Plant sap-feeding insects must dispose of excess fluid, yet at millimeter scales droplet release is constrained by capillary adhesion and contact-line pinning. How phloem-feeding insects solve this puzzle, particularly as the excretory apparatus changes in size and form from nymph to adult, has remained unclear. Combining micro-CT, high-speed imaging, measurements of honeydew properties, and reduced-order modeling, we show that the spotted lanternfly (Lycorma delicatula) uses distinct release mechanics across ontogeny. Nymphs release honeydew with an anal stylus that acts as a capillary rectifier, imposing a curvature asymmetry that biases the attached droplet toward detachment through a Laplace-pressure difference. Adults use a longer stylus associated with an elastic basal region, maintain stylus-droplet contact through a finite compression phase, and release droplets with greater translational and rotational momentum. In both stages, stylus rotation is ultrafast, with peak angular accelerations of order 10^7 rad/s^-2 and release unfolding on millisecond timescales, yet droplet ejection speed remains below stylus tip speed. Weber-Bond scaling based on measured honeydew properties places both stages at We_d<1 and Bo_d<1 at the outlet, but distinguishes their post-release states: nymphal droplets remain surface-tension dominated, whereas adult droplets enter deformation- and spin-influenced regimes. Development therefore maintains waste clearance across ontogeny under the same outlet-scale capillary constraint by changing how stylus motion is coupled to the droplet at release, linking life-stage biomechanics to honeydew placement in this invasive phloem feeder and suggesting bioinspired strategies for droplet ejection, antifouling, and self-cleaning surfaces.

IntroductionThe tea shot hole borer (TSHB), Euwallacea perbrevis (Schedl 1951) (Coleoptera: Curculionidae) is an invasive ambrosia beetle that carries multiple symbiotic fungi and vectors Fusarium spp. to avocado (Persea americana Mill.). This study investigated the role of six fungal species (Fusarium sp. FL-1, Fusarium sp. AF-8, Fusarium sp. AF-6, Graphium sp., Acremonium sp., and Acremonium murorum) as nutritional symbionts of TSHB, and the role of Fusarium species in plant pathogenicity.MethodsFour experimental approaches were used: (1) testing each of the six symbionts as a food source for TSHB larvae, (2) examining the stability of symbiotic associations by rearing TSHB on substrates previously colonized by individual fungi, (3) establishing TSHB colonies with single Fusarium symbionts (Mono-Fusarium Lines, MFL), (4) testing disease development in avocado trees infested with MFL.ResultsFusarium sp. FL-1 and Fusarium sp. AF-8 supported the highest percentage of larval development among the tested fungi. These two fungi persisted in the mycangia of beetles reared on a substrate pre-inoculated with other symbionts. In addition, both fungal species caused the largest lesions in avocado branches. TSHB feeding on the other tested symbionts (Fusarium sp. AF-6, Graphium sp., Acremonium sp. or Acremonium murorum) resulted in poor larval development and/or overall reduced reproduction compared to feeding upon Fusarium sp. FL-1 and AF-8 and the symbiont blend (control).DiscussionThese findings demonstrate the dual role of Fusarium sp. FL-1 and AF-8 as nutritional symbionts of TSHB and as key drivers of pathogenicity in avocado.

The spotted lanternfly, Lycorma delicatula (Hemiptera: Fulgoridae) (SLF), is a damaging invasive pest and generalist phloem feeder that has been found in 18 states in the United States. It has a complex multimodal communication system involving semiochemicals, emitted both from their honeydew and their bodies, and substrate-borne vibrations. Sensitive and effective traps for detection and survey are essential management tools, but no potent lures for SLF exist yet. We sought to test an alternative that relies on live-trapped SLF acting as lures to improve trap efficacy after the first SLF is captured. SLF circle traps were modified by replacing the commonly used plastic collection bag with a mesh bag pinned to the tree trunk. These allowed the trapped SLF to remain alive and generate signals through the mesh bag, thus leveraging their natural modes of communication to draw additional SLF into the traps. We compared mesh and plastic bags over three years targeting fourth instars and adults and found that prior to oviposition, circle traps with mesh bags captured significantly more fourth instar (70% mesh: 30% plastic) and adult SLF (59% mesh: 41% plastic) compared to plastic bags, but during oviposition time, the results were mixed.

