CONFIRMED AND POTENTIAL WILD HOSTS OF THE SPOTTED WING
DROSOPHILA (DROSOPHILA SUZUKII) IN SLOVENIA
Maarten d e Gr o o t
1
, Andreja KAv č ič
1
& Jaka r Az in Ge r
2
1
Gozdarski inštitut Slovenije, v ečna pot 2, 1000 Ljubljana, Slovenia
2
Kmetijski inštitut Slovenije, Hacquetova ulica 17, 1000 Ljubljana, Slovenia
Abstract  t he spotted wing drosophila (Drosophila suzukii (Matsumura)) is a highly
invasive species and attacking different species of berry carrying hosts. Much research
has already been done on the crop hosts over the world and in Slovenia, but for wild
hosts less is known. o n basis of literature and fieldwork we prepared a list of potential
and actual known species of wild hosts for Slovenia. in 2019, berries of different
species were collected and D. suzukii was either reared from these berries or berries
were dissected. in total we found in the literature for e urope 99 species which were
used as host for D. suzukii. For Slovenia we found 71 potential hosts and 14 hosts
which were actually infested. in Slovenia there was a broad range of potential hosts
from 41 genera. t he genera with the most potential hosts were Prunus, Lonicera and
Vaccinium. Among the potential hosts were also many species which were invasive
alien or alien species. t he list was discussed in the context of management implications
and further research on D. suzukii in Slovenia.
Ke y w o r d S: Spotted wing drosophila, Drosophila suzukii, berries, potential hosts,
wild hosts, Slovenia, forests, pest control
Izvleček – Po t e n CiALn i in Po t r Je n o n APAd e n i d iv Ji Go St it e LJi
PLo d o v e v in SKe MUŠiCe (DROSOPHILA SUZUKII) v SLo v e n iJi
Plodova vinska mušica (Drosophila suzukii (Matsumura)) je invazivna tujerodna
žuželka, ki napada številne rastline s sočnimi plodovi, zlasti jagodičje. n arejenih je
bilo že veliko raziskav v povezavi z gostitelji D. suzukii, ki so gojeni kot ekonomsko
pomembne kmetijske rastline, malo pa je znanega o divje rastočih, t.j. negojenih go-
stiteljih te vrste. v naši raziskavi smo na podlagi pregleda literature in lastnih raziskav
izdelali seznam potencialnih in znanih divjih gostiteljev plodove vinske mušice v
Sloveniji. v sezoni 2019 smo nabrali plodove različnih vrst divjih gostiteljev, ki smo
jih v laboratoriju analizirali na prisotnost D. suzukii. Literatura navaja, da je v e vropi
121
ACTA ENTOMOLOGICA SLOVENICA
LJUBLJANA, DECEMBER 2020    Vol. 28, øt. 2: 121–130

101 vrsta rastlin primerna kot gostiteljska za D. suzukii. o d teh je v Sloveniji prisotnih
72 vrst, ki spadajo v 41 rodov. D. suzukii je bila pri nas potrjena na 16 vrstah. n ajpo-
gostejši gostitelji plodove vinske mušice v Sloveniji so iz rodov Prunus, Lonicera in
Vaccinium. Med potencialnimi divjimi gostitelji za D. suzukii v Sloveniji so tudi
rastline, ki so tujerodne ali invazivne tujerodne vrste. č lanek obravnava seznam
divjih gostiteljev plodove vinske mušice v Sloveniji v luči iskanja novih načinov za-
tiranja plodove vinske mušice in daje smernice za nadaljnje raziskave na tem po-
dročju.
