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Biological Control 63 (2012) 40–47

Contents lists available at SciVerse ScienceDirect

Biological Control

Ability of European parasitoids (Hymenoptera) to control a new invasive Asiatic pest, Drosophila suzukii a a a,b b a,⇑ Stan Chabert , Roland Allemand , Mathilde Poyet , Patrice Eslin , Patricia Gibert   a Université de Lyon, F-69000, Lyon; Université Lyon 1; CNRS, UMR 5558, Laboratoire de Biométrie et Biologie Evolutive, F-69622, Villeurbanne, France b Laboratoire de Bio-écologie des Insectes phytophages et Entomophages, EA 4698 EDYSAN, Université de Picardie Jules Verne, 33, rue Saint Leu, F-80089 Amiens Cedex, France

h i g h l i g h t s g r a p h i c a l a b s t r a c t

" We test the effectiveness of five major European Parasitoids against Drosophila suzukii. . " We found that only the two pupal parasitoids were able to develop on D. suzukii. " The three larval parasitoids exhibited no parasitism success. " D. suzukii exhibits high immune resistance capacity against Leptopilina species.

a r t i c l e i n f o a b s t r a c t

Article history: Understanding the ecological factors involved in successful invasions is essential for choosing appropriate Received 5 March 2012 management measures. One mechanism recognized as often being essential for invasion success is for the Accepted 22 May 2012 invasive species to be less subject to attack by natural enemies. The spotted-wing drosophila, Drosophila Available online 30 May 2012 suzukii (Matsumura, 1931) is an Asian pest of fruit crops that has recently appeared simultaneously in North America and Europe (2008). Here we investigate the effectiveness of European parasitoids of Dro- Keywords: sophila in parasitizing D. suzukii. Of the five main European parasitoid species, only two pupal parasitoids Enemy release hypothesis with wide host ranges develop on D. suzukii. Two specialized larval parasitoids were unable to develop, Drosophila suzukii presumably because of a strong immune response. The third specialized larval parasitoid rarely ovipos- Host-parasitoid interactions Leptopilina heterotoma ited in D. suzukii. This confirms that host switching is often difficult for specialist parasitoids. Asobara tabida 2012 Elsevier Inc. All rights reserved.

1. Introduction that have accelerated the movement of species, breaking down biogeographic barriers that had isolated populations for thousands Recent years have seen drastic changes in the composition of of years (Williamson and Fitter, 1996; Mooney and Cleland, 2001). biotic communities largely as a result of human activities (develop- These changes can play an important role in the erosion of biodi- ment of international trade and intercontinental transportation) versity and the disruption of the invaded ecosystems (Lodge, 1993). Some newly-introduced species become invasive, and have

a considerable economic impact (Pimentel et al., 2000). A biologi- ⇑Corresponding author at: CNRS, UMR 5558, Laboratoire de Biométrie et Biologie Evolutive, UCB Lyon 1, 43 bd du 11 novembre 1918, F-69622 Villeurbanne, France. cal invasion can be defined as the successful establishment, devel- Fax: +33 4 72431388. opment and maintenance of a species outside its native geographic E-mail addresses: [email protected] (S. Chabert), Roland.Allemand@ range (Facon et al., 2006). Three phases are typically distinguished univ-lyon1.fr (R. Allemand), [email protected] (M. Poyet), patrice. between the introduction of species and the stage of biological [email protected] (P. Eslin), [email protected] (P. Gibert). invasion (Sakai et al., 2001; Williamson, 2006): (1) the

1049-9644/$ - see front matter 2012 Elsevier Inc. All rights reserved. http://dx.doi.org/10.1016/j.biocontrol.2012.05.005

S. Chabert et al. / Biological Control 63 (2012) 40–47 41 introduction of a number of founding individuals, (2) establish- (Hymenoptera Diapriidae) (Allemand et al., 1999; Fleury et al., ment (sometimes after a latency period), and finally (3) population 2009). Most studies agree that Drosophila parasitoids induce a high growth. rate of mortality in their host populations despite the fact that the During the various phases of the invasion, individuals encoun- level of parasitism varies depending on the breeding site, the local ter new environmental conditions, and are generally subjected to situation, and the season. The natural average rate of parasitism strong selective pressures. Among the various key factors for a suc- can reach about 90% at some sites in Southern France (Fleury cessful biological invasion, the ‘‘Enemy Release Hypothesis’’ (ERH) et al., 2004). These high parasitism levels indicate that parasitoids is commonly recognized as an essential mechanism. This hypothe- may be one of the main factors of mortality in fly populations, and sis suggests that the success of an exotic species outside its native thus constitute selective force acting on their hosts. range is due to the absent or reduced effectiveness of its natural The aim of this paper is to test the ability of European frugivo- enemies (pathogens, predators, parasitoids) (Keane and Crawley, rous Drosophila parasitoids to develop in D. suzukii under labora- 2002; Blumenthal, 2006; Facon et al., 2006; Liu and Stiling, tory conditions. The results should allow to estimate the 2006). Within the community, each species has to face enemies potential role that these parasitoids may play to control this new that can be specialists or generalists with regard to other compet- invasive Asiatic Drosophila pest. ing species. It is assumed that natural enemies in the area where the species is introduced need time to adapt before they can suc- 2. Materials and methods cessfully reduce the ecological threat of a successful invader. Para- sitoids are of particular interest for studying the mechanisms 2.1. Populations used involved in eco-evolutionary processes in , because they de- velop in close association with their hosts, and are known to coe- Two D. suzukii females were wild-caught in October 2010 using volve with them (Quicke, 1997). banana traps in Sainte-Foy-lès-Lyon (lat 45.74 N, France) and used Parasitoids constitute a large group of insects (consisting to establish a mass strain. This strain was mass reared at 21 C, fed mainly of Hymenoptera and Diptera), which develop inside or on an artificial diet (David and Clavel, 1965), and maintained with a the surface of other (usually insects), and consume 12:12 LD photoperiod. the host tissues during their development. In communities, Five European Drosophila parasitoid species were used in this they play a critical role in controlling phytophagous populations, experiment: three larval parasitoids, L. heterotoma and L. boulardi, such as pest populations in agricultural and forest environments either as a natural control (Hawkins et al., 1997), or when they A. tabida, and two pupal parasitoids P. vindemmiae and Trichopria cf drosophilae. Parasitoids were trapped in October 2010 in France are used in biological control strategies (Greathead, 1986). Recently, the spotted-wing drosophila, Drosophila suzukii (Mat- (Rhône valley) using banana traps, and mass strains were kept in the laboratory at 21 C on D. melanogaster with a 12:12 LD photo- sumura, 1931), a species belonging to the melanogaster group orig- period. To avoid any possible sampling effect, two populations of inally reported in Japan, has been observed elsewhere: in 2008 in each parasitoid species were tested (see Table 1). Each population North America and in Europe (Spain) (Calabria et al., 2010; Hauser, was founded from 20 to 30 inseminated females. 2011). In 2010, its presence was confirmed in southern France, and The additional experiment on the infestation behavior of A. tab- it has subsequently spread rapidly throughout the country. D. suzu- ida was done using the A1 strain collected in 1994 in Sainte Foy- kii is a pest of fruit crops with a very wide host range, and lives les-Lyon and reared on D. melanogaster larvae at 20 C and LD mainly on red fruits (raspberry, strawberry, cherry) (Lee et al., 13:11. 2011). Although in the vast majority of fruit flies, larvae develop As a control, we used a Japanese thelytokous strain of A. japon- only in damaged or rotting fruit, D. suzukii is the one of rare Dro- ica reared under the conditions described above. This strain was sophila species which lays its eggs in sound fruit using its serrated graciously provided by Professor J. van Alphen of Leiden University ovipositor (Mitsui et al., 2006). Damage is then caused by larvae (The Netherlands). feeding on the pulp inside the fruits and berries. Subsequently, sec- ondary fungal or bacterial infections may contribute to further fruit deterioration. Significant losses have been reported, particularly in 2.2. Parasitoid effectiveness the United States (California, Oregon, Washington), and following the first damage observed in Italy, D. suzukii was added to the The temperature used for development was 25 C, except for A. watch list of the Organization for European Plant Protection Orga- tabida, which was kept at 21 C because of its thermal preference. nization (EPPO) in January 2010. The control, A. japonica, was allowed to develop at both 21 and Despite the scale of the damage that can be produced by this 25 C. species, there is a surprising lack of information available about D. suzukii females, between 5 and 18 days old, were allowed to its biology and its natural enemies, and in particular about parasit- oviposition at 21 C (LD 12:12) on a banana medium (mix and oids that could limit its expansion. To date, only one pupal ectopar- heat: [20 g agar, 500 mL H2O], 400 g mashed banana, 50 g brewers’ asioid, Pachycrepoideus vindemmiae (=dubius, Hymenoptera: yeast, 30 g flour, 20 g sugar, [4 g nipagine, 30 mL alcohol 70 ],

Pteromalidae) (Brown et al., 2011) has been successfully reared 550 mL H2O). Eggs were then counted and transferred in a vial con- on D. suzukii. In a field-sampling study in Japan, three larval endo- taining standard artificial medium (David and Clavel, 1965). parasitoids were reported to develop on D. suzukii, Ganaspis xan- Because of the low fertility of D. suzukii in our laboratory condi- thopoda (Hymenoptera: Figitidae) and two Asobara species, tions, for each parasitoid population, we set up two blocks of five Asobara tabida (Hymenoptera: Braconidae) and Asobara japonica vials each containing 60 host eggs (LD 12:12). After 24 h (to allow at an extremely low rate (Mitsui et al., 2007). the eggs to hatch) or 5 days (to obtain pupae) for the larval and pu- In France, the communities of frugivorous Drosophila and their pal parasitoids respectively, we added one mated female wasp parasitoids involve several Drosophila species (mainly Drosophila (aged 4–7 days), fed with honey, with no previous experience of melanogaster, Drosophila simulans, Drosophila immigrans, Drosophila parasitism to each vial for 5 days. The vials were maintained until subobscura), and five species of parasitoids, including three larval the flies and wasps had emerged. koinobiont and solitary endoparasitoids A. tabida, Leptopilina het- Parasitoid development was estimated on two groups of 10 erotoma and Leptopilina boulardi (Hymenoptera Eucoilidae), and vials (two blocks of five vials for each population) each containing two pupal ones P. vindemmiae and Trichopria cf drosophilae 60 Drosophila eggs and one parasitoid female, as described above. S. Chabert et al. / Biological Control 63 (2012) 40–47 42

Table 1 Geographical description of the different populations of parasitoids tested on D. suzukii, with indices (% ± SE) summarizing host-parasitoid interactions per population. DI: Degree of infestation, this measures the proportion of hosts that were successfully parasitized, SP: Success rate of parasitism, this measures the probability that an infested host will give rise to an adult wasp, TER: encapsulation rate after complete development this estimates the immune capacity of D. suzukii which allows it to survive to adulthood. NA: data non available.

Species Temperature/Population Lat. N Long. E Distance between the two No of replicates DI SP TER populations tested (km) A. japonica 21 C 10 98 ± 0.8 41 ± 8 NA 25 C 5 91 ± 3.1 71 ± 10.5 NA A. tabida Igé (IG) 46.39 4.74 120 10 0 0 – Sablons (SA) 45.32 4.77 10 0 0 – P. vindemmiae Serrières (SE) 45.32 4.76 70 9 68 ± 8.5 60 ± 11.2 – Maison Neuve (MN) 44.43 4.30 5 50 ± 9.4 53 ± 16.8 – Trichopria cf drosophilae Ste Foy (SF) 45.74 4.79 46 10 85 ± 5.1 38 ± 11.8 – Sablons (SA) 45.32 4.77 10 69 ± 8.7 76 ± 14.2 – L. boulardi Sablons (SA) 45.32 4.77 180 10 70 ± 4.9 0 45 ± 6.9 Eyguières (EY) 43.69 5.03 10 63 ± 6.9 0 59 ± 10.8 L. heterotoma St Etienne/Chalaronne (ST) 46.15 4.88 340 10 52 ± 7.7 0 68 ± 14.9 Antibes (AN) 43.58 7.12 10 83 ± 5.7 0 80 ± 8.9

From this known number of eggs, some of the Drosophila would overestimated and the success rate of parasitism and the encapsu- die as a result of ‘natural’ mortality and an unknown proportion lation rate after complete development underestimated. would be parasitized. There were three possible outcomes of this parasitism: the death of the drosophila, that of the parasitoid if 2.3. Behavioral experiment on A. tabida the drosophila were able to resist to this parasitism, or the death of both species as a result of physiological inadequacy between In the first experiment, a null Degree of infestation being ob- both partners. served for A. tabida, we performed a separate experiment on A. tab- To assess the quality of the development of D. suzukii in the ab- ida alone to explore this further. Each A. tabida female (7 days old) sence of parasitoids (control vials), we estimated the developmen- used for this experiment was stimulated for 5 min with larvae of D. tal success from egg to adult at both 25 C and at 21 C. The values subobscura. D. subobscura is one of main natural hosts of A. tabida obtained at 25 C were not significantly different whatever the age (van Alphen and Janssen, 1981). This protocol allowed us to select of the Drosophila parents (between 4 and 52 days old) females that exhibited host-seeking behavior. The five female par- (F = 2.69, p = 0.105). We therefore pooled the values to in- (1,83) asitoids selected were placed for one hour in a vial containing 20 crease the number of individuals (85 vials corresponding to 5100 Drosophila larvae (stage L2), either D. melanogaster or D. suzukii. eggs), and obtained a value of 48.2% ± 1.3 (n = 85) for T. The value Eight replicates were used. The behavior of the females (probing, of T used for the experiment with A. japonica at 21 C, 62.7% ± 3.3 oviposition) was recorded, and the number of larvae parasitized (n = 10), was obtained in another experiment with young drosoph- was determined after dissection. ila mothers. We counted the total numbers of adults of Drosophila (di) and of parasitoids (pi) emerging from the vials for each female parasitoid. 2.4. Statistical analysis We scored two indices that summarize the host-parasitoid interac- tions (see for example Boulétreau and Fouillet, 1982; Boulétreau To compare the values of Degree of infestation, Success rate of and Wajnberg, 1986; Gibert et al., 2010). parasitism and Encapsulation rate after complete development in First, the ‘Degree of infestation’ (DI) measures the proportion of the different populations (different temperatures for A. japonica) hosts that were successfully parasitized, and is estimated as and blocks for each species separately, we used a hierarchical anal- (T di)/T; T being the average number of emerging flies in the ab- ysis of variance (blocks in populations) on arcsine (square root) sence of the parasitoid. transformed proportions with linear models on the statistical soft- Second, the ‘Success rate of parasitism’ (SP) measures the prob- ware R (version 2.10.1), after checking the normality of residuals ability that an infested host will give rise to an adult wasp, and this and homoscedasticity with Shapiro’s and Bartlett’s test respec- is estimated as pi/(T di). In some cases, pi > (T di); for these we tively. The transformed data enable to homogenize variances of set SP = 1. proportions. The biological significance of these two parameters is quite clear since the Degree of Infestation represents the probability of 3. Results a given host’s being parasitized, and the Success rate of parasitism the probability that a parasitized host would give rise to a wasp. Estimations of the Degree of infestation, Success rate of parasit- Moreover, when parasitized by parasitoids, many larvae of Dro- ism and encapsulation rate after complete development (TER) for sophila species are able to defend themselves by surrounding for- each species are summarized in Table 1. eign bodies with blood cells that will melanize and form a black capsule that kills the parasitoid by asphyxiation. In order to esti- mate the immune capacity of D. suzukii against parasitoids, we 3.1. A. japonica counted the number of adult flies with a capsule (dc). This enabled us to estimate the ‘encapsulation rate after complete development’ Our results for A. japonica (Fig. 1A and B) confirmed that this (TER) as the ratio dc/(T di + dc) (see Eslin and Prévost 2000 for species was very effective at both temperatures, and of the six instance). wasps tested it was the one that exhibited the highest level of De- Because it is impossible to exclude the possibility that a gree of infestation (DI > 90%). It was very successful (SP about 40– Drosophila larva could be killed by the parasitoid’s oviposition 70%), and performed slightly but significantly better at 25 C than without receiving a wasp egg, the degree of Infestation might be at 21 C (F(1,12) = 7.04, p = 0.021 ; see Fig. 1). S. Chabert et al. / Biological Control 63 (2012) 40–47 43

