Olfactory attractants constitute the foundation of monitoring and control strategies for D. suzukii. However, this section focuses exclusively on the odors emitted by fruits at different developmental stages—ripening, mature, and decaying—and by the microorganisms associated with them. The underlying premise is that flies seek host fruits at various stages for feeding, mating, and oviposition (Tochen et al., 2016; Cloonan et al., 2018; Young et al., 2018; Clymans et al., 2019; Galland et al., 2020; Piñero et al., 2022; Elsensohn and Burrack, 2023; Kim et al., 2023). Traditional attractants, including apple cider vinegar (ACV), rice vinegar, wine vinegar, acetoin, octanoate, acetic acid, ethanol, and isoamyl acetate compounds alone or in mixtures, have long been used as reference or control (Cloonan et al., 2018; Renkema et al., 2018; Lasa et al., 2020; Cruz-Esteban et al., 2021a, 2024a, 2024b). However, their efficacy varies depending on the composition and ratio of volatile compounds (Abraham et al., 2015; Cha et al., 2015; Mazzetto et al., 2015, 2016a; Akasaka et al., 2017; Cloonan et al., 2018; Feng et al., 2018; Jaffe et al., 2018; Kirkpatrick et al., 2018a; Lasa et al., 2019; Willbrand and Pfeiffer, 2019; Lasa et al., 2020; Toledo-Hernández et al., 2021a, 2021b; Urbaneja-Bernat et al., 2021; Keene-Snickers et al., 2025). In Mexico, several commercial products are available that are based on the four most attractive compounds—acetoin, methionol, acetic acid, and ethanol—previously reported by Cha et al. (2014, 2015, 2017, 2018). One such product is Z-Kinol^® (Squid Biological and Pheromones S.A. de C.V., Texcoco, Mexico) (Cruz-Esteban et al., 2021a), which uses water with 5% soap as the retention solution. Another commercially available product manufactured outside Mexico is Pherocon^® SWD (Trécé Inc., Adair, OK, USA) (Burrack et al., 2015; Frewin et al., 2017; Tonina et al., 2018), as well as Scentry^® (Scentry Biologicals Inc., USA), which uses ACV as the retention solution (Kirkpatrick et al., 2017; Lethmayer and Egartner, 2017; Briem et al., 2018; Whitener et al., 2022). These four volatile organic compounds (VOCs), identified as highly attractive and incorporated into the formulation of these commercial attractants, have also been detected and confirmed in homemade fermentations evaluated in Mexico (Cruz-Esteban et al., 2024b; Alavez-Rosas et al., 2024). Other food-based attractants, such as SuzukiiTrap^® (Bioibérica, Barcelona, Spain) (Tonina et al., 2018; Cruz-Esteban et al., 2021a), Fruit Fly Attractant (Koppert), and SuzukiiLURE-Max (Dinusa, Oaxaca, Mexico), also employ these components as retention solutions and claim to be effective for monitoring D. suzukii. However, these products should be evaluated under local conditions, as not all of them were specifically developed for Mexican populations. Furthermore, some products are described as scientifically developed, yet the supporting references demonstrating their efficacy are often not available, possibly due to geographical variation in D. suzukii populations (Jaffe et al., 2018; Wollmann et al., 2019; Larson et al., 2021; Whitener et al., 2022). Another important factor is their relatively high cost, which highlights the need to identify and develop formulations that are more effective, affordable, long-lasting, and environmentally friendly—safe for pollinators and, above all, for farmers—and that can be readily adapted to local populations.

