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Pterygota
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Pterygota
Temporal range: Serpukhovian–Recent
Clockwise from top left:
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Dicondylia
Subclass: Pterygota
Gegenbaur, 1878[1]
Subdivisions

Pterygota (/ˌtɛrəˈɡoʊtə/ terrə-GOH-tə[2] Ancient Greek: πτερυγωτός, romanized: pterugōtós, lit. 'winged') is a subclass of insects that includes all winged insects and groups which lost them secondarily.[3]

Pterygota group comprises 99.9% of all insects.[4] The orders not included are the Archaeognatha (jumping bristletails) and the Zygentoma (silverfishes and firebrats), two primitively wingless insect orders. Unlike Archaeognatha and Zygentoma, the pterygotes do not have styli or vesicles on their abdomen (also absent in some zygentomans), and with the exception of the majority of mayflies, are also missing the median terminal filament which is present in the ancestrally wingless insects.[5][6][7]

The oldest known representatives of the group appeared during the mid-Carboniferous, around 328–324 million years ago, and the group subsequently underwent rapid diversification. Claims that they originated substantially earlier during the Silurian or Devonian based on molecular clock estimates are unlikely based on the fossil record, and are likely analytical artefacts.[8]

Systematics

[edit]

Traditionally, this group was divided into the infraclasses Paleoptera and Neoptera.[9] The former are nowadays strongly suspected of being paraphyletic, and better treatments (such as dividing or dissolving the group) are presently being discussed[citation needed]. In addition, it is not clear how exactly the neopterans are related among each other. The Exopterygota might be a similar assemblage of rather ancient hemimetabolous insects among the Neoptera like the Palaeoptera are among insects as a whole. The holometabolous Endopterygota seem to be very close relatives, indeed, but nonetheless appear to contain several clades of related orders, the status of which is not agreed upon.[citation needed]

The following scheme uses finer divisions than the one above, which is not well-suited to correctly accommodating the fossil groups.

Classification

[edit]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Pterygota

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Pterygota is the subclass of winged s within the class Insecta, encompassing all that possess functional wings as adults or have secondarily lost them through evolution, while deriving from winged ancestors. This diverse group includes nearly all of the approximately 1.06 million described insect species worldwide as of 2025, excluding only the primitively wingless orders and . The name Pterygota originates from the Greek "pteron," meaning , reflecting the defining presence of veined wings attached to the meso- and metathoracic segments in most members. Key characteristics include the ability to undergo metamorphosis—either incomplete (hemimetabolous) or complete (holometabolous)—and the absence of pregenital abdominal appendages in adults, with mandibles typically adapted for chewing unless modified. Unlike the wingless , pterygotes exhibit a adapted for flight or derived from flying forms. Pterygotes dominate global ecosystems, occupying terrestrial, freshwater, and some marine habitats. They play crucial ecological roles, such as (e.g., by bees and ), predation (e.g., dragonflies), and decomposition (e.g., beetles). Economically, they are vital for agriculture through pollination services valued at billions annually, but also significant as pests (e.g., locusts, damaging crops) and vectors of diseases (e.g., mosquitoes transmitting ). Many contribute to , forensics, and industry, like silk production from silkworms. Their supports food webs and nutrient cycling, underscoring their importance to environmental stability. Taxonomically, Pterygota is broadly divided into the infraclasses Paleoptera (primitive winged forms like mayflies and dragonflies) and the more diverse Neoptera (including insects with foldable wings, subdivided by metamorphosis type). This framework encompasses over 30 orders, reflecting extensive evolutionary radiation. Detailed classification is covered in the Taxonomy section. Pterygota originated in the mid-Carboniferous period around 325 million years ago from wingless ancestors, enabling flight and adaptive success across environments. Further evolutionary details are discussed in the Evolution section.