3.2. Local pupal parasitoids oviposit inside the larvae: oviposition was observed only three times (out of a total of 160 larvae), so that only 1.25% of D. suzukii As we have already said, two populations were tested for each larvae were parasitized by A. tabida. Under the same conditions, in species, and experiments were done on two successive blocks (five a control with D. melanogaster as host 225 ovipositions were ob- vials per block). Both pupal parasitoids effectively parasitized D. served, resulting in 81.25% of larvae infested by A. tabida, most of suzukii, but the outcomes were very variable. which were superinfested. For P. vindemmiae, a significant block effect was found for the Degree of infestation for both populations (t = 2.25, p = 0.048 for MN ; t = 3.18, p = 0.01 for SE ; see Fig. 2A and C). The values of Suc- 3.4. Leptopilina species cess rate of parasitism averaged 57%, and there was no significant difference between blocks or populations. For the two Leptopilina species, we observed a high level of De- For Trichopria cf drosophilae, the Degree of infestation was sig- gree of infestation that averaged 67% in L. boulardi, and reached nificantly different between the different blocks (F < 29.5, (1,17) 95% in one population of L. heterotoma (AN). We found a significant p < 0.001) and populations (F < 5.92, p = 0.027), with higher (1,16) block effect (t = 2.58, p = 0.02) for this population. No significant values obtained for SF than for SA (84.8% vs 69.1%). Success rate difference was found between the populations for L. boulardi. For of parasitism was found to be significantly different between the L. heterotoma, the two populations were significantly different, two populations (F < 5.74, p = 0.029; see Fig. 2b), with SF hav- (1,16) with higher values obtained in AN than in ST. ing a lower Success rate of parasitism than SA (38.2% vs 76.3%) (see Nonetheless, the Success rate of parasitism was null for both Fig. 2B and D). populations of both species, showing that these species could not develop on D. suzukii (Fig. 3). This unexpected result could be ex- 3.3. A. tabida plained by the high immune response produced by D. suzukii against these two species, as shown by the high values of the A. tabida seemed to be unable to parasitize D. suzukii (DI = 0). In encapsulation rate after complete development, which averaged the laboratory, we noticed that on D. melanogaster the A. tabida fe- 74% for L. heterotoma and 52% for L. boulardi. For this parameter, males displayed more oviposition behavior when they were not we observed a significant block effect for L. boulardi in the EY pop- isolated. We therefore performed another experiment on D. suzukii ulation (t = 2.54, p = 0.02). No significant differences were found following the same procedure, but this time with four females between populations for the two species. placed together in the vial. This confirmed the null Degree of infes- We performed another experiment following the same proce- tation observed in the previous experiment in three replicates. dure, but placing together four Leptopilina females in the vial (three In Japan A. tabida has been found emerging from D. suzukii pu- replicates). For L. boulardi, we confirmed the null Success rate of pae (Mitsui et al., 2007), so we performed a separate behavioral parasitism, but for L. heterotoma (AN) three adults emerged (from experiment to confirm our results. We observed that when in close 180 eggs), indicating that although it is difficult for this species contact with D. suzukii, A. tabida females almost never tried to to develop on D. suzukii, it is physiologically possible.

Fig. 1. Results of the parasitism of D. suzukii by A. japonica at two developmental temperatures. (A) Percentage of emerging Drosophila that escaped parasitism (di) and the percentage of parasitoids emerging as adults (pi). (B) Mean ± se of the Degree of infestation (DI) and Success rate of parasitism (SP). At 21 C two blocks of five replicates were used while only one block of five replicates was used at 25 C. For each replicate, 60 eggs were exposed to one parasitoid female. Different letters indicate a significant difference between blocks (p < 0.05); and NS indicate significant (p < 0.05) and non-significant differences between the two temperatures, respectively. S. Chabert et al. / Biological Control 63 (2012) 40–47 44

Fig. 2. Results of the parasitism of D. suzukii by the two pupal parasitoids: (A and C) parasitism by P. vindemmiae – (B and D) parasitism by Trichopria cf drosophilae. For each species, two populations were tested at 25 C with two blocks per population, and five replicates of 60 eggs and one parasitoid female per block. (A and B) shows the proportion of emerging Drosophila that escaped being parasitized (di) and the percentage of parasitoids emerging as adults (pi). (C and D) shows the mean ± se of Degree of infestation (DI) and Success rate of parasitism (SP). Different letters indicate a significant difference between blocks (p < 0.05); and NS denote significant (p < 0.05) and non- significant differences between the two populations, respectively.

4. Discussion whereas the larval parasitoids were unable to parasitize this new host, showing that host switching is difficult for the more special- The aim of this paper was to investigate the effectiveness of ized local parasitoids. French local Drosophila parasitoids against the pest D. suzukii under The successful parasitism obtained with pupal parasitoids is standard laboratory conditions. We showed that the three larval not surprising, especially in the case of P. vindemmiae, which is parasitoid species tested are not able to parasitize successfully D. the most generalist of the five parasitoids used in this experi- suzukii. Only two parasitoid pupae were able to grow in this new ment, and has been reported to attack over 60 fly species, includ- host. ing many tephritid fruit flies and several Drosophila species In nature, female parasitoids respond to a hierarchy of physical (Wang and Messing, 2004). This species is widely distributed, and/or chemical stimuli that lead them to their potential host and has been found in America, Africa and Europe (Carton (Doutt,1959; Godfray, 1994), and so this study constitute a first ap- et al., 1986). The successful parasitism of the two French popula- proach in laboratory of the importance of the Enemy Release tions of Pachycrepoideus found on D. suzukii (57%) in this study is Hypothesis which predicts that (i) specialized enemies of the intro- of the same magnitude as had already been reported on D. mela- duced species are absent in the invaded region, (ii) host switching nogaster (Delpuech et al., 1994). of specialized enemies from the native species to the introduced Trichopria cf drosophilae is a more specialist species and is able species is rare and (iii) finally, generalist enemies are more effec- to develop on many frugivorous Drosophila (Carton et al., 1986). tive against native species than invasive ones (Keane and Crawley, For this species, we found significant differences between popula- 2002). tions with regard both to the degree of infestation and the success Our results match under laboratory conditions the predictions of parasitism. The two populations used were collected at the same of the Enemy Release Hypothesis as only pupal parasitoids, which time and at a distance of only 50 km, so these differences are diffi- are known to be generalists, were effective against D. suzukii, cult to explain, but suggest possible genetic variability. S. Chabert et al. / Biological Control 63 (2012) 40–47 45

Fig. 3. Results of the parasitism of D. suzukii by the two Leptopilina species. (A and C) Parasitism by L. boulardi – (B and D) Parasitism by L. heteroroma. For each species, two populations were tested at 25 C with two blocks per population, and five replicates of 60 eggs and one parasitoid female per block. (A and B) shows the proportion of emerging Drosophila that escaped being parasitized (di–dc) and of infested flies that exhibited an immune response (dc). (C and D) shows the mean ± se of Degree of infestation (DI) and encapsulation rate after complete development (TER). Each letter indicates significant differences between blocks (p < 0.05); and NS denote significant (p < 0.01) and non-significant differences between the two populations, respectively.

In France, these two pupal parasitoids correspond to less than busckii, Drosophila funebris, Drosophila hydei, Drosophila kuntzei, D. 20% of the parasitoid community (Fleury et al., 2009). However, melanogaster, Drosophila obscura, Drosophila phalerata, D. simulans, this could have been underestimated as a result of the experimen- D. subobscura and Drosophila willistoni) and related genera (Chy- tal procedures used, which depends on the time of exposure in the momyza or Scaptomyza) (Jenni, 1951; Carton et al., 1986; Janssen, field and very little is known about their capacity to control natural 1989; Gibert et al., 2010). L. heterotoma exhibits a wide holarctic populations of Drosophila. distribution (Nordlander, 1980; Carton et al., 1986; Janssen et al., With regard to the larval parasitoids, our results demonstrate 1988; Hardy and Godfray, 1990; van Alphen et al., 1991; Mitsui that the French populations of the two Leptopilina species were et al., 2007; Fleury et al., 2009; Gibert et al., 2010), and is thus sym- not able to develop in D. suzukii (no Leptopilina adult parasitoids ob- patric with D. suzukii in Japan. For these two larval parasitoids, we tained from 1200 D. suzukii eggs for both species). This result is not observed a significant rate of effective encapsulation by D. suzukii, very surprising for L. boulardi, which is able to develop on only a few reaching 52% for L. boulardi and 74% for L. heterotoma. frugivorous Drosophila species, including D. melanogaster and D. A. tabida is able to develop in a few Drosophila species, includ- simulans (Barbotin et al., 1979; Carton et al., 1987; Carton and ing D. subobscura, D. obscura and D. melanogaster, but can also de- Nappi, 1991; Fleury et al., 2004), and also in the tropical species velop on D. tristis (Janssen, 1989; Kraaijeveld and Van Alphen, Drosophila yakuba (Dubuffet et al., 2008). This parasitoid species 1995). Its geographic range includes the northwest of America has been recorded in Mediterranean and intertropical climates (Hoang, 2002), Japan (Mitsui et al., 2007) and Europe (Carton (Barbotin et al., 1979; Chabora et al., 1979; Hertlein, 1986; et al., 1986). For this species, we found a null degree of infestation. Nordlander, 1980; Carton et al., 1991; Allemand et al., 2002). In This was rather surprising because this species has been reported contrast, L. heterotoma is clearly the most generalist parasitoid of amongst emerging parasitoids from field sampling of D. suzukii in the three larval parasitoids investigated. Its successful development Japan, but at a very low rate: in only one individual out of over has been recorded on numerous Drosophila species (Drosophila 1152 D. suzukii pupae collected in the field (Mitsui et al., 2007). 

Additional experiments that we performed on the behavior of A. tics used in this paper and Theotim Colin who took the photo of the tabida females showed that this species almost never tried to ovi- graphical abstract. We also thank Professor J. van Alphen who gra- posit on D. suzukii. Overall, only three of the 160 D. suzukii larvae ciously provides us the A. japonica strain. were infested by A. tabida. One possible explanation of the differ- ence between our results and those of Mitsui et al. is that local References European populations of A. tabida have still not adapted to this exotic new host. One way to test this hypothesis, which would fit Allemand, R., Fleury, F., Lemaitre, C., Boulétreau, M., 1999. Dynamique des in with the predictions of the enemy release hypothesis, would populations et interactions compétitives chez deux espèces de Leptopilina, be to compare the parasitism efficiency of Asian and European parasitoïdes de drosophiles, dans la vallée du Rhône (Hymenoptera: Figitidae). populations of A. tabida under the same conditions. Annales de la Société Entomologique de France 35, 97–103. Allemand, R., Lemaitre, C., Frey, F., Boulétreau, M., Vavre, F., Nordlander, G., van In our experiment, we used two French populations for each Alphen, J., Carton, Y., 2002. Phylogeny of six African Leptopilina species parasitoid species in order to avoid a possible strain or sampling (Hymenoptera :Cynipoidea, Figitidae), parasitoids of Drosophila, with description of three new species. Annales de la Société Entomologique de effect that could have interfered with our conclusions. In most France 38, 319–332. cases we found concordant results for the two populations. A Barbotin, F., Carton, Y., Kelner-Pillault, S., 1979. Morphologie et biologie de Cothonaspis boulardi n. sp. parasite de drosophiles. Bulletin de la Société slight but significant difference was observed for the Degree of Entomologique de France 84, 20–26. Infestation and SP in Trichopria cf drosophilae, which is difficult Blumenthal, D.M., 2006. Interactions between resource availability and enemy to explain since the two populations used were separated by only release in plant invasion. Ecology Letters 9, 887–895. Boulétreau, M., Fouillet, P., 1982. Genetic variability of suitability for a 46 km. More interestingly, a highly significant difference was Hymenopteran parasite in a natural population of Drosophila melanogaster. found for the two populations of L. heterotoma, separated by Comptes rendus de l’Academie des sciences. Serie III 295, 775–778. 340 km, with the higher Degree of infestation found for the popu- Boulétreau, M., Wajnberg, E., 1986. Comparative responses of two sympatric parasitoid cynipids to the genetic and epigenetic variations of the larvae of their lation from further south. This result confirms in the findings of host, Drosophila melanogaster. Entomologia Experimentalis et Applicata 41, previous studies revealing considerable latitudinal genetic differ- 107–114. entiation between populations of this area, with greater invest- Brown, P.H., Shearer, P.W., Miller, J.C., Thistlewood, H.M.A., 2011. The discovery and rearing of a parasitoid (Hymenoptera: Pteromalidae) associated with spotted ment in reproduction in southern populations (Fleury et al., wing drosophila, Drosophila suzukii in Oregon and British Columbia. ESA annual meetings, Reno NV, USA. 2004; Ris et al., 2004). Calabria, G., Maca, J., Bächli, G., Serra, L., Pascual, M., 2010. First records of the Finally we must highlight a significant block effect in many potential pest species Drosophila suzukii (Diptera: ) in Europe. Journal of Applied Entomology 136, 139–147. cases that could be related to a problem in the quality of the devel- Carton, Y., Boulétreau, M., Van Alphen, J.J., VanLanteren, J.C., 1986. The Drosophila opment of D. suzukii. As stated in Section 2, we used a modified Parasitic Wasps. In: Ashburner, M., Carson, H.L., Thompson, D.J. (Eds.), The Genetics and Biology of Drosophila. Academic Press, London, pp. 347–393. medium enriched with banana for rearing the flies, because this Carton, Y., Chibani, F., Haouas, S., Marrakchi, M., 1987. Egg-laying strategy under species has very low fecundity on standard medium (but with an natural conditions of Leptopilina boulardi, a Hymenopteran parasitoid of average of four eggs laid per day and per female on banana med- Drosophila spp. Entomologia Experimentalis et Applicata 43, 193–201. ium). However, development was carried out on a standard med- Carton, Y., Haouas, S., Marrakchi, M., Hochberg, M., 1991. 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$,%0.!jT.+/+,$%(/161'%%,.+"%0!W0W!((! Z1*.!(9 $!)!*0 !(,.!//%+* !/!/!**!)%/*01.!(/!* Physiological Entomology (2013) 38, 45 – 53 DOI: 10.1111/phen.120021.+,!Y

Resistance of Drosophila suzukii to the larval parasitoids Leptopilina heterotoma and Asobara japonica is related to haemocyte load

M A T H I L D E P O Y E T 1,3, S E B A S T I E N H A V A R D 2, G E N E V I E V E P R E V O S T 1, O L I V I E R C H A B R E R I E 1, G E R AL DI NE DO UR Y 1, 3 1 P A T R I C I A G I B E R T and P A T R I C E E S L I N 1 Unite´ Ecologie et Dynamique des Syste´mes Anthropise´s, Universite´ de Picardie Jules Verne, Amiens, France, 2 Laboratoire Diversite´, Ge´nomes et Interactions Microorganismes-Insectes, Universite´ de Montpellier 2, Montpellier, France and 3 Laboratoire de Biome´trie et Biologie Evolutive, Universite´ Lyon, Lyon 1, France

Abstract. Unlike other Drosophila species, the invasive Drosophila suzukii Mat- sumura (Diptera: Drosophilidae) shows a remarkable pest status. Among the physio- logical traits that may explain the high level of resistance to parasitoids of Drosophila larvae, the haemocyte load is shown repeatedly to play an important role. To determine whether haemocyte load can explain immunity resistance of D. suzukii to parasitoids, the haemocytes of parasitized and healthy larvae are quantified in two Japanese and three French populations of D. suzukii. Parasitization tests are conducted with two larval parasitoids: the paleartic Leptopilina heterotoma Thomson (Hymenoptera: Fig- itidae) and the Asian Asobara japonica Belokobylskij (Hymenoptera: Braconidae). Based on morphological and functional criteria, D. suzukii has classes of haemocytes similar to those described in Drosophila melanogaster. However, healthy larvae of the five populations tested possess particularly large numbers of haemocytes com- pared with D. melanogaster. Haemocyte load is also higher in larvae from the French populations than in the Japanese strains. The ability of D. suzukii larvae to encapsu- late eggs of L. heterotoma is associated with a particularly high load of circulating haemocytes. However, it is notable that A. japonica induces a strong depression of the haemocyte population in this resistant host associated with an inability to encapsulate parasitoid eggs. The results show that the cellular immune system plays a major role in the failure of larval parasitoids to develop in most instances in larvae of D. suzukii, possibly contributing to the success of this species as an invader.

