Invasive Species Compendium

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Datasheet

Bactrocera oleae
(olive fruit fly)

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Datasheet

Bactrocera oleae (olive fruit fly)

Pictures

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PictureTitleCaptionCopyright
Bactrocera oleae (olive fruit fly); adult. Museum set specimen.
TitleAdult
CaptionBactrocera oleae (olive fruit fly); adult. Museum set specimen.
Copyright©Ian M. White
Bactrocera oleae (olive fruit fly); adult. Museum set specimen.
AdultBactrocera oleae (olive fruit fly); adult. Museum set specimen.©Ian M. White
Bactrocera oleae (olive fruit fly); infested olive fruits on tree. Chania, Kolymvari, Crete, Greece. October, 2014.
TitleInfested olive fruits on tree
CaptionBactrocera oleae (olive fruit fly); infested olive fruits on tree. Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); infested olive fruits on tree. Chania, Kolymvari, Crete, Greece. October, 2014.
Infested olive fruits on treeBactrocera oleae (olive fruit fly); infested olive fruits on tree. Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); healthy olive fruit (a) and, three infested fruits (b-d). Chania, Kolymvari, Crete, Greece. October, 2014.
TitleHealthy and infested olive fruits
CaptionBactrocera oleae (olive fruit fly); healthy olive fruit (a) and, three infested fruits (b-d). Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); healthy olive fruit (a) and, three infested fruits (b-d). Chania, Kolymvari, Crete, Greece. October, 2014.
Healthy and infested olive fruitsBactrocera oleae (olive fruit fly); healthy olive fruit (a) and, three infested fruits (b-d). Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); larval feeding tunnel (arrowed) in olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
TitleLarval feeding tunnel
CaptionBactrocera oleae (olive fruit fly); larval feeding tunnel (arrowed) in olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); larval feeding tunnel (arrowed) in olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
Larval feeding tunnelBactrocera oleae (olive fruit fly); larval feeding tunnel (arrowed) in olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); third stage larva (arrowed) consumining olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
TitleThird stage larva
CaptionBactrocera oleae (olive fruit fly); third stage larva (arrowed) consumining olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); third stage larva (arrowed) consumining olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.
Third stage larvaBactrocera oleae (olive fruit fly); third stage larva (arrowed) consumining olive mesocarp. Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); exit hole (arrowed) of Bactrocera oleae in an olive fruit. Chania, Kolymvari, Crete, Greece. October, 2014.
TitleLarval exit hole
CaptionBactrocera oleae (olive fruit fly); exit hole (arrowed) of Bactrocera oleae in an olive fruit. Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); exit hole (arrowed) of Bactrocera oleae in an olive fruit. Chania, Kolymvari, Crete, Greece. October, 2014.
Larval exit holeBactrocera oleae (olive fruit fly); exit hole (arrowed) of Bactrocera oleae in an olive fruit. Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); McPhail trap loaded with protein attractant for monitoring olive fruit fly populations. Chania, Kolymvari, Crete, Greece. October, 2014.
TitleMcPhail trap for monitoring fly populations
CaptionBactrocera oleae (olive fruit fly); McPhail trap loaded with protein attractant for monitoring olive fruit fly populations. Chania, Kolymvari, Crete, Greece. October, 2014.
Copyright©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); McPhail trap loaded with protein attractant for monitoring olive fruit fly populations. Chania, Kolymvari, Crete, Greece. October, 2014.
McPhail trap for monitoring fly populationsBactrocera oleae (olive fruit fly); McPhail trap loaded with protein attractant for monitoring olive fruit fly populations. Chania, Kolymvari, Crete, Greece. October, 2014.©Kiki Varikou-2014
Bactrocera oleae (olive fruit fly); aculeus, dorsal view (optical section) of apex.
TitleAculeus
CaptionBactrocera oleae (olive fruit fly); aculeus, dorsal view (optical section) of apex.
Copyright©CAB International
Bactrocera oleae (olive fruit fly); aculeus, dorsal view (optical section) of apex.
AculeusBactrocera oleae (olive fruit fly); aculeus, dorsal view (optical section) of apex.©CAB International

