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Read Ebook: The Life-Story of Insects by Carpenter George H George Herbert
Font size: Background color: Text color: Add to tbrJar First Page Next Page Prev PageEbook has 448 lines and 32636 words, and 9 pagesOn the other hand, the digestive system of insects that undergo a metamorphosis, passes through a profound crisis of dissolution and rebuilding. This is not surprising when we remember that there is often a great difference between larva and imago in the nature of the food. The digestive canal of a caterpillar runs a fairly straight course through the body and consists of a gullet, stomach , intestine, and rectum; it is adapted for the digestion of solid food. In the butterfly there is one outgrowth of the gullet in the head--a pharyngeal sac adapted for sucking liquids; and another outgrowth at the hinder end of the gullet --a crop or food-reservoir lying in the abdomen. The intestine of the butterfly also is longer than that of the larva, being coiled or twisted. Towards the end of the last larval stage, the cells of the inner coat lining the stomach begin to undergo degeneration, small replacing cells appearing between their bases and later giving rise to the more delicate epithelium that lines the mid-gut of the imago. The larval cells are shed into the cavity of the stomach and become completely broken down. J. Anglas , describing these microscopic changes in the transformations of wasps and bees, has shown that the tiny replacing cells can be recognised in sections through the digestive canal of a very young larva; they may be regarded as representing imaginal buds of the adult gastric epithelium. In the transformations of two-winged flies of the bluebottle group, A. Kowalevsky has shown that these replacing cells are aggregated in little masses scattered at different points along the stomach and thus corresponding rather closely to the imaginal discs of the legs and wings. The gullet, crop, and gizzard of an insect, which lie in front of the stomach, are lined by cells derived from the outer skin which is pushed in to form what is called the 'fore-gut.' Similarly the intestine and rectum, behind the stomach, are lined with ectodermal cells which arise from the inpushed 'hind-gut.' The larval fore- and hind-guts are broken down at the end of larval life and their lining is replaced by fresh tissue derived from two imaginal bands which surround the cavity of the digestive tube, one at the hinder end of the fore-gut, and the other at the front end of the hind-gut. The larval salivary glands in connection with the gullet are also broken down, and fresh glands are formed for the imago. A large part of the substance of an insect larva consists of muscular tissue, surrounding the digestive tube, and forming the great muscles that move the various parts of the body, and of fat, surrounding the organs and serving as a store of food-material. Very many of the muscle-fibres and the fat-cells also become disintegrated during the late larval and pupal stages, and the corresponding tissues of the adult are new formations derived from special groups of imaginal cells, though some muscles may persist from the larva to the adult. Similarly the complex air-tube or tracheal system of the larva is broken down and a fresh set of tubes is developed, adapted to the altered body-form of pupa and imago. The destruction of larval tissue and the development of replacing organs from special groups of cells, derived of course from the embryo, and carrying on the continuity of cell-lineage to the adult, are among the most remarkable facts connected with the life-story of insects. The process of tissue-destruction is known as 'histolysis'; the rebuilding process is called 'histogenesis.' Considerable difference of opinion has existed as to factors causing histolysis, and for a summary of the conflicting or complementary theories, the reader is referred to the work of L.F. Henneguy . In the histolysis of the two-winged flies, wandering amoeboid cells--like the white corpuscles or leucocytes of vertebrate blood--have been observed destroying the larval tissues that need to be broken down, as they destroy invading micro-organisms in the body. But students of the internal changes that accompany transformation in insects of other orders have often been unable to observe such devouring activity of these 'phagocytes,' and attribute the dissolution of the larval tissues to internal chemical changes. The fact that in all insect transformation a part, and in many a large part, of the larval organs pass over to the pupa and imago, suggests that only those structures whose work is done are broken down through the action of internally formed destructive substances, and one function of the phagocytes is to act as scavengers by devouring what has become effete and useless. LARVAE AND THEIR ADAPTATIONS Among the insects that undergo a complete transformation, there is, as we have seen in the preceding chapter, an amount of inward change, of dissolution and rebuilding of tissues, that varies in its completeness in members of different orders. It is now advisable to consider the various outward forms assumed by the larvae of these insects, or rather by a few examples chosen from a vast array of well-nigh 'infinite variety.' In comparing the transformations of endopterygote insects of different orders, it is worthy of notice that in some cases all the members of an order have larvae remarkably constant in their main structural features, while in others there is great variety of larval form within the order. For example, the caterpillars of all Lepidoptera are fundamentally much alike, while the grubs of beetles of different families diverge widely from one another. A review of a selected series of beetle-larvae will therefore serve well to introduce this branch of the subject. In the larvae of the little timber-beetles and their allies , including the 'death-watches' whose tapping in old furniture is often heard, a marked shortening of the legs and reduction in the size of the head accompany the whitening and softening of the cuticle. This shortening of the legs is still more