Effects of nonnutritive sugar inclusion in laboratory diets and attracticidal spheres on survivorship and mobility of 2 Dipteran species, Rhagoletis pomonella (Diptera: Tephritidae) and Drosophila suzukii (Diptera: Drosophilidae).

Native apple maggot fly, Rhagoletis pomonella, and invasive spotted-wing drosophila, Drosophila suzukii, are key pests of apple and small fruit, respectively, in the United States. Both species are typically managed with standard insecticide applications. However, interest in alternative strategies that result in insecticide reductions has led to evaluations of nonnutritive sugars as toxicants for Drosophila species and development of attracticidal spheres for both species. Here, we evaluated the survivorship of R. pomonella and D. suzukii when provided with standard diets that substituted saccharin, sucralose, aspartame, erythritol, dextrose, or mannitol for the sucrose component and compared them with standard diets and water-only controls for up to 15 days. Presence of erythritol and mannitol significantly decreased survivorship of R. pomonella and erythritol significantly decreased the survivorship of D. suzukii. However, mobility trials following a 2 h exposure to aqueous solutions of each sugar treatment resulted in no strong impact on either species. Survivorship after 30 min exposure to erythritol or mannitol alone, or in combination with varying concentrations of sucrose (serving as a phagostimulant) at 30 min and 24 h were evaluated for both species. Only D. suzukii survivorship was affected with decreased survivorship on erythritol:sucrose solutions of 20:0% and 15:5% for 24 h. Based on all results, erythritol appeared most promising, and was integrated into attracticidal spheres as a toxicant but even at the highest concentration, survivorship remained unaffected for either species, thus making this nonnutritive sugar impractical and ineffective as a toxicant substitute in attracticidal spheres.

Development and survivorship of Lycorma delicatula (Hemiptera: Fulgoridae) on cultivated and native Vitis spp. (Vitales: Vitaceae) of the Eastern United States.

As Lycorma delicatula (White) continues to spread across the United States, more winegrapes are potentially susceptible to damage from this pest. Lycorma delicatula, spotted lanternfly, is primarily associated with Ailanthus altissima (Mill.) Swingle, a tree from its native range that is now globally distributed. While L. delicatula is a known pest of cultivated Vitis spp. in South Korea, its relationship with the specific grape species grown in the United States is unclear. This study assessed L. delicatula survivorship and development on 5 Vitis species, including 2 winegrape V. vinifera L. varieties, 'Pinot Noir' and 'Chardonnay', Concord grape, Vitis labrusca L., River grape, Vitis riparia Michx., and muscadine grape, Vitis rotundifolia Michx. var. 'Carlos'. A diet of A. altissima served as a positive control. Lycorma delicatula provided with a diet of V. riparia or V. vinifera 'Pinot Noir' yielded the highest survivorship and fastest rates of development among grape diets and were statistically equivalent to those provided with A. altissima. Vitis rotundifolia did not support L. delicatula growth past the third-instar life stage, indicating this species is a poor host for the early development of this pest. Our results indicate that both V. riparia and V. vinifera are favorable hosts for L. delicatula and may provide the means for this insect to invade and establish in new regions.

Do adult Magicicada (Hemiptera: Cicadidae) feed? Historical perspectives and evidence from molecular gut content analysis.

The periodical cicadas in the genus Magicicada are remarkable for their unusual life histories and dramatic synchronized emergences every 13 or 17 years. While aspects of their evolution, mating behaviors, and general biology have been well-characterized, there is surprising uncertainty surrounding the feeding habits of the short-lived adult stage. Despite a tentative scientific consensus to the contrary, the perception that adult Magicicada do not feed has persisted among the general public, and recent studies are lacking. We directly investigated the feeding behavior of Magicicada spp. through high-throughput sequencing (HTS)-based dietary analysis of nymphs, freshly molted (teneral) adults, and fully sclerotized adults collected from orchard and wooded habitats during the 2021 emergence of Brood X. Identifiable plant DNA (trnF, ITS amplicons) was successfully recovered from nymphs and adults. No plant DNA was recovered from teneral adults, suggesting that all DNA recovered from sclerotized adults was ingested during the post-teneral adult stage. Both nymphs and adults were found to have ingested a range of woody and herbaceous plants across 17 genera and 14 families. Significantly more plant genera per individual were recovered from adults than from nymphs, likely reflecting the greater mobility of the adult stage. We hypothesize that the demonstrated ingestion of plant sap by Magicicada adults is driven by a need to replace lost water and support specialized bacteriome-dwelling endosymbionts that cicadas depend upon for growth and development, which constitutes true feeding behavior.