Key words. Asobara japonica, Drosophila suzukii, haemocyte concentration, immune resistance, Leptopilina heterotoma.

Introduction intercontinental transportation, local causes of the success of individual species are still a matter of debate. Two major and The increase of invasive insect species has been recorded complementary hypotheses contribute to our understanding of worldwide within the past decade (Huang et al., 2011). the success of invasive species: (i) the enemy release hypoth- Although invasions are known to be largely a result of human esis (Keane & Crawley, 2002) considers that the success of a activities through the development of international trade and species outside its native range is the result, at least partly, of

the absence or the reduced efficiency of natural enemies, and Correspondence: Patrice Eslin, Laboratoire de Bio-e´cologie des (ii) facing this lack of enemy, the evolution of increased com- Insectes Phytophages et Entomophages, Universite´ de Picardie Jules petitive ability hypothesis (Blossey & No¨ tzold, 1995) predicts Verne, 33 rue Saint Leu, F-80039 Amiens Cedex, France. Tel.: +33 that non-indigenous species will be more productive and thus (0)3 22 82 75 47; e-mail: [email protected] successful in habitats into which they are introduced compared

© 2013 The Royal Entomological Society 45

Resistance to parasitoids in D.suzukii with their native habitats. Therefore, the success of an invasion exhibited by lepidopteran plasmatocytes (Ribeiro & Brehe´lin, depends not only on environmental factors of the colonized 2006). Lamellocytes are considered as the major capsule- habitat (invasibility), but also on the biological characteristics forming haemocyte type in Drosophila (Brehe´lin & Duvic, of the invasive species (invasiveness). These two hypotheses 1999; Havard et al., 2009). These cells are large and round are tested mainly in plants (Keane & Crawley, 2002; Blumen- flattened, and represent approximately 6% of the total haemo- thal, 2006; Facon et al., 2006; Liu & Stiling, 2006) but are less cyte count in non-immune-challenged Drosophila larvae (Eslin studied in , especially insects. According to the Euro- & Pre´vost, 1996, 1998; Lanot et al., 2001). They participate in pean DAISIE (Delivering Alien Invasive Species Inventories in the encapsulation reaction against intruders that exceed the size Europe) programme conducted between 2005 and 2008, 85% limit for phagocytosis by plasmatocytes (Eslin et al., 2009). of the 1517 exotic invertebrate species already established on Both the number and maturity of the haemocytes circulating at the European continent are insects (DAISIE, 2009). Never- the time of parasitization may be important factors influencing theless, the biological traits determining the success of these the success of the encapsulation process in Drosophila species invasions are still poorly understood. (Eslin & Pre´vost, 1998, 2000). Drosophila suzukii Matsumura (Diptera: Drosophilidae) is Recent studies show that different strains of D. suzukii [a a Drosophila species belonging to the melanogaster group. French one (Chabert et al., 2012) and two American ones Native to Asia, it has recently colonized the American (Hauser (Kacsoh & Schlenke, 2012)] are resistant to the majority of et al., 2009) and European (Calabria et al., 2010) continents. the larval parasitoids tested. Increased resistance against larval The infests a variety of commercial fruits, including rasp- parasitoids also appears to be associated with a high haemocyte berry, strawberry, grape and tree fruits (cherry, kiwi, fig, apple, load in D. suzukii (Kacsoh & Schlenke, 2012). Kacsoh plum, peach) (Hauser et al., 2009; Steck et al., 2009). Its inva- & Schlenke (2012) compare the amounts of constitutive sion is extremely fast and raises many concerns from Amer- haemocytes between D. suzukii and D. melanogaster larvae ican and European producers of red fruit facing important either after a wound inflicted by a sterile needle or after economic losses (Goodhue et al., 2011; Walsh et al., 2011). parasitization by a strain of the parasitoid Leptopilina boulardi Drosophila suzukii represents an excellent biological model (Hymenoptera: Figitidae) that is considered avirulent to for understanding better the conditions and factors allowing D. melanogaster host. a successful biological invasion. Researchers already benefit In the present study, the haemocytes of D. suzukii larvae par- from a huge knowledge on Drosophila biology, genetics and asitized by the Paleartic parasitoid species Leptopilina hetero- ecology regarding competitive interactions and the selective toma Thompson (Hymenoptera: Figitidae) or the Asian species pressure exerted by natural enemies (Hemmat & Eggleston, Asobara japonica Belokobylskij (Hymenoptera: Braconidae) 1988; Fleury et al., 2009; Verspoor & Haddrill, 2012). The are described and quantified. Both of these larval parasitoid same data on D. suzukii would provide a greater insight into the species are known to be highly virulent to D. melanogaster and mechanisms underlying successful biological invasions. From their effects on the regulation of the host larval immune system a more applied perspective, this would allow the opportunities are known (Rizki & Rizki, 1984; Mabiala-Moundoungou et al., available to halt the spread of this pest to be better identified. 2010). Five strains of D. suzukii (three collected in France Among natural enemies, parasitoids constitute a group recently and two others from Japan) are used to test whether whose impact on insect communities is now well described. D. suzukii suffers a decline of its circulating haemocyte popu- They play a key role by controlling insect pests and affect- lation after parasitization, and also whether this may be linked ing invasive populations (Magal et al., 2008). In host insects, to the success or failure of the development of the parasitoid. the process of encapsulation is the main physiological defence described against endoparasitoids. The parasite is enclosed within a capsule, which undergoes a progressive blacken- Materials and methods ing as a result of melanization (Carton & Nappi, 1997). The interaction between frugivorous Drosophila, especially D. Insects melanogaster, and their parasitoids is the subject of many stud- ies (Pre´vost, 2009). In the larval stages of D. melanogaster, Five strains of D. suzukii were tested. The three French three main types of haemocytes are described that play a role strains were collected using banana traps. One strain was in the formation of capsules. The plasmatocytes represent the collected in Sainte-Foy-les-Lyon (latitude: 45◦ 7 ) in 2010, most abundant haemocyte type circulating in the haemolymph. whereas the other two were collected in 2011 in Gotheron These small rounded cells, also called Drosophila plasmato- (latitude: 44◦ 4 ) and Compie`gne (latitude: 49◦ 24 ). The two cytes (Ribeiro & Brehe´lin, 2006), are not only involved in Japanese strains were collected in Tokyo (strain E-15014 phagocytosis, but also in the initial and terminal steps of encap- OGH06-03) in 2006 and in Tsushima (strain E-15017 TSM92) sulation (Russo et al., 1996). Crystal cells, the least abundant in 2008, and were provided by Masayoshi Watada (Ehime haemocyte type, contain elements (substrate and enzymes) of University, Japan). For phagocytosis assay, one strain of the phenoloxidase cascade. They are involved in the melaniza- D. melanogaster originating from Sainte-Foy-les-Lyon, France tion of the capsules, as well as in clotting and the produc- (collected in 1994), was used. All strains were mass reared tion of cytotoxic radicals (Evans et al., 2003; Meister, 2004; at 20 ◦ C and were fed a regular banana Drosophila diet Carton et al., 2008). The third haemocyte type is the lamel- (Chabert et al., 2012) and maintained under an LD 13 : 11 h locyte, whose structure and function are very similar to those photocycle.

Resistance to parasitoids in D.suzukii

Two species of Drosophila larval parasitoids were used number of cells. A total haemocyte count (i.e. total number of in this experiment: the cynipid L. heterotoma Thomson immune cells) and a differential haemocyte count (i.e. number (Hymenoptera: Figitidae) was collected in October 2010 of plasmatocytes, lamellocytes and crystal cells) were per- in France (Rhoˆ ne valley) using banana traps, and the formed. Haemocyte counts were expressed as the number of Japanese strain of A. japonica Belokobylshij (Hymenoptera: cells per mm3 of haemolymph. Braconidae) was graciously provided by Professor J. van Alphen (Leiden University, The Netherlands). Both parasitoid species were mass reared in the laboratory at 20 ◦ C on D. Development experiment melanogaster under a LD 13 : 11 h photocycle. In this experiment, parasitized and nonparasitized larvae were allowed to complete their development at 20 ◦ C until Procedure for controlled parasitization emergence as an adult Drosophila or parasitoid. Parasitization was achieved under controlled conditions as described above. To ensure that Drosophila larvae would develop syn- In each strain of D. suzukii, 100 repetitions were established chronously, eggs of D. suzukii and D. melanogaster were for each treatment. Adult Drosophila and parasitoids were collected after a 4-h oviposition period. The second-instar lar- counted at emergence. The mean number of emerging vae were then exposed to females of either L. heterotoma or from the unparasitized control gave an estimate of the via- A. japonica for 96 h. Parasitization was observed thereafter bility of each D. suzukii strain in the absence of parasitism. under a stereomicroscope. After a single oviposition by a par- Drosophila adults carrying a capsule were counted as hosts asitoid female, each parasitized D. suzukii larva was removed resistant to parasitism. The number of emerged A. japonica or individually and placed in a tube containing artificial banana L. heterotoma adults gave an estimate of the percentage of sus- Drosophila diet. ceptible larvae hosting a developing parasitoid. The difference between the number of flies emerging from the unparasitized control and the total number of insects emerging from the para- Phagocytosis assay sitized treatment gave an estimate of the number of parasitized larvae that died during development. The production of haemocytes was stimulated by controlled The total encapsulation rate (TER), the successful parasitism parasitization of third-instar larvae (7 days old at 20 ◦ C) by development rate (SPR) and the mortality rate of parasitized L. heterotoma 72 h before phagocytosis assay. For phagocy- larvae (MR) were estimated as follows: tosis assays, 20 nL of fluorescent polystyrene latex micro- spheres (approximately 11 000 beads; Fluoresbrite® – YG • TER = (number of Drosophila with a capsule/number of Microspheres, Polysciences Europe GmbH, Germany) were parasitized Drosophila ) × 100 microinjected into each larva (Havard et al., 2012). Thirty min- • SPR = (number of Drosophila permitting parasitoid devel- utes after the injection of the latex beads, the haemolymph opment/number of parasitized Drosophila ) × 100 was collected and the haemocytes were observed using phase • MR = (number of parasitized larvae that died during contrast microscopy under indirect ultraviolet illumination. development/number of parasitized Drosophila ) × 100

Haemocyte counts Statistical analysis

Ninety larvae per Drosophila strain were bled. Thirty of For each strain of D. suzukii, the number of haemocyte cells the larvae were parasitized by A. japonica and another group was compared between parasitized and unparasitized (control) of 30 larvae were parasitized by L. heterotoma. The remain- larvae using analysis of variance (anova; P < 0.05). anova ing 30 larvae were unparasitized (control). Haemolymph was was also used to compare the number of haemocytes between collected from Drosophila larvae by sectioning the body pos- control and parasitized larvae for each of the five strains of teriorly to the mandibules with opthalmic scissors (Eslin & D. suzukii. Normality and homosedasticity assumptions were Pre´vost, 1996). After bleeding, each parasitized larva was dis- checked before the analyses. The relationship between encap- sected to check for the presence of a parasitoid egg or larva sulation rates and mean total haemocyte counts was assessed and the status of parasitization success in terms of encap- using linear regressions. Statistical analyses were performed sulation. Hosts containing an encapsulated parasitoid (resis- using Statistica, version 6.1 (StatSoft, Inc., Tulsa, Oklahoma). tant hosts) or a non-encapsulated egg or larva (susceptible hosts) were separated. The haemocyte counts were made 72 h after parasitization when parasitized and nonparasitized lar- Results vae (7-day-old third-instar larvae raised at 20 ◦ C) provided sufficient haemolymph (approximately 0.2 μL) to be applied D. suzukii plasmatocytes show phagocytosis ability without dilution onto a Thoma haemocytometer slide (Eslin & Pre´vost, 1996). Haemocyte counts were obtained by applying Larvae previously immunostimulated by L. heterotoma haemolymph onto this slide and then immediately counting the parasitization produced a large quantity of haemocytes. As

Resistance to parasitoids in D.suzukii

Fig. 1. Phagocytosis by haemolymph plasmatocytes in third-instar larvae of Drosophila suzukii at 72 h post-parasitization by the parasitoid Leptopilina heterotoma and 30 min post-injection of 20 nL fluorescent polystyrene latex microspheres into each larva. Phase contrast images are shown on the left. Although lamellocytes are devoid of latex beads, plasmatocytes have engulfed the foreign bodies. Black arrowhead, plasmatocyte; white arrow, lamellocyte. Scale bar = 20 μm.

in D. melanogaster, most of the plasmatocytes of D. suzukii show that the haemocyte loads of the French D. suzukii strains larvae were found to engulf the injected fluorescent latex are eight-fold higher than the mean for the D. melanogaster microscopic beads, whereas no fluorescence was observed in strain. The results from the haemocyte counts (Table 1) showed the lamellocytes (Fig. 1). that these observations were consistent for the three categories of haemocytes: plasmatocytes, lamellocytes and crystal cells.

