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Simple Summary: The spotted lanternfly (SLF) is an invasive generalist that is spreading throughout the eastern United States. They feed in large aggregations, but the mechanisms they use to form these aggregations and find each other for mating are poorly understood. Laboratory evidence that they use pheromones is based on volatiles from their bodies and from honeydew excretions, but, due to their broad sensory capabilities, the pheromone components have been difficult to isolate and identify. This is the first evidence gathered in a field study that demonstrates, using naturally collected SLF odors as lures, that volatiles from SLF bodies and honeydew are both required to attract males at mating time and females during oviposition time. 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. [ABSTRACT FROM AUTHOR]

Simple Summary: Giant pine scale, Marchalina hellenica Gennadius (Hemiptera, Marchalinidae), is a sap sucking insect native to the Eastern Mediterranean Basin. In 2014, giant pine scale was detected for the first time in Victoria, Australia, feeding on a new host, Pinus radiata. We studied the life cycle and feeding activity of giant pine scale in Victoria over 32 months, with the aim of drawing comparisons between exotic and native populations. Australian life stages of this pest emerged during similar months to Greek seasonal equivalents, although the timing of Australian life stages differed between years. Insect density and feeding activity on infested trees differed among locations and between generations. Strong evidence was found in support of density and feeding intensity being explained by climatic conditions. The value of Pinus radiata as a food source for this insect may fluctuate with climate conditions. Our findings aim to inform future management efforts for this scale insect, including surveillance strategies and optimal seasons for release of biocontrol agents. Our findings also suggest that the impact of this pest in Australia may be exacerbated by climate change. Invasive insects pose an increasing risk to global agriculture, environmental stability, and public health. Giant pine scale (GPS), Marchalina hellenica Gennadius (Hemiptera: Marchalinidae), is a phloem feeding scale insect endemic to the Eastern Mediterranean Basin, where it primarily feeds on Pinus halepensis and other Pinaceae. In 2014, GPS was detected in the southeast of Melbourne, Victoria, Australia, infesting the novel host Pinus radiata. An eradication program was unsuccessful, and with this insect now established within the state, containment and management efforts are underway to stop its spread; however, there remains a need to understand the insect's phenology and behaviour in Australia to better inform control efforts. We documented the annual life cycle and seasonal fluctuations in activity of GPS in Australia over a 32 month period at two contrasting field sites. Onset and duration of life stages were comparable to seasons in Mediterranean conspecifics, although the results imply the timing of GPS life stage progression is broadening or accelerating. GPS density was higher in Australia compared to Mediterranean reports, possibly due to the absence of key natural predators, such as the silver fly, Neoleucopis kartliana Tanasijtshuk (Diptera, Chamaemyiidae). Insect density and honeydew production in the Australian GPS population studied varied among locations and between generations. Although insect activity was well explained by climate, conditions recorded inside infested bark fissures often provided the weakest explanation of GPS activity. Our findings suggest that GPS activity is strongly influenced by climate, and this may in part be related to changes in host quality. An improved understanding of how our changing climate is influencing the phenology of phloem feeding insects such as GPS will help with predictions as to where these insects are likely to flourish and assist with management programs for pest species. [ABSTRACT FROM AUTHOR]

