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Neuropterida

Neuropterida is a superorder of holometabolous insects within the subclass Endopterygota, encompassing the three extant orders Raphidioptera (snakeflies), Megaloptera (alderflies and dobsonflies), and Neuroptera (lacewings and allies).^[1]^[2]^[3] This clade, comprising over 7,800 described species, represents a diverse group with a global distribution, excluding Antarctica, and is recognized for its predatory lifestyles and ecological roles in biological control.^[4]^[3] Adults of Neuropterida are typically characterized by large, membranous wings with extensive venation forming a net-like pattern, often held roof-like at rest, and chewing mouthparts adapted for predation or pollen feeding; body sizes vary widely, from small forms under 5 mm to large dobsonflies exceeding 150 mm in wingspan.^[3] Larvae exhibit a campodeiform (elongate, flattened) body plan with prognathous heads and are predominantly predaceous, feeding on other arthropods via specialized mouthparts that form a sucking tube; larval habitats range from terrestrial (most Neuroptera and Raphidioptera) to aquatic (all Megaloptera and select Neuroptera families like Sisyridae).^[2]^[3] The group undergoes complete metamorphosis, featuring pharate pupae enclosed in silken cocoons or leaf litter, with adults emerging to continue predatory behaviors.^[2] Neuropterida occupies a near-basal phylogenetic position among holometabolous insects, serving as the sister group to the clade comprising Coleoptera (beetles) and Strepsiptera (stylopids), with fossil records extending back to the Permian period and significant diversification during the Mesozoic.^[1]^[3] Ecologically, the superorder plays a key role in pest management, particularly through Neuroptera families like Chrysopidae (green lacewings), which are widely used in integrated pest management programs against aphids and other soft-bodied insects.^[3] Diversity is highest in tropical and subtropical regions, with major centers in the Oriental and Afrotropical realms, reflecting ongoing evolutionary radiations in families such as Myrmeleontidae (antlions) and Chrysopidae.^[3]

Systematics

Taxonomy

Neuropterida is a superorder of holometabolous insects classified within the subclass Pterygota and the infraclass Holometabola, encompassing three extant orders: Neuroptera, Megaloptera, and Raphidioptera.^[3] The order Neuroptera includes approximately 7,000 species, Megaloptera about 400 species, and Raphidioptera around 260 species, for a total of over 7,800 described species worldwide (as of 2024).^[4] Within Neuroptera, major families include Chrysopidae (approximately 2,000 species), known for green lacewings used in biological control, and Myrmeleontidae (approximately 2,000 species), which encompasses antlions.^[5] Megaloptera is represented primarily by Sialidae (approximately 80 species, alderflies) and Corydalidae (approximately 320 species, including dobsonflies and fishflies).^[6] Raphidioptera consists of two families: Raphidiidae (approximately 220 species, snakeflies) and Inocelliidae (approximately 40 species).^[7] The superorder Neuropterida was originally proposed in the early 20th century to unite these orders based on shared wing and larval traits, with historical classifications treating them as suborders under a broad Neuroptera; it formerly encompassed Mecoptera in some schemes like Mecopterida until the latter was reclassified within Mecopteroidea.^[1]^[3] These species counts are dynamic and updated regularly in catalogs such as the Lacewing Digital Library. Neuropterida is defined by synapomorphies including complete metamorphosis (holometaboly) and specific wing venation patterns featuring numerous crossveins that create a reticulate, lace-like structure.^[3]^[8]

Phylogeny

Neuropterida constitutes a monophyletic clade within the subclass Holometabola, positioned in the subgroup Neuropteroidea, with phylogenomic analyses indicating it as the sister group to Coleopterida (encompassing Coleoptera and Strepsiptera).^[8] This placement is supported by transcriptome-based studies integrating morphological data, highlighting shared evolutionary innovations such as complete metamorphosis and endopterygous development. Within Neuropterida, cladistic analyses consistently recover the monophyly of its three extant orders, with Raphidioptera forming the sister group to the clade (Megaloptera + Neuroptera). This topology is corroborated by phylogenomic investigations utilizing transcriptomes from 92 Neuropterida species, alongside broader sampling from 141 holometabolous taxa, which employed maximum likelihood and Bayesian inference to resolve internal relationships with high support.^[8] Key synapomorphies defining the clade include a reduced complement of four Malpighian tubules (compared to higher numbers in some basal holometabolans) and distinctive larval head structures, such as a prognathous, flattened capsule with a prominent gula and specialized hypopharyngeal sclerites adapted for predatory feeding.^[9] Phylogenetic controversies persist regarding the precise branching order among the orders, with alternative hypotheses—such as (Megaloptera + Raphidioptera) sister to Neuroptera—emerging from some mitogenomic and anchored hybrid enrichment datasets.^[10] A 2023 reanalysis of conflicting phylogenomic data (including transcriptomes and anchored loci from over 100 taxa) using site-heterogeneous models of compositional heterogeneity reconciled these discrepancies in favor of Raphidioptera sister to (Megaloptera + Neuroptera), underscoring the impact of modeling assumptions on inference.^[11] Additionally, extinct taxa like Glosselytrodea, known from Permian to Jurassic deposits, play a crucial role in clarifying deeper nodes by potentially representing a stem-group to Neuropterida or an early-diverging offshoot, providing morphological evidence for ancestral wing venation and genitalic traits that bridge gaps in the fossil record.^[12]

