Drosophila suzukii (spotted wing drosophila)
- 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
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- 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
- Drosophila suzukii Matsumura
Preferred Common Name
- spotted wing drosophila
Other Scientific Names
- Drosophila (Sophophora) suzukii (Matsumura)
- Drosophila suzukii (Matsumura) Kanzawa
International Common Names
- English: cherry fruit fly; spotted wing drosophila; spotted-wing drosophila
Local Common Names
- English: cherry vinegar fly
- Germany: Kirschessigfliege
- Japan: oto shojobae; outou shoujyou bae
Summary of InvasivenessTop of page
The fruit fly D. suzukii is a fruit crop pest and is a serious economic threat to soft summer fruit. A polyphagous pest, it infests a wide range of fruit crops, included grape, as well as an increasing number of wild fruits. D. suzukii is an economically damaging pest because the females are able to infest thin-skinned fruits before harvest and the larvae destroy the fruit pulp by feeding. The species is endemic in Asia. It was first recorded as invasive in Hawaii in 1980 and then simultaneously in California and in Europe in 2008. Since 2008 it has spread rapidly throughout the temperate regions of North America and Europe, due to global trade and the initial lack of regulation over the spread of any Drosophila. This species has a high reproductive rate and short generation time; D. suzukii can theoretically have up to 13 generations per year, which may contribute towards rapid spread, given available suitable hosts. D. suzukii is listed on the EPPO alert list.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Diptera
- Family: Drosophilidae
- Genus: Drosophila
- Species: Drosophila suzukii
Notes on Taxonomy and NomenclatureTop of page
In June, 1916, maggots were found to be infesting pre-harvest cherries (Prunus avium) in Yamacho, Higashi Yamanashi County, Yamanashi Prefecture, Japan (Kanzawa, 1935). Infested fruit was collected and the adult flies that emerged were confirmed as a species of Drosophila (Kanzawa, 1935). The species was later described in 1931 by Dr Shounen Matsumura as Drosophila suzukii Matsumura, and he gave it the common name of cherry drosophila (Kanzawa, 1935).
D. suzukii has also been described in the Kashmir region of India as the D. suzukii subsp. indicus (Parshad and Paika, 1965).
D. suzukii belongs to the melanogaster species group of the subgenus Sophophora. The melanogaster group is further divided into species subgroups, one of which (the suzukii subgroup) comprises, together with 6 other subgroups, the ‘oriental lineage’ (Kopp and True, 2002; Schawaroch, 2002; van der Linde et al., 2010).Relationships between and within these subgroups are still far from being resolved, and the suzukii subgroup itself is commonly regarded as polyphyletic (Kopp and True, 2002). Recent papers suggested D. biarmipes is the sister taxon of D. suzukii (Yang et al., 2011; Chiu et al., 2013; Ometto et al., 2013; Rota Stabelli et al., 2013), in accordance with previous findings (Kopp and True, 2002; Barmina and Kopp, 2007); however, Prud’homme et al. (2006) and van der Linde and Houle (2008), instead supported D. subpulchrella as the sister species of D. suzukii (with D. biarmipes being the sister species of D. subpulchrella + D. suzukii). Genome scale data may help explore the relationship between D. suzukii and D. subpulchrella.
DescriptionTop of page
D. suzukii adults are 2-3 mm long with red eyes, a pale brown or yellowish brown thorax and black transverse stripes on the abdomen. The antennae are short and stubby with branched arista. Sexual dimorphism is evident: males display a dark spot on the leading top edge of each wing and females are larger than males and possess a large serrated ovipositor.
The eggs are oval (minor axis is 0.2 mm), milky-white, with two filaments (aeropyle or spiracle) at one end, 0.4 to 0.6 mm long.
The maggot-like larvae are white with visible internal organs and black mouthparts. They grow throughout three larval stages and when fully grown can reach 5.5 mm long and 0.8 mm wide.
The pupae are spindle-shaped, reddish-brown and bear two stalks with small finger-like projections, 3.5 mm long and 1.2 mm wide).
DistributionTop of page
D. suzukii is thought to be native of eastern and southeastern Asia, including China, Japan and Korea (Walsh et al., 2011), although little is known about its geographical origin. According to references reported by Hauser (2011) there is the possibility that the species is not native to Japan, but had been introduced into the country at the turn of the century.
D. suzukii has been introduced to several Hawaiian islands, including Oahu (Hauser, 2011). It has also been introduced into North America and Europe. D. suzukii has recently been recorded in Iran, indicating expansion of its territory into the Middle East (Parchami-Araghi et al., 2015). It has also been recorded in Réunion (EPPO, 2018).
