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Neuroptera

Neuroptera is an order of holometabolous insects comprising approximately 5,800 described species worldwide, commonly known as lacewings, antlions, mantidflies, and their relatives, and characterized by two pairs of similar-sized, membranous wings with a dense network of veins that give them a lacy appearance.^[1]^[2] These insects are predominantly predatory throughout their life stages, with larvae that are active hunters using specialized mouthparts to capture and liquefy prey, and adults that often feed on other insects, nectar, or pollen.^[2]^[3] The adults of Neuroptera typically have soft bodies, long filiform antennae, large compound eyes, and chewing mouthparts, with wings held roof-like over the abdomen at rest; they are generally weak fliers and nocturnal or crepuscular in activity.^[2]^[4] Larvae exhibit a campodeiform body form—elongated, flattened, and sclerotized—with prominent sickle-shaped mandibles and maxillae forming pincers for grasping prey, and most species undergo complete metamorphosis, pupating in silken cocoons or soil.^[5]^[2] Eggs are often laid on stalks to protect them from predators, a notable adaptation seen in many lacewing species.^[5] Neuroptera is classified into three suborders: Nevrorthiformia (spongeflies), Hemerobiiformia, which includes lacewings and mantidflies, and Myrmeleontiformia, encompassing antlions and owlflies, with a total of 16 extant families such as Chrysopidae (green lacewings), Myrmeleontidae (antlions), and Mantispidae (mantidflies).^[2]^[6]^[7] The order belongs to the superorder Neuropterida, alongside Megaloptera and Raphidioptera, and fossils indicate a much greater past diversity, with Neuroptera dating back to the Permian period.^[2]^[6] Ecologically, Neuroptera play a significant role as predators in terrestrial and, in some cases like Sisyridae, aquatic habitats, where larvae control populations of pests such as aphids, mites, and small arthropods.^[2]^[3] Many species, particularly green lacewings in the family Chrysopidae, are commercially reared and released as biological control agents in agriculture and horticulture to manage insect pests, reducing the need for chemical pesticides.^[8]^[2] Their presence also serves as an indicator of environmental health in natural and semi-natural ecosystems.^[9]

Classification and Diversity

Taxonomy

Neuroptera is an order of insects within the class Insecta, characterized by holometabolous development and wings with a dense network of veins, often described as net-veined. The order was originally described by Linnaeus in 1758 under the name "Neuroptera" in his Systema Naturae, encompassing a broad assemblage of insects with net-veined wings. Major taxonomic revisions followed, including Handlirsch's comprehensive work in 1906-1908, which incorporated fossil evidence to refine familial boundaries and phylogenetic relationships within the group. Subsequent efforts by Aspöck et al. in 1980 provided a detailed catalog of European species, emphasizing morphological characters for classification. Cladistic analyses by Aspöck et al. in 2001 further revised the taxonomy using 36 morphological characters from adults and larvae, supporting a monophyletic Neuroptera while questioning broader inclusions. In modern taxonomy, Neuroptera is divided into three suborders: Nevrorthiformia (spongillaflies), Myrmeleontiformia (including antlions and owlflies), and Hemerobiiformia (including green lacewings and mantidflies). Historically, the order included Raphidioptera (snakeflies) and sometimes Megaloptera (alderflies and dobsonflies) as suborders under a broader "Neuroptera sensu lato," but contemporary classifications recognize these as separate orders within the superorder Neuropterida, based on morphological and molecular evidence clarifying their distinct evolutionary lineages. Approximately 16 extant families are recognized, with more than 6,500 described species; notable examples include Chrysopidae (type genus Chrysopa) and Myrmeleontidae (type genus Myrmeleon).^[10]^[11] Taxonomic challenges persist, particularly regarding the monophyly of Neuroptera and the historical inclusion of Megaloptera, with debates centering on whether cladistic approaches fully resolve interfamilial relationships or if traditional morphology-based groupings better reflect evolutionary history. Recent phylogenomic studies post-2020, incorporating transcriptomic and genomic data, have integrated molecular evidence to affirm Neuroptera's monophyly and its position within Neuropterida, where it forms the sister group to Megaloptera, and the entire Neuropterida clade is sister to Coleoptera + Strepsiptera in Holometabola.^[12]

