Bactrocera tryoni (Queensland fruit fly)

Pictures

Picture	Title	Caption	Copyright
Title Adult Caption B. tryoni adult. Copyright CABI	Adult	B. tryoni adult.	CABI

Identity

Preferred Scientific Name

• Bactrocera tryoni (Froggatt)

Preferred Common Name

• Queensland fruit fly

Other Scientific Names

• Bactrocera (Bactrocera) tryoni (Froggatt)
• Chaetodacus sarcocephali Tryon
• Chaetodacus tryoni (Froggatt)
• Dacus ferrugineus tryoni (Froggatt)
• Dacus tryoni (Froggatt)
• Strumeta melas Perkins & May
• Strumeta tryoni (Froggatt)
• Tephritis tryoni Froggatt

International Common Names

• English: qfly; Queensland fruitfly
• Spanish: mosca de la fruta de Queenslandia
• French: mouche des fruits de Queenslande

Local Common Names

• Germany: Fruchtfliege, Queensland-

EPPO code

• DACUTR (Bactrocera tryoni)

Summary of Invasiveness

B. tryoni, the Queensland fruit fly, is the most costly horticultural pest in Australia and has invaded several countries in the surrounding region (White and Elson-Harris, 1994). It has the potential to spread to many places around the world because of its wide climatic and host range (Meats 1989b; Sutherst et al., 2000) and a tendency to be carried by human travellers at the larval stage inside infested fruit. B. tryoni is a very serious pest of a wide variety of fruits throughout its range. Damage levels can be anything up to 100% of unprotected fruit.

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

B. tryoni was originally described as Tephritis tryoni by Froggatt in 1897 and two little-used synonyms are attributable to Tryon. The status of B. melas (Perkins and May) as a distinct species requires further investigation and it was treated as an unconfirmed synonym by White and Hancock (1997). B. tryoni could be confused with B. aquilonis (May), a species known only from northern Western Australia and the Northern Territory. There are some other generic combinations, most notably Dacus tryoni. It is a member of subgenus Bactrocera and can therefore sometimes be cited as Bactrocera (Bactrocera) tryoni.

Description

Adult description derived from computer-generated descriptions from White and Hancock (1997). Larval description from White and Elson-Harris (1994).

Adult

Head: Pedicel+1st flagellomere not longer than ptilinal suture. Face with a dark spot in each antennal furrow; facial spot large, round to elongate. Frons - 2 pairs frontal setae; 1 pair orbital setae.

Thorax: Predominant colour of scutum red-brown. Postpronotal (=humeral) lobe entirely pale (yellow or orange). Notopleuron yellow. Scutum with lateral postsutural vittae (yellow/orange stripes), which do not extend anterior to suture, are tapered, and reach to the posterior supra-alar seta. Scutum without a medial vitta. Scutellum entirely yellow (except for narrow basal band). Anepisternal stripe not reaching anterior notopleural seta. Yellow marking on both anatergite and katatergite. Postpronotal lobe (=humerus) without a seta. Notopleuron with anterior seta. Scutum with anterior supra-alar setae and prescutellar acrostichal setae. Scutellum without basal setae.

Wing: length 4.8-6.3 mm. With a complete costal band which may extend below R2+3, but not to R4+5; not expanded into a spot at apex. With an anal streak. Cells bc and c coloured. No transverse markings. Cell bc without extensive covering of microtrichia. Cell c with extensive covering of microtrichia. Cell br (narrowed part) with extensive covering of microtrichia.

Legs: All femora yellow / pale.

Abdomen: Predominant colour red-brown. Tergites not fused. Abdomen not wasp waisted. Pattern on abdomen diffuse to distinct. Tergite 3 darkened basally and laterally. Tergite 4 dark laterally. Medial longitudinal stripe on T3-5.

Terminalia and secondary sexual characters: Male wing without a bulla. Male tergite 3 with a pecten (setal comb) on each side. Male sternite 5 V-shaped posteriorly. Surstylus (male) without a long posterior lobe. Wing (male) with a deep indent in posterior margin. Hind tibia (male) with a preapical pad. Aculeus apex pointed.