Simple Summary: Like many aphids, the sorghum aphid (SA) Melanaphis sorghi produces honeydew, a waste product that can increase parasitoid retention and elicit foraging behaviors on plants. We determined the potential of SA honeydew to retain a common aphid parasitoid, Aphelinus nigritus, to assess the practicality of SA biological control on grain sorghum, Sorghum bicolor. Since SAs also feed on Johnson grass, Sorghum halepense, an alternative plant host, we characterized the composition of honeydew from aphids feeding on either grain sorghum or Johnson grass and evaluated A. nigritus preference for honeydew produced on either plant species. Our study found that A. nigritus often remained on honeydew produced by SAs feeding on both grain sorghum and Johnson grass. However, despite sharing similar sugar, amino acid, and organic acid profiles, honeydew produced on Johnson grass was preferred by A. nigritus over honeydew produced on grain sorghum. Our results suggest that SA honeydew could facilitate A. nigritus parasitoid retention on Johnson grass to lower SA populations before the grain sorghum growing season. How aphid parasitoids of recent invasive species interact with their hosts can affect the feasibility of biological control. In this study, we focus on a recent invasive pest of US grain sorghum, Sorghum bicolor, the sorghum aphid (SA), Melanaphis sorghi. Understanding this pest's ecology in the grain sorghum agroecosystem is critical to develop effective control strategies. As parasitoids often use aphid honeydew as a sugar resource, and honeydew is known to mediate parasitoid–aphid interactions, we investigated the ability of SA honeydew to retain the parasitoid Aphelinus nigritus. Since SAs in the US have multiple plant hosts, and host–plant diet can modulate parasitoid retention (a major component in host foraging), we measured SA honeydew sugar, organic acid, and amino acid profiles, then assessed via retention time A. nigritus preference for honeydew produced on grain sorghum or Johnson grass, Sorghum halepense. Compared to a water control, A. nigritus spent more time on SA honeydew produced on either host plant. Despite similar honeydew profiles from both plant species, A. nigritus preferred honeydew produced on Johnson grass. Our results suggest the potential for SA honeydew to facilitate augmentation strategies aimed at maintaining A. nigritus on Johnson grass to suppress SAs before grain sorghum is planted. [ABSTRACT FROM AUTHOR]

Simple Summary: Planthopper outbreaks in rice are associated with excessive fertilizer applications. Public research has focused on developing resistant rice to combat these outbreaks. However, to preserve ecosystem resilience, natural enemy efficacy should be maintained on resistant and susceptible rice. We examined the impact of egg parasitoids on planthoppers (Nilaparvata lugens (Stål) [BPH] and Sogatella furcifera (Horváth) [WBPH]) and a leafhopper (Nephotettix virescens (Distant) [GLH]) in field plots of resistant (IR62) and susceptible (IR64) rice under low and high nitrogen. GLH and WBPH were more abundant in low-nitrogen plots during dry (GLH) and wet (GLH, WBPH) season sampling at an early crop stage. GLH were also more abundant on IR64. Parasitoids killed between 24 and 52% of planthopper eggs during exposures in trap plants. Parasitism by Oligosita and Anagrus wasps was higher on IR64 (BPH eggs) and in high-nitrogen plots (Oligosita spp. on BPH and WBPH eggs; Anagrus spp. on BPH eggs). Parasitism by Anagrus spp. was associated with the presence of honeydew and was highest where honeydew was derived from BPH feeding on IR62; these effects were only observed under high nitrogen. Results suggest that honeydew from IR62 favors parasitoids when plants are most vulnerable (i.e., under high nitrogen), thereby countering nitrogen-induced declines in host resistance. Host plant resistance is the most researched method for the management of planthoppers and leafhoppers in tropical rice. For optimal effects, resistance should be resilient to fertilizer inputs and work in synergy with natural enemies. In field plot experiments, we examined how rice resistance and fertilizer inputs affect mortality of planthopper and leafhopper eggs by hymenopteran parasitoids. We used IR62 as a variety with resistance to Nilaparvata lugens (Stål) [BPH], Sogatella furcifera (Horváth) [WBPH] and Nephotettix virescens (Distant) [GLH], and IR64 as a susceptible control. The herbivores were more abundant during wet season sampling in low-nitrogen plots. During this study, parasitoids killed between 31 and 38% of BPH eggs and 24 and 52% of WBPH eggs during four days of field exposure. Parasitism, mainly due to Oligosita spp., was generally higher in high-nitrogen and IR64 plots. Similar densities of eggs in exposed plants suggest that these trends were mediated by semiochemicals and therefore support the Optimal Defense Hypothesis. Honeydew from BPH on IR62 had more xylem-derived wastes than honeydew on IR64. We applied honeydew from both varieties to sentinel plants. Parasitism by Anagrus spp. was higher on plants of either variety treated with honeydew derived from IR62; however, the effect was only apparent in high-nitrogen plots. Results suggest that Anagrus spp., by responding to honeydew, will counter the nitrogen-induced enhancement of planthopper fitness on resistant rice. [ABSTRACT FROM AUTHOR]