marked in the larvae of the Longhorn Beetles burrowing in the wood of trees or felled trunks; here the legs are reduced to small vestiges. A fact of much importance in the transformations of beetles as pointed out by Brauer is that in a few families, the first larval instar is campodeiform, while the subsequent instars are eruciform. We may take as an example of such 'hypermetamorphosis' the life-story of the Oil or Blister-beetles as first described by J.H. Fabre , and later with more elaboration by H. Beaur?gard . From the egg of one of these beetles is hatched a minute armoured larva, with long feelers, legs, and cerci, whose task is, for example, to seize hold of a bee in order that the latter may carry it, an uninvited guest, to her nest. Safely within the nest, the little 'triungulin' beetle-grub moults; the second instar has a soft cuticle and relatively shorter legs, which, as the larva, now living as a cuckoo-parasite, proceeds to gorge itself with honey, soon appear still further abbreviated. Later comes a stage during which legs are entirely wanting, the larva then resting and taking no food. The last larval instar again has short legs like the grub of the second period. In connection with this life-history we notice that the newly-hatched larva is not in the neighbourhood of its appropriate food. Hence the preliminary armoured and active instar is necessary in order to reach the feeding place; this journey accomplished, the eruciform condition is at once assumed. In all cases indeed we may say that the particular larval form is adapted to the special conditions of life. A few examples from other orders of endopterygote insects will illustrate this point. The campodeiform type is relatively unusual, but most of the Neuroptera have larvae of this kind, active, armoured creatures with long legs, though devoid of the tail-processes often associated with similar larvae among the Coleoptera. Such are the 'Ant-lions,' larvae of the exotic lacewing flies, which hunt small insects, digging a sandy pit for their unwary steps in the case of the best-known members of the group, some of which are found as far north as Paris. In our own islands the 'Aphis-lions,' larvae of Hemerobius and Chrysopa, prowl on plants infested with 'green-fly' which they impale on their sharp grooved mandibles, sucking out the victims' juices, and then, in some cases, using the dried cuticle to furnish a clothing for their own bodies. Among these insects, while the mouth of the imago is of the normal mandibulate type adapted for eating solid food, the larval mouth is constricted and the slender mandibles are grooved for the transmission of liquid food. From Medea to Berrouaghia the mountain range is broken up into large rounded hills, in part cultivated, but almost wholly open, scantily covered with shrubs or trees and mainly grazed over by large flocks of sheep and goats. About midway between the two places we pass through an open forest of oaks, from whose boles the bark has been removed. A chance acquaintance told us that the cork was removed about once in four years, but this is probably not the true cork oak , which grows under more moist conditions, as between Tunis and Constantine, or between the latter place and Algiers, along the littoral. The altitude of this region is somewhat under 4,000 feet. The first representatives of the cedar forest were encountered as stragglers in the dry wash at the north base of the mountain on which the forest is situated. In part these trees were shapely, with a taper summit, and in part they were short, with a summit broad and flat, in effect like an inverted cone. When the main forest was entered the trees were mainly of the type first characterized, with widely reaching lower branches and slender summits. In exposed places or in older parts of the forest the trees of the second type were often seen; and on the crest of the mountain the most bizarre shapes , the trunks nearly parallel to the ground and the branches hugging the ground. In the upper portion of the forest the trees were more widely separated than in the lower portion, and here and there we met with really large specimens, which must have been very ancient. One of the large trees had a bole which 1 meter above the ground was about 5 meters in circumference. The trees were fruiting freely, but we did not see many seedlings. Why, was not apparent. There were no indications that fires had swept over the mountain recently. The route followed across the High Plateau ran from Boghari to Ain Ossera, Guelt-es-Stel, and Djelfa, which is in the midst of the Saharan Atlas, and from thence to Laghouat. Boghari, situated at the place where the Oued Chelif, having come across the High Plateau, enters the Tellian Atlas on its way through them to the Mediterranean, lies on the northern edge of the High Plateau and in what appears to be a fairly arid region. The oued at this place is rather narrow and has low banks. In its bed, in October, were a few pools of water. On either side is the flood-plain of the oued, several meters in width, sometimes partly under cultivation. Tilled fields are to be seen to the west and not far from the town. From the low mountains immediately to the west of the town the steppes stretch unbroken to mountains bounding the southern horizon, possibly 100 miles straight away. The mountains are the Dj. Sahari, the Saharan Atlas, beyond which lies the desert. The alfa, or bunch-grass, covers large areas in Algeria as also in Spain. In November the long leaves of the grass are dry, tightly rolled, and rush-formed, in place of being flat as during the rainy season or period of growth. The species reproduces largely by means of much-branched rhizomes, from which spring the young, fleshy leaves, enlarged at the base. In Algeria, "situ?es en territoire civil," there are 543,620 acres of alfa, mostly on the High Plateau, but a part along the littoral in the province of Oran, west of Algiers. The leading environmental influence upon the peculiar distribution of the species is apparently that of rainfall, reacting in this respect very like plants with storage organs, avoiding alike regions where the rainfall is