Overview

Definition and Scope

Pterygota is a major within the class Insecta, encompassing all winged as well as secondarily wingless forms that have evolved from winged ancestors. This subclass is defined primarily by the presence of wings in its members, typically two pairs arising from the meso- and metathoracic segments, which support a remodeled adapted for flight. Pterygota constitutes approximately 99.9% of all known , highlighting its dominance in insect diversity. The name Pterygota derives from "pterygo," meaning "," reflecting the defining characteristic of wing development in this group. In taxonomic scope, Pterygota excludes the primitively wingless , which comprises only the orders (jumping bristletails) and (silverfish and firebrats), but includes all remaining orders such as Ephemeroptera, , and Coleoptera. Over one million species of Pterygota have been formally described, with estimates suggesting millions more remain undescribed, underscoring the vast unexplored biodiversity within this clade. Phylogenetically, Pterygota occupies a central position within the subphylum Hexapoda as the primary winged lineage of Insecta.

Ecological and Economic Importance

Pterygota, encompassing the vast majority of insect species with wings, play pivotal roles in terrestrial and freshwater ecosystems as pollinators, predators, decomposers, and prey. Many species, such as bees in the order Hymenoptera, are essential pollinators that facilitate plant reproduction and maintain biodiversity by transferring pollen between flowers; insects support pollination for approximately 75% of global flowering plants and 35% of the world's food crops. Predatory pterygotes, including lady beetles and lacewings, regulate herbivore populations, preventing outbreaks that could devastate vegetation and crops, thus stabilizing food webs. Decomposers like dung beetles in Coleoptera break down organic matter, accelerating nutrient recycling and soil fertility; for instance, these beetles bury dung to process vast quantities, reducing parasite transmission and enhancing pasture productivity. As abundant prey, pterygotes form the base of many food chains, providing nutrition for birds, bats, reptiles, and other insects, which underscores their integral position in trophic dynamics. Economically, pterygota contribute both benefits and challenges to human societies. Positive impacts include the production of honey by bees, a global industry valued at USD 9.73 billion as of 2024, and silk from silkworms (Lepidoptera), which supports a textile market valued at USD 20 billion as of 2024. Conversely, negative effects arise from pests like aphids (Hemiptera) that damage crops by feeding on sap, causing billions in agricultural losses yearly, and disease vectors such as mosquitoes (Diptera), which transmit pathogens like malaria and dengue, leading to over 700,000 human deaths annually. The ecological services provided by pterygota, including pollination, pest control, and decomposition, are estimated to be worth at least $57 billion per year in the United States alone, highlighting their indispensable value to economies reliant on agriculture and natural resources. Pterygota dominate global biomass and , comprising about two-thirds of terrestrial biomass in soils and achieving peak diversity in tropical , where they account for over 80% of species in canopy and layers. These hotspots, such as the Amazon, host millions of pterygote species that drive functioning, though recent studies indicate biomass declines of up to 60-fold in some rainforest sites due to environmental pressures. Populations of pterygota face severe threats from habitat loss and , which exacerbate declines and risk cascading disruptions. Deforestation and land-use changes have fragmented habitats, reducing pterygote abundance by up to 45% in affected areas, while rising temperatures and events increase extinction risks for 25-66% of studied by altering life cycles and distributions. These pressures threaten vital services like and nutrient cycling, potentially undermining and worldwide.