Major Families and Species

The order Neuroptera encompasses more than 6,500 described species distributed across 16 families, with the highest diversity concentrated in tropical and subtropical regions worldwide.^[10] This diversity reflects adaptations to varied habitats, from arid deserts to humid forests, though species richness declines toward polar and high-altitude zones.^[13] The suborder Myrmeleontiformia includes the small suborder Nevrorthiformia, represented by the single family Nevrorthidae (spongillaflies) with approximately 15–20 species, which are aquatic or semi-aquatic and restricted to specific regions like Europe, Asia, and Australia.^[14] Within the larger Myrmeleontiformia, the family Myrmeleontidae, commonly known as antlions, represents one of the most species-rich groups with approximately 1,700 species; their larvae are renowned for constructing pit traps in sandy soils to capture prey.^[15]^[13] The Ascalaphidae, or owlflys, comprise around 450 species characterized by large, conspicuous eyes and crepuscular, fast-flying adults that resemble dragonflies.^[13] Nemopteridae, the threadwings or spoonwings, include approximately 150 species distinguished by their dramatically elongated hindwings, which can exceed body length in some taxa.^[13] The suborder Hemerobiiformia features several prominent families, including Chrysopidae, the green lacewings, with roughly 1,400 species noted for their bright green coloration and predatory larvae that target aphids and other soft-bodied insects.^[13] Hemerobiidae, or brown lacewings, account for about 590 species; these smaller, less vividly colored insects have wings typically 4–10 mm long and are adapted to temperate woodland environments.^[13]^[16] Mantispidae, the mantidflies, encompass around 400 species with raptorial forelegs that mimic those of praying mantises, enabling them to grasp prey effectively.^[13] Coniopterygidae, known as dustywings, include approximately 460 species; these minute insects (2–5 mm) have wings dusted with a powdery coating and reduced venation.^[13]^[17] Although sometimes historically grouped with Neuroptera, the order Raphidioptera (snakeflies) is now recognized as a distinct lineage within the superorder Neuropterida due to differences in head structure and life history, and is excluded from modern Neuroptera classifications. Distribution patterns vary by family: many, such as Myrmeleontidae and Chrysopidae, exhibit pantropical ranges with extensions into temperate zones, while others like Nemopteridae are predominantly Old World arid-dwellers; notable endemism occurs in Australia, particularly in Psychopsidae with 26 species restricted to that continent.^[13] Conservation concerns affect certain taxa, including some Nemopteridae species classified as endangered due to habitat loss in arid ecosystems, such as those in southern Africa and China.^[18]

Morphology and Physiology

External Anatomy

Adult Neuroptera possess a tripartite body plan comprising a distinct head, thorax, and abdomen, with the overall form ranging from small and delicate to larger and more robust depending on the family. The head features large compound eyes that provide wide visual fields, and ocelli may be present or absent across suborders, such as their absence in Hemerobiiformia. Mouthparts are of the chewing type, adapted for nectar and pollen feeding in many adults.^[2]^[19] The thorax is typically short, with the prothorax often enlarged in families like Chrysopidae, supporting three pairs of legs that are generally ambulatory but show raptorial modifications in larvae rather than adults. Wings are a defining feature, with four membranous, net-veined structures of similar size, the costal vein thickened for reinforcement, and extensive branching venation including numerous crossveins and parallel radial sector branches that give the order its name ("nerve-winged"). Wing coupling occurs via a frenulum on the hind wing engaging a jugal fold or lobe on the forewing in many species, facilitating coordinated flight, while specific venation patterns like the recurrent humeral vein characterize families such as Chrysopidae.^[2]^[19]^[20] The abdomen lacks cerci and is segmented, with females typically bearing a short ovipositor for depositing stalked eggs, a common adaptation in many families for protecting eggs from predation. Sexual dimorphism is generally subtle, though males in some Ascalaphidae exhibit larger wings relative to body size, potentially aiding in mate attraction or display.^[21] Larval Neuroptera display diverse external morphologies, predominantly campodeiform—elongate, flattened bodies with well-developed thoracic legs and prognathous heads suited for active predation—though variations occur, such as more robust forms in some Myrmeleontidae with less mobile bodies. The head is prognathous with prominent sickle-shaped mandibles and maxillae that interlock to form piercing-sucking stylets for extraintestinal digestion of prey, alongside short antennae and stemmata for sensory input. Thoracic legs are cursorial with single-segmented tarsi and paired claws, while the abdomen is multi-segmented, often bearing tubercles or setae for camouflage or mobility, and lacks true cerci.^[2]^[22]^[19] Unique to larval Neuroptera is the production of silk from modified Malpighian tubules, extruded through anal spinnerets to form cocoons for pupation, rather than dedicated silk glands found in other insects. This silk originates as liquid in the tubules, passes through the hindgut, and solidifies upon extrusion, enabling construction of protective pupal cases in soil or silk.^[2]

