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The Life-Story of Insects
by Geo. H. Carpenter
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The broad end of the maggot is the tail, while the narrow extremity marks the position of the mouth. Above this are a pair of very short feelers (fig. 21 c), while from the aperture project the tips of the mouth-hooks (fig. 21 e, f), formidable, black, claw-like structures, articulated to the strong pharyngeal sclerites and moved by powerful muscles, tearing up the fibres of the flesh. On either side of the prothorax is an anterior spiracle, a curious branching or fan-like outgrowth (fig. 21 b), with a variable number of tiny openings which are probably of little use for the admission of air to the tubes. In many maggots the mouth-hooks and the front spiracles become more and more complex in form in the successive instars. The cuticle, white and smooth to the unaided eye, is seen on microscopic study to be set with rows of tiny spines which assist the maggot's movements through its food-mass. At the tail-end the large hind spiracles are conspicuous on a flattened dorsal area of the ninth abdominal segment; each shows a hard brown plate, traversed by three slits. And as we watch this curious degraded larva thrusting its narrow head-end into the depths of its ofttimes loathsome food-supply, we understand the advantage of access to the air-tube system being mainly confined to the hinder end of the body.

Maggots, differing from that of the Bluebottle only in minor details, are the larval forms of a vast multitude of allied species and display great variation in the nature of their food. Most, however, hide their soft defenceless bodies in some substance which affords shelter as well as food. The Bluebottle maggot burrows into flesh, that of the House-fly into horse-dung or vegetable refuse. The maggot of the Cabbage-fly eats its way into the roots of cruciferous plants, that of the Mangel-fly works out a broad blister between the two skins of a leaf, into which the newly-hatched larva crawls directly from the egg. A large number of species, forming an entire subfamily (the Tachininae) have larvae that feed as parasites within the bodies of other insects.

The habit of parasitism by maggots in back-boned animals has led to some remarkable modifications of the larva and to curious adventures in the course of the life-story. The Bot-fly of the Horse (Gastrophilus equi) and the Warble-fly of the Ox (Hypoderma bovis, fig. 22) lay eggs attached to the hairs of grazing animals, which, at least in the case of Gastrophilus, lick the newly-hatched larvae into their mouths. The 'bot,' or maggot of Gastrophilus, comes to rest in the horse's stomach; often a whole family attach themselves by their mouth-hooks to a small patch of the mucous coat of that organ. The maggot is relatively short and stout, with rows of strong spicules surrounding the segments, and with spiracles capable of withdrawal through a cup-like inpushing of the tail-region of the body, so that the parasite is preserved from drowning when the host drinks water. The young maggot of Hypoderma (fig. 22 e) is elongate and slender, spends its first two stages burrowing in the gullet wall and then wandering through the dorsal tissues of its host; ultimately it arrives beneath the skin of the back and assumes for its third and fourth instars a broad barrel-like form (fig. 22 b). The supply of free oxygen within the ox's tissues being now insufficient, the warble-maggot bores a circular hole through the skin and rests with the tail spiracles directed upwards towards the outer air. When fully grown the maggot works its way through the hole in the host's skin, and falling to the ground pupates in some sheltered spot, the life cycle occupying about a year. Similarly the Horse-bot escapes from the host's intestine with the excrement, and pupates on the ground.

A curious modification of the maggot is noticeable in the larva of the Hover-flies (Syrphus). These, unlike most of their allies, live exposed on the foliage of plants, where they feed by preying on aphids.



In agreement with this manner of life, the cuticle is roughly granulated, often greenish or reddish in hue, and the maggot, despite its want of definite head and sense organs, moves actively and purposefully about, often rearing up on its broad tail-end with an aphid victim impaled on its mouth-hooks.

In a previous chapter reference was made to the exopterygote insects, stone-flies, dragon-flies, and may-flies, whose preparatory stages live in the water. Among the endopterygote orders many Neuroptera and Coleoptera, all Trichoptera, a very few Lepidoptera and many Diptera, have aquatic larvae. One or two examples of the adaptations of dipteran larvae to life in the water may well bring the present chapter to a close. Many members of the hover-fly family (Syrphidae) have maggots with the tail-spiracles situated at the end of a prominent tubular process. Among the best-known of syrphid flies are the drone-flies (Eristalis), often seen hovering over flowers, and presenting a curious likeness to hairy bees. The larva of Eristalis is one of the most remarkable in the whole order, the 'Rat-tailed maggot' found in the stagnant water of ditches and pools. It has a cylindrical body with the hinder end drawn out into a long telescopic tube, a more slender terminal section being capable of withdrawal into, or protrusion from, a thicker basal portion. At the extremity of the slender tube is a crown of sharp processes, forming a stellate guard to the spiracles. These processes can pierce the surface-film of the water, and place the tracheal system of the maggot in touch with the pure upper air; while its mouth may be far down, feeding among the foul refuse of the ditch, it can still reach out to the medium in which the end of its life-story must be wrought out.

Reverting to the first great division of the Diptera, we find varied adaptations to aquatic life among many grubs that possess a definite head. The larva of a Gnat (Culex[9]) has projecting from the hind region of the abdomen a long tubular outgrowth, at the end of which are the spiracles, guarded by three pointed flaps forming a valve. When closed these pierce the surface-film of the water in which the larva lives; when opened a little cup-like depression is formed in the surface-film, from which the larva hangs. Or having accumulated a supply of air, it can disengage itself from the surface-film and dive through the water, its tracheal system safely closed. Another mode of breathing is found in the 'Blood-worms' and allied larvae of the Harlequin-midges (Chironomidae) whose transformations are described in detail by Miall and Hammond (1900). These larvae have two pairs of cylindrical, spine-bearing pro-legs—one on the prothorax and the other on the hindmost abdominal segment; the latter structures serve to fix the larva in the muddy tube which it inhabits at the bottom of its native pond. The penultimate abdominal segment has four long hollow outgrowths, which contain blood, and have the function of gills, while the hindmost segment has four shorter outgrowths of the same nature. Enabled thus to breathe dissolved air, the Chironomus larva needs not, like the Culex or the Eristalis, to find contact with the atmosphere beyond the surface-film.

[9] See Frontispiece, A.

Most remarkable, in many respects, of all aquatic larvae are the grubs of the Sand-midges (Simulium). These live entirely submerged and, having no special gills, carry out an exchange of gases through the general surface of the cuticle between the dissolved air in the water and the cavities of the air-tube system. The body is shaped like a flask swollen slightly at the hinder end and possesses a median pro-leg just behind the head, also another at the tail, which serves to attach the larva to a stone or to the leaf of an aquatic plant. The head has, in addition to feelers and jaws, a pair of processes with wonderful fringes which by their motion set up currents in the water, and bring food particles within reach of the mouth. A number of the larvae usually live in a community. Their power of spinning silken threads by which they can work their way back when accidentally dislodged from their resting-place, has been vividly described by Miall (1895).

Examples might be multiplied, but enough have been given to enforce the conclusion that the forms of insect-larvae are wondrously varied, and that frequently, within the limits of the same order or even family, modifications of type may be found which are suited to various modes of life adopted by different insects. A survey of the multitudes of insect larvae—grubs, caterpillars, maggots—living on land, on plants, underground, in the water; feeding on leaves, in stems, on roots, on carrion, on refuse; by hunting or by lurking after prey; as parasites or as scavengers, brings home to us most strongly the conclusion that each larva is fitted to some little niche in the vast temple of life, each is specially adapted to its part in the great drama of being.



CHAPTER VII

PUPAE AND THEIR MODIFICATIONS

The pupal stage is characteristic of the life-story of those insects whose larvae have wing-rudiments in the form of inpushed imaginal discs, and in all these insects there is, as we have seen, considerable divergence in form between larva and imago. In the pupa the wings and other characteristically adult structures are, for the first time, visible outwardly; it is the instar which marks the great crisis in transformation. The pupa rests, as a rule, in a quiescent condition, and during the early period of this stage the needful internal changes, the breaking down of many larval tissues, and their replacement by imaginal organs, go on. Both outwardly and inwardly therefore, the insect undergoes, at the pupal stage, a reconstruction necessitated by the differences in form and often in habit, between the larva and the winged adult; and the greater these differences, the more profound must be the changes that mark the pupal stage.

From the prominence of imaginal structures in the pupa, it is at once seen that the pupa of any insect must resemble the adult more nearly than it resembles the larva. But in different groups of insects we find different degrees of likeness between pupa and imago. In a beetle pupa (see fig. 16 c), the appendages—feelers, jaws, legs, wings—stand out from the body as do those of the perfect insect. This type is called a free pupa. The pupal cuticle has to be shed for the emergence of the imago, but the pupa is already a somewhat reduced model of the final instar, with abbreviated wings and doubled-up legs. A free pupa is characteristic of the Coleoptera, Neuroptera, Trichoptera, Hymenoptera and many Diptera. In some cases the pupa requires to be specially adapted for a peculiar mode of life; for example, a special arrangement of breathing organs may be necessary for life under water, and there must needs be temporary pupal structures, not represented in the imago.

On the other hand, in the pupae of most Lepidoptera and of some Diptera, there is more or less coalescence between the cuticle of the appendages and the cuticle of the body generally, so that the appendages do not stand out, but being, as it were, glued down to the body, are somewhat masked (see fig. 1 e and fig. 23). Consequently the obtect pupa, as this type is called, does not resemble its imago as fully as a free pupa does. The outline of the wings for example in a butterfly's pupa can in some cases be traced only with difficulty. T.A. Chapman has shown (1893) that the completely obtect pupa characterises the more highly developed families of Lepidoptera, while in the more primitive families the pupa is incompletely obtect. If the pupa of a butterfly or moth be lifted and held in the hand, a bending or wriggling motion of the abdomen can be observed. In the incompletely obtect pupa, this motion is evident in a greater number of segments than in the completely obtect, the number concerned varying from five to two in different families. In the nymphalid butterflies, the pupa is often called a 'chrysalis' on account of the golden hue displayed by the cuticle, and the term 'chrysalis' is sometimes bestowed indiscriminately on any kind of pupa. It has been shown by Poulton (1892) and others, that the colour of a butterfly pupa is to some extent affected by the surroundings of the caterpillar just before its last moult.