Morphology

Adult features

Adult Neuropterida are characterized by their elongate, soft-bodied form, ranging from about 2 mm to 75 mm in body length, with a well-developed head featuring large compound eyes that provide wide-field vision essential for predation or navigation.^[13]^[3] The mouthparts are mandibulate, adapted for chewing solid prey or imbibing nectar and pollen, though some species exhibit raptorial forelegs for capturing insects.^[14] Antennae are generally long and multisegmented, either filiform (thread-like) or pectinate (comb-like) in some taxa, bearing chemosensory sensilla that detect pheromones and host volatiles for mate location and prey detection.^[15] Ocelli are variably present or absent across the group, aiding in light detection where developed.^[16] The wings dominate adult morphology, consisting of two pairs of large, membranous structures with dense, net-like venation forming a characteristic lace-like pattern; for instance, Neuroptera often feature numerous costal veinlets along the leading edge.^[17] Wingspans range from 1 to 15 cm, with the fore- and hindwings similar in size and held roof-like over the abdomen at rest.^[3] Flight is generally weak due to underdeveloped muscles, resulting in clumsy, fluttering motion, and most species are nocturnal or crepuscular, minimizing exposure to diurnal predators.^[18] Order-specific traits further distinguish adults within Neuropterida. In Neuroptera, the prothorax is reduced and membranous, contributing to a compact thoracic profile.^[14] Megaloptera exhibit a more robust build, reflecting adaptations from their aquatic larvae, with sclerotized thoracic nota and projecting mandibles.^[16] Raphidioptera are notable for their elongated prothorax, which forms a flexible "neck" allowing greater head mobility, paired with a prognathous head and prominent pterostigma on the wings.^[19]

Larval features

Larvae of Neuropterida exhibit a predominantly campodeiform body form, characterized by an elongate, dorsoventrally flattened shape with well-developed thoracic legs for active locomotion and a heavily sclerotized head capsule housing sensory structures and mouthparts.^[3] This morphology contrasts with the winged adults and supports a predatory lifestyle, where larvae crawl through soil, litter, bark, or aquatic substrates to ambush prey. In some Neuroptera families, such as Ithonidae and Polystoechotidae, older instars adopt a more eruciform, grub-like form with a C-shaped body, reduced leg functionality, and less elongation, resembling scarab beetle larvae.^[20] Predatory adaptations are prominent, featuring powerful, sclerotized mandibles that form a specialized piercing-sucking apparatus in conjunction with the maxillae. In Neuroptera, these mandibles are typically hollow with internal venom canals, allowing larvae to inject liquefying enzymes into prey and extract fluids, enabling predation on small arthropods like aphids or other insects.^[3] Certain Neuroptera, including antlions (Myrmeleontidae), possess silk glands associated with Malpighian tubules, which produce silk for constructing pit traps in sandy substrates to capture prey via vibrational cues.^[3] Megaloptera and Raphidioptera larvae retain simpler chewing mouthparts suited for solid food consumption, lacking the advanced stylets seen in Neuroptera.^[2] Order-specific traits reflect diverse habitats and lifestyles within Neuropterida. Megaloptera larvae are aquatic, bearing lateral abdominal gills for respiration and often exhibiting flattened heads with closed stigmata, adapted for life in streams and rivers as active predators.^[2] Neuroptera larvae are generally terrestrial or semi-aquatic, with exceptions like Sisyridae (bearing gills in later instars for sponge-dwelling) and Nevrorthidae (lacking gills but using cuticular respiration); many possess abdominal adhesive structures or scoli for camouflage or mobility in litter.^[2] Raphidioptera larvae are terrestrial and corticolous, with an elongated body, compact head, and lithopsid-like camouflage on tree bark, facilitating predation in arboreal microhabitats.^[3] Larval sizes range from approximately 2 mm in early instars to 50 mm in mature forms, varying by order and family; for instance, Megaloptera like Corydalidae can reach up to 50 mm, while smaller Neuroptera such as Ithonidae top out at around 10-15 mm in head width equivalents.^[20]^[3]

Biology

Life cycle

Neuropterida undergo holometabolous (complete) metamorphosis, characterized by four distinct life stages: egg, larva, pupa, and adult, with profound morphological changes between the larval and adult forms. The egg stage typically lasts from a few days to several weeks, depending on species and environmental conditions; females lay eggs in clusters or singly, often on vegetation or near water.^[21] The larval stage is the longest in the life cycle for most Neuropterida, comprising multiple instars and often occupying 1–3 years, though durations vary by order and species. In Megaloptera, such as dobsonflies, larvae (known as hellgrammites) undergo 10–14 instars and may persist for up to 5 years in cooler aquatic environments.^[22] Raphidioptera larvae typically complete 9–11 instars over 2–3 years in terrestrial habitats under bark or litter, while Neuroptera larvae generally have 3 instars (up to 5 in some families like Ithonidae), lasting weeks to months in predatory species like lacewings but extending to years in larger forms. Larvae are active predators, with body forms ranging from campodeiform (elongate, mobile) to eruciform (scarab-like). Pupation follows, lasting 1–4 weeks; Neuroptera pupae form silken cocoons, whereas Megaloptera and Raphidioptera construct soil or bark cells, often requiring a chilling period to initiate in Raphidioptera.^[23] Adults emerge short-lived, typically surviving days to weeks, though some Megaloptera and Raphidioptera may persist for months in captivity. Development is temperature-dependent, with warmer conditions accelerating progression through stages, while cooler temperatures prolong larval durations. Many species enter diapause as larvae or pupae to overwinter, induced by short photoperiods in Neuroptera like Chrysopa lacewings, allowing survival through unfavorable seasons.^[24] In Raphidioptera, low temperatures are essential to break larval diapause and trigger pupation.^[21] A unique feature in some Neuroptera families, such as Mantispidae, is hypermetamorphosis, where the first larval instar is highly mobile and planidial (adapted for host-seeking), contrasting with more sedentary later instars that resemble scarab grubs. This specialization aids in parasitoid or kleptoparasitic lifestyles but is absent in Megaloptera and Raphidioptera.^[25]