When discussing fermented products, it is essential to consider another group of VOCs produced and emitted by microorganisms, such as bacteria, fungi, and yeasts, that are associated with fruits (Hamby and Becher, 2016). These organisms participate directly or indirectly in the fermentation process. Among symbiotic bacteria, the most common species found in microbial communities associated with Drosophila suzukii are Tatumella spp. (Enterobacteriaceae) (Chandler et al., 2014), Gluconobacter spp (Chandler et al., 2014; Mazzetto et al., 2016a; Bueno et al., 2020), Komagataeibacter spp (Mazzetto et al., 2016a), and Acetobacter spp. (Acetobacteraceae) (Chandler et al., 2014; Mazzetto et al., 2016a; Bueno et al., 2020). Other bacteria, such as Oenococcus oeni, commonly involved in malolactic fermentation during wine production (Alawamleh, 2021; Ðurović et al., 2021; Maddalena, 2016, 2021), and Lactobacillus kunkeei and other lactic acid bacteria strains (Alawamleh, 2021; Alawamleh et al., 2021), have also been associated with D. suzukii. Yeasts are particularly important because they play a key role in the attraction of D. suzukii (Barata et al., 2012; Scheidler et al., 2015). The most studied species include Saccharomyces cerevisiae, a yeast widely used in baking (Batista et al., 2017; Lasa et al., 2017; Spitaler et al., 2020); Saccharomycopsis vini (Spitaler et al., 2020; Castellan et al., 2024); Hanseniaspora uvarum (Scheidler et al., 2015; Batista et al., 2017; Mori et al., 2017; Noble et al., 2019; Bianchi et al., 2020; Bueno et al., 2020; Spitaler et al., 2020; Huang and Gut, 2021; Kleman et al., 2022; Rehermann et al., 2022; Castellan et al., 2024); H. opuntiae (Bueno et al., 2020); Issatchenkia terricola; Metschnikowia; Metschnikowia pulcherrima (Spitaler et al., 2020; Jones et al., 2021); Candida zemplinina (Scheidler et al., 2015; Spitaler et al., 2020; Jones et al., 2021); Clavispora santaluciae; Saccharomycopsis vini; Issatchenkia terricola; Metschnikowia pulcherrima (Castellan et al., 2024), and Pichia pijperi (Jones et al., 2021), a yeast frequently found in the alimentary canals of flies and in larval excreta (Hamby et al., 2012; Solomon et al., 2019; Bellutti et al., 2018). These species are highly attractive to D. suzukii, enhance fecundity, and often exhibit positive synergistic effects when combined (Jones et al., 2025), as well as with commercial or homemade attractants, and with bioinsecticides or conventional insecticides (Jones et al., 2022; Barone and Hartbauer, 2024; Duménil et al., 2025; Molokwu et al., 2025). Gas chromatography–electroantennographic detection (GC-EAD) analyses have identified isobutyl acetate and isoamyl acetate emitted by H. uvarum as key volatiles perceived by D. suzukii antennae (Scheidler et al., 2015). Isoamyl acetate is also found in several attractive fruits, including strawberries and blueberries (Dekker et al., 2015). Studies have shown that mated females are more strongly attracted to H. uvarum and fruit odors but may reduce oviposition when feeding on yeast, suggesting a trade-off between feeding and reproduction (Mori et al., 2017). Yeast-associated volatiles can also enhance oviposition in other fruits, such as cherries (Bellutti et al., 2018), although their effects appear to depend on the background fruit odors (Cloonan et al., 2019). Furthermore, acetic acid bacteria such as Gluconobacter spp. emit volatiles including ethanol, acetic acid, and aldehydes, which also elicit antennal responses in D. suzukii (Mazzetto et al., 2016a). More recently, Castellan et al. (2024) and Duménil et al. (2025) reported that in electrophysiological assays, antennae of D. suzukii responded to ethyl acetate, propanoate, isoamyl acetate, β-myrcene, benzaldehyde, and linalool, compounds that may underlie the strong attractiveness of S. vini and H. uvarum to this species.