Reference has been made (p. 58) to the power of spinning silk possessed by many larvae; often the principal use of this silk is to form some protection for the pupa, the larva before its last moult constructing a cocoon within which the pupa may rest safely. Many larvae bury themselves in the earth, and the pupa lies in an earthen chamber, the lining particles of soil fastened together by fine silken threads. Larvae that feed in wood, like the caterpillar of the Goat-moth (Cossus) make a cocoon of splinters spun together, while hairy caterpillars, such as those of the Tiger-moths, work some of their hairs in with the silk to make a firm cocoon (fig. 17 b). On the other hand, those caterpillars known as 'silkworms' make a dense cocoon of pure silk, consisting of two layers, the outer of coarse and the inner of fine threads. Silken cocoons very similar in appearance are spun by the larvae of small Ichneumon-flies. Many pupae lie in a loose cocoon formed of a few interlacing threads, as for example the conspicuous black and yellow banded pupa of the Magpie-moth (Abraxas grossulariata) and the pupae of various leaf-beetles. Others again spin together the edges of leaves with connecting silken threads. The grubs of bees and wasps which are reared in the comb-chambers of their nests seal up the opening of the chamber with a lid, partly silk (fig. 18 co) and partly excretion, when ready to pass into the pupal state. An additional external 'capping' may be also supplied by the workers.

The pupae of butterflies are especially interesting, as illustrating the extreme reduction of the silken cocoon. The pupa of a 'swallowtail' (Papilionid) or a 'white' (Pierid) butterfly (fig. 23) may be found attached to a twig of its food-plant or to a wall, in an upright position, its tail fastened to a pad of silk and a slender silken girdle encircling its thorax. The pupa of a 'Tortoiseshell' or 'Admiral' (Nymphalid) butterfly hangs head downwards from a twig, supported only by the tail-pad of silk, which, useless as a shelter, serves only for attachment. The pupa is fastened to this pad by a spiny hook or process, the cremaster (fig. 23 cr), on the last abdominal segment. The cremaster is a characteristic structure in the pupa of a moth or butterfly. C.V. Riley (1880) and W. Hatchett-Jackson (1890) have shown that it corresponds with a spiny area, the suranal plate, which lies above the opening of the caterpillar's intestine. The means by which the suspended pupa of a nymphalid butterfly attaches its cremaster to the silken pad which the larva has spun in preparation for pupation, is worthy of brief attention. The caterpillar, hanging head downwards, is attached to the silken pad by its hindmost pair of pro-legs or claspers and by the suranal plate, and the cuticle is slowly worked off from before backwards, so as to expose the pupa. Were the process of moulting to be simply completed while the insect hangs by the claspers, the pupa would of course fall to the ground. But there is enough adhesion between the pupal and larval cuticles at the hinder end of the body, especially by means of the everted lining of the hind-gut, for the pupa to be supported while it jerks its cremaster out of the larval cuticle and works it into the meshes of the silken pad. The moult is thus completed and the pupa hangs securely all the time. In the numerous cases where the pupa is enclosed in a cocoon, the cremaster serves to fix the pupa to the surrounding silk. Chapman (1893) has drawn attention to the fact that among the more highly organised moths the pupa remains in the cocoon, the emergence being entirely left to the imago, while the pupae of the more primitive moths work their way partly out of the cocoon before the final moult begins. In the latter case, the cremaster is anchored by a strand of silk which allows a certain degree of emergence, and the pupa has rows of spines on its abdominal segments, of which a greater number retain the power of mutual motion than in those pupae which do not come out of their cocoons.



While the pupa on the whole resembles the imago that is to emerge from it, there are not a few cases in which a special structure necessary for some contingency in pupal life is retained or adopted in this stage. A butterfly pupa, like the imago, has no mandibles, but in the case of the Caddis-flies (Trichoptera) and two families of small moths, the most primitive of all Lepidoptera, the pupa, like the larva, has well-developed mandibles. These enable the caddis pupa to bite its way out of the shortened larval case in which it has pupated, and then to swim upwards through the water ready for the caddis-fly's emergence into the air. Pupae that are submerged require special breathing-organs. In the previous chapter (p. 77) mention was made of the gnat's aquatic larva with its tail-spiracles adapted for procuring atmospheric air through the surface-film. The pupa of the gnat[10] also has 'respiratory trumpets' serving the same purpose, but these are a pair of processes on the prothorax, so that the pupa, which is fairly active, hangs from the surface-film with its abdomen pointing downwards through the water. This change of position is correlated with the necessity for the imago to emerge into the air; were the pupa to hang head downwards as the larva does, the gnat would perforce have to dive into the water. With the beautifully adapted transfer of the functional spiracles, their position is appropriately arranged for the gnat's emergence at the surface, and the empty pupal cuticle floats serving the insect as a raft. On this it rests securely and the crumpled wings have opportunity to expand and harden before the insect takes to flight.

[10] See Frontispiece, B.

The aquatic pupae of other Diptera, many species of the midges Chironomus and Simulium for example, breathe dissolved air by means of tufts of thread-like gills, which arise on either side of the prothorax. The pupae of Simulium rest in their curious little cup-like dwellings, attached to submerged stones or plants. The Chironomus pupa is usually found in an elongate gelatinous case adhering to a stone. From this case the pupa rises to the surface of the water, that the midge may emerge into the air. Miall and Hammond (1900) describe the arrangement by which, when the pupal stage ends, and these gills are no longer required, their connection with the air-tube system is severed 'without undue violence.' The walls of the fine air-tubes that pass into the gills are specially strengthened, but just below the pupal cuticle these walls are exceedingly thin and delicate. Thus when the pupal cuticle is cast, they are readily broken there, and the cuticle of the midge forming beneath has a spiracular opening into the main air-trunk, ready for use during the insect's aerial life.

Among those Diptera whose larva is the headless maggot a most remarkable arrangement for protecting the pupa is to be found. The last larval cuticle, instead of being as usual worked off and cast, after separation from the underlying structures, becomes hard and firm, forming a protective case (puparium) within which by the processes of histolysis and histogenesis already described the organs of the pupa and imago are built up. This puparium (fig. 22 d) is usually dark in colour, often brown and barrel-shaped, and a subcircular lid splits off from it at the head-end to allow the emergence of the fly[11]. While the maggot breathes by its tail-spiracles, the functional spiracles of the puparium (connected with the tracheal system of the enclosed pupa) are far forward, and these may be situated at the tips of long sometimes branching processes, which recall the thoracic gills of the aquatic pupae mentioned a few pages above. Adaptations, various and beautiful, to special modes of life, are thus seen to characterise pupae as well as larvae.

[11] The presence of this sub-circular lid characterises Brauer's suborder Cyclorrhapha. Those Diptera in which the pupal cuticle splits in the normal, longitudinal manner are included in the Orthorrhapha (see p. 67).



CHAPTER VIII

THE LIFE-STORY AND THE SEASONS

A number of interesting questions are associated with the seasonal cycle of an insect's life-history. In a previous chapter (IV. pp. 30, 34) reference has been made to the contrast between the long aquatic life of the larval dragon-fly or may-fly, extending over several years, and the short aerial existence of the winged adult restricted in the case of the may-flies to a few hours. Here we see that the feeding activities of the insect are carried on during the larval stage only; the may-fly in its winged condition takes no food, pairing and egg-laying form the whole of its appointed task. A similar though less extreme shortening of the imaginal life may be noticed in many endopterygote insects. For example, the bot- and warble-flies have the jaws so far reduced that they are unable to feed, and the parasitic life of the maggot (see p. 74) extending over eight or nine months in the body of the horse or ox, prepares for a winged existence of probably but a few days. Again in many moths the jaws are reduced or vestigial so that no food can be taken in the winged state, as for example in the 'Eggars' (Lasiocampidae) and the 'Tussocks' (Lymantriidae). It is noteworthy that in these short-lived insects the male is often provided with elaborate sense-organs which, we may believe, assist him to find a mate with as little delay as possible; the male may-fly has especially complex eyes, while the feelers of the male silk-moth or eggar are comb-like or feathery, the branches bearing thousands of sensory hairs. A box with a captive living female of one of these moths, if taken into a wood haunted by the species becomes rapidly surrounded by a swarm of would-be suitors, attracted by the odour emitted from the prisoner's scent-glands.

Very exceptionally the imaginal stage may be omitted from the life-story altogether. Nearly fifty years ago N. Wagner (1865) made the remarkable discovery that in the larvae of certain gall-midges (Cecidomyidae) the ovaries might become precociously mature and unfertilised eggs might be developed into small larvae observable within the body of the mother-larva; ultimately these abnormally reared young break their way out. In this case therefore there may be a series of larval generations, neither pupa nor imago being formed. Extended observations on the precocious reproductive processes of these midges have lately been published by W. Kahle (1908). A less extreme instance of an abbreviated life-story was made known by O. Grimm (1870) who saw pupae of Harlequin-midges (Chironomus) lay unfertilised eggs, which developed into larvae. Here the imaginal stage only is omitted from the life-history. Not always however is it the imaginal stage of the life-history which is shortened. Reference (p. 18) has already been made to the case of the virgin female aphids, whose eggs develop within the mother's body, so that active, formed young are brought forth. Among the Diptera it is not unusual to find similar cases, the female fly giving birth to young maggots instead of laying eggs. Such is the habit of the great flesh-fly (Sarcophaga), of some allied genera (Tachina, etc.) whose larvae live as parasites on other insects, and occasionally of the Sheep Bot-fly (Oestrus). In such cases we recognise the beginning of a shortened larval period, and Brace's investigations in 1895, summarised by E.E. Austen (1911), have shown that females of the dreaded African Tsetse flies (Glossinia) bring forth nearly mature larvae, which pupate soon after birth. In another group of Diptera, the blood-sucking parasites of the Hippoboscidae and allied families, the whole larval development is passed through within the mother's body, and a full-grown larva is born the cuticle of which hardens and darkens immediately to form a puparium; hence these flies are often called, though incorrectly, Pupipara. Still more astonishing is the mode of reproduction in the allied family of the Termitoxeniidae, curious, degraded, wingless 'guests' of the termites, or 'white ants,' lately made known through the researches of E. Wasmann (1901). Here the individual is hermaphrodite—a most exceptional condition among insects—and lays a large egg, whence is usually hatched a fully-developed adult! Here then we find that all the early stages, usual in the higher insects, are omitted from the life-story.