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: 29 Apr 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|-Anhui||Present||Cited as recorded from Kai et al. 1993|
|-Jammu and Kashmir||Present||Native|
|Japan||Present, Widespread||Introduced||Invasive||The four main Islands of Japan, Ryukyu, Bonin, Kume-jima and Iriomote-jima Islands (DAFF, 2013)|
|Myanmar||Present, Widespread||The central northern regions and the highlands|
|North Korea||Present, Widespread||Native|
|South Korea||Present, Widespread||Native|
|Turkey||Present, Few occurrences|
|Bosnia and Herzegovina||Present, Localized|
|Greece||Present, Few occurrences|
|-Crete||Present, Few occurrences|
|Hungary||Present, Few occurrences|
|Poland||Present, Few occurrences|
|Portugal||Present, Few occurrences||Introduced||2012|
|Russia||Present, Few occurrences|
|-Russian Far East||Present, Few occurrences|
|Slovakia||Present, Few occurrences|
|United Kingdom||Present, Few occurrences||Introduced||2012|
|-England||Present, Few occurrences|
|-British Columbia||Present, Widespread||Introduced||2009|
|-Newfoundland and Labrador||Present|
|-Prince Edward Island||Present|
|Costa Rica||Absent, Unconfirmed presence record(s)|
|Mexico||Present, Localized||Introduced||2011||Invasive||Aguascalientes, Baja California, Colima, Guanajuato, Jalisco, Michoacán, Estado de México and Veracruz|
|United States||Present, Localized||Introduced||2008|
|French Polynesia||Present, Localized|
|-Distrito Federal||Present, Few occurrences|
|-Espirito Santo||Present, Localized|
|-Rio de Janeiro||Present, Few occurrences|
|-Rio Grande do Sul||Present|
|-Sao Paulo||Absent, Intercepted only|
|Chile||Present, Localized||Present: but managed|
|Ecuador||Absent, Unconfirmed presence record(s)|
History of Introduction and SpreadTop of page
Although considered native to East Asia, it is possible that D. suzukii is not native to Japan but was introduced into the country at the turn of the century. A detailed historic account of the dispersion of D. suzukii outside of its native area is given in Hauser (2011).
North America and Hawaii
In 1980 it was collected in Hawaii without any report of it causing economic damage. In September 2008 a sample of flies collected in a raspberry field in Santa Cruz County, California, USA, was the first detection of D. suzukii in mainland USA. The following spring several records of maggots were found in cherries. In 2009 D. suzukii had spread to more than 20 counties in California, the other Pacific Coast states of Oregon, Washington and British Columbia (Canada), as well as in Florida, USA.
In 2010 D. suzukii was detected in six other states in the USA (Utah, North Carolina, South Carolina, Wisconsin, Michigan and Mississippi). D. suzukii rapidly spread into other 15 states in USA in 2011, an additional nine States in 2012 and a further two in 2013. By late 2013, only eight US states were not invaded by D. suzukii: Arizona, Nevada, New Mexico, Oklahoma, Kansas, Nebraska, South Dakota and Wyoming (Burrack et al., 2012).
In Canada, D. suzukii spread rapidly in 2010. Already present in British Columbia, in that year it was also recorded in Alberta, Manitoba, Ontario and Quebec (Hauser, 2011).
The first detection and spread of D. suzukii in Europe was detailed by Cini et al. (2012). The first adults of D. suzukii were caught at the same time in Rasquera Province, Spain (Calabria et al., 2012) and in the Tuscany region, Italy (Raspi et al., 2011) in 2008. In 2009 D. suzukii adults were recorded in traps in other regions of Spain (Bellaterra, near Barcelona), France (Montpellier and Maritimes Alpes) and Italy (Trentino) (Grassi et al., 2009; Mandrin et al, 2010; Calabria et al., 2012). In Trentino, both first oviposition on wild hosts (Vaccinium, Fragaria and Rubus spp.) and economically significant damage on several species of cultivated berries were reported (Grassi et al., 2009).
By 2010-2011, the range of D. suzukii enlarged further, including other regions in Italy and France (Cini et al, 2012; Weydert et al, 2012) as well as expanding north and east, invading Switzerland (Baroffio and Fisher, 2011), Slovenjia (Seljiak, 2011), Croatia (Milek et al., 2011), Austria (Lethmayer, 2011), Germany (Vogt et al., 2012), Belgium (Mortelmans et al., 2012), the Netherlands (NPPO, 2012), the UK (EPPO, 2012) and Hungary (Kiss et al, 2013).
Risk of IntroductionTop of page
The global fresh fruit trade, coupled with the cryptic nature of the larvae to hide inside the fruit undetected until after transportation, facilitate the increasing distribution of this pest. Given its very rapid spread in Europe and North America in recent years, it seem likely that D. suzukii will continue to expand its range in these continents to some extent. Calabria et al. (2012) estimated that D. suzukii was able to spread approximately 1400 km a year.
HabitatTop of page
D. suzukii development is fostered by widespread cultivation of susceptible crops (mainly soft fruits and cherry) (Lee et al., 2011; Bellamy et al., 2013), distribution of cultivated land at different altitudes (offering a differentiated and extended fruit ripening period), proximity of forests and uncultivated or marginal areas with susceptible wild fruits. D. suzukii seems to have important relationships with forests and woodland, where it can find a suitable microclimate and host plants year-round (Grassi et al, 2011).
The establishment of D. suzukii in more northern regions, where there are harsh winters, is likely to depend on the presence of favourable overwintering sites that are generally associated with human habitation (Dalton et al., 2011; EPPO, 2013a).