KLJUč n e b e Se d e : Plodova vinska mušica, Drosophila suzukii, jagodičje, potencialni
gostitelji, divji gostitelji, Slovenija, gozdovi, zatiranje 
Introduction
t he Spotted wing drosophila (Sw d ) (Drosophila suzukii (Matsumura, 1931)),
originating from Asia, is highly invasive and damaging of economical crops in e U
and USA (Cini et al., 2012; Asplen et al., 2015) (Figure 1). in e urope, it was first
found in Spain in 2008 and rapidly spread throughout e urope (Asplen et al., 2015). it
was first observed in Slovenia in 2011 (Seljak, 2011). o ne of the reasons for its inva-
siveness is the fact that it has multiple generations and a large variety of host plants
(Asplen et al., 2015). o ne of the difference between e uropean d rosophilidae fruit fly
species and the Sw d is that the Sw d female is able to damage healthy, undamaged
fruits with its serrated ovipositor, while the females of other species can only feed on
rotting fruits (Sasaki in Sato, 1995). 
Sw d has a strong negative influence on the yield of the fruit crops. As it is highly
polyphagous most of the fruit crops have problems with this species (Cini et al.,
2012; Asplen et al., 2015). y ield losses ranging from 30-40% to 100% have been es-
timated, depending on the crop and the area. t he costs of the Sw d damage are esti-
mated on 500 million dollars per year for only the USA (b olda et al., 2010). in italy,
the costs were estimated on 500.000 e uro in 2010 to 3 million e uro in 2011 (d e r os
et al., 2012). in the recent years there has been an increase of management options
which increased the outcome but increased the management costs which was estimated
to 1857 CHF per hectare (Mazzi et al., 2017). t ill now, the number of known hosts
of Sw d in Slovenia were 23 species, most of which are actually crop hosts (Seljak et
al., 2015). All berry crops in Slovenia were strongly negatively affected with in some
case more than 50% of the crops (Seljak et al., 2015). d ue to the increasing damage
caused by the Sw d , new approaches and the development of new methods are needed
for the control of the populations of this species to avoid large economic damages
(Asplen et al., 2015). 
it was observed that the species has strong fluctuations over the years, where dry
years have low population densities, while wet years have high densities (Seljak et
al., 2015). interestingly, in the wet years also the dynamics change – the pest’s flight
starts earlier (Kenis et al., 2016).  
Acta entomologica slovenica, 28 (2), 2020
122

t he main focus of the research on Sw d management is on crops. However, many
wild fruits are taxonomically related to the crop fruits and therefore it is logical to
assume that many wild hosts are also infested. t wo studies in e urope by Poyet et al.
(2015) and Kenis et al. (2016), showed that there were respectively 33 and 84 wild
hosts found. in total this comes to 88 non crop hosts of the Sw d in e urope till now.
t he importance of the wild hosts is shown by the invasive species Prunus serotina
which was almost 70% infested and is assumed to be a strong factor of the Sw d dis-
persal (Poyet et al., 2014). Although Seljak et al. (2015) have performed a preliminary
study, in Slovenia the situation with crop/wild hosts has not yet been studied in detail.
Landscape is known to affect the population dynamics of the Sw d (Santoiemma
et al., 2019). it has been shown that there is a strong spill-over effect from non-crop
areas to the crops (Santoiemma et al., 2018; t onina et al., 2018). e specially with the
fact that Sw d is a strong disperser (up to 9000 m) also forest a bit further away can
have strong impacts on the Sw d dynamics in crops (t ait et al., 2018) it was found
that forests had higher densities of Sw d than meadows (Santoiemma et al., 2019).
Factors affecting the population densities in e urope are the forest cover (Haro-b archin
Maarten de Groot, Andreja Kavœiœ, Jaka Razinger: Confirmed and potential wild hosts of the spotted wing drosophila
123
Figure 1: r ecognition of the spotted wing drosophila. a) Males have a dark spot
on the tip of the wings. b) t arsomere i and ii of male forelegs bear a set of spines
each (sex combs). Females (c) have clear wings, and a strongly sclerotized ovipositor
with black teeth (d). (photos: Jaka r azinger)

et al., 2018) and the forest edges (Santoiemma et al., 2019). Forests are an important
habitat for source population, because of overwintering (many optimal microclimatic
conditions) (Santoiemma et al., 2018) and feeding on wild hosts which can provide
breeding material during the whole flying season of Sw d (Poyet et al., 2015; Kenis et
al., 2016). t he function of forests should therefore be an important focus for research
on the management of Sw d . Forests contain most of the wild hosts, however it was
shown that there are strong regional differences in the occupancy of hosts by Sw d
(Kenis et al., 2016).
t he aim of this study was to investigate the range of wild hosts on which Drosophila
suzukii can (potentially) reproduce in Slovenia. 