Identity

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Preferred Scientific Name

  • Bactrocera oleae Gmelin, 1790

Preferred Common Name

  • olive fruit fly

Other Scientific Names

  • Bactrocera (Daculus) oleae
  • Daculus oleae (Gmelin)
  • Dacus oleae (Gmelin)
  • Musca oleae (Gmelin)

International Common Names

  • English: fruit fly, olive; olive fly; olive fruit fly
  • Spanish: mosca de las aceitunas; mosca del olivo; mosca olearia
  • French: mouche de l'olive; mouche des olives; ver de l'olive

Local Common Names

  • Germany: Fliege, Oliven-; Olivenfliege
  • Israel: zvuv hazayit
  • Italy: mosca delle olive
  • Turkey: zeytin sinegi

EPPO code

  • DACUOL (Bactrocera oleae)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Diptera
  •                         Family: Tephritidae
  •                             Genus: Bactrocera
  •                                 Species: Bactrocera oleae

Notes on Taxonomy and Nomenclature

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This is the only species which definitely belongs to subgenus Bactrocera (Daculus), although Drew (1989) included it in B. (Polistomimetes) together with species which are here placed in B. (Tetradacus).

Description

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Adult description derived from computer generated description from White and Hancock (1997). Larval description based on Phillips (1946), and Kandybina (1977), as given in White and Elson-Harris (1994).

Head

Pedicel + 1st flagellomere not longer than ptilinal suture. Face with a dark spot in each antennal furrow. Facial spot, round/elongate, small.

Thorax

Predominant colour of scutum, orange-brown to black. Postpronotal (=humeral) lobe entirely pale (yellow or orange). Scutum without lateral and medial postsutural vittae (yellow/orange stripes). Scutellum with a deep basal band and often deepened to form a black triangle. Anepisternal stripe not reaching anterior notopleural seta. Yellow marking on hypopleural calli restricted to lower callus (katatergite) only. Postpronotal lobe (=humerus) without a seta. Scutum without anterior supra-alar setae and without prescutellar acrostichal setae. Scutellum without basal setae.

Wing

Length, 4.3-5.2 mm. Cells bc and c without extensive covering of microtrichia. Cell br (narrowed part) without extensive covering of microtrichia. Without a complete costal band; marked at end of R4+5 only; without an anal streak. Cells bc and c not coloured.

Legs

yellow / pale.

Abdomen

Predominant colour orange-brown to black. Tergites not fused. Abdomen not wasp waisted. Pattern on abdomen distinct; tergite 3 and 4, dark laterally. No medial longitudinal stripe on T4. Ceromata round/ovoid.

Terminalia and secondary sexual characters

Male wing without a bulla. Male tergite 3 with a pecten (setal comb) on each side. Surstylus (male) without a long posterior lobe. Wing (male) with a deep indent in posterior margin. Wing (male) with microtrichia area around cell cup. Hind tibia (male) with a preapical pad. Aculeus apex pointed.

Larvae

Medium sized, length 6.5-7.0 mm; width 1.2-1.7 mm.

Head. Oral ridges in 10-12 shallow, short rows; mouthhooks heavily sclerotized, each with a short, slender, curved apical tooth.

Thoracic and abdominal segments. Anterior portion of T1-T3, A1 and A2 with 3-5 rows of small spinules encircling each segment; A3-A5 with a few spinules dorsally and a heavier concentration ventrally; A6-A8 with spinules ventrally, none dorsally and laterally; spinules in creeping welts smaller in central rows.

Anterior spiracles. 8-12 short tubules.

Posterior spiracles. Spiracular slits 3.5-4.0 times as long as broad, with a thick rima; spiracular hairs about half the length of a spiracular slit, frequently branched, dorsal and ventral bundles of 7 hairs, lateral bundles of 2-4 hairs.

Anal area. Lobes small, slightly protuberant, surrounded by several discontinuous rows of small spinules.

