
Abstract Biodiversity surveys are essential for discovering species that provide ecosystem services which may assist with sustainable pest management. Surveys were conducted for North American egg parasitoids of native Fulgoridae to detect potential biological control agents for Lycorma delicatula (White, 1845) (Hemiptera: Fulgoridae). Two parasitoid species, Anastatus reduviopsis sp. nov. and A. ruficollis (Cameron, 1905), were collected in Arizona from Scaralina Yanega (Hemiptera: Fulgoridae) eggs, representing the first record of egg parasitoids of native US Fulgoridae and supporting the hypothesis that native Anastatus Motschulsky (Hymenoptera: Eupelmidae) may have biocontrol potential for L. delicatula. Laboratory colonies of A. reduviopsis sp. nov. were established from field-collected parasitoids, and preliminary observations demonstrated successful parasitism of L. delicatula eggs, providing a basis for future evaluation of its potential as a biocontrol agent for this invasive pest. Morphological and molecular analyses confirmed species identities and revealed a close relationship between A. reduviopsis sp. nov. and A. reduvii (Howard, 1880), previously shown to reduce the survival of L. delicatula eggs through parasitism and host-feeding in laboratory trials. Molecular barcodes of three Anastatus, two Merostenus Walker (Hymenoptera: Eupelmidae), and two Scaralina species are provided for the first time, supporting accurate species identifications for future research. By linking taxonomy, ecology, and applied entomology, this research is the first attempt to advance Anastatus taxonomy in North America after nearly 60 years. This work demonstrates that targeted biodiversity surveys can reveal native parasitoids potentially contributing to biotic resistance to invasive pests, thereby emphasizing the value of investing in biodiversity research.
Abstract The symbiosis of fungus-growing (attine) ant agriculture is nestled within an entangled bank of multipartite complex species interactions. In the decades since the field first arose, with a focus on the attine ants and the fungi they cultivate, there have been notable shifts in what is considered a “canonical” interaction within attine gardens. The study of ant–cultivar interactions broadened to describe tripartite interactions between ants, cultivars, and fungal parasites and continued expanding to include parasite-inhibiting bacteria. Beyond this quadripartite network, additional invertebrates and a great diversity of “non-canonical” microbes are frequently present in fungus-growing ant gardens. While providing an overview of the members of these complex, interacting communities, we summarize their functional importance from observations, experiments, genomics, and chemistry. We suggest that the ecological and evolutionary dynamics of attine agriculture are closely intertwined with multipartite interactions that occur within and around attine gardens. The field’s next steps will require interdisciplinary collaboration and integrative approaches in taxonomy, genomics, chemistry, and experimentation at the community level. These multiple lines of evidence will allow for a better understanding of the diversification and evolution of the ant–fungal agriculture as a multitrophic and multisymbiotic network.
Abstract Despite the overwhelming diversity and ecological impact of insects across most ecosystems, their responses to environmental stress remain underexplored. To survive temperate seasonal stressors, particularly winter cold conditions and nutrient and water shortage, insects undergo physiological changes and overwinter in a diapause state. During diapause, cessation of development is coupled with reduced metabolic activity, analogous to hibernation in mammals, in which gut microbiota help manage waste and recycle nitrogen. Similarly, many insects have obligate symbiotic relationships with microbes that support nutrient acquisition. However, unlike mammals, many insects have open or transient gut communities, making it more likely that microbial dynamics are strongly shaped by environmental microbes acquired through food or habitat. This review synthesizes recent literature on the roles of host-associated microbial communities in insect diapause, spanning nutrient provisioning, nitrogen recycling, and host immunity, including how associated microbes contribute to pathogen defense during dormancy. We propose future research avenues for more mechanistic understanding of host–microbe interactions involved in diapause, drawing on conceptual parallels established from mammalian hibernation research.
