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

THE RESPIRATORY SYSTEM

While land vertebrates breathe by inhaling the air through the mouth into the lungs, insects respire by internal air-tubes (tracheæ), which ramify throughout every part of the body and its appendages. The air enters these tubes through a few openings, called spiracles or stigmata, arranged segmentally in the sides of the body. These tracheæ are everywhere bathed by the blood, and thus the latter is constantly aërated or kept fresh; the blood not, as in vertebrates or as in molluscs, seeking the lungs or gills, or any specialized respiratory portion of the body where the oxygen combines with the hæmoglobin, but the respiratory tubes, so to speak, themselves seek out the blood and the blood-tissue in every part of the insect body, penetrating to the tips of the antennæ and of the legs, entering the most delicate tissues, even perhaps passing through the walls of epithelial cells. As Lang remarks, the want of an arterial vascular system is compensated for as well as conditioned by the extremely profuse branching of the tracheæ.

Fig. 389.—Rat-tailed larva of Eristalis.

The aquatic larvæ of certain dragon-flies (Agrionidæ), may-flies, case-worms, etc., respire by means of tracheal gills or branchiæ, which are either filamental or leaf-like appendages containing tracheæ. Somewhat similar structures appended to the thorax of pupal aquatic Diptera, as in the mosquito and its allies, enable them to breathe while stationed a little beneath the surface of the water. Other larvæ, as the rat-tail larva of Eristalis, etc., lying at the bottom of shallow pools or in ditches, etc., can breathe by raising slightly above the surface a long appendage with two spiracles at the end, through which the air enters the tracheal system. (See p. 461.)

Although Aristotle, as well as the natural philosophers of the Middle Ages, supposed that insects did not breathe, one can easily see that they do by holding a grasshopper or dragon-fly in one’s hand and observing the rhythmical rise and fall of the upper and lower walls of the abdomen, during which the air enters and passes out of the air-openings or spiracles on each side of the body.

It is plain that insects consume very little air, since caterpillars may be confined in very small, almost air-tight tin boxes, and continue to eat and undergo their transformations without suffering from the confinement. According to H. Müller an insect placed in a small, confined space absorbs all the oxygen. Insects can survive for many hours when placed in an exhausted receiver, or in certain irrespirable gases. “Cockroaches in carbonic acid speedily become insensible, but after twelve hours’ exposure to the pure gas they survive and appear none the worse.” (Miall and Denny, p. 165.) Insects of the swiftest flight breathe most rapidly, their great muscular activity requiring the absorption of an abundance of oxygen.

Fig. 390.—Section of Sphinx embryo, showing at s the ectoderm invaginated, and forming the germ of a stigma and trachea (t).—After Kowalevsky.

Warmth, plenty of food, besides muscular activity, increases the demand for oxygen and the quantity of carbonic acid exhaled.

a. The tracheæ

Fig. 391.—Portion of a trachea of a caterpillar, with its branches B, C, D: a, peritracheal membrane; b, nucleus.—After Leydig, from Gegenbaur.

Fig. 392.—Structure of a trachea, diagrammatic: portions of the peritracheal membrane (hy) and chitinous intima (cc) removed to show the structure; in the chitinous intima or endotrachea (cc) can be seen the spiral thickenings or tænidia.—After Lang.

It will much simplify our conception of the nature of the air-tubes when we learn that they originate in the embryo as tubular ingrowths of the integument (ectoderm), these branching and finally reaching every part of the interior of the body. They are elastic tubes, and being filled with air are silvery in color, though at their origin near the spiracles they are reddish or violet bluish; or, in the larva of Æschna, reddish brown, this tint being due to a finely granular pigment situated in the peritoneal membrane.

Fig. 393.—Longitudinal section of the trachea of Hydrophilus piceus: ep, epithelium; cu, cuticula; f, spiral threads.—After Minot.

In their essential structure the tracheæ consist of the chitinous intima, which is a continuation of the cuticle of the integument, and of a cellular membrane or outer layer of cells (a continuation of the hypodermis) called the peritoneal membrane, or ectotrachea (Figs. 392, 393).

Leydig discovered that the spiral filaments are not distinct and separate, but intimately connected with the inner membrane (intima), and he detected the outer or peritoneal membrane, which Chun afterwards found to be epithelial in its nature, Minot stating that it is a true pavement epithelium.

Figure 393 represents a longitudinal section of a large trachea of Hydrophilus, showing the peritoneal membrane (ectotrachea, ep) and the intima or endotrachea, divided into the cuticula (cu), with the darker colored inner layer, in which are embedded the dark-colored tænidia (f).

Fig. 394.—Testis of Anabrus, showing the ramifications of the tracheæ.—After Minot.

Distribution of the tracheæ.—The distribution of the air-tubes, as Lubbock and also Minot state, depends first upon the shape of the organs, and upon the size of those whose size is variable. Around the large, hollow organs (digestive canal, sexual organs) the tracheæ ramify in all directions, forking so that the branches diverge at a wide angle. In the organs which have muscular walls, like the oviduct, the tracheæ run straight when the walls are distended, but have a sinuous course when the walls are contracted. (Minot.)

“Around the organs of more elongated form the branches of the tracheæ run more longitudinally, as is shown by the air-tubes of the muscles, which also present some peculiarities worthy of especial notice.

“A short, thick trunk arrives at the muscular bundle, and dividing very rapidly, breaks up into a large number of delicate tubes, which penetrate between the muscular fibres, then terminating in tubes of exceeding fineness, which at first sight seem to form a network that might well be called a rete mirabile. A closer examination, however, reveals that it is not a real network, but rather an interlacing confusing to the eye. The longitudinal direction of the tracheæ of the muscles presents a striking contrast to the system of divarication represented in Figs. 13 and 14. The course of the tracheæ of the Malpighian tubes is also very curious. There is one large trachea which winds around the tube in a long spiral, giving off numerous small branches which run to the surface of the tube, upon which they form delicate ramifications. Each tube has but a single main trachea, and I think the trachea continues the whole length of the tube, but of this last point I am not quite sure.” (Minot.)

While in the nymphs of Orthoptera the tracheæ very closely resemble those of the adult, in larvæ of insects with a complete metamorphosis the tracheæ differ very much in distribution from those of the adult. The larval tracheæ are also more generalized and more like those of the original type than the tracheæ of perfect insects. (Lubbock.)

In general there are two main tracheæ, one passing along each side of the body, near the digestive canal, connected with its mate by a few transverse anastomosing branches, and sending off a branch to each spiracle, this arrangement being most simple and apparent in the maggots of Diptera. From these two main branches smaller twigs branch off into every part of the body with its appendages, passing among the different organs, often serving as cables to hold them loosely in place; they also penetrate into the component parts of the organ themselves, passing into the fat-bodies, and among the fibres of muscles, where they become finely attenuated and refined like the capillaries of the vascular system of vertebrates. (Figs. 395, 396.)

Fig. 395.—Melanoplus femur-rubrum, showing distribution of air-tubes (tracheæ) and air-sacs; V, main ventral trachea (only one of the two shown); S, left stigmatal trachea, connecting by vertical branches with D, the left main dorsal trachea; c, left cephalic trachea; oc, ocular dilated trachea. From the first, second, third, and fourth spiracles arise the first four abdominal air-sacs, which are succeeded by the plexus of three pairs of dilated tracheæ, I, II, III, in Fig. 396. Numerous air-sacs and tracheæ are represented in the head and thorax. The two thoracic spiracles are represented, but not lettered.

Fig. 396.—D, left dorsal trachea; S, left stigmatal trachea; I, II, III, first, second, and third pairs of abdominal dilated tracheæ, forming a plexus behind the ovaries; 1, pair of enormous thoracic air-sacs; 2, pair of smaller air-sacs; 3–7, abdominal air-sacs; oc, ocular dilated trachea and air-sacs; c, cephalic trachea. The relations of the heart to the dorsal tracheæ are indicated.—Drawn by Emerton from dissections by the author.

In the youngest larva of Corethra plumicornis Weismann ascertained the thickness of the longitudinal stem to be 0.0017 mm. That of the finest tracheal endings in the silk-glands of the silkworm was found by Von Wistinghausen to be 0.0016 mm. (Zeits. f. Wiss. Zool. xlix, 1890, p. 575.) Weismann states that in the larvæ of Corethra and Chironomus the tracheal system is only incompletely developed; the tracheæ are not united with each other, and in the youngest larvæ they do not contain air.

Fig. 397.—Tracheal system of the right side of Machilis maritima: k, head; I, II, III, thoracic segments; 1–10, abdominal segments; s, stigma.—After Oudemans, from Lang.

Each of the two main tracheæ, as Kolbe states, sends off into each segment of the body three branches.

1. An upper or dorsal branch, which supplies the muscles of the dorsal region.

2. A middle (visceral) branch, whose twigs pass to the digestive canal and back to the organs of reproduction.

3. A lower (ventral) branch, whose twigs are distributed to the ganglia and to the muscles of the ventral region.

In certain Thysanura, as a species of Machilis (Fig. 397), we probably have the primitive condition of the tracheal system, the longitudinal and transverse anastomoses being absent, but in other Thysanura (Japyx, Nicoletia, Lepisma, and a few species of Machilis) they are present.

