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Neoptera

Neoptera is an infraclass within the subclass Pterygota of the class Insecta, encompassing the vast majority of winged insect orders and representing over 95% of all described insect species.^[1]^[2] This clade is defined by a key morphological innovation: the presence of a wing-folding mechanism, or flexon, that allows the forewings to fold back over the abdomen when at rest, distinguishing Neoptera from the Paleoptera, whose wings remain extended.^[3]^[4] Originating in the early Carboniferous period, Neoptera evolved from ancient pterygote ancestors and diversified rapidly, with phylogenetic analyses supporting its monophyly based on shared traits like oblique wing articulation and indirect flight musculature.^[5] The group is subdivided into two major superorders: Exopterygota (hemimetabolous insects with gradual metamorphosis, including orders such as Orthoptera and Hemiptera) and Endopterygota (holometabolous insects with complete metamorphosis, encompassing dominant orders like Coleoptera, Lepidoptera, Diptera, and Hymenoptera).^[3] Neopterans play critical ecological roles worldwide, serving as pollinators, predators, decomposers, and vectors for diseases, while also impacting agriculture through both beneficial services and pest activities.^[3]

Introduction

Definition and Scope

Neoptera is a major clade within the subclass Pterygota of the class Insecta, comprising winged insects characterized by their ability to flex their wings back over the abdomen when at rest.^[6] The name derives from the Greek words neos (new) and pteron (wing), literally meaning "new wings," reflecting the evolutionary innovation of wing folding relative to more basal winged insects. This clade encompasses the vast majority of extant winged insect orders, excluding the basal Palaeoptera such as Odonata (dragonflies and damselflies) and Ephemeroptera (mayflies), and spans a temporal range from the Late Carboniferous to the present day.^[6]^[4] The defining morphological feature of Neoptera is the presence of an oblique superior pleural suture in the thoracic pleuron, which separates the episternum and epimeron and facilitates wing flexion through articulation with indirect flight muscles.^[7] In contrast, Palaeoptera lack this oblique orientation, with their pleural suture running more vertically, preventing the wings from folding over the abdomen and instead holding them outstretched.^[8] This wing-flexing mechanism, involving specialized axillary sclerites at the wing base and associated muscles, enables greater maneuverability and protection of the wings during rest.^[6] Within the broader phylogeny of Insecta, Neoptera represents the derived sister group to Palaeoptera under the traditional division of Pterygota, though molecular studies continue to refine these relationships.^[4]

Historical Recognition

The concept of Neoptera was formally proposed by the Russian entomologist and paleontologist Andrei Vasilievich Martynov in his 1923 publication, where he distinguished two primary types of insect wings based on their articulation and folding mechanisms, separating Neoptera—characterized by the ability to flex wings over the abdomen—from the Palaeoptera.^[9] Martynov expanded on this in subsequent works, including 1924 and 1925 papers, emphasizing the evolutionary significance of the oblique wing articulation in Neoptera as a key synapomorphy for classifying the majority of pterygote insects.^[10] This proposal marked a foundational shift in insect systematics by prioritizing functional wing base structure over earlier criteria.^[9] Early adoption of the Neoptera framework encountered debates regarding the placement of certain orders with ambiguous wing traits, such as Thysanoptera (thrips), whose reduced and fringed wings initially raised uncertainty about their alignment with neopterous flexion or palaeopterous rigidity.^[9] These uncertainties stemmed from pre-Martynov classifications that often positioned Thysanoptera as transitional or osculant between groups like Orthoptera and Neuroptera, complicating their integration into the new division. Over time, morphological analyses confirmed Thysanoptera's neopterous affinity, resolving the debate by incorporating them into the Paraneoptera subgroup alongside Hemiptera and Psocodea, based on shared features like mandibular stylets and wing base modifications.^[10] Martynov's emphasis on wing articulation profoundly influenced entomological systematics, redirecting attention from detailed venation patterns—previously dominant in 19th- and early 20th-century classifications—to the biomechanical properties of wing attachment and flexion.^[9] This paradigm shift facilitated more robust phylogenetic hypotheses for Pterygota, underpinning later subdivisions like Polyneoptera and Oligoneoptera, and remains a cornerstone of insect evolutionary studies despite subsequent molecular refinements.^[10]