Abstract The arthropods and microbes found in trees attacked by the southern pine beetle (Dendroctonus frontalis Zimmermann) have long been referred to as “southern pine beetle associates.” This community includes predators, parasitoids, competitors, mutualists, symbionts, and taxa whose relationship to southern pine beetle may be unclear or not well-described. We present the first systematic review of the southern pine beetle-associated community. We performed a systematic search using the Web of Science Core Collection to identify sources addressing some aspect of southern pine beetle symbiosis or facilitation. Our search returned 231 results; of these, 174 papers fit our criteria for analysis. We also incorporated 198 additional sources from field-specific journals and government publications. From each paper, we extracted taxon names for all organisms identified as southern pine beetle-associates, any description of their ecological role, and the year, location, and study methodology. Our review reveals patterns in the literature and tells the history of this line of research. We present 760 taxa that have been reported as southern pine beetle associates. Not surprisingly given its role as a key natural enemy, the predatory clerid, Thanasimus dubius (Fabricius), was the most reported southern pine beetle associate. The number of publications increased markedly in the 1970s, coinciding with major federal funding for southern pine beetle research. Using the resulting database of southern pine beetle associates and their reported roles, we explored the central question, “what is an associate?” We categorize each southern pine beetle-associate pairing as direct or indirect: these range from observed predation and parasitism (direct association) to co-occurrence in southern pine beetle-attacked trees or in funnel traps (indirect association).
Abstract Insect remains recovered from ancient or historical graves provide data that can be used for studying past populations and their environment. A growing interest in archaeoentomology of mortuary contexts requires an accessible accounting of the current methods and available resources to promote quality outcomes for an inherently destructive process. This review of 76 unique results synthesized the methods and analyses used in existing funerary archaeoentomology research. Software used to identify, collate, track duplicates, and analyze search results included Publish or Perish and Covidence. Our search protocol yielded 1,242 original titles that were each screened for relevance and then included or excluded by 2 reviewers. Results support our hypothesis that the collection of funerary archaeoentomology material is broadly consistent with sampling techniques used by forensic entomologists. Researchers were found to collect directly from remains, hand-sort various biological grave materials, typify insect trace impressions on human bone, incorporate experimental approaches, and use dry or wet sieving when appropriate. Flexibility allowed context or material type to dictate the specific techniques or methods employed. Most papers clearly reported species richness, which was found to contain natural history observations pertinent to insect species’ native or historical ranges. As predicted, beetles (Coleoptera) and fly puparia (Diptera) were the most frequently reported entomofaunal groups. We identified specific titles that report detailed methods, which future researchers may leverage for guidance. Clear reporting on abundance data, identification criteria, and use of analytical tools are areas that require development.
Abstract Insect wings commonly have 2 types of veins: primary and secondary. Primary veins have fixed positions across members of a given species, while the number, position, and connections of secondary veins vary from wing to wing even within individuals. Relative to primary veins, very little is known about the patterning and development of secondary venation, in part because they cannot be analyzed with traditional morphometric approaches, which require the identification of homologous landmarks across wings. We present a landmark-free morphometric approach for analyzing secondary wing venation, and we use it to describe the effects of temperature on secondary vein patterns in the common cricket Acheta domesticus (Linnaeus 1758). Our principal components-based approach identified 3 major compartments to the wing within which we observed correlations among the shapes of domains formed by secondary veins. Most interestingly, the effects of temperature varied across these compartments: higher temperatures were associated with larger domains in one compartment, smaller domains in another, and had no marginal correlation with domain shape in a third. This suggests that secondary vein development may involve multiple independent patterning systems in different regions, or more likely, a system whose parameters vary across the wing field.