D. suzukii larvae have a very high load of circulating haemocytes Encapsulation of L. heterotoma eggs is associated with high total haemocyte count in D. suzukii larvae The three types of D. suzukii haemocytes (plasmatocytes, lamellocytes and crystal cells) were morphologically similar After parasitization by L. heterotoma, the larval development to those of D. melanogaster. By contrast to D. melanogaster of the host D. melanogaster was never completed, whereas lar- (total haemocyte count of approximatelly 5000 cells per mm3 vae of all strains of D. suzukii showed the ability to encapsulate of haemolymph; Eslin & Pre´vost, 1998), D. suzukii showed the eggs of this parasitoid species (Table 2). In particular, the a very high haemocyte load (Fig. 2). Total haemocyte counts French strains of D. suzukii showed a significantly higher abil- revealed that the Japanese strains of D. suzukii produced up ity to encapsulate this parasitoid than the Japanese strains. to five-fold more haemocytes (up to 25 000 cells) than D. Unexpectedly, L. heterotoma, a virulent parasitoid of melanogaster (data for unparasitized larvae). Also, the French D. melanogaster was unable to parasitize D. suzukii larvae strains of D. suzukii had almost twice as many haemocytes successfully. The total haemocyte counts recorded in the three (over 40 000 cells per mm3 of haemolymph) as the Japanese French D. suzukii strains were much higher than those recorded strains (F = 11.89, d.f. = 145, P < 0.0001). The data also in the two Japanese strains (Fig. 3). The three French strains showed very high counts of circulating haemocytes after par- asitization by L. heterotoma (up to 160 000 cells per mm3 of haemolymph), and counts that were significantly higher (F = 5.68, d.f. = 145, P < 0.0001) than for the two Japanese strains (up to 91 000 cells). One of the more remarkable observa- tions is that parasitization by L. heterotoma induced a decrease in the number of circulating haemocytes in D. melanogaster, whereas it led to a large increase in the total haemocyte counts of D. suzukii. Figure 4 shows that total haemocyte concentrations recorded in larvae 72 h post-parasitization by L. heterotoma and encap- sulation rates were highly correlated (r 2 = 0.93, P = 0.008).

Observations on lamellocytes of D. suzukii showed that lar- Fig. 2. Total haemocyte counts (mean ± SE) in unparasitized Drosophila suzukii larvae from three French strains (Compie`gne, vae parasitized by L. heterotoma were characterized by bipolar Gotheron and Sainte-Foy-les-Lyon) and two Japanese strains (Ogh 06 or elongated cells, whereas unparasitized (control) larvae and and Tsm 92). Different letters indicate a significant difference between those parasitized by A. japonica exhibited discoidal shaped strains (P < 0.05). For comparison, data showing haemocyte numbers cells. Also, the percentage of bipolar lamellocytes was lower in unparasitized Drosophila melanogaster larvae were obtained from in the Japanese strains (Table 1). Finally, in all strains of D. Eslin & Pre´vost (1998) (*). suzukii, only the numbers of crystal cells were significantly

Resistance to parasitoids in D.suzukii

Table 1. Differential and total haemocytes counts (THC) (mean ± SE) in Drosophila suzukii larvae from three French strains (Compie`gne, Gotheron and Sainte-Foy-les-Lyon) and two Japanese strains (Ogh 06 and Tsm 92) unparasitized (control) or parasitized by Leptopilina heterotoma or Asobara japonica.

Total number of haemocytes per mm3 of haemolymph

Lamellocytes Number Drosophila suzukii strains Conditions Plasmatocytes (% bipolar lamellocytes) Crystal cells of larvae Compie`gne Controls 36 369 ± 3621 2281 ± 502 1703 ± 166 30 Parasitized by Leptopilina heterotoma 128 489 ± 8122 33 458 ± 2416 1031 ± 174 30 (10.1 ± 1.5) Parasitized by Asobara japonica 13 989 ± 1411 1093 ± 285 229 ± 76 30 Gotheron Controls 33 151 ± 3667 1265 ± 151 1114 ± 175 30 Parasitized by Leptopilina heterotoma 100 234 ± 7416 29 515 ± 2968 671 ± 109 30 (11 ± 1.4) Parasitized by Asobara japonica 13 479 ± 1042 1604 ± 257 406 ± 105 30 Ste Foy Controls 36 489 ± 2128 3229 ± 713 2072 ± 288 30 Parasitized by Leptopilina heterotoma 130 572 ± 6180 29 750 ± 1180 1260 ± 175 30 (8.8 ± 1.8) Parasitized by Asobara japonica 10 385 ± 1359 1489 ± 623 333 ± 79 30 Ogh 06 Controls 18 671 ± 1847 1940 ± 228 1520 ± 181 30 Parasitized by Leptopilina heterotoma 63 580 ± 3244 22 005 ± 1705 39 ± 31 30 (7.8 ± 2.1) Parasitized by Asobara japonica 13 345 ± 676 2187 ± 189 427 ± 89 30 Tsm 92 Controls 20 812 ± 1682 1484 ± 333 1854 ± 194 30 Parasitized by Leptopilina heterotoma 68 872 ± 5967 22 008 ± 1536 489 ± 103 30 (6.5 ± 2.1) Parasitized by Asobara japonica 11 875 ± 1024 2137 ± 295 350 ± 82 30 Drosophila melanogaster Controlsa 4528 ± 856 28 ± 18 337 ± 88 21 Parasitized by Leptopilina heterotoma b 4129 ± 726 111 ± 31 127 ± 34 20 c Parasitized by Asobara japonica 2552 ± 320 158 ± 25 22 ± 10 52 For comparison, data for D. melanogaster larvae in the same conditions were obtained from: a Eslin & Pre´vost (1998), b Havard et al. (2012) and c Mabiala- Moundoungou et al. (2010) (analysis of variance; P < 0.05).

Table 2. Total encapsulation rate, successful parasitism rate and mortality D. suzukii were able to encapsulate them, although encapsu- rate in Drosophila suzukii larvae from three French strains (Compie`gne, lation occurred at different rates (Table 3). Among the five Gotheron and Sainte-Foy-les-Lyon) and two Japanese strains (Ogh 06 and D. suzukii strains tested, the Compie`gne strain showed the Tsm 92) parasitized by Leptopilina heterotoma. greatest ability to encapsulate A. japonica eggs (ER = 26%)

(Table 3). Host larvae of both D. melanogaster and D. suzukii Successful Drosophila Total encapsulation parasitism rate Mortality rate were also able to support the successful development of suzukii strains rate (%) (%) (%) A. japonica, although the success of parasitoid development (SPR) was higher in D. melanogaster (76%) than in D. suzukii Compie`gne 87 ± 0.2 0 13 ± 0.2 (50 – 63%) (Table 3). Gotheron 77 ± 0.4 0 23 ± 0.4 Ste Foy 80 ± 0.7 0 20 ± 0.7 In all five strains of D. suzukii, parasitization by A. japonica Ogh 06 62 ± 0.7 0 38 ± 0.7 generated a significant drop in the total number of haemocytes Tsm 92 59 ± 0.4 0 41 ± 0.4 (Fig. 3). This was observed for the three haemocyte types: Drosophila 0 85 15 plasmatocytes, lamellocytes and crystal cells. No significant melanogaster difference in the number of haemocytes was observed between the five strains of D. suzukii infested by A. japonica (approxi- Results were recorded after the insects (parasitoids and resistant hosts) had completed their development. Percentages (mean ± SE) were estimated mately 12 000 and 15 000 cells). However, the number of cir- from 100 larvae in each Drosophila strain. For comparison, data for culating haemocytes was five-fold greater in D. suzukii than in D. melanogaster larvae were obtained from Havard et al. (2012). D. melanogaster when larvae were parasitized by A. japonica. reduced in larvae parasitized by L. heterotoma compared with unparasitized larvae (F = 9.903, d.f. = 145, P < 0.0001). Discussion

The haemocytes of D. suzukii are morphologically and Parasitism by A. japonica generates a significant drop of the functionally similar to those described in D. melanogaster total haemocyte count in D. suzukii (Brehe´lin, 1982; Kacsoh & Schlenke, 2012). As in D. melanogaster (Lanot et al., 2001; Lavine & Strand, 2002; Although larvae of D. melanogaster were totally unable Meister, 2004), plasmatocytes represent the main, most to encapsulate eggs of A. japonica, the five strains of abundant haemocytic cell type in unparasitized D. suzukii

Resistance to parasitoids in D.suzukii

Fig. 3. Total haemocytes counts (mean ± SE) in Drosophila suzukii larvae from three French strains (Compie`gne, Gotheron and Sainte-Foy-les- Lyon) and two Japanese strains (Ogh 06 and Tsm 92) unparasitized (control) or parasitized by Leptopilina heterotoma or Asobara japonica. Different letters indicate a significant difference between blocks (P < 0.05). For comparison, haemocyte numbers in Drosophila melanogaster larvae under the same conditions were obtained from Eslin & Pre´vost (1998) (*), Havard et al. (2012) (**) and Mabiala-Moundoungou et al. (2010) (***) (analysis of variance; P < 0.05).

Table 3. Total encapsulation rate, successful parasitism rate and mortality rate in Drosophila suzukii larvae from three French strains (Compie`gne, Gotheron and Sainte-Foy-les-Lyon) and two Japanese strains (Ogh 06 and Tsm 92) parasitized by Asobara japonica.

Successful Drosophila Total encapsulation parasitism rate Mortality suzukii strains rate (%) (%) rate (%)

Compie`gne 26 ± 0.5 50 ± 1.1 24 ± 1.4 Gotheron 10 ± 1.1 63 ± 0.5 27 ± 1.3 Ste Foy 8 ± 0.4 56 ± 0.6 36 ± 0.8 Ogh 06 7 ± 0.2 56 ± 0.9 37 ± 0.8 Tsm 92 6 ± 0.3 60 ± 2.1 34 ± 1.9 Drosophila 0 76.8 ± 3.6 23.2 ± 3.6 Fig. 4. Correlation between encapsulation rate (TER) and the melanogaster total haemocyte count (THC; mean ± SE) in three French strains (Compie`gne, Gotheron and Sainte-Foy-les-Lyon) and two Japanese Results were recorded after the insects (parasitoids and resistant hosts) had strains (Ogh 06 and Tsm 92) of Drosophila suzukii larvae parasitized completed their development. Percentages (mean ± SE) were estimated by Leptopilina heterotoma (r 2 = 0.9267, P = 0.008). from 100 larvae in each Drosophila strain. For comparison, data for D. melanogaster larvae were obtained from Mabiala-Moundoungou et al. (2010). larvae, although crystal cells and lamellocytes are also found. In unparasitized larvae of D. suzukii, the lamellocytes range Pre´vost, 1998). The Japanese strains of D. suzukii carry four- to from 4% of the total number of circulating haemocytes (in the five-fold more haemocytes than D. melanogaster, whereas the Gotheron strain) to 10% (in the OGH 06 strain), a result that French strains have eight-fold higher haemocyte counts than is consistent with what is already known for the melanogaster D. melanogaster. Such differences between the French and the subgroup of Drosophila (Eslin et al., 2009) but different from Japanese or American geographical strains may be related to the results reported in other subgroups (Havard et al., 2009, genetic variations, which may contribute to the success of bio- 2012). By contrast, the total haemocyte counts measured in logical invasions (Lindholm et al., 2005), at the same time as D. suzukii are quite unusual. To the best of current knowl- reflecting the rapid evolutionary changes likely to occur among edge, the numbers of circulating haemocytes in D. suzukii invasive species (Sakai et al., 2001). larvae are the highest ever recorded for Drosophila larvae Larvae of the five D. suzukii strains show a prompt of the melanogaster group (for a comparison, see Eslin & haemocytic response after parasitization by L. heterotoma.

Resistance to parasitoids in D.suzukii

The high haemocyte load and the ability to trigger an The exceptionally high haemocyte load of D. suzukii larvae at increase in circulating plasmatocytes and lamellocytes quickly the time of parasitization is considered as likely contributing when challenged by L. heterotoma are correlated with the to their resistance towards the highly virulent A. japonica. The immune resistance to parasitoids. A correlation between the same phenomenon is reported for the interaction between the concentration of circulating haemocytes and the aptitude to closely-related parasitoid Asobara citri and host Drosophila form a haemocytic capsule is demonstrated for species in the simulans (Moreau et al., 2005). melanogaster subgroup (Eslin & Pre´vost, 1998). The present Cynipids from the Leptopilina genus and Braconids from study shows that such a correlation not only applies to another the Asobara genus are the most studied larval endoparasitoids host-parasitoid association (D. suzukii – L. heterotoma ) but of Drosophila to date. Over the last 20 years, an extensive also is true at the intraspecific level as well, considering knowledge has accumulated on the genetics and physiology variations among local strains of D. suzukii. Crystal cells are of the interactions between hosts of the melanogaster sub- the only haemocyte type to show a decline in number after group and their larval parasitoids. Molecular data regarding the parasitization by L. heterotoma. These cells are considered to virulence strategies of these parasitoids have also been charac- be main carriers of some of the enzymes of the phenoloxidase terized (Chiu et al., 2006; Colinet et al., 2007; Dubuffet et al., system (Meister, 2004). A quantitative decrease in the 2009; Pre´vost et al., 2011). This knowledge should facilitate circulating crystal cells could either be the result of cell our understanding of the interactions between D. suzukii and lysis releasing opsonizing factors in the early stages of the potential larval parasitoids. Ongoing studies on the molecular defence reaction or a parasitoid-induced disruption of the basis of the virulence of A. japonica towards D. suzukii are functioning of the phenoloxidase system (Nappi et al., 2009). expected to bring some insight to both fundamental and applied Bipolar lamellocytes range from approximately 6.5% (Tsm92 research, in particular regarding the control of this invasive strain) to 11% (Gotheron strain) in parasitized larvae of D. fruit fly. suzukii. Virulent strains of L. boulardi and L. heterotoma exert In conclusion, the results of the present study support the an active depressive effect on the immune system of their hypothesis that the cellular immune system of D. suzukii plays Drosophila hosts (Rizki & Rizki, 1991; Dubuffet et al., 2009). a major role in the failure, in most instances, of larval para- The proportion of affected lamellocytes can represent up to sitoids to develop in larvae of this species. The results suggest 68% of haemocytes in D. melanogaster larvae parasitized by that D. suzukii has evolved a more potent cellular immune L. heterotoma (Rizki & Rizki, 1991). The molecular factors response than any other Drosophila species (i.e. tested to responsible for this effect in D. melanogaster are located date), providing resistance to the guild of parasitoids poten- in the wasp’s venom and have been identified (Chiu et al., tially present in its habitat. High numbers of haemocytes may 2006; Dubuffet et al., 2009; Gueguen et al., 2011). The results allow better protection from several physiological constraints of the present study suggest that the immunosuppressive and could improve the ability to respond quickly to envi- factors of L. heterotoma could act on some haemocytes of ronmental changes. The rapid dissemination of D. suzukii in D. suzukii larvae, in particular the lamellocytes and possibly North America and Europe demonstrates the remarkable inva- the crystal cells, although this effect may not be sufficient to sion capability of this species. In addition to other life-history prevent the encapsulation reaction. One possible explanation traits that are presently under study, the high haemocyte load for the observed difference between the immune response of D. suzukii suggests that this species is particularly well towards L. heterotoma in D. suzukii and D. melanogaster armed for successful invasion. Identifying the biological fac- (Rizki & Rizki, 1991) could be that European populations of tors involved in the success of an invasion is an important step L. heterotoma are not adapted to this new exotic host. This towards defining suitable measures for invader management. hypothesis is consistent with the predictions of the enemy According to the evolution of increased competitive ability release hypothesis. hypothesis (Blossey & No¨ tzold, 1995), a host could reallocate Drosophila suzukii appears to be the first species of resources from defence mechanisms into growth and develop- the melanogaster subgroup to resist the immunosuppressive ment in the absence of effective natural enemies. Populations properties of the parasitoid L. heterotoma by encapsulating might be evolving towards an opposite direction. By contrast, its eggs. Encapsulation of A. japonica by D. suzukii remains it appears that D. suzukii, which is invading Europe, avoids at a low level, as also reported by Chabert et al. (2012) and the competitor species that it may have in its exotic range Kacsoh & Schlenke (2012). The present study shows that the because it is the only Drosophila species using unripe fruits. number of circulating haemocytes is consistently reduced in the Such a gain of energy could be reallocated into the production five D. suzukii strains 72 h post-parasitization by A. japonica. of haemocytes, therefore improving the resistance of the fly to Parasitism by A. japonica also affects haemocyte load and parasitoids, a condition that would certainly contribute to the the phenoloxidase activity of D. melanogaster (Mabiala- success of its invasion. Moundoungou et al., 2010). Therefore, A. japonica, which exists in sympatry with D. suzukii in Japan, appears to exert immunosuppressive effects on larvae of D. suzukii that appear Acknowledgements to be similar to those described in D. melanogaster. However, D. suzukii is able to encapsulate eggs of A. japonica eggs at This work was supported by the ANR (project CLIMEVOL – rates ranging from 6% (Tsm 92 strain) to 26% (Compie`gne ANR-08-BLAN-0231) and the GDR-CNRS 2153. James strain), which are never observed in D. melanogaster larvae. Langan is thanked for help with the English language.