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Principal habitat|
|Terrestrial||Managed||Protected agriculture (e.g. glasshouse production)||Principal habitat|
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat|
|Terrestrial||Managed||Urban / peri-urban areas||Secondary/tolerated habitat|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat|
|Terrestrial||Natural / Semi-natural||Scrub / shrublands||Principal habitat|
Hosts/Species AffectedTop of page
D. suzukii is predisposed towards infesting and developing in undamaged, ripening fruit. Fruits become susceptible to D. suzukii as they start to change colour, which coincides with softening skins and higher sugar levels (Burrack et al., 2013). There are differences in fruit susceptibility within species and among varieties within the same fruit species (Lee et al., 2011). Fruit penetration force is one potential measure of host susceptibility, but host attractiveness will likely depend upon additional factors, such as soluble sugar content (Burrack et al., 2013). If there is no suitable fruit available, then D. suzukii will attack damaged or deteriorating fruit (Kanzawa, 1935; Lee et al., 2011). Non-commercially marketed fallen fruit or damaged fruit of the following plant hosts may also be attacked: Prunus persica, Malus pumila var. domestica, Prunus triflora, Prunus armeniaca, Pyrus pyrifolia, Pyrus sinensis, Eriobotrya japonica, Lycopersicum esculentum (Kanzawa, 1939) and Rubus microphyllus (Mitsui et al., 2010), as well as over-ripe figs still on the tree (Ficus carica) (Yu et al., 2013).
D. suzukii has been reared from rotting strawberry guava fruits (Psidium cattleianum) collected from trees and on the ground (Kido et al., 1996). It has been observed feeding upon injured or culled fruit including apple and oranges (Walsh et al., 2001).
A recently extensive study on seasonal life cycles and food resources of D. suzukii from low to high altitudes in central Japan (Mitsui et al., 2010) confirmed that D. suzukii emerges almost only from fruits. Some D. suzukii individuals emerged from the fruits of Rubus crataegifolius, Alangium platanifolium, Cornus kousa, Torreya nucifera and Viburnum dilatatum. Grassi et al. (2011) reared D. suzukii also on Prunus laurocerasus and Mann and Stelinski (2011) reported Ribes spp. as host plant of D. suzukii, but this latest observation has not been confirmed in Europe. D. suzukii adults also emerged from the flowers of Styrax japonicus (Mitsui et al., 2010), and in early spring in southern Japan it was also observed to breed on the flowers of Camellia japonica (Nishiharu, 1980).
This field of work is not well described, and so the list of Host Plants and Other Plants Affected contains probable as well as reported hosts.
Host Plants and Other Plants AffectedTop of page
SymptomsTop of page
D. suzukii larvae cause damage by feeding on the pulp inside fruit and berries. The infested fruit begins to collapse around the feeding site causing a depression or visible blemish on the fruit. The oviposition scar exposes the fruit to secondary attack by pathogens and other insects, which may cause rotting (Hauser et al., 2009; Walton et al., 2010).
List of Symptoms/SignsTop of page
|Fruit / internal feeding|
|Inflorescence / external feeding|
Biology and EcologyTop of page
The D. suzukii genome is comparable in size and repeat content to other Drosophila species. Genome-scale relaxed clock analyses indicate a late Miocene origin of D. suzukii, concomitant with paleo-geological and climatic conditions that suggest an adaptation to temperate climates. Furthermore, all the analyses support a close genetic relationship between D. suzukii and D. biarmipes but a low nucleotide substitution rate in comparison with the lineage leading to D. biarmipes (Yang et al., 2012; Chiu et al., 2013; Ometto et al., 2013).
Detailed information about the biology of D. suzukii is available in Kanzawa (1935). D. suzukii overwinter as adults (Dalton et al., 2011). Flies emerge in spring, but some adults may be active during warm winter days. Eggs are laid in ripening fruits and the number of eggs per fruit ranges from one to several, scattered over the fruit. D. suzukii host selection under field conditions may differ among species and among varieties within a species, and laboratory observations suggest that fruit firmness may be one driver of this selection (Burrack et al., 2013). Egg-laying lasts 10-65 days with up to 21 eggs laid per day. Each female can lay 195 eggs during her lifetime (Kanzawa, 1939; Tochen et al., 2014). Eggs hatch in 1-3 days, larvae mature in 3-13 days and most of them pupate in the fruit, but some drop and creep into the soil. Pupae period lasts between 4 and 43 days. The minimum, optimal and maximum intrinsic rate of natural increase was estimated at 13.4, 21.0 and 29.4°C by Tochen et al. (2014).
Mating of new adults can happen any time of the day, but it can be observed more often during the day when the temperature is relatively high. Males are always active, but females are passive. Courtship was described by Kanzawa (1939) and the role of the visual stimulus in the courtship was investigated by Fuyama (1979). The crucial role of specific substrate-borne vibrations during courtship in D. suzukii was demonstrated by Mazzoni et al. (2013). Females oviposit after mating and repeat mating days later (Kanzawa, 1939). Oviposition generally occurs from April to November. Mitsui et al. (2010) reported that D. suzukii individuals collected in autumn were reproductively immature, suggesting a winter reproductive diapause. No reproductive behaviour was observed during laboratory experiments where D. suzukii was kept for the entire life cycle at temperatures below 10°C (Mitsui et al., 2010; Tochen et al., 2014). Mitsui et al. (2010) assumed that the males which were emerging in those experimental conditions were rendered sterile and were unable to mate successfully with emerged females. Low levels of reproduction or no reproduction were found at temperatures above 30°C (Tochen et al., 2014).