Taxonomy

Higher Classification

Neoptera is classified as an infraclass within the subclass Pterygota, which comprises all winged insects, under the class Insecta in the subphylum Hexapoda and phylum Arthropoda.^[11]^[12] This placement reflects the Linnaean hierarchy, where Neoptera represents a major division of pterygote insects distinguished by advanced wing mechanics from more basal winged groups.^[13] The primary synapomorphy defining Neoptera is the specialized articulation of the wing base, including axillary sclerites and flexion lines, that allows the fore- and hindwings to fold flat against the abdomen at rest.^[4] This neopterous folding mechanism, first described by Martynov in 1924, enables greater maneuverability and protection compared to the rigid-winged Paleoptera, and it is supported by associated musculature such as the first tergosternal muscle.^[14] Additional shared traits include modifications in the thoracic pleural regions that facilitate this wing flexion.^[15] In contemporary taxonomy, Neoptera is accepted as a monophyletic clade encompassing the principal subgroups Polyneoptera, Paraneoptera, and Holometabola, based on morphological and molecular evidence confirming their common ancestry.^[16] This classification underscores Neoptera's dominance among winged insects, accounting for the vast majority of extant pterygote diversity.^[4]

Major Subgroups

Neoptera is primarily divided into three major subgroups: Polyneoptera, Paraneoptera, and Holometabola, each distinguished by unique morphological and developmental traits that reflect their evolutionary divergence within the clade. These correspond to the traditional superorders, with Polyneoptera and Paraneoptera comprising Exopterygota (hemimetabolous neopterans) and Holometabola equivalent to Endopterygota (holometabolous neopterans).^[17] These divisions encompass the vast majority of neopteran diversity, with Holometabola representing the most species-rich branch.^[18] Polyneoptera comprises a diverse assemblage of orders characterized by incomplete metamorphosis and wing structures featuring symmetrical fore- and hindwings, often with the hindwings expanded into a broad, fan-like anal region for enhanced flight capabilities.^[19] Key orders include Orthoptera (grasshoppers and crickets), Dermaptera (earwigs), and Plecoptera (stoneflies), which exhibit chewing mouthparts and cerci on the abdominal tip as common features.^[20] This subgroup accounts for a moderate portion of neopteran species, emphasizing terrestrial and riparian habitats.^[21] Paraneoptera includes orders with predominantly hemimetabolous development and specialized piercing or sucking mouthparts adapted for fluid feeding, alongside reduced wing venation and a tendency toward an elongate head due to postclypeal enlargement.^[22] Representative orders are Hemiptera (true bugs), Thysanoptera (thrips), and Phthiraptera (lice), with Psocodea (booklice) often grouped closely; these insects are notable for their ecological roles in herbivory, parasitism, and sap-sucking.^[20] The subgroup's monophyly is supported by molecular and morphological synapomorphies, such as specific antennal and maxillary structures.^[23] Holometabola forms the largest subgroup, defined by complete metamorphosis involving distinct larval, pupal, and adult stages, where adult appendages develop internally via imaginal discs.^[18] Major orders include Coleoptera (beetles), Lepidoptera (butterflies and moths), and Hymenoptera (bees, ants, and wasps), which dominate global insect diversity with over 80% of described species.^[18] This group's success is linked to specialized larval feeding strategies and pupal remodeling.^[20] These subgroups constitute the core branches of Neoptera, with Polyneoptera branching basally, followed by the sister clades Paraneoptera and Holometabola, as inferred from ribosomal RNA and protein-coding gene analyses.^[20]