Short- and long-term effects of season-long infestation of Lycorma delicatula (Hemiptera: Fulgoridae) on young apple (Malus domestica) and peach (Prunus persica) trees.

Lycorma delicatula White, commonly called spotted lanternfly, is an invasive fulgorid that was first detected in the United States in Pennsylvania in 2014 and has spread across Eastern states. Lycorma delicatula is a phloem feeder, and feeding of persistent populations can cause wilt and dieback in host plants. Vineyards in infested regions have experienced vine loss, where damage has been attributed to L. delicatula. Nymphal and adult lifestages have been observed on cultivated apple and peach trees, but the feeding effects of this pest on these crops have not been quantified. Here, we caged young potted apple and peach trees and infested them with 50 nymphal or 25 adult L. delicatula per plant for 3 months and tracked plant health. Horticultural measurements including trunk diameter, height, leaf drop, and photosynthesis rates were recorded before, during, and after the infestation. To evaluate the long-term effects of infestation, all apple and peach trees were removed from pots and planted in an orchard block where their phenology, growth, and physiology could be evaluated the following growing season. Short- and long-term measurements showed no significant differences in apple tree health between infested and noninfested trees. There was a significant reduction in the growth of trunk diameter in peach trees during exposure to early-instar nymphs; infested peach trees also had significantly increased frost damage to buds the following spring. However, there were no long-term impacts on peach growth and physiology. These results indicate that L. delicatula likely will not be a major threat to these deciduous fruit crops.

Assessing acceptability of wild and cultivated hosts of Lycorma delicatula (Hemiptera: Fulgoridae) under semifield conditions.

Lycorma delicatula White (Hemiptera: Fulgoridae) is an invasive phloem feeder with a broad host range that includes both hardwood trees and cultivated temperate fruit crops. Here, we evaluated acceptability of wild hosts, A. altissima and Juglans nigra L. (Fagales: Juglandaceae) and cultivated hosts Vitis vinifera, Malus domestica (Rosales: Rosaceae), and Prunus persica L. Batsch (Rosales: Rosaceae) to L. delicatula under field conditions. Fluorescent-marked early instar nymphs, late instar nymphs, or adult L. delicatula were released at the base of single potted host plants and the number of individuals retained was recorded over 24 h. Paired choice trials with A. altissima and another host plant were conducted. Individuals retained on or moving between plants were recorded over 24 h. Sentinel A. altissima, J. nigra, V. vinifera, M. domestica, and P. persica potted plants were deployed at 5 sites and the number of L. delicatula present on each plant was recorded weekly. In single and paired host trials, early instars and adults were generally retained in higher numbers on A. altissima and V. vinifera, and late instars were retained on A. altissima and J. nigra. Significantly more L. delicatula were present on sentinel A. altissima compared with other host plants, except J. nigra during the period when late instars were the most prevalent lifestage in the field. These results indicate that wild hosts such as A. altissima and J. nigra are likely supporting establishment of L. delicatula populations, and that presence of cultivated V. vinifera may contribute to population establishment and growth.

Evaluating deployment strategies for spotted lanternfly (Lycorma delicatula Hemiptera: Fulgoridae) traps.

The spotted lanternfly, Lycorma delicatula (White) (Hemiptera: Fulgoridae), is an invasive planthopper that was first detected in the United States in Berks County, PA, in 2014, and has since spread to 13 states in the Eastern United States. This phloem-feeding pest has a broad host range, including economically important crops such as grapevine, Vitis spp. Monitoring presence and relative abundance of L. delicatula is essential to develop pest management tools. Here, we compared deployment strategies to optimize use of L. delicatula monitoring traps. Standard circle traps, sticky bands, and circle traps with replaceable bag tops were deployed at sites with either high or low populations present. Trap deployment at different heights and on different host tree species and trap sampling intervals were evaluated for standard circle traps only. Circle traps captured significantly more L. delicatula adults at low-density sites compared with other trap types in 2021, and no differences were detected at high-density sights. Traps deployed 1 m from the ground captured significantly more adults than those deployed at 0.5 m; no differences were detected for nymphs. While no significant differences in captures were found among intervals, weekly or biweekly sampling prevented specimen degradation. Although traps deployed on Ailanthus altissima (Mill.) Swingle (Sapindales: Simaroubaceae) captured significantly or numerically more L. delicatula at most sites, traps deployed on other hosts also yielded consistent captures. We were also able to alter the construction of circle trap skirts to allow for deployment on different sized tree trunks.

