A Text-book of Entomology / Including the Anatomy, Physiology, Embryology and Metamorphoses of Insects for Use in Agricultural and Technical Schools and Colleges as Well as by the Working Entomologist

by A. S. Packard

SUMMARY OF THE FACTS AND SUGGESTIONS AS TO THE CAUSES OF METAMORPHISM

An explanation of the causes of metamorphosis is one of the most difficult undertakings in biology, and the phenomenon has been considered as one of the chief difficulties in the way of the acceptance of the theory of descent.

A review, however, of the facts of hypermetamorphism, particularly the life-history of Mantispa, throws much light on the subject, since it is very probable that the supernumerary stages and marked changes of form characterizing them are due to changes of environment, of habits, and of food, causes which have exerted such a profound influence on organic beings throughout all time. Besides these, as the result of changes in the environment and nature of the food, we have the results brought about by the use or disuse of structures brought into existence by the action of stimuli from without, the class of insects abounding in examples of temporary structures which perform a certain function, and then disappear.

Again, if the origin of a hypermetamorphosis can thus be explained, it follows that normal metamorphosis is most probably due to changes of habitat, of seasons, of food, and to accelerated growth resulting from the approach of sexual maturity.

The following facts and conclusions appear to be well established:—

1. The apterous insects (Synaptera) are ametabolous, only the winged insects undergoing a metamorphosis.

2. The complete metamorphosis was not inherited from the primitive ancestor of all insects, but acquired at a later period (F. Müller). The eruciform type is a secondary, adaptive form, derived from the earlier, campodeoid type of larva.

3. The earliest, most primitive pterygote insects passed through only a slight metamorphosis. In other words, as soon as the wings were evolved and insects became adapted to live or take refuge in a new medium, the air, at the approach of the period of adult life, with the ripening or perfection of the reproductive organs, a metamorphosis began to take place, and the number of species greatly multiplied. On the other hand, the Arachnida and Myriopoda, in which as a rule there is no metamorphosis, being confined to a creeping life, with no change of medium, remained poor in number of species.

4. At first the nymphs mainly differed from the adults in lacking wings, though having the same habits; in holometabolous insects, the larva became adapted to entirely different habits and environments, so that in Hymenoptera, and especially Diptera, the larva became remarkably unlike the imago.

5. Until the Mesozoic age, or late in the Carboniferous period, there were, so far as we now know, only ametabolous and heterometabolous insects, and these orders (Orthoptera, Dermaptera, Hemiptera, Plectoptera, Odonata, and Neuroptera) were not numerically rich in genera and species, while since early Mesozoic times geological extinction has reduced their numbers.

6. During the Mesozoic age, and since then, the number of species, genera, families, and orders has greatly increased, and insects have become more and more holometabolous. The orders of Coleoptera, Lepidoptera, Hymenoptera, and Diptera are many fold greater in number of species and variety of form than the heterometabolous orders.

The rapid increase in the number and variety of types of insects evidently is correlated with the profound geological changes which took place at the end of the Paleozoic age, involving the appearance of larger continental masses, or a greater land area, thus opening new regions for settlement. Also the origin of flowering plants at about this time undoubtedly had much to do with the genesis of new adaptive structures, such as the changes in the mouth-parts and wings.

7. The process of metamorphosis, at least in the subtropical, temperate, and polar regions, is largely dependent on the change from summer to winter, and, in the tropics, from the rainy to the dry season.

As regards the organization of larval (nepionic) as compared with imaginal forms, the nymphs and larvæ of insects are, with the exception of many Diptera, nearly as perfectly developed as the adult. In this respect the immature insect differs fundamentally from the larvæ of certain worms (for example, the pilidium of Nemerteans) and from the pluteus and brachiolaria stages of echinoderms, which possess only digestive and water-vascular organs.

Insect nymphs and larvæ also differ from the nauplius and zoëa of Crustacea in having at birth all the most important systems of organs (digestive, circulatory, respiratory, nervous, muscular, with sometimes a nearly perfected reproductive system) of the imago, also the same number of cephalic, thoracic, and abdominal segments and appendages. Metamorphism in insects involves (except in the Diptera) rather modifications in the form and functions of organs and appendages already present than the formation of new ones. In larval Crustacea, the thoracic and abdominal appendages do not arise until some time after hatching from the egg.