Extracellular vesicles (EVs) are nanoscale, lipid-bound structures released by cells across all domains of life. Once viewed as a means for discarding unwanted cellular components, they are now understood to be central mediators of intercellular communication. Much of what is known about EVs comes from mammalian systems, where extensive work has defined the major EV subtypes and the mechanisms that generate exosomes. These findings continue to serve as the primary reference for interpreting EV biology in other organisms. EVs have been isolated from diverse arthropods, including a few insect and tick species, yet the molecular pathways that produce them remain less characterized. Current evidence from Drosophila, mosquito cell lines, and tick systems shows that many of the core components driving exosome biogenesis in mammals, including ESCRT complexes, tetraspanins, lipid-modifying enzymes, Rab GTPases, and SNARE proteins, are present and, in several cases, experimentally validated in arthropods. These findings point to broad conservation of exosome biogenesis across taxa, while also highlighting key caveats, with most conclusions relying on only a small number of model systems. The definitions of EV subtypes remain unclear in many arthropods, and alternative EV biogenesis pathways have received little attention. Future studies that incorporate non-model species, apply rigorous EV characterization standards, and explore the roles of various EV subtypes will clarify how these pathways operate across arthropod lineages and how they differ from well-studied mammalian systems.
Leafhoppers (Hemiptera: Cicadellidae) use substrate-borne vibrations to communicate, and characteristics of these vibrational signals help identify, locate, and assess suitability of potential mates. The strength and frequency composition of signals influence behavioral responses in conspecifics. Leafhoppers are known to produce these vibrational signals while probing, but it is unclear whether this dual activity degrades signal transmission, inducing a trade-off between nutrient acquisition and mate attraction. In contrast, we hypothesized that inserted mouthparts could provide an additional point of contact with the plant, improving signal transmission. Here, we combine simultaneous electropenetrography (EPG) and accelerometer recordings to assess how beet leafhopper [Neoaliturus tenellus (Baker)] vibrational signals vary with probing behavior, including xylem ingestion, phloem salivation/ingestion, and probing of epidermis and mesophyll cells. Signals documented during 6-h recordings of male leafhoppers had significantly higher amplitude and dominant frequency when leafhoppers were probing than when they were not probing. However, the dominant frequency of signals was similar when leafhoppers were engaged in pathway phase, phloem ingestion, and xylem ingestion. Of the different probing behaviors, xylem ingestion had the strongest positive effect on signal amplitude, but phloem ingestion did not influence signal amplitude. Additional contact between the leafhopper and the plant surface, provided by leafhopper mouthparts, may improve vibration transmission, potentially increasing signal active space. In light of our finding that the act of probing plant tissues influences information conveyed in leafhopper vibrational signals, we suggest further research to evaluate the impacts these signal changes have on the behavior of other mates and natural enemies.
There has been limited information when it comes to understanding the nutritional intake and distribution of vitamins in insects. This is especially true for how vitamins affect insect foraging behavior. Vitamins C, B1, and B3 are essential micronutrients that many insects require for development and growth as larvae and for reproduction as adults. This study used the geometric framework design to examine the relative food preference for vitamins C, B1, and B3 in the eastern subterranean termite, Reticulitermes flavipes (Kollar). Three experiments were performed in which termites were presented with 2 artificial foods enriched with an opposing 3:1 ratio of 2 of the 3 vitamins tested in this study. They were allowed to feed on the foods for 2 wk. The amount of food consumed was measured periodically over the 2-wk period, and the total amount of each vitamin consumed was calculated. Food with a higher level of vitamin C was consumed significantly more than food with a higher level of vitamin B1 or B3. There was no apparent consumption preference between food with higher levels of vitamin B1 and B3; however, more vitamin B3 was consumed than vitamin B1 in the experiment that compared these 2 vitamins.
