


General features

In numbers of species and individuals and in adaptability and wide distribution, insects are perhaps the most eminently successful group of all animals. They dominate the present-day land fauna with about 1,000,000 described species. This represents about three-fourths of all described animal species. Entomologists estimate the actual number of living insect species could be as high as 5,000,000 to 10,000,000. The orders that contain the greatest numbers of species are Coleoptera (beetles), Lepidoptera (butterflies and moths), Hymenoptera (ants, bees, wasps), and Diptera (true flies).

Appearance and habits






Distribution and abundance


If the insects (including the young and adults of all forms) are counted on a square yard (0.84 m2) of rich moist surface soil, 500 are found easily and 2,000 are not unusual in soil samples in the north temperate zone. This amounts to roughly 4,000,000 insects on one moist acre (0.41 ha). In such an area only an occasional butterfly, bumblebee, or large beetle, supergiants among insects, probably would be noticed. Only a few thousand species, those that attack man’s crops, herds, and products and those that carry disease, interfere with human life seriously enough to require control measures.


Role in nature


Commercial significance


Insects as a source of raw materials



Insect damage to commercial products



Agricultural significance
Ecological factors



Damage to growing crops



Medical significance

Insect damage to man and livestock also may be direct or indirect. Direct injury to man by insect stings and bites is of relatively minor importance, although swarms of biting flies and mosquitoes often make life almost intolerable, as do biting midges (sand flies) and salt-marsh mosquitoes. Persistent irritation by biting flies can cause deterioration in the health of cattle. Some blowflies, in addition to depositing their eggs in carcasses, also invade the tissue of living animals and man, a condition known as myiasis. An example of an insect that causes this condition is the screwworm fly (Cochliomyia) of the southern U.S. and Central America. In many parts of the world various blowflies infest the fleece and skin of living sheep. This infestation, called sheep-strike, causes severe economic damage.

Many major diseases of man are produced by micro-organisms conveyed by insects, which serve as vectors of pathogens. Malaria is caused by the protozoan Plasmodium, which spends part of its developmental cycle in Anopheles mosquitoes. Epidemic relapsing fever, caused by spirochetes, is transmitted to man by the human louse Pediculus. Leishmaniasis, caused by the protozoan Leishmania, is carried by the sand fly Phlebotomus. Sleeping sickness in man and a group of cattle diseases that are widespread in Africa and known as nagana are caused by protozoan trypanosomes transmitted by the bites of tsetse flies (Glossina). Under nonsanitary conditions the common housefly Musca can play an incidental role in the spread of human intestinal infections (e.g., typhoid, bacillary and amebic dysentery) by contamination of human food. The tularemia bacillus can be spread by deer fly bites, the bubonic plague bacillus by fleas, and the epidemic typhus rickettsia by the louse Pediculus. Various mosquitoes spread viral diseases (e.g., several encephalitis diseases; dengue and yellow fever in man and other animals).

The relationships among the various organisms are complex. Malaria, for example, has a different epidemiology in almost every country in which it occurs, with different Anopheles species responsible for its spread. These same complexities affect the spread of sleeping sickness. The relationships between man and some diseases are indirect. Plague, a disease of rodents transmitted by flea bites, is dangerous to man only when heavy mortality among domestic rats forces their infected fleas to attack man, thereby causing an outbreak of plague. Typhus, tularemia, encephalitis, and yellow fever also are maintained in animal reservoirs and spread occasionally to man.

Control of insect damage

The historical objective of the entomologist was primarily to develop and introduce modifications into the environment in such ways that diseases will not be spread by insects, and crops will not be damaged by them. This objective has been achieved in numerous cases. For example, in many cities flies no longer play a major role in spreading intestinal infections, and improved land drainage, housing of man and animals, and insecticide use has eliminated malaria in many parts of the world.