Taxonomy

Historical Classification

The classification of Pterygota, encompassing all winged insects and those that secondarily lost wings, began with early taxonomic efforts that recognized winged forms as a distinct group within Insecta but without formal subclass status. In the 18th century, laid foundational work in his (10th edition, 1758), dividing insects primarily based on wing characteristics into orders such as Aptera (wingless), Diptera (two wings), and several quadri-winged groups like Coleoptera, , and , effectively lumping winged insects together under broader Insecta categories without distinguishing their . This approach reflected an artificial system focused on morphology rather than phylogeny, influencing 19th-century entomologists who expanded on wing-based groupings but struggled with paraphyletic arrangements that mixed primitive and derived forms. By the late 19th century, Friedrich Moritz Brauer advanced the framework in his 1885 work, formally proposing the division of into the subclasses Apterygogenea for primitively wingless and Pterygogenea for winged or secondarily wingless . Brauer recognized around 16 orders within Pterygogenea, such as Ephemeroptera and Isoptera, though his system retained challenges in resolving evolutionary relationships due to reliance on superficial traits like wing venation. These divisions highlighted ongoing debates over , as some groups appeared transitional between wingless and winged lineages, prompting refinements in the early 20th century. Significant progress occurred in the 1920s through the paleontological and morphological analyses of Andrey Martynov, who between 1923 and 1938 proposed the key subdivision of Pterygota into Paleoptera and based on wing articulation mechanisms. Paleoptera, including orders like Ephemeroptera and , featured wings incapable of folding over the abdomen due to the absence of a jugum and fixed articulation, while encompassed the majority of winged with flexible wings that could fold. Martynov's scheme addressed earlier issues by emphasizing functional and structural homologies, though it sparked debates on the of Paleoptera, with some fossil evidence suggesting alternative arrangements. In the late 20th century, Niels Kristensen's 1991 cladistic analysis marked a pivotal shift toward phylogenetic , integrating morphological characters to hypothesize relationships among extant hexapod orders within Pterygota. Kristensen supported Martynov's Paleoptera-Neoptera dichotomy but refined it with synapomorphies like structure and genital segmentation, while critiquing unresolved polytomies in basal lineages and advocating for character polarization to resolve paraphyletic groupings. This work facilitated the transition from phenetic to cladistic methods, setting the stage for later molecular refinements.

Modern Phylogenetic Framework

The modern phylogenetic framework of Pterygota is primarily based on integrative analyses combining morphological traits with extensive molecular datasets, particularly from transcriptomic and genomic sequencing efforts. Pterygota is divided into two main infraclasses: Paleoptera, comprising Ephemeroptera (mayflies) and (dragonflies and damselflies), and , which encompasses all remaining winged insect orders. While Paleoptera is traditionally defined by the inability to fold wings over the , recent phylogenomic studies indicate it may be paraphyletic, with Ephemeroptera diverging basally and Odonata positioned closer to Neoptera. Within , three major clades are consistently supported: (including like grasshoppers and like stick insects), (encompassing such as true bugs and like booklice), and (featuring Coleoptera beetles and butterflies). These clades emerged through successive divergences, with branching first, followed by the clade uniting and . This structure underscores Neoptera's and highlights adaptive radiations, such as the complete in . Phylogenomics has been pivotal in resolving these relationships, leveraging large-scale data from the 1KITE project and similar initiatives since the . For instance, analyses of over 1,400 single-copy nuclear genes across hundreds of species have confirmed Eumetabola's and dated key divergences, such as Neoptera's origin around 350 million years ago. These studies integrate Bayesian and maximum likelihood methods to overcome limitations of earlier morphology-based or small-gene phylogenies, providing robust support for the overall framework. Despite these advances, some issues remain unresolved, including the precise placement of (webspinners) within and lingering debates on Strepsiptera's (twisted-wing parasites) affinity, often positioned near Coleoptera but with variable support across datasets. Ongoing genomic sampling, especially for understudied lineages, is expected to clarify these positions.

