3(B):411-424
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Badr-Elsabah AF, Afia YI, 2004. Preliminary biological aspects of the new recorded parasitoid, Dirhinus luzonesis, (Hymenoptera: Chalcididae) of cucurbit fruit fly, Dacus ciliatus (Loew.) and peach fruit fly, Bactorcera (=Dacus) zonata (Saunders) (Diptera: Tephritidae) in Egypt. Egyptian Journal of Biological Pest Control, 14(1):73-76
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Vayssières JF, Carel Y, Coubes M, 2002. Demographic parameters and biotic factors of two Dacini species, Bactrocera cucurbitae and Dacus ciliatus, on Réunion Island. In: Proceedings of the 6th International Symposium on fruit flies of economic importance, Stellenbosch, South Africa, 6-10 May 2002 [ed. by Barnes BN], 91-95
Vayssières JF, Carel Y, Coubes M, Duyck PF, 2008. Development of Immature Stages and Comparative Demography of Two Cucurbit-Attacking Fruit Flies in Reunion Island: Bactrocera cucurbitae and Dacus ciliatus (Diptera Tephritidae). Environmental Entomology, 37((2)):307-314
Vayssières JF, Dal F, 2002. Responses of the Ethiopian fruit fly, Dacus ciliatus Loew (Diptera: Tephritidae), to coloured rectangles, spheres and ovoids. In: Proceedings of the 6th International Symposium on fruit flies of economic importance, Stellenbosch, South Africa, 6-10 May 2002 [ed. by Barnes BN], 111-116
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Kambura C, Tanga C M, Kilalo D, Muthomi J, Salifu D, Rwomushana I, Mohamed S A, Ekesi S, 2018. Composition, host range and host suitability of vegetable-infesting tephritids on cucurbits cultivated in Kenya. African Entomology. 26 (2), 379-397. DOI:10.4001/003.026.0379
Maklakov A, Ishaaya I, Freidberg A, Yawetz A, Horowitz A R, Yarom I, 2001. Toxicological studies of organophosphate and pyrethroid insecticides for controlling the fruit fly Dacus ciliatus (Diptera: Tephritidae). Journal of Economic Entomology. 94 (5), 1059-1066. DOI:10.1603/0022-0493-94.5.1059
Okolle J N, Ntonifor N N, 2005. Field ovipositional behavior and laboratory studies on development of Dacus punctatifrons (Diptera: Tephritidae) on tomato. Insect Science. 12 (5), 393-398. DOI:10.1111/j.1005-295X.2005.00049.x
Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435
Umeh V C, Garcia L E, Meyer M de, 2008. Fruit flies of sweet oranges in Nigeria: species diversity, relative abundance and spread in major producing areas. Fruits (Paris). 63 (3), 145-153. http://www.fruits-journal.org/ DOI:10.1051/fruits:2008004
Vayssières J F, Carel Y, Coubes M, Duyck P F, 2008. Development of Immature Stages and Comparative Demography of Two Cucurbit-Attacking Fruit Flies in Reunion Island: Bactrocera cucurbitae and Dacus ciliatus (Diptera Tephritidae). Environmental Entomology. 37 (2), 307-314.
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11/08/2008 Updated by:
Ana Larcher Carvalho, Biologika consulting, Ag & Rural Development Tv do Terreirinho, 11, 2-D, 1100-600 Lisboa, Portugal
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