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Originally appearing in Volume V06, Page 670 of the 1911 Encyclopedia Britannica.
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COLEOPTERA, a term used in zoological classification for the true beetles which form one of the best-marked and most natural of the orders into which the class HexaDoda (or Insecta) has been divided. For the relationship of the Coleoptera to other ordersof insects see HEXAPOD. The name (Gr. KoXeos, a sheath, and 7rmepa, wings) was first used by Aristotle, who noticed the firm protective sheaths, serving as coverings for the hind-wings which alone are used for flight, without recognizing their correspondence with the fore-wings of other insects. These firm fore-wings, or elytra (fig. 1, A), are usually convex above, with straight hind margins (dorsa); when the elytra are closed, the two hind margins come together along the mid-dorsal line of the body, forming a suture. In many beetles the hind-wings are reduced to mere vestiges useless for flight, or are altogether absent, and in such cases the two elytra are often fused together at the suture; thus organs originally intended for flight have been transformed into an armour-like covering for the beetle's hind-body. In correlation with their heavy build and the frequent loss of the power of flight, many beetles are terrestrial rather than aerial in habit, though a large proportion of the order can fly well. Aristotle's term was adopted by Linnaeus (1758), and has been universally used by zoologists. The identification of the elytra of beetles with the fore-wings of other insects has indeed been questioned (188o) by F. Meinert, who endeavoured to compare them with the tegulae of Hymenoptera, but the older view was securely established by the demonstration in pupal elytra by J. G. Needham (1898) and W. L. Tower (1903), of nervures similar to those of the hind-wing, and by the proof that the small membranous structures present beneath the elytra of certain beetles, believed by Meinert to represent the whole of the true fore-wings, are in reality only the alulae. Structure.—Besides the conspicuous character of the elytra, beetles are distinguished by the adaptation of the jaws for biting, the mandibles (fig. 1, Bb) being powerful, and the first pair of maxillae (fig. 1, Bc) usually typical in form. The maxillae of the second pair (fig. 1, Bd) are very intimately fused together to form what is called the " lower lip " or labium, a firm trans-verse plate representing the fused basal portions of the maxillae, which ma.y carry a small median " ligula," representing apparently the fused inner maxillary lobes, a pair of paraglossae (outer maxillary lobes), and a pair of palps. The feelers of beetles differ greatly in the different families (cf. figs. 2 b, 9b and 26 b, c); the number of segments is usually eleven, but may vary from two to more than twenty. The head is extended from behind forwards, so that the crown (epicranium) is large, while the face (clypeus) is small. The chin (gula) is a very characteristic stlerite in beetles, absent only in a few families, such as the weevils. There is usually a distinct labrum (fig. 1, Ba). The prothorax is large and " free," i.e. readily movable on the mesothorax, an arrangement usual among insects with the power of rapid running. The tergite of the prothorax (pronotum) is prominent in all beetles, reaching back to the bases of the elytra and forming a substantial shield for the front part of the body. The tergal regions of the mesothorax and of the meta-thorax are hidden under the pronotum and the elytra when the latter are closed, except that the mesothoracic scutellum is often visible—a small triangular or semicircular plate between the bases of the elytra (fig. 1, A). The ventral region of the thoracic skeleton is complex, each segment usually possessing a median sternum with paired episterna (in front) and epimera (behind). The articular surfaces of the haunches (coxae) of the fore-legs are often conical or globular, so that each limb works in a ball-and-socket joint, while the hind haunches are large, displacing the ventral sclerites of the first two abdominal segments (fig. 1, C). The legs themselves (fig. 1, A) are of the usual insectan type, but in many families one, two, or even three of the five foot-segments may be reduced or absent. In beetles of aquatic habit the intermediate and hind legs are modified as swimming-organs (fig. 2, a), while in many beetles that burrow into the earth or climb about on trees the fore-legs are broadened and strengthened for digging, or lengthened and modified for clinging to branches. The hard fore-wings (elytra) are strengthened with marginal ridges, usually inflected ventrally to form epipleura which fit accurately along the edges of the abdomen. The upper surface of the elytron is sharply folded inwards at intervals, so as to give rise to a regular series of external longitudinal furrows (striae) and to form a set of supports between the two chitinous layers forming the elytron. The upper surface often shows a number of impressed dots (punctures). Along the sutural border of the elytron, the chitinous lamella forms a tubular space within which are numerous glands. The glands occur in groups, and lead into common ducts which open usually so much reduced that the foremost apparent ventral sclerite of the abdomen represents the third sternite. From this point backwards the successive abdominal segments, as far as the seventh or eighth, can be readily made out. The ninth and tenth segments are at most times retracted within the eighth. The female can protrude a long flexible tube in connexion with the eighth segment, carrying the sclerites of the ninth at its extremity, and these sclerites may carry short hairy processes A C Under side. 3-foxlike. FIG. I.