As Kolbe remarks, whether the fine ends of the tracheæ are closed or open, whether after the analogy of the blood capillaries of vertebrates they anastomose with each other, whether the ends of the air-tubes pass between the cells or penetrate into them, these questions are not fully settled. According to Leydig’s[^[61]] latest views the tracheæ penetrate into the cells and unite with the hyaloplasma. Hence the process of respiration in the last instance takes place in the hyaloplasma. This assumption accords with the fact that in the tracheate Arthropods the terminations of the tracheæ carry the atmospheric air into the space bounded by the cellular network, also to the hyaloplasma filling the spaces. Leydig[^[62]] also thinks that the finest tracheal endings penetrate into the muscular tissue and unite with the primitive muscular fibres.

Kupffer is likewise of the opinion that the fine tracheæ penetrate into the cells, and Lidth de Jeude asserts that they enter the epithelial cells, “each cell containing several branches.” Kölliker, Emery, etc., maintain, however, that the tracheal endings lie between the cells. Wielowiejski,[^[63]] in describing the line tracheæ of the phosphorescent organs, thinks that the tracheal endings (tracheal capillaries) rarely end blindly, but anastomose with one another, forming an irregular network. The latest observer, Gilson (1893), asserts that tracheal twigs penetrate deeply into the epithelial cells of the silk glands of larval Trichoptera as well as of caterpillars, passing through their protoplasm.

Fig. 398.—Tracheal network of the male glands of Lampyris splendidula: tec, tracheal end-cells; cap, tracheal capillaries; at a, an expanded matrix.—After Wielowiejski.

Fig. 399.—Tracheal capillary end-network (tr. c. n.) of silk glands of Ocneria dispar: p, peritoneal (peritracheal) membrane.—After Wistinghausen.

A late investigator, C. von Wistinghausen, finds in the tracheæ of the spinning-glands of caterpillars a completely formed network between the terminal branches of two or several tracheal groups. The tracheal tubes of this series of terminal branches pass into this network, which he calls the tracheal capillary end-network (Figs. 398, 400). This last varies in thickness and spreads out under the membrana propria of the glandular mass over the entire surface of the large gland-cells and on a level with the tracheal capillaries. The tracheal endings do not penetrate into the cells, but are separated from the plasma of the cells by a thin membrane. The tracheal capillary end-network appears as a system of fine tubes like the tracheal capillaries, consisting of a peritoneal layer and a chitinous intima (Fig. 400). The walls of these tubes are homogeneous, not porous, though readily permeable by the parenchymatous fluid. The interchange of gases consequently may go on easier and more vigorously in a system of richly anastomosing tubules of the net-like mass of tracheal capillaries, than in tubes ending blindly.

While the diameter of the tracheal capillaries is 0.0016 mm. or 1 µ, that of the tubules composing the tracheal capillary end-network is scarcely measurable, but is less than 1 µ.

Fig. 400.—Tracheal end-cells of Lampyris splendidula: tr, trachea with tænidia; tre, tracheal capillaries.—After Wielowiejski.

These tracheal capillaries also occur on the seminal and other sexual tubes, on the intestine, on the urinary tubes, on the fat-bodies, but are most easily detected on the silk-glands.

The latest researches are those of E. Holmgren, who has studied the branching of the tracheæ in the spinning-glands of caterpillars. He prefers to call the end-cells “transition cells,” as they lead from the tracheal tubes proper to the capillary network. This latter is formed by slender nucleated cells, often with an intracellular lumen, and, according to the author, probably constituting a respiratory epithelium. He finds that both large and small tracheæ may penetrate the gland-cells. (Anat. Anzeiger, xi, 1895, pp. 340–6, 3 figs.; Jour. Roy. Micr. Soc., 1896, p. 182.)

b. The spiracles or stigmata

The spiracles are segmentally arranged openings in the sides of the thorax and abdomen, through which the air passes into the air-tubes. In its essential structure a spiracle, or stigma, is a slit-like opening surrounded by a chitinous ring, the lips or edges of the opening being membranous and closed by a movable valve of the spiracle attached by its lower edge, which is closed by an occlusor muscle (Fig. 401). The aperture when open forms a narrow oval slit; and in most insects the slit is within guarded by a row of projecting spines or setæ, which form a lattice work or grate to keep out dust, dirt, fluids, etc.

Fig. 401.—Horizontal section of left third stigma and trachea of Melolontha vulgaris, showing the chamber or drum leading into the trachea: a, a, external frame or valve protecting the outer opening of the stigma; b, c, c, inner frame closing the entrance into the trachea (l, k); m, occlusor muscle closing the inner orifice.—After Straus-Dürckheim.

Krancher[^[64]] has described five leading types of stigmata, not, however, taking into account those of the Synaptera.

I. Stigmata without lips (Primitive or generalized stigmata).

a. The simplest stigma is an aperture which is kept open by a chitinous ring (Acanthia). The opening may be round or elliptical. There are no lips nor any movement of the edges to be observed. Such air-holes occur in the abdomen of bugs (Hemiptera) and beetles (Coleoptera); within the opening of the stigmata in the same insects is a funnel-like contraction. Also in the Diptera the abdominal stigmata are of the same type.[^[65]] The stigmata of the Pulicidae (Siphonaptera) are more complicated, as the edges of the openings are provided with setæ (Fig. 402).

Fig. 402.—First abdominal spiracle with a part of the trachea of the cat-flea: sp, spiracle; t, trachea.

Fig. 403.—Stigma of Melolontha larva, seen from without: b, bulla; s, sieve-like plate; o, curved slit-like opening.—After Boas.

b. The stigma consists of a series of minute single stigmata, which are usually surmounted by a common chitinous ring, and whose tubular continuations unite within in a common trachea, so that the single tubes pass off from the stigma like the fingers on the hand. This form is found in the larvæ and puparia of Diptera.

II. Stigmata with lips (Secondary more specialized stigmata).

c. The lips are represented by a single chitinous ring, with sparse spines. One side of the stigma is a little higher, and partly overlaps the other posteriorly; this form is peculiar to the Orthoptera and Libellulidae.

d. The lips are roof-like, bent inwards and densely hairy, forming a peculiar kind of felting. The setæ of the lips are in most beetles and many Lepidoptera separate, and more or less branched. In caterpillars, the setæ are so finely branched as to form a loose felt, or sieve-like arrangement.

e. The stigmata are round, with a very broad border and a concentric middle portion, the structure being complicated. The concentric middle portion is pouch-like and bears the occlusor muscle. This form occurs in the larvæ of lamellicorn beetles, and can be seen with the naked eye, or with a lens, in Oryctes, Cetonia, and Melolontha (Fig. 403).

f. Over the outer opening of the spiracle is an incurved chitinous projection, on one side of which the trachea takes its origin. It is thus in the Hymenoptera.

The remarkable grate-like stigma of the lamellicorn larvæ has the appearance as if the outer closing plate or valve were impenetrable. The earlier observers considered these stigmata to be open, but Meinert regards them as closed; Schiödte, however, has observed by pressing a preserved specimen of a Melolontha larva the alcohol within passing out in drops, through the grate-like plate, and hence he considers this a proof that the stigma is permeable (Kolbe).

More recently (1893) Boas has examined the same structure in the same species of larva as examined by Schiödte, and he finds it to be open only during the process of moulting. He finds that on each side of the larva there are nine short and wide stigmatic branches, each of which is shut off from the exterior by a brown plate; this consists of a reniform sieve-plate, and of a curved bulla which fits into the cavity of the plate. The stigmatic branch, however, is provided with a large external opening, which is homologous with the stigma, but which is usually closed by the plate and bulla, and is only open during the moulting; at first it is circular, but later becomes a cleft. A transverse section shows that the bulla is a simple tegumentary fold, the outer chitinous layer of which has become especially firm. The plate forms a horizontal half-roof, which springs from one side of the tracheal orifice, and is supported by obliquely set bases, which spring from the adjoining part of the inner side of the tracheæ. The plate and bars are purely cuticular structures. (Zool. Anz., 1893; also Journ. Roy. Micr. Soc., p. 54.)

The tracheal system of libellulid nymphs is not closed; on the other hand, in the fully-grown nymphs the anterior stigmata occurring on the dorsal side are large, and the tracheæ arising from them are thick. These stigmata are permeable by the air. In half-grown and still younger stages of Æschna the two anterior thoracic stigmata are undeveloped. In order to breathe, the fully-grown nymph either rises up on the upper side and elevates the end of the body to the surface in order to take the air into the rectum, or it rests with the back of the thorax at the surface in order to breathe through the large stigmata. The young nymphs take in air only through the rectum. The young nymphs of Libellula and its allies, on the other hand, possess large thoracic stigmata, but they prefer to breathe through the rectum. The fully-grown nymphs of Agrion breathe through the thoracic stigmata. (Dewitz, in Kolbe.)

The position and number of pairs of stigmata.—The spiracles are usually situated in the soft membrane between the tergites and pleurites, but their exact position varies in different groups. In the Coleoptera they occupy on the thorax a more ventral position, and on the abdomen are placed near the edge of the dorsal side, under the elytra. In the dragon-flies, the first pair is situated much more dorsally than the second and third pairs; the following seven pairs are almost wholly ventral and lie concealed in the membranous fold near the external plate. In the Hemiptera, also, the abdominal stigmata, though entirely free and visible, are situated ventrally.

Primarily, in the embryo a pair of stigmata appear on each segment of the thorax and abdomen, except the 10th and 11th, and even possibly in the head, for a pair of stigmata are said to occur in the head of Podurids (Smynthurus) (Lubbock), though this statement needs confirmation. Scolopendrella, however, is known to possess a pair of cephalic spiracles.