Another important group of VOCs that influence the attraction of D. suzukii are produced and emitted by fruits, particularly across different developmental stages. For instance, it has been reported that virgin or protein-deficient females, as well as males, are attracted to fermented food sources, where females can feed and wait for egg maturation. These same sites also serve as mating grounds, where a possible pheromone and numerous co-occurring compounds are likely involved (Revadi et al., 2015; Tochen et al., 2016; Clymans et al., 2019; Khan et al., 2019). Once mated, females are guided by VOCs emitted from ripening or fully ripe fruits in good condition to locate suitable oviposition sites and ensure offspring survival (Tochen et al., 2016; Cloonan et al., 2018; Young et al., 2018; Clymans et al., 2019). However, this behavior in gravid females has also been shown to vary with seasonal conditions (Clymans et al., 2019). Collectively, these findings indicate that host-search behaviour in D. suzukii is not static but changes throughout the female reproductive cycle. The behavioural ecology of host searching has therefore been conceptualized as a sequential process in which female responses shift according to their physiological state (Cloonan et al., 2018). Newly emerged virgin females primarily orient toward nutritional resources that maximize survival and reproductive maturation, particularly fermenting fruits and surrounding foliage. These substrates emit long-range volatile cues associated with microbial fermentation, including ethanol, acetic acid, acetoin, methionol, and other yeast-derived compounds, while surrounding vegetation may contribute additional cues such as β-cyclocitral. Besides functioning as feeding sites, these substrates also likely serve as mating arenas because recently emerged males aggregate in similar resource patches. Although several studies have suggested the possible involvement of intraspecific chemical signals, including cis-vaccenyl acetate and contact cuticular hydrocarbons, no major long-range sex pheromone has yet been conclusively demonstrated for D. suzukii (Bartelt et al., 1985; Lebreton et al., 2017; Cloonan et al., 2018; Kwadha, 2022; Lima et al., 2023). Following mating, gravid females exhibit a marked behavioural shift toward intact ripening fruits suitable for oviposition. At this stage, females appear to integrate multiple fruit- and microorganism-derived semiochemicals rather than responding to individual compounds alone. Attractive volatiles identified from host fruits include E-2-hexenal, 3-methyl-1-butanol, ethyl acetate, isoamyl acetate, butyl acetate, methyl 2-methylbutyrate, and ethyl propionate, together with several fermentation-derived compounds emitted by yeasts and associated bacteria. Interestingly, isoamyl acetate occurs in both fruit- and microorganism-derived odour sources, suggesting that some behaviourally active compounds originate from multiple biological substrates. Once females approach the fruit, short-range visual and tactile cues—including fruit colour, reflectance, firmness, surface integrity, and contrast with the surrounding vegetation—probably contribute to the final acceptance of oviposition sites. Other compounds, such as hexanol, 3-methyl-2-butenyl acetate, 3-methyl-2-butanone, 2-heptanone, isobutyl acetate, and additional fermentation volatiles, have also been proposed as candidate cues involved in host location or oviposition. However, current evidence suggests that these compounds should be regarded as context-dependent secondary cues whose behavioural relevance varies with concentration, blend composition, female physiological condition, and environmental context, rather than as universally attractive semiochemicals. Several studies have demonstrated host preferences for economically important fruits, classifying them according to attraction levels observed in laboratory bioassays. For example, Abraham et al. (2015) found that D. suzukii exhibited the strongest preference for raspberry, followed by strawberry, blueberry, and cherry—consistent with the findings of Cai et al. (2019). These results align with field observations by Cruz-Esteban et al. (2021a), who reported higher fly captures in raspberry crops, followed by blackberry, strawberry, and blueberry. However, when cultivated alone, any of these fruits can be highly susceptible to D. suzukii infestation (Little et al., 2017; Cruz-Esteban et al., 2021a). Analyses of VOCs from these mature fruits, both intact (healthy) and mechanically damaged, revealed that alcohols, aldehydes, and ketones are the predominant compounds (Abraham et al., 2015; Revadi et al., 2015; Cloonan et al., 2018). Gas chromatography–electroantennographic detection (GC-EAD) analyses have shown that D. suzukii antennae respond to several volatiles from raspberry and strawberry, including esters, alcohols, and ketones. An eleven-compound blend identified from raspberry was attractive to both sexes, although less so than the natural fruit extract (Abraham et al., 2015; Keesey et al., 2015). The volatile profile of apples consists mainly of esters, alcohols, and ketones, among which acetoin and octanoate have been identified as the most attractive compounds to D. suzukii (Feng et al., 2018). Similarly, four volatile compounds from bayberry—methyl (E)-3-hexenoate, methyl (E)-2-hexenoate, ethyl (E)-2-hexenoate, and α-humulene, in a ratio of 1:1.3:1:6.4—were reported to be attractive under field conditions (Liu et al., 2018). Approximately 33 compounds have been identified from blackberry, with acetaldehyde, hexyl acetate, linalool, and myrtenol being the most attractive (Dewitte et al., 2021). Overall, VOCs emitted by host fruits are highly attractive to this invasive fly species, as confirmed by recent studies comparing natural fruit odors with various commercial attractants (Babu et al., 2022). Therefore, ongoing research efforts remain focused on identifying and developing volatile blends that are more attractive under field conditions.

In Mexico, Cruz-Esteban et al. (2021a) demonstrated that the type of attractant, rather than the trap design, significantly influences the capture efficiency of D. suzukii in berry crops. In their study, two commercial attractants—Z-Kinol^® and SuzukiiTrap^®—as well as apple cider vinegar (ACV) and a previously reported highly attractive blend (ACV + EtOH + CO₂) (Lasa et al., 2017) were compared under a factorial design that included raspberry, blackberry, strawberry, and blueberry crops. Significant differences in capture efficiency were observed among attractants but not between trap designs, highlighting the importance of selecting appropriate baits for effective monitoring and population control. Recent studies conducted outside Mexico support these findings. Larson et al. (2021) evaluated various attractant blends for early detection and monitoring, identifying specific compounds such as acetoin that enhance selectivity and detection sensitivity. The recent work of Cruz-Esteban et al. (2024b) further advanced this field by testing fermented baits formulated from blueberries, raspberries, baker’s yeast, and sugar. In field trials, these baits showed superior attractiveness compared to commercial ACV or homemade vinegar blends, resulting in higher capture rates of D. suzukii adults in both blueberry and blackberry crops. Notably, these fermentative blends also demonstrated consistent performance across multiple harvests, high attractiveness to Zaprionus indianus (Gupta) (Diptera: Drosophilidae)—particularly during seasons when both pests overlap—and low attraction of non-target drosophilids, thereby improving selectivity while maintaining cost-effectiveness and local producibility. Similar findings were reported in Michigan, where a mixture of wine, vinegar, and yeast proved more attractive in the field than each component alone (Huang et al., 2017). Likewise, in the United States, laboratory assays revealed that fruit VOCs became more attractive when combined with β-cyclocitral or a blend of β-cyclocitral, isoamyl acetate, and methyl butyrate (Bolton et al., 2019; Piñero et al., 2019; Bolton et al., 2022). Despite their advantages, fermentative attractants present several challenges, including variable duration of efficacy, sensitivity to environmental conditions, and the potential to attract non-target species. In contrast, synthetic baits with controlled release offer longer persistence, greater selectivity when properly formulated, and reduced maintenance requirements, although they are generally more expensive. Another critical aspect is the capture threshold, as trap detections often occur when populations have already reached levels capable of causing economic damage. For instance, monitoring in Michigan estimated that capturing a single individual in a trap corresponded to approximately 192 flies within a 2.7 ha area, underscoring the need for prompt action upon first detection (Larson et al., 2021).