Internal Systems and Sensory Biology

The digestive system of Neuroptera is adapted to their predominantly carnivorous diet, with larvae and many adults preying on small arthropods such as aphids and mites. The foregut includes a prominent crop for temporary food storage, as observed in species like Chrysoperla externa, where the crop forms the largest portion of the foregut and connects to the midgut via a stomodeal valve. The midgut serves as the primary site for enzyme secretion and nutrient absorption, featuring an enlarged sac-like structure in larvae; in antlion larvae (Myrmeleontidae), the midgut is notably discontinuous from the hindgut, preventing solid waste elimination until pupation, when accumulated feces are expelled as a meconial pellet. The hindgut facilitates water reabsorption and waste compaction, supporting efficient processing of liquefied prey contents injected via larval mandibles. Neuroptera possess an open circulatory system typical of insects, characterized by a hemocoel cavity where hemolymph bathes internal organs directly. The heart functions as a dorsal vessel, pumping hemolymph anteriorly through an aorta and posteriorly via ostia, distributing nutrients, hormones, and immune factors without distinct blood vessels. Respiratory exchange occurs through a tracheal system, with spiracles distributed along the thorax and abdomen for direct oxygen delivery to tissues; tracheal gills are absent in most families, though aquatic Sisyridae larvae feature abdominal gills for underwater respiration. This system supports high metabolic demands during predation and flight in adults. Reproductive physiology in Neuroptera aligns with their holometabolous development, involving complete metamorphosis from egg to larva, pupa, and adult. Mating often involves spermatophore transfer, where males deposit a nutrient-rich packet of sperm externally or internally, as seen in Coniopterygidae and Chrysopidae, facilitating indirect insemination and providing females with supplemental nutrition. Eggs develop with chorionic micropyles—specialized pores allowing sperm entry for fertilization—ensuring precise oviposition on vegetation or soil; females typically lay stalked eggs to deter predation. The nervous system comprises a centralized brain with prominent optic lobes for visual processing, connected to a subesophageal ganglion that integrates sensory inputs from mouthparts and coordinates feeding responses. This configuration enables rapid sensory integration for prey detection, with neural pathways linking antennal and visual cues to mandibular strikes in larvae. The ventral nerve cord extends posteriorly, supporting coordinated locomotion and predatory behaviors. Sensory organs are highly specialized for detecting prey and environmental cues. Antennae bear mechanoreceptors and chemosensory sensilla, including trichoid types sensitive to vibrations and pheromones, aiding in mate location and host detection. Compound eyes provide panoramic vision with ultraviolet (UV) sensitivity, particularly in dorsofrontal regions of Ascalaphidae (owlflies), where photoreceptors peak at approximately 350 nm for enhanced contrast in foliage. Some taxa possess auditory organs, such as tympanal structures in adults of Chrysopidae and Ascalaphidae, tuned to ultrasonic bat echolocation for evasion.^[23] Biochemical defenses include mucus or silk production for protection; antlion larvae spin silken cocoons during pupation, while some chrysopids secrete adhesive mucus from mandibular glands to deter attackers. Certain larvae produce alkaloids or neurotoxins in midgut fluids to immobilize prey or discourage predators, as in Myrmeleontidae, where injected venom contains paralytic compounds for predation deterrence.

Life History and Ecology

Life Cycle

Neuroptera undergo complete metamorphosis, consisting of four distinct life stages: egg, larva, pupa, and adult. This holometabolous development is characteristic of the order, with each stage adapted for specific functions such as protection, feeding, transformation, and reproduction.^[2] The egg stage begins with females laying eggs that are typically stalked or pedunculated to deter predation, often in clusters on vegetation or near prey sources; for example, in Chrysopidae, eggs are attached to slender stalks secreted by the female. Incubation lasts 5-20 days, depending on temperature, with optimal hatching at 25-30°C; eggs of green lacewings (Chrysopidae) hatch in 4-5 days under warm conditions. Predation risks are high during this vulnerable phase, prompting protective adaptations like the stalks.^[22]^[24]^[25] Larvae progress through three instars, emerging as active predators with piercing-sucking mouthparts formed by sickle-shaped mandibles and maxillae, which they use to feed on small arthropods like aphids. In some families, such as Mantispidae, hypermetamorphosis occurs, with the first instar being highly mobile (campodeiform) and later instars more sedentary or grub-like; however, most larvae, including those of antlions (Myrmeleontidae), maintain a consistent campodeiform shape across instars. Development duration varies widely: 2-3 weeks for lacewing larvae under optimal conditions, but up to 1-3 years for soil-dwelling antlion larvae, influenced by prey availability and temperature. Larvae lack a complete digestive system, accumulating waste as meconium that is expelled during pupation.^[2]^[22]^[26] The pupal stage is non-feeding and occurs within a silken cocoon spun by the larva using secretions from the Malpighian tubules, often in a soil cavity or attached to vegetation; the pupa is exarate (appendages free) and decticous (functional mandibles). This stage lasts 1-4 weeks, with lacewing pupae requiring 5-7 days and antlion pupae about 4 weeks, again modulated by temperature; ecdysis to the adult happens within the cocoon.^[2]^[22]^[24] Adults are short-lived, typically surviving days to weeks (e.g., 20-40 days for green lacewings at 75°F or 4-6 weeks for others), with a primary focus on reproduction and dispersal via flight; they feed on nectar, pollen, honeydew, or small prey. In temperate species, diapause may occur in adults or late larval stages to overwinter. Incomplete metamorphosis is absent in the order, and while parthenogenesis is rare, it has been noted in some Chrysopidae species. Developmental timing across stages is heavily influenced by temperature, with warmer conditions accelerating the cycle to as little as 4 weeks total for many species.^[25]^[24]^[22]^[27]