Egg

The egg of B. oleae was described in detail by Margaritis (1985) and those of other species are probably very similar. Size, 0.8 mm long, 0.2 mm wide, with the micropyle protruding slightly at the anterior end. The chorion is reticulate (requires scanning electron microscope examination). White to yellow-white in colour.

Third instar larva

Larvae medium-sized, length 8.0-11.0 mm; width 1.2-1.5 mm.
Head: Stomal sensory organs large, rounded, each with 3 sensilla and surrounded by 6 large unserrated preoral lobes; oral ridges with 9-12 rows of deeply serrated, bluntly rounded teeth; 8-12 small, serrated accessory plates; mouthhooks large, heavily sclerotised, without preapical teeth. Thoracic and abdominal segments: a band of small posteriorly directed spinules encircling anterior portion of each thoracic segment. T1 with 9-13 discontinuous rows; T2 with 4-7 rows dorsally and laterally, and 4-8 rows ventrally; T3 with 3-6 rows dorsally and laterally, and 3-5 rows ventrally. Creeping welts with 2-3 anteriorly directed and 3-8 posteriorly directed rows of spinules. A8 with well defined intermediate areas and large sensilla. Anterior spiracles: 9-12 tubules. Posterior spiracles: placed just above midline; each spiracular slit about 3 times as long as broad. Dorsal and ventral spiracular hair bundles of 12-17, broad, stout, often branched hairs; lateral bundles of 5-9 similar hairs. Anal area: lobes well defined, surrounded by 3-5 discontinuous rows of spinules, becoming longer and stouter below anal opening.

Puparium

Barrel-shaped with most larval features unrecognisable, the exception being the anterior and posterior spiracles which are little changed by pupariation. White to yellow-brown in colour. Usually about 60-80% length of larva.

Distribution

B. tryoni is found throughout the eastern half of Queensland, eastern New South Wales, and the extreme east of Victoria. In 1989 it became established in the Perth area of Western Australia and it was declared eradicated by 1991. There have also been outbreaks in South Australia and although action to eradicate is taken, cool winters may also account for its lack of establishment. The record for Tasmania in CABI/EPPO (1998) is an error. B. tryoni has never been found in Tasmania.

There is a preliminary report of an isolated population of B. tryoni detected in an urban residential area of central Auckland, New Zealand (IPPC, 2015).

A few males have been trapped in Papua New Guinea but it is unlikely to be established there (Drew, 1989). It is also adventive in French Polynesia (Austral and Society Islands) and New Caledonia and has twice been adventive in Easter Island, but eradicated (Bateman, 1982). The distribution of this species was mapped by Drew (1982) and IIE (1991).

B. tryoni has a distribution almost entirely sympatric with B. neohumeralis, and both species attack a similar range of hosts, although B. tryoni is by far the more damaging. These two species mate at different times of day (B. tryoni at dusk; B. neohumeralis at midday). There is genetic evidence that the two species hybridize (Morrow et al., 2000). Reports of hybridization between B. tryoni and B. aquilonis (EPPO, 2002) (a similar species in the Northern Territory) are almost certainly erroneous as those two species lack sympatry.

See also CABI/EPPO (1998, No. 31).

Distribution Table

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.

Introductions

Risk of Introduction

The major risk is from the importation of fruit containing larvae, either as part of cargo, or through the smuggling of fruit in airline passenger baggage or mail. For example, in New Zealand Baker and Cowley (1991) recorded 7-33 interceptions of fruit flies per year in cargo and 10-28 per year in passenger baggage. Private individuals who successfully smuggle fruit are likely to discard it when they discover that it is rotten. An isolated catch of B. tryoni in a cue lure baited trap in California (Foote et al., 1993) probably had an origin of this sort.

Habitat List

Hosts/Species Affected

B. tryoni is the most serious insect pest of fruit and vegetable crops in Australia, and it infests all commercial fruit crops, other than pineapple (Drew, 1982). Most of the data given here are from the host catalogue of Hancock et al. (2000), much of which derives from host data gathered in a major survey in the Cairns area. That revised list recorded B. tryoni from 49 families of plants, represented by 234 species.