Results regarding the use of visual cues (i.e., attractive colors) and trap designs obtained from laboratory and field studies have been quite contradictory. For instance, Rice et al. (2016) reported that black and red spheres captured significantly more D. suzukii than purple, blue, yellow, or white spheres in laboratory assays, and semi-field raspberry trials. Similarly, several studies have concluded that red and black traps are among the most effective for attracting and capturing this invasive fly (Basoalto et al., 2013; Lee et al., 2013; Renkema et al., 2014; Kirkpatrick et al., 2016, 2017; Lasa et al., 2017; Rice et al., 2017). The effectiveness of these visual cues is further enhanced when they act in synergy with food-based attractants (Bolton et al., 2021; Lasa et al., 2024). In contrast, Cahenzli et al. (2018) found that transparent traps fitted with a black lid captured more D. suzukii than traps equipped with a red lid. Similarly, other studies have shown that transparent and yellow traps perform comparably to red and black traps (Lee et al., 2013; Iglesias et al., 2014; Briem et al., 2018; Marjanović and Tanasković, 2019). Overall, these differences were primarily numerical and often not statistically significant. In addition, background color contrast has been shown to influence D. suzukii attraction (Antignus, 2000; Little et al., 2019). For example, under laboratory conditions, Little et al. (2019) found that black, purple, red, or yellow objects presented against a green background did not differ significantly in attractiveness. However, purple objects against a green background were more attractive than red objects against a black background, black objects against a red background, or blue objects against a yellow background. Similar context-dependent effects of visual cues have been reported by Barone and Hartbauer (2024), who demonstrated that Petri dishes covered with red mesh but containing yellow gels attracted more D. suzukii than transparent controls, highlighting the importance of the visual properties of the stimulus itself rather than the external background alone. However, recent studies conducted in Mexico have demonstrated that capture success is not determined by the external color of the trap, but rather by chemical communication and short-range visual stimuli—such as colored cards and reflective surfaces placed inside the trap—which significantly increase the likelihood of fly entry (Cruz-Esteban et al., 2021a, 2021b; Cruz-Esteban, 2021; Figure 1). For example, comparative evaluations of attractants and trap designs showed that chemical cues, particularly fermentation-related compounds, are the primary drivers of D. suzukii attraction. No significant differences among trap designs were detected when all were baited with the same chemical stimulus, indicating that capture efficiency was largely independent of trap architecture under these conditions (Cruz-Esteban et al., 2021a; Figure 1a). However, the potential contribution of visual cues at short range could not be excluded. Subsequent experiments incorporating previously untested colors and transparent traps with internal yellow or violet cards—colors reported as attractive in laboratory assays (Little et al., 2019)—revealed a strong effect of internal visual stimuli. Transparent traps equipped with an internal yellow card captured approximately 400% more D. suzukii than other designs (Cruz-Esteban, 2021; Figure 1b). Further tests using single colors and color combinations inside transparent traps, all baited with the same attractant, confirmed this pattern: yellow cards, alone or combined with green, consistently produced the highest captures (≈350% increase) relative to other colors (Cruz-Esteban et al., 2021b; Figure 2). More recently, variation in yellow reflectance was shown to influence trap performance, with a yellow card exhibiting 67% reflectance and a dominant wavelength of 549.74 nm outperforming higher-reflectance yellow cards in apple cider vinegar–baited traps (Cruz-Esteban et al., 2024a, Figure 2).

Therefore, advances in trap design now emphasize optimizing internal visual stimuli rather than focusing solely on external color (Little et al., 2018, 2019, 2020b; Cruz-Esteban, 2021; Cruz-Esteban et al., 2024a, Figure 2). These findings underscore the importance of integrating visual cues into trap design as complementary components of ethological control strategies, ensuring that adult flies are efficiently guided into traps even when olfactory plumes have a limited range (Figure 2). Overall, visual stimuli represent a critical factor in the performance of monitoring and mass-trapping programs. Optimizing trap color, internal reflectance, and brightness—when combined with high-quality attractants—can maximize capture efficiency and contribute to the sustainable management of D. suzukii in commercial berry systems. When tailored to local crop conditions and fly behavior, these strategies enhance early detection, population suppression, and reduction of crop damage. Likewise, olfactory attractants remain indispensable for both monitoring and population suppression, particularly when integrated with visual stimuli, mass-trapping strategies, and oviposition deterrents. Recent advances in bait formulation, including fermentation-based blends developed from locally available ingredients (e.g., Cruz-Esteban et al., 2024b), further demonstrate that effective and economically accessible attractants can be incorporated into integrated management programs for commercial berry production. Collectively, these studies have substantially advanced our understanding of the chemical ecology of D. suzukii, providing a solid foundation for future research aimed at elucidating host-search behavior, oviposition site selection, and the multimodal integration of olfactory, visual, and tactile cues under field conditions.