Reproduction and development

Reproduction in Neuropterida involves diverse courtship rituals adapted to aerial or terrestrial environments, facilitating mate location and recognition across the orders Neuroptera, Megaloptera, and Raphidioptera. In many Neuroptera, such as green lacewings (Chrysopidae), courtship is mediated by vibrational signals produced through tremulation of the body, where males and females exchange duet-like songs to synchronize mating, often lasting several minutes before copulation.^[26] Wing displays, including fluttering and elevation, accompany these vibrations, as observed in dustywings (Coniopterygidae), where both sexes perform precopulatory abdominal rotations to signal readiness.^[27] Pheromones contribute to long-range attraction in select Neuroptera species, with males releasing volatile compounds to draw conspecific females.^[28] In Megaloptera, courtship behaviors vary by family; in Sialidae, it involves reciprocal wing fanning and abdominal bending, while in Corydalidae, males display by placing their mandibles perpendicularly on the female's wings.^[29]^[30] Internal fertilization predominates, with sperm transfer occurring directly via the male genitalia in most cases, though some Megaloptera species utilize spermatophores—gelatinous packets externally attached to the female during copulation.^[31] This mechanism ensures sperm delivery while allowing males to engage in post-copulatory mate guarding, where they remain attached to the female to deter rival inseminations, enhancing paternity assurance.^[31] Multiple matings are common, particularly in females, supporting sustained egg production over their short adult lifespan. Oviposition strategies emphasize egg protection from predation and environmental hazards, reflecting the predatory nature of larvae. In Neuroptera, females often deposit eggs singly atop elongated silken stalks extruded from the ovipositor, elevating them above foliage to minimize cannibalism by hatching larvae; this is prominent in Chrysopidae, where stalks reach up to 1 cm in length.^[32] Clusters without stalks occur in other families, sometimes coated in gelatinous secretions for added defense. Megaloptera females lay hundreds to thousands of eggs in compact masses on vegetation or rocks overhanging aquatic habitats, frequently enveloping them in a protective gelatinous layer secreted by accessory glands to deter oophagous predators.^[33]^[34] In Raphidioptera, eggs are inserted singly or in small groups into bark fissures or litter using a robust ovipositor, providing concealment without additional coatings.^[35] Parental care remains minimal throughout Neuropterida, with adults typically dispersing after oviposition to focus on feeding or further mating. Exceptions include limited post-copulatory guarding by Megaloptera males and occasional female attendance near egg masses in some species, though no prolonged provisioning or defense occurs. Population sex ratios are generally balanced near 1:1, but female-biased distortions arise in certain Neuroptera due to endosymbiont infections like Rickettsia, which induce male-killing and elevate female proportions to over 90% in affected populations.^[36] Developmental anomalies such as parthenogenesis are exceedingly rare in Neuropterida, with no confirmed cases of thelytoky or arrhenotoky leading to viable offspring; instead, sexual reproduction via outcrossing predominates to sustain genetic diversity amid high larval mortality.^[37]

Ecology

Habitats and distribution

Neuropterida exhibit a cosmopolitan distribution across all continents except Antarctica, encompassing approximately 6,430 extant species. The superorder achieves its highest diversity in tropical regions, where favorable climatic conditions support prolific speciation, particularly within Neuroptera, which comprises the vast majority—around 6,000 species—of the group's total. Polar regions lack representation due to the orders' preferences for temperate to subtropical environments, though some species extend into cooler temperate zones.^[38]^[39]^[40] Habitat preferences differ markedly among the constituent orders, reflecting adaptations to specific ecological niches. Neuroptera are primarily terrestrial, with larvae commonly occupying riparian zones, forest floor litter, and soil substrates where they prey on small arthropods. In contrast, Megaloptera larvae are aquatic, inhabiting clean, flowing streams, rivers, and lentic waters such as ponds and marshes, often in forested watersheds. Raphidioptera favor arboreal settings, with both adults and larvae residing on tree trunks, bark, and foliage in woodland environments. Across Neuropterida, many species are arboreal overall, while others thrive in arid, eremial habitats like deserts and steppes.^[41]^[42]^[21] Biogeographic patterns underscore regional specializations within Neuropterida. Raphidioptera demonstrate Holarctic dominance, with nearly all ~240 species confined to temperate forests of the Northern Hemisphere, including Europe, North America, and Asia. Megaloptera show elevated diversity in the Neotropics, alongside the Indomalayan region, where genera like those in Corydalidae exhibit disjunct distributions across South and Central America. Australia hosts numerous endemics, particularly antlions (Myrmeleontidae) within Neuroptera, with over 90% of the country's ~600 Neuropteran species unique to the continent and adapted to its arid and sclerophyllous landscapes.^[43]^[44]^[45] Contemporary threats to Neuropterida include habitat loss, which disproportionately impacts aquatic Megaloptera through degradation of riparian and freshwater ecosystems via deforestation and urbanization. Climate change exacerbates these pressures by inducing range shifts, as warming temperatures may displace species from historical temperate distributions toward higher latitudes or elevations, potentially disrupting local assemblages.^[46]^[47]