Morphology

Wing Structure and Venation

The wings of Pterygota are primarily membranous structures composed of a double-layered that forms a thin, flexible , supported by a network of tubular veins that provide rigidity, facilitate circulation, and house tracheae and nerves. These veins, which are sclerotized ridges of the , alternate between convex and concave sectors to enhance structural strength and allow deformation during flight. In some orders, such as Coleoptera, the forewings are modified into hardened elytra—sclerotized protective covers with reduced or obscured venation—that shield the delicate hindwings when at rest, while the hindwings remain membranous and folded beneath. Wing venation in Pterygota consists of principal longitudinal veins—costa (C), subcosta (Sc), radius (R), media (M), cubitus (Cu), and anal (A)—interconnected by cross-veins that form enclosed cells, with the arrangement varying by clade for aerodynamic efficiency and taxonomic identification. In primitive Paleoptera, such as , venation is reticulate with numerous cross-veins creating a dense network (archedictyon) for support in powerful flight, including specialized features like the nodus—a transverse and point midway along the leading edge that absorbs stress and enables wing twisting. In contrast, exhibit reduced venation with fewer cross-veins and fused longitudinal veins, facilitating wing folding over the abdomen via an elastic and flexor muscle attached to the third axillary sclerite, as seen in Diptera where only four main longitudinal veins and two cross-veins remain. Hymenoptera display further simplification, with looped anal veins and minimal cross-veins, exemplified by hamuli—tiny hooks along the hindwing's costal margin that interlock with a forewing fold for coupling. To maintain flight stability, Pterygota employ diverse wing mechanisms that synchronize fore- and hindwing motion by linking them during flapping. These include frenulum-retinaculum , where a or spine (frenulum) on the hindwing engages a hook or scale (retinaculum) on the forewing, common in and Trichoptera; jugate , involving overlap of the forewing's jugal lobe with the hindwing, typical in basal ; and amplexiform , relying on broad marginal overlap without specialized structures. Hamulate via hamuli predominates in , ensuring efficient power transfer while allowing detachment at rest. Secondary wing loss has occurred independently in several Pterygota lineages adapted to non-flying lifestyles, such as or social castes, yet vestigial traces of ancestral venation persist in developmental stages. In Siphonaptera (fleas), adults are completely wingless with a highly modified , but pupae retain small mesothoracic vestiges indicative of the lost winged condition. Similarly, in Formicidae (), wingless workers lack functional wings, while alates (reproductive castes) possess them with characteristic reduced venation—such as 16 distinct forewing patterns across subfamilies, featuring fused media and veins—revealing the ancestral pterygote blueprint before caste-specific reduction.

Body Plan Adaptations

The body plan of Pterygota is characterized by a tripartite organization into head, thorax, and abdomen, reflecting an evolutionary specialization for sensory integration, locomotion, and reproduction, respectively. The head capsule encloses the brain and subesophageal , bearing paired compound eyes composed of ommatidia for wide-field vision and filiform or otherwise segmented antennae that function primarily in chemoreception and mechanoreception. Mouthparts, derived from appendages of the head's mandibular, maxillary, and labial segments, exhibit extensive modifications; for instance, orthopterans retain ancestral chewing mandibles for processing solid plant material, while hemipterans have evolved elongate stylets for piercing plant tissues or animal hosts to extract fluids. The thorax, comprising pro-, meso-, and metathoracic segments, serves as the primary locomotor tagma, with robust indirect flight musculature integrated into its endoskeleton; in many neopterans, the prothorax is notably reduced in size and sclerotization compared to the enlarged meso- and metathorax, optimizing weight distribution for flight while retaining three pairs of jointed legs adapted for diverse functions such as cursorial walking or saltatorial leaping. The abdomen, typically formed from 11-12 segments (with the terminal ones often modified or fused), accommodates the visceral organs and external genitalia; it frequently bears cerci as paired, multi-segmented sensory appendages at the posterior end for detecting vibrations or air currents, and in females, valvular ovipositors derived from appendages of segments 8 and 9 facilitate precise egg deposition into substrates. Genital structures display profound interspecific diversity, including asymmetric aedeagi in males and variably shaped spermathecae in females, which enforce mechanical reproductive isolation via lock-and-key or traumatic insemination mechanisms that prevent interspecific mating. Development in Pterygota involves , broadly categorized as hemimetaboly in exopterygotes (e.g., orthopterans and hemipterans), where aquatic or terrestrial nymphs progressively resemble adults with external pad development through gradual molts, and holometaboly in endopterygotes (e.g., lepidopterans and hymenopterans), featuring distinct larval instars specialized for feeding, a non-feeding pupal stage for internal reorganization including eversion, and emergence of the sexually mature adult. These metamorphic strategies enable niche partitioning between juvenile and adult phases, enhancing survival and dispersal. Pterygota encompass an extraordinary size range, from minute in the family Mymaridae with body lengths as small as 0.2 mm, adapted for parasitizing tiny eggs, to expansive moths such as exhibiting wingspans up to 30 cm, which support flight over long distances.