—Structure of Male Stag-Beetle (Lucanus cervus). A, Dorsal view; B, mouth organs; C, under side. in several series along the suture. Sometimes the glands are found beneath the disk of the elytron, opening by pores on the surface. The hind-wings, when developed, are characteristic in form, possessing a sub-costal nervure with which the reduced radial nervure usually becomes associated. There are several curved median and cubital nervures and a single anal, but few cross nervures or areolets. The wing, when not in use, is folded Larva of Dyticus Cybister sp. (Water-Beetle). both lengthwise and transversely, and doubled up beneath the elytron; to permit the transverse folding, the longitudinal nervures are interrupted. Ten segments can be recognized—according to the studies of K. W. Verhoeff (1894–1896)—in a beetle's abdomen, but the tenth sternite is usually absent. On account of the great extension of the metathorax and the haunches of the large hind-legs, the first abdominal sternite is wanting, and the second is—the stylets. This flexible tube is the functional ovipositor, the typical insectan ovipositor with its three pairs of processes (see HnXAPODA) being undeveloped among the Coleoptera. In male beetles, however, the two pairs of genital processes (paramera) belonging to the ninth abdominal segment are always present, though sometimes reduced. Between them is situated, sometimes asymmetrically, the prominent intromittent organ. In the structure of the digestive system, beetles resemble most other mandibulate insects, the food-canal consisting of gullet, crop, gizzard, mid-gut or stomach, intestine and rectum. The stomach is beset throughout its length with numerous small, finger-like caecal tubes. The excretory (malpighian) tubes are few in number, either four or six. Many beetles have, in connexion with the anus, glands which secrete a repellent acid fluid, serving as a defence for the insect when attacked. The " bombardier " ground beetles (fig. 5) have this habit. Oil-beetles (figs. 23 and 24) and ladybirds (fig. 32) defend them-selves by ejecting drops of fluid from the knee-joints. The nervous system is remarkably concentrated in some beetles, the abdominal ganglia showing a tendency to become shifted forward and crowded together, and in certain chafers all the thoracic and abdominal ganglia are fused into a single nerve-centre situated in the thorax,—a degree of specialization only matched in the insectan class among the Hemiptera and some muscid flies. Development.—The embryonic development (see HExAroDA) has been carefully studied in several genera of beetles. As regards growth after hatching, all beetles undergo a " complete " metamorphosis, the wing-rudiments developing beneath the cuticle throughout the larval stages, and a resting pupal stage intervening between the last larval instarl and the imago. The coleopterous pupa (figs. 2d, 3 c) is always " free," the legs, wings and other appendages not being 1 Instar is a convenient term suggested by D. Sharp to indicate a stage in the life-history of an insect between two successive castings of the cuticle. fixed to the body as in the pupa of a moth, and the likeness of pupa to perfect insect is very close. The most striking feature in the development of beetles is the great diversity noticeable in the outward form of the larva in different families. The larva of a ground-beetle or a carnivorous water-beetle (fig. 2 c) is an active elongate grub with well-armoured cuticle. The head—carrying feelers, mandibles and two pairs of maxillae—is succeeded by the three thoracic segments, each bearing a pair of strong five-segmented legs, whose feet, like those of the adult, carry two claws. Ten segments can be distinguished in the tapering abdomen, the ninth frequently bearing a pair of tail-feelers (cerci), and the tenth, attached ventrally to the ninth, having the anal opening at its extremity and performing the function of a posterior limb, supporting and temporarily fixing the tail end of the insect on the surface over which it crawls. Such a typically " campodeiform " grub, moving actively about in pursuit of prey, is the one extreme of larval structure to be noticed among the Coleoptera. The other is exemplified by the white, wrinkled, soft-skinned, legless grub of a weevil, which lives underground feeding on roots, or burrows in the tissues of plants (fig. 3 b). Between these two From Chittenden, Yearbook, 1894, U.S. Dept. of