Distribution

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B. oleae is found throughout the olive-growing zone of the Mediterranean. It is also found (on wild olives) in parts of eastern and southern Africa. The current distribution of the pest includes South and Central Africa, Pakistan, Mediterranean Europe and the Middle East and it has been introduced recently to California, USA, and Mexico (Nardi et al., 2005). B. oleae has been trapped in areas of wild olive in Réunion (White et al., 2000). It is not known if this is a recent introduction or if the flies have been there for a long time and have been overlooked.

The distribution map includes records based on specimens of B. oleae from the collection in the Natural History Museum (London, UK): dates of collection are noted in the list of countries (NHM, various dates).

 

Distribution Table

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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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaAbsent, unreliable recordEPPO, 2014
AzerbaijanAbsent, invalid recordEPPO, 2014
Georgia (Republic of)PresentKandybina, 1977; EPPO, 2014; CABI/EPPO, 2015
IndiaRestricted distributionKapoor, 1993; EPPO, 2014; CABI/EPPO, 2015
-Jammu and KashmirPresentShant, 1999; CABI/EPPO, 2015
IranPresentNouri, 2007; CABI/EPPO, 2015
IsraelPresentFriedberg & Kugler, 1989; EPPO, 2014; CABI/EPPO, 2015
JordanPresentAl-Zaghal and Mustafa, 1987; EPPO, 2014; CABI/EPPO, 2015
LebanonPresentEPPO, 2014; CABI/EPPO, 2015
PakistanPresentKapoor, 1993; EPPO, 2014; CABI/EPPO, 2015
Saudi ArabiaPresentEPPO, 2014; CABI/EPPO, 2015
SyriaPresentEPPO, 2014; CABI/EPPO, 2015
TurkeyPresentGuusay et al., 1990; EPPO, 2014; CABI/EPPO, 2015

Africa

AlgeriaPresentGaouar and Debouzie, 1995; EPPO, 2014; CABI/EPPO, 2015
AngolaPresentCogan & Munro, 1980; EPPO, 2014; CABI/EPPO, 2015
EgyptPresentMunro, 1984; EPPO, 2014; CABI/EPPO, 2015
EritreaPresentMunro, 1984; EPPO, 2014; CABI/EPPO, 2015
EthiopiaPresentNHM, 1975; EPPO, 2014; CABI/EPPO, 2015
KenyaPresentNHM, 1991; EPPO, 2014; CABI/EPPO, 2015
LibyaPresentHammad, 1980; EPPO, 2014; CABI/EPPO, 2015
MauritiusPresentCABI/EPPO, 2015
MoroccoPresentEPPO, 2014; CABI/EPPO, 2015
RéunionPresentNHM, 1997; CABI/EPPO, 2015
SeychellesPresentCABI/EPPO, 2015
South AfricaPresentMunro, 1984; EPPO, 2014; CABI/EPPO, 2015
Spain
-Canary IslandsPresentMerz, 1992; EPPO, 2014; CABI/EPPO, 2015
SudanPresentWhite and Elson-Harris, 1994; EPPO, 2014; CABI/EPPO, 2015
TunisiaPresentEPPO, 2014; CABI/EPPO, 2015

North America

MexicoPresentIntroducedNardi et al., 2005; EPPO, 2014; CABI/EPPO, 2015
USARestricted distributionEPPO, 2014; CABI/EPPO, 2015
-CaliforniaPresentIntroducedRice et al., 2003; EPPO, 2014; CABI/EPPO, 2015