Phylogeny and Evolution

Phylogenetic Position

Neoptera constitutes the sister group to Palaeoptera within the winged insects (Pterygota), a relationship established through cladistic analyses based on shared derived characters (synapomorphies). Key morphological synapomorphies supporting this positioning include the development of a pterothoracic furca, which anchors wing-folding muscles, enabling the characteristic flexion of wings over the abdomen—a feature absent in palaeopterans like dragonflies and mayflies.^[24] This cladistic framework underscores Neoptera's monophyly as a derived lineage adapted for wing articulation, distinguishing it from the more rigid-winged basal pterygotes.^[25] Molecular phylogenetics has robustly confirmed Neoptera's monophyly and its position as sister to Palaeoptera, leveraging large-scale genomic and transcriptomic datasets to resolve deep insect relationships. Seminal studies, such as Kjer et al. (2016), utilized transcriptome sequences from diverse insect orders to reconstruct hexapod phylogeny, yielding strong support for Neoptera as a monophyletic clade encompassing over 99% of pterygote diversity, with bootstrap values exceeding 95% for key nodes.^[26] Similarly, Wipfler et al. (2019) integrated phylogenomic data from 200+ species, including polyneopteran transcriptomes and genomes, to affirm Neoptera's unity, highlighting its divergence from Palaeoptera around 350 million years ago based on calibrated trees.^[25] These analyses employed maximum likelihood and Bayesian methods on thousands of orthologous genes, mitigating long-branch attraction artifacts that plagued earlier molecular efforts.^[25] Within Neoptera, phylogenomic approaches have resolved longstanding debates on subgroup placements, such as the position of Thysanoptera (thrips). Johnson et al. (2018) analyzed 2,395 single-copy nuclear genes from hemipteroid insects, placing Thysanoptera as the sister group to Hemiptera within the Paraneoptera clade, with posterior probabilities near 1.0, thus confirming its integration into Neoptera and refuting prior hypotheses of basal polyneopteran affinity.^[27] This resolution exemplifies how increased taxon sampling and gene coverage in modern phylogenomics have stabilized Neoptera's internal structure, aligning molecular trees with morphological expectations.^[27]

Evolutionary History and Fossil Record

The origins of Neoptera are rooted in the Late Carboniferous period, with the earliest definitive fossils appearing around 315 million years ago during the Namurian B stage. A key example is Baryshnyala occulta, the type species of the family Baryshnyalidae, discovered in the Hagen-Vorhalle locality of Germany; this tiny insect, measuring just 5.5 mm in wingspan, highlights the early morphological diversity and miniaturization within Neoptera, contrasting with the larger-bodied Palaeoptera of the same era.^[28] Further evidence comes from trace fossils in the Wamsutta Formation of Massachusetts, dated to 308–314 million years ago (Westphalian B–C), interpreted as impressions left by a surface-skimming neopteran, likely a stem-group plecopteran (stonefly). This imprint preserves details of wing folding and body posture, indicating that Neoptera had already developed advanced aerial locomotion behaviors, such as low-altitude skimming, potentially as a precursor to more efficient flight.^[29] The emergence of Neoptera is closely tied to the evolution of the wing flexion mechanism, which allowed wings to fold backward over the abdomen for protection during rest and enhanced aerodynamic efficiency in flight. Fossilized nymphs from the Paleozoic era exhibit articular wing bases with V-shaped pteralia (sclerites), a synapomorphy distinguishing Neoptera from contemporaneous Palaeoptera like the griffenflies (family Meganeuridae), giant predatory insects with fixed, non-folding wings and spans up to 70 cm that dominated Carboniferous skies but lacked the adaptability of neopteran flight.^[30]^[31] Diversification intensified in the Permian period (299–252 million years ago), as evidenced by numerous small-bodied neopterans, including early orthopterans such as the Protomeropidae from Early Permian deposits in the USA and Australia. These fossils document a proliferation of exopterygote lineages, with wing venation patterns showing progressive refinement of the flexion apparatus amid shifting terrestrial environments.^[28] A pivotal evolutionary milestone was the major radiation of Neoptera during the Mesozoic era, initiating in the Middle Triassic around 245 million years ago and continuing through the Jurassic and Cretaceous. This expansion, which established much of the modern family-level diversity, temporally coincided with the rise of angiosperms in the Early Cretaceous (~140 million years ago), enabling Neoptera—particularly holometabolous subgroups—to assume dominant ecological roles as herbivores consuming foliage and pollinators facilitating plant reproduction via specialized behaviors like nectar feeding.^[32] The fossil record of early holometabolans within Neoptera reveals significant gaps, particularly between the Late Carboniferous and Triassic; while stem-group forms appear in the Gzhelian stage (~299 million years ago), such as the skleropteran (Stephanastus polinae, order Skleroptera) and stem hymenopterid (Avioxyela gallica) from French deposits, unambiguous records of crown-group orders like Coleoptera appear in the Permian, while those for Hymenoptera remain sparse until the Late Triassic. These early holometabolous fossils suggest an initial diversification driven by environmental stressors like Pennsylvanian glaciations, but the post-Permian Triassic recovery following mass extinctions marked a surge in adaptive radiations.^[33]