Materials and methods
t he preparation of the list of potential hosts of Drosophila suzukii in Slovenia con-
sisted out of three steps: First a review was made of hosts used in e urope. e specially
the list prepared in Kenis et al. (2016), Arnó  et al. (2016) and Poyet et al. (2015) were
used here and additional published literature which were not assessed by mentioned
sources. in the second step, the flora of Slovenia (Martinčič et al., 2010) was used to
determine whether the host actually occurs in Slovenia. in the third step certain hosts
were checked in the field whether they were colonized by Sw d also in Slovenia. For
this literature was checked and fruit from different (potential) host species was collected
in the field. b erries were collected in 14 sites over whole of Slovenia in July 2019.
t he sampling sites were mainly on forest edges. t he berries were put into a rearing
tent in the laboratory in the Slovenian Forestry institute, in order to let the adults of D.
suzukii emerge. w hen after a week the adult did not emerge, the berries were dissected
in order to see if the berries were infested with larvae. 
Results
Literature survey showed that in total 101 species were found to be wild hosts of
D. suzukii in e urope; 72 of these species occur also in Slovenia (t able 1). o f these
72 species, 14 species were found to be colonized by Sw d in Slovenia. t he total e u-
ropean species list contains 44 genera which include on average a bit more than 2
species. in Slovenia, the host plants were coming from 40 genera with on average of
approximately 2 species per genus. t he genera with the most host species were
Prunus, Lonicera and Vaccinium. 14 species were found to be either used as crop but
can also be found in nature.
Discussion
t he results show that there are many wild hosts available which can potentially
sustain Sw d population dynamics in Slovenia. t here were in total 71 potential wild
hosts found during the literature survey which also occur in Slovenia, and from the
field we detected 14 non crop species to be infested by Sw d .
Acta entomologica slovenica, 28 (2), 2020
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Maarten de Groot, Andreja Kavœiœ, Jaka Razinger: Confirmed and potential wild hosts of the spotted wing drosophila
125
Table 1: w ild host species found in e urope and in Slovenia. An asterisk (*)
marks species which are either native or alien and are used as crops, but can also be
found in nature in Slovenia.
Plant species Hosts in Europe
Hosts available in
Slovenia
Found to be infested
in Slovenia
Actinidia chinensis* 1
Amelanchier lamarckii 1
Amelanchier ovalis 1 1
Arbutus unedo 1 1
Arum italicum 1 1
Arum maculatum 1 1
Atropa bella-donna 1 1
Aucuba japonica 1
Bryonia cretica 1
Cornus alba 1
Cornus kousa 1
Cornus mas * 1 1
Cornus sanguinea 1 1
Cornus sericea 1 1
Cotoneaster franchetii 1
Cotoneaster horizontalis 1 1
Cotoneaster lacteus 1
Cotoneaster rehderi 1
Crataegus chrysocarpa 1
Crataegus monogyna 1 1
Daphne mezereum 1 1
Duchesnea indica 1 1
Eriobotrya japonica 1
Fragaria vesca 1 1
Frangula alnus 1 1
Gaultheria x wisleyensis 1
Hippophae rhamnoides 1 1
Ligustrum lucidum 1 1 1
Ligustrum vulgare 1 1 1
Lonicera alpigena 1 1
Lonicera caerulea* 1 1