Agriculture. d, C. oryzae. extremes we find various transitional forms: an active larva, as described above, but with four-segmented, single-clawed legs, as among the rove-beetles and their allies; the body well armoured, but slender and worm-like, with very short legs as in wireworms and mealworms (figs. 18, 21 b) ; the body shortened, with the abdomen swollen, but protected with tubercles and spines, and with longish legs adapted for an active life, as in the predaceous larvae of ladybirds; the body soft-skinned, swollen and caterpillar-like, with legs well developed, but leading a sluggish underground life, as in the grub of a chafer; the body soft-skinned and whitish, and the legs greatly reduced in size, as in the wood-feeding grub of a long-horn beetle. In the case of certain beetles whose larvae do not find themselves amid appropriate food from the moment of hatching, but have to migrate in search of it, an early larval stage, with legs, is followed by later sluggish stages in which legs have disappeared, furnishing examples of what is called hypermetamorphosis. For example, the grub of a pea or bean beetle (Bruchus) is hatched, from the egg laid by its mother on the carpel of a leguminous flower, with three pairs of legs and spiny processes on the prothorax. It bores through and enters the developing seed, where it undergoes a moult and becomes legless. Similarly the newly-hatched larva of an oil-beetle (Meloe) is an active little campodeiform insect, which, hatched from an egg laid among plants, waits to attach itself to a passing bee. Carried to the bee's nest, it undergoes a moult, and becomes a fat-bodied grub, ready to lead a quiet life feeding on the bee's rich food-stores. Distribution and Habits.—The Coleoptera are almost world-wide in their distribution, being represented in the Arctic regions and on almost all oceanic islands. Most of the dominant families—such as the Carabidae (ground-beetles), Scarabaeidae (chafers), or Curculionidae (weevils) have a distribution as wide as the order. But while some large families, such as the Staphylinidae (rove-beetles) are especially abundant on the great northern continents, becoming scarcer in the tropics, others, the Cicindelidae (tiger-beetles), for example, are most strongly represented in the warmer regions of the earth, and becomescarce as the collector journeys far to south or north. The distribution of many groups of beetles is restricted in correspondence with their habits; the Cerambycidae (longhorns), whose larvae are wood-borers, are absent from timberless regions, and most abundant in the great tropical forests. Some families are very restricted in their range. The Amphizoidae, for example, a small family of aquatic beetles, are known only from western North America and Eastern Tibet, while an allied family, the Pelobiidae, inhabit the British Isles, the Mediterranean region, Tibet and Australia. The beetles of the British islands afford some very interesting examples of restricted distribution among species. For example, large and conspicuous European beetles, such as the stag-beetle (fig. i, Lucanus cervus) and the great water-beetle (Hydrophilus piceus, fig. 20), are confined to eastern and southern Britain, and are unknown in Ireland. On the other hand, there are Arctic species like the ground-beetle, Pelophila borealis, and south-western species like the boring weevil, Mesites Tardyi, common in Ireland, and represented in northern or western Britain, but unknown in eastern Britain or in Central Europe. Careful study of insular faunas, such as that of Madeira by T. V. Wollaston, and of the Sandwich Islands by D. Sharp, and the comparison of the species found with those of the nearest continental land, furnish the student of geographical distribution with many valuable and suggestive facts. Notes on habit are given below in the accounts of the various families. In general it may be stated that beetles live and feed in almost all the diverse ways possible for insects. There are carnivores, herbivores and scavengers among them. Various species among those that are predaceous attack smaller insects, hunt in packs crustaceans larger than themselves, insert their narrow heads into snail-shells to pick out and devour the occupants, or pursue slugs and earthworms underground. The vegetable-feeders attack leaves, herbaceous or woody stems and roots; frequently different parts of a plant are attacked in the two active stages of the life-history; the cockchafers, for example, eating leaves, and their grubs gnawing roots. Some of the scavengers, like the burying beetles, inter the bodies of small vertebrates to supply food for themselves and their larvae, or, like the " sacred " beetle of Egypt, collect for the same purpose stores of dung. Many beetles of different families have become the "unbidden guests" of civilized man, and may be found in dwelling-houses, stores and ships' cargoes, eating food-stuffs, paper, furniture, tobacco and drugs. Hence we find that beetles of some kind can hold their own anywhere on the earth's surface. Some climb trees and feed on leaves, while others tunnel between bark and wood. Some fly through the air, others burrow in the earth, while several families have become fully adapted to life in fresh water. A large number of beetles inhabit the deep limestone caves of Europe and North America, while many genera and some whole families are at home nowhere but in ants' nests. Most remarkable is the presence of a number of beetles along the seashore between tide-marks, where, sheltered in some secure nook, they undergo immersion twice daily, and have their active life confined to the few hours of the low ebb. Stridulating Organs.