Resistance to parasitoids in D.suzukii

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The invasive pest Drosophila suzukii may benefit from resource opportunity

in temperate Europe

Mathilde Poyet a,b, Vincent Le Roux a, Patricia Gibert b, Antoine Meirland c, Geneviève

Prévost a, Patrice Eslin a, Olivier Chabrerie a

a Unité Ecologie et Dynamiques des Systèmes Anthropisés (FRE-CNRS 3498), Université de

Picardie Jules Verne, F-80037 Amiens Cedex, France b Laboratoire de Biométrie et Biologie Evolutive (UMR-CNRS 5558), Université Claude

Bernard Lyon1, F-69622 Villeurbanne, France c Gemel-Picardie, Maison de l’Université de Picardie Jules Verne, 115 quai Jeanne d'Arc, F-

80230 Saint-Valery-Sur-Somme, France

Authors’contributions

Conception of the study: P. Eslin, O. Chabrerie.

Field sampling (plants): A. Meirland, O. Chabrerie.

Laboratory tests: M. Poyet, V. Le Roux, P. Eslin.

Statistical analyses: O. Chabrerie.

Writing: M. Poyet, V. Le Roux, P. Gibert, G. Prévost, P. Eslin, O. Chabrerie



Introduction

Biological invaders are considered as one of the most important drivers of biodiversity loss (Sala et al. 2000; Clavero and García-Berthou 2005). Alien species may cause serious impacts on native communities, habitats, ecosystem processes (Simberloff et al. 2013) and, consequently, alter ecosystem services, which are key elements for human well-being

(Charles and Dukes 2007; Pejchar and Mooney 2009; Clarck et al. 2014). In the case where a part of their life cycle occurs in production systems, invasive species may lead to huge economic impacts (Goodhue et al. 2011; Lee et al. 2011). Predicting the success of exotic species and finding common rules in invasion ecology is still challenging, regarding the specificity of each invasion. Indeed, invasion success may depend on a great variety of introduction histories (Jerde and Lewis 2007), on the probability of climate/habitat matching

(i.e. ecosystem invasibility) (Hayes and Barry 2008; Kolar and Lodge 2001) and on a wide set of species-level characteristics (i.e. species invasiveness) (Hayes and Barry 2008; Sakai et al.

2001).

Compared with other phylogenetic groups, insects have been for a long time neglected regarding their potential ecological impact, except those which have sanitary or economic impact, especially related to agriculture or forestry (Roques 2010, Kenis et al. 2009). There are about 1390 insect species introduced in Europe, including 98 Diptera, originating largely from North America or Asia, with an over-representation of phytophagous species, and in particular during the last decade (Roques 2010). Most of introduced Diptera species are largely associated with anthropic environments, such as urban areas and agrosystems

(Skuhravá et al. 2010). Among them, the Drosophila genus experienced a long history of invasion due to the high fecundity, short regeneration cycle and high ability of its members to adapt to changing environments (Rota-Stabelli 2013). Very recently, Drosophila suzukii 

(Matsumura 1931) has become the 9th Diptera of this genus introduced in Europe and is by far the most impacting drosophila species in agricultural areas (Skuhravá et al. 2010).

The geographic origin of D. suzukii is still not clearly known and subjected of debate.

A first scenario suggested a Japanese origin (first record in 1916, and first description by

Matsumura in 1931) and a subsequent spread in different islands of this country and, finally, a progressive spread in South-Est Asia and Eurasia (Calabria et al. 2010). A second scenario using phylogenomic methods suggests a speciation from Drosophila biarmipes group in the

Himalayan/Tibetan plateau followed by a secondary dispersal from temperate climate areas towards Asia (Ometto et al. 2013). In its contemporary exotic range, D. suzukii was first recorded in 2008 in the USA (California) and jointly in South Europe (Spain and Italy;

Hauser et al. 2009; Cini et al. 2012; Rota-Stabelli 2013). Currently, this pest is undergoing a rapid expansion in both North America and Europe (Hauser et al. 2009, Calabria et al. 2010;

Rota-Stabelli 2013), with a so high velocity (about 1000 km per year) that this invasion has few precedents (Rota-Stabelli 2013). In France, it was first recorded in 2009, in the south and reached the north during the following years (Poyet et al. 2013, 2014), probably dispersed both by itself by flying and by passive transport of larvae associated with fruit trade (Liebhold and Tobin 2008; Rota-Stabelli 2013). Dispersal is not a limiting factor to the expansion of this species (Rota-Stabelli 2013) characterized by a migratory behavior in its native and exotic ranges (Mitsui 2010; Rota- Stabelli et al. 2013) and known as a commensal of human

(Lockwood et al. 2007). The species is now largely installed in natural and production systems in which it interacts with resident species and preempts a large part of their fruit production (Poyet et al. 2014).

Drosophila suzukii is currently subjected to intense research because of its huge impact on the red fruit industry in its introduced areas, particularly in Europe and North America (Walsh et al. 2011; Goodhue et al. 2011). Unlike the vast majority of Drosophila flies, which trophic  niches are based on fungi and rotten or over-ripened fallen fruits, the trophic niche of D. suzukii consists of fresh and ripening fruits (Poyet et al. 2014). This fly is able to pierce the skin of fruits using a serrated ovipositor which allows it to lay eggs more deeply in the fruit flesh (Atallah et al. 2014). Oviposition scars give fungi, incl. yeasts, and bacteria access to the fleshy tissues of the fruits which may rot prematurely (Walsh et al. 2011). Commercial small fruits, including blueberry, strawberry, blackberry, raspberry, tomatoes and grape and tree fruits, such as cherry, kiwi, fig, apple, plum, peach and others (Hauser et al. 2009; Steck et al.

2009; Lee et al. 2011; Bellamy et al. 2013; Rota- Stabelli et al. 2013), are suitable hosts potentially damaged by this fly. In 2008 the cost of its invasion was estimated to 500 million dollars for all crops in the USA (Bolda et al. 2010; Bolda et al. 2010). As these economic consequences are the result of the first steps of a very recent and still ongoing invasion, they are likely to increase in the next years, leading to a serious reduction of the revenues of fruit producers (up to 37% according to Goodhue et al. 2011).

Recent advances in invasion ecology offer new concepts that may help understanding the success of D. suzukii in its exotic range. The invasiveness of D. suzukii mainly lays on the presence of its saw-tooth ovipositor which represents an evolutionary innovation (Rota-

Stabelli 2013) and a new weapon (Callaway & Aschehoug 2000; Callaway & Ridenour 2004) in the introduction areas and an advantage on the other drosophila species (see Atallah et al.

2014 for the rapid evolution of that weapon). The evolution of this innovative organ, and the associated behaviour in unmature fruit recognition (Rota-Stabelli 2013), provided a new fundamental function to the fly: the ability to use the flesh of fresh ripening fruits and to preempt their resources before the other species that lay eggs mainly on the surface of rotten fruits (D. melanogaster, D. subobscura). By using this early temporal window in the fruit maturation chronosequence, this new function offered a new trophic niche to D. suzukii. As the ripening fruits are neglected by the other drosophila species (Poyet et al. 2014), D. suzukii  benefits from an empty niche (Stachowicz and Tilman 2005), escapes from major resident competitors and natural enemies (Keane and Crawley 2002) and may consequently increase its performances in the interactions network of the novel ecosystem (Blossey and Nötzold

1995). This increased invasiveness coupled with the absence of drosophila competitor may have also helped the fly to colonize a high diversity of habitats and reproduction/feeding substrates. Indeed, D. suzukii is known to be a polyphagous fly, infesting a great variety of

Prunus stone fruits its native area (Kanzawa 1936; Kanzawa 1939; Dreves et al. 2009; Mitsui et al. 2010; Kasuya et al. 2013) and numerous families of cultivated fruits in the agrosystems of its exotic range (Lee et al. 2011; Cini et al. 2012; Bellamy et al. 2013; Rota-Stabelli 2013).

However, a large screening of wild and ornamental plants hosting the species in its exotic range, especially those occupying temperate Europe, is still lacking from the literature and the mechanisms through which D. suzukii chooses its host is poorly known.

The plasticity of D. suzukii in fruit resource requirement and uptake is a key element of its success that may have led the fly to enlarge its fundamental and realized niche (Berg and Eller 2010) across times. All fruits are not equivalent regarding their suitability to the different stages of D. suzukii life cycle (eggs, larvae, emerged adults). Their suitability mainly depends on a large set of coevolving traits (volatile compounds, pH, shape, structure, firmness, resource quantity and quality, color…) preventing, limiting or favoring the development of the fly (Lee et al. 2011; Bellamy et al. 2013; Burrack et al. 2013). Hence, analyzing relationships between fruits’ functional traits and D. suzukii’s oviposition behaviour or larvae development might help in understanding the mechanisms implied in its invasion success, unrevealed by a taxonomic approach (i.e. species from different families can show similar fruit traits, whilst a high diversity of fruit shape, color can be found in a single family).

In functional ecology (Lavorel and Garnier 2002; Lavorel et al. 1997; Violle et al. 2007;

Weiher et al. 1999), plant species and traits are grouped according to common responses to  the environment (traits termed ‘response traits’) and/or common effects on ecosystem processes (‘effect traits’). For example, spines on leaves, a hairy skin, fruit firmness and color may be a response to herbivores, desiccation in drier environment, granivores and frugivorous seed vectors, respectively, while fruit nutrients and fruit quantity/size may influence ecosystem processes through the quantity and quality of organic matter entering the trophic networks. D. suzukii’oviposition behaviour may respond to fruit color and shape while fruit structure (including the presence of septum within complex fruits and internal partitions like in Rubus polydrupes) and size/diameter may influence the development process of larvae though the limitation of resources availability.

In this study, we examine the relationships between the community of fleshy-fruited plants of a temperate region of Europe and the invader D. suzukii. We tackle two main questions: (1) how many plants may host D. suzukii larvae and lead to full developments of imago? (2) Do fruit traits (structure, color, shape, skin texture, diameter and weight) interact with oviposition choices and development success of D. suzukii?

As D. suzukii is able to migrate across regions, along altitudinal and climatic gradients, within and between ecosystems according to seasons (Kimura 2004, Mitsui et al. 2010, Rota-

Stabelli 2013), fruits of a large set of 67 wild and ornamental plants have been collected during a whole year in the various ecosystems (forest, hedgerow, grassland, garden, coastal dune…) of the North of France and have been submitted to D. suzukii infestation in experimental conditions. As these different plants produce fruits along the four seasons of a year, they may guaranty continuous resource availability along a year and contribute to the persistence of the fly in natural and neighboring cultivated systems.



Materials and methods

Study area and field sampling

The study was carried out in Picardy region in northern France (N 48°50'19'' -

50°21'59''; E 1°22'50'' - 4°15'23''; alt. 0 - 296 m). The climate is of oceanic type, with a mean annual temperature of 10°C and annual rainfall of 700 mm. The geological substrate is mainly composed of Cretaceous chalks covered by clays and/or Quaternary loess. Sand dunes, shingle banks and salt alluvium locally occur along the coast. Landscapes are diversified and consist of mosaics of openfields, bocages, forests, urban areas and coastal vegetations. All these types of landscape have been visited to identify the pool of fleshy-fruited plants that could be potentially used as host plants by D. suzukii during the four seasons of a year.

Between October 2011 and November 2012, fruits from 67 fleshy-fruited plant species

(see details in Appendix 1) were collected in forests, hedgerows, grasslands, coastal shingle banks and dunes, gardens and urban areas in the region. For each plant species, fruits from five individuals separated by a minimum distance of 200 m were randomly collected and stored in different paper bags. Every month, the presence of fruits on the individuals of each species was recorded in the field. Fruits of 10 additional species (information for these additional species is given in Appendix 2) were collected but not included in the analyses because the number of fruits per individual or per species was too low to be tested.

Laboratory tests

For each plant species a total of 150 fruits (1 species x 5 individuals x 3 tests x 10 fruits) were used in laboratory tests. For each individual of each species, 30 fruits were  randomly selected. We carefully examined the fruits with a stereomicroscope (Leica MI 65 C) and excluded those already damaged or attacked by animals or pathogens. The 30 fruits were split into three sets of ten fruits and placed in ventilated transparent plastic boxes (15 cm x 10 cm x 5cm) to perform three types of test.

In the first test (‘adult emergence test’), 10 fruits were exposed to 3 D. suzukii mated females for 24 hours following the protocol of Poyet et al. 2014. After 24 hours, the number of eggs laid in each fruit was counted under a Leica M 165C stereomicroscope. Eggs oviposited in fruits were identified by the holes drilled by the females’ovipositor and by the presence of egg filaments as recommended by Mitsui et al. (2006). In our experiments, every hole that we observed contained one single D. suzukii egg. The numbers of D. suzukii flies emerging from each test were checked daily, flies were counted then removed from the experimental boxes to avoid new oviposition. In the second test (‘larvae development test’), 10 fruits were exposed to 3 D. suzukii females for 24 hours. After 24 hours, the number of eggs laid in each fruit was counted. After one week, fruits were dissected and the total number of larvae present in and on the fruits and in the plastic box of the test was recorded. The third set of 10 fruits was used as a control to monitor the potential fruit contaminations by other insects.

In all tests, we used the strain of D. suzukii collected in 2011 in the Compiègne forest in

Picardy (Poyet et al. 2013, 2014) under an LD 13 : 11 h photocycle at 20°C. D. suzukii strain was mass reared and were fed with a regular banana Drosophila diet (Chabert et al. 2012). We used 5 days old mated females for each experiment, to prevent the delay of eggs laying by young females on non adequate substrate.

