David Sharp and George Herbert Carpenter

“ Much consideration has been given to the nature of metamorphosis in insects, to its value to the creatures and to the mode of its origin. Insect metamorphosis may be briefly described as phenomena of development characterized by abrupt changes of appearance and of structure, occurring during the period subsequent to embryonic development and antecedent to the reproductive state. It is, in short, a peculiar mode of growth and adolescence. ”

“ The earwigs, cockroaches and locusts, which Linnaeus included among the Coleoptera, were early grouped into a distinct order, the Orthoptera. The great advance in modern zoology as regards the classification of the Hexapoda lies in the treatment of a heterogeneous assembly which formed Linnaeus’s order Neuroptera. The characters of the wings are doubtless important as indications of relationship, but the nature of the jaws and the course of the life-history must be considered of greater value. ”

“ The thorax is composed of three segments; each bears a pair of jointed legs, and in the vast majority of insects the two hindmost bear each a pair of wings. From these three pairs of thoracic legs comes the name—Hexapoda—which distinguishes the class. And the wings, though not always present, are highly characteristic of the Hexapoda, since no other group of the Arthropoda has acquired the power of flight. In the more generalized insects the abdomen evidently consists of ten segments, the hindmost of which often carries a pair of tail-feelers, (cerci or cercopods) and a terminal anal segment. ”

“ The embryo of a moth, a dragon-fly or a bug is invaginated into the yolk at the head end, the portion of the blastoderm necessarily pushed in with it forming the amnion. The embryo thus becomes transferred to the dorsal face of the egg, but at a later stage it undergoes reversion to its original ventral position. In some parasitic Hymenoptera there is only a single embryonic membrane formed by delamination from the blastoderm, while in a few insects, including the wingless spring-tails, the embryonic membranes are vestigial or entirely wanting. ”

“ Such a young insect is a larva—a term used by zoologists for young animals generally that are decidedly unlike their parents. It is obvious that the hatching of the young as a larva necessitates a more or less profound transformation or metamorphosis before the perfect state is attained. Usually this transformation comes with apparent suddenness, at the penultimate stage of the insect’s life-history, when the passive pupa (fig. 21, d) is revealed, exhibiting the wings and other imaginal structures, which have been developed unseen beneath the cuticle of the larva. ”

“ Hence we are inclined to look on the imaginal disks as cellular areas that possess in a latent condition the powers of growth and development that exist in the embryo, powers that only become evident in certain special conditions of the organism. ”

“ Growth and Metamorphosis
After hatching or birth an insect undergoes a process of growth and change until the adult condition is reached. The varied details of this post-embryonic development furnish some of the most interesting facts and problems to the students of the Hexapoda. Wingless insects, such as spring-tails and lice, make their appearance in the form of miniature adults. Some winged insects—cockroaches, bugs (fig. 20) and earwigs, for example—when young closely resemble their parents, except for the absence of wings. ”

“ The sub-imago of the Ephemeroptera suggests that a moult, after the wings had become functional, was at one time general among the Hexapoda, and that the resting nymph of the Thysanoptera or the pupa of the Endopterygota represents a formerly active stage in the life-history. Further, although the wing-rudiments appear externally in an early instar of an exopterygotous insect, the earliest instars are wingless and wing-rudiments have been previously developing beneath the cuticle, growing however outwards, not inwards as in the larva of an endopterygote. ”

“ Head Segments.—In the adult insect the head is insignificant in size compared with the thorax or abdomen, but in the embryo it forms a much larger portion of the body than it does in the adult. Its composition has been the subject of prolonged difference of opinion. ”

“ These differences in larval form depend in part on the surroundings among which the larva finds itself after hatching; the active, armoured grub has to seek food for itself and to fight its own battles, while the soft, defenceless maggot is provided with abundant nourishment. But in general we find that elaboration of imaginal structure is associated with degradation in the nature of the larva, eruciform and vermiform larvae being characteristic of the highest orders of the Hexapoda, so that unlikeness between parent and offspring has increased with the evolution of the class. ”

“ As regards the vast majority of insects, the orders proposed by Linnaeus are acknowledged by modern zoologists. His classification was founded mainly on the nature of the wings, and five of his orders—the Hymenoptera (bees, ants, wasps, &c.) , Coleoptera (beetles) , Diptera (two-winged flies) , Lepidoptera (moths and butterflies) , and Hemiptera (bugs, cicads, &c.) —are recognized to-day with nearly the same limits as he laid down. ”