—Many beetles make a hissing or chirping sound by rubbing a " scraper," formed by a sharp edge or prominence on some part of their exoskeleton, over a " file " formed by a number of fine ridges situate on an adjacent region. These stridulating organs were mentioned by C. Darwin as probable examples of the action of sexual selection; they are, however, frequently present in both sexes, and in some families also in the larvae. An account of the principal types of stridulators that have been described has been published by C. J. Gahan (19oo). The file may be on the head—either upper or lower surface—and the scraper formed by the front edge of the pro-thorax, as in various wood-boring beetles (Anobium and Scolytus). Or ridged areas on the sides of the prothorax may be scraped by " files " on the front thighs, as in some ground-beetles. Among the longhorn beetles, the prothorax scrapes over a median file on the mid-dorsal aspect of the mesothorax. In a large number of beetles of different families, stridulating areas occur on various segments of the abdomen, and are scraped by the elytra. It is remarkable that these organs are found in similar positions in genera belonging to widely divergent families, while two genera of the same family may have them in different positions. It follows, therefore, that they have been independently acquired in the course of the evolution of the Coleoptera. Stridulating organs among beetle-larvae have been noted, especially in the wood-feeding grub of the stag-beetles (Lucanidae) and their allies the Passalidae, and in the dung-eating grubs of the dor-beetles (Geotrupes), which belong to the chafer family (Scarabaeidae). These organs are described by J. C. Schiodte and D. Sharp; in the stag-beetle larva a series of short tubercles on the hind-leg is drawn across the serrate edge of a plate on the haunch of the intermediate legs, while in the Passalid grub the modified tip of the hind-leg acts as a scraper, being so shortened that it is useless for locomotion, but highly specialized for producing sound. Whatever may be the true explanation of stridulating organs in adult beetles, sexual selection can have had nothing to do with the presence of these highly-developed larval structures. It has been suggested that the power of stridulation would be advantageous to wood-boring grubs, the sound warning each of the position of its neighbour, so that adjacent burrowers may not get in each other's way. The root-feeding larvae of the cockchafer and allied members of the Scarabaeidae have a ridged area on the mandible, which is scraped by teeth on the maxillae, apparently forming a stridulating organ. Luminous Organs.—The function of the stridulating organs just described is presumably to afford means of recognition by sound. Some beetles emit a bright light from a portion of their bodies, which leads to the recognition of mate or comrade by sight. In the wingless female glow-worm (Lampyris, fig. 15 f ) the luminous region is at the hinder end, the organ emitting the light consisting, according to H. von Wielowiejski (1882), of cells similar to those of the fat-body, containing a substance that undergoes oxidation. The illumination is intermittent, and appears to be under the control of the insect's nervous system. The well-known " fire-flies " of the tropics are large click-beetles (Elateridae), that emit light from paired spots on the prothorax and from the base of the ventral abdominal region. The luminous organs of these beetles consist of a specialized part of the fat-body, with an inner opaque and an outer transparent layer. Its structure has been described by C. Heinemann, and its physiology by R. Dubois (1886), who considers that the luminosity is due to the influence of an enzyme in the cells of the organ upon a special substance in the blood. The eggs and larvae of the fire-flies are luminous as well as the perfect beetles. Fossil History.—The Coleoptera can be traced back farther in time than any other order of insects with complete trans-formations, ii the structures that have been described from the Carboniferous rocks of Germany are really elytra. In the Triassic rocks of Switzerland remains of weevils (Curculionidae) occur, a family which is considered by many students the most specialized of the order. And when we know that the Chrysomelidae and Buprestidae also lived in Triassic, and the Carabidae, Elateridae, Cerambycidae and Scarabaeidae, in Liassic times, we cannot doubt that the great majority of our existing families had already been differentiated at the beginning of the Mesozoic epoch. Coming to the Tertiary we find the Oligocene beds of Aix, of east Prussia (amber) and of Colorado, and the Miocene of Bavaria, especially rich in remains of beetles, most of which can be referred to existing genera. Classification.