Morphology and Physiology

Wing Flexion Mechanism

The wing flexion mechanism in Neoptera is a key anatomical innovation that distinguishes this clade from Palaeoptera, enabling the wings to fold backward along the body at rest. This capability arises from modifications in the pterothoracic structure, particularly the pleuron, which is divided by an oblique pleural suture into an anterior episternum and a posterior epimeron. The suture's oblique orientation creates a flexible hinge line that positions the pleural wing process on the epimeron, allowing the wing base to articulate in a manner that permits downward and backward rotation. Central to this mechanism are the axillary sclerites at the wing base, arranged in a V-shaped configuration unique to Neoptera. The third axillary sclerite (3Ax) serves as the primary flexor element, pivoting around the pleural wing process when pulled by the axillary-pleural muscle (a direct flight muscle originating from the epimeron). This action folds the wing along a basal flexion line, tucking it parallel to the abdomen. In contrast, Palaeoptera lack this pivoting 3Ax and instead have a fixed, straight hinge with fused basivenalia, resulting in wings that remain extended or curved passively backward without active folding.^[34] While the oblique pleural suture and axillary system provide the structural basis for folding, wing movement during flight and rest is powered by a combination of direct and indirect muscles. The dorsal longitudinal muscles contract to elevate the notum (dorsal thorax), and dorso-ventral muscles depress it, deforming the thorax to drive wing beats indirectly—a system preadapted from ancestral leg-rowing motions. The specific folding, however, relies on the targeted contraction of the axillary-pleural flexor to the 3Ax, often supplemented by basalar and subalar muscles for fine control.^[34] Variations in this mechanism occur across Neopteran subgroups, reflecting adaptations to diverse lifestyles. In Polyneoptera (e.g., Orthoptera like crickets), folding is typically complete, with both wing pairs aligning flat against the abdomen for streamlined resting postures. In contrast, some Holometabola exhibit partial or modified folding; for instance, in Coleoptera (beetles), hindwings fold compactly under hardened forewings (elytra), while in Lepidoptera (butterflies), wings often assume a partial roof-like position rather than full flat alignment. These differences stem from elaborations in axillary sclerite morphology and additional folding lines, but all retain the core Neopteran pivot at the 3Ax.^[35] Functionally, the wing flexion mechanism offers significant advantages for survival and adaptation, particularly in terrestrial environments. Folded wings protect delicate membranes from physical damage, abrasion, and predation by allowing insects to navigate narrow crevices and shelters. This configuration also reduces aerodynamic drag and conserves energy during non-flight periods by minimizing body profile against wind and rain. As an evolutionary innovation, it facilitated Neoptera's radiation into diverse habitats, enabling escape behaviors and resource exploitation unavailable to Palaeoptera with their rigid wings.^[36]

Other Key Anatomical Features

Neoptera display two primary modes of postembryonic development: hemimetaboly, which predominates in the Paraneoptera and Polyneoptera subgroups, and holometaboly, characteristic of the Holometabola. In hemimetaboly, nymphs undergo gradual metamorphosis through a series of molts, progressively developing external wing pads and resembling scaled-down adults without an intervening pupal stage, allowing direct transition to reproductive maturity.^[37] Holometaboly, conversely, involves distinct larval feeding stages, a non-feeding pupal phase for histolysis and histogenesis, and emergence of the adult, with the pupal stage serving as a pivotal evolutionary innovation that decouples larval growth from adult form and function, enhancing ecological partitioning.^[37]^[38] Sensory systems in Neoptera feature advanced compound eyes, multifaceted structures composed of ommatidia that provide panoramic vision and acute motion detection, varying in facet number and acuity across orders to suit ecological demands.^[37] Antennae exhibit considerable morphological variation, typically multisegmented appendages equipped with sensilla for olfaction, gustation, and mechanoreception, ranging from filiform in orthopterans to clavate in coleopterans, thereby enabling diverse sensory inputs critical for foraging and mate location.^[37] Reproductive anatomy includes specialized structures such as the ovipositor in Hymenoptera, an appendage derived from gonopods of abdominal segments 8 and 9 that facilitates precise egg insertion into substrates and, in aculeates, modification into a venom-injecting sting for defense and prey subdual.^[37] Males typically bear an aedeagus, a sclerotized intromittent organ that ensures internal fertilization by transferring spermatophores or sperm directly to the female spermatheca, promoting genetic diversity in neopteran lineages.^[37] Key physiological traits support Neoptera's dominance in active lifestyles, particularly through tracheal system enhancements featuring extensive branching tracheoles and dilatable air sacs that optimize oxygen delivery to metabolically demanding flight muscles, facilitating prolonged aerial activity in pterygote forms.^[37] The Malpighian tubules, blind-ended extensions of the hindgut, enable efficient excretion by actively transporting ions and water from hemolymph to form uric acid-rich urine, minimizing water loss and allowing osmoregulation across xeric to humid habitats.^[37]