Habitat and Behavior

Neuroptera exhibit diverse habitat preferences, ranging from arid sandy environments favored by antlion larvae (Myrmeleontidae) that construct pit traps in dry, fine soils, to forested and shrubland areas preferred by lacewings (Chrysopidae), and even aquatic margins where some larvae develop.^[28]^[29]^[30] Species such as green lacewings occupy a broad altitudinal gradient, from sea level to over 4,000 meters in mountainous regions like the Shaluli Mountains in China, allowing adaptation to varied climatic conditions.^[31] Foraging behaviors in Neuroptera are predominantly predatory, with larvae employing ambush strategies; for instance, antlion larvae in the family Myrmeleontidae dig conical pits to trap ants and other small arthropods, while lacewing larvae actively stalk soft-bodied prey like aphids on vegetation.^[32]^[33] Adults, often weak fliers, engage in aerial hunting, detecting prey through visual and vibrational cues, and selectively targeting small insects such as aphids and mites, thereby playing a key role in food webs as mid-level predators.^[34] Mating involves courtship rituals featuring vibrational signals or "songs" produced by abdominal tremulation, particularly in Chrysopidae, with no evidence of nuptial gifts; females select oviposition sites near prey-rich patches, such as aphid-infested plants, to enhance larval survival.^[35]^[36] Most Neuroptera are solitary throughout their life stages, lacking eusociality, though some Chrysopidae larvae form loose aggregations that may provide mutual protection against predators.^[30] Dispersal is limited by their weak flight capabilities, but seasonal migrations occur in temperate zones, as seen in Chrysoperla carnea, which undertake flights to exploit breeding habitats or overwintering sites.^[37] To counter threats from predators like birds and spiders, Neuroptera employ adaptations such as camouflage; many larvae, especially in Chrysopidae, cover themselves with debris or plant material to blend into surroundings and avoid detection.^[38]^[39]

Evolutionary History

Fossil Record

The fossil record of Neuroptera extends back to the Late Permian (approximately 259–252 million years ago), with the earliest known specimens from Upper Permian deposits such as those in the Tunguska Basin, Siberia, including members of the family Permithonidae, which exhibit primitive wing venation patterns distinct from later forms. These fossils, preserved as compressions in fine-grained sediments, represent some of the oldest evidence of the order and suggest an origin tied to early terrestrial ecosystems recovering from the late Paleozoic extinction.^[40] Neuroptera underwent substantial diversification during the Mesozoic era, beginning with Triassic records that indicate initial radiation among early lacewing lineages. Late Triassic fossils from the Momonoki Formation in Japan include five new genera and species, highlighting the order's adaptation to diverse habitats in the aftermath of the Permian-Triassic mass extinction. This diversification accelerated in the Jurassic, particularly at the Karatau locality in Kazakhstan, where compression fossils reveal a rich assemblage of families such as Osmylidae and extinct relatives like Archeosmylidae and Saucrosmylidae, with over 30 species documented from Upper Jurassic strata. In the Cretaceous, the order reached a peak of diversity, with numerous species—exceeding 100 across global deposits—preserved in ambers like those from Myanmar (Burmese amber) and Lebanon; Burmese amber, in particular, yields early Chrysopidae-like forms alongside thorny lacewings (Rhachiberothidae) and other lineages bridging Mesozoic and modern faunas. Recent discoveries, including a venomous larva from Late Cretaceous Kachin amber (as of 2024), continue to reveal details of larval adaptations. Key Cretaceous sites, such as Lebanese amber (Early Cretaceous, ~130–125 million years ago), contribute additional species, including Coniopterygidae and mantidflies, underscoring a period of morphological experimentation and ecological expansion.^[41]^[42]^[43]^[44]^[45]^[46]^[47] The Cenozoic record shows continued prominence in the Eocene and Oligocene, with Baltic amber from Europe preserving diverse taxa such as Hemerobiidae (brown lacewings) and Osmylidae, reflecting dominance in temperate forest ecosystems. In North America, the early Eocene Green River Formation yields fossils of Protosmylinae (Osmylidae relatives) and other forms, including giant lacewings, indicating persistence of Mesozoic lineages into lacustrine environments. Recent findings include a new genus of mantidflies from the earliest Eocene Fur Formation, Denmark (as of 2025). Some Neuroptera lineages experienced declines and local extinctions following the Pleistocene, linked to climatic shifts and habitat fragmentation, though the order as a whole survived into the present. Extinct families like Kalligrammatidae, known primarily from Jurassic compressions in China and Kazakhstan, are notable for their butterfly-like venation and large size, representing a specialized Mesozoic radiation. Preservation biases favor adult wings in compression fossils from sedimentary rocks, while soft-bodied larvae—critical to understanding life cycles—are rare, appearing mostly as exceptional amber inclusions due to their vulnerability to decay.^[48]^[49]^[50]^[51]^[52]^[53]