Interesting comparison may be made between the total duration of various insect life-stories. To some extent at least, the length of an insect's life is correlated with its size, its food, the season of the year when it breeds. Small insects have, as a rule, shorter lives than large ones; those whose larvae devour highly nutritive food generally develop more quickly than those which have to live on dry, poor, substances; life-cycles follow one another most rapidly in summer weather when temperature is high and food plentiful.

In early chapters we have already noticed the long aquatic life of the larva and nymph of a dragon-fly, relatively a large insect, and the rapid multiplication of the repeated summer broods of virgin aphids (p. 18). Within the one order of the Coleoptera it is instructive to compare the small jumping leaf-beetles, the 'turnip-flies' of the farmer, whose larvae mine in the green tissues, and complete their transformations so rapidly that several successive broods appear in the spring and early summer, with the larger click-beetles whose larvae, the equally notorious 'wireworms,' feed on roots for three or four years before they become fully grown. Among the Diptera, the 'leather-jacket' grub of the crane-fly, feeding like the wireworm on roots, has a larval life extending through the greater part of a year, while the maggot of the bluebottle, feeding on a rich meat diet, becomes mature in a few days. As examples of excessively long life-cycles the 'thirteen-year' and 'seventeen-year' cicads of North America, described by C.L. Marlatt (1895), are noteworthy. Certain specially populous 'broods' of these insects are known and localised, so that the appearance of the imagos in future years can be accurately predicted. Here again we have to do with bulky insects whose subterranean larvae and nymphs feed on comparatively innutritious roots.

In our own climate, it is of interest to notice the variation among insects as to the stage which carries the race over the winter. The click-beetles, mentioned just above, emerge from their buried pupae in summer, hibernate under stones or clods, and lay eggs among the herbage next spring. At the same time of course, owing to the extended term of the larval life, many more individuals of the species are wintering underground as 'wireworms' of various ages, and these, except in very severe frosts, can continue their occupation of feeding on roots. But in the case of the 'turnip-flies' the food-supply is cut off in winter, and all those beetles of the latest summer brood that survive hibernate in some sheltered spot, waiting for the return of spring, that they may lay their eggs, and start the life-cycle once again. Among the Diptera, most species pass the winter as pupae, the sheltering puparium being a good protection against most adverse conditions, or as flies. But where there is a prolonged parasitic larval life, as with the bot- and warble-flies, the maggot, warm and well-fed within the body of its mammalian host, affords an appropriate wintering stage.

Among the Hymenoptera an especially interesting seasonal life-cycle is afforded by the alternation of summer and winter generations in many Gall-flies (Cynipidae) as H. Adler (1881, 1896) demonstrated for most of our common species. The well-known 'oak-apples' are tenanted in summer by grubs, which after pupation develop into winged males and wingless females. The latter, after pairing, burrow underground and lay their eggs in the roots, the larvae causing the presence there of globular swellings or root-galls within which they live, pass through their transformations and develop into wingless virgin females. These shelter until February or March in their underground chambers, then climb up the tree and lay on the shoots eggs, from which will be hatched the grubs destined to grow within the oak-apples into the summer sexual brood of flies.

The Lepidoptera afford examples of hibernation in all stages of the life-history. In this order a few large moths with wood-boring caterpillars, the 'Goat' (Cossus) for example, undergo a development extending over several years, while at the other extreme a few small species may have three or more complete cycles within the twelve months. But in the vast majority of Lepidoptera we find either one or two generations, definitely seasonal, within the year; the insect is either 'single-brooded' or 'double-brooded.'

Almost every winter one or more letters may be read in some newspaper recording the writer's surprise at seeing on a sunny day during the cold season, one of our common gaily-coloured butterflies of the Vanessa group, a 'Tortoiseshell' or 'Red Admiral,' flitting about. Surprise might be greater did the observers realise that the imaginal is the normal hibernating stage for these species. Emerging from the pupa in late summer or autumn, they shelter during winter in hollow trees, under thatched eaves, in outbuildings or in similar situations, coming out in spring to lay their eggs on the leaves of their caterpillars' food-plants. The larvae feed and grow through the early summer months, in the case of the Small Tortoiseshell (Vanessa urticae) pupating before midsummer and developing into a July brood of butterflies whose offspring after a late summer life-cycle, hibernate; while for the larger species of the group there is, in our islands, only one complete life-cycle in the year, though the same insects in warmer countries may be double-brooded. C.G. Barrett records (1893, vol. I. pp. 153-4) how in the August of 1879 hundreds and thousands of 'Painted Ladies' (Pyrameis cardui) migrated into the south of England from the European continent where in many places great swarms had been observed early in the summer. 'These August butterflies, the progeny of the June swarms, coming from a warmer climate, had no intention of hibernating, but paired and laid eggs. Some of the larvae were collected and reared indoors [butterflies] emerging in November and December, but out of doors all must have been destroyed by damp or frost, in either the larva or pupa state, for no freshly emerged specimens were noticed in the spring, and no trace of the great migration remained.'

In September and October the pedestrian, even in a suburban square, may see moths with pretty brown, white-spotted wings flying around trees. These are males of the common 'Vapourer' (Orgyia antiqua), in search of the females which, wingless and helpless, rest on the cocoons surrounding the pupae whence they have just emerged, the cocoons being attached to the branches of the trees where the caterpillars have fed. After pairing, the female lays her eggs among the silk of the cocoon, partly covering them with hairs shed from her body, and then dies. The eggs thus protected remain through the winter, the larvae not being hatched till springtide, when the young leaves begin to sprout forth. The caterpillars, adorned and probably protected by their 'tussocks' of black or coloured bristles, feed vigorously. Their activity and habit of occasional migration from one tree to another, compensates, to some extent, as Miall (1908) has pointed out, for the females' enforced passivity; only in the larval state can moths with such wingless females extend their range. The caterpillars spin their cocoons towards the end of summer, and then pupate, the moths emerging in the autumn and the eggs, as we have seen, furnishing the winter stage.

After midsummer, the conspicuous cream, black and yellow-spotted 'Magpie' moth (Abraxas grossulariata) is common in gardens. The female lays her eggs on a variety of shrubby plants; gooseberry and currant bushes are often chosen. From the eggs caterpillars are hatched in autumn, but these, instead of beginning to feed, seek almost at once for rolled-up leaves, cracks in walls, crannies of bark, or similar places, which may afford winter shelters. Here they remain until the spring, when they come out to feed on the young foliage and grow rapidly into the conspicuous cream, yellow and black 'looper' caterpillars mentioned in a previous chapter (p. 60). These, when fully-grown, spin among the twigs of the food-plant a light cocoon, in which the black and yellow-banded wasp-like pupa spends its short summer term before the emergence of the moth.

An equally familiar garden insect, the common 'Tiger' moth (Arctia caia) with its 'woolly bear' caterpillar, affords a life-cycle slightly differing from that of the 'Magpie.' The gaudy winged insects are seen in July and August, and lay their eggs on a great variety of plants. The larvae hatched from these eggs begin to feed at once, and having moulted once or twice and attained about half their full size, they rest through the winter, the dense hairy covering wherewith they are provided forming an effective protection against the cold. At the approach of spring they begin to feed again, and the fully-grown 'woolly bear' is a common object on garden paths in May and June. Before midsummer it has usually spun its yellow cocoon under some shelter on the ground and changed into a pupa.

Another modification with respect to seasonal change is shown by the Turnip moth (Agrotis segetum) and other allied Noctuidae (Owl-moths). These are insects with brown-coloured wings, flying after dark in June. The dull greyish larvae feed on many kinds of low-growing plants, usually hiding in the earth by day and wandering along the surface of the ground by night, biting off the farmer's ripening corn, or burrowing into his turnips or potatoes. On account of the burrowing habits of this insect it can feed throughout the winter, except when a hard frost puts a temporary stop to its activity. By April it has become fully grown and pupates in an earthen chamber a few inches below the surface. The Turnip moth in our countries is partially double-brooded, a minority of the autumn caterpillars growing more rapidly than their comrades so that they pupate, and a second brood of moths appear in September. These pair and lay eggs, the resulting caterpillars going as Barrett suggests (1896, vol. III. p. 291) 'to reinforce the great army of wintering larvae.'

Such underground caterpillars, to a great extent protected from cold, can continue to feed through the winter. With other species we find that the larva becomes fully grown in autumn, yet lives through the winter without further change. This is the case with the Codling moth (Carpocapsa pomonella), a well-known orchard pest, which in our countries is usually single-brooded. The moth is flying in May and lays her eggs on the shoots or leaves of apple-trees, more rarely on the fruitlets, into which however the caterpillar always bores by the upper (calyx) end. Here it feeds, growing with the growth of the fruit, feeding on the tissue around the cores, ultimately eating its way out through a lateral hole, and crawling upwards if its apple-habitation has fallen, downwards if it still remains on the bough, to shelter under a loose piece of bark where it spins its cocoon about midsummer and hibernates still in the larval condition. Not until spring is the pupal form assumed, and then it quickly passes into the imaginal state. In the south of England, as F.V. Theobald (1909) has lately shown, and also in southwestern Ireland, this species may be double-brooded, the usual condition on the European continent and in the United States of America. There the midsummer larvae pupate at once and the moths of an August brood lay eggs on the hanging or stored fruit; in this case, again, however, the full-grown larva, quickly fed-up within the developed apples, is the wintering stage.

Several of the insects mentioned in this survey, like the last-named codling moth, are occasionally double-brooded. As an example of the many Lepidoptera, which in our islands have normally two complete life-cycles in the year, we may take the very familiar White butterflies (Pieris) of which three species are common everywhere. The appearance of the first brood of these butterflies on the wing in late April or May is hailed as a sign of advanced spring-time. They pair and lay their eggs on cabbages and other plants, and the green hairy caterpillars feed in June and July, after which the spotted pupae may be found on fences and walls, attached by the silken tail-pad and supported by the waist-girdle. In August and September butterflies of the second brood have emerged from these and are on the wing; their offspring are the autumn caterpillars which feed in some seasons as late as November, doing often serious damage to the late cruciferous crops before they pupate. The pupae may be seen during the winter months, waiting for the spring sunshine to call out the butterflies whose structures are being formed beneath the hard cuticle.