Simple Summary: The brown planthopper (Niliparvata lugens [Stål]) feeds on rice in Asia. Rice breeders have developed varieties with resistance to the planthopper; however, planthoppers quickly adapt to these varieties (called 'virulence adaptation'). We examined the potential role during adaptation of endosymbionts that are passed by the female through the egg. We adapted planthoppers by exposure to resistant rice (IR62) for >20 generations, and crossed males or females from these adapted populations with non-adapted males and females that were reared on susceptible rice. Virulent males and females contributed to the virulence of their offspring as revealed through faster development rates, higher growth rates, greater numbers of female offspring, and an increased longevity of progeny on IR62 compared to the progeny of avirulent planthoppers on IR62; however, the female contribution to virulence was greater than that of the male. Reducing densities of YLS in adult planthoppers through heat treatment reduced their capacity to gain weight on IR62, irrespective of the virulence of parents, thereby suggesting that YLS derived from the female parent play a role in the vertical transfer of virulence; however, the effects were weak and IR62 seedlings were still more affected by heat-treated planthoppers with a virulent female parent than by heat-treated planthoppers with a virulent male parent or with avirulent parents only. Virulence is a complex state with multiple underlying mechanisms, some of which are derived only from the female parent. The adaptation by planthoppers to feed and develop on resistant rice is a challenge for pest management in Asia. We conducted a series of manipulative experiments with the brown planthopper (Nilaparvata lugens (Stål)) on the resistant rice variety IR62 (BPH3/BPH32 genes) to assess behavioral and bionomic changes in planthoppers exhibiting virulence adaptation. We also examined the potential role of yeast-like symbionts (YLS) in virulence adaptation by assessing progeny fitness (survival × reproduction) following controlled matings between virulent males or females and avirulent males or females, and by manipulating YLS densities in progeny through heat treatment. We found virulence-adapted planthoppers developed faster, grew larger, had adults that survived for longer, had female-biased progeny, and produced more eggs than non-selected planthoppers on the resistant variety. However, feeding capacity—as revealed through honeydew composition—remained inefficient on IR62, even after 20+ generations of exposure to the resistant host. Virulence was derived from both the male and female parents; however, females contributed more than males to progeny virulence. We found that YLS are essential for normal planthopper development and densities are highest in virulent nymphs feeding on the resistant host; however, we found only weak evidence that YLS densities contributed more to virulence. Virulence against IR62 in the brown planthopper, therefore, involves a complex of traits that encompass a series of behavioral, physiological, and genetic mechanisms, some of which are determined only by the female parent. [ABSTRACT FROM AUTHOR]

Simple Summary: Rice in Asia is damaged by a range of sap-sucking planthoppers. For the last 50 years, the main focus for integrated management of these insects was to develop resistant rice varieties. A single, bulk phenotyping test, known as the standard seedling seedbox test (SSST) was used to inform the majority of genetics and breeding programs for planthopper-resistant rice. However, there has been much debate over the efficacy of this test. We compared results from the SSST against those from a range of other phenotyping tests to assess how antibiosis and tolerance (the ability of rice to compensate for damage) affect test results, and how phenotyping might be improved to capture information regarding plant traits that are otherwise overlooked by the SSST. Based on the efficacy and costs of different tests, we recommend that breeding programs adopt a modified seedling seedbox test (MSST) when phenotyping 100s of rice lines but that insect performance tests that assess relative changes in planthopper biomass could be adopted when phenotyping fewer lines (e.g., <100 lines). The days-to-wilt (DTW) test was found to be particularly effective in segregating resistant/tolerant lines from susceptible lines. The standard seedling seedbox test (SSST) is the most prevalent phenotyping test in research on the genetics and breeding of planthopper-resistant rice. Using 16 rice lines that included plants susceptible, resistant and tolerant to the brown planthopper (Nilaparvata lugens), we compared the SSST to modified seedling seedbox tests (MSSTs) and the days-to-wilt (DTW) test. We also conducted a series of performance tests to assess nymph survival and development; adult longevity and egg-laying; egg survival; honeydew production; and plant weight loss. We also assessed the relative costs of the different phenotyping tests to better recommend test protocols that are suitable for high-throughput phenotyping. The SSST was found to be highly robust but fails to identify late-stage resistance; tolerance; or ovicidal responses. MSSTs improved phenotyping by identifying plants with low damage from planthoppers at later growth stages. Herbivore performance tests such as population or biomass build-up tests reduce space requirements and reduce setup and evaluation costs compared with bulk tests. They can also facilitate the assessment of plant tolerance; albeit with added costs. The DTW test most clearly segregates resistant and susceptible plants, thereby facilitating gene discovery and marker-assisted selection. We recommend that bulk testing be improved by switching from the SSST to a suitable MSST and that donor variety and pre-release lines be assessed for the nature of rice–planthopper interactions using biomass build-up tests—including the DTW test. [ABSTRACT FROM AUTHOR]