Survival and development of Lycorma delicatula (Hemiptera: Fulgoridae) on common secondary host plants differ by life stage under controlled conditions.

Host range assessment for emerging invasive insects is a vital step toward fully defining the issues the insect may pose. Spotted lanternfly (SLF) is an invasive species that is rapidly expanding its presence in the United States. The primary hosts facilitating this spread are tree of heaven, a plant from SLF's native range, and the economically important winegrape. Black walnut is also implicated as an important and common host plant. This study investigated the survival and development of SLF on diets that included a variety of crop host plants in the presence or absence of tree of heaven. The following plant species, 'Honeycrisp' apple, 'Reliance' peach, silver maple, and tree of heaven were paired with winegrape or black walnut throughout the study. SLF had strong development and high survival on a diet of winegrape alone, and winegrape or black walnut paired with tree of heaven. Survival parameters were reduced with all other plant pairings. In particular, SLF in the winegrape and peach diet treatment did not develop past the third nymphal instar. A second experiment evaluated the survival of early and late instar nymphs and adult SLF life stages on three specialty crops - 'Cascade' hops, muscadine grapes, and kiwifruit over a two-week period. Nymphs survived longer than adults, with survival of first and second instar nymphs on hops not differing from the control tree of heaven treatment. The adult stage survived best on kiwi and muscadine grape. Our results show tree of heaven and winegrape were the only single plant diets evaluated that are sufficient for complete SLF development, while other host plants may require additional host or hosts of sufficient nutritional quality for SLF survival.

Seasonal Distribution of Halyomorpha halys (Hemiptera: Pentatomidae) Captures in Woods-to-Orchard Pheromone Trap Transects in Virginia.

The invasive brown marmorated stink bug, Halyomorpha halys (Stål), is a pest of numerous economically important crops in the USA. In the Mid-Atlantic region, it is a significant, direct pest in tree fruit orchards, many of which are bordered by woodlots containing a variety of its deciduous tree and shrub hosts. During the growing season, H. halys moves from woodland habitats into crops, but seasonal changes in its relative abundance between these adjacent habitats have not been examined. Using linear transects of six pheromone-baited pyramid traps that extended from 100 m into the interior of woodlots to 100 m into the interior of adjacent commercial apple orchards in Virginia, spatiotemporal changes in H. halys captures were measured during three growing seasons. Captures of H. halys adults and nymphs were recorded weekly from May through October, and annual data were separated into early, mid, and late-season captures. Only adults were captured during the early season, and there was no indication of a spatial trend in captures across traps in the transects among years. Beginning in mid-season and becoming increasingly apparent by late season, captures of H. halys adults and nymphs tended to become most frequent in traps at the woods and orchard edges and at 50 m into the orchard interior. These findings conform with and expand upon previous research documenting an edge effect for H. halys relative abundance and can inform and support the optimization of perimeter-based management strategies for H. halys in Mid-Atlantic apple orchards.

Survivorship and Development of the Invasive Lycorma delicatula (Hemiptera: Fulgoridae) on Wild and Cultivated Temperate Host Plants.