Fruit traits



We targeted fruit characteristics associated to three fundamental challenges faced by

D. suzukii, i.e. egg laying in fruits, larvae development and adult emergence. A set of five biological traits (a total of 16 trait categories) was retained: fruit diameter and weight, type

(i.e. structure), color, shape and skin texture. The information was collected or extracted from flora (Lambinon et al. 2004; Provost 1998; Rameau et al. 1989) and plant databases (Tela botanica, Hortical). Fruit weight and maximum diameter were individually measured before laboratory tests on the 6694 fruits tested for the larvae development and adult emergence experiments and treated as continuous variables. Fruit type, color, shape and skin were treated as categorical variables; the optimal number of categories was defined to be ecologically meaningful and have balanced sizes (Chabrerie et al. 2010; Lavorel et al. 1998). Fruit type included four categories (1: berry; 2: drupe; 3: polydrupe and pseudo-polydrupe; 4: other fruits with complex structures: pseudo-fruit, complex fruit, multilocular capsule, aril), fruit color included six categories (1: black; 2: red; 3: pink; 4: orange-light brown; 5: white; 6: blue), fruit shape included two categories (1: spherical; 2: oval) and fruit skin texture, three categories (1: smooth and waxy, glossy, shiny; 2: smooth and pruinose; 3: rough, irregular).

Plant species nomenclature follows Lambinon et al. (2004) and plant family phylogeny follows APG III.

Data analyses

The effects of fruit traits on the numbers of D. suzukii eggs, larvae and adults emerging from fruits was examined using generalized linear models (GLM) with a Poisson distribution and a log-link term(Mccullagh and Nelder, 1989). In the egg-model (n=67 plant species tested), the number of D. suzukii eggs was the response variable, the fruit diameter was used as a fixed covariate and the fruit type, color, shape, surface type and skin thicknes  were used as fixed factors. In the larva-model, the number of D. suzukii larvae (n=60 plant species tested) was the response variable, the fruit diameter was used as a fixed covariate and the fruit type was used as a fixed factor. Seven species were excluded from this analysis

(Berberis julianae, Gaultheria procumbens, Ligustrum vulgare, Lonicera caprifolium,

Mespilus germanica, Pyrus calleryana 'Chanticleer', Sorbus aria) because we didn’t find enough undamaged fruits per individual to be tested. In the adult-model (n=67 plant species tested), the number of D. suzukii adults emerging from fruits was the response variable, the fruit diameter was used as a fixed covariate and the fruit type as fixed. To fulfil the normality assumption, fruit diameter and weight were log10-transformed. As fruit diameter and weight were highly correlated (R=+0.813; p<0.0001), fruit weight (less correlated with the number of egg laid in the fruits by the flies than the diameter was) was excluded from the analyses to avoid redundant explanatory variables in the models. SPSS v. 17.0 (IBM Corp., Somers, NY,

USA) was used in all analyses.

Results

Description

A very wide range of plant families was used by D. suzukii females and led to adult emergence (2 Adoxaceae, 1 Aracea, 2 Berberidaceae, 3 Caprifoliaceae, 1 Cornacea, 2

Eleagnacea, 1 Garryacea, 2 Grosslariaceae, 1 Moracea, 1 Phytolaccacea, 1 Rhamnacea, 9

Rosaceae, 1 Santalacea, 5 Solanaceae, 1 Taxacea). The plant species showing the highest number of D. suzukii eggs per fruit were Phytolacca americana, Prunus malheb, Rubus fruticosus agg., Viscum album and Prunus lusitanica (figures 1 and 2); those hosting with the highest number of larvae are Rubus fruticosus agg., Prunus mahaleb, Atropa belladonna,

Viscum album and Frangula alnus and those leading to the highest number of imago  emergences were Rubus fruticosus, Atropa belladonna, Prunus mahaleb, Prunus serotina, and Rubus idaeus (figure 2). The highest mean number of eggs per fruit was10.78 (± 1.68) for

Phytolacca americana, the highest mean numbers of larvae per fruits was 6.84 (± 1.37) for

Rubus fruticosus agg. and the highest mean numbers of adult emergences per fruit was 5.20

(± 1.83) for Rubus fruticosus agg..

Among the 67 plant species tested, 33 of them (49.25 %) led to the emergence of

D. suzukii imago, 6 of them (8.96 %) hosted larvae that didn’t reach the adult stage, 11 of them (16.42 %) hosted eggs that didn’t reach the larva stage and 17 of them (25.37 %) were not used by D. suzukii females.

Effects of fruit traits

The GLM showed significant effects of fruit traits on the number of D. suzukii eggs, larvae and adults emerging from fruits (Table 1). The number of eggs, larvae and adults increased significantly with fruit diameter. The number of eggs was higher in berries and drupes than in the other fruit types, higher in white or black fruits than in fruits with other colors, higher in oval than in spherical fruits and higher in rough than in smooth fruits especially in those with a pruinose coating. The numbers of larvae and emerging adults were higher in polydrupes than in the other types of fruits and were also higher in drupes and berries than in fruits with a complex structure.

Species range and fruits phenologies

Among the 33 plant species that led to the emergence of D. suzukii adults, 21 of them

(63.6%) were ornamental or cultivated species, 14 (42.4%) were exotic and 4 (12.1%) were  invasive species in the region. Several naturalized plant species (i.e. observed in the nature but not considered as invasive at present; Lonicera nitida, Ribes sanguineum and

Symhoricarpos albus) also led to the emergence of D. suzukii adults (Appendix 1 and Figure

3). Among the 67 plants tested, the number of larvae per fruit was lower in ornamental (0.49

± 0.12 larvae.fruit-1) than in other plant species (1.57 ± 0.50 larvae.fruit-1; t=-2.799, p=0.007).

The number of emerging adults per fruit was lower in ornamental (0.36 ± 0.09 larvae.fruit-1) than in other plant species (1.06 ± 0.35 larvae.fruit-1; t=-2.548, p=0.013).

The number of plant species with fruits potentially suitable for the development of

D. suzukii offspring was high between September and December with a peak in October

(figure 4). Few host plants suitable for D. suzukii were fruiting in early spring season, but fruits of several potential native and exotic hosts remained hanged on plants during winter

(Viscum album, Aucuba japonica, Hippophae rhamnoides subsp. rhamnoides,

Symphoricarpos albus). These 33 suitable host plants occurred in all the habitats of the region: forest (n=4 species), hedgerow (n=9), grassland (n=3), wetland (n=1), coastal dune and shingle bank (n=2), garden (n=6), urban area and park (n=8).

Discussion

Biological invasions studies have for a long time focused on two separate aspects: the characteristics of invaders and those of the invaded ecosystem. Now, the success of biological invasions is rather seen as a match between the alien and the ecosystem invaded (Facon et al.

2006; Shea and Chesson 2002). According to the Niche opportunity concept (Shea and

Chesson 2002), who can explain the success of biological invasion, Drosophila suzukii could make use of tree promoting factors: the pre-adaptation of physical environment, the decreasing of natural enemies and the increasing of resources availabilities. 

Indeed, the alien D. suzukii shows many physical environment pre-adaptations to invade

Europeen temperate climate. This species is adapted to subtropical and temperate climates, it is able to tolerate a large range of temperatures and to overwinter (Rota-Stabelli 2013; Ometto et al. 2013). This tolerance which not only match with the climate in invaded areas, but also which would be more favorable with climate change. In Europe, D. suzukii established in many ecosystems, such as agrosystems, orchads, bocage, gardens urban areas (pers obs), forests (Poyet et al. 2014) and mountain ecosystems in its native area (Mitsui et al. 2010).

Moreover, it seems that D. suzukii have no natural enemies able to reduce its spread.

According to the enemy release hypothesis (Keane and Crawley 2002) and the Natural enemy opportunity (Shea and Chesson 2002), its specialized enemies are absent from the introduction area and generalists enemies attack preferentially indigenous species. In fact,

Chabert et al. (2012) have showed that of the five main European parasitoid species, only two pupal parasitoids with wide host ranges develop on D. suzukii.

Regarding the resource, none Drosophilidae presents same characteristics of D. suzukii in the invaded areas, and according to Rota- Stabelli et al. (2013) there are virtually no competitors for fresh fruits, which is an argument to say that D. suzukii exploit a vacant niche.

The only potential competitors could be frugivorous birds, but up to now no competition has been demonstrated.

D. suzukii is a very polyphagous species

Unlike the majority of other invasive insects and pests, D. suzukii does not infest a single species or a single family of host plants. Indeed pests are often specialists, as is the case  for two emblematic pests in Europe: the grapevine phylloxera (Daktulosphaira vitifoliae) and the Colorado potato beetle (Leptinotarsa decemlineata) (Omer et al. 1999). Conversely, D. suzukii infested plants belonging to different orders such as Liliopsida (eg arum maculata) and Magnoliopsida.

As agreed D. suzukii lays eggs and grows efficiently on a large number of species of the Rosaceaes family (Kanazawa 1939 Dreves et al 2009; Sasaki and Sato 1995). Indeed this insect is known for the havoc it creates on the red fruits like cherry, strawberry, raspberry, blueberry ....

In 2012, Cini et al. tried to create a list of potential host plant including Rosaceae,

Ericaceae, Grossulariaceae, Moraceae, Rhamnaceae, Actinidiaceae, Ebenaceae, Myrtaceae,

Rutaceae, Caprifoliaceae and Adoxaceae.

However, we have also seen D. suzukii laying on fourteen other families host plants.

And sometimes, on plants known for their toxic (for mammals) or entomotoxic properties.

For example D. suzukii is particularly well developed on the mistletoe (Viscum album), on the

Belladonna (Atropa belladonna), on the Alder Buckthorn (Frangula alnus), or on the

European Black Nightshade (Solanum nigrum).

With this broad polyphagia D. suzukii find host plants throughout the year. The winter phase of the insect may compensate the host plant absence that we can be observed in the spring. Moreover, even if the plant species and their phenology varies according to geographical areas, we can think that D. suzukii always find a potential host plant.



Some fruits features are more attractive

On the attractiveness of host plants, there are limiting factors. Infact D. suzukii refused to lay eggs and feed on some species. However, as the no choice tests were conducted on laboratory, it is important to take a step back from some of them. For example fruits with a larger diameter and thicker skin will be less sensitive to desiccation. While in natura and on the tree fruits are less sensitive to desiccation.

As Burrack et al. (2013) showed fruit penetration strength is one potential measure of host susceptibility. Without redoing their penetration tests, we simply arbitrarily assigned categories of different fruit species to classified them according to the thickness of their skin.

Our results confirm those of Burrack since D. suzukii lays more on fruit whose skin is thinner.

But host attractiveness will likely depend upon additional factors, such as soluble sugar content. So other tests should be done to definer favorable traits in the development of D. suzukii in fruits.

D. suzukii presence will impact plant communities

D. suzukii is a time bomb in natural plant communities. As numerous plants reproduce and disperse with thin-skinned fleshy fruits in temperate European forests and given the damage caused by the fly on cultivated sweet cherries (Beers et al. 2011), we could expect significant impacts of D. suzukii invasion at the plant community scale. The fertility and the dispersal hierarchy between plant species could be modified as well as their frequency and their place in the communities. A shift in the functional composition of forest communities could occur: fleshy-fruited species will be disadvantaged to the benefit of plants reproducing  through vegetative organs or dry seeds unattacked by D. suzukii infestation. Consequently, anemochorous and ectozoochorous processes would be more likely to determine the spatial patterns of species than endozoochorous processes in the future D. suzukii’s word.

Nothing seems to stop the Drosophila suzukii progression in Europe. The number of host plants may be reservoirs for the insect is very important. The presence of host plants as nesting or nutrition does not seem to be a limiting factor to expansion. Also according to

Poyet et al (2013) natural predators such as parasitoids are not able to limit populations.

Only environmental conditions such as temperature (Calabria et al. 2010) may limit the D. suzukii spread. But in Japan this species is known to be able to compensate for the warmer temperatures in the winter and migrate to the altitude (Biosecurity Australia 2010).

But D. suzukii polyphagia and ability to lay eggs on a large number of host plant species could have a negative effect on the insect fitness. Indeed, D. suzukii can lay eggs on eleven species which is not able to develop. Its development is stopped before hatching the egg or in the first larval stage. Among these species, Gaultheria procumbens, Rubia tinctoria and

Pyracantha sp. gives interesting results. On this species D. suzukii lays two to five eggs per fruit and no development ensues. This observation is very interesting, it would allow us to create "trap plants list" to place around crops to protect the red fruit. For this it is necessary to continue this work by conducting tests of choice between grown fruit and this wild species fruits.



References

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Appendix 1: Characteristics of the 67 plant species tested in the study.

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Appendix 2: Emergences of Drosophila suzukii imago from the fruits of an additional set plant species. These additional species were collected but not included in the analyses because the numbers of fruits collected per individual or per species was too low to be tested.

Plant species Number of sampled Number of sampled Mean number of D. ± S.E Mean number of emerging individuals fruits suzukii eggs per fruit adults of D. suzukii per fruit Cotoneaster bullatus 1 10 2.80 0.05 0.00 Cotoneaster watereri 1 10 2.90 0.09 0.00 Duchesena indica 2 20 0.20 0.01 0.00 Fushia sp. 1 10 1.20 0.05 1.20 Iris sp. 1 10 0.00 0.00 0.00 Malus sylvestris 3 24 0.00 0.00 0.00 Rubia peregrina 1 10 0.00 0.00 0.00 Skimmia japonica 1 10 0.00 0.00 0.00 Solanum tuberosum 2 20 0.00 0.00 0.00 Vaccinium uliginosum 1 10 1.20 0.04 0.20



Table 1: Generalized linear models showing the effects of fruit traits on the number of D. suzukii eggs, larvae and adults emerging per 100 fruits. 1: parameter estimate; 2: standard error; 3: degrees of freedom.

1 2 3 Response variables Explanatory variables Category Par. est . β S.E. Wald X² D.F. P -value

Number of eggs Intercept 5.136 0.063 6602.11 1 <0.001

(n =67 species) Fruit diameter (log10) 2.181 0.058 1424.40 1 <0.001 Fruit type 1370.94 3 <0.001 Berry 1.399 0.039 1297.88 1 <0.001 Drupe 1.357 0.042 1033.24 1 <0.001 Polydrupe 0.816 0.053 233.30 1 <0.001 Other (complex structure) 0 - - - - Fruit color 2523.05 5 <0.001 Black 1.291 0.050 675.12 1 <0.001 Red 0.505 0.051 97.72 1 <0.001 Pink 0.354 0.060 35.10 1 <0.001 Orange-light brown -0.791 0.070 128.77 1 <0.001 White 1.503 0.056 718.09 1 <0.001 Blue 0 - - - - Fruit shape 475.66 1 <0.001 Spherical -0.572 0.026 475.66 1 <0.001 Oval 0 - - - - Fruit skin 1601.04 2 <0.001 Smooth, waxy, shiny -1.171 0.041 817.62 1 <0.001 Smooth, pruinose -2.174 0.054 1600.22 1 <0.001 rough 0 - - - -

Number of larvae Intercept 3.929 0.050 6170.05 1 <0.001

(n =60 species) Fruit diameter (log10) 3.746 0.105 1263.09 1 <0.001 Fruit type 633.96 3 <0.001 Berry 0.653 0.055 142.71 1 <0.001 Drupe 0.846 0.056 228.98 1 <0.001 Polydrupe 1.446 0.061 561.24 1 <0.001 Other (complex structure) 0 - - - -

Number of adults Intercept 2.996 0.059 2618.24 1 <0.001

(n =67 species) Fruit diameter (log10) 4.029 0.115 1226.80 1 <0.001 Fruit type 1369.83 3 <0.001 Berry 1.239 0.066 356.70 1 <0.001 Drupe 1.376 0.067 419.30 1 <0.001 Polydrupe 2.291 0.065 1236.05 1 <0.001 Other (complex structure) 0 - - - -



Figure 1: Mean number (± S.E.) of emerging adults, larvae and eggs of D. suzukii per fruit for the 15 plants species showing the highest number of D. suzukii eggs per fuit.