Phylogenetic Relationships

Neuroptera occupies a basal position within the Endopterygota, the holometabolous insects characterized by complete metamorphosis, where it forms a sister group to the clade comprising Coleoptera (beetles) and Strepsiptera (stylopids). This relationship is supported by analyses of 18S rRNA sequences and expanded in comprehensive phylogenomic studies incorporating thousands of genes from transcriptomes and genomes, which place Neuroptera outside the larger Hymenopteran-Dipteran-Lepidopteran clade. Recent updates using anchored hybrid enrichment data and improved models for compositional heterogeneity continue to affirm this topology, with high bootstrap support exceeding 90% for the Neuroptera-(Coleoptera + Strepsiptera) clade.^[54] Internally, the order is divided into two primary suborders: Myrmeleontiformia, which is positioned basally alongside the small Nevrorthiformia, and the more derived Hemerobiiformia.^[55] Myrmeleontiformia includes families like Myrmeleontidae (antlions) and Ascalaphidae (owlflies), characterized by predatory lifestyles and complex larval traps, while Hemerobiiformia encompasses Chrysopidae (green lacewings) and Hemerobiidae (brown lacewings), often with more generalized predation.^[56] Debates persist regarding the placement of Mantispidae (mantidflies) within Hemerobiiformia; molecular and morphological data variably position it as sister to Chrysopidae or within a subclade including Berothidae and Rhachiberothidae, reflecting uncertainties in raptorial leg evolution and host associations.^[14] Key character evolution in Neuroptera involves progressive reductions in wing venation, from the reticulate, primitive patterns in basal lineages like Nevrorthidae to simplified networks in derived Hemerobiiformia, facilitating flight efficiency in diverse habitats.^[57] Unlike sister orders in Neuropterida, such as Megaloptera with aquatic larvae, Neuroptera largely lost aquatic larval stages early in its evolution, shifting to terrestrial predation; only relict basal families like Nevrorthidae retain aquatic forms, highlighting a transition to soil and foliage-based lifestyles.^[56] Molecular evidence underpinning these relationships derives from nuclear genes including CAD (carbamoyl-phosphate synthetase) and wingless, which provide robust support for major clades in multi-gene phylogenies, with recent transcriptomic analyses yielding bootstrap values over 90% for internal branches.^[58] Controversies in Neuropterida phylogeny center on the exclusion of Raphidioptera (snakeflies), whose inclusion was once proposed but rejected by morphological synapomorphies like egg burrowing and wing coupling, favoring instead a Neuropterida clade uniting Neuroptera and Megaloptera based on shared larval trunk-like gills and aquatic adaptations.^[55] Cladistic analyses consistently support Raphidioptera as sister to this Neuropterida clade, resolving earlier ambiguities through integrated genomic data.^[57] The origin of Neuroptera is estimated at approximately 280 million years ago in the early Permian, with stem-group fossils indicating divergence near the Carboniferous-Permian boundary, though definitive crown-group records appear post-Carboniferous.^[59] Major radiations occurred after the end-Permian mass extinction around 252 million years ago, coinciding with the diversification of terrestrial ecosystems and the proliferation of angiosperms in the Mesozoic, driving family-level expansions in both suborders.

Human Interactions

Biological Control and Agriculture

Neuroptera, particularly species in the family Chrysopidae, serve as key agents in biological control due to their predatory larvae, which target aphids and other soft-bodied insect pests in agricultural settings. Green lacewing larvae, such as those of Chrysoperla carnea and C. rufilabris, are generalist predators that consume up to 50 aphids per day, making them effective against a wide range of agricultural pests including mealybugs and mites.^[60] ^[61] Commercial mass rearing of these species began in the late 20th century, with C. carnea being released inundatively in greenhouses across Europe and North America since the 1970s to suppress aphid outbreaks on crops like vegetables and ornamentals.^[62] ^[63] In integrated pest management (IPM) programs, Neuroptera contribute to sustainable agriculture by reducing reliance on chemical pesticides, with demonstrated efficacy against cotton pests such as Heliothis species through larval predation in field crops. Their economic value is embedded within the broader biocontrol market, where natural enemies like lacewings help generate annual benefits estimated at $4.5 billion (as of 2006) for U.S. crop protection alone, supporting cost-effective pest suppression with high benefit-to-cost ratios for classical biological control exceeding 1:250, while augmentative releases show ratios similar to insecticides (around 1:32).^[64] ^[65] ^[66] The global biocontrol agents market, including Neuroptera, was valued at USD 4.01 billion in 2025 and is projected to reach USD 5.62 billion by 2030.^[67] Mass rearing for biocontrol faces challenges, including high rates of cannibalism among C. carnea larvae, which can limit production efficiency when conspecific eggs or young are present, and sensitivity to insecticides like chlorantraniliprole and lambda-cyhalothrin, requiring selective pesticide use to preserve predator populations. Augmentation strategies, such as timed inundative releases, mitigate these issues by introducing larvae directly into pest hotspots while avoiding broad-spectrum sprays.^[68] ^[69] ^[70] The environmental impacts of Neuroptera in agriculture are largely beneficial, with minimal non-target effects on beneficial insects and a role in enhancing biodiversity; for instance, conservation practices in olive orchards increase lacewing abundance, promoting ecosystem stability without disrupting pollinators or parasitoids. In European vineyards, green lacewings have been integrated into IPM to control aphids, reducing pesticide applications and supporting predatory communities, while in U.S. citrus groves, their releases aid in managing citrus aphids, contributing to sustainable grove management.^[71] ^[72] ^[29] Post-2015 studies have focused on selecting genetic strains of C. carnea for enhanced traits, such as insecticide resistance that boosts overall fitness and predatory potential without fitness costs, improving their performance in biocontrol programs. As of 2024, selections for resistance to insecticides like spirotetramat have shown 47-fold increases without fitness costs.^[73] ^[74] ^[75] Emerging research in the 2020s explores CRISPR-Cas9 editing in non-model insects to engineer traits like increased predation efficiency.^[76]