From the foregoing statement it will be seen that while in existing winged insects no more than 10 (in Japyx 11) pairs of stigmata are to be found in any one species, yet that 12 segments of the body, in different groups taken collectively, bear them. The primitive number of pairs of spiracles, therefore, in winged insects, was 12, i.e. a pair in each thoracic segment, and a pair in each of the first nine abdominal segments. Insects were originally all holopneustic, and gradually as the type became differentiated into the different orders they became peripneustic or amphipneustic, and, in certain aquatic forms, apneustic. (See pp. 459, 461.)

In the still more primitive, probably wingless, ancestors of insects there was a larger number of stigmata. Hatschek, in 1877, discovered a pair of tracheal invaginations in each of the three posterior head-segments of the embryo of a moth, with stigmatal openings in the 1st and 2d maxillary segments.

Thus early in embryonic life every segment of the body, except those bearing the eyes and the last abdominal, bore a pair of stigmata, so that the primitive insect had at least 15, and perhaps more, pairs of stigmata.

The position of the stigmata is subject to much variation, the result of adaptation to this or that mode of life. Examples are those insects which live in dusty situations or usually more or less concealed in the earth, as in most beetles, and in the Hymenoptera. In such beetles, the stigmata are situated in the thin membrane between the segments; in the Hymenoptera, on the upper edge of the segments. In the Siphonaptera, Pediculina, bed-bug, and similar forms, which breathe an air freer from dust, the spiracles lie free on the outside of the body.

“When the stigmata are free and without any protection on the abdomen, there are other ways by which the entrance of foreign bodies into the tracheæ is prevented. In such cases the body is covered with dense hairs, as in most Diptera and Neuroptera, as well as many Lepidoptera; or there is situated in front of the stigma either a small fissure which is covered over by a number of hairs arising from the edge, as in many Orthoptera; or, as in most insects, a luxurious growth of hairs on the inside of the stigma forms a thick filter for the air. Thus we see that also in this respect each species of insect is completely adapted to its surroundings.” (Krancher.)

Fig. 404.—A, thoracic stigma of the house-fly: Sb, valve which closes the opening.

Fig. 405.—Diagrammatic figures of the internal apparatus which closes the trachea, in the stag-beetle: A, trachea open; in B, closed; St, the stigma, with its grated lips; Ct, cuticula of the body-walls; Vk, closing pouch; Vbü, closing bow; Vba, closing band; M, occlusor muscle.—From Judeich and Nitsche.

The closing apparatus of the stigma.—Whether the external opening of the stigma is permanently open or closed, communication with the tracheæ may be cut off at pleasure during respiration by an internal apparatus of elastic chitinous bands and rods and the occlusor muscle.

The parts concerned in this operation are: 1. The closing bow; 2. The closing lever or peg; 3. The closing band; 4. The occlusor muscle (Figs. 405, 406).

Fig. 406.—Stigma, with the closing apparatus, of Smerinthus populi (imago), seen from within: b, closing bow; c, closing band: o, stigmatic opening; r, external chitinous ring; l, closing lever; m, occlusor muscle; s, scales which lie like roofing tiles over the stigma.—After Krancher.

“The first three parts are chitinized; they form a ring around the stigmatic opening, and are united to each other by joints. The bow is usually crescentic and as a rule surrounds one-half of the trachea. On the other side is the closing band which, by different contrivances, representing the closing lever or peg, becomes closely pressed against the closing bow. This lever is usually of the shape of a slender chitinous rod, which causes the closure; but it can also bend rectangularly, become converted into a typical lever as in the Lepidoptera, or it may assume the form of two peg-like processes, which press with their base against the closing bow.” (Krancher.)

“The closure of the spiracular opening is effected by the contraction of the muscles, while the opening is due to the elasticity of the chitinous parts. When at rest the spiracle is naturally open, so that the air in the trachea can directly communicate with the external air. Usually one end of the muscle is attached to the closing peg, and the other end to the closing bow. Where, as in Melolontha, the closing apparatus is provided with two levers, then naturally the muscle binds these two together and brings about by powerful contractions a firm closure of the trachea”; but, remarks Krancher, “this is not the only kind; there are numerous modifications. Besides the form just described, the levers assume the form of valves (Sirex), or of a brush (Pulex); or of a ring (larvæ of Diptera) with a circular muscle attached to it; or of a ring which simply becomes compressed (thoracic stigmata of Diptera).”

c. Morphology and homologies of the tracheal system

As first shown by Bütschli, the tracheal system is a series of segmentally arranged tubular invaginations of the ectoderm; a pair of stigmata primitively occurring on every segment of the body except perhaps the most anterior, and the last two or last one, a reduction in their number having since taken place, until in the Podurans none have survived. In the supposed ancestor of myriopods and insects, Peripatus, there are tracheæ; but they are very fine, simple, not-branched chitinous tubes which are united into tufts at the base of a flask-shaped depression of the integument, the outer aperture of which depression is regarded as a stigma. In one species (P. edwardsii) these tufts and their openings are scattered irregularly over the body; but in another kind (P. capensis) some of the stigmata at least show traces of a serial arrangement, being disposed in longitudinal rows—two on each side, one dorsally and one ventrally, those of each row, however, being more numerous than the pairs of legs. (See p. 9 and Fig. 4, D.)

It should be observed that in Peripatus, which does not possess urinary tubes, the segmental organs or nephridia are well developed, hence the tracheal tubes coexisting with them cannot be their homologues. We are therefore compelled to regard the tracheal system as of independent origin, arising in the earliest terrestrial air-breathing arthropod, and not indebted for its origin to any structure found in worms, unless perhaps, as both Kennell and Lang suggest, to dermal glands, since, according to Kennell, certain Hirudinea and many Turbellarian worms possess long, mostly unicellular, glands which spread far through the parenchyma of the body. (Kennell.)

Thus Kennell supposes that the ancestors of the Tracheates had spiracles on every segment of the body where the internal organization allowed them to exist. “The reduction of the breathing holes to a smaller number, and their restriction of a pair only to a single segment, was brought about partly by adaptation to a peculiar mode of life,—as insect larvæ especially teach us,—partly also—I may say mechanically—as a result of the obstruction to their development made by the growth or excessive development of other organs.” Among these he reckons the thick, dense cuticula of the integument, the internal fusion of several segments to form body-regions, and the arrangement and great development of the muscles in the head and thorax, etc. (p. 29.)

Fig. 407.—Section through a tracheal pit and diverging bundles of tracheal tubes taken transversely to the long axis of the body: tr, tracheæ, showing rudimentary spiral fibre; tr. c, cells resembling those lining the tracheal pits, which occur at intervals along the course of the tracheæ; tr. o, tracheal stigma; tr. p, tracheal pit.—After Balfour, from Sedgwick.

Kennell has suggested the origin of the tracheæ of Peripatus from the unicellular dermal glands of annelidan ancestors, since he has found glands in certain land-leaches of tropical America, which are provided with enormously long tubular passages united into bundles and opening externally, these tubes appearing to be slightly chitinized. Fig. 407 will show the appearance of a bundle of fine tracheal tubes of Peripatus ending at the bottom of a follicle formed by a deep invagination of the integument, which may be regarded as a primitive spiracle. (See Kennell, Ueber einige Landblutegel des tropical America, Zool. Jahrb. ii, 1886; also Die Verwandtschaftsverhältnisse der Arthropoden, 1891, p. 25.) We may add that Carrière supposes from his study of the embryology of the wall-bee (Chalicodoma muraria), published in 1890, that not only the salivary glands, but also the tentorium, are homologues of the tracheæ, while other structures than tracheæ may have evolved from unicellular dermal glands, which are widely distributed. It may in this connection be observed that some authors derive the book-lungs or book-leaf tracheæ of Arachnida from the gills of Limulus; hence if those of Arachnida arose from quite different and more specialized organs than dermal glands, it is not impossible that the tracheæ of Peripatus, Myriopods, and insects arose de novo, and then we need not look for any primitive structures in worms from which they arose.

Although Bütschli in 1870 in his embryology of the honey-bee called attention to the “great similarity which the eleven pairs of invaginations in the eleven first trunk-segments in their first indication (anlage) have with the spinning-glands, and also with the segmental organs of Annelids,” he did not go further than this, and it is now known that in the 2d maxillary segment open not only spinning-glands, but in the embryo a pair of stigmata.

Paul Mayer, however, regarded the tracheæ and urinary tubes as homodynamous structures, and this view was advocated by Grassi (1885) for the reason that while in the embryo honey-bee there are ten pairs of stigmata, the first thoracic and two last abdominal segments wanting them, the germs of the urinary tubes arise in a corresponding situation on the two last abdominal segments. To this view Emery (Biol. Centralb., 1886, p. 692) objects that in Peripatus the nephridia and tracheæ “have nothing to do with the segmental organs,” as Peripatus besides nephridia possesses both coxal glands and tracheæ.

Both Kennell and Lang derive the coxal glands of Arthropoda from the setiparous or parapodial glands of annelid worms, and the recent endeavor of Bernard to show that the tracheæ arose from setiparous glands seems to be disproved by the fact that in insects as well as in other Arthropoda coxal glands with their outlets exist in the same segments as those bearing stigmata. Reasoning by exclusion, we are led to regard Kennell’s original view as the soundest.

Patten, however, regards the tracheæ as modified ends of nephridia, remarking: “Since in Acilius some of the abdominal tracheæ at first communicate with the cavities of the mesoblastic somites, it is probable that all the tracheæ represent the ectodermic portions of the nephridia.” (Origin of Vertebrates from Arachnids, p. 355.)