The invasive spotted lanternfly, Lycorma delicatula, (White Hemiptera: Fulgoridae) continues to spread throughout the Eastern United States. This species exhibits a broad host range, with tree of heaven, Ailanthus altissima (Mill.) Swingle, commonly referred to as the preferred host. Here, we evaluated 2-wk survivorship of early nymphal instars, late nymphal instars, and adult L. delicatula on single diets of ten wild and cultivated hosts: tree of heaven; apple, Malus domestica; peach, Prunus persica; black cherry, P. serotina Ehrh; black locust, Robinia pseudoacacia L.; black walnut, Juglans nigra L.; common hackberry Celtis occidentalis L.; mulberry Morus alba L.; sugar maple Acer saccharum Marshall; white oak, Quercus alba L.. Among them, early and late instars had significantly greater survivorship on tree of heaven and black walnut and adults had greatest survivorship on tree of heaven. Additionally, we evaluated development and survivorship of L. delicatula from newly hatched nymphs to adulthood on single diets of tree of heaven, black walnut, grapevine, apple, and peach, and mixed diets of tree of heaven plus one other host. Single host diets that supported L. delicatula development to adulthood were tree of heaven and black walnut. Interestingly, mixed diets also supported development, and reduced development time to adults by up to 12% compared with the single tree of heaven diet. Our results suggest that within agroecosystems and across landscapes, L. delicatula can develop on single hosts such as tree of heaven, but also on multiple host plants, yielding adults earlier in the growing season.

Development of rearing methodology for the invasive Spotted Lanternfly, Lycorma delicatula (Hemiptera: Fulgoridae).

Lycorma delicatula, White (Hemiptera: Fulgoridae), spotted lanternfly, is a univoltine, phloem-feeding, polyphagous and invasive insect in the USA. Although a primary host for this species is Ailanthus altissima, tree of heaven, L. delicatula also feeds on a wide range of hosts important to the USA including cultivated grapevines. Due to the need for classical or augmentative biological control programs to reduce impacts of L. delicatula across invaded areas, we developed a laboratory-based rearing protocol for this invasive species. Here, we evaluated the use of A. altissima apical meristems, epicormic shoots, and fresh foliage cut from A. altissima as a food source for rearing newly hatched L. delicatula. On these sources of plant material <20% of L. delicatula developed into adults and no oviposition occurred. However, when young, potted A. altissima trees were used as a food source, >50% of L. delicatula nymphs developed to the adult stage under natural daylengths and temperatures ranging from 20-25°C. The addition of wild grapevine, Vitis riparia, did not increase survivorship or reduce development time. To elicit mating and oviposition, adults were provided with A. altissima logs as an oviposition substrate and maintained under shortened daylengths and reduced nighttime temperatures (12L:12D and 24°C:13°C). This resulted in 2.12 egg masses deposited per female, which was 4× more than when adults were maintained in standard rearing conditions (16L:8D and 25°C). Based on these experiments, we present a protocol for reliably rearing L. delicatula under laboratory and/or greenhouse conditions.

Refining Pheromone Lures for the Invasive Halyomorpha halys (Hemiptera: Pentatomidae) Through Collaborative Trials in the United States and Europe.

Brown marmorated stink bug, Halyomorpha halys, is native to Asia and has invaded North America and Europe inflicting serious agricultural damage to specialty and row crops. Tools to monitor the spread of H. halys include traps baited with the two-component aggregation pheromone (PHER), (3S,6S,7R,10S)-10,11-epoxy-1-bisabolen-3-ol and (3R,6S,7R,10S)-10,11-epoxy-1-bisabolen-3-ol, and pheromone synergist, methyl (2E,4E,6Z)-decatrienoate (MDT). Here, an international team of researchers conducted trials aimed at evaluating prototype commercial lures for H. halys to establish relative attractiveness of: 1) low and high loading rates of PHER and MDT for monitoring tools and attract and kill tactics; 2) polyethylene lure delivery substrates; and 3) the inclusion of ethyl (2E,4E,6Z)-decatrieonate (EDT), a compound that enhances captures when combined with PHER in lures. In general, PHER loading rate had a greater impact on overall trap captures compared with loading of MDT, but reductions in PHER loading and accompanying lower trap captures could be offset by increasing loading of MDT. As MDT is less expensive to produce, these findings enable reduced production costs. Traps baited with lures containing PHER and EDT resulted in numerically increased captures when EDT was loaded at a high rate, but captures were not significantly greater than those traps baited with lures containing standard PHER and MDT. Experimental polyethylene vial dispensers did not outperform standard lure dispensers; trap captures were significantly lower in most cases. Ultimately, these results will enable refinement of commercially available lures for H. halys to balance attraction and sensitivity with production cost.

Virulence of Entomopathogenic Fungi to Rhagoletis pomonella (Diptera: Tephritidae) and Interactions With Entomopathogenic Nematodes.