8. While cases of the suppression or abbreviation of larval characters and direct development are not uncommon in echinoderms and crustaceans, in insects this phenomenon occurs only so far as yet known in the Diptera. In these insects the polypody in the embryo is outgrown, or lost, the embryos and larvæ not having even the temporary rudiments of abdominal appendages. The campodeoid characters also are entirely suppressed, dropped, or lost in the more specialized holometabolous orders, Lepidoptera, Hymenoptera, and Diptera, though retained in the more primitive and generalized Coleoptera. (This proves that the Coleoptera are lower or more primitive and generalized than the other orders mentioned.) This abbreviation or loss of organs is, as Hyatt and Arms claim, due to the prepotency of acquired characters in phylogeny, and are also the result of homochronous heredity.

“The Insecta of the more specialized orders, x.-xvi., afford, next to some parasites, the most notable examples of this mode of evolution. Their larval or nepionic, and pupal or neanic, stages are prolonged at the expense of the ephebic, winged stage, and the reasons for this prolongation are found in the great number of new features introduced into these stages of development in these orders as contrasted with those of the more primitive, and, in large part, more ancient orders, i.-ix. The law of tachygenesis has been at work here, as in the former cases alluded to above, and it is shown in the encroachments of the adaptive characteristics of the caterpillar, grub, and maggot upon the inherited characteristics of the Thysanuran stage, which loses its ancestral characteristics, until in most cases they are either obsolete or recognizable with difficulty.” (Hyatt and Arms, Natural Science, 1896, p. 400.)

9. In the holometabolous insects there is a resting, quiescent stage during the pupal period, when the insect takes no food. In this respect the more specialized insects differ from other metamorphic animals. The larva has an abundant supply of fat lasting through pupal life, while in the quiescent pupa, respiration and circulation is much lessened, the animal being as a rule motionless. This resting stage is also necessary for the histolysis and formation of the adult body from the imaginal buds present in the larva.

10. The hypermetamorphosis of Mantispa, Meloë, Stylops, etc., indicate very plainly that the eruciform type of larva is derived from the campodeoid, since one and the same insect passes through these stages before reaching sexual maturity.

11. As observed by Miall, the larva of insects differs from that of other invertebrate animals in being larger than the adult.

12. The metamorphoses of insects are in some important respects paralleled by those of the Amphibia. The case of pædogenesis of Chironomus affords a parallel with that of the Siredon, or larva of Amblystoma. Also the organs and appendages of the insects, such as caterpillars, are present, just as the skeleton and other organs of the tadpole are the homologues of those of the adult, although these parts undergo a profound modification, and new structures are added. (See the discussion of this point by Miall, and by Hyatt and Arms.)

Theoretical conclusions; Causes of metamorphosis.—It results from a review of the known facts, together with reasonable inductions from such facts, that so far from opposing the theory of descent, the facts of metamorphosis, and particularly of hypermetamorphosis, seem to afford solid foundation for the theory. While natural selection was not the initiative cause, it plays a part as one of several factors; but the fundamental causes are the same as those which have controlled the origin of species and of the larger groups of animals in general. Owing to the struggle for existence, due to overcrowding, the early insects were forced to take to the air, acquiring wings to enable them to avoid the attacks of creeping and running insects. In the end the insects became, owing to this acquisition of wings, and afterwards to the establishment of a complicated metamorphosis, numerically the most successful type of life in existence, the number of species, extinct and living, mounting into the millions.

All aquatic insects are evidently the descendants of terrestrial forms, and the numberless contrivances and temporary larval organs, particularly of dipterous larvæ, are evidently adaptations to the needs of the insect during its aquatic life, and which are cast aside when the creature passes to a different medium. The sudden or tachygenic appearance of temporary structures, such as hatching spines, various setæ, spines, respiratory organs, so characteristic of dipterous larvæ, and of the protective colors and markings of caterpillars, and which are discarded at pupation, or imagination, are evidently due to the action of stimuli from without, to the primary neolamarckian factors, the characters proper to each larval stadium, and to the pupal and imaginal stadia,—characters probably acquired during the lifetime of the individual,—becoming finally fixed by homochronous heredity.