Europe

AlbaniaPresentHawkes et al., 2005; CABI/EPPO, 2015
CroatiaPresentBrnetic, 1979; EPPO, 2014; CABI/EPPO, 2015
CyprusWidespreadNHM, 1987; EPPO, 2014; CABI/EPPO, 2015
FranceRestricted distributionPanis, 1979; EPPO, 2014; CABI/EPPO, 2015
-CorsicaPresentEPPO, 2014; CABI/EPPO, 2015
GreeceWidespreadNHM, 1984; Fimiani, 1989; EPPO, 2014; CABI/EPPO, 2015
-CretePresentEPPO, 2014; CABI/EPPO, 2015
-Greece (mainland)PresentCABI/EPPO, 2015
ItalyWidespreadFimiani, 1989; EPPO, 2014; CABI/EPPO, 2015
-SardiniaPresentEPPO, 2014; CABI/EPPO, 2015
-SicilyPresentEPPO, 2014; CABI/EPPO, 2015
MaltaPresentEPPO, 2014; CABI/EPPO, 2015
MontenegroPresentPerovic et al., 2007; Perovic et al., 2007; EPPO, 2014; CABI/EPPO, 2015
PortugalWidespreadEPPO, 2014; CABI/EPPO, 2015
-AzoresPresentCABI/EPPO, 2015
-Portugal (mainland)PresentCABI/EPPO, 2015
Russian FederationAbsent, invalid recordEPPO, 2014
SerbiaPresentPerovic et al., 2007
SloveniaPresentJancar, 2003; EPPO, 2014; CABI/EPPO, 2015
SpainWidespreadNHM, 1987; EPPO, 2014; CABI/EPPO, 2015
-Balearic IslandsPresentEPPO, 2014; CABI/EPPO, 2015
-Spain (mainland)PresentCABI/EPPO, 2015
SwitzerlandPresentMerz, 1994; EPPO, 2014; CABI/EPPO, 2015

Risk of Introduction

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B. oleae is not known to attack any fruits outside of the genus Olea and as such can only represent a threat to olive production. As all olive producing countries are already heavily infested by this species it does not represent a significant quarantine threat, although there is a very remote possibility that it may be able to develop on some other related fruits (the family Oleaceae is widespread and some other genera are hosts to specialist fruit flies).

Habitat

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Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial

Hosts/Species Affected

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This species has a very narrow host range, being restricted to Olea spp. In Europe, it attacks cultivated olives but in Africa it is associated with wild olives.

In a study of host trees infested by B. oleae in California, USA, Athar (2005) noted that olives were the preferred host, but trees in the families Rosaceae, Rutaceae, Anacardiaceae, Fabaceae, Lythraceae and Malpighiaceae were also infested. The hosts were mainly fruit trees, with the exceptions of Brazilian pepper tree (Schinus terebinthifolia), carob (Ceratonia siliqua), crape myrtle (Lagerstroemia indica) and ornamental plum (Prunus domestica).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Olea europaea subsp. europaea (European olive)OleaceaeMain

Growth Stages

Top of page Fruiting stage

Symptoms

Top of page Puncture marks and exit holes may be observed. Eggs are laid singly in a small chamber below the oviposition hole.

List of Symptoms/Signs

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SignLife StagesType
Fruit / internal feeding
Fruit / lesions: black or brown
Fruit / premature drop