The life cycle from egg hatching to adult emergence ranges from about 9-10 days to 21-25 days at 25°C and 15°C, respectively (Kanzawa, 1939). Recent laboratory observations recorded the development from egg to egg-laying female as ranging from about a week to 12-15 days at 21.1°C and at 18.3°C, respectively (Walsh et al., 2011).
Observation across a wide geographical range in Asia indicated that the number of generation per year could range from 3 to 13 depending on the climatic conditions (Kanzawa, 1939). According to the degree-day model developed by Coop (2010), D. suzukii is estimated to carry out 3 to 9 generations per year in the West United States, Canada and northern Italy.
The lifespan of adults is 20-56 days, but some overwintering adults lived for more than 200 days (Kanzawa, 1935). Female adult longevity ranged from 35 days at 10°C to 2 days at 30°C (Tochen et al., 2014). Acclimated adult D. suzukii can survive for up to 88 days at a constant 10°C, with no marked change in mortality when flies are subjected to a 7-day freeze period. Adult longevity decreases progressively at a constant temperature below 10°C (Dalton et al., 2011).
Some adults (males and females) overwinter under extended suboptimal cold conditions (Dalton et al., 2011). The lifespan of overwintering adults is considerably longer than non-overwintering adults and many survive until next May or June (Kanzawa, 1939). Females are more likely to overwinter than males.D. suzukii becomes mobile above 5°C, and if the average temperature rises beyond 10°C it starts to become active. If the temperature is high enough during the day, D. suzukii starts to oviposit. It is the most active between 20° and 25°C, but not very active when the temperature reaches 30°C (Kanzawa, 1939; Hamby et al., 2013). Hamby et al. (2013) reported dawn and dusk as the most active periods.
Adults often feed on fruits that has been split or damaged by birds. D. suzukii gathers on fruit that have dropped onto the ground and are spoiled or fermented. If there is no fruit juice available, D. suzukii can feed on sap from wounded oak trees (Kanzawa, 1939; Lee et al., 2011; Bellamy et al., 2013).
D. suzukii has been reported to vector yeasts and bacteria (Hamby et al., 2012; DAFF, 2013). Both larvae and adult of D. suzukii have been reported to be associated with yeast, predominantly Hanseniaspora uvarum (Hamby et al., 2012).
No differences have been observed in thermal tolerance between cool and warm temperate strains of D. suzukii. Their evolutionary capacity to increase cold tolerance seems to be limited (Kimura, 2004). To overcome deficiencies in cold tolerance, it is possible that D. suzukii may be behaviorally adapted to overwinter in man-made protected habitats (Kimura, 2004; Dalton et al., 2011).
ClimateTop of page
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Tolerated||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Cw - Warm temperate climate with dry winter||Preferred||Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)|
|D - Continental/Microthermal climate||Preferred||Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)|
|Dw - Continental climate with dry winter||Preferred||Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Asobara japonica||Parasite||Larvae/Pupae||to genus|
|Asobara rufescens||Parasite||Larvae||to genus|
|Asobara tabida||Parasite||Larvae||to genus|
|Cardiastethus fasciventris||Predator||Arnó et al., 2012|
|Cardiastethus nazarenus||Predator||Arnó et al., 2012|
|Dicyphus tamaninii||Predator||Arnó et al., 2012|
|Ganaspis xanthopoda||Parasite||Larvae||to genus|
|Isaria fumosorosea||Pathogen||Naranjo-Lázaro et al., 2014|
|Leptopilina boulardi||Parasite||Larvae||to genus|
|Leptopilina heterometra||Parasite||Larvae||to genus|
|Metarhizium anisopliae||Pathogen||Naranjo-Lázaro et al., 2014|
|Orius laevigatus||Predator||Arnó et al., 2012|
|Pachycrepoideus vindemmiae||Parasite||Pupae||to genus|
Notes on Natural EnemiesTop of page
Parasitoid wasps target Drosophila spp. and have potential as biocontrol agents of D. suzukii (Kanzawa, 1939). A number of hymenopteran parasitoids have been reported in association with D. suzukii in its native area. In particular, species of the genera Ganaspis and Leptopilina (Hymenoptera: Figitidae) and Trichopria (Hymenoptera: Diapriidae) are reported as parasitoids of D. suzukii in Japan (Cini et al., 2012). Ganaspis species showed the highest rates of D. suzukii parasitism. Ganaspis species lay eggs in larvae that are feeding in fruits and exhibit a high level of specificity for D. suzukii. By contrast, Leptopilina japonica and Asobara japonica (Hymenoptera: Braconidae) were only able to attack D. suzukii larvae and pupae in fallen decaying fruits, and also attacked a wide range of drosophilid hosts (Mitsui et al., 2007; Ideo et al., 2008; Mitsui and Kimura, 2010; Novkovic et al., 2011; Kasuya et al., 2013). Leptopilina heterotoma and Pachecrepoides vindemiae have been found to attack D. suzukii in newly-invaded production regions in Pacific North America and in northern Italy (Rossi-Stacconi et al., 2013).