The objectives of this study were to quantify the virulence of four entomopathogenic fungal species to pupae of Rhagoletis pomonella (Walsh) (Diptera: Tephritidae) and to determine the potential to combine entomopathogenic fungi (EPFs) and entomopathogenic nematodes (EPNs) for biological control of this pest. The four species of EPFs included Beauveria bassiana (strain GHA), Metarhizium brunneum (strain F52), Isaria javanica (wf GA17), and Isaria fumosorosea (Apopka 97 strain). In laboratory assays, all fungi reduced adult emergence but there were no differences between fungal species. Isaria javanica and M. brunneum were examined further in a EPFs and EPNs bioassay that also included the EPNs Steinernema carpocapsae (ALL strain) and S. riobrave (355 strain). All nematodes and fungi were applied either alone or in combination (fungus + nematode). There were no differences between species within the same entomopathogen group (fungi and nematodes). However, the treatment with S. riobrave resulted in lower R. pomonella emergence than either fungal species. The combination of S. riobrave and I. javanica resulted in the lowest R. pomonella emergence (3%) at fourth-week interval, which was significantly lower than any of the single-agent applications, yet virulence of the other three combination treatments was not different from their respective nematode treatments applied alone. Additive interactions were detected for all fungus-nematode combinations. This study suggests that application of entomopathogenic nematodes and fungi could be an effective option to suppress R. pomonella populations.

Development of Behaviorally Based Monitoring and Biosurveillance Tools for the Invasive Spotted Lanternfly (Hemiptera: Fulgoridae).

The spotted lanternfly, Lycorma delicatula White, is an invasive planthopper (Hemiptera: Fulgoridae) that was first detected in the United States in Berks County, PA, in 2014, and has since spread in the mid-Atlantic region. This phloem-feeding pest has a broad host range, including economically important crops such as grape where their feeding causes dieback of infested plants. Monitoring the presence and abundance of L. delicatula is of utmost importance to develop pest management approaches. Current monitoring practices include sticky bands deployed on tree trunks, sometimes paired with commercially available methyl salicylate lures. A drawback associated with sticky bands is the high numbers of nontarget captures. Here, we developed traps for L. delicatula based on a circle trap originally designed for weevils. These traps are comprised of a screen funnel that wraps around the trunk of a tree and guides individuals walking up the trunk into a collection device. In 2018 and 2019, we compared circle trap designs with sticky bands in Pennsylvania and Virginia. In both years, circle trap designs yielded captures that were equivalent to or exceeded captures of L. delicatula on sticky bands. Nontarget captures were significantly lower for circle traps compared with sticky bands. Presence of a methyl salicylate lure in association with traps deployed on host trees or vertical tree-mimicking posts did not increase L. delicatula captures compared with unbaited traps. Circle traps, modified using vinyl screen and a larger collection device, present an alternative to the current approach with reduced nontarget capture for monitoring L. delicatula.

Potential of entomopathogenic nematodes against the pupal stage of the apple maggot Rhagoletis pomonella (Walsh) (Diptera: Tephritidae).

The apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae), is considered a key pest of apples and is native to the eastern United States. The virulence of seven different species of entomopathogenic nematodes (EPN) was assessed against pupae of R. pomonella under laboratory conditions. Nematode species and strains included Steinernema carpocapsae (ALL strain), Steinernema feltiae (SN strain), Steinernema riobrave (355 strain), Steinernema glaseri (VS strain), Heterorhabditis bacteriophora (VS strain), Heterorhabditis indica (HOM1 strain), and Heterorhabditis megidis (UK211 strain). We conducted three bioassays: (i) short-term exposure cup bioassay (7 d), (ii) long-term cup bioassay (30 d), and (iii) pot bioassay (30 d). In the short-term exposure bioassay, all nematode strains (applied at 54 infective juvenile nematodes (IJs) cm-2) significantly reduced (range: 42.9-73.8%) insect survival relative to the control, but no differences were observed among the treatments. For the long-term exposure bioassay, using the same EPN application rate as the short exposure assay, all treatments reduced adult R. pomonella emergence compared with the control. Steinernema riobrave was the most virulent (28.3% survival), and S. glaseri and H. megidis were the least virulent (53.3% survival). In the pot experiment, S. riobrave and S. carpocapsae (applied at 27 IJs cm-2) had the highest virulence (23.3 and 31.7% survival of R. pomonella, respectively), while H. bacteriophora was the least effective (68.33% survival). Our results indicate that S. riobrave, S. carpocapsae, and S. feltiae have substantial potential to attack R. pomonella pupae, and their field application under the tree canopy (prior to adult emergence) in the spring when temperatures are conducive might be a good option for successful IPM of apple maggot fly.The apple maggot, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae), is considered a key pest of apples and is native to the eastern United States. The virulence of seven different species of entomopathogenic nematodes (EPN) was assessed against pupae of R. pomonella under laboratory conditions. Nematode species and strains included Steinernema carpocapsae (ALL strain), Steinernema feltiae (SN strain), Steinernema riobrave (355 strain), Steinernema glaseri (VS strain), Heterorhabditis bacteriophora (VS strain), Heterorhabditis indica (HOM1 strain), and Heterorhabditis megidis (UK211 strain). We conducted three bioassays: (i) short-term exposure cup bioassay (7 d), (ii) long-term cup bioassay (30 d), and (iii) pot bioassay (30 d). In the short-term exposure bioassay, all nematode strains (applied at 54 infective juvenile nematodes (IJs) cm−2) significantly reduced (range: 42.9-73.8%) insect survival relative to the control, but no differences were observed among the treatments. For the long-term exposure bioassay, using the same EPN application rate as the short exposure assay, all treatments reduced adult R. pomonella emergence compared with the control. Steinernema riobrave was the most virulent (28.3% survival), and S. glaseri and H. megidis were the least virulent (53.3% survival). In the pot experiment, S. riobrave and S. carpocapsae (applied at 27 IJs cm−2) had the highest virulence (23.3 and 31.7% survival of R. pomonella, respectively), while H. bacteriophora was the least effective (68.33% survival). Our results indicate that S. riobrave, S. carpocapsae, and S. feltiae have substantial potential to attack R. pomonella pupae, and their field application under the tree canopy (prior to adult emergence) in the spring when temperatures are conducive might be a good option for successful IPM of apple maggot fly.

Influence of Landscape Factors and Abiotic Conditions on Dispersal Behavior and Overwintering Site Selection by Halyomorpha halys (Hemiptera: Pentatomidae).

Since the initial detection of the invasive Halyomorpha halys (Stål) in the United States in the late 1990s, this insect has emerged as a severe agricultural and nuisance pest. Nuisance problems are due to adult dispersal to overwintering sites in the fall at which time they alight onto and eventually settle within human-made structures in addition to natural harborage. This study examined how three factors, elevation, light, and moisture affected overwintering site selection by H. halys in the mid-Atlantic. Observational counts performed along elevational transects revealed elevation was significant predictor of H. halys abundance during both years of the study in 2014 and 2015 with more adults observed at higher elevations. Choice tests examining effects of moisture and light on settling behavior demonstrated H. halys settled within overwintering shelter boxes in significantly greater numbers when shelters were dry compared with those having moist conditions, and in darkened shelters compared with those augmented with LED lights. Our findings indicate that H. halys use cues at both landscape and very localized levels when seeking and selecting overwintering sites.

Identification of volatiles released by diapausing brown marmorated stink bug, Halyomorpha halys (Hemiptera: Pentatomidae).

The brown marmorated stink bug, Halyomorpha halys, is an agricultural and urban pest that has become widely established as an invasive species of major concern in the USA and across Europe. This species forms large aggregations when entering diapause, and it is often these aggregations that are found by officials conducting inspections of internationally shipped freight. Identifying the presence of diapausing aggregations of H. halys using their emissions of volatile organic compounds (VOCs) may be a potential means for detecting and intercepting them during international freight inspections. Headspace samples were collected from aggregations of diapausing H. halys using volatile collection traps (VCTs) and solid phase microextraction. The only compound detected in all samples was tridecane, with small amounts of (E)-2-decenal found in most samples. We also monitored the release of defensive odors, following mechanical agitation of diapausing and diapause-disrupted adult H. halys. Diapausing groups were significantly more likely to release defensive odors than diapause-disrupted groups. The predominant compounds consistently found from both groups were tridecane, (E)-2-decenal, and 4-oxo-(E)-2-hexenal, with a small abundance of dodecane. Our findings show that diapausing H. halys do release defensive compounds, and suggest that volatile sampling may be feasible to detect H. halys in freight.