Biology and Ecology

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Eggs are laid below the skin of the host fruit; a female may lay more than 200 eggs but unlike most other Bactrocera spp. these are laid singly. Eggs hatch within 2-4 days and the larvae feed for another 10-14 days. Pupation is either in the soil under the host plant or, when fruits are attacked early in their development, in the fruit. Pupation takes about 10 days but may be delayed for several weeks under cool conditions. Adults occur throughout the year in Israel (Freidberg and Kugler, 1989) but only during the summer months in cooler areas so the number of generations may be from one to several, with winter passed during pupation. Adults mature after about a week, and may live 1-2 months; most data from Christenson and Foote (1960), Clausen (1978), Mazomenos (1989).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Asecodes erxias Parasite
Belonuchus rufipennis Predator Italy olives
Biosteres longicaudatus Parasite Greece olives
Bracon celer Parasite Larvae Italy olives
Carabus banonii Predator
Cirrospilus variegatus Parasite Larvae Italy olives
Coptera silvestrii Parasite
cricket paralysis virus Pathogen
Cyrtoptyx latipes Parasite
Diachasmimorpha tryoni Parasite Greece olives
Euderus cavasolae Parasite Larvae Italy olives
Eupelmus afer Parasite Italy olives
Eupelmus ater Parasite Larvae Italy olives
Eupelmus urozonus Parasite Larvae Greece; Corfu olives
Eurytoma Parasite Larvae/Pupae
Eurytoma martellii Parasite
Halticoptera daci Parasite Larvae Italy olives
Iridovirus Pathogen
Mesopolobus modestus Parasite Larvae Italy olives
Neochrysocharis formosa Parasite Larvae
Nucleopolyhedrosis virus Pathogen
Opius concolor Parasite Crete; France; Greece; Israel; Italy; Khalki; Mediterranean region; Sicily; Spain; Yugoslavia olives
Opius dacicida Parasite Larvae Italy olives
Opius lounsburyi Parasite
Opius tephritivorus Parasite Larvae Italy olives
Opius trimaculatus Parasite Italy olives
Pnigalio agraules Parasite Greece; Corfu olives
Psyttalia concolor Parasite Larvae Crete;France;Greece;Israel;Italy;Khalki;Mediterranean region;Sicily;Spain;Yugoslavia olives
Pterostichus creticus Predator
Scolopendra cretica Predator
small RNA viruses Pathogen
Tetrastichus Parasite
Triaspis daci Parasite Larvae Italy olives
Trichosteresis glabra Parasite

Notes on Natural Enemies

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Clausen (1978) reviews numerous releases of parasitoids made against B. oleae, primarily in Italy, France and Greece, following extensive searches for parasitoids by P. Silvestri in the early years of this century. Most failed to establish, but Opius concolor has been shown to achieve considerable levels of control when regularly released. Greathead (1976) also reviews the natural enemies of B. oleae. Surveys for biological control agents were made in Africa where there is a greater range of species than in Europe, Supporting the view that the fly originated in Africa. The more important natural enemies were found both in Eritrea and in South Africa, suggesting that they are widespread on B. oleae breeding in wild Olea africana, presumably its original host plant.

Ranaldi and Santoni (1987) reviewed naturally occurring parasitoids in Italy with respect to their time of greatest impact, so as to permit integration with chemical control. However, none of these achieved significant levels of control.

Other sources of parasitoid data are: Mechelany (1969), Fenili and Pegazzano (1971), Arambourg and Pralavorio (1974), Viggiani et al. (1975), Monaco (1978), Bigler and Delucchi (1981), Neuenschwander (1982), Neuenschwander et al. (1983), Bigler et al. (1986) and Mustafa et al. (1987).

A source of virus data is Manousis and Moore (1987).

Means of Movement and Dispersal

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Adult flight and the transport of infected fruit are the major means of movement and dispersal to previously uninfected areas. 

[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 (2016, unpublished data, USDA) 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 Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsFruit in case or handbag. Yes
Containers and packaging - woodOf fruit cargo. Yes
Land vehiclesAeroplanes and boats, with fruit cargo. Yes
MailFruit in post. Yes
Soil, sand and gravelRisk of puparia in soil. Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Fruits (inc. pods) eggs; larvae Yes Pest or symptoms usually visible to the naked eye
Growing medium accompanying plants pupae Yes Pest or symptoms usually visible to the naked eye
Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Leaves
Roots
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
True seeds (inc. grain)
Wood

Impact

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B. oleae is considered the most important pest of cultivated olives, Olea europaea L., in many of the areas of the Mediterranean basin, affecting the quality and quantity of both olive oil and table olives (Michelakis and Neuenschwander, 1983; Manousis and Moore, 1987; Economopoulos, 2002). Unlike the fruits attacked by most other Bactrocera spp., olives containing larvae of B. oleae are frequently included in the harvested crop and subsequent oil production.
 