Simple Summary: Plant sap-sucking insects can secrete saliva into the plant and excrete honeydew on the plant surface, which both interfere with the plant's immune system. However, knowledge about the composition of insect honeydew and the roles it plays in plant defenses is limited. In the present study, a diversity of proteins in the honeydew of two major rice pests, the brown planthopper and green rice leafhopper, were identified by way of proteomic analysis. Results revealed that the honeydew contains some insect proteins that originate from the saliva and guts, proteins that originate from plant sap via the insect digestion system as well as many bacterial proteins from insect or plant symbionts and from plant surfaces. These abundant proteins from insects, microbes and plants in the honeydew might be elicitors, effectors or self-recognized molecules for plant defenses, which provide further insights into their roles in the multitrophic interactions of plants–insects–microbes. Honeydew is a watery fluid excreted by plant sap-feeding insects. It is a waste product for the insect hosts. However, it plays important roles for other organisms, such as serving as a nutritional source for beneficial insects and bacteria, as well as elicitors and effectors modulating plant responses. In this study, shotgun LC–MS/MS analyses were used to identify the proteins in the honeydew from two important rice hemipteran pests, the brown planthopper (Nilaparvata lugens, BPH) and green rice leafhopper (Nephotettix cincticeps, GRH). A total of 277 and 210 proteins annotated to insect proteins were identified in the BPH and GRH honeydews, respectively. These included saliva proteins that may have similar functions as the saliva proteins, such as calcium-binding proteins and apolipophorin, involved in rice plant defenses. Additionally, a total of 52 and 32 Oryza proteins were identified in the BPH and GRH honeydews, respectively, some of which are involved in the plant immune system, such as Pathogen-Related Protein 10, ascorbate peroxidase, thioredoxin and glutaredoxin. Coincidentally, 570 and 494 bacteria proteins were identified from the BPH and GRH honeydews, respectively, which included several well-known proteins involved in the plant immune system: elongation factor Tu, flagellin, GroEL and cold-shock proteins. The results of our study indicate that the insect honeydew is a complex fluid cocktail that contains abundant proteins from insects, plants and microbes, which may be involved in the multitrophic interactions of plants–insects–microbes. [ABSTRACT FROM AUTHOR]

Integrated pest management (IPM) is today a widely accepted pest management strategy to select and use the most efficient control tactics and at the same time reduce over-dependence on chemical insecticides and their potentially negative environmental effects. One of the main pillars of IPM is biological control. While biological control programs of pest insects commonly rely on natural enemies such as predatory insects, parasitoids and microbial pathogens, there is increasing evidence that plant, soil and insect microbiomes can also be exploited to enhance plant defense against herbivores. In this mini-review, we illustrate how microorganisms from diverse origins can contribute to plant fitness, functional traits and indirect defense responses against pest insects, and therefore be indirectly used to improve biological pest control practices. Microorganisms in the rhizosphere, phyllosphere and endosphere have not only been shown to enhance plant growth and plant strength, but also promote plant defense against herbivores both above- and belowground by providing feeding deterrence or antibiosis. Also, herbivore associated molecular patterns may be induced by microorganisms that come from oral phytophagous insect secretions and elicit plant-specific responses to herbivore attacks. Furthermore, microorganisms that inhabit floral nectar and insect honeydew produce volatile organic compounds that attract beneficial insects like natural enemies, thereby providing indirect pest control. Given the multiple benefits of microorganisms to plants, we argue that future IPMs should consider and exploit the whole range of possibilities that microorganisms offer to enhance plant defense and increase attraction, fecundity and performance of natural enemies. [ABSTRACT FROM AUTHOR]