Natural history
Life cycle




Types of metamorphosis


Types of larvae


Role of hormones








Sensory perception and reception













Insect societies




Terrestrial insects




Aquatic insects



Protection from enemies





Population regulation



Form and function
External features

The insect is covered by the cuticle, a layer of inert material laid down by a single sheet of epidermal cells. It consists mainly of chitin, a carbohydrate also known as polyacetylglucosamine, and sclerotin, a hard substance composed of protein tanned by quinones. The cuticle, which has an outer layer of waterproofing wax to prevent loss of water by evaporation, also serves as the skeleton to which the muscles are attached. In insects such as caterpillars, in which the cuticle is soft and flexible, the skeleton is of the hydrostatic type. In this type, body fluid pressure, maintained by muscle tension beneath the body wall, provides the firmness necessary for the function of muscles involved in movement. In insects with hard bodies, the cuticle is made up of hardened areas called sclerites that are connected by flexible joints. At the back of the head and in the thorax, hardened ingrowths of the cuticle, known as apodemes, furnish a kind of internal skeleton for muscular attachment.

Insect colours depend partly on pigments incorporated in the cuticle. However, the most important pigments often occur in epidermal cells below the cuticle. In butterflies and moths, pigments may be deposited in flattened hairs, or scales, covering the wings. Some of the most brilliant insect colours are not the result of pigmentation but are physical interference colours produced by fine laminae (grooves or pits) in the surface of the wing scales or the cuticle itself.


The ancestors of insects most likely had bodies consisting of many similar segments with only minor aggregation of the nervous system in the anterior (head) segment. These primitive insect ancestors probably looked something like modern centipedes, with a pair of appendages on each body segment but without a well-developed head. In present-day insects the primitive segments are grouped into three regions known as the head, thorax, and abdomen.

The first six primitive segments have fused to form the head, and the appendages of these segments have become modified into antennae that bear numerous sense organs and mouthparts that convey food to the mouth. Eyes also are prominent on the head. In most insects the mouthparts, adapted for chewing, consist of several parts; behind the upper lip or labrum is a pair of hard, toothed mandibles. These are followed by a pair of structures called first maxillae, each consisting of a bladelike lacinia, a hoodlike galea, and a segmented palp bearing sense organ. The paired second maxillae are partly fused in the midline to form the lower lip, or labium. Sometimes a median tonguelike structure, called the hypopharynx, arises from the floor of the mouth.

Insect mouthparts have been modified strikingly and reflect particular methods of feeding. The dipterans (true flies) provide instructive examples. In the primitive bloodsucking flies (e.g., the horsefly Tabanus) the mandibles and maxillae form serrated blades that cut through the skin and blood vessels of the host animal. The epipharynx and hypopharynx are elongated and grooved so that, when apposed, they form a tube for sucking blood. The tonguelike labium is used for imbibing exposed fluids. Dipteran mouthparts have evolved in two directions. In the mosquitoes (Culicidae) the mandibles, maxillae, epipharynx, and hypopharynx have become exceedingly slender stylets that form a fine bundle and are used for piercing skin and entering blood vessels. The labium, elongated and deeply grooved, serves only as a sheath for the stylet bundle. In the housefly Musca, however, mandibles and maxillae have been lost; the tonguelike labium alone remains and serves for feeding on exposed surfaces. Certain flies related to Musca have reacquired a capacity to suck blood; however, since they have lost both mandibles and maxillae, a new bloodsucking mechanism has developed. Labial teeth have evolved for cutting through the skin, and the labium itself is plunged into the tissues. The stable fly Stomoxys has an arrangement of this kind. In the tsetse fly Glossina, the labium has become a fine, needlelike structure normally protected by a sheath formed from the palps of the lost maxillae.

Other mouthpart modifications of the mouthpart components provide the cutting and sucking mouthparts of fleas (Siphonaptera), plant-sucking insects (Homoptera), bloodsucking bugs (Heteroptera), honeybees (Hymenoptera), and nectar-feeding butterflies (Lepidoptera).


The insect thorax consists of three segments (called the prothorax, mesothorax, and metathorax), which may be fused but are usually recognizable. Each segment has four groups of hard plates (sclerites); the groups are the notum (upper), the pleura (sides), and the sternum (underside). Thoracic sclerites are located on a given segment by using an appropriate prefix (pro-, meso-, meta-); for example, the notum (upper sclerite) of the prothorax is the pronotum.