B. oleae infestation of olives increases the oxidative and hydrolytic degradation of the oil (Montedoro et al. 1992; Gomez-Caravaca et al. 2008; Tamendjari et al 2009) and negatively affects the flavour, as a consequence of microorganism activity (Bendini et al. 2007). In many cases the main impact is a tainting of the olive oil produced rather than complete destruction of the crop (table olives can be destroyed). Neuenschwander and Michelakis (1978) reported that in batches of olives in which all fruits showed emergence holes, the resulting oil was up to 12 times as acidic as uninfested batches.

Diagnosis

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Simple Diagnosis

This is the only fruit fly likely to be reared from cultivated olives (a few other species of restricted range in Africa and Australia have been reared and these were listed by White and Elson-Harris, 1994). Features to check are the lack of any bright yellow/orange stripes (vittae) laterally on the scutum and the wing markings, which are reduced to an apical dark spot (plus dark cell sc).

Male Lure

This species is not attracted to either cue lure or methyl eugenol.

Diagnosis (minimum characters to differentiate from most other Bactrocera and Dacus spp., from White and Hancock, 1997)

Face with a dark spot in each antennal furrow. Scutum without a medial vitta; without lateral postsutural vittae; without anterior supra-alar setae. Scutellum without basal setae. Wing pattern reduced, costal band reduced to an apical spot. Cell br (narrowed part) without extensive covering of microtrichia. Tergite 3 with dark lateral markings. Male tergite 3 with a pecten (setal comb) on each side.

Detection and Inspection

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B. oleae does not respond to standard fruit fly male lures. Field monitoring must therefore be by sampling susceptible fruits for larvae, or by trapping of adults. Adults may be caught in protein bait traps (see Control) and they are also attracted to the colour yellow (Economopoulos, 1989; Katsoyannos, 1989).

Similarities to Other Species/Conditions

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B. oleae is the only species in subgenus Daculus and as such has no immediate relatives.

Prevention and Control

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Regulatory Control
 
The plant quarantine risk posed by B. oleae is very low and normal anti-fruit fly regulations will seldom be essential. For information, many countries, such as mainland USA, forbid the import of susceptible fruit without strict post-harvest treatment having been applied by the exporter. This may involve fumigation, heat treatment (hot vapour or hot water), cold treatments, insecticidal dipping or irradiation (Armstrong and Couey, 1989). Irradiation is not accepted in most countries and fumigation is a hazardous operation. Heat treatment tends to reduce the shelf life of most fruits. The preferred method of regulatory control is to restrict imports of a given fruit to areas free from fruit fly attack.
 
Cultural Control and Sanitary Methods
 
Fruit wrapping, which applied against many other species, is not applicable to B. oleae due to the small size of fruit. In general, keeping orchards clear of fallen fruit is a common sanitary practice that prevents many larvae from being able to emerge and pupate. However, as most olive harvesting is carried out by encouraging fruit to drop onto ground covering nets, this would be difficult in practise. Rapid collection of the fallen fruit may help reduce fly populations. Delrio et al. (1995) recommended early harvesting (in late November) to minimize tainting.
 
Although B. oleae larvae are dependent on the presence of olive fruit, adults feed on a variety of organic sources including insect honeydews, plant nectar and sugars. B. oleae is attracted by the honeydew secreted by the scale Saissetia oleae (de Andrés 1991). It is therefore very important to control such pests that maintain or attract B. oleae populations within an orchard.
 
Biological Control
 
Biological control was attempted by P. Silvestri, who imported parasitoids to Italy from South Africa in 1911 and Eritrea in 1914. No introduced populations became established. Likewise, attempts to introduce natural enemies of fruit flies from Brazil were unsuccessful. A major effort to establish the Hymenopterid Psyttalia concolor began in Italy in 1914 using material imported from North Africa. This work continued until the end of the 1960s in Italy, and later in France, Greece and Spain. It resulted in establishment of the parasitoid, but although high levels of parasitism were obtained while regular releases were continued, populations dropped to low levels when introductions were stopped (Greathead, 1976). Further attempts to introduce biocontrol agents were made in Greece during an FAO program in the 1970s. Unsuccessful attempts were made to find more effective natural enemies in Africa (Monaco, 1978; Neuenschwander, 1982).
 