Reviewing the small selection of life-stories of various Lepidoptera just sketched, we notice an interesting and suggestive variety in the wintering stage. The vanessid butterflies hibernate as imagos; the 'vapourer' winters in the egg, the magpie as a young ungrown larva, the 'tiger' as a half-size larva; the Agrotis caterpillar feeds through the winter, growing all the time; the codling caterpillar completes its growth in the autumn, and winters as a full-size resting larva; lastly, the 'whites' hibernate in the pupal state. And in every case it is noteworthy that the form or habit of the wintering stage is well adapted for enduring cold.

Our native 'whites' afford illustration of another interesting feature often to be noticed in the life-story of double-brooded Lepidoptera. The butterflies of the spring brood differ slightly but constantly from their summer offspring, affording examples of what is called seasonal dimorphism. All three species have whitish wings marked with black spots, larger and more numerous in the female than in the male. In the spring butterflies these spots tend towards reduction or replacement by grey, while in the summer insects they are more strongly defined, and the ground colour of the wings varies towards yellowish. In the 'Green-veined' white (Pieris napi) the characteristic greenish-grey lines of scaling beneath the wings along the nervures, are much broader and more strongly marked in the spring than in the summer generation, whose members are distinguished by systematic entomologists under the varietal name napaeae. The two forms of this insect were discussed by A. Weismann in his classical work on the Seasonal Dimorphism of butterflies (1876). He tried the effect of artificially induced cold conditions on the summer pupae of Pieris napi, and by keeping a batch for three months at the temperature of freezing water, he succeeded in completely changing every individual of the summer generation into the winter form. The reverse of this experiment also was attempted by Weismann. He took a female of bryoniae, an alpine and arctic variety of Pieris napi, showing in an intensive degree the characters of the spring brood. This female laid eggs the caterpillars from which fed and pupated. The pupae although kept through the summer in a hothouse all produced typical bryoniae, and none of these with one exception appeared until the next year, for in the alpine and arctic regions this species is only single-brooded. Weismann experimented also with a small vanessid butterfly, Araschnia levana, common on the European continent, though unknown in our islands, which is double (or at times treble) brooded, its spring form (levana) alternating with a larger and more brightly coloured summer form (prorsa). Here again by refrigerating the summer pupae, butterflies were reared most of which approached the winter pattern, but it was impossible by heating the winter pupae to change levana into prorsa. Experiments with North American dimorphic species have given similar results. Weismann argued from these experiments that the winter form of these seasonally dimorphic species is in all cases the older, and that the butterflies developing within the summer pupae can be made to revert to the ancestral condition by repeating the low-temperature stimulus which always prevailed during the geologically recent Ice Age. On the other hand, a high temperature stimulus applied to one generation of the winter pupae cannot induce the change into the summer pattern, which has been evolved still more recently by slow stages, as the continental climate has become more genial. In tropical countries where instead of an alternation of winter and summer, alternate dry and rainy seasons prevail, somewhat similar seasonal dimorphism has been observed among many butterflies. Not a few forms of Precis, an African and Indian genus allied to our Vanessa, that had long been considered distinct species are now known, thanks to the researches of G.A.K. Marshall (1898), to be alternating seasonal forms of the same insect. The offspring when adult does not closely resemble the parent; its appearance is modified by the climatic environment of the pupa. The experiments of Weismann just sketched in outline show at least that the same principle holds for our northern butterflies.

We are thus led to see from the life-story of such insects, that the course of the story is not rigidly fixed; the creature in its various stages is plastic, open to influence from its surroundings, capable of marked change in the course of generations. And so the seasonal changes in the history of the individual from egg to imago point us to changes in the age-long history of the race.



CHAPTER IX

PAST AND PRESENT; THE MEANING OF THE STORY

In the previous chapter we recognised how the seasonal changes in various species of butterflies as observable in two or three generations, indicate changes in the history of the race as it might be traced through innumerable generations. The endless variety in the form and habits of insect-larvae and their adaptations to various modes of life, which have been briefly sketched in this little book, suggest vaster changes in the class of insects, as a whole, through the long periods of geological time. Every student of life, influenced by the teaching of Charles Darwin (1859) and his successors, now regards all groups of animals from the evolutionary standpoint, and believes that comparisons of facts of structure and life-history of orders and classes evidently akin to each other, furnish at least some indications of the course of development in the greater systematic divisions, even as the facts of seasonal dimorphism, mentioned in the last chapter, give hints as to the course of development in those restricted groups that we call species or varieties. A brief discussion of the main outlines of the life-story of insects in the wide, evolutionary sense may thus fitly conclude this book.

In the first place we turn to the 'records' of those rocks, in whose stratified layers[12] are entombed remains, often fragmentary and obscure, of the insects of past ages of the earth's history. Compared with the thousands of extinct types of hard-shelled marine animals, such as the Mollusca, fossil insects are few, as could only be expected, seeing that insects are terrestrial and aerial creatures with slight chance of preservation in sediments formed under water. Yet a number of insect remains are now known to naturalists, who are, in this connection, more particularly indebted to the researches of S.H. Scudder (1885), C. Brongniart (1894), and A. Handlirsch (1906).

[12] See Table of Geological Systems, p. 123.

We are now considering insects from the standpoint of their life-histories, and the individual life-story of an insect of which we possess but a few fragments of wings or body, entombed in a rock formed possibly before the period of the Coal Measures, can only be a matter of inference. Still it may safely be inferred that when the structure of these remains clearly indicates affinity to some existing order or family, the life-history of the extinct creature must have resembled, on the whole, that of its nearest living allies. And all the fossil insects known can be either referred to existing orders, or shown to indicate definite relationship to some existing group.

Passing over some doubtful remains of Silurian age, we find in rocks usually regarded as Devonian[13] the most ancient fossils that can be certainly referred to the insects, while from beds of the succeeding Carboniferous period, a number of insect remains have been disinterred. These Palaeozoic insects were frequently of large size, and they show distinct affinities with our recent may-flies, dragon-flies, stone-flies, and cockroaches. In the Permian period, the latest of the divisions of the Palaeozoic, lived Eugereon, an insect with hemipteroid jaws and orthopteroid wings. All these insects must have been exopterygote in their life-history, if we may trust the indications of affinity furnished by their structure. In the Mesozoic period, however, insects with complete transformations must have been fairly abundant. Rocks of Triassic age have yielded beetles and lacewing-flies, while from among Jurassic fossils specimens have been described as representing most of our existing orders, including Lepidoptera, Hymenoptera and Diptera. In Cainozoic rocks fossil insects of nearly six thousand species have been found, which are easily referable to existing families and often to existing genera. We may conclude then, imperfect though our knowledge of extinct insects is, that some of the most complex of insect life-stories were being worked out before the dawn of the Cainozoic era. Some instructive hints as to differences in the rate of change among different insect groups may be drawn from the study of parasites. For example, V.L. Kellogg (1913) points out that an identical species of the Mallophaga (Bird-lice) infests an Australian Cassowary and two of the South American Rheas; while two species of the same genus (Lipeurus) are common to the African Ostrich and a third kind of South American Rhea. These parasites must have been inherited unchanged by the various members of these three families of flightless birds from their common ancestors, that is from early Cainozoic times at latest. On the other hand, the various kinds of such highly specialised parasites as the warble-flies of the oxen and deer, must have become differentiated during those later stages of the Cainozoic period which witnessed the evolution of their respective mammalian hosts.

[13] The 'Little River' beds of St John, New Brunswick, Canada, by some modern geologists however considered as Carboniferous.

The foregoing brief outline of our knowledge of the geological succession of insects shows that the exopterygote preceded, in time, the endopterygote type of life-history. We have already seen that those insects undergoing little change in the life-cycle, and with visible, external wing-rudiments, are on the whole less specialised in structure than those which pass through a complete transformation. These two considerations, taken together, suggest strongly that in the evolution of the insect class, the simpler life-history preceded the more complex. Such a conclusion seems reasonable and what might have been expected, but we are confronted with the difficulty that if the most highly organised insects pass through the most profound transformations, then insects present a remarkable and puzzling exception to the general rules of development among animals, as has already been pointed out in the first chapter of this volume (p. 7). A few students of insect transformation have indeed supposed that the crawling caterpillar or maggot must be regarded as a larval stage which recalls the worm-like nature of the supposed far-off ancestors of insects generally. Even in Poulton's classical memoir (1891, p. 190), this view finds some support, and it may be hard to give up the seductive idea that the worm-like insect-larva has some phylogenetic meaning. But the weight of evidence, when we take a comprehensive survey of the life-story of insects, must be pronounced to be strongly in favour of the view put forward by Brauer (1869), and since supported by the great majority of naturalists who have discussed the subject, that the caterpillar or the maggot is itself a specialised product of the evolutionary process, adapted to its own particular mode of larval life.

The explanation of insect transformation is, in brief, to be found in an increasing amount of divergence between larva and imago. The most profound metamorphosis is but a special type of growth, accompanied by successive castings and renewings of the chitinous cuticle, which envelopes all arthropods. In the simplest type of insect life-story, there is no marked difference in form between the newly-hatched young and the adult, and in such cases we find that the young insect lives in the same way as the adult, has the same surroundings, eats the same food. This is the rule (see Chapters II and III) with the Apterygota, the Orthoptera, and most of the Hemiptera. In the last-named order, however, we find in certain families marked divergence between larva and imago, for example in the cicads, whose larvae live underground, while in the coccids, whose males are highly specialised and females degraded, there succeeds to the larva—very like the young stage in allied families—a resting instar, which in the case of the male, suggests comparison with the pupa of a moth or beetle.