In addition to the hosts listed, Garcinia dulcis, Diplocyclos palmatus, Flaacourtia inermis, Sandoricum indicum, Artocarpus odoratissima, Casimiroa tetrameria, Murraya exotica and Solanum muricatum are economically important hosts of B. tryoni. Other major wild hosts are Annona atemoya, Terminalia aridicola, T. muelleri, T. platyphylla, T. sericocarpa, T. subacroptera, Syzgium suborbiculare, S. tierneyanum and Nauclea orientalis.

Host Plants and Other Plants Affected

Growth Stages

Symptoms

List of Symptoms/Signs

Biology and Ecology

Eggs are laid below the skin of the host fruit. These hatch within 2-3 days and the larvae feed for another 10-31 days. Pupariation is in the soil under the host plant for about 7 days but may be delayed under cool conditions. Adults occur throughout the year in 4-5 overlapping generations and overwinter as adults; up to 70 individuals have been recorded as developing from a single infested fruit (Christenson and Foote, 1960). Adult flight and the transport of infected fruit are the major means of movement and dispersal to previously uninfected areas. Many Bactrocera spp. can fly 50-100 km (Fletcher, 1989). Male B. tryoni are collected in very large numbers in cue lure traps, which will also trap B. neohumeralis in slightly lower numbers in most of its range (Osborne et al., 1997).

Climate

Natural enemies

Notes on Natural Enemies

Bactrocera spp. in general can be attacked as larvae either by parasitoids or by vertebrates eating fruit (either on the tree or as fallen fruit). Mortality due to vertebrate fruit consumption can be very high, as can puparial mortality in the soil, either due to predation or environmental mortality (see White and Elson-Harris, 1994, for brief review). Parasitoids appear to have little effect on the populations of most fruit flies and Fletcher (1987) noted that 0-30% levels of parasitism are typical. To date, complete biological control in the classical sense, has never been achieved for any Bactrocera or Dacus spp. (Wharton, 1989). Three opiine parastoids (Hymenoptera: Braconidae), Fopius arisanus (Sonan), Diachasmimorpha tryoni (Cameron) and D. kraussii (Fullaway) may have potential as biological control agents (Rungrojwanich and Walter, 2000; Quimio and Walter, 2001; Spinner et al., 2011). These species have established following introduction in Australia. Their ecology throughout their ranges requires study and no augmentative releases have been made. In some places frugivorous birds and rodents can destroy a large percentage of wild fruit that would be otherwise available to fruit flies or may have fruit fly larvae already in them (Drew, 1987).

Due to difficulties in verifying the identifications of both parasitoids and (in some cases) the fruit fly hosts, no attempt has been made to catalogue all natural enemy records; see White and Elson-Harris (1994) for major sources. Consequently, no comprehensive list of parasitoid records is given here; those listed were extracted from Waterhouse (1993) and Wharton and Gilstrap (1983).

Means of Movement and Dispersal

Natural Dispersal

This is normally limited to about 1 km.

Accidental Introduction

Jump dispersal, such as hitch-hiking in infested fruit in luggage, cargo and vehicles is common. Adventitious introduction by human agency does not always lead to establishment; in South Australia 71% of incipient incursions did not establish to a stage that warranted insecticidal or other treatments (Meats et al., 2003). Most released B. tryoni do not disperse far from their point of origin (~45% <100 m; ~95% < 1 km) (Meats and Edgerton, 2008) and this is consistent with the finding that the spread of incipient populations is also limited to ~1 km (Maelzer et al., 2004).

Intentional Introduction is unlikely.

Pathway Causes

Pathway Vectors

Plant Trade

Impact Summary

Impact

This is a very serious pest of a wide variety of fruits throughout its range. Damage levels can be anything up to 100% of unprotected fruit. In Australia potential losses if fruit flies were not controlled have been estimated at A$100 million a year (Anonymous, 1986), and most of this would be attributable to B. tryoni.