Infestation of willow plants by the giant willow aphid Tuberolachnus salignus (Hemiptera: Aphididae) is associated with copious deposition of sugar-rich honeydew under the plant canopy. We explored the effect of aphid honeydew on the soil biota and biochemical indicators in a two-year field trial. Soil samples from under aphid-infested and control willow trees, as well as samples from black sooty mould spots under the aphid-infested willows were compared; soil samples before aphid inoculation were used as a baseline. The honeydew deposition had a positive effect on the total soil carbon (C), but not on the total soil nitrogen content or soil pH. Microbial biomass C, basal respiration, number of yeast colony forming units, and the geometric mean of activities for six enzymes were significantly higher in honeydew-affected soils than in the control treatment on both years. The honeydew deposition also increased soil meso-fauna abundance, especially in the black sooty mould spots. The soil biochemical properties, which differed before and after aphid infestation, showed considerable overlap between the first and second year post-infestation. The results highlight the cascading effects of T. salignus on soil biological activity and the importance of using a multitrophic approach to explore similar scenarios. [ABSTRACT FROM AUTHOR]

Plant sugar is an essential dietary constituent for mosquitoes, and hemipteran honeydew is one of the many forms of plant sugar that is important to mosquitoes. Many insects rely on volatile honeydew semiochemicals to locate aphids or honeydew itself. Mosquitoes exploit volatile semiochemicals to locate sources of plant sugar but their attraction to honeydew has not previously been investigated. Here, we report the attraction of female yellow fever mosquitoes, Aedes aegypti, to honeydew odorants from the green peach aphid, Myzus persicae, and the pea aphid, Acyrthosiphon pisum, feeding on fava bean, Vicia faba. We used solid phase micro-extraction and gas chromatography-mass spectrometry to collect and analyze headspace odorants from the honeydew of A. pisum feeding on V. faba. An eight-component synthetic blend of these odorants and synthetic odorant blends of crude and sterile honeydew that we prepared according to literature data all attracted female A. aegypti. The synthetic blend containing microbial odor constituents proved more effective than the blend without these constituents. Our study provides the first evidence for anemotactic attraction of mosquitoes to honeydew and demonstrates a role for microbe-derived odorants in the attraction of mosquitoes to essential plant sugar resources. [ABSTRACT FROM AUTHOR]

Honeydew is considered a cornerstone of the interactions between aphids and their natural enemies. Bacteria activity occurring in aphid honeydew typically results in the release of volatile organic compounds (VOCs) that are used by the natural enemies of aphids to locate their prey. Because atmospheric carbon dioxide (CO2) concentration directly impacts the physiology of plants, we raise the hypothesis that elevated CO2 concentrations impact the quantity of honeydew produced by aphids, as well as the diversity and quantity of honeydew VOCs, leading to cascade effects on the foraging behavior of aphids’ natural enemies. Using solid-phase microextraction, we analyzed the VOCs emitted by honeydew from pea aphids (Acyrthosiphon pisum Harris) reared under 450 ± 50 ppm of CO2 (aCO2) or 800 ± 50 ppm CO2 (eCO2). While the total amount of honeydew excreted was only slightly reduced by eCO2 concentrations, we detected qualitative and quantitative differences in the semiochemistry of aphid honeydew between CO2 conditions. Three VOCs were not found in the honeydew of eCO2 aphids: 3-methyl-2-buten-1-ol, 2-methyl-1-butanol, and isobutanol. However, no difference was observed in the searching and oviposition behaviors of hoverfly (Episyrphus balteatus (De Geer)) females exposed to plants covered with honeydew originating from the different CO2 conditions. The present work showed the effect of a particular aspect of atmospheric changes, and should be extended to other abiotic parameters, such as temperature. [ABSTRACT FROM AUTHOR]