It is probable, therefore, that the tracheæ first arose as modifications of dermal glands, as in mites and Peripatus, and that at first they were not provided with tænidia (as in Chilopoda), while in later forms tænidia were developed. In the earliest tracheate forms the stigmata were not segmentally arranged, probably appearing irregularly anywhere in the body, but afterwards in the myriopods and insects became serially arranged.

d. The spiral threads or tænidia

It is generally supposed that the so-called “spiral thread” forms a continuous thread from one end of a tracheal branch to the other. This was first shown not to be the case by Platner in 1844. Minot has proved that “there is not a single spiral thread, but several, which run parallel to one another and end after making a few turns around the trachea.”

The tænidia we have found to be in some cases separate, independent, solid rings, though when there is more than one turn the thread necessarily becomes spiral. The tænidia of a main branch stop at the origin of the smaller branches, and a new set begins at the origin of each branch. The tænidia at the origin of the branch do not pass entirely around the inside of the peritoneal membrane; in the axils they are short, separate, spindle-shaped bands (Fig. 409).

At one point in the main trachea of the larva of Datana the tænidia were seen to end singly on one side (at a considerable distance from any branch or axil) at intervals, with a tænidium situated between them, making four or five turns; then there is only one band situated between two ends; this band or thread is succeeded by a set with five turns between the two ends, this set being succeeded by one complete ring situated between two ends; in all cases the ends vary in length, some threads being short and others long, so that they apparently end anywhere along the circumference of the trachea, and this arrangement is seen to apparently extend along the whole length of the trachea. Hence it is seen that as a rule the tænidia vary much in length, and never, as generally supposed, pass continuously from one end to another of a tracheal branch, for there are many spirals in a branch, each making only from one to five turns, most usually four turns. Fig. 408, part of a trachea of Dyticus marginatus, shows that at a slight bend in a trachea the tænidia is interrupted, and short, incomplete, wedge-shaped tænidia (e) are interpolated; at A, d is seen a split in one of the tænidia (compare also MacLeod, Pl. 1, Fig. 9). The threads are quite irregular in width. In the axils of the branches there is, as seen in Fig. 409, a basketwork of independent, short, often spindle-shaped tænidia; these are succeeded by longer ones, until we have threads passing entirely around near the base of each new branch; these being succeeded by others which make from two to five spiral turns.

Fig. 408.—Tænidia of Dyticus: d, a split tænidium; e, e, ends of tænidia.

The shape of the tænidia appears to vary to a great extent. In lepidopterous insects we have observed them to be in their general shape rather flat and slightly concavo-convex, the hollow looking towards the centre of the trachea. Minot’s section (Fig. 393) shows that in Hydrophilus they are cylindrical and solid, and Chun states that those of Stratiomys are round, while in Eristalis they are round, with a ridge projecting into the cavity of the trachea; in Æschna the thread is quadrangular. MacLeod states that sometimes it is cylindrical, in other cases flat, likewise prismatic; Macloskie believes that the spiral threads of the centipede are “fine tubules, externally opening by a fissure along their course.”

Fig. 409.—Tænidia of Dyticus in an axil of two branches: e, e, ends of tænidia.

Stokes confirms Macloskie’s statements, stating that in the hemipterous Zaitha fluminea “the tænidia are fissured tubules formed within and from chitinized folds of the intima, the convexity of the folds looking towards the lumen of the tracheæ.” In Fig. 414, 1, are represented portions of several tænidia showing the fissure, which is sometimes interrupted; at 2 are seen “the formation of what may be called apertures in a chitinous bridge.” Stokes regards the tænidia as “inwardly directed folds of the membrane.” Near the spiracles the tracheal membrane is externally studded with minute papillæ, as shown at 3, where are represented three broad and incomplete tænidia, with the tapering end, or the beginning, of another. Stokes adds, “Here they are only broad grooves, with no appearance of the narrow fissure of the completed tænidium. At 4 is figured a portion of the internal surface of a large trachea near the external orifice, the tænidia being in an incipient stage, evidently forming more or less of a network, as is usually the case next to the stigma” (compare p. 451, and Fig. 414).

Fig. 410.—End of salivary duct in base of proboscis of Stomoxys calcitrans: a, incomplete and irregular tænidia; b, two tænidia making incomplete rings near the distal end of the duct.

The tracheæ of chilopod myriopods appear to be like those of insects. A number of authors have failed to detect the spiral threads in the Julidæ. As to the Arachnida, several observers, including Menge and Bertkau, have denied the existence of the spiral thread in the spiders with the exception of the Attidæ; and MacLeod finds them “scarcely visible” in Argyroneta.

Besides the tracheæ, the salivary duct is kept permanently distended by tænidia, which, however, are not spiral. They usually form incomplete rings, as in Stomoxys, arranged as shown in Fig. 410.

The labella (proboscis) of flies are supported by incomplete chitinous tubes or “pseudo-tracheæ,” the ends of which form the scraping teeth, this being, according to Dimmock, their primary function. Dimmock describes them as cylindrical channels opening on the surface in zigzag slits. These channels are held open by incomplete rings, one end of which is forked. “These rings are apparently arranged so that one has its fork on one side of the opening of the channel, the next ring the fork on the opposite side of the channel, and so on, in alternation. Their true structure is revealed when flattened out.”

Fig. 411.—Abdominal spiracle (left side) of cockroach (P. americana), side view, showing the bow: p, lateral pouch of spiracle (in centre) seen from within. The tessellated structure of spiracle and trachea shown at A, and the margin of the external aperture at B.—After Miall and Denny.

The use of the elastic tænidia is to render the tracheæ elastic, and to keep them permanently open, as is the case with the parallel rings of the trachea of the higher vertebrates. The tracheæ are thus rendered firm and solid, at the least expense of chitinous material. The spiral thread, as MacLeod remarks, “is the realization in nature of what engineers call a form of the greatest resistance.”

The tænidia are wanting in the fine endings of the tracheæ (tracheal capillaries); also in the cockroach, according to Miall and Denny, they are not developed in the large tracheæ close to the spiracles, and the intima or wall of the tube has a tessellated instead of a spiral marking (Fig. 411). The same structure is seen in the Perlidæ (Nemoura, Gerstaecker, Zeit. f. wissen. Zool. xxiv, Taf. xxiii, Figs. 5 and 7); also in Æschna (Hagen, Zool. Anz. 1880, p. 159). In certain fine tracheæ of the eyes of the fly no spiral threads are developed. (Hickson.) The air-sacs or dilated tracheæ are also without tænidia.

While in the living insect the main and smaller tracheæ are filled, with air, it is stated by Von Wistinghausen that the fine capillary ends contain a fluid.

e. Origin of the tracheæ and of the “spiral thread”

While we owe to Bütschli the discovery of the mode of origin and morphology of the tracheæ, which as he has shown[^[66]] arise by invaginations of the ectoblast; there being originally a single layer of epiblastic cells concerned in the formation of the tracheæ; we are indebted to Weismann[^[67]] for the discovery of the mode of origin of the “intima,” from the epiblastic layer of cells forming the primitive foundation of the tracheal structure.

Weismann did not observe the earliest steps in the process of formation of the stigma and main trunk of the tracheæ, which Bütschli afterwards clearly described and figured.

Weismann, however, thus describes the mode of development of the intima; after describing the cells destined to form the peritoneal membrane, he says: “The lumen is filled with a clear fluid and already shows a definite border in a slight thickening of the cell-wall next to it.

“Very soon this thickening forms a thin, structureless intima, which passes as a delicate double line along the cells, and shows its dependence on the cells by a sort of adherence to the rounded sides of the cells (Taf. vii, 97 A, a b c). Throughout the mass, as the intima thickens, the cells lose their independence, their walls pressing together and coalescing, and soon the considerably enlarged hollow cylinder of the intima is surrounded by a homogeneous layer of a tissue, whose origin from cells is recognized only by the regular position of the rounded nuclei (Taf. vii, Fig. 97, B).

“Then as soon as the wavy bands of the intima entirely disappear, and it forms a straight, cylindrical tube, a fine pale cross-striation becomes noticeable (vii, 97, B, int), which forms the well-known ‘spiral thread,’ a structure which, as Leydig has shown, possesses no independence, but arises merely from a partial thickening of the originally homogeneous intima.

“Meyer’s idea that the spiral threads are fissures in the intima produced by the entrance of air is disproved by the fact that the spiral threads are present long before the air enters. Hence the correctness of Leydig’s view, based on the histological structure of the tracheæ, is confirmed by the embryological development, and the old idea of three membranes, which both Meyer and Milne-Edwards maintain, must be given up.”

Weismann also contends that the elastic membrane bearing the “spiral thread” is in no sense a primary membrane, not corresponding histologically to a cellular membrane. On the contrary, the “peritoneal membrane comprises the primary element of the trachea; it is nowhere absent, but envelops the smallest branches, as well as the largest trunks, only varying in thickness, which in the embryo and the young larva of Musca stands in relation to the thickness of the lumen.”

The trachea, then, consists primarily of an epithelial layer, the “peritoneal membrane,” or the invaginated epiblast; from this layer an intima is secreted, just as the skin or cuticle is secreted by the hypodermis. We may call the peritoneal membrane the ectotrachea, the intima or inner layer derived from the ectotrachea the endotrachea. The so-called “spiral threads” are a thickening of the endotracheal membrane, sometimes arranged in a spiral manner. For these chitinous bands we have proposed the name tænidia (Greek, little bands).

As to the origin of the spiral thread our observations[^[68]] have been made on the caterpillar of a species of Datana, which was placed in alcohol, just before pupation, when the larva was in a semipupal condition, and the larval skin could be readily stripped off. At this time the ectotrachea of the larva had undergone histolysis, nothing remaining but the moulted endotrachea, represented by the tænidia, which lay loosely within the cavity of the trachea. The ectotrachea or peritoneal membrane of the pupa is meanwhile in process of formation; the nuclear origin of the tænidia is now very apparent.