Feeding and behavior

Neuropterida larvae are predominantly predatory, employing specialized mouthparts to capture and subdue arthropod prey such as aphids, small insects, and other invertebrates. They inject venom and digestive enzymes through their hollow mandibles, which liquefy the prey's internal tissues for extraintestinal digestion, allowing the larvae to suck up the resulting fluid.^[48] This predatory strategy is common across the clade, with larvae of families like Myrmeleontidae (antlions) and Chrysopidae (lacewings) exhibiting fierce hunting behaviors adapted to their environments.^[49] Adult Neuropterida display more varied feeding habits, often shifting from predation to phytophagy. In Chrysopidae, such as green lacewings, adults primarily consume pollen, nectar, and honeydew from plants, though some species opportunistically prey on soft-bodied insects like aphids. This dietary flexibility supports reproduction, as nectar and pollen provide essential nutrients for egg production, contrasting with the strictly carnivorous larvae.^[50] Predatory behaviors in Neuropterida emphasize ambush tactics, particularly among larvae. Antlion larvae (Myrmeleontidae) construct conical pits in loose sand or soil, positioning themselves at the bottom to detect and capture falling prey through substrate vibrations; this pit-building involves meticulous excavation and maintenance to optimize trap efficiency.^[51] Similarly, larvae of other groups, such as owlflies (Ascalaphidae) and some Chrysopidae, adopt sit-and-wait strategies on foliage or ground litter, relying on immobility and sensory cues for prey detection.^[52] Mating behaviors incorporate vibrational communication, especially in Neuroptera. Male green lacewings produce species-specific courtship songs by vibrating their bodies against substrates, generating complex signals that attract females and prevent hybridization; these tremors are detected via subgenual organs and play a key role in pair formation.^[53] Snakefly (Raphidioptera) adults exhibit similar substrate-borne signals during courtship, though less studied, facilitating mate location in forested habitats.^[54] Most Neuropterida are solitary throughout their life cycle, with larvae typically avoiding conspecifics to minimize cannibalism risks. However, some Neuroptera larvae, particularly in Chrysopidae, show limited communal tendencies, such as gregarious aggregation around prey patches for shared feeding, though without true social structure.^[55] Defensive behaviors are primarily passive or evasive. Snakefly larvae often rely on camouflage, blending with bark or litter through their elongated, campodeiform bodies to avoid detection by predators.^[56] Activity patterns align with predatory and reproductive needs. Adult Neuropterida are predominantly crepuscular or nocturnal, emerging at dusk for flight and foraging to evade diurnal predators and capitalize on reduced competition.^[3] Larval ambush strategies reinforce this, with many species active during low-light periods when prey is more vulnerable.^[57]

Diversity

Extant groups

Neuropterida encompasses over 6,700 extant species worldwide, with the order Neuroptera accounting for more than 85% of this diversity, comprising approximately 6,000 species across 17 families.^[58]^[59] The remaining species are distributed among the orders Megaloptera and Raphidioptera, which together represent a smaller but ecologically significant portion of the clade. Megaloptera includes 425 species in two families, Sialidae (alderflies) and Corydalidae (dobsonflies and fishflies), primarily inhabiting riparian and aquatic environments.^[6] Raphidioptera, known as snakeflies, consists of 253 species in two families, Raphidiidae and Inocelliidae, with a more restricted distribution mainly in temperate regions of the Northern Hemisphere.^[60] Within Neuroptera, family-level diversity varies widely, reflecting adaptations to diverse habitats from forests to arid zones. For instance, the family Hemerobiidae (brown lacewings) includes about 500 species, many of which are generalist predators effective in temperate woodlands and agricultural settings.^[61] Other prominent families include Chrysopidae (green lacewings, over 1,200 species) and Myrmeleontidae (antlions, approximately 2,000 species), which dominate in terms of species richness and are key components of terrestrial ecosystems.^[58] Recent taxonomic efforts have documented numerous new species, particularly from tropical regions like Southeast Asia and South America, enhancing our understanding of neuropteran biodiversity.^[62]^[63] Conservation concerns for Neuropterida are generally low, as most species are widespread and not currently listed as threatened; however, habitat degradation poses risks to localized populations, such as certain dobsonfly species in North America that depend on clean, unpolluted streams.^[64] Lacewings, especially from families like Chrysopidae and Hemerobiidae, play a vital role in biological control within agriculture, where their predatory larvae target pests such as aphids and mealybugs, reducing the need for chemical pesticides in integrated pest management programs.^[65] Ongoing research using molecular barcoding techniques has increased diversity estimates by uncovering cryptic species and undescribed taxa, particularly in tropical forests where sampling remains limited, suggesting the true number of extant Neuropterida species may exceed current figures.^[66]^[67] This approach highlights the potential for further discoveries in biodiverse hotspots like the Amazon and Southeast Asian rainforests.

Fossil record

The fossil record of Neuropterida spans from the Late Permian to the Cenozoic, with the oldest known specimens attributed to the family Permithonidae from deposits in the Tunguska Basin of Siberia, dating to approximately 259–252 million years ago (Ma). These early fossils, characterized by simple wing venation, represent stem-group Neuroptera and mark the initial appearance of the clade near the end of the Permian period. Diversity remained low through the Triassic, with only sporadic records, before expanding significantly during the Mesozoic. Peak diversity occurred in the Jurassic and Cretaceous periods, when Neuropterida exhibited a wide array of forms, including large-bodied taxa and specialized predators, reflecting an adaptive radiation in terrestrial ecosystems.^[68]^[69]^[70] Major fossil deposits have yielded exceptional insights into Neuropterida paleodiversity, particularly amber inclusions from the mid-Cretaceous (Albian–Cenomanian, ~110–94 Ma). Burmese (Myanmar) amber from Kachin State preserves over 100 genera and 135 species across 23 families in all three orders (Neuroptera, Megaloptera, Raphidioptera), representing nearly 15% of all described fossil Neuropterida and highlighting the clade's richness during the height of the dinosaur era. Dominican amber (Eocene, ~20–15 Ma) contains diverse Raphidioptera, including snakefly species in genera like Agulla and Raphidia, often preserved with fine details of body structures and associations with other arthropods. Compression fossils from lacustrine and lagoonal sediments, such as those in the Jurassic Daohugou Beds of China (~165 Ma), provide additional records through wing imprints.^[71]^[72]^[73] Several extinct families underscore the morphological experimentation within Neuropterida during the Mesozoic. The Kalligrammatidae, known from Jurassic deposits in Eurasia (~165–145 Ma), included butterfly-like lacewings with colorful wing patterns and elongated palps, adapting to flower-visiting behaviors in early angiosperm ecosystems. Rafaelidae, an enigmatic group from the Lower Cretaceous Crato Formation of Brazil (~110 Ma), featured highly specialized, lacewing-like forms with unique raptorial forelegs, possibly representing a short-lived lineage bridging Neuroptera and other holometabolans. To date, approximately 930 fossil species of Neuropterida have been described, with the majority from Mesozoic amber and compression sites.^[74]^[75]^[71] Preservation in amber has captured rare behavioral evidence, such as oviposition and hatching in Neuroptera. In Early Cretaceous Lebanese amber (~130 Ma), green lacewing (Chrysopidae) larvae are preserved alongside split eggshells, demonstrating the first fossil record of eclosion mechanics where neonates use egg bursters to emerge. Mid-Cretaceous Burmese amber includes aphidlion-like larvae still attached to chorionic egg cases, indicating predatory specialization from the outset of development. Compression fossils, prevalent in Permian and Jurassic shales, excel at revealing intricate wing venation patterns, which are critical for taxonomic identification and phylogenetic analyses of extinct lineages.^[76]^[77]^[70]