Since its introduction into the USA, the spotted lanternfly (SLF), Lycorma delicatula, (White) (Hemiptera: Fulgoridae) has spread across the landscape relatively unchecked. With a wide host range, it is considered a serious pest of native forest species, as well as agricultural crops. Circle traps placed on Ailanthus altissima (Miller) Swingle (Sapindales: Simaroubaceae) are passive traps collecting SLF as they walk up and down the tree trunk. These traps are successful at detecting new populations of SLF, but this can be challenging to implement at a large scale due to costs and host availability. To improve and facilitate SLF trapping practices, we investigated three key trapping components: improved collection containers, placement on alternative hosts, and lure (methyl salicylate) impact. In initial trials comparing collection jars to removable plastic bags, the adult SLF catch was four times higher using the bag design. In a multi-state survey at varying population densities, the bag traps were comparable to the jar traps but were significantly more effective than BugBarrier® tree bands, especially during the adult stage. Catch and detection in circle traps placed on alternative hosts, Acer spp. L. (Sapindales: Sapindalaceae) and Juglans nigra L. (Fagales: Juglandaceae), were comparable to those placed on the preferred host A. altissima, especially in the earlier life stages. Additionally, detection rates of methyl salicylate-baited traps on all three hosts were comparable to those on non-baited traps. These results suggest that circle traps fitted with bags provide higher trap catch and an improvement in sample quality. In addition, circle traps were equally effective when placed on maple and black walnut, while methyl salicylate lures do not enhance trap catch or detection.

Survey and detection of the spotted lanternfly, Lycorma delicatula (White), rely either on traps that exploit the insect’s behavior as it navigates its environment, or on visual surveys of either its mobile life stages or egg masses. A recently described egg mass trap, coined the “lampshade” trap, can assist with early detection in newly infested areas, provide egg masses for researchers, and potentially facilitate spotted lanternfly population reduction by removal of egg masses from the environment. Here, we describe a modified lampshade trap, the sconce trap, that uses less material, can be pre-cut prior to deployment, and can be deployed by one person, representing potential cost, labor, and time savings. Both traps were comparable at detecting populations of spotted lanternflies, and while females deposited more eggs on the larger lampshade traps, they deposited more eggs on sconce traps as a function of trap area.

Using semiochemicals collected from spotted lanternflies Lycorma delicatula (Hemiptera: Fulgoridae) (SLF) and deployed in the field with circle traps, we demonstrated that SLF responded to SLF pheromones: in particular, this was the case for males while seeking mates and for females while ovipositing. The attractants consisted of SLF body extract emitted from diffuser lures and SLF honeydew on burlap ribbons, collected from heavily infested locations. Traps with attractants were deployed in field sites with very light SLF infestations to avoid competing signals of pre-existing aggregations. The number of SLF equivalents emitted by each diffuser per trapping period was used in a dose–response analysis. Three trees per block received either (1) a control hexane lure and a clean ribbon, (2) a lure containing SLF extract and a clean ribbon, or (3) a lure containing SLF extract and a honeydew-laden ribbon. Ten blocks were sampled three times per week for twelve weeks. We found a significant positive dose–response by males to SLF body extract only in the presence of SLF honeydew, indicating a synergistic effect between honeydew volatiles and body volatiles. This dose–response occurred for five weeks after mating started, after which males no longer responded. Subsequently, females had a significant positive dose–response to SLF extract only in the presence of honeydew when oviposition was their primary activity, continuing for two weeks, suggesting that females may use pheromones to aggregate for oviposition. The extract in the absence of honeydew did not result in a positive dose–response, nor did the hexane control. These findings suggest that SLF respond synergistically to the combination of pheromones present in both SLF honeydew and SLF bodies. Thus, combining key components from both sources may aid the development of semiochemical lures for SLF.

In herbivorous insects, antennae play a crucial role in chemical communication and orientation when locating host plants and mates. To evaluate antennal sensitivity in response to odor stimuli, electroantennography (EAG) has been a practical technique. In the current study of the invasive spotted lanternfly (SLF), Lycorma delicatula (Hemiptera: Fulgoridae), we evaluated and compared their antennal sensitivity to a series of volatile chemicals collected from their bodies, honeydew, and host plants. To do this, we exposed the antennae of SLF fourth-instar and adult males and females to individual chemicals at a fixed dose of 50 ng. Further, a series of dose–response tests were carried out within a range of 0.5 to 100 ng. Although the amplitude of antennal responses varied among stages and sexes, adult males generated the strongest antennal responses in both experiments. In dose–response experiments, increased doses of chemicals up to 50 ng revealed the saturation points except in adult females which required a higher dose (100 ng) to reveal the saturation point. Although EAG does not provide any information on behavioral responses, our results are consistent with the olfactory bioassays in previous publications in which adult males, not females, were attracted to natural volatiles of their conspecifics. EAG indicated a higher sensitivity of adult male antennae to odor stimuli, particularly conspecific volatiles, than female antennae and highlighted sexual differences in the perception of chemical cues in SLF.