Turning to the stone-flies, dragon-flies and may-flies, whose life-stories have been sketched in Chapter IV, we find that the early stages are passed in water, whence before the final moult, the insects emerge to the upper air. Except for the possession of tufted gills, adapting them to an aquatic life, the stone-fly nymphs differ but slightly from the adults; the grubs of the dragon-flies and may-flies, however, are markedly different from their parents. In connection with these comparisons, it is to be noted that the dragon-flies and may-flies are more highly specialised insects than stone-flies, divergent specialisation of the adult and larva is therefore well illustrated in these groups, which nevertheless have, like the Hemiptera and Orthoptera, visible external wing-rudiments.

From the vast array of insects that show internal wing-growth and a true pupal stage, a few larval types were chosen for description in Chapter VI, and a review of these suggests again the thought of increasing divergence between larva and imago. Reference has been made previously to the many instances in which the former has become pre-eminently the feeding, and the latter the breeding stage in the life-cycle. It seems impossible to avoid the conclusion that the active, armoured campodeiform grub differing less from its parent than an eruciform larva differs from its parent, is as a larval type more primitive than the caterpillar or maggot. A. Lameere has indeed, while admitting the adaptive character of insect larvae generally, argued (1899) with much ingenuity that the eruciform or vermiform type must have been primitive among the Endopterygota, believing that the original environment of the larvae of the ancestral stock of all these insects must have been the interior of plant tissues. He is thus forced to the necessity of suggesting that the campodeiform larvae of ground-beetles or lacewings must be regarded as due to secondarily acquired adaptations; 'they resemble Thysanura and the larvae of Heterometabola only as whales resemble fishes.' There are two considerations which render these theories untenable. The Neuroptera and Coleoptera among which campodeiform larvae are common, are less specialised than Lepidoptera, Hymenoptera, and Diptera, in which they are unknown. And among the Coleoptera which as we have seen (pp. 50 f.) display a most interesting variety of larval structure, the legless, eruciform larva characterises families in which the imago shows the greatest specialisation, while in the same life-story, as in the case of the oil-beetles (pp. 56-7), the newly-hatched grub may be campodeiform, changing to the eruciform type as soon as it finds itself within reach of its host's rich store of food.

A certain amount of difficulty may be felt with regard to the theory of divergent evolution between imago and larva, in the case of those insects with complete transformation whose grubs and adults live in much the same conditions. By turning over stones the naturalist may find ground-beetles in company with the larvae of their own species. On the leaves of a willow tree he may observe leaf-beetles (Phyllodecta and Galerucella) together with their grubs, all greedily eating the foliage; or lady-bird beetles (Coccinella) and their larvae hunting and devouring the 'greenfly.' All of these insects are, however, Coleoptera, and the adult insects of this order are much more disposed to walk and crawl and less disposed to fly than other endopterygote insects. Their heavily armoured bodies and their firm shield-like forewings render them less aerial than other insects; in many genera the power of flight has been altogether lost. It is not surprising, therefore, that many beetles, even when adult, should live as their larvae do; since the acquirement of complete metamorphosis they have become modified towards the larval condition, and an extreme case of such modification is afforded by the wingless grub-like female Glow-worm (Lampyris).

With most insects, however, the larva must be regarded as the more specially modified, even if degraded, stage. Miall (1895) has pointed out that the insect grub is not a precociously hatched embryo, like the larvae of multitudes of marine animals, but that it exhibits in a modified form the essential characters of the adult. Comparison for example can be readily made between the parts of the caterpillar and the butterfly, whose story was sketched in the first chapter of this book, widely different though caterpillar and butterfly may appear at a superficial glance. And the survey of variety in form, food, and habit of insect larvae given in Chapter VI enforces surely the conclusion that the larva is eminently plastic, adaptable, capable of changing so as to suit the most diverse surroundings. In a most suggestive recent discussion on the transformation of insects P. Deegener (1909) has claimed that the larva must be regarded as the more modified stage, because while all the adult's structures are represented in the larva, even if only as imaginal buds, there are commonly present in the larva special adaptive organs not found in the imago, for example the pro-legs of caterpillars or the skin-gills of midge-grubs. The correspondence of parts in butterfly and caterpillar just referred to, may still be traced, though less easily, in bluebottle and maggot. The latter is an extreme example of degenerative evolution, and its contrast with the elaborately organised two-winged fly marks the greatest divergence observable between the larva and imago. With this divergence the resting pupal stage, during which more or less dissolution and reconstruction of organs goes on, becomes a necessity, and it has already been pointed out how the amount of this reconstruction is greatest where the divergence between the larval and perfect stages is most marked. Whatever differences of opinion may prevail on points of detail, the general explanation of insect metamorphosis as the result of divergent evolution in the two active stages of the life-story must assuredly be accepted. No other explanation accords with the increasing degree of divergence to be observed as we pass from the lower to the higher insect orders.

The successive incidents of the life-story of most insects are largely connected with the acquisition of wings. Wings, and the power of flight wherewith they endow their possessors, are evidently beneficial to the race in giving power of extending the range during the breeding period and thus ensuring a wide distribution of the eggs. In no case are wings fully developed until the closing stage of the insect's life, they are always acquired after hatching or birth. We have already noticed (p. 40) how Sharp (1899) has laid stress on the essential difference between the exopterygote and endopterygote insects, the wing-rudiments of the former growing outwards throughout life while those of the latter remain hidden until the pupal instar. Sharp considers that there is some difficulty in bridging, in thought, the gap between these two methods of wing-growth, and has put forward an ingenious suggestion to meet it (1902). Reference has already been made to insects of various orders in which one sex is wingless, the Vapourer Moth (p. 96) for example, or all the individuals of both sexes are wingless, as the aberrant cockroaches mentioned in Chapter II (p. 15), or certain generations of virgin females are wingless, for example aphids (pp. 18-19) and gall-flies (pp. 94-5). Insects may thus become secondarily wingless, that is to say be manifestly the offspring of winged parents, and such wingless forms may on the other hand give rise to offspring or descendants with well-developed wings. Frequently, as in the case of the aphids, many wingless generations intervene between two winged generations. A striking illustration of this fact is afforded by an aquatic bug, Velia currens, commonly to be seen skating over the surface of running water. The adults of Velia are nearly always wingless, but now and then the naturalist meets with a specimen provided with functional wings, the possession of which enables the insect to make its way to a fresh stream. Moreover there are whole orders of parasitic insects, such as the lice and fleas, which, showing clear affinity to orders of winged insects, are believed to be secondarily wingless. These orders are designated by Sharp 'Anapterygota.' And from the analogy of the periodic loss and recovery of wings in various generations of the same species, he has concluded that the gap between the exopterygote and the endopterygote method of development may have been bridged by an anapterygote condition; that the ancestors of those insects with complete transformations were the wingless descendants of primitive insects which grew their wings from visible external rudiments, and that in later times re-acquiring wings, they developed these organs in a new way, from inwardly directed rudiments or imaginal buds.

This theory of Sharp's is original, daring, and ingenious, but the loss and re-acquisition of wings which it presupposes is difficult to imagine in large groups during a prolonged evolutionary history, while the sudden appearance of a totally new mode of wing-growth in the offspring of wingless insects would be an extreme example of discontinuity in development.

On the whole the most probable suggestion which can be made as to the origin of 'complete' transformation in insects is that the instar in which wings were first visible externally became later and later in the course of the evolution of the more highly organised groups. In this way a gradual transition from the exopterygote to the endopterygote type of life-story is at least conceivable. It will be remembered that a may-fly (p. 33) undergoes a moult after acquiring functional wings, emerging into the air as a 'sub-imago.' In not a few endopterygote insects, the pupa shows more or less activity, swimming through water intermittently (gnats) or just before the imago has to emerge (caddis-flies); working its way out of the ground (crane-flies) or coming half-way out of its cocoon (many moths). The pupa of the higher insects almost certainly corresponds with the may-fly's sub-imago, and the facts just recalled as to remnants of pupal activity suggest that in the ancestors of endopterygote insects what is now the pupal instar was represented by an active nymphal or sub-imaginal stage, possibly indeed by more than one stage, as Packard and other writers have stated that pupae of bees and wasps undergo two or three moults before the final exposure of the imago. Such an early pupal instar has been defined as a 'pro-nymph' or a 'semi-pupa.' Examples have been given of the exceptional passive condition of the penultimate instar in Exopterygota. The instars preceding this presumably had originally outward wing-rudiments in all insect life-histories, and the endopterygote condition was attained by the postponement of the outward appearance of these to successively later stages. The leg and wing rudiments of the male coccid (pp. 20-1) beneath the cuticle of the second instar are strictly comparable to imaginal buds, and these are present in one instar of what is generally regarded as an exopterygote life-history. The first instar in all insects has no visible wing-rudiments, but when they grow outwardly from the body, they necessarily become covered with cuticle, so that they must be visible after the first moult. There is no supreme difficulty in supposing that the important change was for these early rudiments to become sunk into the body, so that the cuticle of the second, and, later, of the third and succeeding instars, showed no outward sign of their presence. This suggestion is confirmed by Heymons' (1896, 1907) observation of the occasional appearance of outward wing-rudiments on the thoracic segments of a mealworm, the larva of the beetle Tenebrio molitor, and by F. Silvestri's discovery (1905) of a 'pro-nymph' stage with short external wing-rudiments between the second larval and the pupal instars of the small ground-beetle Lebia scapularis. Whatever may be the exact explanation of these abnormalities, they show that in the life-story of the higher insects outward wing-rudiments may even yet appear before the pupal stage, confirming our belief that such appearance is an ancestral character. The inward growth of these wing-rudiments may well have been correlated with a difference in form between the newly-hatched insect and its parent. As this difference persisted until a constantly later stage, and the pre-imaginal instar became necessarily a stage for reconstruction, the present condition of complete metamorphosis in the more highly organised orders was finally attained.

To explain satisfactorily these complex life-stories is however admittedly a difficult task. The acquisition of wings is, as we have seen, a dominating feature in them all, but if we try to go yet a step farther back and speculate on the origin of wings in the most primitive exopterygote insects, the task becomes still more difficult. Many years ago Gegenbaur (1878) was struck by the correspondence of insect wings to the tracheal gills of may-fly larvae, which are carried on the abdominal segments somewhat as wings are on the thoracic segments. But Boerner has recently (1909) brought forward evidence that these abdominal gills really correspond serially with legs. Moreover Gegenbaur's theory suggests that the ancestral insects were aquatic, whereas the presence of tubes for breathing atmospheric air in well-nigh all members of the class, and the fact that aquatic adaptations, respiratory and otherwise, in insect-larvae are secondary force the student to regard the ancestral insects as terrestrial. It is indeed highly probable that insects had a common origin with aquatic Crustacea, but all the evidence points to the ancestors of insects having become breathers of atmospheric air before they acquired wings. How the wings arose, what function their precursors performed before they became capable of supporting flight, we can hardly even guess.