A possible explanation for the poor performance of parasitoids concerns the relative size of commercial olive fruit compared to the wild olive fruits. Wild fruit are small, usually 1 cm in diameter, whereas cultivated varieties can be thicker-skinned and are significantly larger, usually 2-3 cm in diameter (Bartolini and Petruccelli, 2002; Tzanakakis, 2003). Parasitoids penetrate the fruit with their ovipositors, and large cultivated olives have a pulp thickness that greatly exceeds the maximum ovipositor length, which is very short for most Psyttalia species <2mm). Due to the feeding of the second instar of the olive fruit fly larvae close to the pit of the fruit, the ability to successfully parasitize certain hosts is limited, a problem that has been well documented for other fruit fly parasitoids (Sivinski et al., 2001; Sivinski and Aluja, 2003; Wang et al., 2008b). Species that parasitize B. oleae in natural environments, though evidently well adapted to attacking small wild olives, may have difficulty reaching close to the pit of the fruit in larger cultivated olives.
 
A classical biological control program initiated in California, USA, in 2002 studied five braconid parasitoids from Africa or Central Asia for the control of the B. oleae (Sime et al., 2006a,b,c, 2007; Daane et al., 2008). In order of increasing ovipositor length, these were: Utetes africanus (Silvestri), P. lounsburyi, P. ponerophaga, P. concolor (Szιpligeti) and Bracon celer Szιpligeti (Wang et al., 2008b). Of these, U. africanus, P. lounsburyi, and P. ponerophaga appeared to be the most specialized on B. oleae (Neuenschwander, 1982; Wharton and Gilstrap, 1983; Sime et al., 2007; Daane et al., 2008). P. concolor females with longer ovipositors showed higher levels of parasitism of olive fruit than females with shorter ovipositors (Wang et al., 2009).
 
Chemical Control
 
Although cover sprays of entire crops are sometimes used, the use of bait sprays is both more economical and more environmentally acceptable. A bait spray consists of a suitable insecticide mixed with a protein bait. For spraying, no more than 300 ml of spraying solution per tree is applied into the olive canopy (more practical details of bait sprays are given by Bateman, 1982). Cover sprays consist of insecticide at a dose of ten times less than that of the recommended dose for bait sprays (Haniotakis, 2005).
 
Bait sprays consisting of 2% bait (protein) mixed with registered insecticides (Roessler, 1989) have been used for many years against B. oleae (Nadel, 1966; Manousis and Moore, 1987). The protein most widely used is hydrolysed, but some are acid hydrolysed and so highly phytotoxic. Smith and Nannan (1988) developed a system using autolysed protein, but this method has not been tested with B. oleae. See Roessler (1989), Jervis and Kidd (1993) and Delrio (1995) for reviews.
 
Bait sprays have the advantage over cover sprays in that the former can be applied as a spot treatment (Mangan, 2009), and there is a minimal impact on natural enemies (OEPP/EPPO, 2005). Bait sprays work mainly on female Tephritidae fruit flies, which are strongly attracted to the ammonia given off by protein sources. However, Fabre et al. (2003) reported that protein-based attractants can also attract males.
The effectiveness of bait sprays may depend on the proteins used (Varikou et al., 2014).
 
Control (larvicide) is recommended if over 15% of fruits destined for oil production are infested, and for table olives if 5% are infested (Fimiani, 1989).
 
Sterile Insect Technique (SIT)
 
SIT is an environmentally-friendly and species-specific method of pest control based on the release of large numbers of sterilised insects (Dyck et al., 2005). Competition for mating between wild and sterile males results in a decrease in the number of fertile matings and a decline in the overall population size. SIT has been successfully implemented against various pest insect species including several Tephritidae. However, despite decades of research aimed at developing an olive fly SIT programme using radiation-sterilised flies (Economopoulos et al., 1977), consistently poor results led to the eventual abandonment of trials. Key issues included low quality of the radiation-sterilised mass-reared flies, economical production of sufficient numbers of sterilised flies, and assortative mating of released and wild populations because of different preferred mating times (Economopoulos et al., 1977; Zevas and Economopoulos, 1981; Economoppoulos, 2001). Laboratory-reared wild-type flies were found to mate several hours earlier than wild flies (Zevas and Economopoulos, 1981), presumably due to differential selective pressures in the artificial laboratory-rearing environment. Male-only release was proposed as a solution (Economoppoulos, 2001). 
 