Lycorma delicatula (White) (Hemiptera: Fulgoridae), spotted lanternfly (SLF), is an invasive pest that feeds and oviposits on numerous woody and herbaceous plants important to agricultural, forest, ornamental, and nursery industries. Describing and understanding SLF movements is key to implementing surveillance and control strategies for this pest and projecting population spread. We used radio telemetry (RT) and harmonic radar (HR) to track the movements of individual SLF at field sites in eastern Pennsylvania and northwestern New Jersey. SLF equipped with HR or RT tags were tracked in 2019 and 2020 from adult emergence until oviposition time, and their movements are described. Although the bulkier RT tags disproportionately affected the distance traveled by males, which are smaller than females, both males and females were more likely to be lost due to signal attenuation when affixed with the lighter-weight HR tags. Females were tracked moving longer distances than males, with maximum distances of 434 m by a single female and 57 m by a single male. A significant positive relationship was found between their height in trees and the distance of subsequent movement. Adult SLF were found in trees predominantly at heights between 6–9 m high. For the fraction of SLF found at eye level, males, but not females, significantly moved above eye level in the weeks prior to mating, likely resulting in the observed sex ratio shift that defines the Early-2 stage. During mating time, tracked SLF were significantly higher than 8 m and oriented to trees where tight aggregations of SLF were present. This orientation towards tight aggregations started when mating began and peaked in the following 2.5 weeks for males in Late-1 and the beginning of Late-2 (after oviposition began), whereas females started this orientation behavior a half-week after males, and this activity peaked for two weeks. Male and female SLF adults exhibited slight differences in host preference, and strong preferences for wild grape, black walnut, sweet birch, and tree-of-heaven were observed. The HR-tagged nymphs moved up to 27.6 m over a five-day period in a cornfield. Nitinol wire HR tags performed better than Wollaston process or tungsten wire tags. SLF movement parameters in the field are described.

Since its discovery in North America in 2014, the spotted lanternfly (SLF), Lycorma delicatula, has become an economic, ecological, and nuisance pest there. Developing early detection and monitoring tools is critical to their mitigation and control. Previous research found evidence that SLF may use pheromones to help locate each other for aggregation or mating. Pheromone production necessitates specific conditions by the insects, and these must be investigated and described. A chemical process called photo-degradation has been described as a final step in the production of pheromones in several diurnal insect species, in which cuticular hydrocarbons were broken down by sunlight into volatile pheromone components. In this study, photo-degradation was investigated as a possible pheromone production pathway for SLF. Extracts from SLF mixed-sex third and fourth nymphs and male or female adults were either exposed to simulated sunlight to produce a photo-degradative reaction (photo-degraded), or not exposed to light (crude), while volatiles were collected. Behavioral bioassays tested for attraction to volatiles from photo-degraded and crude samples and their residues. In third instars, only the volatile samples from photo-degraded mixed-sex extracts were attractive. Fourth instar males were attracted to both crude and photo-degraded residues, and volatiles of photo-degraded mixed-sex extracts. Fourth instar females were attracted to volatiles of crude and photo-degraded mixed-sex extracts, but not to residues. In adults, only males were attracted to body volatiles from crude and photo-degraded extracts of either sex. Examination of all volatile samples using gas chromatography coupled with mass spectrometry (GC-MS) revealed that most of the identified compounds in photo-degraded extracts were also present in crude extracts. However, the abundance of these compounds in photo-degraded samples were 10 to 250 times more than their abundance in the crude counterparts. Results from behavioral bioassays indicate that photo-degradation probably does not generate a long-range pheromone, but it may be involved in the production of a short-range sex-recognition pheromone in SLF. This study provides additional evidence of pheromonal activity in SLF.

A mark-release-recapture experiment was conducted to evaluate the orientation of spotted lanternfly (SLF) Lycorma delicatula White (Hemiptera: Fulgoridae) nymphs when released equidistant between two trees. The experiment was repeated weekly for eight weeks in a heavily infested area with mature tree-of-heaven Ailanthus altissima (Mill.) Swingle (Sapindales: Simaroubaceae) planted in rows as ornamental street trees in Beijing, China. One tree in each pair received a methyl salicylate lure, and the lure was rotated between trees every week as it aged. Two additional independent variables for each tree were also analyzed: size and SLF population density. Marked-released SLF significantly chose trees with higher SLF population density over trees with lower density populations, and they also chose larger trees significantly more than smaller trees. Population density and tree size were better predictors of attraction than lures, but when those factors were controlled, SLF significantly chose trees with methyl salicylate lures over control trees for the first 4 weeks of lure life. Wild SLF distribution was assessed weekly, revealing strong aggregation in first and second instars that diminished with development to the third and fourth instars. Thus, nymphal SLF aggregate, and orientation is strongly guided by the presence of other SLF and tree size.