Entomopathogenic fungi such as Isaria fumosorosea and Metarhizium anisopliae are also being assessed as potential biological control agents (Naranjo-Lázaro et al., 2014).
Means of Movement and DispersalTop of page
Natural Dispersal (Non-Biotic)
D. suzukii, as a fruit-specialist species among drosophilid flies, performs seasonal migration between low altitudes, which can be resource-poor in the summer, and high altitudes, where it is thought to exploit further resources (Mitsui et al., 2010). However, additional data needed in order to support this hypothesis.
The key pathway for the introduction of D. suzukii into new areas is by traded host fruits. Its rapid dispersal worldwide is in part due to increasing global fresh fruit trade and the cryptic nature of larvae hidden inside fruit, which means they are often undetected until after transportation (Gerdeman and Tanagoshi, 2011). Consequently, passive diffusion is likely the main cause of the spread of D. suzukii (Westphal et al., 2008; Cini et al, 2012; EPPO, 2013a). Even though additional analyses on a larger number of specimens are needed, the similarities of the introduction dates in North America and in Europe, along with the same COI haplotype found in both areas, suggest that the two invasions could be related (Calabria et al. 2012; Freda and Braverman, 2013). Calabria et al. (2012) stated that D. suzukii was able to spread approximately 1400 km in one year, but they could not say if the dispersion was active or passive via infested fruits. D. suzukii may also be introduced to new areas via the transport of flowers, although this is less likely.
Pathway CausesTop of page
Pathway VectorsTop of page
Impact SummaryTop of page
|Environment (generally)||Positive and negative|
Economic ImpactTop of page
The damage caused by D. suzukii larvae renders the fruit unmarketable (Bolda et al., 2010). Assessments of the economic impact of D. suzukii are relatively scarce and most focus on California, USA (Bolda et al., 2010; Goodhue et al., 2011), or the Trentino region in Europe (De Ros et al., 2013).
In 2008 economic losses (based on maximum reported yield losses) for California, Oregon and Washington were estimated at 40% for blueberries, 50% for caneberries, 33% for cherries and 20% for strawberries. Production in these three states could sustain $511 million in damages annually because of D. suzukii (Bolda et al., 2010). In California alone, the estimated decrease of the gross revenue due to D. suzukii infestation in the absence of management has been estimated at 37% for raspberry and 20% for processed strawberries (Goodhue et al., 2011).
Crop losses of 20-40% were reported from both Washington and Oregon states’ 2009 late season blueberries and caneberries (Gardeman and Tanigoshi, 2011). Growers in small fruit production regions in coastal Pacific Northwest, USA, currently apply pesticides from 5-7 times per season on average (J Flake, pers comm.). When taking into consideration current crop levels, input costs, fruit drop due to machinery rubbing against fruit canopies and loss due to D. suzukii infection, the annual costs to control D. suzukii in Oregon’s affected small fruit industries ranged between $12 and$16 million dollars annually, depending on the level of infestation that season (Julian et al., 2011). These costs are only associated with management techniques and do not take into consideration market loss or losses associated with altered processing practices, or downgrading of fruit.
Chemical control of D. suzukii may also lead to the rejection of exported fruits due to residual pesticide levels exceeding the maximum residue limits (Haviland and Beers, 2012).
Risk and Impact FactorsTop of page
- Parasitism (incl. parasitoid)
- 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
UsesTop of page
Apart from the use of laboratory cultures for research purposes, no human uses of D. suzukii have been described.
Uses ListTop of page
- Research model
DiagnosisTop of page
Molecular identification is possible by amplification of the barcode COI gene with universal primers (Folmer et al., 1994, Grassi et al., 2011; Calabria et al., 2012; Freda and Braverman, 2013; Chiu et al., 2013). DNA barcoding is the only fully reliable identification method (Freda and Braverman, 2013).
Detection and InspectionTop of page
Detailed morphological description of each stage is given by Kanzawa (1935). A more recently updated description, including references for additional morphological details, is given by Hauser (2011), and another by Vlach (2010), who published a dichotomous key for easy identification. An easy-to-use description of the combination of diagnostic characters that could be used for tentative identification of D. suzukii within its subgroup is given by both Hauser (2011) and Cini et al. (2012). Fruit infestation symptoms are described by Walton et al. (2010).
The dark spots on the male wings together with two sets of black tarsal combs make the identification of the males relatively easy, although the males of some other species do also have wing spots. The wing spots of D. subpulchrella are particularly similar in shape and position to those of D. suzukii. Males without dark wing spots can occur, as it takes two full days before the spots become obvious, although they start to appear within 10 hours of emergence at high temperatures.
The situation is complex for the eggs, larvae and pupae, as no reliable morphological diagnostic features have been identified (Okada, 1968). The eggs of D. suzukii have two respiratory appendages but this character is not species-specific. Instar stages can be estimated by the size of larvae and the colour of the mouthparts, but it is most accurately judged by pre-respiratory ducts (Kanzawa, 1935; Walsh et al., 2011).