Economic Impact

There are about 4,500 species of tephritid flies (Diptera: Tephritidae). Approximately one third are frugivorous and around 250 are considered economic pests, with 23 of these known to be serious pests in Australia, Oceania and tropical Asia (White and Elson-Harris, 1992; Vijaysegaran, 1997). Adults of frugivorous Tephritidae lay their eggs beneath the skin of sound ripening fruit; the larvae feed within the fruit and cause direct damage and induce decay and premature fruit drop (Allwood and Leblanc, 1997). The percentage of produce lost has been estimated to be 10-50% in tropical Asia and Oceania and higher levels can occur in other parts of the world if control measures are not in place (Allwood and Leblanc, 1997). B. tryoni has a permanent presence in the eastern Australian states as well as the Northern Territory and the north of Western Australia (Meats, 2006; Cameron et al., 2010). Various statutory authorities have estimated economic losses in Australia due to B. tryoni to be between $28.5 million and $100 million per annum (Sutherst et al., 2000).

Environmental Impact

Impact on Natural Habitats

Impacts on natural habitats are unlikely because B. tryoni is a generalist and is mainly abundant in crops, villages and towns, and in natural habitats it would be only one of several fruit fly species present (Drew et al., 1984; Raghu et al., 2000).

Impact on Biodiversity

Impacts on biodiversity are also unlikely for the same reasons as for impacts on natural habitats. However, as far as fruit flies are concerned an unequivocal answer to the question - whether there is an impact of a pest species on other species in a district - should be assessed only by experiment or by incubating field-sampled fruit individually in order to rear out and identify surviving adult insects (see for example Gibbs, 1967; Fitt, 1986). Conversely, frugivorous birds and rodents can destroy a large percentage of wild fruit in some places that would be otherwise available to fruit flies or have fruit fly larvae already in them (Drew, 1987).

Risk and Impact Factors

• Invasive in its native range
• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Capable of securing and ingesting a wide range of food
• Highly mobile locally
• Has high reproductive potential
• Has high genetic variability

• Host damage
• Negatively impacts agriculture
• Negatively impacts livelihoods
• Damages animal/plant products
• Negatively impacts trade/international relations

• Highly likely to be transported internationally accidentally

Uses

No known positive value.

Detection and Inspection

Similarities to Other Species/Conditions

B. tryoni is separated from most of the other pest species by the coloured cells bc and c (i.e. the costal band extends from the wing base, not just from cell sc [the stigma]). However, it occurs sympatrically with B. neohumeralis, which also has that feature but from which it differs in having yellow postpronotal (=humeral) lobes. In Australia both species attack a similar range of hosts and can even be reared from the same individual specimens of field-collected fruit (Gibbs, 1967). These two species mate at different times of day (B. tryoni at dusk; B. neohumeralis ~ 10 AM–4 PM. There is no genetic evidence that the two species hybridize (Gilchrist and Ling, 2006).

B. tryoni is allopatric from B. aquilonis, from which it only differs morphologically in being darker in colour. Previous arguments about distinguishing B. tryoni from B. aquilonis in northern Australia are well discussed in Morrow et al. (2000; see also CABI/EPPO, 1998, No. 31) but the evidence and analysis provided by Cameron et al. (2010) favours the conclusion that B. tryoni is found in allopatric populations across northern Australia from north Queensland to the northwest coast of Western Australia.

The minimum characters which differentiate B. tryoni from all other Bactrocera and Dacus spp. (White and Hancock, 1997) are as follows: postpronotal lobe entirely yellow. Scutum predominantly red-brown; with lateral vittae (yellow stripes) not extended anterior of suture, posteriorly reaching to the posterior supra-alar setae; with prescutellar acrostichal setae. Anepisternal stripe not reaching as far as anterior notopleural seta. Wing cell c covered in microtrichia; cell bc devoid of microtrichia. Tergite 3 dark laterally and basally.

B. tryoni is larger than a house fly (wing length 4.8-6.3 mm).

Prevention and Control

Biological Control

Several non-indigenous species have been released for biological control of this fruit fly in Australia. Of these, only Fopius arisanus became established, and although it reduced the number of flies per fruit it had little effect on the percentage of fruits damaged (Waterhouse, 1993).