Evolution

Origins and diversification

The origins of Neuropterida trace back to the early holometabolan radiation during the late Carboniferous, approximately 300 million years ago (Ma), when the group emerged as a stem lineage within Holometabola following the divergence from clades like Panorpida and Hymenopterida.^[78] The crown group of Neuropterida is estimated to have originated around 321 Ma in the mid-Carboniferous, with initial diversification linked to the broader adaptive radiation of endopterygote insects amid expanding terrestrial ecosystems.^[8] Stem-group fossils, including extinct families such as Permoberothidae and Permithonidae from the Early Permian, represent early eidoneuropterans and mark the appearance of characteristic net-veined wings, a defining feature that facilitated flight and dispersal in Paleozoic forests. Major diversification events unfolded during the Mesozoic era, particularly from the Triassic to Jurassic, as Neuropterida underwent an explosive radiation that established the modern orders. Order-level splits occurred by the Early Triassic, with Megaloptera diverging around 239 Ma and developing aquatic larval adaptations that allowed exploitation of freshwater habitats, as evidenced by Jurassic fossils of sialids and corydalids.^[8] Neuroptera followed with a crown age of about 281 Ma in the Early Permian, but saw peak family-level diversification in the Jurassic around 197 Ma, coinciding with the Mesozoic "golden age" for the group.^[10] Raphidioptera emerged later, around 132 Ma in the Early Cretaceous. Some lineages experienced a decline toward the end of the Cretaceous, with reduced diversity in certain families following the Cretaceous-Paleogene (K-Pg) extinction event at 66 Ma, though core groups persisted into the Cenozoic. Key evolutionary drivers included co-evolution with emerging angiosperms during the Cretaceous, where the radiation of flowering plants provided pollen as a supplementary food source for adults in families like Chrysopidae, enhancing survival and reproductive strategies amid shifting vegetation. Predation pressures also profoundly shaped larval morphology, favoring diverse predatory forms—such as the active hunters in Neuroptera and the amphibious predators in Megaloptera—that specialized in capturing soft-bodied prey like aphids and small invertebrates, thereby driving niche partitioning and adaptive bursts throughout the Mesozoic.^[10] These factors, combined with habitat expansions, underpinned the group's resilience and order-level distinctions by the Triassic.^[8]

Relationships to other insects

Neuropterida belongs to the superorder Endopterygota, the holometabolous insects characterized by complete metamorphosis, where larvae and adults occupy distinct ecological niches. Within this diverse group, which encompasses over 80% of all insect species, Neuropterida is positioned as the sister clade to Coleopterida—a lineage comprising the orders Coleoptera (beetles) and Strepsiptera (twisted-wing parasites)—together forming the higher taxon Neuropteroidea.^[8] This relationship has been consistently supported by recent phylogenomic analyses, contrasting with earlier morphological hypotheses that placed Neuropterida as sister to the bulk of Endopterygota excluding Hymenoptera (sawflies, bees, wasps, and ants).^[8] In some phylogenies, this Neuropteroidea clade appears basal to a larger assemblage including Hymenoptera and the mecopteroid lineages (Mecoptera, Siphonaptera, Diptera, Trichoptera, and Lepidoptera), highlighting Neuropterida's role in early holometabolan diversification.^[79] Neuropterida shares certain ancestral traits with the mecopteroid superorder Mecopteroidea, particularly in the morphology of their pupal stages; both groups exhibit relatively active, exarate pupae capable of limited locomotion and defensive behaviors, unlike the more immobile pupae in advanced holometabolans such as Diptera.^[80] Fossil evidence further underscores these connections, with Permian and Mesozoic specimens showing scorpionfly-like (Mecoptera) venation patterns in early neuropteridan wings, suggesting a shared evolutionary history in the development of flexible, predatory adult forms.^[81] However, Neuropterida diverges markedly in other features; its holometaboly is considered primitive, retaining a more direct larval-pupal transition with free appendages and active pupae, in contrast to the highly derived, adecticous pupae of Diptera where appendages are glued to the body and emergence relies on ecdysis without movement.^[80] Wing venation in Neuropterida also echoes that of the palaeopterous order Odonata (dragonflies and damselflies), featuring a dense, net-like pattern of crossveins that supports broad, gliding flight, though neuropteridan wings possess reduced indirect flight musculature compared to the powerful, direct musculature of odonates.^[82] Molecular evidence from transcriptome-based phylogenomics has firmly established Neuropterida as monophyletic, resolving internal relationships such as Raphidioptera as sister to Megaloptera + Neuroptera, while excluding extinct taxa like Miomoptera, which morphological reanalyses now classify as stem-group cercarians outside the clade.^[8] Key studies from 2020 to 2023, utilizing datasets of thousands of orthologous genes (e.g., 3,983 clusters of orthologous groups across 1.5 million amino-acid sites), have corroborated this monophyly with high bootstrap support, mitigating long-branch attraction artifacts common in earlier mitochondrial genome analyses.^[8]^[83] These findings exclude Miomoptera based on the absence of defining synapomorphies like raptorial forelegs and complete metamorphosis in molecularly informed trees, emphasizing Neuropterida's integrity as a crown holometabolan group.^[84] The phylogenetic position of Neuropterida offers critical insights into the evolution of insect flight, as its retained primitive wing structures—such as extensive venation and tracheation patterns—provide a morphological bridge between early pterygote innovations and the specialized elytra of Coleoptera or halteres of Diptera.^[82] This placement suggests that neuropteridan wings, while effective for sustained gliding in predatory contexts, represent a conservative design that may inform the origins of powered flight in Endopterygota, predating more derived modifications.^[85] Nonetheless, ongoing re-evaluation of extinct taxa, such as potential miomopteran relatives, raises the possibility of paraphyly if additional fossil synapomorphies link them to basal neuropteridans, potentially reshaping understandings of early holometabolan radiation.^[84]