EDITORIAL article Front. Insect Sci., 03 October 2023Sec. Invasive Insect Species Volume 3 - 2023 | https://doi.org/10.3389/finsc.2023.1292590

Species belonging to the Euwallacea fornicatus Eichhoff (Coleoptera: Scolytinae) species complex have invaded the continental U.S. since at least 2003. Three species of this complex are known to have established, two in California (E. fornicatus; and Euwallacea kuroshio), and a third in Florida (Euwallacea perbrevis). Their native ranges are spread across southern and southeast Asia. In Taiwan, all three species occur in sympatry. They attack healthy trees of widely varied species and cause severe damage and death to the trees. The attractant quercivorol is commonly used to promote their detection by passive trapping. Recent studies in Florida have shown that trapping of E. perbrevis can be further improved by adding a synergist, α-copaene, alongside the quercivorol lure. Thus, we were interested in testing the effectiveness of α-copaene for trapping the other invasive members of the complex in California and in an area of Taiwan where all three species co-occur. We found that α-copaene marginally enhanced the trapping of E. perbrevis in Taiwan, but had no effect on the trapping of E. fornicatus or E. kuroshio in either California or Taiwan. We conclude that any enhancing effect of α-copaene is specific to E. perbrevis. This highlights the economic importance of accurate species identification in developing and implementing an efficient, and yet cost-effective, monitoring program for the management of E. fornicatus and E. kuroshio in California and elsewhere.

The spotted lanternfly, Lycorma delicatula (Hemiptera: Fulgoridae) is a polyphagous, phloem-feeding invasive forest, agricultural, and nuisance pest that is rapidly spreading through the U.S. Little is known about how fulgorids locate each other to mate. To determine if conspecific semiochemicals such as pheromones occur, whole body extracts (WBE) of adult spotted lanternflies from different physiological states were tested for attraction in a dual-choice olfactometer. In olfactometer assays, prior to mating, males were oriented to WBE from males and females. During their mating period, males were attracted to WBE of females, but not to that of males. After mating and oviposition had taken place, males were not attracted to either male or female extracts. Conversely, females did not orient to any WBE from either sex during any physiological state. These behavioral responses by males but not females to WBE from conspecifics in different physiological states suggest the possible presence of an aggregation-sex pheromone in the spotted lanternfly.

The spotted lanternfly Lycorma delicatula White (Hemiptera: Fulgoridae) is a polyphagous insect pest that invaded the United States in 2014, in Berks County, Pennsylvania. It has since spread to several northeastern states and poses a significant threat to northeastern grape production. Most studied species of Hemiptera are known to communicate intraspecifically using some form of substrate-borne vibrational signals, although such behavior has not yet been reported in L. delicatula. This report demonstrates that adult and fourth-instar L. delicatula were attracted towards broadcasts of 60-Hz vibroacoustic stimuli directed to a laboratory arena and test substrate, which suggests that both adults and fourth instar nymphs can perceive and respond to vibrational stimuli.

The spotted lanternfly (SLF),Lycorma delicatula(Hemiptera: Fulgoridae), is a generalist phloem feeder that produces copious amounts of honeydew, which in turn coats the understory. These insects form large aggregations covering the trunks of some trees, while similar trees nearby mysteriously seem unattractive. We investigated whether volatiles from SLF honeydew are attractive to conspecifics by collecting honeydew from the field and testing it for SLF attraction in a two-choice olfactometer. We found that honeydew excreted by adult male SLF was significantly attractive to male SLF, but not female SLF. Although the honeydew excreted by adult female SLF did not significantly attract male or female SLF, both sexes showed a positive trend towards attraction in response to female honeydew in the olfactometer. Analysis of the headspace volatiles of honeydew was conducted, and numerous semiochemicals were identified. Five of which, 2-heptanone, 2-octanone, 2-nonanone, benzyl acetate, and 1-nonanol, were tested in two-choice behavioral assays against a blank control. Benzyl acetate and 2-octanone were attractive to both sexes, whereas 2-heptanone was only attractive to males, and 2-nonanone only to females. The remaining compound, 1-nonanol, repelled females, but not males. Although honeydew has been reported as a source of kairomones for some natural enemies, this may be the first report of sex-specific attractants for conspecific insects found in the honeydew volatiles of a planthopper.