Lonicera caprifolium 1 1
Lonicera ferdinandii 1
Lonicera nigra 1 1
Lonicera nitida 1
Lonicera periclymenum 1
Lonicera xylosteum 1 1
Mahonia aquifolium 1 1
Malus baccata 1
Morus alba 1 1
Morus nigra 1 1 1
Paris quadrifolia 1 1
Parthenocissus quinquefolia 1 1
Photinia beauverdiana 1
Photinia villosa 1
Photinia prunifolia 1

Acta entomologica slovenica, 28 (2), 2020
126
Physalis alkekengi 1 1
Phytolacca americana 1 1
Phytolacca esculenta 1
Polygonatum multiflorum 1 1
Prunus avium * 1 1 1
Prunus cerasifera* 1 1
Prunus cerasus * 1 1 1
Prunus domestica* 1 1 1
Prunus laurocerasus 1 1
Prunus lusitanica 1
Prunus mahaleb 1 1
Prunus padus 1 1
Prunus serotina 1 1
Prunus spinosa 1 1
Pyracantha sp. 1 1
Pyrus calleryana 1 1
Rhamnus cathartica 1 1
Rhamnus fallax 1 1 1
Ribes alpinum 1 1
Ribes rubrum * 1 1
Rosa acicularis 1
Rosa canina * 1 1
Rosa glauca 1 1
Rosa pimpinellifolia 1 1
Rosa rugosa 1
Rubus caesius 1 1 1
Rubus fruticosus agg. * 1 1 1
Rubus idaeus * 1 1
Rubus spp. 1 1 1
Rubus phoenicolasius * 1 1
Rubus saxatilis 1 1
Rubus ulmifolius 1 1
Sambucus ebulus 1 1
Sambucus nigra * 1 1 1
Sambucus racemosa 1 1
Solanum chenopodioides 1
Solanum dulcamara 1 1
Solanum nigrum 1 1
Sorbus aria 1 1
Sorbus aucuparia * 1 1
Symphoricarpos albus 1 1
Tamus communis 1 1
Taxus baccata 1 1
Vaccinium myrtilloides 1
Vaccinium myrtillus 1 1 1
Vaccinium oldhamii 1
Vaccinium praestans 1
Vaccinium vitis-idea 1 1
Viburnum lantana 1 1
Viburnum opulus 1 1 1
Viburnum rhytidophyllum 1
Viscum album 1 1
Vitis vinifera * 1 1 1
Total 99 71 14

w e found a large number of potential host species for Slovenia over a large range
of genera. t his is in principle not new as it is already shown in previous studies that
it is a polyphagous species (Asplen et al. 2015, Kenis et al. 2016). However, such a
study was not yet done for Slovenia. t he fact that there was a large number of
potential hosts is probably also one of the reasons that the species could invade
Slovenia so fast and can be found in large abundances throughout all of Slovenia
(Seljak et al., 2015).
Many of the host species listed in t able 1 are autochthonous species in e urope,
but some are invasive alien species (Kenis et al., 2016). it was shown that invasive
alien species can be an important food source for the Sw d (Kenis et al., 2016). P.
serotina was shown to have 70% of damage (Poyet et al., 2014), while Phytolacca
americana had the highest number of eggs on the fruits in a survey of 33 host species
(Poyet et al., 2015). w ith the increasing disturbance by wind and bark beetles in the
forests of Slovenia (de Groot & o gris, 2019; z GS, 2019), many more forest gaps
will develop. t hese gaps are suitable places for invasive alien species to grow when
spread by wind or birds and support the Sw d populations. o n the other hand, orna-
mental plants which become invasive are introduced into Slovenia via the trade (d e
Groot et al., 2017). t hese species, which might escape into the forests and other
habitats can also become host plants for the Sw d and therefore support the populations
of Sw d . 
t he e uropean list of wild hosts mentioned in t able 1 were mainly sampled in
France, italy, Switzerland and the n etherlands (Poyet et al., 2015; Kenis et al., 2016).