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Figure 3: Fruiting periods of studied plant species. Plant species are grouped into four categories according their suitability for the different development stages of Drosophila suzukii.

60

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   Agricultural and Forest Entomology (2014), DOI: 10.1111/afe.12052

Invasive host for invasive pest: when the Asiatic cherry fly (Drosophila suzukii) meets the American black cherry (Prunus serotina) in Europe

Mathilde Poyet∗† , Patrice Eslin∗, Marie He´ raude∗, Vincent Le Roux∗, Genevie` ve Pre´ vost∗, Patricia Gibert† and Olivier Chabrerie∗‡ ∗Unite´ Ecologie et Dynamiques des Syste`mes Anthropise´s – FRE 3498, CNRS – Universite´ de Picardie Jules Verne, 1 rue des Louvels, F-80037, Amiens, Cedex, France and † Laboratoire de Biome´trie et Biologie Evolutive – UMR 5558, CNRS – Universite´ Lyon 1, 43 boulevard du 11 novembre 1918, F-69622, Villeurbanne, France

Abstract 1 The vinegar fly Drosophila suzukii Matsumara (Diptera: Drosophilidae), native to Asia, recently invaded Europe and North America. By contrast to other frugivorous D rosophila species, D. suzukii lays eggs on ripening fruits, heavily reducing fruit production. Although cultivated host plants of D. suzukii are well documented, very little is known about wild hosts in the invaded areas. 2 The American black cherry Prunus serotina Ehrh., a tree species native to North America, became one of the main woody forest invaders in Europe. One cause of its invasion success is the huge amount of fruit produced by P. serotina trees. 3 A field survey showed that P. serotina is a suitable reservoir for the development and persistence of D. suzukii populations in European natural systems (on the forest area investigated, up to 70% of all the fruits of P. serotina were infested in one of the sampling sites). 4 Laboratory tests demonstrated that D. suzukii prefers ripening cherries to ripe ones, therefore increasing the chance of the larvae to fully develop and reach maturity before the mesocarp of the fruit totally decays. 5 Infestation of P. serotina cherries could reduce the life span of fruits, as well as their attractiveness to seed consumers and dispersers, yet P. serotina could represent a suitable plant reservoir promoting D. suzukii invasion in Europe and North America.

Keywords Drosophila suzukii, Prunus serotina, exotics, fleshy fruits, introduced species.

Introduction (i.e. the exocarp). Oviposition scars give pathogens (fungi, yeast and bacteria) access to the fleshy tissues of the fruit that Drosophila suzukii (Matsumura, 1931) (Fig. 1a,b) is a fruit will rot prematurely (Walsh et al., 2011). This ‘fruit predator’ fly that is native to Asia. It was recorded for the first time strategy does not occur in other European Drosophila species in North America and Southern Europe in 2008 (Calabria characterized by opportunistic egg laying on damaged and et al., 2010) and then spread rapidly across Europe (Cini et al., 2012). Unlike most other Drosophila species using over-ripe fermented fruit. Drosophila suzukii is thus considered as an and rotting fruit, D. suzukii lays eggs on healthy, ripening invasive pest in agricultural production systems (Beers et al., fruit: females possess a serrated ovipositor (Fig. 1c) (Hauser, 2011; Walsh et al., 2011) because it attacks fleshly fruits of 2011) allowing them to ‘drill’ holes through the fruit skin cultivated plants and drastically reduces fruit production. Cultivated host plants of D. suzukii are well documented in Correspondence: Olivier Chabrerie. Tel.: +33 (0)3 22 82 79 09; fax: the invaded area and include blueberry, strawberry, blackberry +33 (0)3 22 82 74 69; e-mail: [email protected] and raspberry (Lee et al., 2011; Bellamy et al., 2013). Among ‡ Present address: Ecologie et Dynamique des Syste`mes Anthropise´s the wide range of hosts, the Prunus genus appears to be a (EDYSAN, CNRS, FRE 3498), Universite´ de Picardie Jules Verne, taxonomic group that is particularly important to D. suzukii , 1 rue des Louvels, 80037 Amiens, Cedex 1, France. with a large number of species potentially favourable to the fly’s

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When D. suzukii meets P. serotina 2

(a) (b)

(c) (d)

(e) (f)

Figure 1 (a) Spotted-wing adult male Drosophila suzukii and (b) adult female Drosophila suzukii trapped in Compie` gne Forest (scale bar = 1 mm); (c) serrated ovipositor of Drosophila suzukii (scale bar = 300 μm); (d) Drosophila suzukii trap deployed in a Prunus serotina stand in Compie` gne Forest (scale bar = 10 cm); (e, f) infested cherries of Prunus serotina showing oviposition scars (white arrows) and egg filaments (scale bar = 600 μm). development both in native and invaded areas. For example, Deckers et al., 2005, 2008; Godefroid et al., 2005; Verheyen the cherry (Prunus avium ) is widely recognized as one of the et al., 2007; Chabrerie et al., 2008). Prunus serotina was most suitable host for D. suzukii (Kanzawa, 1935, 1939; Beers first introduced in Europe near Paris in the 17th Century et al., 2011), although other species such as Prunus domestica for ornamental purpose. It was planted throughout Western (plum), Prunus persica (peach) or Prunus armeniaca (apricot) Europe during the first half of the 20th Century, mainly in pine may also serve as suitable hosts (Cini et al., 2012; Bellamy plantations and on drained substrates for soil improvement and et al., 2013), and Prunus nipponica (Mitsui et al., 2010) or timber production (Starfinger, 1997; Starfinger et al., 2003). Prunus donarium (Mitsui et al., 2006; Kasuya et al., 2013) are One of the major causes of P. serotina invasion success two major resources for D. suzukii in Japan. is the huge amount of fruit produced in its exotic range, The American black cherry Prunus serotina Ehrh. is a gap- combined with a particular strategy called the ‘Oskar syndrome’ dependent tree species native to North America and, currently, (Closset-Kopp et al., 2007), which leads to the formation it is widely spread throughout temperate forests and hedgerow of a dense carpet of long-living saplings at the forest floor networks of Western and Central Europe (Starfinger, 1997; level. For example, a mean of 6011 drupes per tree was

k



reported in a French forest (Closset-Kopp et al., 2007), up anaesthesia. The identification and nomenclature of Drosophila to 10 253 drupes per tree were reported in a Belgian pine species followed that described by Markow and O’Grady plantation (Pairon et al., 2006) and up to 53 048 drupes (2006). per tree were counted in a Belgian agricultural landscape To investigate which factor(s) may condition the number of (Deckers et al., 2008). Currently, P. serotina has become one D. suzukii adults emerging from P. serotina fruits, we measured of the main woody forest invaders in Europe. This exotic for each of the 20 sampling sites: (i) the distance (m) between species is able to modify soil properties (Plichta et al., 1997; the P. serotina tree and the forest edge; (ii) the height (m) of the Chabrerie et al., 2008), the species composition of forest sampled tree; (iii) the total canopy cover (%); (iv) P. serotina communities (Chabrerie et al., 2007; Verheyen et al., 2007) density (number of individuals with a diameter at breast height and forest management practices (Muys & Lust, 1992) and > 5 cm; stems per hectare); and (v) P. serotina cover (%) in also reduces plant functional diversity (Chabrerie et al., 2010). shrub (1 – 6 m) and tree (> 6 m) layers in a 20 m × 20 m plot The development of P. serotina seedlings is constrained by centred on the sampled tree. The density of P. serotina fruits indigenous pathogens in its native range in North America fallen on the ground (fruits per m2 ) was also estimated using (Packer & Clay, 2000), although very few enemies have been five 25 cm × 25 cm quadrats randomly distributed under the reported in Europe, with the exception of the bird-cherry weevil canopy of each sampled P. serotina tree. (Furcipus rectirostris ), which has a low infestation rate (Pairon The number of Drosophila larvae developing in P. serotina et al., 2006). Prunus serotina thus benefits from the absence or fruits from the Compie`gne forest was also estimated: 30 fruits reduced efficiency of enemies in its exotic range, as do most per tree were randomly collected in the canopy of a subset of invasive species (Keane & Crawley, 2002). 10 P. serotina trees and 20 fruits per tree were collected on the In the present study, we investigated whether P. serotina soil under the canopy of a subset of 11 P. serotina trees. Each represents a suitable host for D. suzukii . A field survey was fruit was dissected under a Leica M 165C stereomicroscope conducted to evaluate the infestation rate of P. serotina fruits by and larvae were identified and counted. All D. suzukii larvae D. suzukii in natura and laboratory tests were run to determine were raised to be double checked after adult emergence. the susceptibility of the different ripeness stages of the fruits Drosophila suzukii populations captured in traps or emerging of P. serotina . from collected fruits were reared for one generation on a standard Drosophila medium (Chabert et al., 2012) for subsequent laboratory tests. Female adult flies reared in the Materials and methods laboratory were maintained between 5 and 7 days with males in mixed containers before the oviposition experiments. Field sampling The Compie`gne forest (49◦ 22 N, 2◦ 54 E; 32 – 148 m a.s.l., 14 417 ha, Picardie region, Northern France) is the forest most Laboratory tests on oviposition and development

invaded by P. serotina in France (Chabrerie et al., 2007). A Observations from field sampling strongly suggested that D. high density of P. serotina fruits is commonly observed on the suzukii oviposition behaviour may depend on fruit ripeness. forest floor in October, at the peak of the seed rain period for We also know that colour changes in Prunus fruits can be this tree species (Pairon et al., 2006). We chose to evaluate well pronounced during maturation. For example, P. donarium the infestation rate of P. serotina fruit by D. suzukii in the fruits turn from yellow to black before falling (Mitsui et al., Compie`gne forest (width 15 km, length 20 km) in the autumn 2006), whereas P. serotina fruits change from green (unripe) of 2011. Over this area, 25 traps were randomly distributed in to red (ripening) and finally to black (ripe) during maturation different stands separated by a minimum distance of 200 m. (Willson & Melampy, 1983). In the Compie`gne forest, it Banana traps are classical tools for sampling frugivorous takes 3 – 5 weeks for a red fruit to turn black. Therefore, for Drosophila in nature (Carson & Heed, 1983). Plastic traps these oviposition choice experiments, we adopted the fruit containing half a banana cut into 4 cm sections on sawdust colour categories given by Mitsui et al. (2006) to assess the were left for 8 days under the canopy of P. serotina trees attractiveness to D. suzukii of two ripeness stages of P. (Fig. 1d). Traps were then collected and brought back to the serotina fruits: entirely red (ripening) and entirely black (ripe) laboratory where fruit flies were identified and counted under fruits were collected on 20 P. serotina trees scattered in the CO2 anaesthesia using a stereomicroscope (Leica M 165C; Compie`gne Forest. Leica, Germany). Fruits were carefully examined with a stereomicroscope and The same P. serotina trees were used for both D. suzukii those already damaged or attacked by animals or pathogens trapping and fruit sampling (n = 20 trees; five trees that did were excluded. Healthy red and black fruits were then placed not produce enough fruits were excluded from fruit sampling). in separated plastic bags and conserved at 5 ◦ C. To test the For each sampled tree, 100 fruits were randomly collected attractiveness of fruits of P. serotina to D. suzukii flies, we in the canopy and 100 fruits were collected on the ground conducted no-choice and choice replicated tests. under the canopy. Almost all collected fruits were ripe (black), although some from the ground had started to rot. Fruit samples were weighed then kept in ventilated transparent plastic boxes No-choice test. Ten fruits of P. serotina were exposed to three (15 × 10 × 5 cm) at laboratory temperature (20 ◦ C). The number D. suzukii mated females for 24 h in cylindrical transparent of D. suzukii flies emerging in each box was checked daily; test tubes (diameter 2.5 cm, height 9.5 cm). In this experiment, flies were removed then identified and counted under CO2 we offered three separate substrates to D. suzukii females:



(i) 10 red fruits; (ii) 10 black fruits of P. serotina ; and (iii) with a Poisson distribution and a log-link term (McCullagh & ripe banana (for an equivalent volume of 10 P. serotina Nelder, 1989). The distribution of the values of the response fruits), which was used as a control because banana is a variable (number of Drosophila individuals) conformed to a common standardized substrate to trap and rear Drosophila Poisson distribution (Kolmogorov – Smirnov test: Z = 1.157, spp. A total of 20 replicates per treatment were maintained at P > 0.05) and the model residuals were normally distributed laboratory temperature (20 ◦ C). After 24 h, the number of eggs (Shapiro – Wilk test, W = 0.9408, P > 0.05). Only significant laid in each fruit was counted under a stereomicroscope. As variables (distance to forest edge and P. serotina fruits weight) reported by Mitsui et al. (2006), eggs laid in fruits are easily were retained in the final model. The number of D. suzukii detectable from the holes drilled by the females’ ovipositor adults emerging from fruits collected in the canopy and on associated with the presence of egg filaments (Fig. 1e,f). In this the soil was compared using paired t -tests. In the laboratory experiment, every hole that we observed contained one single tests, the numbers of eggs laid in the different categories of fruit D. suzukii egg. were compared using analysis of variance and Tukey’s honestly To monitor the ‘full development test’, fruits infested during significant difference comparisons of means (P < 0.05) for no- the no-choice test were maintained at room temperature until choice tests, paired t -tests for choice tests and t -tests for full fly emergence. The number of D. suzukii flies emerging from development tests. The number of D. suzukii adults emerging each set of 10 fruits of each category of black and red fruits from the different categories of fruit in full development was recorded. tests was compared using t -tests with spss, version 17.0 (SPSS Inc., Chicago, Illinois). Normality and homoscedasticity assumptions were tested before analyses using the Levene Choice test. In 45 replicates, one D. suzukii mated female was test and dagoTest-functions of the car and fBasics libraries, placed in a transparent glass cage (18 × 12 × 7.5 cm) containing respectively, in r, 2.11.1 (R Development Core Team, 2010). one red P. serotina fruit, one black P. serotina fruit and a banana substrate (of a volume equivalent to one P. serotina cherry). The three fruits were positioned at the corners of a Results virtual equilateral triangle (7 × 7 × 7 cm). After 3 h, the eggs Drosophila community deposited in each fruit were counted. A total of 196 flies from six Drosophila species were col- lected in the 25 traps (Table 1). The number of species varied Statistical analysis between 0 and 6 per trap, with a mean ± SD of 1.56 ± 1.90 species per trap. The Drosophila community was dominated The influence of environmental variables on the number of by Drosophila immigrans (41.8% of the total individuals) and D. suzukii adults emerging from fruit collected in the canopy of D. suzukii was the second most abundant species in the traps P. serotina trees was examined using generalized linear models (14.8%). The sex ratio (males/females) of D. suzukii popula-

tions in the traps was lower (sex ratio = 0.81) than the sex ratio Table 1 Drosophila species present in the 25 traps of D. suzukii populations emerging from fruits collected either in the canopy (sex ratio = 0.96; n = 709 D. suzukii individuals Relative Mean ± SD number in 2000 fruits) or under the canopy of P. serotina trees (sex n abundance (%) of individuals ratio = 0.89; n = 310 D. suzukii individuals in 2000 fruits).