Rearing silkworms on artificial diets (AD) is crucial for sustainable sericulture, but their oligophagous nature makes adapting to non-mulberry leaf (ML) diets challenging. To date, the issues of reduced resistance and diminished vitality in silkworms reared on AD remain unresolved, and there is a lack of comprehensive research on alterations in their intestinal immune responses. In response to this knowledge gap, this study systematically investigated the impact of AD on intestinal stress and antibacterial capacity in silkworms. Compared to ML-fed silkworms, those reared on an AD exhibited a significant reduction in the antibacterial capacity of hemolymph and digestive fluids. Furthermore, Toll/IMD immune pathway genes from intestinal cells and antimicrobial peptide genes from fat bodies and hemolymph displayed signs of baseline immune stress, yet elicited a weak immune response following infection. Mechanistically, silkworms reared on an AD experienced a decline in intestinal antioxidant capacity, accompanied by elevated levels of reactive oxygen species. Additionally, the secretory function of midgut epithelial cells was impaired, the pH of digestive juices was decreased, the peritrophic membranes were damaged, the density of longitudinal muscle fibers in the intestine was lower, and intestinal motility was compromised. In conclusion, the intestinal barrier and functionality of silkworms reared on an AD were compromised, leading to a reduction in antibacterial capacity. These impairments were associated with pre-existing oxidative and immune stress conditions in the intestinal tissue under baseline conditions. These findings offer a novel perspective for improving silkworm feed formulations and developing immune-enhancing agents, extending beyond considerations of nutrient absorption.
Insect pollination represents one of the most consequential mutualisms in terrestrial ecosystems, yet its early evolutionary history remains poorly understood. The mid-Cretaceous, a period characterized by the rapid radiation of angiosperms, reshaped pollination networks, as insects either shifted from gymnosperm hosts to angiosperms or evolved in close association with them. Here, we report the first male of Pelretes bicolor (Kateretidae) from mid-Cretaceous Myanmar amber (similar to 99 Ma), preserved alongside three thrips (Thysanoptera). The beetle has enlarged mandibles and distinctive pronotal morphology, providing direct evidence of sexual dimorphism in this extinct lineage. The associated thrips, assignable to Parallelothrips (Stenurothripidae), carry Eucommiidites-type pollen, diagnostic of Erdtmanithecales, an enigmatic group of Mesozoic gymnosperms. This represents direct evidence of thrips pollination of Erdtmanithecales and establishes stenurothripid thrips as a generalist pollinator lineage in the Cretaceous. Reexamination of pollen associated with Pelretes vivificus, another Myanmar kateretid, indicates that these grains also correspond to the Eucommiidites type. Collectively, our study not only documents sexual dimorphism in fossil Kateretidae but also provides fossil evidence that both beetles and thrips were integral components of complex Cretaceous pollination networks during the rise of angiosperms. These findings highlight the versatility of early pollinators and shed light on the dynamics of insect-plant interactions during a critical interval in terrestrial ecosystem evolution.
Biological control through arthropod predation is one of the most economically important ecosystem services in agriculture. Because of this, documenting predator communities' ecological and behavioral responses to the environment and the resultant changes to predator activity and predation are crucial to agricultural sustainability. Temporal variability in resources, environmental conditions, and arthropod behavior is increasingly recognized for its ecological importance. However, this recognition is largely focused on shifts in diurnal aspects of the ecosystem. Nocturnal predation can match or exceed diurnal predation across predatory taxa, so ignoring nocturnal predation risks overlooking species that contribute significantly to predation services. This review assembles current knowledge about nocturnal predation in annual row crops, outlines its importance, and advocates for further work in this understudied area. Comprehensive identification of predator communities is quite rare, with nocturnal communities even more woefully under-researched than their diurnal counterparts. This represents a major blind spot in entomology, especially for predation ecology, and even persists where predation is known to primarily occur nocturnally, allowing severe underestimation of pest control by nocturnal predators. The more ephemeral nature of predation and the difficulty of direct documentation preclude many passive collection methods, especially for evaluating diel activity of the predatory complex in situ. Further, changing climates may differentially affect diurnal and nocturnal predators, leading to significant impacts on predation outcomes. Both nocturnal and diurnal predators face spiking temperatures and their associated desiccation threat and phenological shifts though different activity patterns will likely alter the impact of the changing climate on these predatory complexes.