Mass trapping is a key strategy for the suppression of D. suzukii populations, going beyond its traditional role in monitoring. This approach relies on deploying attractant-baited traps to capture adults, thereby reducing reproductive output and limiting dispersal. The effectiveness of mass trapping is strongly influenced by trap density and spacing, as closely spaced traps can experience overlapping attraction radii, leading to intra-trap competition and potential saturation of the target area (Clymans et al., 2022). Based on this study, recommended trap densities range from 75 to 200 traps ha^-¹ in spring and 90 to 300 traps ha^-¹ in summer, depending on bait type and environmental conditions. Controlled-release synthetic baits, which maintain attractiveness over longer periods, can allow reductions to approximately 25 traps ha^-¹ without compromising population suppression (Clymans et al., 2022). Trap spacing can range between 3 and 5.5 m (Hampton et al., 2014; Kirkpatrick et al., 2018a; Spies and Liburd, 2019; Clymans et al., 2022).

Field trials in cherry orchards have further demonstrated the utility of mass trapping. The Decis™ Trap, which combines a persistent attractant with an insecticide-treated interior, was highly effective at reducing D. suzukii populations when deployed at 100 traps ha^-¹, particularly when placed early in the growing season, resulting in significant reductions in fruit damage (De Maeyer et al., 2018). These findings highlight that the timing of trap placement is as critical as trap density in maximizing suppression and preventing early-season infestations.

Mass-trapping strategies have been successfully applied in berry crops. For example, in Tiripetío, municipality of Morelia, Michoacán, Mexico (19°31′55″ N, 101°22′10″ W), transparent multi-hole traps equipped with yellow cards (67% reflectance at 549.74 nm) and baited with a fermented solution containing blueberries, raspberries, baker’s yeast, and sugar have been deployed during 2023–2024 in blueberry and blackberry crops (Cruz-Esteban et al., 2024b; Figure 2). Traps were installed 6 m apart, totaling 289 traps ha^-¹. This approach proved highly effective, maintaining infestation levels below 5% and enabling all harvest lots to meet export standards. It has been observed that when fruits in ripening, mature, and overripe stages are abundant, fly captures decrease, as previously reported by other authors (Harris et al., 2014; Burrack et al., 2015; Joshi et al., 2017). However, we have found a way to stimulate fly activity so that they leave the host fruit and are attracted to and captured by the traps. Applications of garlic extract (an insect repellent, 1 kg/100 L H₂O) have been made every 15 days in the center of each crop row, raising the sprayer half a meter above the ground (Allium, AGROARSA, Mexico). For monitoring purposes, random samples of approximately 100 fruits per hectare per crop are collected, placed in a Ziploc bag, and gently macerated by hand to avoid damaging the larvae. The fruits are then submerged in a sugar-water solution (180–200 g/L) to facilitate larval detection by flotation (SENASICA (Servicio Nacional de Sanidad, Inocuidad y Calidad Agroalimentaria), 2014; Shaw et al., 2019; Babu et al., 2023). This information is complemented with a direct physical inspection of each fruit using a magnifying glass or a stereomicroscope (Van Timmeren et al., 2021).

Several critical factors influence the success of mass-trapping programs. Optimizing trap spacing is essential to avoid intra-trap competition caused by overlapping attraction radii. Bait selection also plays a major role, as controlled-release synthetic baits provide longer-lasting attraction and greater selectivity compared to traditional fermentative baits such as apple cider vinegar. Additionally, environmental factors—including temperature, crop phenology, and pest population density—strongly affect trap performance and must be considered when planning deployment. The integration of mass trapping with visual stimuli, oviposition deterrents, and complementary biological control can further enhance population suppression, highlighting the importance of multi-tactic approaches for sustainable management of D. suzukii in commercial berry production.

