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

PART III.—THE METAMORPHOSES OF INSECTS

We have seen that the embryo rapidly passes through extraordinary changes of form, and now, after hatching, especially in the insects with a complete metamorphosis, the animal continues to undergo striking changes in form, in adaptation to different modes of life.

The life of a winged insect, such as a butterfly, fly, or bee, may be divided into four stages: the embryo, or egg state, the larva, pupa, and imago,—the term metamorphosis being applied to the changes after birth, or post-embryonic stages of life. The transformations of the more specialized orders of insects involve wonderful changes of form, which are only paralleled in other types of animals by the metamorphoses of the echinoderms, of certain worms, and of the Crustacea, as well as by those of the frog. An insect, such as a butterfly or bee, during its post-embryonic life lives, so to speak, three different lives, having distinct bodily structures and existing under quite dissimilar surroundings and habits; so that a caterpillar is practically a different animal from the pupa, and the latter from the imago, with different organs, the appendages and other structures being so modified as to be, so far as regards their functions, radically different. These changes of functions or of habits have also been plainly enough the exciting cause of the divergency in structure of what fundamentally is one and the same organ, the change having been brought about by adaptation of the same organs to quite different uses.

The changes are not only observable in the body and its appendages, but also in the internal organs, and consequently are both structural and physiological. The term larva, as applied to the first stage of animals, is a very variable and indefinite one, that of insects in general being a much more highly organized animal than the larva of a worm, starfish, or crustacean.

a. The nymph as distinguished from the larval stage

As there is no marked difference between the different stages of the young in the insects with an incomplete metamorphosis (Heterometabola), the chief difference being the possession of the rudiments of wings and the absence of a resting stage, the terms larva and pupa are in reality scarcely applicable to them, and we much prefer the term nymph, first proposed by Lamarck for the active “pupa” of Orthoptera, Hemiptera, the Odonata and Ephemeridæ, and adopted in part by many. Indeed, in the more generalized and older orders, the larval and pupal stages are not differentiated, though the term larval, in its general sense, will probably always be used; just as we speak of the larval stages of worms, echinoderms, or Crustacea.

Eaton in his elaborate work on the Ephemeridæ employs the term nymph to designate all the aquatic or early stages in the development of the young after hatching, and he urges that the old-fashioned usage of larva and pupa seem scarcely worth retention. “Nymphs are young which live an active life, quitting the egg at a tolerably advanced stage of morphological development and having the mouth-parts formed after the same main type of construction as those of the adult insect.” The word nymph is used in the same sense by McLachlan, and by Cabot. Calvert also applies the term nymph “to the stage of odonate existence between the egg and the transformation into the imago.” On the other hand, Brauer applies the term nymph to the pupa of holometabolous insects. For larval Hyatt proposes the term nepionic.

b. Stages or stadia of metamorphosis

The intervals or periods between the moults or ecdyses of caterpillars and other eruciform larvæ are called stages or stadia; thus, as most caterpillars moult four times, we have five stages or stadia, or stage (stadium) I to V. As observed by Sharp, there is, unfortunately, no term in general use to express the form of the insect at the various stadia; “entomologists say, ‘the form assumed at the first moult,’ and so on.” Hence he adopts a term suggested by Fischer,[^[86]] and calls the insect as it appears after leaving the egg the first instar, and what it is after the first moult the second instar, and so on; hence the pupa, or chrysalis, which assumed that condition after moulting five times would be the sixth instar, and the butterfly itself would be the seventh instar.

c. Ametabolous and metabolous stages

In the Synaptera development is direct, the young differing neither in form, structure, or habits from the adult. Hence they are said to be ametabolous. Since there is an absence of even a tendency to a partial metamorphosis, it is evident that the insects have not inherited a tendency to undergo a transformation, but that it is an adaptation induced in the hexapod type after the first winged insects appeared, and which became more marked in the more specialized insects and at a period comparatively late in geological history, i.e. perhaps at or soon after the beginning of the Carboniferous period.[^[87]]

The transformations of the pterygote insects vary greatly in degree, and it is difficult to draw the line between the grades. Those in which the adults differ from the freshly hatched young only or mainly in having wings are generally said to have an incomplete or gradual metamorphosis. There is no inactive, resting, or pupal stage, and the wings are acquired only after successive moults. Insects with an incomplete metamorphosis are the Orthoptera, Dermaptera, Platyptera (Mallophaga, Plecoptera, Corrodentia, Embidæ), Ephemeridæ, Odonata, Thysanoptera, and Hemiptera, with the exception of the male Coccidæ, in which there is a resting or sub-nymph stage. As regards the number of moults in the Synaptera, Grassi states that in Campodea there is a single fragmentary ecdysis, while Sommers tells us that Macrotoma plumbea sheds its skin throughout life, even after attaining its full size.

As an example of the partial metamorphosis of the hemimetabolous insects we may select that of the locust, in which there are five moults and six stages (instars), as seen in Fig. 558, five of which are nymphal. In the first two stages there are no rudiments of wings, these appearing after the second moult. Besides the acquisition of wings there are slight differences after each moult, both in structure and color, besides size, so that we may always recognize the comparative age and the particular stage of growth of any individual.[^[88]]

We have watched the development of Melanoplus spretus from the egg to the imago, and examined thousands of specimens which show the six stages. On the other hand, European authors differ as to whether there are three, four, or five moults in the migratory locust.[^[89]] It is not improbable that, as is the case with many other insects, the number of moults may vary according to the temperature and food, variation in these agencies causing either retardation or rapidity in development.