Each segment bears a pair of legs, and, in the mature insect, the mesothorax and metathorax typically carry a pair of wings. Each leg always consists of five parts: a coxa articulated to the thorax, a small trochanter, a femur, a tibia, and a tarsus with one to five segments. The tarsal segments often carry claws with adhesive pads between them (arolia or pulvilli); these enable the insect to hold onto smooth surfaces. The legs may be modified for leaping, burrowing, grasping prey, or swimming in various ways.

The wings at rest may be extended permanently on each side, as in some dragonflies (Odonata), or held erect above the body, as in mayflies (Ephemeroptera); in most insects, however, they are folded against the abdomen. The wing consists of cuticular sacs that bud out from the wall of the thorax; the sacs become flattened during development, and the two membranes, pressed together, are stiffened by thickenings of the cuticle that form cylindrical veins carrying tracheae, nerves, and circulating blood to all parts of the wing. Wings utilized for flight commonly are made of thin membranous cuticle. In some insects, notably beetles (Coleoptera), the wings of the middle segment of the thorax have become thick and horny and serve as protective sheaths (elytra) of the membranous hindwings.

The locomotion of insects is effected by muscles acting on the external skeleton. In leaping insects (e.g., grasshoppers, fleas) the force of muscle contraction is used to compress a pad of an elastic protein, resilin; when the catch mechanism is released, the stored energy in the protein molecule is used to project the insect into the air. Insect flight is achieved by flapping the wings; during these movements the wing blade, twisted as it passes from elevation to depression, produces the same effect as the rotating propeller of an aircraft. Muscles capable of changing this inclination control the direction of flight. The chief flight muscles control flight in one of two ways: in dragonflies, directly on a lever at the base of each wing; but, in most insects, indirectly by deforming the shape of the thorax. The longitudinal muscles of the thorax depress the wings that are articulated with it; the vertical muscles elevate them.

In butterflies, the number of wing beats per second may be as low as 8 to 12, while the rate in mosquitoes may exceed 600. These rates can exceed the frequency of contraction and relaxation of muscles responding to nerves because the muscles, after they have begun contracting and relaxing, respond to the alternating elastic tension in the thoracic wall, where the frequency is determined by the natural periodic oscillation of the thorax. The flight of insects, despite their small size, conforms to the aerodynamic laws that regulate the flight of aircraft.


The abdomen consists of a maximum of 11 segments, although this number commonly is reduced by fusion. Appendages are usually absent except in caterpillars, which use up to five pairs of abdominal prolegs in walking, and in adult insects where the appendages at the hind end have become transformed into external genitalia. In the male these genitalia are paired claspers used to hold the female; in the female, three pairs of valvulae are used to manipulate eggs during oviposition. In some insects, notably crickets and cockroaches, two feelers, or cerci, at the hind end of the abdomen bear sense organs.

Internal features
Digestive system

The nutritive requirements of insects are much the same as those of mammals—water, inorganic ions, and essential amino acids (i.e., those that cannot be synthesized by the animal). The requirements for preformed fat and carbohydrate vary with the species. Although vitamins of the B group are needed by insects, neither vitamins A nor D are required, and many insects can synthesize ascorbic acid (vitamin C). On the other hand, insects cannot synthesize adequate quantities of cholesterol; thus, in effect, cholesterol can be defined as a vitamin for insects.

Insects that feed solely on some restricted diet (e.g., sterile blood, plant juices, refined flour) have special cells termed mycetocytes that harbour symbiotic micro-organisms; these organisms, transmitted through the egg to the next generation, benefit their host by furnishing it with an internal source of vitamins and perhaps other essential nutrients. If the symbiotic micro-organisms are removed experimentally, an insect fails to grow if not provided with a diet rich in vitamins.

The digestive system consists of a foregut formed from the mouth region (stomodaeum), a hindgut formed similarly from the anal region (proctodaeum), and a midgut (mesenteron). The foregut and hindgut are lined by cuticle continuous with that on the body surface. The mouth is followed by the muscular pharynx, which functions in sucking and swallowing, and the esophagus, which may be enlarged to form a crop. The crop discharges into the midgut, sometimes, as in cockroaches, by way of a muscular gizzard or proventriculus. The termination of the midgut is marked by the attachment of the malpighian tubules, the chief organs of excretion. The hindgut commonly consists of a narrow ileum followed by a larger and often thick-walled rectum, which discharges at the anus.