Larvae are often undetected inside the fruit. The infested fruits can be detected only by visual inspection under optical magnification (15-20 x magnification). Detection of larvae inside the fruits can also be performed by immersion of fruit samples in sugar or salt solution. Sugar solution can be prepared using approximately 1 part sugar to 6 parts water in order to reach at least 15°Brix. Gently crush the fruits and wait for 10 minutes until the larvae in the sample float to the surface. The same procedure can also be followed using a salt solution, adding 1 part salt to 16 parts water (BCMA, 2013).
Traps baited with different baits have been proposed for detecting adults in the field. Traps can be installed around a site where fruits for shipment are stored, and for early detection in potentially newly-invaded areas, such as near fruit markets, warehouses of food retailers and sites where rotten fruits are disposed. For more information on traps and baits, see the Monitoring and Surveillance section in Prevention and Control.
Similarities to Other Species/ConditionsTop of page
The distinguishing features of the two sexes (serrated ovipositor and black wing spots) are also present in 150 other Drosophila species, making species identification difficult in areas where they are sympatric. D. subpulchrella Takamori and Watabe males’ black spots are very similar in shape and position to those of D. suzukii (Takamori et al., 2006). The occasional lack of wing black spots in teneral male D. suzukii could lead to misidentification with other closely related Drosophila species whose males do not have spots on the wing, including: D. ashburneri Tsacas, D. immacularis Okada, D. lucipennis Lin, D. mimetica Bock and Wheeler, D. oshimai Choo and Nakamura and D. unipectinata Duda.
Other characteristics can instead be used for identification, such as the sex combs on the foretarsi; D. suzukii has one row of combs on the first and one row on the second tarsal segment while D. biarmipes has two combs on the first tarsomere,
Females can also pose problems with identification. On the basis of the shape and length of the ovipositor, D. suzukii can be easily discriminated from related species, such as D. biarmipes, but not easily from other species such as D. immigrans Sturtevant and D. subpulchrella (Takamori et al., 2006), which possess very similar ovipositors (Hauser, 2011). In such cases, a final determination should be made by a taxonomist, based on the relative size of spermatheca compared to the size of the ovipositor (Hauser, 2011).
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.
Emergency measures to prevent the introduction of D. suzukii via host fruits (and, to a lesser extent, flowers) included carbon dioxide/sulphur dioxide fumigation. According to the preliminary data available the treatment causes 100% mortality of D. suzukii. Verification of the treatment efficacy by inspection of fruit cuts under optical magnification is an additional emergency measures (DAFF, 2013).
Early Warning Systems
Contrary to some other potentially invasive pests, D. suzukii is not subject to regulation either in Europe or the United States. Consequently there are neither official limits on the movement of host crops from infested areas, nor coordinated actions for monitoring its presence in new areas. An early warning system with baited traps is sometimes established as a volunteer-based monitoring network (Burrack et al., 2012).
Due to its high reproductive capacity and dispersal abilities of this pest, early warning systems should be considered vital in areas currently free of D. suzukii. By the time D. suzukii was detected in both Europe and USA it had already established itself to such an extent that eradication was deemed impossible in both continents (EPPO, 2013).
Cultural Control and Sanitary Measures
Sanitation measures include the removal and destruction of both infested fruit and any ripe, overripe and rotten fruit at the crop site that could serve as a host. Research is underway to evaluate solarizing, burying, bagging, crushing, and spraying infested fruit to discourage D. suzukii colonization (Walsh et al., 2011).
A fly-screen with 0.98-1.0 mm mesh prevents D. suzukii fruit damage on blueberry (Kawase and Uchino, 2005). Physical crop protection by using anti-insect nets are under experimental evaluation and seem to be a promising alternative control strategies for use in the near future (Ioriatti et al., 2012).
Early experiments tested the efficacy of Phaenopria spp. (Hymenoptera: Diapriidae) under laboratory conditions, but results were unsatisfactory (Kanzawa, 1939).
Studies to determine the current presence of indigenous parasitoid biological control agents and their efficacy in controlling D. suzukii were undertaken both in North America and in Europe by different research groups (Brown et al., 2011; Chabert et al., 2012; Rossi Stacconi et al., 2013). Under laboratory conditions several naturally occurring parasitoids of drosophilids in France were able to successfully parasitize D.suzukii. These included two larval parasitoids, Leptopilina heterotoma and Leptopilina boulardi, and two pupal parasitoids, Pachycrepoideus vindemiae (Hymenoptera: Pteromalidae) and Trichopria drosophilae (Hymenoptera: Diapriidae). Both Leptopilina parasitoids displayed high parasitism rates on D. suzukii, but because of the strong immune response of the host larvae, they did not give rise to an adult wasp (Chabert et al., 2012).
D. suzukii produces up to five times more hemocytes than D. melanogaster, making it significantly more resistant to wasp parasitism (Kacsoh and Schlenke, 2012) and making it less likely for indigenous specialized parasitoids to shift host onto it. While parasitization by L. heterotoma induced a decrease in the number of circulating haemocytes in D. melanogaster, it led to a large increase in the total haemocyte counts of D. suzukii (Poyet et al., 2013).
