201. Embryo of Diplax.
Having assumed the creation of our Leptus by evolutional laws, we must now account for the appearance of tracheæ and those organs so dependent on them, the wings, which, by their presence and consequent changes in the structure of the crust of the body, afford such distinctive characters to the flying insects, and raise them so far above the creeping spiders and centipedes. Our Leptus at first undoubtedly breathed through the skin, as do most of the Poduras, since we have been unable to find tracheæ in them, nor even in the prolarva of a genus of minute ichneumon egg parasites, nor in the Linguatulæ and Tardigrades, and some mites, such as the Itch insect and the Demodex, and other Acari. In the Myriopod, Pauropus, Lubbock was unable to find any traces of tracheæ. If we examine the embryo of an insect shortly before birth, as in the young Dragon fly (figure 201, the dotted line t crosses the rudimentary tracheæ), we find it to consist of two simple tubes with few branches, while there are no stigmata, or breathing holes, to be seen in the sides of the body. This fact sustains the view of Gegenbaur[26] that at first the tracheæ formed two simple tubes in the body-cavity, and that the primary office of these tubes was for lightening the body, and that their function as respiratory tubes was a secondary one. The aquatic Protoleptus, as we may term the ancestor of Leptus, may have had such tubes as these, which acted like the swimming bladder of fishes for lightening the body, as suggested by Gegenbaur. It is known that the swimming bladder of fishes becomes developed into the lungs of air-breathing vertebrates and man himself. As our Leptus adopted a terrestrial life and needed more air, a connection was probably formed by a minute branch on each side of the body with some minute pore (for such exist, whose uses are as yet unknown) through the skin, which finally became specialized into a stigma, or breathing pore; and from the tracheal system being closed, we now have the open tracheal system of land insects.
The next inquiry is as to the origin of the wings. Here the question arises if wingless forms are exceptional among the winged insects, and the loss of wings is obviously dependent on the habits (as in the lice), and environment of the species (as in beetles living on islands, which are apt to lose the hinder pair of wings), why may not their acquisition in the first place have been due to external agencies; and, as they are suddenly discarded, why may they not have suddenly appeared in the first place? In aquatic larvæ there are often external gill-like organs, being simple sacs permeated by tracheæ (as in Agrion, Fig. 129, or the May flies). These organs are virtually aquatic wings, aiding the insect in progression as well as in aërating the blood, as in the true wings. They are very variable in position, some being developed at the extremity of the abdomen, as in Agrion, or along the sides, as in the May flies, or filiform and arranged in tufts on the under side of the body, as in Perla; and the naturalist is not surprised to find them absent or present in accordance with the varying habits of the animal. For example, in the larvæ of the larger Dragon flies (Libellula, etc.) they are wanting, while in Agrion and its allies they are present.
Now we conceive that wings formed in much the same way, and with no more disturbance, so to speak, to the insect's organization, appeared during a certain critical period in the metamorphosis of some early insect. As soon as this novel mode of locomotion became established we can easily see how surrounding circumstances would favor their farther development until the presence of wings became universal. If space permitted us to pursue this interesting subject farther, we could show how invariably correlated in form and structure are the wings of insects to the varied conditions by which they are surrounded, and which we are forced to believe stand in the relation of cause to effect. Again, why should the wings always appear on the thorax and on the upper instead of the under side? As this is the seat of the centre of gravity, it is evident that cosmical laws as well as the more immediate laws of biology determine the position and nature of the wings of an insect.
Correlated with the presence of wings is the wonderful differentiation of the crust, especially of the thorax, where each segment consists of a number of distinct pieces; while in the spiders and Myriopods the segments are as simple as in the abdominal segments of the winged insect. It is not difficult here to trace a series leading up from the Poduras, in which the segments are like those of spiders, to the wonderful complexity of the parts in the thoracic segments of the Lepidoptera and Hymenoptera.
In his remarks "On the Origin of Insects,"[27] Sir John Lubbock says, "I feel great difficulty in conceiving by what natural process an insect with a suctorial mouth like that of a gnat or butterfly could be developed from a powerfully mandibulate type like the Orthoptera, or even from the Neuroptera." Is it not more difficult to account for the origin of the mouth-parts at all? They are developed as tubercles or folds in the tegument, and are homologous with the legs. Figure 186 shows that the two sorts of limbs are at one time identical in form and relative position. The thought suggests itself that these long, soft, finger-like appendages may have been derived from the tentacles of the higher worms, but the grounds for this opinion are uncertain. At any rate, the earliest form of limb must have been that of a soft tubercle armed with one, or two, or many terminal claws, as seen in aquatic larvæ, such as Chironomus (Fig. 202), Ephydra (Fig. 203 a, b, c, pupa) and many others. As the Protoleptus assumed a terrestrial life and needed to walk, the rudimentary feet would tend to elongate, and in consequence need the presence of chitine to harden the integument, until the habit of walking becoming fixed, the necessity of a jointed structure arose. After this the different needs of the offspring of such an insect, with their different modes of taking food, vegetable or animal, would induce the diverse forms of simple, or raptorial, or leaping or digging limbs. A peculiar use of the anterior members, as seen in grasping the food and conveying it to the mouth (perhaps originally a simple orifice with soft lips, as in Peripatus), would tend to cause such limbs to be grouped together, to concentrate around the mouth-opening, and to be directed constantly forwards. With use, as in the case of legs, these originally soft mouth-feet would gradually harden at the extremities, until serviceable in biting, when they would become jaws and palpi. Given a mouth and limbs surrounding it, and we at once have a rude head set off from the rest of the body. And in fact such is the history of the development of these parts in the embryo. At first the head is indicated by the buds forming the rudiments of limbs; the segments to which they are attached do not form a true head until after the mouth-parts have attained their jaw-like characters, and it is not until the insect is about to be hatched, that the head is definitely walled in.
202. Foot of Chironomus.
203. Ephydra.