Fig. 412.—Longitudinal section of a trachea, showing the origin of the tænidia.

Fig. 413.—Origin of the tænidia from nuclei.

Fig. 412 represents a longitudinal section through a secondary tracheal branch, showing the origin of the chitinous bands, or tænidia. At t′ are pieces of six tænidia which have been moulted; ectr indicates the nuclei forming the outer cellular layer, the ectotrachea or peritoneal membrane. These nuclei send long slender prolongations around the inside of the peritoneal membrane; these prolongations, as may be seen by the figure, become the tænidia. The tænidia, being closely approximate, grow together more or less, and a thin endotracheal membrane is thus produced, of which the tænidia are the thickened band-like portions. The endotracheal membrane is thus derived from the ectotrachea, or primitive tracheal membrane, and the so-called “spiral thread” is formed by thickenings of the nuclei composing the secondary layer of nuclei, and which become filled with the chitin secreted by these elongated nuclei. The middle portion of the tænidia, immediately after the moult, is clear and transparent, with obscure minute granules, while the nuclear base of the cell is filled as usual with abundant granules, and contains a distinct nucleolus.

Fig. 414.—Tænidia and internal hairs of Zaitha.—After Stokes.

The origin of the tænidia is also well shown by Fig. 413, which is likewise a longitudinal section of a trachea at the point of origin of a branch. The peritracheal membrane or ectotrachea (ectr) is composed of large granulated nuclei; and within are the more transparent endotracheal cells; at t′ are fragments of the moulted tænidia. The new tænidia are in process of development at t; at base they are seen to be granulated nuclei, with often a distinct nucleolus, each sending a long, slender, transparent, pointed process along the inside of the trachea. These unite to form the chitinous bands or spiral threads.

Internal hair-like bodies.—In the large tracheæ of Lampyris very fine chitinous bristles project free into the cavity of the tube (Gerstaecher), while according to Leydig there are similar chitinous points in the tracheæ of the Carabid beetle Procrustes. Dugardin had previously (1849) called attention to such hairs, giving a list of the insects in which he observed them. Emery figures a section of the tracheæ of Luciola, “in wendig behaart.”[^[69]] Stokes has described those of Zaitha fluminea (Fig. 414) as “internal chitinous, hair-like bodies arising from the fold of the tænidia and projecting into the lumen of the tubes.” They are hollow, their minute cavity distinctly communicating with that of the tænidium, from which they arise by an enlarged base. They end in an exceedingly fine point which is occasionally bifid or trifid. In Fig. 414, 4, several are shown attached to the wrinkles of the tracheæ near a spiracle, and at 5 is represented a transverse section of a trachea with three hairs projecting into its cavity.[^[70]]

Stokes has also described “certain minute, elliptical bodies in the tænidia, each with an internal, presumably glandular, appendage, to all appearance forming part of the tænidium from which it springs.” These are shown in Fig. 414, at 1, 3, and, more in detail, at 6; those at 7, whose thickness is about 1
8000 of an inch, appear as collections of exceedingly minute, rounded apertures in a cushion-like mass. Although not commonly occurring on the tracheal membrane between the tænidia, they may be found there, as at 4.

f. The mechanism of respiration and the respiratory movements of insects

By holding a locust in the hand one may observe the ordinary mode of breathing in insects. During this act the portion of the side of the body between the stigmata and the pleurum contracts and expands; the contraction of this region causes the spiracles to open. The general movement is caused by the sternal moving much more decidedly than the tergal portion of the abdomen. When the pleural portion of the abdomen is forced out, the soft pleural membranous region under the fore and hind wings contracts, as does the tympanum, or ear, and the membranous portions at the base of the hind legs. When the tergum or dorsal portion of the abdomen falls, and the pleurum contracts, the spiracles open; their opening is nearly but not always exactly coördinated with the contractions of the pleurum, but as a rule they are. There were 65 contractions in a minute in a locust which had been held between the fingers about ten minutes. It was noticed that when the abdomen expanded, the air-sacs in the first abdominal ring contracted.

For expanding the abdomen no special muscles are required, since it expands by the elasticity of the parts. For contracting its walls there are two sets of muscles, viz., special vertical expiratory muscles serving to compress or flatten the abdomen (Figs. 415–418), and other muscles which draw together or telescope the segments.

It was formerly supposed that when the abdomen contracted the air was expelled from the body and the tracheæ emptied; that, when the abdomen again expanded by its own elasticity, the air-tubes were refilled, and that no other mechanism was needed. But Landois insisted that this was not enough; as Miall and Denny state: “Air must be forced into the furthest recesses of the tracheal system, where the exchange of oxygen and carbonic acid is effected more readily than in tubes lined by a dense intima. But in these fine and intricate passages the resistance to the passage of air is considerable, and the renewal of the air could, to all appearance, hardly be effected at all if the inlets remained open. Landois accordingly searched for some means of closing the outlets, and found an elastic ring or spiral, which surrounds the tracheal tube within the spiracle.” By means of the occlusor muscle this ring compresses the tube, “like a spring clip upon a flexible gas-pipe.” “When the muscle contracts, the passage is closed, and the abdominal muscles can then, it is supposed, bring any needful pressure to bear upon the tracheal tubes, much in the same way as with ourselves, when we close the mouth and nostrils, and then, by forcible contraction of the diaphragm and abdominal walls, distend the cheeks or pharynx.”

Thus an important point in the respiration of tracheate animals, whether insects, myriopods, or arachnids, is, as Landois claimed, the closure of the spiracles, in order that pressure may be brought upon the air in the tubes, so that it may pass onward into the finest terminations.

The injection of air by muscular pressure into a system of very fine tubes may, as Miall and Denny remark, appear extremely difficult or even impossible. Graham (Researches, p. 44) applies the law of diffusion of gases to explain the respiration of insects, but until physical experiments have been made, we may, with Miall and Denny, “be satisfied that an appreciable quantity of air may be made by muscular pressure to flow along even the finer air-passages of an insect.”

As to the respiratory movements of insects, Plateau is the principal authority, and the following account of the process is taken from his elaborate memoir, and from the statements afterwards contributed by him to Miall and Denny’s “The Cockroach.”

Although many observers have superficially described the respiratory movements of various insects, Rathke was the first one to state precise views as to the mechanism of respiration. His posthumous work, treating of the respiratory movements of the movable chitinous plates of the abdomen, and of the respiratory muscles characteristic of all the principal groups, filled an important blank in our knowledge. But, notwithstanding the skill displayed in this research, many questions still remain unanswered which require more exact methods than mere observations with the naked eye or the simple lens.

Plateau, who was followed a year later by Langendorff, conceived the idea of studying, by such graphic methods as are now familiar, the respiratory movements of perfect insects.

“He has made use of two modes of investigation. The first, or graphic method, in the strict sense of the term, consisted in recording, upon a revolving cylinder of smoked paper, the respiratory movements, transmitted by means of very light levers of Bristol board attached to any part of the insect’s exoskeleton. Unfortunately, this plan is only applicable to insects of more than average size. A second method, that of projection, consisted in introducing the insect, carried upon a small support, into a large magic lantern fitted with a good petroleum lamp. When the amplification does not exceed 12 diameters, a sharp profile may be obtained, upon which the actual displacements may be measured, true to the fraction of a millimetre. Placing a sheet of white paper upon the lantern screen, the outlines of the profile are carefully traced in pencil so as to give two superposed figures, representing the phases of inspiration and expiration respectively. By altering the position of the insect so as to obtain profiles of transverse sections, or of the different parts of the body, and, further, by gluing very small paper slips to parts whose movements are hard to observe, the successive positions of the slips being then drawn, complete information is at last obtained of every detail of the respiratory movements; nothing is lost.”

“This method, similar to that employed by the English physiologist, Hutchinson,[^[71]] is valuable, because it enables us, with a little practice, to investigate readily the respiratory movements of very small arthropods, such as flies or lady-birds. It has this advantage over all others, that it leaves no room for errors of interpretation.”

“Not satisfied with mere observation by such means as these, of the respiratory movements of insects, the writer has also studied the muscles concerned, and, in common with other physiologists (Faivre, Barlow, Luchsinger, Dönhoff, and Langendorff), has examined the action of the various nervous centres upon the respiratory organs. The result at which he has arrived may be summarized as follows:—

Fig. 415.—Muscles of right half of the abdomen of Forficula auricularia: A, a, longitudinal tergal and sternal muscles; D, E, oblique muscles; a (in upper figure) vertical expirator muscles.

“1. There is no close relation between the character of the respiratory movements of an insect and its systematic position. Respiratory movements are similar only when the arrangement of the abdominal segments, and especially when the disposition of the attached muscles, are almost identical. Thus, for example, the respiratory movements of the cockroach are different from those of other Orthoptera, resembling those of the heteropterous Hemiptera. Those of the Trichoptera are like those of the aculeate Hymenoptera, while the Locustidæ ally themselves in respect to these movements with the Neuroptera and Lepidoptera.

“2. The respiratory movements of insects, when at rest, are localized in the abdomen. As graphically stated by Graber, in insects the chest is placed at the hinder end of the body. If thoracic respiratory movements exist, they do not depend on the action of special muscles.

“3. In most cases the thoracic segments do not share in the respiratory movements of an insect at rest. The respiratory displacements of the posterior segments of the thorax are, however, less rare than Rathke believed. Plateau has observed them in certain Coleoptera (Staphylinus, Chlorophanus, Corymbites), and they are more feebly manifested in Hydrophilus, Carabus, and Tenebrio. Among the singular exceptions to this rule is the cockroach (Periplaneta orientalis), in which the terga of the meso- and metathoracic segments perform movements exactly opposite in direction to those of the abdomen (Fig. 419).