Mass Trapping
 
Mass trapping involves attracting adult B. oleae to a trap device by a trophical, sexual or colour attractant (Economopoulos, 1989). Traps can be treated by insecticide or filled with an attractant-insecticide solution (Haniotakis et al., 1983; Barcley and Haniotakis, 1991; Bjeliš, 2006). The main advantage of mass trapping is the exclusion of chemicals from the whole tree canopy.
 
A sex pheromone released by virgin females attracts male B. oleae (Haniotakis, 1974; Haniotakis et al., 1977). The principal component of this sex pheromone was identified as 1,7-dioxaspiro [5.5] undecane (Baker et al., 1980). 1,7 dioxaspiro [5.5] undecane is also strongly attractive to males (Jones et al., 1983).
 
Mass trapping of both sexes was developed by Haniotakis et al. (1991) using a combination of a food attractant, a phagostimulant, a male sex pheromone and a female aggregation pheromone.
In practice, mass trapping cannot fully control B. oleae and often complementary bait sprays are necessary in order to reduce fly populations and avoid infestation (Varikou et al., 2009).
 
Field Monitoring
 
Monitoring may be carried out by i) installation of protein bait/McPhail traps loaded with an attractant solution (see image), for recording the adult population in an olive orchard, and ii) by dissection of fruit to look for larvae and estimate infection levels. 
 
i) Suitable traps were described by White and Elson-Harris (1994). McPhail traps baited with 2% ammonium sulfate solution or hydrolyzed protein have been used for over 50 years in several Mediterranean countries for monitoring B. oleae (Rössler, 1989; Katsoyannos, 1992; Haniotakis, 2005). McPhail traps are plastic or usually glass containers with a reservoir for liquid baits; flies enter from the bottom of the trap through an opening and then drown in the solution. Every five to seven days the attractant solution is replaced with fresh solution and the captured adult flies are counted and their sex distinguished (Varikou et al., 2013). Protein hydrolysates have proved more attractive than ammonium salts (Orphanidis et al., 1958; Prokopy and Economopoulos, 1975; Prophetou et al., 2003). However, traps baited with ammonium sulfate can be inefficient for recording adult B. oleae populations (Neuenschwander and Michelakis, 1979). Sometimes yellow sticky panel traps are used for monitoring an adult population, though they give a poor representation of B. oleae population density in an olive grove and can indicate a significantly lower B. oleae population than McPhail traps indicate (in September) (Varikou et al., 2013). Though considered low-tech by some (Bueno and Jones, 2002), McPhail traps are still in use today and give useful information, especially regarding female B. oleae activity as well as including female fecundity levels, which is obtained by dissecting females caught in these traps and assessing the stage of development of the ovaries.
 
ii) In order to estimate the percentage alive, dead and total infestation by immature stages inside the olive fruit, olive fruit samples should be randomly taken at least twice a month in the olive orchard, from the time the pit hardens until the harvest period. The following can be determined as follows:
 
% Live infestation = live (eggs + larvae (L1 +L2 +L3) + pupae)
% Dead infestation = dead (eggs + larvae (L1 +L2 +L3) + pupae) + parasitized ((larvae (L2 +L3) + pupae))
% Total infestation = % alive infestation + % dead infestation + exit holes
 
The use of such information of both traps and olive fruit samples can be combined with climatic data in order to make predictions and apply preventive measures.

References

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29/09/14 updated by:

K Varikou, Institute for Olive tree and Subtropical Plants, Crete, Greece

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