Generalist insect vectors are notoriously difficult to manage due to their ability to use a range of host plants within and across seasons. Improving vector management requires novel approaches that assess host use across space and time to predict pathogen transmission dynamics. Molecular gut content analysis of vector insects has been instrumental in identifying host use but has often been poorly linked with hosts related to pathogen transmission. Here, we integrate gut content analysis of vector insects with monitoring pathogen incidence to determine the role of various plant hosts in Neoaliturus tenellus (Baker) (beet leafhopper) movement and pathogen spread. We tested 226 beet leafhopper adults collected from 24 potato fields and nearby weeds over 3 growing seasons to assess host use and acquisition of beet curly top virus and 'Candidatus Phytoplasma trifolii'. We observed seasonal variation in pathogen acquisition from host plants with N. tenellus acquiring pathogens from Sisymbrium spp. and Brassica spp. (wild mustards) in spring, whereas Salsola/Kali spp. (Russian thistle) and Bassia spp. (kochia) serve as the primary insect hosts during summer. Congruent with other recent research, we detected tree DNA in the guts of N. tenellus confirming tree probing or feeding, including Tilia spp. (linden), Prunus spp./Pyrus spp./Citrus spp (fruit), and Tsuga spp./Pinus spp. (pine). Our study refines our understanding of N. tenellus ecology and highlights the importance of host use patterns in predicting pathogen transmission, ultimately improving pathogen risk assessments.
Dryinus sinicus Olmi (Hymenoptera: Dryinidae) is an important nymphal parasitoid of the spotted lanternfly, Lycorma delicatula White (Hemiptera: Fulgoridae), in its native range in China. As part of evaluations of this parasitoid as a biological control agent for invasive L. delicatula in the United States, this study evaluated its immature developmental morphology and reproductive potential. Adult D. sinicus exhibits sexual dimorphism; females possess a chelate protarsus. Larvae are ectoparasitic, with the head partially embedded in the host's coelom and the body protruding between 2 sclerites, covered by the larval exuviae (thylacium). The larva develops through 5 instars before crawling out to pupate in a silken cocoon, killing its host in the process. Dryinus sinicus is highly synovigenic (ovigeny index: 0.0843), with females emerging with 15.6 +/- 2.2 mature oocytes and ready for oviposition. Host feeding and oviposition likely prompt egg production. At 20 degrees C, adult females lived 61.3 +/- 9.9 d, parasitized 137.2 +/- 34.9 nymphs, laid 175.8 +/- 46.1 eggs, and host-fed on 20.8 +/- 3.8 nymphs during their lifetime. Self-superparasitism occurred frequently in the laboratory setting, and multiple offspring occasionally developed from a single host. The lifetime fecundity and predation of D. sinicus were positively correlated with longevity. In the laboratory trials, D. sinicus more effectively parasitized and host-fed upon first than second instars and killed approximately 7 times more nymphs through parasitism than through host feeding. Overall, D. sinicus demonstrates high fecundity and strong potential as a biological control agent for L. delicatula.
Artificial intelligence (AI) is rapidly transforming the practice of science, including the field of entomology. Once viewed primarily as a tool for data analysis, AI is now reshaping publishing, research, and teaching in ways that raise both opportunities and concerns. To capture this pivotal moment, we organized a symposium at the 2024 Annual Meeting of the Entomological Society of America titled "Artificial Intelligence (AI) in Entomology: An Aid to Publishing, Research, and Teaching." Presentations highlighted diverse applications of AI, from generative models that translate Extension materials for underserved audiences, to computer vision systems detecting stored product pests, to tools accelerating the digitization of insect collections, and to experimental manipulations of dominance hierarchies in social insects. Collectively, these case studies illustrate how AI can enhance efficiency, expand the scope of questions that can be asked, and democratize access to advanced research tools. At the same time, they emphasize the importance of careful oversight, given risks such as mistranslations, misclassifications, and overreliance on generative outputs. Beyond research, AI is already reshaping scientific publishing. Drawing on symposium discussions and a review of current policies, we propose baseline recommendations for authors, reviewers, and editors that are now integrated into Entomological Society of America publishing policies. These include transparent disclosure of AI use in acknowledgments and methods, alongside clear boundaries on acceptable applications. Our synthesis underscores that while AI can accelerate entomological science, ongoing dialogue, validation, and ethical guardrails are essential to ensuring trust, integrity, and creativity in the discipline.