Fig. 416.—Muscles of the left half of abdomen of Staphylinus olens; A, B, longitudinal dorsal muscles; D, E, oblique fascia; a, longitudinal sternal muscles; d, respiratory muscles (vertical expirators).

“4. Leaving out of account all details and all exceptions, the respiratory movements of insects may be said to consist of the alternate contraction and recovery of the figure of the abdomen in two dimensions, viz. vertical and transverse. During expiration both diameters are reduced, while during inspiration they revert to their previous amounts. The transverse expiratory contraction is often slight, and may be imperceptible. On the other hand, the vertical expiratory contraction is never absent, and usually marked. In the cockroach (P. orientalis) it amounts to one-eighth of the depth of the abdomen (between segments 2 and 3); in Eristalis tenax to one-ninth (at the 2d segment).

“5. Three principal types of respiratory mechanism occur in insects, and these admit of further subdivision:

“a. Sterna usually short and very convex, yielding but little. Terga mobile, rising and sinking appreciably. To this class belong all Coleoptera, heteropterous Hemiptera, and Blattina (Fig. 420).

“In the cockroach (Periplaneta), the sterna are slightly raised during expiration (Fig. 421).

“b. Terga well developed, overlapping the sterna on the sides of the body, and usually concealing the pleural membrane, which forms a sunken fold. The terga and sterna approach and recede alternately, the sterna being almost always the more mobile. To this type belong Odonata, Diptera, aculeate Hymenoptera, and acrydian Orthoptera (Fig. 422).

Fig. 417.-Muscles of right half of abdomen of Phryganea striata, ♀: A, B, longitudinal dorsal muscles; a, b, longitudinal sternal muscles; D, e, oblique muscles; 1, 2, inspirator muscles.

“c. The pleural membrane, connecting the terga with the sterna, is well developed and exposed on the sides of the body. The terga and sterna approach and recede alternately, while the pleural zone simultaneously becomes depressed, or returns to its original figure. To this type, Plateau assigns the Locustidæ, Lepidoptera, and the true Neuroptera (excluding Trichoptera) (Fig. 423).

Fig. 418.—Muscles of left half of abdomen of Melolontha, ♀: A, B, longitudinal muscles (prétracteurs of Straus); a, a, true respiratory muscles (expirators).—This and Figs. 415–417, after Plateau.

“6. Contrary to the opinion once general, changes in length of the abdomen, involving protrusion of the segments and subsequent retraction, are rare in the normal respiration of insects. Such longitudinal movements extend throughout one entire group only, viz. the aculeate Hymenoptera. Isolated examples occur, however, in other zoölogical groups.

“7. Among insects, such as large beetles, Locustidæ, dragon-flies, etc., sufficiently powerful to give good graphic tracings, it can be shown that the inspiratory movement is slower than the expiratory, and that the latter is often sudden.

Fig. 419.—Profile of trunk of cockroach (P. orientalis). The black surface represents the expiratory contour, while the inspiratory is indicated by a thin line. The arrows show the direction of the expiratory movement: Ms. th, mesothorax; Mt. th, metathorax. Reduced from a magic-lantern projection.—After Plateau.

“8. In most insects, contrary to what obtains in mammals, only the expiratory movement is active; inspiration is passive, and effected by the elasticity of the body-wall.

“9. Most insects possess expiratory muscles only. Certain Diptera (Calliphora vomitoria and Eristalis tenax) afford the simplest arrangement of the expiratory muscles. In these types, they form a muscular sheet of vertical fibres, connecting the terga with the sterna, and underlying the soft, elastic membrane which unites the hard parts of the somites. One of the most frequent complications arises by the differentiations of this sheet of vertical fibres into distinct muscles, repeated in every segment, and becoming more and more separated as the sterna increase in length. Special inspiratory muscles occur in Hymenoptera, Acridiidæ, and Trichoptera.

“10. The abdominal, respiratory movements of insects are wholly reflex. Like other physiologists who have examined this side of the question, Plateau finds that the respiratory movements persist in a decapitated insect, as also after destruction of the cerebral ganglia or œsophageal connectives; further, that in insects whose nervous system is not highly concentrated (e.g. Acridiidæ and dragon-flies), the respiratory movements persist in the completely detached abdomen; while all external influences which promote an increased respiratory activity in the uninjured animal, have precisely the same action upon insects in which the anterior, nervous centres have been removed, upon the detached abdomen, and even upon isolated sections of the abdomen.

“The view formerly advocated by Faivre, that the metathoracic ganglia play the part of special, respiratory centres, must be entirely abandoned. All carefully performed experiments on the nervous system of Arthropoda have shown that each ganglion of the ventral chain is a motor centre, and, in insects, a respiratory centre, for the somite to which it belongs. This is what Barlow calls the ‘self-sufficiency’ of the ganglia.” (Miall and Denny.)

Fig. 420.—Transverse section of abdomen of a lamellicorn beetle. The position of the terga and sterna after an inspiration is indicated by the thick line; the dotted line shows their position after an expiration; and the arrow marks the direction of the expiratory movement.

Fig. 421.—Cross-section of abdomen of cockroach.

Fig. 422.—Cross-section of abdomen of bee (Bombus).

Fig. 423.—Cross-section of abdomen of Sphinx.—This and Figs. 420–422 after Plateau.

Plateau has made similar observations upon the respiration of spiders and scorpions; but, to his great surprise, he was unable, either by direct observation, or by the graphic method, or by projection, to discover the slightest respiratory movement of the exterior of the body. This can only be explained by supposing that inspiration and expiration in pulmonate Arachnida are “intrapulmonary,” and affect only the proper, respiratory organs. The fact is less surprising because of the wide zoölogical separation between Arachnida and insects.

g. The air-sacs

In flying insects the tracheæ are in certain parts of the body enlarged into sacs of various sizes. These air-sacs were first observed by Swammerdam in a beetle (Geotrupes) and afterwards by Sir John Hunter in the bee, Sprengel subsequently discovering them in other insects. Those of the cockroach were described and illustrated in a very elaborate and detailed way by Straus-Dürckheim (Figs. 424 and 425). These vesicles are without tænidia. In the locust (M. femur-rubrum) there is a pair of very large vesicles in the prothorax (Fig. 396). The five pairs of large abdominal air-sacs arise, independently of the main tracheæ, directly from branches originating from the spiracles. All these large sacs are superficial, lying directly beneath the hypodermis, while the smaller ones are buried among the muscles. We have detected 53 of these vesicles in the head.

In the honey-bee (Fig. 426) and humble bee (Fig. 427) as well as the flies there are two enormous air-sacs at the base of the abdomen. In larval and wingless insects these sacs are entirely absent.

Fig. 424.—Thorax and abdomen of the cockchafer (Melolontha vulgaris), showing the tracheæ and air-sacs.—This and Fig. 425 after Straus-Dürckheim.

The use of the air-sacs.—It was supposed by Hunter as well as by Newport, and the view has been generally held, that the use of these sacs is to lighten the weight, i.e. lessen the specific gravity of the body during flight. It has, however, been suggested to us by A. A. Packard that this view from the standpoint of physics is incorrect. It is evident that the wings have to support just as much weight when the insect is flying, whether the tracheæ and vesicles are filled with air or not, the body of the insect during flight not being lightened by the air in the sacs. The use of these numerous sacs, some of them very spacious, is to afford a greater supply of air or oxygen than that contained in the air-tubes alone, and thus to afford a greater breathing capacity. The sacs are largest in dragon-flies, moths, flies, and bees, which are swift of flight. When we compare the active movements of these insects on the wing with those of a caterpillar or maggot, it will be seen that the far greater muscular exertions of the volant insect create a demand for a sudden and abundant supply of air to correspond to the increased rapidity of respiration; and the enlargements of the air-tubes, rapidly filled with air at each inspiration, render it possible to supply the demand.

Fig. 425.—Head of Melolontha vulgaris, showing the numerous air-sacs, represented only on the left side, front view.

Fig. 426.—Tracheal, nervous, and digestive systems of the honey-bee (the tracheal system on the right side only partially drawn): tb, the large vesicles in the abdomen; st, stigmata; hm, honey stomach; cm, chyle stomach; vm, urinary tubes; rd, rectal glands; ed, rectum; a, antenna; an, eye; b₁-b₃, legs.—After Leuckart, from Lang.

The case is thus seen to be very different from that of those fishes which, having a swimming-bladder, can in the water change the specific gravity of their bodies. The case of insects is almost exactly paralleled by that of birds, where, as stated by Wiedersheim, the air-sacs appear to form integral parts of the respiratory apparatus: “a greater amount of air can by their means pass in and out during inspiration and expiration, especially through the larger bronchi, and consequently there is less necessity for the expansion of the lung parenchyma.” In other words, the supply of air in these sacs, as in insects, increases the breathing capacity of the bird during flight. Wiedersheim’s retention of the old idea that the specific gravity of the body is lessened (p. 262) seems, however, to be incorrect, as the weight of the bird’s body is not diminished by the air contained in the sacs.

h. The closed or partly closed tracheal system

Fig. 427.—The lateral and lower series of sacs of Bombus terrestris, ♂: a, c, longitudinal tracheæ;, connected by b, and dilated at f, and again in the succeeding segments; i, k, funnel-shaped dilatations passing over the dorsal surface of the abdomen and anastomosing (g) with their fellows opposite; at l, communicating directly by a large branch.—After Newport.