“ These later stages, comprising the greater part of the larval history, are adapted for an inquiline or a parasitic life, where shelter is assured and food abundant, while the short-lived, active condition enables the newly-hatched insect to make its way to the spot favourable for its future development, clinging, for example, in the case of an oil-beetle’s larva, to the hairs of a bee as she flies towards her nest. ”

“ The number of limb-bearing somites in the insectan head is thus seen to be seven. All of these are to be regarded as primitively post-oral, but in the course of development the mouth moves back to the mandibular segment, so that the first three somites—ocular, antennal and intercalary—lie in front of it. In Lankester’s terminology, therefore, the head of an insect is “triprosthomerous.” The maxillae of the hinder pair become more or less fused together to form a “lower lip” or labium, and the segment of these appendages is, in some insects, only imperfectly united with the head-capsule. ”

“ When the wing is spread, its long axis is more or less at a right angle to the body axis. A wing is an outgrowth from the dorsal and pleural regions of the thoracic segment that bears it, and microscopic examination shows it to consist of a double layer of cuticularized skin, the two layers being in contact except where they are thickened and folded to form the firm tubular nervures, which serve as a supporting framework for the wing membrane, enclose air-tubes, and convey blood. ”

“ The appendages of the abdomen are called cerci, stylets and gonapophyses. They differ much according to the kind of insect, and in the adult according to sex. Difference of opinion as to the nature of the abdominal appendages prevails. The cerci, when present, appear in the mature insect to be attached to the tenth segment, but according to Heymons they are really appendages of the eleventh segment, their connexion with the tenth being secondary and the result of considerable changes that take place in the terminal segments. ”

“ The possession of a variable number of excretory tubes (Malpighian tubes) , which are developed as outgrowths of the hind-gut and pour their excretion into the intestine, is also a distinctive character of the Hexapoda.
The wings of insects are, in all cases, developed after hatching, the younger stages being wingless, and often unlike the parent in other respects. In such cases the development of wings and the attainment of the adult form depend upon a more or less profound transformation or metamorphosis. ”

“ They differ from other Endopterygota in the multiplication of their Malpighian tubes, and from all other Hexapoda in the union of the first abdominal segment with the thorax. Specialized as they are in form, development and habit, they retain mandibles for biting, and in their lower sub-order—the Symphyta—the maxillae are hardly more modified than those of the Orthoptera. From the evidence of fossils it seems that the higher sub-order—Apocrita—can be traced back to the Lias, so that we believe the Hymenoptera to be more ancient than the Diptera, and far more ancient than the Lepidoptera. ”

“ Reproductive Organs.—In the cockroach embryo, before the segmentation of the germ-band has begun, the primitive germ-cells can be recognized at the hinder end of the mesoderm, from whose ordinary cells they can be distinguished by their larger size. At a later stage further germ-cells arise from the epithelium of the coelomic pouches from the second to the seventh abdominal segments, and become surrounded by other mesoderm cells which form the ovarian or testicular tubes and ducts (fig. 19, g) . ”

“ In reference to the structure of the head-capsule in the imago, it appears that the clypeus and labrum represent, as already said, an unpaired median outgrowth of the oral piece. According to W. A. Riley (1904) the epicranium or “vertex,” the compound eyes and the front divisions of the genae are formed by the cephalic lobes of the embryo (belonging to the ocular segment) , while the mandibular and maxillary segments form the hinder parts of the genae and the hypopharynx. ”

“ Accessory glands are commonly present in connexion both with the male and the female reproductive organs. The poison-glands of the sting in wasps and bees are well-known examples of these.
Embryology
The Egg.—Among the Hexapoda, as in Arthropods generally, the egg is large, containing an accumulation of yolk for the nourishment of the growing embryo. Most insect eggs are of an elongate oval shape; some are globular, others flattened, while others again are flask-shaped, and the outer envelope (chorion) is often beautifully sculptured (figs. 20, d; 21, a, b) . ”