References

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[12]
[PDF] The first holistic SEM study of Coniopterygidae (Neuroptera)
The sensilla- like structures situated on the plicatures are not consid- ered to be sensory organs as their openings are not large enough to serve as terminal ...
[13]
Phylogeny and Evolution of Neuropterida: Where Have Wings of ...
Aside from molecular studies, the presence of plesiomorphic characters in female genitalic morphology and the number of Malpighian tubules place them outside of ...
[14]
Unraveling the evolutionary history of the snakefly family Inocelliidae ...
Mar 29, 2022 · Currently, the order is placed as the sister group of Megaloptera + Neuroptera in a number of recent phylogenetic studies based on different ...
[15]
[PDF] Grub-like larvae of Neuroptera (Insecta)
Potential morphological synapomorphies of mature larvae of Ithonidae and Polystoechotidae are the mandibles with exceptionally broad base and markedly thickened ...
[16]
[PDF] The Biology of Raphidioptera: A Review of Present Knowledge
Jul 30, 2002 · and lasts a few days up to about three weeks (life cycle type I). In some (or all) species of a few genera of Raphidiidae (life cycle type II:.
[17]
Urban Entomofauna | Pinning Block - U.OSU
Jun 20, 2023 · Upon arrival they will spend from one to five years in this aquatic larval stage, depending on water temperature, latitude, and the time of ...
[18]
dobsonflies & fishflies - University of Kentucky
Dec 19, 2006 · Dobsonflies and fishflies live most of their lives in the larval stage, often 2 or 3 years. Larval dobsonflies and fishflies, often called ...
[19]
Diapause in the neuropteran Chrysopa oculata - ScienceDirect.com
The results suggest that under natural conditions diapause is primarily induced by the short-days of late summer, that it is maintained by the short-days of ...Missing: Neuropterida | Show results with:Neuropterida
[20]
Hypermetamorphosis - ScienceDirect.com
Only the Mantispidae have true hypermetamorphosis. Most of the species feed on spider eggs associated with a single egg sac. The first instar either enters a ...
[21]
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 ...
[22]
Mating Behavior of Three Species of Coniopterygidae (Neuroptera)
Aug 10, 2025 · ... ... Mating is preceded by precopulatory behaviour of both sexes by rotating or elevating their abdomina, fluttering their wings and walking ...
[23]
(PDF) Chemical Ecology of Neuroptera - ResearchGate
Mar 17, 2016 · mating system involving male-produced pheromones may be characteristic of many neuropterans. In Neuroptera, which is an ancestral lineage in ...
[24]
Megaloptera - Lacewings and Allies Recording Scheme
Males use vibrations or scent glands to attract a mate. The females lay their eggs on rocks or leaves near water. They have predaceous larvae that capture other ...
[25]
Bundled sperm and post-copulatory mate guarding by male fishflies ...
Insemination through an externally attached spermatophore: Bundled sperm and post-copulatory mate guarding by male fishflies (Megaloptera: Corydalidae).
[26]
[PDF] Protective role of the egg stalk in Chrysopidae (Neuroptera)
Some species lay eggs in clusters without stalks, others in clusters in which the stalks are glued together and others singly on stalks either in loose groups ...
[27]
Oviposition of the Dobsonfly (Corydalus cornutus, Megaloptera) on a ...
This coating appeared to protect the eggs from excessive heating and predation. Some 4428 egg masses were observed at 260 sites in the two annual surveys. Trees ...
[28]
Functions of Egg-Coating Substances Secreted by Female ...
Aug 25, 2022 · The accessory gland substances of most species of Megaloptera also function to protect eggs from attack by oophagous predators, such as ladybird ...Missing: guarding | Show results with:guarding
[29]
Order Raphidioptera - UBC Zoology
Females use the ovipositor to deposit eggs singly or in small batches in bark, crevices, in debris or under the body of mature scale insects. The elongate ...Missing: oviposition | Show results with:oviposition
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Green lacewings (Neuroptera: Chrysopidae) are commonly ...
Aug 14, 2017 · In arthropods, Rickettsia may cause sex-ratio distorting phenotypes, i.e., male-killing [5, 6] or parthenogenesis [7, 8], but has also been ...
[31]
Parthenogenesis in Hexapoda: holometabolous insects
Sep 14, 2017 · Parthenogenesis is recorded within six orders of holometabolous insects, namely Hymenoptera, Strepsiptera, Coleoptera, Trichoptera, ...
[32]
Neuropterida from South America: large diversity, largely unknown
Dec 15, 2019 · Neuropterida comprises about 6 430 extant and 930 fossil species distributed all over the world, except in Antarctica. In South America ...<|control11|><|separator|>
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Biodiversity of the Neuropterida (Insecta: Neuroptera, Megaloptera ...
Neuroptera is a group of mainly terrestrial insects with approximately 6000 species, 603 genera, and 15 families [18] , with a worldwide distribution.
[34]
Neuroptera - an overview | ScienceDirect Topics
Neuropterida comprises three orders with holometabolous development, with approximately 6500 species of Neuroptera in almost 20 families, 300 of Megaloptera ...Missing: taxonomy | Show results with:taxonomy
[35]
Fauna Europaea: Neuropterida (Raphidioptera, Megaloptera ...