excessively heavy or where it is so little as to cause marked desertic conditions. It is apparently confined to sandy soils and is replaced by others wherever the soil of a region is of clay or is charged with any considerable amounts of salts. It is an important article of export from Algeria. Its total tonnage is said to amount to 80,000 each year, bringing approximately ,500,000. It is sent to England, Belgium, and France, and used in the manufacture of fine grades of paper, light, strong, and of a silky texture; also in making baskets, hats, and mats, for which a superior grade of the grass, commanding especially high prices, is employed . Soon after passing the dunes the way lay through a country with low mountains, almost bare of vegetation, where scattering oaks and junipers constitute the only species of plants, until we reached the walled town of Djelfa. The bleakness and the bareness of the environs of Djelfa come with a surprise when one considers that the rainfall of the place is not inconsiderable, about 375 mm., and that the altitude is about 1,110 meters, which insures a fairly low temperature and hence a relatively low evaporation rate. The sparseness of the vegetation is probably partly due to the fact that the rainfall does not occur at one or at two seasons, as nearer the coast, but is distributed fairly evenly between the four seasons, and also to the long occupancy by the Arabs and the French, by which possibly most of the useful native plants, large and small, have long since been destroyed. Somewhat removed from the town, particularly on the mountains to the west, is a forest of pines. Along the streets are many shade trees, as Lombardy poplar, ash, locust, and others, and within the town limits is a small but fine public park and experimental garden with a large variety of shrubs and trees. From Djelfa to Laghouat the road runs through barren mountain passes, and is dreary and of little interest. Tristram's description of the approach to Laghouat, written about 1860, gives very well the present condition of things: The next day's journey was through a rocky desert country. . . . We afterwards passed a low-lying strip of sand-hills on the west, with the marks of an ancient ocean beach; on the east a high range of mountains, with the stratification regular and horizontal. . . . Our next day's ride was by a base of a continuous chain of steep ridges, again with an even water-line very near the crest, and presenting a singular serrated appearance . I counted no less instars a broad barrel-like form . 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 . 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 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 , 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 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 whose transformations are described in detail by Miall and Hammond . 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. Most remarkable, in many respects, of all aquatic larvae are the grubs of the Sand-midges . 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 . 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. 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. 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 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 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 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. 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 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 within which by the processes of histolysis and histogenesis already described the organs of the pupa and imago are built up. This puparium 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. While the maggot breathes by its tail-spiracles, the functional spiracles of the puparium 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. 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 . 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 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 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' and the 'Tussocks' . 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 made the remarkable discovery that in the larvae of certain gall-midges 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 . A less extreme instance of an abbreviated life-story was made known by O. Grimm who saw pupae of Harlequin-midges 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 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 , of some allied genera whose larvae live as parasites on other insects, and occasionally of the Sheep Bot-fly . In such cases we recognise the beginning of a shortened larval period, and Brace's investigations in 1895, summarised by E.E. Austen , have shown that females of the dreaded African Tsetse flies 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 . 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 . 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 , 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 as H. Adler 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' 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 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 how in the August of 1879 hundreds and thousands of 'Painted Ladies' 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 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' , 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 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 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 . 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 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. 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 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. 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. 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 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 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 , C. Brongniart , and A. Handlirsch . 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. Add to tbrJar First Page Next Page Prev Page |
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