The observed difference between the immune response towards L. heterotoma in D. suzukii and D. melanogaster could suggest that European populations of L. heterotoma are not adapted to this new exotic host (Poyet et al., 2013); however, this hypothesis disagrees with the recent observations of a European-wide strain of L. heterotoma that is able to develop and emerge from D. suzukii. (Rossi Stacconi et al., 2013). It is probable that the European-wide strain of L. heterotoma has more effective venom, or that the strain of L. heterotoma used in the original study had lost its ability to develop on D. suzukii because of continued laboratory rearing on D. melanogaster.
Pupal parasitoids seem less susceptible to the high hemocyte levels of D. suzukii and they appear to have the highest potential for use in biocontrol of D. suzukii (Kacsoh and Schlenke, 2012). This was confirmed by the successful parasitism rate obtained with a pupal parasitoid by Chabert et al. (2012).
Predators of D. suzukii include several species of the bug genus Orius, a generalist predator, which were observed feeding on D. suzukii larvae in backyard raspberries in the autumn of 2009 (Walsh et al., 2011). Preliminary laboratory studies with O. insidiosus (Walsh et al., 2011), O. laevigatus and O. maiusculus (V. Malagnini, personal comm.) indicated that they can feed on D. suzukii larvae infesting blueberries, but their effective control of the pest population have not been proved yet.
The activity of microorganisms, as well as the intimate association of the pest species with endosymbionts, has not yet been exploited for biocontrol purpose.
Recently, DNA viruses have been isolated in Drosophila species (Unkless, 2011) and were found to be related to other viruses used for pest control.
Strains of endosymbiotic bacterium Wolbachia associated with D. suzukii populations have been collected in both the USA and Italy (Siozios et al., 2013; Tochen et al., 2014). These findings suggest the possibility of control of D. suzukii based on pathogens.
Current control efforts for D. suzukii rely heavily on the use of insecticides. The range of insecticides available for use on D. suzukii includes spinosyns, organophosphates, pyrethroids and neonicotinoids. However, the active ingredients are not very persistent. In addition, the fast generation turnover of D. suzukii requires many chemical interventions at the ripening stage, which can increase the risk of residues in fruits, promote insect resistance and negatively affect pollinators and other beneficial species. Moreover, the efficacy of the current available insecticides against D. suzukii larvae within fruits is limited, and D. suzukii control is focused on treatments based on chemicals targeting adults (Cini et al., 2012).
Significant adult D. suzukii mortality resulted from bioassays performed using formulated products of spinosyns, organophosphates and pyrethroids when directly applied on the insect (Bruck et al., 2011). In the same studies, neonicotinoids organic pyrethroid (pyrethrin) and azadiractin provided from moderate to low control, with significantly higher levels of male mortality (Bruck et al., 2011; Beers et al., 2011). High level of mortality was also obtained when D. suzukii adult were exposed to fresh residue of spinosyns, organophosphates and pyrethroids on fruits (Bruck et al., 2011). Malathion, bifenthrin and spinetoram also provided high mortality levels when D. suzukii adults were exposed to one-day field aged residue (Bruck et al., 2011; Beers et al., 2011). Tolfenpyrad had relatively good activity by topical exposure, but residual activity has yet to be determined. Mortality of flies exposed to cyazypyr was relatively low after 16 hours but caused intermediate mortality after 40 hours. The low level of mortality of the flies exposed to residues of imidacloprid, acetamiprid and cyazypyr on fruit seems to be compensated by a reduced adult emergence due to the systemic effect (Beers et al., 2011; Van Timmeren and Isaacs, 2013). Exposure to spinetoram, lambda-cyhalothrin and carbaryl reduced the number of eggs laid in cherries (Beers et al., 2011).
Timely field applications of lambda-cyhalothrin, deltamethrin, dimetoate and phosmet provided good control of the fruit damage with a residual activity lasting up to two weeks, whereas unsatisfactory efficacy was obtained with neonicotinoids (Grassi et al., 2011; Profaizer et al., 2012). Despite the high adult mortality measured in the semi-field bioassay, malathion did not provide satisfactory effective control of D. suzukii infestation in field trials (Profaizer et al., 2012). Van Timmeren and Isaacs (2013) reported that its effectiveness dropped quickly over time because of its sensitiveness to breakdown from exposure to ultraviolet light.
Organic crop production is seriously threatened by D. suzukii as only a few natural insecticides are allowed. The efficacy of these pesticides against D. suzukii is lower than organophosphates and pyrethroids. Field trails with pyrethrins and spinosad have a degree of efficacy and short pre-harvest interval, but residual impact is limited to a few days (Walsh et al., 2011; Grassi et al., 2011; Profaizer et al., 2012). Spinosyns, formulated as a bait, are also available for both conventional and organic fruit production, but are not highly effective for D. suzukii (Walsh et al., 2011). The addition of sugar-yeast bait to spinosyns significantly increased fly mortality (Knight et al., 2013). In blueberry production the few available insecticides for the control of D. suzukii have provided protection against infestation, but the need for repeated treatments with limited insecticide options increases the chance of resistance developing in the future (Van Timmeren and Isaacs, 2013).