Neuropterida is a fairly small group of Insecta, with about 6,500 described species, most of which live in arboreal habitats, but many species are eremial and ...
[36]
[PDF] the aquatic neuropterida of iowa
Jul 29, 2020 · Larvae of this species occur in lentic habitats, including ponds, marshes, and swamps (Cuyler 1958,. Tarter et al. 1976, Bowles and Sites. 2015) ...
[37]
World distribution of the order Raphidioptera. - ResearchGate
Recent snakeflies comprise ~240 species in only two families, i.e. Raphidiidae and Inocelliidae, and they mainly occur in the Holarctic region with a few ...
[38]
Megaloptera Biogeographical History: Mitochondrial Phylogenomics
Nov 24, 2021 · Here we present a comprehensive phylogeny of Megaloptera based on a large-scale mitochondrial genome sequencing of 99 species representing >90% of the world ...
[39]
Neuroptera (Net-winged Insects)
Neuropterans can be found across most habitats in Australia and 90% of Australian species are endemic (New, 1992). Most of them are terrestrial.
[40]
GIS-based modelling reveals the fate of antlion habitats in ... - Nature
Mar 24, 2020 · While significant erosion control is a feasible positive effect, habitat loss is likely to be a negative consequence of afforestation in ...
[41]
[PDF] Aquatic-insects, their biodiversity, importance and conservation
Feb 18, 2025 · However, anthropogenic factors such as industrial pollution, climate change, pesticide runoff, and habitat destruction threaten their ...
[42]
Divergent venom effectors correlate with ecological niche in ...
Aug 13, 2024 · Neuropteran larvae are fierce predators that use venom to attack and feed on arthropod prey. Neuropterans have adapted to diverse and ...
[43]
A first higher-level time-calibrated phylogeny of antlions (Neuroptera
They also possess potent digestive enzymes, which are used to liquefy the internal components of their prey (Griffiths, 1980, Griffiths, 1982, Van Zyl et al., ...
[44]
Common Green Lacewing Biocontrol Agent Factsheet - Cornell CALS
Adult lacewings need nectar, honeydew, or pollen as food before egg laying. ... Lacewing larva feeding on an aphid. Green leaf with black spots and holes ...
[45]
Predatory Behavior of a Pit-Making Antlion, Myrmeleon ... - jstor
The larvae of antlions (Neuroptera: Myrmeleontidae) are renowned for their pred- atory tactic: the construction of funnel-shaped pitfall traps in sandy ...
[46]
The Morphological Diversity of Antlion Larvae and Their Closest ...
The larvae of owlflies and antlions (here shortly embraced by the term “owllions”) are ambush predators. Their mouthparts are transformed into teeth-bearing ...Missing: communal sociality
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Courtship Songs of Green Lacewings Filmed in Slow Motion: How a ...
Feb 5, 2015 · Courtship Songs of Green Lacewings Filmed in Slow Motion: How a Simple Vibrating Structure can Generate Complex Signals (Neuroptera: Chrysopidae ...Missing: mating | Show results with:mating
[48]
(PDF) Acoustic Communication in Neuropterid - ResearchGate
Nov 17, 2016 · ... The specimen emits the vibration that travel through the substrate and is received by the other nearby individuals through the subgenual ...
[49]
[PDF] Not quite social – possible cases of gregarious behaviour of ...
Jan 23, 2022 · Character combinations that were found in neuropteran larvae, also particularly in myrmeleontiformian larvae, cannot be found in any extant form ...
[50]
Declining morphological diversity in snakefly larvae during last 100 ...
Apr 18, 2022 · Snakeflies retain many plesiomorphies as adults, such as a prominent ovipositor in females. In addition, the larvae appear to have a rather ...
[51]
reconstructing the ancestral state of autotomy and regeneration - PMC
Using an ancestral trait reconstruction, we find that autotomy and regeneration were present at the base of the arthropod and reptile trees.Missing: Neuropterida | Show results with:Neuropterida
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Further Studies on the Daily Activity Pattern of Neuroptera with some ...
Aug 7, 2025 · The results of the experiments were described by activity diagram and were categorized into Duelli-type flight activity pattern. During the ...
[53]
Neuroptera - Royal Entomological Society
Worldwide there are around 6,000 known species in 17 families; in Britain there are 70 species in 6 families. Identification help. See also. Insect Week ...
[54]
Bibliometric Analyses of Web of Science Illuminate Research ...
Neuropterida is one of the earliest ancient groups of Holometabola that consists of Neuroptera, Raphidioptera and Megaloptera. There are more than 6500 extant ...<|control11|><|separator|>
[55]
A world checklist of extant and extinct species of Megaloptera (Insecta
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[56]
Chromosome-level genome assembly of the snakefly ... - Nature
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[57]
Family Hemerobiidae - Brown Lacewings - BugGuide.Net
Classification. Kingdom Animalia (Animals). Phylum Arthropoda (Arthropods) · Explanation of Names. Hemerobiidae Latreille, 1802 · Numbers. >60 spp. · Size. 6-15 mm ...Missing: count | Show results with:count
[58]
New dustywings (Neuroptera, Coniopterygidae) from mid ... - ZooKeys
Mar 5, 2019 · Two new genera and species of Coniopterygidae (Neuroptera) are described and illustrated from mid Cretaceous (Cenomanian) amber of Myanmar.