Digestive enzymes, secreted not only by the salivary glands but also by the cells of the midgut and its diverticula, vary with the diet of the insect. The most important enzyme secreted by the salivary glands is amylase; the midgut secretes several enzymes including protease, lipase, amylase, and invertase. The products of digestion are absorbed chiefly in the midgut.

The hindgut receives food residues from the midgut as well as waste products from the malpighian tubules. The end products of nitrogen metabolism are uric acid, small amounts of amino acids, and urea; in aquatic insects, ammonium salts may be a major form for nitrogen excretion. In the rectum, the epithelial cells lining the gut wall often are enlarged, particularly in restricted areas where they form rectal glands. The epithelial cells of these glands are supplied richly with tracheae and function in the reabsorption of water and ions. The rectal contents of insects that inhabit dry environments commonly are reduced to dry fecal pellets prior to discharge. In many insects, particularly those which feed on relatively dry foods (e.g., beetles infesting stored grain), the upper segments of the malpighian tubules are bound by a sheath to the rectal surface and form a cryptonephridial system that serves to increase the capacity of the rectum for reabsorbing water and salts. The products of digestion, discharged into the hemocoele, or general body cavity, are transported by the circulatory fluid, or hemolymph, to the organs.

Circulatory system

The circulatory system is an open one, with most of the body fluid, or hemolymph, occupying cavities of the body and its appendages. The one closed organ, called the dorsal vessel, extends from the hind end through the thorax to the head; it is a continuous tube with two regions, the heart or pumping organ, which is restricted to the abdomen, and the aorta, or conducting vessel, which extends forward through the thorax to the head. Hemolymph, pumped forward from the hind end and the sides of the body along the dorsal vessel, passes through a series of valved chambers, each containing a pair of lateral openings called ostia, to the aorta and is discharged in the front of the head. Accessory pumps carry the hemolymph through the wings and along the antennae and legs before it flows backward again to the abdomen.

The circulating hemolymph, or blood, is not important in respiration but functions in transporting nutrients to all parts of the body and metabolic waste products from the organs to the malpighian tubules for excretion. It contains free cells called hemocytes, most of which are phagocytes that help to protect the insect by devouring micro-organisms. An important tissue bathed by the hemolymph is the fat body, the main organ of intermediary metabolism. It serves for the storage of fat, glycogen, and protein, particularly during metamorphosis. These materials are set free as required by the tissues for energy production or for growth and reproduction.

Respiratory system

The respiratory system consists of air-filled tubes or tracheae, which open at the surface of the thorax and abdomen through paired spiracles. The muscular valves of the spiracles, closed most of the time, open only to allow the uptake of oxygen and the escape of carbon dioxide. The tracheal tubes are continuous with the cuticle of the body surface. The tracheae are stiffened by spiral thickenings or threadlike ridges called taenidia, which branch repeatedly, becoming reduced in cross section and ending in fine thin-walled tracheoles less than one micron in diameter. The tracheoles insinuate themselves between cells, sometimes appearing to penetrate into them, and push deeply into the plasma membrane.

Although movements of oxygen and carbon dioxide occur solely by gaseous diffusion in sedentary insects, the system is ventilated mechanically in active species. Pumping movements of the abdomen provide the force necessary to drive out streams of air at some spiracles and suck them in at others. The taenidia keep the tracheae distended, thus allowing free passage of air. In addition, the most active insects have large thin-walled dilatations of the tracheae called air sacs, which serve to increase the volume of air displaced during respiratory movements. Both lack of oxygen and accumulation of carbon dioxide provide stimuli to nerve centres that induce increased respiration during muscular activity.

