fixed to the body as in the pupa of a moth, and the likeness of pupa to perfect insect is very close.
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| From Chittenden, Yearbook, 1894, U.S. Dept. of Agriculture. |
| Fig. 3.—Grain Weevils. a, Calandra granaria; b, larva; c, pupa; d, C. oryzae. |
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 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 longhorn 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 become scarce 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. 1, 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 (1900). The file may be on the head—either upper or lower surface—and the scraper formed by the front edge of the prothorax, 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