There are two chief morphological tracheal systems: 1. The open or normal and primitive (holopneustic) type, and 2. The closed, or secondary and adaptive, i.e. apneustic, type. The open system is characterized by the presence of the stigmata. Through them the air directly enters into the tracheal tubes, whose delicate walls allow the exchange of gases in the blood. This type occurs in all sexually mature individuals, and also in the greater number of larvæ.

The closed or apneustic tracheal system is distinguished either by the want of stigmata, or, if present, they are not open, and do not function, so that the tracheæ cannot communicate with the air. In such cases the direct oxygenation of the blood is effected through the delicate integument, especially over the surface of the body in general, or in certain specialized places where the gill-like expansions of the skin are rich in tracheæ; such outgrowths, generally tubular or leaf-like, are called by Palmén tracheal gills.

This closed form of the tracheal system only occurs in the larval stage of aquatic or parasitic insects, as in the Plectoptera (Ephemeridæ), Perlidæ, Odonata, and Trichoptera, besides single genera of other orders, i.e. among Coleoptera, Gyrinus, Pelobius, Cnemidotus, and the young larva of Elmis; in the aquatic caterpillar of Paraponyx; in certain Diptera (Corethra, Chironomus, etc.), and some of the parasitic Hymenoptera (Microgaster).

Palmén has discovered that in the nymphs of Ephemeridæ, Perlidæ, Odonata, and the larvæ of most Trichoptera the tracheal branch (stigmatal branch) sent from the longitudinal trachea to where the thoracic stigmata would be situated if present, or where their vestiges only exist, are aborted, becoming simple solid cords not filled with air (Fig. 436, vf, and 447, f, funiculus or stigmatic cord). In the imago, however, they resume their function, connecting with the open functional stigmata. In Corethra, in its earliest stages, the entire tracheal system is, like the stigmatic branch, a system of solid cords and empty of air. (Palmén.)

Embryology shows that these stigmatal branches are well developed, and are formed at the same time as the stigmata. It was also shown by Dewitz, in a posthumous paper (1890), that in the young larval stage of the Odonata and Ephemeridæ the tracheal system is at first an open one, and in some of the families (Libellulidæ, Agrionidæ, and Ephemeridæ) thoracic stigmata are seen at a very early stage. From numerous experiments Dewitz concludes that in the young stages of Odonata and Ephemeridæ there is an open tracheal system; certainly in very young nymphs the thoracic spiracles allow the air to pass out. Fully grown nymphs of Æschnidæ, Libellulidæ, and Agrionidæ are capable not only of forcing the air out, but also, like the perfect insect, of inhaling it. Moreover, he proved that the gills of Ephemeridæ and Agrionidæ are not indispensable for the maintenance of life, as the insects can live without them, breathing either through the skin or by the rectum, or in both ways. It would seem that while in freshly hatched or very young larvæ of aquatic insects of different orders the skin is so delicate as to allow of dermal respiration, in after life, when the skin becomes thicker and denser, these expansions (gills), provided with a very thin and delicate skin, of a necessity grow out from the walls of the body.

It thus appears that the closure and total or partial abolition of the stigmata are in adaptation to aquatic life, and that such insects have descended from terrestrial air-breathing winged forms. This is an important argument against the view that the wings are modified tracheal gills.

In this connection may be noticed the closure of the 2d and 3d thoracic stigmata in holopneustic insects. We have found on laying open the body of a Sphinx larva that a large number of tracheal branches are seen to arise from the prothoracic and from the first pair of abdominal stigmata. Now between these points there are no spiracles or any external signs of them, there being in Lepidoptera no mesothoracic or metathoracic spiracles. Yet the main lateral trachea between the prothoracic and first abdominal segments deviates from its course and bends down to send off a small shrivelled stigmatal branch or cord to a place where, did a spiracle exist, we should look for it. In the larva of Platysamia cecropia, a similar vestigial stigmata branch is present.

In the larva of Corydalus, also, a trachea as large as the main longitudinal one takes its origin and passes directly under the main trachea. Now both tracheæ send a stigmatal branch opposite to where the mesothoracic stigma should be, if present, i.e. on the hind edge of the segment.

Verson, moreover, has found in the freshly hatched silkworm vestiges of meso- and metathoracic stigmata, each consisting of a circle of high hypodermal cells radially arranged around a common centre. The stigmatal branch is long, but shrivelled; its peritoneum is widened out into several berry-like saccules filled with cell-elements. In profile these rudimentary stigmata appear as a series of high hypodermal cells, which form the basis of a short blind tube.

Lydia M. Hart del.
Plate I.—Examples of metapneustic insects: 1, Bittacomorpha clavipes, larva; 1 a, false foot; 1 b, its pupa; 2, Limnophila luteipennis; 2 a, end of larva; 2 b, its pupa; 3, end of larva of Tipula eluta.—After C. A. Hart.

After the second moult there begins a peculiar transformation of the rudimentary stigmata. The stigmatal branch connected with them sends off at various points thick tufts of capillary tracheæ which press against the base of the blind tube. Gradually lengthening, they form a fold which continues to increase in length. The numerous tufts of tracheal capillaries extend beyond the inner surface of the two layers of which the developing wing consists, the berry-like saccules are drawn into the wing and converted into more or less thick tubes, which finally form the “veins.” It is clear, therefore, says Verson, as Landois claimed, that the wings of Lepidoptera must be regarded as in the fullest sense organs of respiration. (Zool. Anz., 1890, p. 116.)

The number of pairs of stigmata varies, especially in maggots or larval Diptera, in adaptation to their varied modes of life. The larvæ of most flies (Muscidæ) have a pair of peculiarly shaped processes on the prothoracic segment bearing spiracular openings, and two anal spiracles, while in Ctenophora atrata L. only the anal pair are present. In the rat-tailed maggots (Eristalis) the long caudal process ends in two stigmata forming a respiratory tube, which can be thrust out of the water for the reception of air. In the larval mosquito (Fig. 433) and its ally, Mochlonyx, a short thick dorsal tube arises from the penultimate segment of the body, in which the two main tracheæ end, opening outward by a single spiracular aperture. Other dipterous larvæ (Simulium, Tanypus, and Ceratopogon) possess no spiracles, the tracheal system being a closed one.

The larvæ of most water beetles (Dyticidæ, Hydrophilidæ) possess but two spiracles, which, as in maggots, are situated at the end of the body. The aquatic larva of Amphizoa, according to Hubbard, breathes much as in the Dyticidæ, by means of two large valvular spiracles placed close together at the end of the body; “closed or rudimentary stigmata also occur on the mesothorax and on abdominal segments one to seven inclusive.”

Hubbard adds: “The larva of Pelobius is wholly aquatic and breathes by branchiæ, but the obsolete stigmata are indicated precisely as in Amphizoa, with the exception of the last pair, which in Amphizoa are open spiracles, but in Pelobius are suppressed; the terminal eight segments being prolonged in a swimming stylet.”

From a review of the distribution of spiracles, and their atrophy, partial or total, it will be seen that there are intermediate stages between the open (holopneustic) and closed (apneustic) systems. These, following Schiner, Brauer, and Palmén, may be defined thus:

1. Metapneustic type.—The larvæ possess only a single pair of open stigmata situated at the end of the body. (The dipterous Eristalis, Tipula, Culex, Ptychoptera, Bittacomorpha (Plate I.) with certain Tachinidæ, and in Coleoptera, the larvæ of Dyticus, and allies of Hydrophilus and Cyphon.)

2. Propneustic type.—The pupæ of Corethra, Culex, etc., in which only the most anterior pair of spiracles are open.

Fig. 428.—Visceral tracheal system of the nymph of Æschna maculatissima: o, œsophagus; E, stomach; M, urinary tubes; R, rectum; A, anus; tv, visceral tracheal trunks; td, dorsal trunks.—After Oustalet.

3. Amphipneustic type.—Larvæ with a pair of open spiracles situated at each end of the body, the intermediate spiracles being closed. (Most dipterous larvæ, Musca, after the first moult, Œstridæ, Asilidæ, and Syrphus.)

Fig. 429.—Branchial tuft of nymph of Æschna.

4. Peripneustic type; with prothoracic and abdominal spiracles, the mesothoracic pair atrophied or closed. (The larvæ of Neuroptera, Mecoptera, Trichoptera, Lepidoptera, of most Coleoptera,[^[72]] of most Diptera, and of most of the Hymenoptera.[^[73]])

Fig. 430.—Part of three rows of respiratory folds from cuticular living rectum of Æschna. The shaded parts are abundantly supplied with tracheal tubes. The leaflets appear to be connected with a central trachea, but this is not really the case.—After Miall.

These differences in the number of functional spiracles are in direct relation with the surroundings of the insects, the physical conditions of existence evidently determining the position of the active functional open spiracles and the closure of those useless to the organism.

i. The rectal tracheal gills, and rectal respiration of larval Odonata and other insects

The remarkable mode of respiration by tracheal gills situated within the intestine of the nymphs of dragon-flies was first described by Swammerdam and afterwards by Réaumur. The most complete and best illustrated modern account is that of Oustalet. In these insects the large rectum is lined with six double longitudinal ridges, in Æschna bearing numerous delicate tubes or papillæ, each of which contains very numerous (by estimate 24,000) tracheal branches (Fig. 431); while in Libellula the gills are lamellate (Fig. 432). The tracheæ arise both from the main dorsal and visceral longitudinal trunks, which give rise to secondary branches passing into the walls of the rectum and sending into the branchial papillæ fine twigs, which, extending to the distal end of the papilla or lamella, recurve and anastomose with the efferent twigs.