Reproductive system

The reproductive system consists of the sex glands, or gonads (male testes and female ovaries), the ducts through which the sexual products are carried to the exterior, and the accessory glands. The two testes are made up of a variable number of follicles in which the spermatocytes mature and form packets of elongated spermatozoa. Spermatozoa, liberated in bundles with heads held in a cap of gelatinous material, accumulate in the vesicula seminalis, a dilated section of the male sexual duct (vas deferens).

Each of the two ovaries consists of a number of ovarioles. The ovarioles converge upon the two oviducts, and the oviducts unite to form a common oviduct down which the ripe eggs are discharged. Each ovariole consists of a germarium and a series of ovarial follicles. The germarium is a mass of undifferentiated cells that form oocytes, nurse cells, and follicular cells. The nurse cells provide nourishment for the oocytes during the early stages of their growth; follicular cells, which invest the enlarging oocyte as a continuous epithelium, provide the materials for yolk formation and, in the final stages, lay down the eggshell or chorion. The ovarial follicles increase progressively in size as the oocytes grow to form ripe eggs.

During copulation, bundles of spermatozoa are sometimes introduced directly into the female vagina by means of the male copulatory organ, or aedeagus. Secretions from the accessory glands of the female activate the sperm, the sperm bundles disperse, and the free spermatozoa make their way up to the receptaculum seminis, or spermatheca, where they are stored, ready to fertilize the eggs. In most insects, the male accessory glands secrete materials that form a tough capsule, or spermatophore; spermatozoa are encased in this spermatophore, which is inserted into the entrance of the vagina. The spermatophore walls commonly contain a gelatinous substance that swells upon exposure to secretions of the female and forces out the spermatozoa. The vagina serves both for receiving sperm and for laying eggs.

The terminal segments of the abdomen of females sometimes are modified to form an ovipositor used for depositing eggs. In butterflies and moths (Lepidoptera) a second copulatory canal independent of the vagina has been evolved, so that the sperm enter by one route, and the eggs are deposited by another.

The eggshell, or chorion, commonly provided with an air-filled meshwork, provides for respiration of the developing embryo. The chorion is also pierced by micropyles, fine canals that permit entry of one or more spermatozoa for fertilization. As the egg passes down the oviduct before egg laying, the micropyles come to lie opposite the duct of the spermatheca; at this stage fertilization occurs. Eggs must be waterproof to prevent desiccation; each egg has a layer of waterproofing wax, sometimes over the entire shell surface, more often lining the inside.

Nervous system

The central nervous system consists of a series of ganglia that supply nerves to successive segments of the body. The three main ganglia in the head (protocerebrum, deutocerebrum, and tritocerebrum) commonly are fused to form the brain, or supraesophageal ganglion. The rest of the ganglionic chain lies below the alimentary canal against the ventral body surface. The brain is joined by paired connectives to the subesophageal ganglion, which is linked in turn by paired connectives to the three thoracic and eight abdominal ganglia (numbered according to segment). In most insects the number of separate ganglia has been reduced by fusion. The last abdominal ganglion always serves several segments. In homopterans and heteropterans all the abdominal ganglia usually fuse with mesothoracic and metathoracic ganglia; and in the larvae of higher flies (Cyclorrhapha), the ganglia of the brain, thorax, and abdomen form one mass.

Each ganglion is made up of nerve-cell bodies that lie on the periphery and a mass of nerve fibres, the neuropile, that occupies the centre. There are two types of nerve cells, motor neurons and association neurons. Motor neurons have main processes, or axons, that extend from the ganglia to contractile muscles, and minor processes, or dendrites, that connect with the neuropile. Association neurons, usually smaller than motor neurons, are linked with other parts of the nervous system by way of the neuropile.

Cell bodies of the sense organs, called sensory neurons, lie at the periphery of the body just below the cuticle. Sensory neurons occur as single cells or small clusters of cells; the distal process, or dendrite, of each cell extends to a cuticular sense organ (sensillum). The sensilla are usually small hairs modified for perception of specific stimuli (e.g., touch, smell, taste, heat, cold); each sensillum consists of one sense cell and one nerve fibre. Although these small sense organs occur all over the body, they are particularly abundant in antennae, palps, and cerci. The sense cell of each sensillum gives off a proximal process, or sensory axon, which runs inward to the central nervous system, where it enters the neuropile and makes contact with the endings of association neurons. Bundles of both sensory axons and motor axons, which are enclosed in protective membranous sheaths, constitute the nerves.