Those with a complete metamorphosis are said to be metabolous or holometabolous. (Lang.)

Leach[^[90]] in 1815 gave the name of Ametabola to insects without, and Metabola to insects with a metamorphosis.

Fig. 558.—Partial metamorphosis of Melanoplus femur-rubrum, showing the five nymph stages, and the gradual growth of the wings, which are first visible externally in 3, 3b, 3c.—Emerton del.

Latreille (1831) called insects with an incomplete metamorphosis homotenous (which means similar to the end of life), and those with a complete metamorphosis, polymorphous. For the different degrees of metamorphosis of insects he employed two terms: for the incomplete degree, metamorphosis dimidia, and for the total or pupal, metamorphosis perfecta.

Westwood in his Introduction to the Modern Classification of Insects (1839), taking into account the relation of the larva with the imago, divided insects into two divisions: the Heteromorpha, or those in which there is no resemblance between the parent and its offspring, and Homomorpha, in which the larva resembles the imago, except in the absence of wings.

Fig. 559.—Manometabolous metamorphosis of the cockroach (Phyllodromia germanica) with its four nymphal stadia a-d; e, h, adult; f, female with egg-case; g, egg-case.—From Riley.

From the point of view of the degree of metamorphosis, insects have been divided into Heterometabola and Metabola.

I. Heterometabola.—This group may be divided as follows:

1. Manometabola,[^[91]] embracing those forms with a slight or gradual metamorphosis, but which are active in all the stages, without any resting stage. The orders passing through this degree of metamorphosis are the following: Orthoptera, Dermaptera, Platyptera, Thysanoptera, and Hemiptera (Coccidæ excepted).

In all these groups, the only external differences of importance between the freshly hatched nymph and the adult is the presence of wings. The chief difference internally is the complete development of the sexual glands.

It should be observed, however, that in the last nymph stage of the Thysanoptera the articulations of the limbs are enveloped by a membrane and the wings enclosed in short fixed sheaths; the antennæ are turned back on the head, and the insect, though it moves about, is much more sluggish than in the other state. (Haliday.) Hence here we have a close approach to the following degree.

2. Heremetabola,[^[92]] including those forms with a gradual though slight or incomplete metamorphosis, but with a quiescent or resting stage at the close of the nymph life. Lang has emphasized this stage, calling attention to the fact that the fore legs of the nymph of the 17–year Cicada, which lives underground on the roots of trees, are thick and adapted for digging. The transition from the nymph to the winged adult is signalized by the decided change in form of the fore legs, as well as by the acquisition of the wings. “The last larval stage is, then, what is called quiescent, i.e. the organization of the imago develops within the chrysalis at the expense of the accumulated reserve material.” (Lang.) There seems to be a resting stage, when the insect does not perhaps suck the sap from the roots, and awaits in its chamber its approaching change to the imago; but we should scarcely apply the term pupa to this stage, though the antennæ of the freshly hatched larva are larger and longer than in the fully grown nymph and are distinctly 8–jointed.

3. Hemimetabola.—In this division, so named by Brauer, the changes are more marked, though there is no truly inactive pupa-like stage. The orders are Perlaria (Plecoptera), Odonata, and Plectoptera (Ephemeridæ). The freshly hatched nymphs of these three groups are much alike in shape, that of Perlidæ, and indeed most of the Platyptera, being more generalized, unless we except that of Chloëon; all closely recall Campodea, and are therefore in the Campodea-stage. These nymphs are indeed more generalized than the freshly hatched nymph of Blattidæ, or any other of the orders mentioned except the Platyptera, to which perlids belong. They all have feet, and the body is more or less flattened. (Fig. 560.)

II. Holometabola.—In this division we have for the first time a true larva, and a pupa stage as distinguished from the imago. Moreover, the insect at each stage is distinguished by radical differences in form, surroundings, and in the nature of the food, while the pupa is inactive, usually immovable, and incapable of taking any food, and is often protected by a cocoon spun by the larva. The holometabolous orders are the Neuroptera, Coleoptera, Mecoptera, Trichoptera, Lepidoptera, Siphonaptera, Diptera, and Hymenoptera.

As we have among worms, echinoderms, and Crustacea certain exceptional species in a metamorphic group whose metamorphosis is suppressed, their development being direct, so there is in pterygote insects, though in a very much less degree, cases of direct development. In the wingless cockroaches such as Pseudoglomeris, etc., of the tribe of Periphæriides, in some of which, however, the males are winged, and in the Hemiptera, occur wingless forms such as the lice and bed-bug. The Mallophaga are all wingless, while certain Dermaptera (Chelidura, Anisolabis) are also apterous. The absence of wings in such cases is due to disuse from parasitism, or to a life under stones or in cracks and fissures, where the insects are driven to avoid their enemies, and hence do not need wings. The growth of wings and consequently the development of a metamorphosis is suppressed, so that, as Lang says, “in contrast to the original ametabola of the Apterygota, we have here an acquired ametabola.”

It is rare that, after the rudiments of wings have once appeared in the very young, they should disappear in the late nymph stage; this is, however, said by Walsh to be the case with the Ephemerid Bætisca (Fig. 440). This is a case of retardation in an acquired ametabolesis.