—The Coleoptera have been probably more assiduously studied by systematic naturalists than any other order of insects. The number of described species can now hardly be less than 100,000, but there is little agreement as to the main principles of a natural classification. About eighty-five families are generally recognized; the difficulty that confronts the zoologists is the arrangement of these families in ` superfamilies or " sub-orders." Such obvious features as the number of segments in the foot and the shape of the feeler were used by the early entomologists for distinguishing the great groups of beetles. The arrangement dependent on the number of tarsal segments—the order being divided into tribes Pentamera, Tetramera, Heteromera and Trimera—was suggested by E. L. Geoffroy in 1762, adopted by P. A. Latreille, and used largely through the 19th century. W. S. Macleay's classification (1825), which rested principally on the characters of the larvae, is almost forgotten nowadays, but it is certain that in any systematic arrangement which claims to be natural the early stages in the life-history must receive due attention. In recent years classifications in part agreeing with the older schemes but largely original, in accord with researches on the comparative anatomy of the insects, have been put forward. Among the more conservative of these may be mentioned that of D. Sharp (1899), who divides the order into six great series of families: Lamellicornia (including the chafers and stag-beetles and their allies with five-segmented feet and plate-like terminal segments to the feelers); Adephaga (carnivorous, terrestrial and aquatic beetles, all with five foot-segments); Polymorpha (including a heterogeneous assembly of families that cannot be fitted into any of the other groups); Heteromera (beetles with the fore and intermediate feet five-segmented, and the hind-feet four-segmented); Phytophaga (including the leaf-beetles, and longhorns, distinguished by the apparently four-segmented feet), and Rhynchophora (the weevils and their allies, with head prolonged into a snout, and feet with four segments). L. Ganglbauer (1892) divides the whole order into two sub-orders only, the Caraboidea (the Adephaga of Sharp and the older writers) and the Cantharidoidea (including all other beetles), since the larvae of Caraboidea have five-segmented, two-clawed legs, while those of all other beetles have legs with four segments and a single claw. A. Lameere (1900) has suggested three sub-orders, the Cantharidiformia (including the Phytophaga, the Heteromera, the Rhynchophora and most of the Polymorpha of Sharp's classification), the Staphyliniformia (including the rove-beetles, carrion-beetles and a few allied families of Sharp's Polymorpha), and the Carabidiformia (Adephaga). Lameere's classification is founded on the number of abdominal sterna, the nervuration of the wings, the number of malpighian tubules (whether four or six) and other structural characters. Preferable to Lameere's system, because founded on a wider range of adult characters and taking the larval stages into account, is that of H. J. Kolbe (1901), who recognizes three sub-orders: (i.) the Adephaga; (ii.) the Heterophaga, including the Staphylinoidea, the Actinorhabda (Lamellicornia), the Heterorhabda (most of Sharp's Polymorpha), and the Anchistopoda (the Phytophaga, with the ladybirds and some allied families which Sharp places among the Polymorpha); (iii.) the Rhynchophora. Students of the Coleoptera have failed to agree not only on a system of classification, but on the relative specialization of some of the groups which they all recognize as natural. Lameere, for example, considers some of his Cantharidiformia as the most primitive Coleoptera. J. L. Leconte and G. H. Horn placed the Rhynchophora (weevils) in a group distinct from all other beetles, on account of their supposed primitive nature. Kolbe, on the other hand, insists that the weevils are the most modified of all beetles, being highly specialized as regards their adult structure, and developing from legless maggots exceedingly different from the adult; he regards the Adephaga, with their active armoured larvae with two foot-claws, as the most primitive group of beetles, and there can be little doubt that the likeness between larvae and adult may safely be accepted as a primitive character among insects. In the Coleoptera we have to do with an ancient yet dominant order, in which there is hardly a family that does not show specialization in some point of structure or life-history. Hence it is impossible to form a satisfactory linear series. In the classification adopted in this article, the attempt has been made to combine the best points in old and recent schemes, and to avoid the inconvenience of a large heterogeneous group including the vast majority of the families.
End of Article: COLEOPTERA

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