Tactile hairs may be sensitive enough to perceive air vibrations and thus serve as organs for sound reception. Tympanal organs (eardrums) are present in certain butterflies and grasshoppers. Mechanical sensilla (chordotonal organs) below the surface of the cuticle serve for perception of internal strains and body movements.


The eyes are of two kinds, simple eyes, or ocelli, and compound eyes. In the adults of higher insects both types are present. The visual sense cells are derived from the epidermis, as are those of other sense organs, and are connected to the optic ganglia (a part of the brain) by sensory axons. Each visual sense cell has a zone at its surface, which, on exposure to light, gives rise to chemical products that stimulate the sense cell, called the retinula cell, and initiate the nerve impulse in the sensory axon. The light-receptive zone, or rhabdom, of the retinula cell commonly has a rodlike form; because it lies perpendicular to the surface, light passes lengthwise along it. In the simple eyes (ocelli) a lens-shaped area of cuticle lies over the group of retinula cells that form the retina. Since the optical structure is primitive, the visual image received is crude; ocelli can perceive only light, darkness, and movement.

The compound eye, made up of a number of facets, resembles a honeycomb; each facet overlies a group of six or seven retinal cells that surround the rhabdom. Each of the retinal units below a single facet is termed an ommatidium. The number of facets varies. For example, there are only a few dozen facets in the eye of the primitive apterygote Collembola, while the eye of the housefly Musca has some 4,000, and the highly developed eye of the dragonfly may contain up to 28,000.

During light reception, rays from a small area of the field of view fall on a single facet and are concentrated upon the rhabdom of the retinula cells below. Since each point of light differs in brightness, all the ommatidia that form the retina receive a crude mosaic of the field of view. Unlike the image in a camera or in human eyes, the mosaic image in the compound eye is not inverted but erect. The fineness of the mosaic and, therefore, the degree of resolution improves with increasing numbers of facets. It is estimated that the eye of the honeybee has visual acuity equal to 1 percent that in man.

Each ommatidium commonly is shielded by a curtain of pigmented cells that prevent the spread of light to neighbouring ommatidia. This is termed an apposition eye. In the eyes of insects that fly at night or in twilight, however, the pigment can be withdrawn so that light received from neighbouring facets overlaps to some extent. This is termed a superposition eye. The image formed is brighter but not as sharp as that formed by the apposition eye. In addition to perceiving brightness, the eyes of insects can perceive colour as well as some other properties of light.

Evolution and paleontology
Origin of insects

The most primitive insects known are found as fossils in rocks of the Middle Devonian Period and lived about 350,000,000 years ago. The bodies of those insects were divided then, as now, into a head bearing one pair of antennae, a thorax with three pairs of legs, and a segmented abdomen. Those insects originated with the terrestrial branch of the phylum Arthropoda. The Arthropoda, whose origin is thus far unknown, probably arose in Precambrian times, perhaps as much as 1,000,000,000 years ago. Some arthropods colonized the open sea and have become the present-day class Crustacea (crabs, shrimps) and the now-extinct Trilobita. Other arthropods colonized the land. This terrestrial line persists chiefly as the classes Onychophora, Arachnida (spiders, scorpions, ticks), the myriapods (consisting of Diplopoda [millipedes], Pauropoda, Symphyla, and Chilopoda, or centipedes), and finally the class Insecta.

The most primitive insects today are found among the wingless (apterous) hexapods; sometimes known collectively as apterygotes, they include proturans, thysanurans, diplurans, and collembolans. It is agreed generally that insects are related most closely to the myriapod group, among which the Symphyla exhibit most of the essential features required for the ancestral insect form (i.e., a Y-shaped epicranial suture, two pairs of maxillae, a single pair of antennae, styli and sacs on the abdominal segments, cerci, and malpighian tubules). There is, therefore, general agreement that the insects probably arose from an early symphylan-like form.