“ The Neuroptera, with their similar fore- and hind-wings and their campodeiform larvae, seem to stand nearest to the presumed isopteroid ancestry, but the imago and larva are often specialized. The campodeiform larvae of many Coleoptera are indeed far more primitive than the neuropteran larvae, and suggest to us that the Coleoptera—modified as their wing-structure has become—arose very early from the primitive metabolous stock. ”

“ If it should be objected that the wings so developed would be rudimentary, and that there would be nothing to encourage their development into perfect functional organs, we may remind the reader that we have already pointed out that imperfect wings of Exopterygota do, even at the present time under certain conditions, become perfect organs; and we may also add that there are, even among existing Endopterygota, species in which the wings are usually vestiges and yet sometimes become perfectly developed. ”

“ K. Escherich (1901) , after a new research on the embryology of the muscid Diptera, claims that the fore and hind endodermal rudiments arise from the blastoderm by invagination, and are from their origin distinct from the mesoderm. On the whole it seems likely that the endoderm is represented in part by the yolk, and in part by those anterior and posterior rudiments which usually form the mesenteron, but that in some Hexapoda the whole digestive tract may be ectodermal. ”

“ In other insects the imaginal disks are less completely disconnected from the superficies of the larval hypodermis, and may indeed be merely patches thereof. The number of imaginal disks in an individual is large, upwards of sixty having been discovered to take part in the formation of the outer body of a fly. With regard to the internal organs, we need only say that transformation occurs in an essentially similar manner, by means of a development from centres distributed in the various organs. ”

“ In the adult state no insect possesses more than six legs, and they are always attached to the thorax; in many Thysanura there are, however, processes on the abdomen that, as to their position, are similar to legs. In the embryos of many insects there are projections from the segments of the abdomen similar, to a considerable extent, to the rudimentary thoracic legs. ”

“ The Australian fauna is rich in characteristic and peculiar genera, and New Zealand, while possessing some remarkable insects of its own, lacks entirely several families with an almost world-wide range—for example, the Notodontidae, Lasiocampidae, and other families of Lepidoptera. Interesting relationships between the Ethiopian and Oriental, the Neotropical and West African, the Patagonian and New Zealand faunas suggest great changes in the distribution of land and water, and throw doubt on the doctrine of the permanence of continental areas and oceanic basins. ”

“ Parthenogenesis.—A number of cases are known among the Hexapoda of the development of young from the eggs of virgin females. In insects so widely separated as bristle-tails and moths this occurs occasionally. In certain gall-flies (Cynipidae) no males are known to exist at all, and the species seems to be preserved entirely by successive parthenogenetic generations. In other gall-flies and in aphids we find that a sexual generation alternates with one or with many virgin generations. The offspring of the virgin females are in most of these instances females ”

“ Most of the families and a large proportion of the genera of insects are exceedingly widespread, but a study of the genera and species in any of the more important families shows that faunas can be distinguished whose headquarters agree fairly with the regions that have been proposed to express the distribution of the higher vertebrates. Many insects, however, can readily extend their range, and a careful study of their distribution leads us to discriminate between faunas rather than definitely to map regions. ”

“ On this view the entire food-canal in most Hexapoda must be regarded as of ectodermal origin, the “endoblast” represents mesoderm only, and the median furrow whence it arises can be no longer compared with the blastopore. According to Heymons, the yolk-cells must be regarded as the true endoderm in the hexapod embryo, for he states (1897) that in the bristle-tail Lepisma and in dragon-flies they give rise to the mid-gut. ”

“ But in many highly organized insects a remarkable concentration of the trunk-ganglia takes place, all the nerve-centres of the thorax and abdomen in the chafers and in the Hemiptera, for instance, being represented by a single mass situated in the thorax. The legs, wings and other organs of the trunk receive their nerves from the thoracic and abdominal ganglia, and the fusion of several pairs of these ganglia may be regarded as corresponding to a centralization of individuality. ”

“ The outer cellular layer (ectoderm or “hypodermis”) of insects as of other Arthropods, secretes a chitinous cuticle which has to be periodically shed and renewed during the growth of the animal. The regions of this cuticle have a markedly segmental arrangement, and the definite hardened pieces (sclerites) of the exoskeleton are in close contact with one another along linear sutures, or are united by regions of the cuticle which are less chitinous and more membranous, so as to permit freedom of movement. ”