Species Drosophila immigrans 82 41.8 3.28 ± 6.80 Drosophila suzukii 29 14.8 1.16 ± 2.43 Infestation rates in the fruits sampled in the field Drosophila subobscura 24 12.2 0.96 ± 2.19 Drosophila phalerata 14 7.1 0.56 ± 1.58 The weight of P. serotina drupes varied in the range Drosophila melanogaster 9 4.6 0.36 ± 0.57 0.155 – 0.615 g, with a mean ± SD of 0.405 ± 0.125 g per fruit Drosophila simulans 9 4.6 0.36 ± 0.81 over the 20 sampling sites (2000 fruits collected in the canopy Total 196 100.0 7.84 ± 14.93 of 20 trees; Table 2). The number of D. suzukii emerging from these fruits was 2.3-fold higher in the samples collected in the

Table 2 Drosophila adults emerged from Prunus serotina fruits

Drosophila adults for 100 fruits

Number of Number of Number of Relative sampled trees sampled fruits Drosophila abundance (%) Minimum Maximum Mean ±SD Fruits on the trees Drosophila suzukii 20 2000 709 100.0 0 72 35.5 ± 17.2 Fruits on the ground Drosophila suzukii 20 2000 310 76.5 6 37 15.5 ± 9.2 Drosophila subobscura 20 2000 95 23.5 0 19 4.8 ± 4.7



Table 3 Generalized linear model showing the effects of distance to forest edge (variable ‘DIST’) and Prunus serotina fruits weight (‘WEIGHT’) on the number of Drosophila suzukii adults (‘SUZUKII’) emerging from fruits collected in the canopy of Prunus serotina trees

Dependent variables Explanatory variables Parameter estimate SE d.f. χ 2 P SUZUKII Model constant: 3.008 0.144 1 435.16 < 0.001 DIST 0.010 0.002 1 21.36 < 0.001 WEIGHT 1.001 0.325 1 9.46 0.002

Table 4 Abundance of Drosophila suzukii larvae in the fruits

D. suzukii larvae for 100 fruits

Number of Number of Number of Number of sampled trees dissected fruits infested fruits Drosophila suzukii Minimum Maximim Mean ± SD Fruits collected in the canopy 10 300 124 127 13 70 42.3 ± 17.9 Fruits collected on the ground 11 220 49 49 5 45 22.3 ± 16.4

canopy (35.5 ± 17.2 adults per 100 fruits) than in the samples 80.0

from the ground (15.5 ± 9.2 adults per 100 fruits; t = 5.137; a P < 0.0001). Drosophila subobscura accounted for 25.5% of 70.0 the Drosophila adults emerging from the fruits collected on the ground but was not found in the fruits collected in the 60.0 canopy. None of the other four Drosophila species captured in the traps emerged from any P. serotina drupes. 50.0 More D. suzukii adults (emerging from fruits collected in 40.0 the canopy of P. serotina trees) (Table 3) were found in the b

interior of the forest than in the edge. The number of D. suzukii 30.0 adults emerging from fruits also increased with the weight of

P. serotina fruits (Table 3). S.D.) number of eggs per 10 fruits 20.0 The dissections of P. serotina fruits collected in the canopies ± showed that a mean of 42.3% was infested by D. suzukii larvae, 10.0 c with percentages varying in the range 13.3 – 70% depending on ( Mean the collecting site (Table 4). The mean number of D. suzukii 0.0 larvae was 1.9-fold higher in the fruits collected in the canopy Red fruits Black fruits Banana (42.3 ± 17.9 adults per 100 fruits) than in the fruits from the Figure 2 Mean ± SD number of eggs laid by Drosophila suzukii in ground (22.3 ± 16.4 adults per 100 fruits). Among the 124 fruits no-choice t each test, 10 Prunus serotina fruits of each type collected in the canopy and infested by D. suzukii larvae, 122 ests. In fruits (98.4%) hosted one single larva, whereas only one fruit (ripe/ripening) and an equal volume of banana were used. Different hosted two larvae and another fruit hosted three. The 49 fruits letters indicate a significant difference. collected on the ground and infested by D. suzukii hosted only one larva each. All of the fruits collected from the ground and the tree canopy were ripe (black) fruits and some of the fruits 4.7-fold more eggs in red fruits than in black ones in the no- collected on the ground were starting to rot. choice and choice tests, respectively. In the full development test, the number of D. suzukii adults emerging from the fruits (between 9 and 13 days after oviposition) was significantly higher in red cherries (4.78 ± 0.38 adults per fruit) than in black Choice and no-choice tests ones (2.68 ± 0.13 adults per fruit; t = 11.59; P < 0.001).

In the no-choice test, the number of eggs laid was signifi- cantly higher in red cherries (5.35 ± 1.64 eggs per fruit) than in black ones (2.46 ± 1.17 eggs per fruit) or banana substrate Discussion

(F = 95.276; P < 0.001) (Fig. 2). In the choice test, D. suzukii Drosophila suzukii infests ripening, red fruits of the invasive females preferred to lay eggs in P. serotina red fruits rather than P. serotina in black ones (t = 6.586; P < 0.001) (Fig. 3). No egg was ever observed on the banana substrate in the choice tests. The num- The results from our field sampling indicate that P. serotina is ber of eggs laid per red fruit of P. serotina was similar between a suitable host for the cherry vinegar fly D. suzukii in European no-choice (5.35 ± 1.64 eggs per fruit) and choice (4.62 ± 4.00 temperate forests: in the Compie`gne forest, where the invasive eggs per fruit) tests. Drosophila suzukii females laid 2.2- and P. serotina presents a large population (Chabrerie et al., 2007), P. serotina fruits can show high rates of infestation by



10.0 The preference of D. suzukii for ripening fruits is presently

poorly documented. This preference probably increases chances 9.0 a that its progeny will reach maturity before total degradation of 8.0 the fruit mesocarp. The D. suzukii preference for fresh fruits on trees (as opposed to over-ripe and fallen fruits) has been 7.0 observed in cultivated plants (Walsh et al., 2011). However, 6.0 unlike the results of the present study, the number of D. suzukii

5.0 eggs oviposited per cherry was reported to be higher on black fruits than on red or light red ones in its native range (for 4.0 P. donarium , see Mitsui et al., 2006). In the case of P. serotina b 3.0 cherry, D. suzukii oviposits in the very early stages of fruit

2.0 generations during fruit maturation. Similar densities of eggs 1.0 (approximately five eggs per fruit) were laid in red cherries in Mean (± S.D.) number of eggs per 1 fruit 0.0 oviposition behaviour of D. suzukii . Any threshold in the Red fruits Black fruits Banana number of eggs oviposited in each fruit could contribute to limit Figure 3 Mean ± SD number of eggs laid by Drosophila suzukii in choice intraspecific competition between larvae feeding on limited tests. In each test, one Prunus serotina fruit of each type (ripe/ripening) resources (Atkinson, 1979). A great variety of mechanisms has and an equal volume of banana were used. Different letters indicate a been proposed to explain host selection by D. suzukii (Lee significant difference. et al., 2011; Bellamy et al., 2013; Burrack et al., 2013). For example, the volatile compounds or the pH of ripening fruits D. suzukii (up to 70% of infested fruits in one of the sampling of P. serotina may be more attractive than those of ripe ones. sites). Regarding the firmness of P. serotina ripening fruits, it is The mean rate of infestation was higher when estimated at worth noting that this is not a barrier to D. suzukii oviposition, the larval stage (42.3 larvae from 100 dissected fruits) than whereas the fly is mainly known as a pest of soft-skinned fruits when based on rearing out adult flies (35.5 adults per 100 with a low surface penetration force (Burrack et al., 2013). The fruits). This appears to indicate that a high mortality (16.1%) role of colour in host selection remains questionable because may occur during D. suzukii development in fruits. The results the fly is able to infest plant species with a wide range of fruit of the present study showed a positive effect of the fruit colours and its colour preference is different in its native range weight on the number of D. suzukii emerging flies and we also (Mitsui et al., 2006).

observed several eggs laid per fruit during our laboratory tests. However, 98.4% of the sampled fruits hosted one larva only, suggesting that larval competition may occur when several The ecological niche of D. suzukii and its threat larvae grow in a same fruit. Each P. serotina fruit may thus on cultivated plants play the role of an individual development cell for D. suzukii Drosophila suzukii was the only species emerging from the larvae after competitive exclusion of the congeners. Therefore, fruits collected in P. serotina canopy. This could mean that the dynamics of the fly population from the Compie`gne forest infesting red fruits hanging on the tree is an efficient strategy for could be constrained by the amount of resources in P. serotina escaping from competition with other fruit flies. By developing fruits and by the size of the P. serotina host population. on fresh fruits, D. suzukii will certainly reduce the overlap of The mean diameter of P. serotina fruits (8 mm in Compie`gne its temporal niche with other flies. Drosophila subobscura flies forest) is close to the one of P. donarium fruits (5 mm in emerged from P. serotina fruits collected on the ground but Tokyo), a Japanese host commonly infested by D. suzukii never from fruits from the canopy, which is coherent with (Mitsui et al., 2006). With P. serotina fruits, D. suzukii larvae this hypothesis. In addition, it is possible that D. subobscura may have met hosts of a suitable size providing resources close (and perhaps undetected other species), which feeds mainly on to those of their native range. rotting substrates, could benefit from the activity of D. suzukii . Both (choice and no-choice) oviposition tests conducted in As observed in cultivated fruits (Cini et al., 2012), D. suzukii the laboratory show that D. suzukii prefers to lay eggs in red, females damage fresh fruits by leaving scars after oviposition, ripening P. serotina drupes rather than in ripe fruits. In a choice thus leading to premature rotting. situation, females can lay over four-fold more eggs in ripening Only 23 D. suzukii flies were counted out of 25 traps, fruits than in ripe ones. This finding is reinforced by the whereas 709 individuals emerged from fruits sampled on observation that, in the Compie`gne forest, D. suzukii larvae the same sites. This clearly indicates that the banana-trap mainly develop in the fresh fruits hanging on trees. More than commonly used to collect fruit flies is not attractive to two-fold more D. suzukii flies emerged from fruits collected in D. suzukii , as confirmed by the experiments conducted in the the tree canopy than from fruits sampled on the ground. This laboratory where no egg was ever observed on banana substrate also indicates that D. suzukii is successfully developing on P. in choice tests. Using banana-traps also could have contributed serotina red fruits, as confirmed by the development experiment to the low sex ratio (i.e. a lower proportion of males) of the conducted in the laboratory. trapped populations. Drosophila suzukii females driven by their



oviposition need may be more likely than males to investigate a in natura and in controlled environments, are needed to new substrate, even though it is less attractive than baits. This disentangle the fly’s aptitude from the habitat characteristics emphasizes the need to develop new collecting features and facilitating D. suzukii invasion. A next step will be to more species-specific traps to enable a proper evaluation of the investigate the attractiveness and susceptibility of fruits of other dispersion of D. suzukii populations. Work on trap improvement fleshy-fruited plants to D. suzukii with the aim of evaluating is still ongoing (Landolt et al., 2012; Lee et al., 2012; Cha their possible contribution to its successful invasion. et al., 2013).

During the fruiting period of P. serotina in the Compie`gne forest, the distance from the collecting site to the forest Possible effects of D. suzukii on the invasive P. serotina edge may be an environmental factor increasing the number of D. suzukii flies emerging from the fruits of the invasive Two important consequences of D. suzukii occurrence on the tree. This result is consistent with those reported by Martins invader P. serotina can be predicted, or at least expected. (2001), showing that the Drosophilid fruit-fly guild is strongly First, by damaging ripening cherries, D. suzukii larvae may structured by the edge effect in forest habitat and that many cause premature fall of fruits containing immature seeds. This Drosophila species increase in abundance from the forest edge premature abscission of infested fruits has long been known in to the forest interior. Edge may affect organisms in a forest cultivated systems (Levine & Hall, 1977) and the fruits from a fragment by causing changes in the biotic and abiotic conditions wide range of plant species damaged by D. suzukii drop or split (Murcia, 1995). Thus, the difference of fly density between (Lee et al., 2011). This mechanism may reduce the viability edge and forest interior may be a response to changes in of P. serotina fruits. Second, D. suzukii may also change the habitat quality and resources (Martins, 2001), such as changes dispersal efficiency of P. serotina cherries by affecting the fruit in microclimate conditions, light, disturbance frequency, fruit attractiveness to dispersers (Manzur & Courtney, 1984). availability or plant species composition (Murcia, 1995; Foggo Indeed, birds prefer to eat P. serotina fruits that have not been et al., 2001; Chabrerie et al., 2013). Therefore, the forest attacked by insects (Pairon et al., 2006). Being rejected by interior appears suitable for the development of D. suzukii birds, numerous seeds may fall under the canopy of mother populations. The ecological niche of D. suzukii in its exotic trees, increasing the competition between P. serotina offspring range is still poorly known. However, the species has been and their parents.

trapped in European forested areas (forested sites neighbouring crops in Le Centre Technique Interprofessionnel des Fruits et Le´gumes, C. Weydert, personal communication) and we Conclusion know that it is associated with tree populations in Japan (Mitsui et al., 2006, 2010). Prunus serotina could thus become In conclusion, the present study is the first report on D. a plant reservoir for the development and the persistence of suzukii infestation behaviour on fruits of a European forest tree, D. suzukii populations in European natural ecosystems (the P. serotina , which itself is an invasive species in European presence of D. suzukii was recorded a second then a third temperate forest. Our results demonstrate that the invasive year, in 2012 and 2013 during our survey campaigns; P. Eslin P. serotina may be an important plant reservoir for D. suzukii and O. Chabrerie, unpublished data). Such a reservoir would populations in the European forests. This potential refuge for certainly be a potential source of contamination for fields of D. suzukii may be a source of contamination and a threat cultivated fruits surrounding forests and hedgerows already for cultivated fruits surrounding European temperate forests invaded by P. serotina . For example, we estimated that, in invaded by P. serotina . On the other hand, D. suzukii may the Compie`gne forest (5704 hectares invaded by P. serotina play a role in affecting the dispersion of P. serotina lacking trees) where the mean density of P. serotina is 687 stems/ha natural enemies in its exotic range. These results on P. serotina (Chabrerie et al., 2007, 2008) with a mean fruit production hosting D. suzukii in European forests are also of interest to of 6011 drupes per tree (Closset-Kopp et al., 2007), and with the North American continent because P. serotina is native to each drupe potentially giving rise to the development of a this area and D. suzukii has now been detected throughout the mean of 0.355 D. suzukii (35.5 adults per 100 fruits) (Table 2), western and the eastern U.S.A. and in Quebec, and populations the whole P. serotina population living in this forest could have been observed for several years in many locations (Bolda release a mean of 8.36 × 109 flies per year. This potential et al., 2010; Walsh et al., 2011; Burrack, et al., 2013; Saguez value may vary between years as a result of changes in climate et al., 2013).

conditions and fruit production. We suggest that, in regions of temperate forests, fruits sampled from P. serotina trees are used as indicators to reveal the presence and the risk of dispersion Acknowledgements of this invasive cherry fly. The number of eggs per P. serotina cherry recorded in our We thank the French Office National des Foreˆts for providing tests is clearly higher than that reported by Mitsui et al. (2006) assistance during the field work. in a Japanese wooded area. Increased investment in offspring production between native and exotic ranges is one of the main References characteristics of invader species (Blossey & No¨ tzold, 1995). Further investigations comparing the oviposition behaviours Atkinson, W.D. (1979) A field investigation of larval competition in between native and exotic D. suzukii populations, both domestic Drosophila . Journal of Animal Ecology , 48, 91 – 102.



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