Our study of the life-story of insects, therefore, while it has taught us something of what is going on around us to-day, and has given us hints of the course of a few threads of that long life-story which runs through the ages, brings us face to face with the most instructive, if humbling fact that 'there are many more things of which we are ignorant.' The passage from creeping to flight, as the caterpillar becomes transformed into the butterfly, was a mystery to those who first observed it, and many of its aspects remain mysterious still. Perhaps the most striking result of the study of insect transformation is the appreciation of the divergent specialisation of larva and imago, and it is a suggestive thought that of the two the larva has in many cases diverged the more from the typical condition. The caterpillar crawling over the leaf, or the fly-grub swimming through the water, may thus be regarded as a creature preparing for a change to the true conditions of its life. It is a strange irony that the preparation is often far longer than the brief hours of achievement. But the light which research has thrown on the nature of these wonderful life-stories, the demonstration of the unseen presence and growth within the insect, during its time of preparation among strange surroundings, of the organs required for service in the coming life amid its native air, confirm surely the intuition of the old-time students, who saw in these changes, so familiar and yet so wonderful, a parable and a prophecy of the higher nature of man.



OUTLINE CLASSIFICATION OF INSECTS

Class INSECTA or HEXAPODA.

Sub-class A, APTERYGOTA.

Order 1. Thysanura (Bristle-tails). 2. Collembola (Spring-tails).

Sub-class B, EXOPTERYGOTA.

Order 1. Dermaptera (Earwigs). 2. Orthoptera (Cockroaches, Grasshoppers, Crickets). 3. Plecoptera (Stone-flies). 4. Isoptera (Termites or 'White Ants'). 5. Corrodentia (a) Copeognatha (Book-lice). (b) Mallophaga (Biting-lice). 6. Ephemeroptera (May-flies). 7. Odonata (Dragon-flies). 8. Thysanoptera (Thrips). 9. Hemiptera (a) Heteroptera (Bugs, Pond-skaters) (b) Homoptera (Cicads, 'Greenfly,' Scales). 10. Anoplura (Lice).

Sub-class C, ENDOPTERYGOTA.

Order 1. Neuroptera (Alder-flies, Ant-lions, Lacewings). 2. Coleoptera (Beetles). 3. Mecaptera (Scorpion-flies). 4. Trichoptera (Caddis-flies). 5. Lepidoptera (Moths and Butterflies). 6. Diptera (Two-winged flies) (a) Orthorrhapha (Crane-flies, Midges, Gnats) (b) Cyclorrhapha (Hover-flies, House-flies, Bot-flies, &c). 7. Siphonaptera (Fleas). 8. Hymenoptera (a) Symphyta (Saw-flies) (b) Apocrita (Gall-flies, Ichneumon-flies, Wasps, Bees, Ants).



TABLE OF GEOLOGICAL SYSTEMS

These names, given by geologists to the various divisions of rocks, as indicated by the fossils entombed in them, are arranged in 'descending' order, the more recent formations above, the more ancient below, as newer deposits necessarily lie over older beds.

CALNOZOIC OR TERTIARY GROUP.

Pleistocene. Pliocene. Miocene. Eocene.

MESOZOIC OR SECONDARY GROUP.

Cretaceous. Jurassic. Triassic.

PALAEOZOIC OR PRIMARY GROUP.

Permian. Carboniferous. Devonian. Silurian. Cambrian.



BIBLIOGRAPHY

The following list of some books and papers, referred to in this little volume or of especial service to the author in its preparation, is needless to say very far from exhaustive. To save space, titles are often abbreviated. Most of the works in the general list (A) contain extensive lists of literature on insects and their transformations, these should be consulted by the serious student.

A. GENERAL WORKS.

1909. C. Boerner. Die Verwandlungen der Insekten. Sitzb. d. Gesellsch. naturforsch. Freunde, Berlin.

1869. F. Brauer. Betrachtung ueber die Verwandlung der Insekten. Verhandl. der K.K. zool.-bot. Gesellschaft in Wien. XIX.

1899. G.H. Carpenter. Insects, their Structure and Life. London.

1859. C. Darwin. The Origin of Species. London.

1909. P. Deegener. Die Metamorphose der Insekten. Leipzig.

1906. J.W. Folsom. Entomology. London.

1878. C. Gegenbaur. Grundriss der Vergleichende Anatomie. Leipzig.

1906. A. Handlirsch. Die fossilen Insekten. Leipzig.

1904. L.F. Henneguy. Les Insectes. Paris.

1907. R. Heymons. Die verschiedenen Formen der Insectenmetamorphose. Ergebnisse der Zoologie. I.

1899. A. Lameere. La raison d'etre des Metamorphoses chez les Insectes. Ann. Soc. Entom. Bruxelles. XLIII.

1874. J. Lubbock. The Origin and Metamorphoses of Insects. London.

1895. L.C. Miall. (a) The Transformations of Insects. Nature. LIII.

1895. —— (b) The Natural History of Aquatic Insects. London.

1908. —— Injurious and Useful Insects. 2nd edition. London.

1839. G. Newport. Insects. Todd Cyclopaedia. II. London.

1898. A.S. Packard. Text book of Entomology. New York.

1734-42. R.A.F. de Reaumur. Memoires pour servir a l'Histoire naturelle et a l'anatomie des Insectes. Paris.

1895-8. D. Sharp. The Cambridge Natural History, V, VI. London.

1899. —— Some points in the Classification of Insects. IV. Internat. Zoolog. Congress.

1902. —— Insects in Encycl. Brit. 10th Edition, XXIX. London.

1910. —— and G.H. Carpenter. Hexapoda in Encycl. Brit. 11th Edition. Cambridge.

1737. J. Swammerdam. Biblia Naturae. Leyden (incorporates works on Insects published during the author's lifetime 1669-75).

1909. F.V. Theobald. Insect Pests of Fruit. Wye.

B. SPECIAL WORKS.

1881. H. Adler. Ueber den Generationswechsel den Eichen-Gallwespen. Zeitsch. f. wissensch. Zoologie. XXXV.

1896. —— and C.R. Straton. Alternating Generations. Oxford.

1902. J. Anglas. Nouvelles Observations sur les Metamorphoses Internes. Arch. d'Anat. Microscop. IV.

1911. E.E. Austen. Handbook of the Tsetse-Flies. London (Brit. Museum).

1909. F. Balfour-Browne. Life-History of Agrionid Dragonfly. Proc. Zool. Soc. Lond.

1893, &c. C.G. Barrett. Lepidoptera of the British Islands. London.

1890. H. Beauregard. Les Insectes Vesicants. Paris.

1909. C. Boerner. Die Tracheenkiemen der Ephemeriden. Zoolog. Anz. xxxiii.

1863. F. Brauer. Monographie der Oestriden. Wien.

1894. C. Brongniart. Recherches pour servir a l'histoire des Insectes fossiles des Temps Primaires. St Etienne.

1893. T.A. Chapman. Structure of Pupae of Heterocerous Lepidoptera. Trans. Entom. Soc. Lond.

1891. H. Dewitz. Das geschlossene Tracheensystem bei Insektenlarven. Zoolog. Anz. xiii.

1857-8. J.H. Fabre. L'Hypermetamorphose et les Moeurs des Meloides. Ann. Sci. Nat. (Zool.), (4). VII. IX.

1869. M. Ganin. Die Entwicklungsgeschichte bei den Insekten. Zeitsch. f. wissensch. Zoolog. xix.

1894. J. Gonin. La Metamorphose des Lepidopteres. Bull. Soc. Vaud. Sci. Nat. xxx.

1870. O. Grimm. Die ungeschechtliche Fortpflanzung einer Chironomus. Mem. Acad. Imper. St Petersbourg (7). xv.

1890. W. Hatchett-Jackson. Morphology of the Lepidoptera. Trans. Linn. Soc. (Zool.) Lond. (2). v.

1896. R. Heymons. Fluegelbildung bei der Larve von Tenebrio molitor. Sitzb. d, Gesellsch. Naturforsch. Freunde, Berlin.

1906. —— Ueber die ersten Jugendformen von Machilis alternata. Ib.

1908. W. Kahle. Die Paedogenesis der Cecidomyiden. Zoologica. IV.

1913. V.L. Kellogg. Distribution and Species-forming of Ectoparasites. Amer. Naturalist. XLVII.

1887. A. Kowalevsky. Die nachembryonale Entwicklung der Musciden. Zeitsch. f. wissensch. Zool. XLV.

1904. O.H. Latter. Natural History of Common Animals (chaps. III, IV, V). Cambridge.

1890-95. B.T. Lowne. The Blowfly, 2 vols. London.

1863. J. Lubbock. Development of Chloeon. Trans. Linn. Soc. Lond. XXIII.

1762. P. Lyonet. Traite anatomique de la Chenille. Haag.

1669. M. Malpighi. De Bombyce. London.

1898. C.L. Marlatt. The periodical Cicada. Entom. Bull. 14, U.S. Dept. Agric.

1898. G.A.K. Marshall. Seasonal Dimorphism in Butterflies. Ann. Mag. Nat. Hist. (7). II.

1900. L.C. Miall and A.B. Hammond. The Harlequin Fly. Oxford.

1901-3. R. Newstead. Coccidae of the British Isles. London.

1877. J.A. Palmen. Zur Morphologie des Tracheensystems. Leipzig.

1891. E.B. Poulton. External Morphology of the Lepidopterous Pupa. Trans. Linn. Soc. Zool. (2). V.

1892. —— Colour-relation between Lepidopterous Larvae &c. and their surroundings. Trans. Entom. Soc. Lond.