Diversity and Ecology

Major Orders and Species Richness

Neoptera encompasses the vast majority of insect diversity, accounting for over 99% of the approximately 1.05 million described insect species worldwide as of 2025. The infraclass is primarily composed of the superorder Holometabola, which undergoes complete metamorphosis and represents about 80% of all known insects. This subgroup includes several hyperdiverse orders that drive much of the overall richness of Neoptera. Among these, Coleoptera (beetles) stands as the most species-rich order, with nearly 400,000 described species as of 2025, showcasing immense morphological and ecological variation from soil-dwelling scarabs to predatory ground beetles. Lepidoptera (butterflies and moths) follows with around 180,000 species, noted for their scaled wings and roles in pollination and as indicators of environmental health. Hymenoptera (bees, wasps, and ants) comprises over 154,000 species, including eusocial forms like honeybees and army ants that exhibit complex behaviors such as division of labor and nest construction. Diptera (flies) also totals approximately 160,000 species, encompassing medically significant groups like mosquitoes alongside decomposers and pollinators such as hoverflies. Additionally, within Exopterygota, Hemiptera (true bugs) includes about 107,000 species, featuring piercing-sucking mouthparts adapted for plant sap-feeding in aphids or predatory habits in assassin bugs.^[39] This extraordinary species richness is concentrated in tropical regions, where environmental stability fosters high endemism and speciation rates. However, pollinator taxa within Hymenoptera, particularly bees, confront pressing conservation challenges from habitat fragmentation, pesticide exposure, and climate change, underscoring the need for targeted protection to sustain ecosystem services like pollination.

Habitats and Global Distribution

Neoptera, encompassing the vast majority of insect species, predominantly occupy terrestrial habitats worldwide, including forests, grasslands, and deserts, where adults and many immature stages thrive in diverse microenvironments such as soil, foliage, and bark. Although over 94% of insect species are primarily terrestrial, approximately 5% undergo aquatic larval stages in freshwater systems, notably in orders like Plecoptera (stoneflies), whose nymphs inhabit cool, oxygen-rich streams and rivers, contributing to nutrient cycling and serving as indicators of water quality.^[40]^[41] Within these habitats, neopterans fulfill critical ecological roles as herbivores (e.g., feeding on plant tissues), predators (e.g., targeting other arthropods), decomposers (e.g., breaking down organic matter), and pollinators (e.g., facilitating plant reproduction).^[42] The global distribution of Neoptera is cosmopolitan, spanning all continents and major biomes, with the highest species diversity concentrated in tropical regions due to the latitudinal diversity gradient, where stable climates and resource availability support elevated richness compared to temperate or polar zones.^[43] Neopterans exhibit remarkable adaptations to environmental extremes; for instance, certain butterflies (Lepidoptera) have evolved physiological and genetic traits enabling persistence at high altitudes above 4,000 meters, such as enhanced oxygen uptake and UV-protective wing pigmentation.^[44] In polar regions, non-biting midges (Chironomidae, Diptera) dominate, with species tolerating subzero temperatures and short growing seasons in Arctic and Antarctic tundras, where they form key components of aquatic and terrestrial food webs.^[45] Neopterans engage in significant ecological interactions that shape ecosystems and human societies. Symbiotic relationships, such as ant-plant mutualisms where ants defend plants from herbivores in exchange for nectar or shelter, enhance plant fitness and biodiversity in tropical forests.^[46] Conversely, outbreaks of locusts (Orthoptera: Acrididae) devastate agriculture and rangelands across arid and semi-arid regions, consuming vast vegetation and disrupting food security for millions.^[47] Beneficial interactions include the domestication of silkworms (Bombyx mori, Lepidoptera), which provide silk for global textile industries, supporting economic livelihoods in sericulture-dependent communities.^[48] Ongoing climate change is altering these distributions, driving poleward and upslope range shifts in many neopteran species, potentially leading to novel assemblages and increased extinction risks in montane and boreal habitats.^[49]

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