Evolution

Origin of Wings

The origin of wings in Pterygota remains a subject of ongoing , with two primary hypotheses dominating the discussion: the paranotal theory and the exite theory. The paranotal theory posits that wings evolved as flattened outgrowths or projections from the tergum (dorsal body wall) of the , initially serving as planar structures for or before enabling powered flight. In contrast, the exite theory suggests wings arose from gill-like appendages (exites) on the proximal segments of thoracic legs, homologous to structures in ancestors, which were co-opted for aerial locomotion. Developmental genetic evidence, particularly the expression patterns of the Distal-less (Dll) gene, supports the exite hypothesis, as Dll is activated in both leg branch primordia and wing discs across diverse insect species, indicating a shared developmental origin. Earliest potential evidence for winged insects comes from Devonian trace fossils dating to approximately 400 million years ago (Mya), but these are highly debated and often interpreted as non-insectan tracks rather than definitive pterygote remains. Additional debated body fossils, such as the early hexapod Strudiella devonica from ~370 Mya, suggest pre-Carboniferous insect diversification but lack preserved wings and remain controversial. The scientific consensus places the origin of true Pterygota in the mid-Carboniferous period, around 328–324 Mya, based on the first unambiguous fossils of winged from that era. The evolution of wings conferred significant adaptive advantages, primarily enhancing escape from predators through rapid flight, facilitating dispersal to new habitats, and improving access to patchy resources such as food and mates. These benefits likely drove the rapid diversification of Pterygota, transforming insects into highly mobile organisms capable of exploiting three-dimensional space. Molecular clock analyses, calibrated against constraints, estimate the divergence of crown-group Pterygota between 400 and 350 Mya, predating the oldest confirmed and suggesting an initial phase of cryptic evolution in the Late .

Fossil Record and Timeline

The record of Pterygota begins in the mid-Carboniferous period, with the oldest unambiguous winged dating to approximately 328–324 million years ago (Mya) during the Namurian stage of the early Pennsylvanian subperiod. These early specimens, such as an Archaeorthopteran from the Ostrava Formation in the Ostrava-Karviná Basin in the , represent primitive flying with elongate wings exhibiting simple venation patterns, providing the first direct evidence of pterygote flight capabilities. This discovery marks a significant gap in the pre-Carboniferous record, as earlier traces and body remain equivocal and likely pertain to non-pterygote hexapods or other arthropods. During the late and Permian (roughly 307–252 Mya), Pterygota underwent substantial diversification, particularly in the Paleozoic of Euramerica and . Notable among these are the giant griffenflies of the extinct order , stem-group relatives of modern , which achieved remarkable sizes with wingspans reaching up to 70 cm in species like Meganeuropsis permiana from Permian deposits in . These predators, preserved in sites such as the Mazon Creek Lagerstätte in , illustrate early adaptations for aerial predation and highlight a radiation of paleodictyopteroids, odonatoids, and orthopteroids, with over 5,500 pterygote occurrences documented from Pennsylvanian–Permian strata. This era saw the emergence of key lineages like early and , though many orders, including , declined toward the end-Permian mass extinction. The era (252–66 Mya) witnessed the explosive expansion of , the endopterygote encompassing over 60% of extant species, with diversification accelerating from the into the . evidence from compression deposits like the in reveals modern-like forms, including early and Coleoptera, adapted to angiosperm-pollinated ecosystems. Amber inclusions from sites, such as Burmese () and Lebanese ambers (~99–66 Mya), preserve detailed specimens of holometabolous insects like bees and flies, indicating behavioral and morphological similarities to living taxa amid rising global temperatures and plant diversity. In the (66 Mya–present), Pterygota exhibited resilience and recovery following the Cretaceous–Paleogene (K–Pg) , with minimal lineage turnover compared to vertebrates; most orders persisted across the boundary, leading to renewed dominance in forests. Eocene amber from the (~44 Mya), containing over 30,000 insect inclusions across 6,000 species, documents a hyperdiverse assemblage of Diptera, , and Coleoptera that closely mirrors contemporary faunas, underscoring rapid recolonization and adaptation to post-extinction niches. This period's record, enriched by sites like the in , reflects ongoing radiations driven by climatic warming and ecological opportunities, culminating in the group's current representation of over one million described species.