1880. C.V. Riley. Pupation of Butterflies. Proc. Amer. Assoc. XXVIII.

1902. E.D. Sanderson. Report of Entomologist. Delaware. U.S.A.

1885. E.O. Schmidt. Metamorphose und Anatomie des maennlichen Aspidiotus. Archiv f. Naturgeschichte. LI.

1885. S.H. Scudder. Insekten in Zittel's Paleontologie. II.

1907. A.J. Siltala. Die postembryonale Entwicklung der Trichopteren-Larven. Zoolog. Jahrb. Suppl. IX.

1905. F. Silvestri. Metamorfosi e Costumi della Lebia scapularis. Redia. II.

1900. J.B. Smith. The Apple Plant-louse. New Jersey Agric. Exp. Station Bull. 143.

1888. J. Van Rees. Die innere Metamorphose von Musca. Zoolog. Jahrb. Anat. III.

1911. K.W. Verhoeff. Ueber Felsenspringer, Machiloidea. Zoolog. Anz. XXXVIII.

1865. N. Wagner. Die viviparen Gallmueckenlarven. Zeitsch. f. wissensch. Zoolog. XV.

1901. E. Wasmann. Termitoxenia. Zeitsch. f. wissensch. Zoolog. LXX.

1864. A. Weismann. Die nachembryonale Entwicklung der Musciden. Zeitsch. f. wissensch. Zoolog. XIV.

1865. —— Die Metamorphose von Corethra. Ib. XVI.

1876. —— Studien zur Descendenz-Theorie. Leipzig. (English Translation by R. Meldola, London, 1882.)



INDEX

Abraxas grossulariata, 60, 83, 97-8

Adaptation of larvae, 57, 79, 114

Adephaga, 51

Adler, H., 94

Aeschnidae, 27, 29, 31

Agrionidae, 27, 28

Agrotis segetum, 98

Air-tubes, 2, 11, 23, 47, 70, 77, 87, 120

Alternation of generations, 17, 94

Ametabola, 11, 35

Anapterygota, 116

Anglas, J., 46

Ant-lions, 57

Ants, 64, 66

Aphidae, 17-20, 116

Aphis pomi, 18-19

Aphis-lion, 57

Apterygota, 41, 110

Aquatic insects, 23-34, 76-9, 120

Araschnia levana and var. prorsa, 103

Arctia caia, 98

Arctiadae, 59

Arthropoda, 9

Austen, E.E., 91

Avebury, Lord, see Lubbock, J.