Lycorma delicatula White, commonly known as spotted lanternfly, is an invasive phloem-feeding insect which was first detected in the United States in 2014. It is now present in 19 states and is a serious pest in specialty crops (eg cultivated vineyards) and a nuisance pest in residential areas. Despite ongoing studies into methods for monitoring and biosurveillance, an effective attractant has not been identified. In this study, we recorded adult L. delicatula on the trunks of mature Ailanthus altissima (Mill.) from late August to mid-November in 2020, 2021, and 2023. We observed adults on mature trees from the ground to 2 m above every 2 wk between 7 AM and 11 AM, and recorded every aggregation, individual, and pair of L. delicatula present. For each aggregation, the number of adults (aggregation size), sex of each adult, aggregation area, distance from ground level, and notable behaviors were recorded (feeding, resting, walking up or down, courtship and mating). Our data on aggregation size, individuals, and male-female pairs found no evidence of a relationship with day of year beginning in September. However, male-female sex ratio approaches 1 in late September; reproductive behaviors occur around the same sex ratio. Feeding was the most frequently observed behavior within and between all years and continued throughout the mating and oviposition period. Our data indicate targeting the period when courtship and mating begin may provide opportunities for identification of novel olfactory or vibroacoustic stimuli for monitoring or biosurveillance.
Human activities are reshaping ecosystems worldwide, facilitating the spread of invasive insect species beyond their native ranges. The lesser banded hornet Vespa affinis Linnaeus (Vespidae:Vespa), a known predator of honeybees, poses risks to public health and apiculture outside its native range. However, whether suitable habitats for V. affinis overlap with human population hotspots and international ports are poorly understood. Here, we used an ensemble modeling approach to assess the probability of V. affinis establishment and invasion risks. Specifically, we considered global habitat suitability for this species under baseline (current) conditions (1970-2000) and in 2081-2100 (2090s) under the socioeconomic pathway (ssp585) scenario. Our models suggest that suitable areas for V. affinis will increase by over similar to 21% from now (2.06 & times; 107 km2) to the 2090s (2.49 & times; 107 km2). High-risk international ports are mainly concentrated in Africa, Southeast Asia, the Caribbean, and central parts of the Americas. Our findings also suggest there is high likelihood for human-hornet interactions and public health concerns in South and Southeast Asia, West Africa, and northern South America. These findings provide key insights into the hornet's potential distribution and risk areas supporting targeted monitoring efforts and the development of biosecurity strategies to mitigate its spread.
Understanding virus-vector-host interactions in the context of mixed infections is crucial to predict virus spread and develop effective control measures in economically important crops. The virus transmission process plays a central role in plant disease epidemiology and is particularly influenced by vector behavior and life-history traits. However, how mixed virus infections shape vector fitness and host preference remains poorly understood. Here, we determined the effects of double infections with the noncirculative cucumber mosaic virus (CMV, Cucumovirus) and the circulative cucurbit aphid-borne yellows virus (CABYV, Polerovirus) on the fitness and host plant preference of the aphid Aphis gossypii Glover on the host plant, melon (Cucumis melo L.). We revealed that aphids feeding on melon infected only with CMV (single-infected) weighed less and had lower fecundity than those feeding on uninfected plants or CABYV single- or double-infected plants. The mean relative growth rate and, consequently, the intrinsic rate of natural increase were also lower in aphids on CMV single-infected plants, indicating a negative impact of CMV single-infection on aphid fitness. In choice experiments, aphids were initially attracted to CABYV/CMV double-infected plants, but this was followed by a shift in host preference toward mock-inoculated plants over time. These findings suggest that mixed infections can modulate vector behavior in complex and dynamic ways. The pattern of early attraction followed by dispersal appears to promote CMV spread while having limited consequences for CABYV transmission, indicating an asymmetrical benefit between these 2 viruses.