Diversity

Major Orders and Clades

Pterygota encompasses approximately 28 extant orders of insects, representing the vast majority of insect diversity through its major clades: Paleoptera, Polyneoptera, Paraneoptera, and Holometabola. These clades are defined primarily by phylogenetic relationships and developmental patterns, with Paleoptera and the subgroups of Neoptera (Polyneoptera, Paraneoptera, and Holometabola) forming the core structure of winged insect evolution. The Holometabola alone accounts for over 80% of all described insect species, highlighting its dominance in species richness. The Paleoptera clade includes two extant orders: Ephemeroptera (mayflies), characterized by aquatic nymphs that undergo incomplete metamorphosis, and (dragonflies and damselflies), known for their predatory adult flight and aquatic larval stages. These orders retain primitive wing articulation that prevents folding over the , distinguishing them from more derived neopteran groups. comprises around 10 orders, including (grasshoppers and crickets), (stick insects), Dermaptera (earwigs), (cockroaches), Mantodea (mantises), (stoneflies), and others such as Embioptera, Zoraptera, Grylloblattodea, and Mantophasmatodea. This clade is united by shared traits like direct wing articulation and hemimetabolous development, with a primarily terrestrial inferred from phylogenetic analyses. Paraneoptera, also known as the hemipteroid assemblage, includes three orders: (true bugs, , and related groups with piercing-sucking mouthparts), (lice and booklice), and (thrips). These orders exhibit hemimetabolous and specialized reductions in mouthpart structure, contributing to their adaptation for fluid-feeding and parasitic lifestyles. The , or Endopterygota, is the most species-rich with 11 orders, featuring complete involving distinct larval, pupal, and adult stages. Key orders include Coleoptera (beetles, the largest order by species count), (butterflies and moths), Diptera (flies), and (bees, , and wasps), which together represent over 99% of holometabolan diversity. This clade's success is attributed to its developmental flexibility, enabling diverse ecological roles from to predation.

Global Species Distribution

Pterygota, the of winged and secondarily wingless , accounts for approximately 1 million described worldwide, comprising nearly all known insect diversity, with total estimates ranging from 5.5 million to 10 million when including undescribed taxa. The orders Coleoptera and dominate this diversity numerically, with Coleoptera encompassing about 400,000 described and around 180,000, highlighting their roles as the most speciose groups within the clade. These figures underscore the immense scale of Pterygota , driven largely by adaptive radiations in varied habitats. Biogeographic patterns reveal a strong concentration of Pterygota in tropical regions, which harbor over 80% of global insect diversity, including high in areas like the where exhibit exceptional local exceeding 100 ichneumonid wasps per site. In temperate zones, diversity shifts toward dominance by orders such as , which achieve peak in grasslands, and Diptera, which maintain broad ecological versatility across cooler climates. These patterns reflect adaptations to regional environmental conditions, with tropical forests supporting complex multilayered canopies that foster , while temperate ecosystems favor more generalized herbivores and decomposers. A pronounced latitudinal diversity gradient characterizes Pterygota distribution, with peaking in equatorial zones and steadily declining toward higher latitudes due to constraints like and stability. Notable exceptions occur in polar regions, such as , where endemic Diptera like the Belgica antarctica represent the only native , thriving in extreme conditions through physiological adaptations. Endemism hotspots amplify Pterygota diversity through isolated radiations, as seen in the Hawaiian archipelago, where over 500 species of Drosophila have evolved endemically since a single colonization event more than 25 million years ago. Similarly, hosts extraordinary insect , with freshwater insects showing levels up to 100% in certain lineages, yet these populations face acute threats from and , which have reduced forest cover by over 80% in recent centuries. Such vulnerabilities highlight the need for targeted conservation to preserve these irreplaceable distributions.

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