Cultural and Symbolic Significance

In various cultures, antlions (family Myrmeleontidae) have appeared in folklore and mythology, often symbolizing patience and cunning due to the predatory pit-building behavior of their larvae. In North American traditions, the antlion larva is commonly known as a "doodlebug," and children recite charms such as "Doodlebug, doodlebug, fly away home, your house is on fire and your children are gone" to coax it from its burrow, reflecting a playful interaction with its hidden, waiting nature.^[77] This folklore highlights the insect's patient ambush strategy, evoking themes of perseverance in the face of adversity. The mythical ant-lion, a hybrid creature with a lion's face and an ant's body, features prominently in medieval European bestiaries, where it walks backward to conceal its tracks and avoid detection, symbolizing deceit or hypocrisy.^[78] Derived from classical texts and biblical references to the "mirmecoleon" (ant-lion), this beast was interpreted allegorically as a representation of the devil luring souls into temptation, underscoring themes of moral vigilance.^[79] Such depictions influenced later literary works, including Jorge Luis Borges' essays exploring the creature's evolution from myth to entomological reality. Their ephemeral adult lifespan and graceful flight contribute to broader symbolic interpretations of fragility and renewal across cultures, often contrasting the predatory prowess of their larvae. In literature, Neuroptera have inspired entomological narratives since classical antiquity, with antlions noted for their predatory habits in ancient texts.^[79] The 19th-century naturalist Jean-Henri Fabre detailed observations of antlions in his Souvenirs Entomologiques, portraying their life cycles as marvels of instinct and adaptation, influencing popular perceptions of insects as subjects of wonder rather than mere pests. During the Victorian era, Neuroptera specimens were prized in entomology collections, featured in cabinets of curiosity alongside other insects as symbols of scientific progress and aesthetic beauty.^[80] The British Museum's 1852 catalogue of neuropterous insects exemplifies this enthusiasm, with mounted lacewings and antlions displayed to educate and captivate audiences amid the era's fascination with natural history. In contemporary contexts, Neuroptera appear in eco-art and conservation efforts, where lacewings are depicted as icons of biodiversity and natural pest control in murals and installations addressing environmental decline.^[81] These representations emphasize their role in ecological balance, fostering public awareness of insect conservation in the face of habitat loss.