Spotted lanternflies (SLF) Lycorma delicatula are economically important invasive planthoppers discovered in North America in 2014. SLF are gregarious, but how they locate each other, or who finds whom and when, is poorly understood. Here we describe adult SLF behavior and phenology on their preferred host, Ailanthus altissima, under field conditions, in the context of both aggregation and mate-location, since SLF demonstrated aggregation prior to mating. We documented aggregation behavior of adults and found we could manipulate free-living SLF populations in both number and sex ratio by the placement of confined populations of SLF males or females on trees. Trap capture of arriving SLF was significantly higher on trees with confined SLF aggregations than on control trees, and was corroborated with photographic data, demonstrating the manipulation of attraction and aggregation behavior. Sex ratios of trapped SLF arrivals were significantly more male-biased on trees with confined males and more female-biased on trees with confined females, evidence that the male- and female-biased sex ratios observed on trees naturally can be explained by sex-specific conspecific signals. SLF sex ratios shifted over time in the same pattern over two consecutive years. A mark-release-recapture study over time found that 1) SLF behavior is density dependent and strongly influenced by natural populations, 2) released females were captured significantly more on trees with caged females, particularly prior to mating, and 3) released males were captured significantly more on trees with caged females starting at mating time. Photographic data revealed that most clustering behavior (a measure of courtship) of free-living SLF began on trees with caged females during mating time, but not on trees with caged males or controls. We describe adult male and female SLF phenology whereby 1) aggregation behavior occurs, 2) males and females arrive at different times, 3) females began to aggregate several weeks prior to mating, 4) males subsequently joined aggregations at the time of mating, and 5) aggregation continued into oviposition. Population density and aggregation behavior were found to be key factors in their natural history which can be manipulated, providing a foothold for future research. Possible mechanisms for future exploration are discussed.

The spotted lanternfly, Lycorma dehcatula, (Hemiptera Fulgoridae) is an invasive pest in Korea and the United States, but originating from mainland Asia. Both tree of heaven, Ailanthus altissima, and grapevine (Vitis) are known to be preferred hosts of adults. However, much is unknown about its adult behavior with respect to mating or dispersal patterns. In 2015, just after it was discovered in the United States, we performed an observational study to elucidate movement by adults in relation to host plants. We noted weekly changes in their presence and sex ratio on host plants at four sites in a quarantine zone in Berks County, PA. Just after adult emergence, the trunks of large ( > 15 cm diameter at breast height) A. altissima hosted nearly exclusively females, while smaller A. altissima, Vitis, Salix nigra, and other less commonly inhabited tree species, had high proportions of males. By October 1st, greater proportions of males were observed on the trunks of the larger A. altissima trees. Pairings of males and females were observed most frequently on the smaller A. altissima and Vitis. At the very end of the season, a large mixed-sex population was found on Salix nigra, which was the only deciduous tree species in the area still bearing green leaves.

Studies were conducted from 2015 to 2018 to evaluate spotted lanternfly (SLF) distribution and developmental suitability of different plant species in the U.S. Tree bands on 283 trees spanning 33 species captured 21,006 SLF in 2 yr. More SLF per tree were trapped on tree-of-heaven Ailanthus altissima (Mill.) Swingle (Sapindales: Simaroubaceae) than on other species, on average, and most adults were captured on tree-of-heaven. Frequency of detection of adult SLF was higher on tree-of-heaven than on other species but was actually equal or lower on tree-of-heaven than on all other species combined for younger SLF stages in 2015. An enclosed choice test between tree-of-heaven and black walnut Juglans nigra L. (Fagales: Juglandaceae) revealed nymphs showed little consistent preference, whereas adults consistently and significantly preferred tree-of-heaven. No-choice field sleeve studies evaluated SLF survivorship on 26 host plant species in 17 families. Ten plant species supported SLF for an average of ≥45 d, with the rest unable to support SLF for >30 d. Eight species were able to support development from first instar to adult: black walnut, chinaberry Melia azedarach L. (Sapindales: Meliaceae), oriental bittersweet Celastrus orbiculatus Thunb. (Celastrales: Celastraceae), tree-of-heaven, hops Humulus lupulus L. (Rosales: Cannabaceae), sawtooth oak Quercus acutissima Carruthers (Fagales: Fagaceae), butternut Juglans cinerea L, and tulip tree Liriodendron tulipifiera L. (Magnoliales: Magnoliaceae). The ability of SLF to develop to adult on hosts other than tree-of-heaven may impact pest management decisions.