[59]
New Cretaceous antlion-like lacewings promote a phylogenetic ...
Aug 12, 2021 · Here we report on three new genera and species of Babinskaiidae from the mid-Cretaceous Kachin amber of Myanmar, namely Calobabinskaia xiai gen.
[60]
Dobsonflies, Fishflies, and Alderflies: Megaloptera - Encyclopedia.com
CONSERVATION STATUS. No megalopterans are listed as endangered or threatened, but like all species that live only in small geographic areas, their populations ...
[61]
Green lacewings - UC IPM
Adults are soft-bodied insects with golden eyes and 4 membranous wings held rooflike over the body at rest. The wings and body are commonly green. Adults (head ...
[62]
Application of DNA barcodes in wildlife conservation in Tropical East ...
Jun 24, 2016 · DNA barcodes are short standardized DNA sequences used for taxonomic purposes and have the potential to lessen the challenges of biodiversity inventory and ...
[63]
Elevational Diversity Patterns of Green Lacewings (Neuroptera
Nov 14, 2021 · We investigate the species diversity patterns of green lacewings (an important predatory group of insects) along the gradient of elevation from the Shaluli ...
[64]
Permithonidae) from the Guadalupian of the Lodève Basin ...
Neuropterida together with sister-group Coleopterida represent a holometabolan insect clade so called Neuropteroida, well supported by phylogenetic analyses ...
[65]
New lacewings (Neuroptera) from the terminal Permian and basal ...
Aug 5, 2025 · The only other neuropteran family recorded from the Permian is Archeosmylidae (one species in the terminal Permian of Australia, other species ...
[66]
A Remarkable New Family of Jurassic Insects (Neuroptera) with ...
The family comprises large neuropterans with most primitive wing venation in the order indicated by the presence of ScA and AA1+2, and the dichotomous branching ...
[67]
The Neuropterida from the mid-Cretaceous of Myanmar
Neuropterida are a holometabolous superorder, represented by Megaloptera, Raphidioptera and Neuroptera. The adults of Neuropterida, known as green lacewings, ...
[68]
Diverse Neuropterida in Cretaceous amber, with particular reference ...
They are relatively abundant in the fossil record of various Cretaceous ambers across the Northern Hemisphere, but all the Cretaceous rhachiberothids belong to ...
[69]
The Neuropterid Fauna of Dominican and Mexican Amber ...
Sep 6, 2007 · Although the fossil lacks one of the four synapomorphies of the genus (i.e., apomorphic loss of forewing 4m-cu), we have placed the species here ...
[70]
The evolutionary convergence of mid-Mesozoic lacewings and ...
Mid-Mesozoic kalligrammatid lacewings (Neuroptera) entered the fossil record 165 million years ago (Ma) and disappeared 45 Ma later.Missing: Rafaelidae | Show results with:Rafaelidae
[71]
A new extraordinary neuropterid family from the Lower Cretaceous ...
Aug 10, 2025 · A new, highly specialized family, Rafaelidae, is proposed for two new lacewing-like species from the Lower Cretaceous Crato Formation in the ...
[72]
Newborn insects trapped in amber show first fossil evidence of how ...
Dec 20, 2018 · Trapped together inside 130 million-year-old Lebanese amber, or fossilised resin, researchers found several green lacewing newborn larvae, the ...
[73]
A hatching aphidlion‐like lacewing larva in 100 million years old ...
Dec 17, 2022 · We report a fossil aphidlion-like larva preserved with its egg case in 100 million year old Kachin amber, Myanmar.
[74]
Single-copy nuclear genes resolve the phylogeny of ... - BMC Biology
Jun 24, 2009 · Palaeontological and phylogenetic evidence suggest an origin for Holometabola in the late Carboniferous (318-300 Ma) [1, 32, 33], but ...
[75]
9-Genes Reinforce the Phylogeny of Holometabola and Yield ...
Figure 3. ML phylogenetic tree showing the placement of Strepsiptera sister to Neuropterida and interrelationships of other holometabolous insects based on the ...
[76]
Holometabola - an overview | ScienceDirect Topics
The pupae of many primitive holometabolans (Neuropterida, Trichoptera, known Mecoptera) are relatively active (and, in particular, allow various activities of ...
[77]
[PDF] The First Mecopteroids (Insecta: Papilionidea = Mecopteroidea) and ...
The first mecopteroids are close to neuropteroids. (Myrmeleontidea = Neuropteroidea) and originate from primitive members of the latter superorder (Mar- tynova, ...Missing: Neuropterida | Show results with:Neuropterida
[78]
Wing Tracheation in Chrysopidae and Other Neuropterida (Insecta)
Dec 18, 2017 · In the present study, we review the homology statements of wing venation among Neuropterida, with an emphasis on Chrysopidae (green lacewings).
[79]
[PDF] Improved modelling of compositional heterogeneity ... - Biotaxa
Feb 28, 2023 · Recent phylogenomic studies of Neuropterida based on mitogenomes, anchored hybrid enrichment (AHE) data, and transcriptomes have yielded a well- ...<|control11|><|separator|>
[80]
Redefining the extinct orders Miomoptera and Hypoperlida as stem ...
Aug 25, 2017 · Epimastax Martynov, 1928 [1] (type genus of the Epimastacidae Martynov ... Neuropterida and Panorpida); no distinct angle of radius at base ...
[81]
Theoretical morphospace analysis of neuropteran wings reveals ...
The harmonics and functional optimality data were split into five time bins: Permian and Triassic, Jurassic, Cretaceous, Paleogene and Neogene, and Present.