Push–pull strategies and oviposition deterrents are designed to reduce egg laying on host crops while simultaneously attracting adults to traps or non-crop buffer zones (Figure 3). Recent studies have demonstrated the potential of both chemical and biological signals to manipulate D. suzukii behavior under field conditions. For instance, the presence of live Drosophila melanogaster (Meigen) (Diptera: Drosophilidae) larvae has been shown to reduce oviposition by D. suzukii on artificial substrates, with the live larvae acting as deterrent cues, whereas eggs and adults of D. melanogaster did not elicit the same response (Tungadi et al., 2022). The mechanism is not fully explained by cuticular hydrocarbon cues, suggesting the involvement of volatile signals, microbial interactions, or physical cues. Chemical deterrents have also been evaluated as key components of push–pull systems. Roh et al. (2023) demonstrated that a two-component deterrent blend—ethyl anthranilate and methyl salicylate—applied to raspberry fruits significantly reduced D. suzukii infestation in field trials, confirming its practical potential. Other studies have identified volatile organic compounds (VOCs) such as ethyl anthranilate, methyl salicylate, green leaf volatiles, and monoterpenes as oviposition deterrents that signal unsuitable or previously occupied substrates (Burrack et al., 2015; Iglesias and Liburd, 2017; Tonina et al., 2020). These compounds can be applied via coatings, sprays, or controlled-release dispensers, offering versatility in field applications. Particle films, including kaolin, diatomaceous earth, and mica, have also been tested as oviposition deterrents, particularly in vineyards and berry crops. These films act as physical barriers, reducing ovipositor penetration and altering surface properties such as reflectance, which further discourages egg-laying (Rossi-Stacconi et al., 2016; Tonina et al., 2020). Field evaluations in Austria and other regions demonstrated substantial reductions in oviposition, especially when applied during early fruit developmental stages. Essential oils such as geranium, ginger, peppermint (menthol), thyme (thymol), clove (eugenol), citronella (citronellol, geraniol), lavender, 2-pentylfuran and neem, among others, have shown repellent, oviposition-deterring, and even lethal effects against D. suzukii (Renkema et al., 2016, 2017; Gowton, 2019; Galland et al., 2020; Wernicke et al., 2020; Wang Q, et al., 2021; Baleba et al., 2025; Shrestha et al., 2025). Moreover, mixtures of these oils are commercially available under the names KeyPlex Ecotrol^® PLUS (rosemary, geraniol, and mint oils) and KeyPlex Sporan^® EC2 (rosemary, clove, thyme, and mint oils), which have been evaluated and shown promising results in raspberry and blueberry crops in Forest Lake, MN, USA (Gullickson et al., 2020). Compounds including 1-octen-3-ol (Wallingford et al., 2016, 2017), 2-n-pentylfuran (Cha et al., 2021; Stockton et al., 2021), thymol (Reher et al., 2019), p-menthane-3,8-diol (Wernicke et al., 2020), ethyl butanoate, and ethyl (E)-but-2-enoate (Quadrel et al., 2025), appear to be promising repellents and oviposition deterrents for future field evaluation. One of the promising plants as a repellent, although currently available only commercially, is hop (Humulus lupulus L.), which also deserves consideration in future studies (Reher et al., 2019). In contrast, geosmin has shown inconsistent results (Stockton et al., 2021). These active ingredients have been formulated in a slow-release SPLAT^® matrix (Specialized Pheromone & Lure Application Technology, ISCA Technologies, Inc.), which does not require water or adjuvants. The SPLAT matrix allows for controlled release of labile semiochemicals (Stelinski et al., 2005) and has been used in formulations for mating disruption, attract-and-kill, and repellency systems targeting both pest insects and pollinators (e.g., Vargas et al., 2008; Rodriguez-Saona et al., 2010; Mafra-Neto et al., 2013). Another formulation, HOOK SWD^®, a sprayable attract-and-eliminate (A&E) bait, was evaluated over three growing seasons in raspberry and other small fruit crops in New Jersey and California. Residual activity trials showed adult mortality rates exceeding 78–93% after 35 days of field aging under plastic tunnels (Klick et al., 2019). Under laboratory conditions, lavender oil, Ecotrol^®, and Sporan^® deterred SWD from diet substrates, while field trials indicated that Ecotrol^® reduced infestation in raspberries, though not in blueberries (Gullickson et al., 2020). Recently, a two-component deterrent blend consisting of octanoic and decanoic acids was developed as an alternative to spatial repellents for the behavioral control of D. suzukii infestations (Movva et al., 2025). Other promising natural extracts include attract-and-eliminate formulations derived from mandarin (Citrus reticulata) and tea tree (Melaleuca alternifolia) oils, though fruits treated with the latter were classified as non-edible (Bedini et al., 2020). Integration of push–pull systems with mass trapping and attractant-baited devices enhances overall suppression of D. suzukii. By redirecting adult flies away from crops while simultaneously removing them from the population, these tactics reduce both current and future infestations. However, effective deployment requires careful consideration of compound selection, timing, and spatial configuration to maximize deterrent efficacy without reducing trap attractiveness. Future research should focus on identifying synergistic effects among chemical, visual, and biological deterrents to optimize multi-tactic management programs, ensuring sustainable and practical implementation in commercial berry production. For example, in the United States, methyl benzoate was tested as an in-crop repellent in blueberries, while attractant-baited traps were deployed along the field perimeters. However, these compounds alone showed limited synergistic efficiency (Gale et al., 2024). Conversely, mass trapping using RIGA^® AG traps (Zürich, Switzerland) baited with 80 mL of RIGA^® AG lure (apple cider vinegar, red wine, sugar, and cranberry juice) (Figure 2), combined with perimeter insecticide applications, proved effective for D. suzukii control in blueberry fields (Spies and Liburd, 2019; Kehrli et al., 2022).