Insect fossil record

The insect fossil record has many gaps. Among the primitive apterygotes, only the collembolans (springtails) have been found as fossils in the Devonian Period. Ten insect orders are known as fossils, mostly of Late Carboniferous and Permian times. No fossils have yet been found from the Late Devonian or Early Carboniferous periods, when the key characters of present-day insects are believed to have evolved; thus, early evolution must be inferred from the morphology of extant insects.

It has become evident that insect evolution, like that of other animals, was far more active at some periods than at others. There have been geological epochs of “explosive” evolution during which many new forms have appeared. Those epochs may have followed some modification or innovation in body function, or new developments favoured by climatic changes or evolutionary advances of other animals and plants. During those periods of evolutionary change, new methods of feeding and living led to diversity of insect mouthparts and limbs, the origin of metamorphosis, and other changes.

Insect phylogeny

Figure 1 is a simplified family tree of the presumed evolutionary history of winged insects (Pterygota) throughout the geological periods from the Devonian to the Recent. The apterygotes, which are regarded as survivors of primitive insect stock, are omitted from the family tree. Dark lines indicate the periods during which the various orders have been found as fossils. Some lines stop at the names of orders now extinct and known only as fossils. Light lines indicate the hypothetical origin of various orders. Many insect types, traces of which have not yet been discovered, must have been produced during the explosive periods of evolution in Carboniferous and Permian times.

The primitive wingless insects (Figure 1) gave rise to a paleopterous stock. Descendants of this stock included ancient fossil types that flourished in Permian times, such as the giant dragonflies or Protodonata (some of which had a wing span of more than half a metre) and dragonflies and damselflies (Odonata) and mayflies (Ephemeroptera), both of which have persisted with little change to the present. The primitive insect stock also gave rise to a neopterous stock, believed to include the progenitors of the remaining insect orders. The Orthoptera (grasshoppers) and the Plecoptera (stoneflies) have been found as fossils even in late Carboniferous times. The Isoptera (termites, sometimes placed in the order Blattodea), Embioptera (webspinners), and Dermaptera (earwigs), though doubtless of ancient origin, have not been found yet as fossils dated earlier than the Mesozoic Era.

The evolutionary radiation (Figure 1), believed to have given rise to the orders listed above in the Middle Carboniferous Period, is thought to have produced also a paraneopterous stock, which formed the base for a new evolutionary radiation during the Permian Period. Present-day derivatives of this stock evolved into the Psocoptera (psocids), Mallophaga (chewing lice), Anoplura or Siphunculata (sucking lice), Thysanoptera (thrips), Heteroptera (true bugs), and Homoptera (e.g., aphids).

Several phylogenetic lines (Figure 1) are exopterygote (i.e., insects with simple metamorphosis) some of which, such as Mallophaga and Anoplura, are secondarily wingless. The remaining orders are endopterygote (insects with complete metamorphosis). They are shown in Figure 1 as derivatives of an oligoneopterous stock, which gave rise to Neuroptera (lacewings), Hymenoptera (ants, wasps, and bees), and Coleoptera (beetles) in the Early Permian Period; the early ancestry of these orders is obscure, however, and the earliest fossils closely resemble present-day forms. One line from the evolutionary radiation (Figure 1) at the beginning of the Permian gave rise to a mecopteroid stock, and there is good evidence that a sub-radiation of these mecopteroid orders (sometimes called the panorpoid complex) provided the origin for the present Mecoptera (scorpionflies), Diptera (true flies), Siphonaptera (fleas), Trichoptera (caddisflies), and Lepidoptera (butterflies and moths).

Wings and flight

Insect wings develop as paired outgrowths from the thorax, stiffened by ribs, or veins, in which run tracheae. These tracheae follow a consistent pattern throughout the Pterygota, and their specific modifications (known as venation) are important in classification and in estimations of the degree of relationship between groups. The basic consistency of venation suggests that wings have been evolved only once among the insects, that is, all the Pterygota (as shown in Figure 1) arose from a single stem. By the time (toward the end of the Carboniferous) fossil insects are found, wings are developed fully. In the Paleoptera (Figure 1) the wings are held aloft above the back, as in mayflies, or held extended permanently on each side of the body, as in dragonflies. Throughout the Neoptera there is a wing-flexing mechanism (secondarily lost in butterflies) that enables the wings to be folded back to rest on the surface of the abdomen.