Traps baited with apple cider vinegar (ACV) were initially used for crop risk assessment and treatment timing in IPM. Insecticide formulations are selected according to their efficacy, residual activity, pre-harvest interval, and the presence of other pests that could be controlled at the same time (Beers et al., 2011). Follow-up applications of pesticides should be applied when monitoring traps indicate the presence of D. suzukii (Bruck et al., 2011). ACV-baited traps are not always a reliable indicator of relative crop risk and it raises the possibility that traps baited with ACV are less attractive than natural ripe hosts. Improved estimation of the seasonal phenology of D. suzukii has been obtained by adding wine and sugar to ACV (Grassi and Maistri, 2013).
For an effective IPM strategy, chemical control has to be coupled with cultural management tactics such as sanitation (proper removal and disposal of unharvested or infested fruits) (Thistlewood et al., 2012). The amount and timing of rainfall may also negatively impact the longevity of insecticides due to wash off (Van Timmeren and Issacs, 2013). For short-residual insecticides, evening applications may be recommended. Due to the ability of D. suzukii to move up to several kilometers from infested fields, it is essential that management practices are carried out over a wide area (EPPO, 2013a). Scattered fruit trees, abandoned orchards, unmanaged host plants in private gardens or in nearby woodland should be considered potential sources of infestation and the associated risk of crop damage should be included in the management program.
As an alternative to chemical control, netting may be useful in keeping flies from attacking fruit on cane berries and cherry, provided they are installed before the fruit begins to ripen (Caprile et al., 2013). Netting with mesh size of 1 x 1 mm and 1 x 1.6 mm have been applied on blueberry and provided a good level of protection, but Grassi and Pallaoro (2012) suggested using smaller mesh sizes of 1 x 1 mm in order to maximize fruit protection. Netting must be secured at the ground and two layers of netting should be applied at the entrance of the tunnel (Grassi and Maistri, 2013).
Monitoring and Surveillance (incl. Remote Sensing)
The presence of adult D. suzukii in the field can be monitored by using traps baited with different attractants. Although field captures of D. suzukii in traps indicate their presence, trapping does not appear to accurately predict infestation in all crops (Lee et al., 2012; Wilson et al., 2013; Tochen et al., 2014). Any 250-750 ml plastic container with a closely fitting lid can be used as a trap. 0.5 – 1 mm diameter holes should be drilled in the side in order to enable the flies to enter.
A variety of trap prototypes made by researchers and commercial traps are available to monitor adult D. suzukii. Comparisons among different trap design (size, colour, volatilization area, entry area) have been performed across different regions and crops in North America (Lee et al., 2012; 2013). The number of captures increased as the entry area of traps increased, but small size of the holes slowed evaporation and increased the selectivity against the larger insects. Red, yellow and black traps were preferable over clear or white, but there was an interaction between the trap colour and the crop type. Trap colour had no effect on the selectivity towards other drosophlids (Lee et al., 2013). Laboratory bioassays found that flies were attracted to dark colours, ranging from red to black, and that the use of three alternating red, black and red coloured strips significantly increased the number of flies caught (Basoalto et al., 2013). Bait is needed to attract the flies to the trap.
Apple cider vinegar was one of the first baits used because it was effective and practical to use (EPPO, 2013a). This lure has been lately improved by adding wine (Landolt et al., 2012 ) and wine and sugar (Grassi and Maistri, 2013).
The fly response to the combination of vinegar and wine was greater than the response to acetic acid or the combination of acetic acid and ethanol, which are the principal volatile chemical components of vinegar and wine respectively (Landolt et al. 2012). This finding indicates that other volatile chemicals emitted by vinegar and wine, in addition to acetic acid and ethanol, may also be attractive to male and female D. suzukii. A sugar-yeast bait has been used successfully and was found to out-perform apple cider vinegar (Knight et al., 2013). A small drop of dish soap added to the liquid bait as a surfactant, or the placement of a sticky card within the trap, results in more fly captures.
More recently, multi-component volatile blends had been identified (Cha et al., 2012; 2013) that may provide a more selective lure and may reduce the time for trap servicing. Additionally, a synthetic chemical lure provided from a controlled release dispenser should remain an attractive bait for longer and would be more selective against non-target insects (Landolt et al. 2012; Cha et al., 2013).
Emergency mitigation measures include cold treatment or carbon dioxide/sulphur dioxide fumigation of host fruit when exporting from an infested country to an area free of D. suzukii (DAFF, 2013).
Gaps in Knowledge/Research NeedsTop of page
Some work has been conducted on the normal temperature ranges at which D. suzukii can develop and reproduce (Tochen et al., 2014). This information needs to be integrated into automated remotely-sensed weather data to create more accurate real-time and automated seasonal and regional risk maps. The effects of fluctuating temperatures have not yet been studied and need further investigation, as do the lower and upper temperature tolerance limits of D. suzukii in all of its invasive areas.
It will be important to explore the relationships between D. suzukii and D. subpulchrella using genome scale data.
Research into the possibility of controlling D. suzukii using viral pathogens is urgently needed.
Making lures more attractive to D. suzukii together with optimizing trap design are major objectives of different research teams aiming for an effective tool for mass trapping D. suzukii (Lee et al., 2012; 2013).
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30/11/13 Original text by:
Claudio Ioriatti, Center for Technology Transfer, Italy; Marco Stacconi, Fondazione Edmund Mach, Italy; Gianfranco Anfor, Fondazione Edmund Mach, Italy
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