Balfour-Browne, F., 28

Bark-beetles, 55

Barrett, C.G., 96, 99

Beauregard, H., 56

Bees, 40, 46, 64, 83

Beetles, 40, 50-7, 80, 107, 112-3, 119

Bell Moths, 62

Bird-lice, 108

Birth, 18, 91

Blatta orientalis, 15

Blister-beetles, 56

Blowfly or Bluebottle, 43, 44, 46, 67, 71-3, 93, 114

Boerner, C., 32, 120

Bot-flies, 73-4, 89, 91

Brain, 44

Brauer, F., 6, 52, 56, 67, 109

Bristle-tails, 11

Brongniart, C., 106

Butterflies, 1, 83, 95-6, 114

Cabbage-butterflies, 39, 41, 85, 100-1

Cabbage-fly, 73

Caddis-flies, 62-3, 86, 117

Cainozoic insects, 107

Calliphora, 43. See also Blowfly

Campodeiform larvae, 52, 56, 111

Carabidae, 52

Carboniferous insects, 107

Carpocapsa pomonella, 99-100

Carrion-beetles, 50

Caterpillar, 4, 36, 49, 58-62, 95-101, 109, 114

Cecidomyidae, 68-70, 90

Cerambycidae, 55

Cercopods, 12, 15

Chafers, 52

Chapman, T.A., 81, 84

Chironomus, 43, 77, 87, 91

Chloeon, 33

Chrysalis, 82. See also Pupa

Chrysomelidae, 53. See also Leaf-beetles

Chrysopa, 57

Cicads, 22, 93, 110

Classification, 122

Clearwing Moths, 62

Click-beetles, 52, 93

Clothes-moths, 62

Coccidae, 20, 110, 118

Coccinella, 113

Cockroaches, 11, 14, 15, 107, 115

Cocoons, 82

Codling Moth, 62, 99

Coleoptera, 50-6, 80, 112, 119

Collembola, 11

Complete transformation, 35, 107, 119. See also Endopterygota

Corethra, 43

Cossus, 38, 62, 82, 95

Crane-flies, 67, 70, 93, 117

Cremaster, 83

Crustacea, 7, 120

Culex, 43, 77, 86

Curculionidae, 55

Cuticle, 2, 9, 29, 37, 40, 50, 81, 87, 110

Cynipidae, 94. See also Gall-flies

Daddy-long-legs, 69-70

Darwin, C., 105

Deegener, P., 6, 114

Devonian insects, 107

Dewitz, H., 28

Digestive system, 10, 45-7

Diplosis pyrivora, 70

Diptera, 42, 64, 67-79, 81, 86-8, 91, 94, 107

Divergence between larva and imago, 110, 114, 121

Double-brooded Lepidoptera, 95, 100-4

Dragon-flies, 26-31, 107, 110

Drone-flies, 76

Duration of life, 34, 89, 92-3, 95

Dyticus, 51

Ecdysis, 10. See also Moult

Ectoderm, 9, 11, 47

Eggar Moths, 59, 89

Eggs, 6, 17-18, 26, 34, 65-7, 71, 90, 94-5, 97

Elateridae, 52

Endopterygota, 41, 49, 108, 112, 115-6

Ephemeroptera, 24. See also May-flies

Epidermis, 9, 40

Eristalis, 76

Eruciform larvae, 56, 58-70, 111

Evolution, 16, 103, 105-21

Exopterygota, 41, 108, 115-6, 118

Exoskeleton, 9

Fabre, J.H., 56

Fat-body, 47

Feeding-period, 27, 32, 36, 89, 111

Feelers, 1, 4, 42, 71

Fleas, 116

Fore-gut, 47

Free pupa, 80

Gall-flies, 64-6, 94, 115

Gall-midges, 68-70, 90

Ganin, M., 66

Gastrophilus equi, 73-4

Gegenbaur, C., 120

Geological history, 106-8, 123

Geometridae, 59

Gills, 24, 27, 32, 78, 87, 114, 120

Glossinia, 91

Glow-worm, 50, 113

Gnats, 43, 77, 86

Goat Moth, 38, 62, 82, 95

Gonin, J., 38, 41

Grasshoppers, 11, 14, 15

Grimm, O., 90

Ground-beetles, 52, 112

Growth, 9

Grub, 63-70. See also Caterpillar, Larva

Hairs, 59, 82, 98

Hammond, A.R., 43, 77, 87

Handlirsch, A., 106

Harvey, William, 7

Hatchett-Jackson, W., 83

Hawk Moths, 60

Heart, 45

Helodes, 50

Hemerobius, 57

Hemimetabola, 35

Hemiptera, 17, 110

Henneguy, L.F., 45, 48

Heymons, R., 6, 11, 119

Hibernation. See Wintering stages

Hind-gut, 47

Hippoboscidae, 91

Histogenesis and Histolysis, 48

Holometabola, 35

House-fly, 67, 71, 73

Hover-flies, 74-6

Hymenoptera, 58, 64, 94, 107

Hypermetamorphosis, 56

Hypoderma bovis, 73-5

Hypodermis, 9

Ichneumon-flies, 64, 66, 82

Imaginal buds or discs, 34-48, 114, 117-8

Imago, 24, 34, 114

Instar, 13, 33, 56, 117-9

Jaws of imago and larva, 2, 4, 5, 32, 42, 89

Jurassic insects, 107

Kahle, W., 90

Kellogg, V.L., 108

Kowalevsky, A., 46

Labium, 2, 27

Lacewing-flies, 57, 107

Ladybirds, 113

Lameere, A., 111

Lampyris, 113

Larva, 4, 22, 26-7, 32, 49-79, 110-15

Larval reproduction, 90

Lasiocampidae, 59, 89

Latter, O.H., 28

Leaf-beetles, 53, 83, 92-3, 113

Lebia scapularis, 119

Lepidoptera, 1, 36, 38, 49, 58, 81, 95-104, 107

Libellulidae, 27

Lice, 116

Lipeurus, 108

Longhorn Beetles, 55

Looper caterpillars, 59, 61

Lowne, B.T., 42

Lubbock, J., 6, 32

Lymantriidae, 90

Lyonet, P., 38

Machilis, 11

Maggot, 44, 67, 71-6, 109, 114

Magpie Moth, 60, 82, 97-8

Mallophaga, 108

Mandibles, 4, 17, 26, 58, 67, 86

Mangel-fly, 73

Marlatt, C.L., 93

Marshall, G.A.K., 104

Maxillae, 2, 17, 37, 42

May-flies, 31-4, 107, 110, 117, 120

Meloidae, 56

Mesozoic insects, 107

Metabola, 35

Metamorphosis (in general), 6, 109; (degrees of in insects) 8, 35, 109, 117-19

Miall, L.C., 6, 28, 33, 43, 77, 78, 87, 97, 113

Mosquito. See Culex, Gnats

Moths, 1, 58-62, 84, 95-100, 117

Moult, 10, 32, 36, 41

Musca domestica, 71

Muscidae, 44

Muscles, 47

Nervous system, 44-5

Neuroptera, 57, 80, 112

Newport, G., 41, 44

Noctuidae, 60, 98

Nymph, 15, 28, 33

Oak-apples, 94

Obtect pupa, 81

Odonata, 24. See also Dragon-flies

Oestrus ovis, 91

Oil-beetles, 56, 112

Orgyia antiqua, 96-7

Orthoptera, 17, 35, 110

Owl Moths, 60, 98

Packard, A.S., 56, 118

Paedogenesis. See Larval reproduction

Painted Lady Butterfly, 96

Palaeozoic insects, 107

Palmen, J.A., 25

Parasitic insects, 73-4, 108, 116

Parental care, 64-6

Parthenogenesis, 18

Partial transformation, 35, 37

Perla, 24

Permian insects, 107

Phagocytes, 48

Phyllodecta, 53, 113

Phyllotreta, 53

Pieris brassicae, 39, 41, 85, 100

Pieris napi and var. bryoniae, 102-3

Platygaster, 66

Plecoptera, 24. See also Stone-flies

Pompilidae, 66-7

Poulton, E.B., 61, 82, 109

Precis, 104

Proctotrypidae, 66

Pro-legs, 4, 58-9, 84, 114

Pro-nymph, 118, 119

Protective coloration, 60-1

Psylliodes chrysocephala, 54

Ptinidae, 54

Pupa, 4, 37, 40, 79-88, 114, 117

Puparium, 88

Pupipara, 91

Pyrameis cardui, 96

Rat-tailed maggot, 76

Reaumur, R.A.F. de, 8, 28, 33, 41

Reproductive larvae, 90; pupae, 91

Reproductive organs, 45

Rhabdophaga heterobia, 70

Riley, C.V., 83

Sanderson, E.D., 17

Sand-midges, 78

Sarcophaga, 91

Saw-flies, 58-9

Scale-insects, 20. See also Coccidae

Scarabaeidae, 52

Schmidt, E.O., 21

Scolytidae, 55

Scudder, S.H., 106

Seasonal changes, 89-104

Seasonal dimorphism, 102

Semi-pupa, 118

Sesiidae, 62

Sexual differences, 15, 20-1, 90

Sharp, D., 13, 36, 40, 115

Silk-spinning, 58, 62-3, 82

Silkworms, 82

Silpha, 50

Siltala, A.J., 63

Silvestri, F., 119

Simulium, 78, 87

Smith, J.B., 17

Sphegidae, 66-7

Sphingidae, 60

Spinneret, 58

Spiracles, 2, 23, 70, 72, 77, 86, 87

Spring-tails, 11

Stone-flies, 24, 107, 110

Sub-imago, 33, 117

Sucking insects, 17

Swammerdam, J., 33

Syrphus, 74-6

Tachininae, 73, 91

Tenebrio molitor, 119

Termitoxeniidae, 92

Theobald, F.V., 100

Thysanura, 11

Tiger Moths, 59, 82, 98

Timber-beetles, 54

Tineidae, 62

Tipulidae, 70

Tortoiseshell Butterfly, 45, 95

Tortricidae, 62

Tracheal system. See Air-tubes, Spiracles

Transformation. See Metamorphosis

Triassic insects, 107

Trichocera, 70

Trichoptera, 62-3, 76, 80, 86

Tsetse Flies, 91

Turnip-fly, 53, 92, 94

Turnip Moth, 98-9

Tussock Moths, 90, 97

Vanessa urticae, 45, 95

Van Rees, J., 42

Vapourer Moth, 96-7, 115

Velia currens, 116

Verhoeff, K.W., 11

Vermiculiform larvae, 67, 71-6, 111

Virgin stem-mothers, 18

Viviparous reproduction. See Birth

Wagner, N., 90

Warble-fly, 73-4, 89, 108

Warning coloration, 60

Wasmann, E., 92

Wasps, 46, 64, 66-7, 83

Water-insects. See Aquatic insects

Weevils, 55

Weismann, A., 38, 42, 102

White Butterflies, 41, 83, 85, 100-3

Willow-beetles, 53

Wingless insects, 15, 18, 20, 96, 115

Wing-rudiments, 13, 18, 20, 22, 24, 28, 33, 36-8, 40, 111, 115, 117-19

Wings, 1, 14, 115, 119-20

Winter broods, 102-3

Wintering stages, 93-101

Wireworms, 52, 93

Wood-wasps, 65



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THE CAMBRIDGE MANUALS OF SCIENCE AND LITERATURE

Published by the Cambridge University Press

GENERAL EDITORS P. GILES, Litt.D. Master of Emmanuel College and A.C. SEWARD, M.A., F.R.S. Professor of Botany in the University of Cambridge

70 VOLUMES NOW READY

HISTORY AND ARCHAEOLOGY

Ancient Assyria. By Rev. C.H.W. Johns, Litt.D. Ancient Babylonia. By Rev. C.H.W. Johns, Litt.D.

A History of Civilization in Palestine. By Prof. R.A.S. Macalister, M.A., F.S.A.

China and the Manchus. By Prof. H.A. Giles, LL.D.

The Civilization of Ancient Mexico. By Lewis Spence.

The Vikings. By Prof. Allen Mawer, M.A.

New Zealand. By the Hon. Sir Robert Stout, K.C.M.G., LL.D., and J. Logan Stout, LL.B. (N.Z.).

The Ground Plan of the English Parish Church. By A. Hamilton Thompson, M.A., F.S.A.

The Historical Growth of the English Parish Church. By A. Hamilton Thompson, M.A., F.S.A.

English Monasteries. By A.H. Thompson, M.A., F.S.A.

Brasses. By J.S.M. Ward, B.A., F.R.Hist.S.

Ancient Stained and Painted Glass. By F.S. Eden.

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Cash and Credit. By D.A. Barker.

The Theory of Money. By D.A. Barker.

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The Early Religious Poetry of Persia. By the Rev. Prof. J. Hope Moulton, D.D., D.Theol. (Berlin).

The History of the English Bible. By John Brown, D.D.

English Dialects from the Eighth Century to the Present Day. By W.W. Skeat, Litt.D., D.C.L., F.B.A.

King Arthur in History and Legend. By Prof. W. Lewis Jones, M.A.

The Icelandic Sagas. By W.A. Craigie, LL.D.

Greek Tragedy. By J.T. Sheppard, M.A.

The Ballad in Literature. By T.F. Henderson.

Goethe and the Twentieth Century. By Prof. J.G. Robertson, M.A., Ph.D.

The Troubadours. By the Rev. H.J. Chaytor, M.A.

Mysticism in English Literature. By Miss C.F.E. Spurgeon.

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The Idea of God in Early Religions. By Dr F.B. Jevons.

Comparative Religion. By Dr F.B. Jevons.

Plato: Moral and Political Ideals. By Mrs A.M. Adam.

The Moral Life and Moral Worth. By Prof. Sorley, Litt.D.

The English Puritans. By John Brown, D.D.

An Historical Account of the Rise and Development of Presbyterianism in Scotland. By the Rt Hon. the Lord Balfour of Burleigh, K.T., G.C.M.G.

Methodism. By Rev. H.B. Workman, D.Lit.

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Life in the Medieval University. By R.S. Rait, M.A.

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The Administration of Justice in Criminal Matters (in England and Wales). By G. Glover Alexander, M.A., LL.M.

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The Coming of Evolution. By Prof. J.W. Judd, C.B., F.R.S.

Heredity in the Light of Recent Research. By L. Doncaster, M.A.

Primitive Animals. By Geoffrey Smith, M.A.

The Individual in the Animal Kingdom. By J.S. Huxley, B.A.

Life in the Sea. By James Johnstone, B.Sc.

The Migration of Birds. By T.A. Coward.

Spiders. By C. Warburton, M.A.

Bees and Wasps. By O.H. Latter, M.A.

House Flies. By C.G. Hewitt, D.Sc.

Earthworms and their Allies. By F.E. Beddard, F.R.S.

The Wanderings of Animals. By H.F. Gadow, F.R.S.

ANTHROPOLOGY

The Wanderings of Peoples. By Dr A.C. Haddon, F.R.S.

Prehistoric Man. By Dr W.L.H. Duckworth.

GEOLOGY

Rocks and their Origins. By Prof. Grenville A.J. Cole.

The Work of Rain and Rivers. By T.G. Bonney, Sc.D.

The Natural History of Coal. By Dr E.A. Newell Arber.

The Natural History of Clay. By Alfred B. Searle.

The Origin of Earthquakes. By C. Davison, Sc.D., F.G.S.

Submerged Forests. By Clement Reid, F.R.S.

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Plant-Animals: a Study in Symbiosis. By Prof. F.W. Keeble.

Plant-Life on Land. By Prof. F.O. Bower, Sc.D., F.R.S.

Links with the Past in the Plant-World. By Prof. A.C. Seward.

PHYSICS

The Earth. By Prof. J.H. Poynting, F.R.S.

The Atmosphere. By A.J. Berry, M.A.

Beyond the Atom. By John Cox, M.A.

The Physical Basis of Music. By A. Wood, M.A.

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The Psychology of Insanity. By Bernard Hart, M.D.

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The Modern Locomotive. By C. Edgar Allen, A.M.I.Mech.E.

The Modern Warship. By E.L. Attwood.

Aerial Locomotion. By E.H. Harper, M.A., and Allan E. Ferguson, B.Sc.

Electricity in Locomotion. By A.G. Whyte, B.Sc.

Wireless Telegraphy. By Prof. C.L. Fortescue, M.A.

The Story of a Loaf of Bread. By Prof. T.B. Wood, M.A.

Brewing. By A. Chaston Chapman, F.I.C.



SOME VOLUMES IN PREPARATION

HISTORY AND ARCHAEOLOGY

The Aryans. By Prof. M. Winternitz.

Ancient India. By Prof. E.J. Rapson, M.A.

The Peoples of India. By J.D. Anderson, M.A.

The Balkan Peoples. By J.D. Bourchier.

Canada of the present day. By C.G. Hewitt, D.Sc.

The Evolution of Japan. By Prof. J.H. Longford.

The West Indies. By Sir Daniel Morris, K.C.M.G.

The Royal Navy. By John Leyland.

Gypsies. By John Sampson.

A Grammar of Heraldry. By W.H. St John Hope, Litt.D.

Celtic Art. By Joseph Anderson, LL.D.

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Women's Work. By Miss Constance Smith.

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Early Indian Poetry. By A.A. Macdonell.

The Book. By H.G. Aldis, M.A.

Pantomime. By D.L. Murray.

Folk Song and Dance. By Miss Neal and F. Kidson.

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The Natural Sources of Energy. By Prof. A.H. Gibson, D.Sc.

The Sun. By Prof. R.A. Sampson.

Roentgen Rays. By Prof. W.H. Bragg, F.R.S.

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The Life-story of Insects. By Prof. G.H. Carpenter.

The Flea. By H. Russell.

Pearls. By Prof. W.J. Dakin.

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Soil Fertility. By E.J. Russell, D.Sc.

Coast Erosion. By Prof. T.J. Jehu.

INDUSTRIAL AND MECHANICAL SCIENCE

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