Although ethological strategies primarily aim to manipulate pest behavior, the integration of complementary biological control has become an essential component for the sustainable management of D. suzukii, given the compatibility of both approaches (Daane et al., 2016; Becher et al., 2017; Koźbiał and Lewandowski, 2025).

A recent systematic review found that biological-control research for D. suzukii is heavily dominated by parasitoids (64% of publications) compared to entomopathogens (26%) and predators (7%) (Morales-Abeijon et al., 2025). Within parasitoids, the families Figitidae (46%; n = 21), Braconidae (33%; n = 15) and Pteromalidae (13%; n = 6) were most frequently studied (Morales-Abeijon et al., 2025). Among the ten most investigated species are the pupal parasitoids Trichopria drosophilae (Perkins) (Kremmer et al., 2017; Rossi-Stacconi et al., 2018, 2019; Gonzalez-Cabrera et al., 2019; Trivellone et al., 2020; Wolf et al., 2020; Collatz and Romeis, 2021; Wang X, et al., 2021; Daane et al., 2025; Morales-Abeijon et al., 2025; Sun et al., 2025), Trichopria anastrephae (Lima) (Hymenoptera: Diapriidae) (Krüger et al., 2019; Vieira et al., 2020; da Costa Oliveira et al., 2021; Wang X, et al., 2021; Krüger et al., 2024), Pachycrepoideus vindemmiae (Rondani) (Kremmer et al., 2017; Collatz and Romeis, 2021; da Costa Oliveira et al., 2021; Wang X, et al., 2021; Daane et al., 2025; Morales-Abeijon et al., 2025), and Spalangia erythromera (Förster) (Hymenoptera: Pteromalidae) (Trivellone et al., 2020), along with the larval parasitoids Ganaspis kimorum Buffington formerly also known as Ganaspis brasiliensis (Ilhering) (Daane et al., 2016; Girod et al., 2018; Stahl et al., 2024; Morales-Abeijon et al., 2025), Leptopilina japonica (Girod et al., 2018; Morales-Abeijon et al., 2025; Rossi-Stacconi et al., 2025), L. japonica japonica, L. j. formosana (Novković and Kimura), L. boulardi (Barbotin) (Hymenoptera: Figitidae) (Daane et al., 2016; Mazzetto et al., 2016b; Daane et al., 2025), L. heterotoma (Thomson) (Hymenoptera: Eucoilidae) (Beckwith et al., 2025; Daane et al., 2025; Schöfer et al., 2025), Carton and Keiner-Pillault (Hymenoptera: Figitidae) (Huang et al., 2023), Asobara japonica (Belokobylskij) (Girod et al., 2018), A. rufescens (Förster), A. brevicauda (Guerrieri & van Achterberg), and A. leveri (Nixon) (Hymenoptera: Braconidae) (Daane et al., 2016; Mazzetto et al., 2016b). These species have demonstrated significant parasitism potential under laboratory and field conditions, particularly G. brasiliensis and L. japonica, which have established naturally in several regions of Europe and North America with parasitism rates reaching up to 30% (Girod et al., 2018; Sanchez-Gonzalez et al., 2020; Morales-Abeijon et al., 2025).

Beyond parasitoids, several other biological control agents (BCAs) have been identified targeting D. suzukii, including 23 bacterial species, viruses spanning eight families, nine entomopathogenic nematodes and three major entomopathogenic fungi—Beauveria bassiana Bals. (Vuill.), Metarhizium anisopliae (Metschn.) Sorokin and Isaria fumosorosea Wize (Alnajjar et al., 2017; Hiebert et al., 2020; Bing et al., 2021; Galland et al., 2023; Morales-Abeijon et al., 2025). Among bacterial agents, Xenorhabdus nematophila (Poinar and Thomas), Bacillus thuringiensis Berliner (Bt) and Brevibacillus laterosporus (Laubach) have shown promising pathogenicity, while the La Jolla virus (Iflaviridae) was one of the most frequently reported viral agents (Linscheid et al., 2022; Bruner-Montero et al., 2023; Garriga et al., 2023; Morales-Abeijon et al., 2025). Although these findings demonstrate a rich potential of microbial and entomopathogenic resources for D. suzukii control, field efficacy remains variable, often depending on environmental conditions, formulation stability, and the developmental stage of the pest (Sial et al., 2019; Morales-Abeijon et al., 2025). Field evaluations indicate that B. bassiana and M. anisopliae achieve moderate reductions of adult populations when applied in shaded, humid environments, while B. thuringiensis and B. laterosporus are more effective against larvae inside fruits or substrates (Cuthbertson and Audsley, 2016; Gutierrez-Palomares et al., 2021; Mastore et al., 2021; Morales-Abeijon et al., 2025). Combining these agents with attractant-based systems—such as bait stations or attract-and-kill traps—enhances exposure and infection rates, thereby increasing overall pest suppression. Furthermore, integrating microbial control with semiochemical-based strategies (e.g., pheromone traps, kairomone lures, or oviposition deterrents) enables more targeted control while reducing reliance on synthetic insecticides (Wallingford et al., 2017; Ðurović, 2021; Morales-Abeijon et al., 2025). Such integrated approaches not only improve management efficiency but also promote the long-term sustainability of berry production systems by preserving beneficial organisms and minimizing ecological risks.