A majority of the host plant species which are found in these countries have related
species in the countries of the Southern e urope. t his would mean that the shown
number of potential hosts can be still increased by species which are not yet surveyed.
it is therefore expected that the total number of 72 wild host species for Slovenia and
101 species for e urope will still increase.
o ne of the advantages to be a polyphagous species is that the host species are spread
in time. Kenis et al. (2016) and Poyet et al. (2015) showed already that with the com-
bination of plant species the berries are available  throughout the year. Most species are
fruiting during spring and summer. t he winter is a period with not so many berries
available. Plant species like Duchesnea indica, Prunus laurocerasus, Rosa canina,
Lonicera nitida, Viscum album and Aucuba japonica contain or can contain berries
also during the winter which could sustain the population till the next year (Poyet et al.,
2015; Kenis et al., 2016). t he number of wild hosts of the Sw d is expected to grow in
the future. Given the trends, this could have detrimental effects on crops and economy
due to the expected positive influence on the Sw d population levels. 
Implications for management
Slovenia is known for its large forest cover (almost 60%). Most of the forest con-
tains host species which are presented in this study. Knowing that there is a lot of
host availability in the forest, it becomes clear that the population of Sw d could be
sustained also from the forests. it remains therefore a question what we could do to
Maarten de Groot, Andreja Kavœiœ, Jaka Razinger: Confirmed and potential wild hosts of the spotted wing drosophila
127

minimize the spillover effect from the forest to the orchards. Kenis et al. (2016) pro-
posed to control the amount of wild host in the vicinity of the fruit orchards, but in
light of recent research the Sw d flies can migrate over long distances (t ait et al.,
2018); therefore this action will not have much effect. instead, it is important to be
aware, if there are many wild hosts in the vicinity of orchards and adapt management
strategies accordingly. For instance, one could try to use early ripening fruit as crops,
as the highest abundance of Sw d is reached in the late summer (Seljak et al., 2015;
t onina et al., 2018). Another option is to start growing less susceptible fruit (w ang
et al., 2019). A third option would be that in areas with a high amount of wild hosts,
monitoring would be intensified and used to time the application of insecticides on
the crops to prevent damage by the Sw d . A fourth option is to use appropriate nets
(1 mm mesh or finer) covering entire orchards. t his is mostly applicable for newly
established orchards, whereas older orchards could be partially protected by the use
of lateral netting (Cini et al., 2012; Leach et al., 2016; w eber et al., 2016).
o ur study shows a list of potential host species which can be or already are attacked
by the Sw d in Slovenian forests and other habitats of wild hosts. w e show that many
wild hosts are available in Slovenia for the Sw d outside crop areas. However, there
are concerns that invasive alien plants’ abundance and distribution will increase in the
coming years and therefore facilitate the population of Sw d . o n the other hand,
climate change can increase disturbance in the forest and increase the amount of hosts
in gaps like Rubus. Furthermore, it can also decrease the Sw d development time and
therefore increase the number of Sw d generations. in this study the host species of
Sw d are pin pointed, however to understand the distribution of the host plant would
give a better insight on the distribution, abundance and risk of Sw d in Slovenia. inte-
grating wild hosts of the Sw d in regard of risk maps and planning of orchards, and
the use of very fine protective netting in the development of management strategies
for this pest are becoming important aspects in the control of this pest in the future.  
Acknowledgments
w e would like to thank z avod za Gozdove Slovenije (z GS) for the help for col-
lecting berries in the field and dr. d arinka Koron, dr. n ika w eber and dr. Lado
Kutnar for their constructive comments. t he research was performed within the
project “o bvladovanje plodove vinske mušice (Drosophila suzukii) z metodami z
nizkim tveganjem” financed by the Slovenian r esearch Agency (Ar r S) (grant v 4-
1802; Agrobiodiversity program group, grant P4-0072) and the Administration of
the r epublic of Slovenia for Food Safety, v eterinary Sector and Plant Protection
(Uv Hv v r ), Ministry of Agriculture, Forestry and Food (MKGP).
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