Regulatory Control

Many countries, such as the 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). Recent work on hot water dipping was reported by Waddell et al. (2000). Irradiation is not accepted in most countries and many have now banned methyl bromide fumigation. Heat treatment tends to reduce the shelf life of most fruits and so the most effective method of regulatory control is to preferentially restrict imports of a given fruit to areas free of fruit fly attack.

Cultural Control and Sanitary Methods

One of the most effective control techniques against fruit flies in general is to wrap fruit, either in newspaper, a paper bag, or in the case of long/thin fruits, a polythene sleeve. This is a simple physical barrier to oviposition but it has to be applied well before the fruit is attacked. Little information is available on the attack time for most fruits but few Bactrocera spp. attack prior to ripening.

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 (e.g. malathion) mixed with a proteinaceous bait (usually termed ‘protein’). Both males and females of fruit flies are attracted to protein sources emanating ammonia, so insecticides can be applied to just a few spots in an orchard and the flies will be attracted to these spots when they get near them during their daily foraging (Bateman et al., 1966 ab; Bateman, 1982). The protein most widely used in Australia was acid-hydrolysed yeast. This was neutralised by sodium hydroxide yielding a concentrate with a salt content of up to 50%. In South Australia an effective concentration was found to be strongly phytotoxic due to its high salt content. Thus from 1983 yeast autolysate was used instead (Madge et al., 1997). This product can be made cheaply from brewery waste (Umeh and Garcia, 2008). Horticultural mineral oil (HMO) is strongly repellent to female B. tryoni and can be used successfully to protect fruit in small crops, including home gardens (Nguyen et al., 2007; Meats et al., 2012).

Male Suppression/Annihilation Techniques and SIT.

The males of most pest species of Bactrocera are attracted to either cue lure (4-(p-acetoxyphenyl)-2-butanone) or to methyl eugenol (4-allyl-1,2-dimethoxybenzene). Males of B. tryoni are attracted to cue lure, sometimes in very large numbers. Combined with an insecticide it can be impregnated into small caneite blocks or other absorbent material. If these are distributed at sufficient density (~ 30m spacing) most males can be annihilated (Bateman, 1982). This has been termed the ‘male annihilation technique’ (MAT). Bateman et al. (1966a,b) pioneered combined MAT and bait spray in Australian coastal and inland towns and on Easter Island (Bateman et al.,1973; Bateman, 1982). This tactic is now used in are-wide management programmes.

The sterile insect technique (SIT) has been used for localised outbreaks in quarantined areas (Jessup et al., 2007).

Early Warning Systems

Many countries that are free of Bactrocera spp., such as the USA (California and Florida) and New Zealand, maintain a grid of methyl eugenol and cue lure traps, at least in high risk areas (ports and airports) if not around the entire climatically suitable area. The trap used will usually be modelled on the Steiner trap (White and Elson-Harris, 1994) or Lynfield (pot) trap (Cowley et al., 1990).

Field Monitoring

Monitoring is largely carried out by traps (as above) set in areas of infestation. However, there is evidence that some fruit flies have different host preferences in different parts of their range and host fruit surveys should also be considered as part of the monitoring process.

IPM

The control of tephritid fruit flies is practised in two ways. The first is area-wide control that requires quarantine regulations and expensive technology such as SIT in a restricted and defendable area, but may require grower and community participation (Jessup et al., 2007). Features include trap arrays for early warning and prompt responses, border inspections, community awareness programmes as well as bait-spraying and the male annihilation technique (MAT) (Jessup et al., 2007). A good example and case study is given by Lloyd et al. (2010).

The second is farmer-operated local or ‘crop by crop’ control and is generally suited to local economies with local (non-export) distribution and is particularly relevant to areas with naturally high endemic pest populations and to village horticulture in tropical Asia and the South Pacific islands (Allwood & Leblanc 1997; Vijaysegaran 1997), where high infestation rates would damage local economies and cause migration to towns.

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31/03/14 Updated by:

Alan Meats, University of Sydney, Australia

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