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Antlions prefer dry, fine-sand, warm environments, which are optimal for pit construction. They are found in arid habitats.Missing: ecology | Show results with:ecology
[24]
Habitat Diversity Increases Chrysoperla carnea s.l. (Stephens, 1836 ...
Feb 12, 2024 · These habitats included shrublands, “montado,” grasslands, eucalyptus and pine forests, vineyards, and olive groves. The findings revealed C.
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General Information | SpringerLink
Apr 20, 2017 · Some authors treat the taxon as an order encompassing three suborders: Megaloptera, Raphidioptera, and Planipennia. ... Raphidioptera, Megaloptera ...
[26]
Elevational Diversity Patterns of Green Lacewings (Neuroptera
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[27]
[PDF] Preference of antlion and wormlion larvae (Neuroptera ...
Mar 20, 2015 · Larvae of antlions (Neuroptera: Myrmeleontidae) are predators living in a variety of habitats, ranging from ar- boreal and rocky habitats, ...<|control11|><|separator|>
[28]
Sedentary antlion larvae (Neuroptera: Myrmeleontidae) use ... - NIH
Specifically, antlion larvae rapidly learn to differentiate between the vibrational cues associated with prey of different sizes, and they save resources by ...Missing: characteristics | Show results with:characteristics
[29]
Divergent venom effectors correlate with ecological niche in ... - Nature
Aug 13, 2024 · Our results indicate that molecular venom evolution plays a role in the adaptation of antlions to their unique ecological niche.
[30]
THE ROLE OF COURTSHIP SONGS IN REPRODUCTIVE ...
Male and female lacewings tremulate during courtship, establishing duets that always precede copulation. Three distinct courtship songs are found in populations ...
[31]
Acoustical Communication during Courtship and Mating in the ...
Courtship and mating of the green lacewing Chrysopa carnea Stephens is described. Copulation is preceded by 5 well defined activities: search, antennal contact, ...
[32]
Abundance and Seasonal Migration Patterns of Green Lacewings ...
May 1, 2024 · Many insects, including green lacewings, migrate seasonally to exploit suitable breeding and winter habitats. Green lacewings are important ...
[33]
Early evolution and ecology of camouflage in insects - PNAS
Dec 26, 2012 · This trash packet camouflages the larva, preventing de- tection by predators and prey and constituting a defensive shield in instances where the ...
[34]
Neuropterans - Encyclopedia of Arkansas
Sep 16, 2021 · There are three suborders (Osmyloidea, Hemerobiiformia, and Myrmeleontiformia) and nine extinct and fifteen extant families. There are about ...
[35]
(PDF) The Lower Permian Insects of Kansas. Pan 12. Protorthoptera ...
Aug 10, 2025 · The Lower Permian Insects of Kansas. Pan 12. Protorthoptera (continued), Neuroptera, Additional Palaeodictyoptera, and Families of Uncertain ...<|separator|>
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Late Triassic lacewings (Insecta: Neuroptera) from Japan
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[37]
Jurassic beaded lacewings (Insecta: Neuroptera: Berothidae) from ...
Feb 1, 2015 · auliensis sp. nov) are described from the Upper Jurassic of Karatau (Kazakhstan). Sinosmylites hotgoricus sp. nov. is described from the Upper ...
[38]
Lance lacewings of the world (Neuroptera: Archeosmylidae ...
Apr 9, 2019 · The genera of the lance lacewing family Osmylidae (Neuroptera) and extinct allied families (Archeosmylidae, Saucrosmylidae) are reviewed.
[39]
The Neuropterida from the mid-Cretaceous of Myanmar
A catalogue and a faunal analysis on Neuropterida recorded from world amber deposits are provided. The Myanmar amber Neuropterida highlights the Cretaceous ...<|separator|>
[40]
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Mar 22, 2018 · The first green lacewing (Insecta: Neuroptera: Chrysopidae) from the mid-Cretaceous amber of Myanmar. XIUMEI LU; BO WANG; MICHAEL OHL ...
[41]
Lebanese amber: A time capsule from the dawn of modern ...
Sep 15, 2025 · To date, 286 biological inclusions (including Insecta, Archnida, Myriapoda and vertebrata) have been described so far from the Early Cretaceous ...
[42]
(PDF) A new genus of Hemerobiidae (Neuroptera) from Baltic amber ...
Aug 6, 2025 · PDF | A new genus and two new species of Hemerobiidae (Neuroptera) are described from the late Eocene Baltic amber, i.e., ...
[43]
[PDF] New Protosmylinae (Neuroptera: Osmylidae) from the early Eocene ...
May 31, 2021 · Key words: Neuroptera, Osmylidae, Protosmylinae, Okanagan Highlands, Green River Formation, Baltic amber,. Eocene. Introduction. The extant ...
[44]
Quantitative analysis of lacewing larvae over more than 100 million ...
Apr 14, 2023 · Overview on morphological diversity of lacewing larvae from the Cretaceous to today, illustrated with selected larval morphologies. Several ...
[45]
(PDF) Two New Species of Kalligrammatidae (Neuroptera) From the ...
Aug 5, 2025 · Two new fossil species of Kalligrammatidae (Neuroptera) are described from the Middle Jurassic locality at Daohugou, Inner Mongolia, ...
[46]
Diverse Cretaceous larvae reveal the evolutionary and behavioural ...
Aug 22, 2018 · Mostly soil dwellers with a soft cuticle, their larvae fossilize only as amber inclusions, and thus their fossil record is remarkably sparse.
[47]
Improved modelling of compositional heterogeneity reconciles ...
Feb 28, 2023 · Recent phylogenomic studies of Neuropterida based on mitogenomes, anchored hybrid enrichment (AHE) data, and transcriptomes have yielded a well- ...
[48]
The phylogeny of the Neuropterida: long lasting and current ...
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Evolution of lacewings and allied orders using anchored ...
Nov 23, 2017 · A phylogeny of Neuropterida is presented using anchored hybrid enrichment genomic data for 136 species across all families.<|separator|>
[50]
The Mitochondrial Genomes of Neuropteridan Insects and ... - NIH
Feb 1, 2019 · Based on the morphological characters, the taxon Neuroptera is divided into three suborders Nevrorthiformia, Myrmeleontiformia and ...