Winged insects must have made their appearance very early in the Carboniferous, more than 300,000,000 years ago; but there is no fossil evidence to show the way they evolved. One hypothesis is that wings arose as fixed planes extending sideways from the thorax and that these planes were used, perhaps in some large leaping insect, for gliding. Later muscles developed, first to control inclination and then to move the wings in flapping flight. Another hypothesis is that wings may have originated from large thoracic tracheal gills, similar to the movable tracheal gills along the abdomen of some mayfly larvae. Such outgrowths could have been useful to insects exposed by the drying up of a temporary aquatic habitat and might have carried them in rain-bearing winds to a new watery home. It is likely that the most primitive symphylan-like insects were terrestrial; throughout insect evolution, however, independent adaptations to aquatic habitats have occurred. Usually the pattern is one in which the adults leave the water and disperse. Many pterygote insects have become secondarily wingless, sometimes as single species or groups of species within large orders, sometimes as entire orders (the parasitic lice, Mallophaga and Anoplura, and the fleas, Siphonaptera).


It generally is agreed that insect metamorphosis evolved as adult insects gradually adopted different modes of life from those of larvae. The characters of larva and adult became genetically independent; in response to natural selection, therefore, each was able to evolve independently of the other. Mouthparts, limbs, and other morphological features were modified in different directions and in higher groups. Where these differences were extreme, an intermediate pupal stage evolved to bridge the morphological gap between larva and adult. It seems quite probable that the development of metamorphosis occurred more than once during the evolution of insects.

Feeding methods

Insects did not evolve in a constant environment. Throughout geological time there were prodigious changes in climate; in addition, evolution was continuous among all other animals and plants. Geologically the selection pressures among insects were changing continuously. At the end of the Mesozoic Era the first flowering plants appeared. Insect evolution has paralleled that of the flowering plants; they have evolved together. As Lepidoptera (butterflies and moths), Hymenoptera (ants, bees, and wasps), Diptera (true flies), and Coleoptera (beetles) began to feed upon flowers, nectar, or pollen, flowering plants came to rely more and more upon insects—rather than upon the wind—for transferring their pollen. Flowers evolved nectaries, scents, and conspicuous colours as attractants for those insects that could effect cross-pollination. Insects likewise evolved appropriate mouthpart modifications for extracting nectar from flowers.

During the Mesozoic warm-blooded animals (mammals and birds) first appeared; by the dawn of the Tertiary Paleogene Period, they had become predominant among the earth’s large animals. The warm fermenting excrement and the decaying dead bodies of mammals furnished excellent nutrient media for many insect larvae, notably among the Diptera and Coleoptera. The adults in both groups found their nourishment in flowers. Some heteropterans (true bugs) and dipterans pierce the skin of birds and mammals and feed on their blood. The Anoplura (sucking lice) and the Siphonaptera (fleas) have become so specialized for this type of parasitic existence that their relationships to other insects are not yet known with certainty.

Continuing evolution

Evolution is occurring among present-day insects. They exhibit a balanced genetic polymorphism; in other words, in response to small environmental changes, one genetic form, more successful than another, will become more plentiful. Sometimes there is no visible difference between these forms, the advantage presumably lying in some physiological change. It is advantageous for a species to have a gene pool from which favourable characters can be selected so that the species can respond to environmental changes. Changes within a species may occur progressively over a large geographical area. Such a progressive genetic change is termed a cline; in some cases insects at the extremes of the cline are so unlike that they are taken as separate species and may be infertile when crossed.

One well-known example of evolution in action among insects is industrial melanism (accumulation of the black pigment melanin); many butterflies inhabiting industrial areas have become almost black during the past century; black forms are more tolerant of pollution and less conspicuous to predators. Another example of this cline type of evolution is the development of insect strains resistant to an insecticide that has been applied heavily in an area for several years. In many parts of the world houseflies have become highly resistant to DDT.