In addition to parasitoids and pathogens, predatory arthropods contribute to the natural regulation of D. suzukii populations. Fifteen predatory species across eight families have been reported (Morales-Abeijon et al., 2025). The families Anthocoridae (Hemiptera) and Carabidae (Coleoptera) are the most represented, each with four species. Among them, the rove beetle Dalotia coriaria (Kraatz) (Coleoptera: Staphylinidae) has been the most extensively studied due to its capacity to prey upon both D. suzukii eggs and larvae in the soil or fruit substrate. Other notable predators include the pirate bug Orius insidiosus (Say) (Hemiptera: Anthocoridae), the green lacewing Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae), and the European earwig Forficula auricularia L. (Dermaptera: Forficulidae), which exhibit opportunistic predation on eggs and early larval stages (Renkema et al., 2015; Englert and Herz, 2019; Morales-Abeijon et al., 2025). Less frequently studied but potentially valuable species include the ground beetles Bembidion quadrimaculatum (L.), Limodromus assimilis (Paykull), Poecilus cupreus (L.), and Pterostichus melanarius (Illiger) (Coleoptera: Carabidae); the true bugs Dicyphus hesperus Knight, Macrolophus pygmaeus (Rambur), and Nesidiocoris tenuis (Reuter) (Hemiptera: Miridae); Orius laevigatus (Fieber) and O. majusculus (Reuter) (Hemiptera: Anthocoridae); as well as the predatory stink bug Podisus maculiventris (Say) (Hemiptera: Pentatomidae) and the field cricket Gryllus pennsylvanicus Burmeister (Orthoptera: Gryllidae). Although these species have shown low specificity toward D. suzukii, their presence in fruit ecosystems suggests a relevant role within broader functional guilds of natural enemies, contributing indirectly to pest suppression through intraguild interactions and disturbance of oviposition behavior (Ballman et al., 2017; Bonneau et al., 2019; Siffert et al., 2021; Wang X et al., 2021; Morales-Abeijon et al., 2025).

Overall, the synergistic integration of parasitoids, entomopathogens, and generalist predators—combined with behavioral manipulation and habitat management—represents one of the most promising directions for ecologically sustainable suppression of D. suzukii. Future research should focus on optimizing the spatiotemporal compatibility among these natural enemies, developing formulations for microclimatic stability, and designing multi-trophic attract-and-kill systems that maximize contact and infection rates in the field.

In Mexico, pupal parasitoids such as Pachycrepoideus vindemmiae and Trichopria drosophilae have been evaluated, with T. drosophilae showing a higher intrinsic rate of increase and a shorter development time, making it a promising agent for natural control, particularly during pupal stages (Garcia-Cancino et al., 2020). Field and semi-field studies in Europe and North America have demonstrated that these pupal parasitoids can significantly reduce adult emergence from infested fruits, especially when integrated with habitat management practices that provide refuges or alternative hosts for parasitoid persistence (Wang et al., 2016; Girod et al., 2018; Renkema et al., 2020; Rehermann et al., 2022; Hogg and Daane, 2025).

The integration of biological control with mass trapping and push–pull systems can enhance suppression by reducing adult emergence while simultaneously disrupting host-seeking and oviposition through the use of attractants, visual cues, and deterrent elements. Furthermore, the combination of parasitoid releases with selective, low-risk insecticides has shown potential to control D. suzukii effectively while preserving beneficial arthropod populations (Hamby et al., 2016; Van Timmeren et al., 2025). However, the field efficacy of parasitoids is strongly influenced by environmental conditions, crop structure, and synchronization between parasitoid release and pest population dynamics. Interactions with other drosophilid species, such as Z. indianus, may also affect host preference and parasitism success. Consequently, optimizing release strategies, habitat modifications, and compatibility with other tactics remains a priority for improving biological control outcomes within integrated pest management frameworks.

To fully exploit the synergistic potential of behavioral and biological control, further field-scale evaluations under diverse climatic and cropping systems are needed. Future research should focus on refining parasitoid deployment timing, enhancing formulation stability, and assessing multitrophic interactions among parasitoids, entomopathogens, and semiochemical-based tools. Such integration will strengthen eco-efficient management strategies for D. suzukii, ensuring sustainable protection of commercial berry crops.