[51]
Phylogeny and Evolution of Neuropterida: Where Have Wings of ...
Abstract. The last 25 years of phylogenetic investigation into the three orders con- stituting the superorder Neuropterida—Raphidioptera, Megaloptera, and.<|control11|><|separator|>
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Phylogeny of the Neuropterida: a first molecular approach - 2004
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(PDF) Molecular Phylogeny of Neuropterida with Emphasis on the ...
Systematic Biology 46: 1–68. Neuropterida. Understanding the phylogeny of this basal holometabolan. superorder comprising the orders Raphidioptera,. Megaloptera ...<|control11|><|separator|>
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Chrysoperla (=Chrysopa) carnea, C. rufilabris - Biological Control
Green lacewings are used for biological control, with larvae being active predators, especially of aphids, and are considered generalist beneficials.
[55]
Chrysopa-System - Aphid control - Biobest
Feeds on most agriculturally important aphid species - including large prey ; Highly voracious – Chrysopherla carnea larvae feed on up to 50 aphids a day ...<|separator|>
[56]
Chrysopidae) used for biological pest control - ScienceDirect.com
Today, both Chrysoperla carnea (Stephens) and C. rufilabris (Burmeister) are mass-produced for aphid control on various crops in Europe and North America (van ...
[57]
Green Lacewings: Biological Control Agents of Greenhouse Insect ...
Jun 1, 2024 · Green lacewings are predatory insects used to control pests in greenhouses. Their larvae eat aphids, mealybugs, mites, and other pests. They ...
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[PDF] Neuroptera and Biological Control (Neuropterida) - Zobodat
Neuroptera, mainly lacewings, are used in biological control as predators of pests, especially in field crops, and are considered 'aphido-phagous'.
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(PDF) Economic Value of Biological Control in Integrated Pest ...
Here we discuss approaches and methods available for valuation of biological control of arthropod pests by arthropod natural enemies and summarize economic ...
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Biological control and sustainable food production - Journals
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Conspecific neighbors and kinship influence egg cannibalism in the ...
Nov 1, 2021 · Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) is an illustrious predator that performs cannibalism upon facing small and defenseless conspecifics.
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The Effects of Alternative Foods on Life History and Cannibalism of ...
Suitable foods are essential for the successful mass rearing of natural enemies and can directly affect their quality and performance, which are the ...Missing: challenges | Show results with:challenges
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[PDF] Comparing Effects of Insecticides on Two Green Lacewings Species ...
ABSTRACT This study compared lethal and sublethal effects of ﬁve insecticides, chlorantraniliprole, cyantraniliprole, spinetoram, novaluron, and lambda- ...
[64]
Agricultural Management Systems Affect the Green Lacewing ...
Agricultural Management Systems Affect the Green Lacewing Community (Neuroptera: Chrysopidae) in Olive Orchards in Southern Spain | Environmental Entomology | ...
[65]
Attraction of green lacewings (Neuroptera: Chrysopidae) to native ...
The expansion and intensification of agroecosystems have led to a decrease in insect population and diversity, resulting in a decline in ecosystem services such ...<|separator|>
[66]
Selection of the predator green lacewing Chrysoperla carnea for ...
Oct 22, 2025 · Instead of harmful effects, insecticide resistance increased the fitness and predatory potential of C. carnea. In a biological control program, ...Missing: post- | Show results with:post-
[67]
Resistance of green lacewing, Chrysoperla carnea (Stephens), to ...
Its excellent predatory potential against a wide range of pests, high searching aptitude, vast geo-graphical distribution and enhanced field adaptability than ...
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CRISPR-Cas9 Techniques for Editing Non-Model Insects
Sep 5, 2025 · When applied to eukaryotic organisms, including insects, this technology allows for the targeted editing of genes with unprecedented accuracy.
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Folklore - Antlion Pit
The most common type of U.S. antlion folklore takes the form of a charm about the colloquial "doodlebug." Perhaps the most well-known example of a doodlebug ...Missing: Native American
[70]
Beasts : Ant-lion - Medieval Bestiary
Jun 18, 2024 · It is a beast that is the result of a mating between a lion and an ant. It has the face of a lion and and the body of an ant, with each part having its ...
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The Mermecolion: From Bible to Bestiary to Borges - Antlion Pit
The earliest antlions in literature were not the insects that we know today, but rather mythical animals that possessed qualities of both ants and lions.
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Lacewing Spiritual Meaning, Symbolism, and Totem (2025)
Jul 27, 2022 · Lacewings are considered to be a good omen by many Native American tribes. They often represent new beginnings, change, and transformation.
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Catalogue of the specimens of neuropterous insects in the collection ...
Mar 2, 2007 · Title. Catalogue of the specimens of neuropterous insects in the collection of the British museum. By. British Museum (Natural History).Missing: Victorian | Show results with:Victorian
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Insects in Art during an Age of Environmental Turmoil - PMC
May 9, 2022 · We surveyed relevant work by 73 artists and found a bias favoring insect art addressing habitat destruction or climate change.