Fig. 431.—A small part of one leaflet, highly magnified, showing many fine tracheal branches. The portion shown is marked by a small circle in Fig. 430, lower left-hand corner.—After Miall.

Fig. 432.—Leaves, mh, from a lamellate tracheal gill of Libellula: t, trachea.—This and Fig. 429, after Oustalet.

The anal opening is externally protected by the suranal and lateral triangular chitinous plates, three to five in all. When open, the water passes into the rectum and bathes the rectal gills, where it may be forcibly expelled as if shot out from a syringe, thus propelling the insect forward. In Libellula the anus affords direct access to the intestinal cavity, but in Æschna Oustalet describes “a sort of vestibule separated from the rectum by a circular valvule.” He also states that the inspiration and the repulsion of water is produced at irregular intervals, and rather by the movements of the dorsal and sternal arches of the abdomen than by the contractions of the rectum, since the walls of this organ are less muscular than is supposed.

Fig. 433.—Larva of a mosquito (Culex nemorosus) of middle age, seen from above, the tracheal system omitted: at, antennæ; ab, their middle joint; eg, elastic articular membrane; atm, antennal muscle; atn, antennal nerve; zau, compound; eau, simple eye; os, brain; oex, extensor; ofl, flexor of labrum; ha, neck; œ, œsophagus; spd, salivary gland; mau, cœca; ch, chyle stomach; di, contents of intestine; mg, urinary tubes; dd, ileum; ed, rectum; a, anus; s, sipho; z″, its bristles; kb, tracheal gills: k₁, k₂, k₃, closing lobes of the sipho; kn, basal tubercle of tactile hair; g, its ganglion cell; th, tactile hair of the siphon valve.

The nymph of Calopteryx (and probably of all the group Calopteryginæ) possesses rectal gills besides external caudal tracheal gills. There are three double rectal longitudinal folds or ridges, interpenetrated by tracheal twigs. (Dufour, denied by Poletaiew, but confirmed by Hagen.)

Dewitz claims that the caudal gills of the Agrionidæ are not their sole means of respiration, since he cut off the caudal tracheal gills of an Agrionid nymph, which continued to live for a week. Hence he thinks that there may be a rectal respiration, since under the microscope he saw a stream of water pass in and out of the end of the intestine.

Dewitz’ experiments prove that in young Ephemerids there may be besides branchial, both rectal and skin respiration. He saw under the microscope the anus for a while opened and then closed, causing the rectum to move; powdered carmine mixed with water was drawn into and then expelled from the rectum. There was, however, no enlargement and contraction of the abdomen as in the rectal respiration of Æschna. (Zool. Anz. 1890, p. 500.)

Fig. 434.—End of the body of the same larva as in Fig. 431, seen from the side, the branches of the main tracheæ (htr) omitted: kbl, excrementitial pellet in rectum; kb, tracheal gills; b, funnel of the closing apparatus; hz, hollow tooth of the closing apparatus; k₁, k₂, k₃, siphonal lobes; th, tactile hair; as, chitinous plate; str, rudder; l, its thickened edge; sch, its shank; z′, z″, bristles.—This and Fig. 433, after Raschke.

Eaton states that there is a rectal respiration in the nymphs of may-flies, and Palmén observed in young larvæ of Bætis and Cloëon that the rectum took in “by gulps” water colored by carmine and expelled the whole of it at once, in order to fill it again in the same way. “This rectal respiration therefore corresponds to that of Libellulid larvæ.”

Fig. 435.—Thorax and anterior abdominal segments of the nymph of a may-fly (Cloëon dimidiatum) with tracheal gills (tk₁, tk₂, tk₃) and the rudiments of the fore wings (VF) and hind wing (HF): tl, tracheal longitudinal trunks.—After Graber, from Lang.

Fig. 436.—Gills on the middle abdominal segments of larva of Bætis binoculatus: trl, longitudinal tracheal trunks; vf, stigmatic cord; ktr, gill-tracheæ; trk, tracheal gills.—After Palmén, from Lang.

Besides breathing by spiracles, by tracheal gills, as well as through the integument, the larva of Culex has been observed by Raschke to have a rectal respiration. At the anterior end of the rectum arises a countless number of fine tracheæ, which pass through the walls and, subdividing, end in numberless very fine twigs in the papilla-like folds situated within the rectum. The supply of tracheal twigs is greatest where the papillæ are largest. (Figs. 433, 434.)

j. Tracheal gills of the larvæ of insects

In many aquatic insects respiration is carried on by tracheal gills. These are delicate, hollow, leaf-like or tubular outgrowths of the integument usually attached to the sides or end of the hind-body, and containing a trachea which usually sends off numerous minute branches, so that the exchange of gases readily takes place in them.

Fig. 437.—A, nymph of Ephemerella ignita, with gills of left side removed; g, gills. B, nymph of Tricorythrus (sp), with gill-cover of right side removed; gc, gill-cover; g, g′, gills.—After Vayssière.

Fig. 438.—Left maxilla of Jolia weselii, with the cephalic tracheal gill (h) inserted at the base on the under side.—After Vayssière.

Palmén has shown that these tracheal gills, as he calls them, are not developed on the same segments as the stigmata, and that the two structures have no genetic connection with each other. It is evident that these gills are secondary, adaptive organs.

In some cases (see p. 475) the tracheæ are wanting, but as such gills are filled with blood, the air contained in the water must pass in through their delicate walls.

In the Plectoptera (Ephemeridæ) the tracheal gills are either foliaceous or filamentous; when foliaceous they form simple or double leaves, with or without branches, or with a fringe of tubules, or under the leaf-like cover-bearing tufts of filaments. They are situated on the (usually) basal seven abdominal segments, at their hinder edge (Figs. 435, 436). In Oligoneuria and Jolia a pair occurs on the under side of the head, attached to the maxillæ, while in Jolia there is a pair on the under side of the first thoracic segment at the insertion of each of the legs. In certain genera (Heptagenia, Oligoneuria, and Jolia), they are in the form of a flat cover, under which lies a tuft of respiratory tubes, or (Ephemerella) a small bifid cluster of very delicate leaves (Fig. 437, A). In Cœnis and Tricorythus the tracheal gills of the second pair are modified to form plates covering all the succeeding pairs, those of the first pair being nearly atrophied and well-nigh functionless. (Fig. 437, B.)

Fig. 439.—Inner side of a gill-cover of the first pair, of Ephemerella, with the tracheal gills.—After Vayssière.

Fig. 440.—Nymph of Bætisca: III, section of abdomen; a, gills; b, flap; 1–9, abdominal segments.—After Walsh.

Fig. 441.—Nymph of Prosopistoma punctifrons: o, upper orifice of the respiratory chamber.—After Vayssière.

Fig. 442.—Filamentous tracheal gill and part of a trachea of Pteronarcys.—After Newport from Sharp.

Finally, in the highly modified forms Bætisca and Prosopistoma the tracheal gills are entirely concealed and protected by mesothoracic projections so as to form a true respiratory chamber, to which the water has access either by an opening behind, as in Bætisca, or by three openings, two ventral and one dorsal (Fig. 441), as in Prosopistoma.

The slender cylindrical tracheal gills of Heptagenia in the third or fourth nymphal stage are 2–jointed, and the first abdominal pair in Cænis are said by Palmén to be finger-shaped and 2–jointed. In Polymitarcys virgo the gills do not appear until the eighth or tenth day after hatching.

Dewitz found that young nymphs of Ephemerids will well endure the amputation of their gills, while fully grown ones die. Amputation of the lateral gills hastens ecdysis. After the change of skin, the gills are smaller than before, and at first contain no tracheæ, but in a few weeks they develop as completely as in normal individuals. The caudal gills were also renewed.

Fig. 443.—A, larva of Sisyra, enlarged. B, one of the hinder gills, with its tracheæ.—After Westwood, from Sharp. C, a gill, showing the branched tracheæ.—After Grube.

In the nymphs of Perlidæ the tracheal gills are usually present, and are either foliaceous (Nemoura) or more commonly filamentous in shape (Fig. 442). They are situated either on the prosternum (Nemoura and Pteronarcys), or on each side of the thorax, or on the sides of the abdomen, or are restricted to a tuft on each side of the anus at the base of the caudal stylets (Pteronarcys and Perla). Unlike the Ephemeridæ the gills persist in certain genera throughout life.

The larvæ of the aquatic Neuroptera, Sisyra, Sialis, and Corydalus possess lateral pointed bristle-like tracheal gills, which in Sisyra are 2–jointed; those of Sialis are, in the living larva, curved upwards and backwards (Fig. 444). Corydalus is also provided with a ventral tuft of delicate filamentous gills, which, however, according to Riley, do not appear until after the first moult.

While the nymphs of Agrionidæ (which have rectal gills) respire chiefly by the large caudal foliaceous gills (Fig. 445), there are, according to Hagen, two genera of the Calopteryginæ (Euphæa, Fig. 445, and Anisopleura) whose nymphs possess seven pairs of external lateral tracheal gills, in shape like those of Sialis, besides three caudal and three rectal tracheal gills.[^[74]]

Fig. 444.—Larva of Sialis lutarius.—After Miall.

Fig. 445.—Caudal tracheal gill of nymph of Agrion.

Hagen has also detected in the under side of the 5th abdominal segment of Epitheca and Libellula a pair of sacs of the shape of a Phrygian bonnet, each of which contains a smaller sac lined with epithelium,—as in Æschna they occur in the 5th and 6th, and in Gomphus in the 4th, 5th, and 6th segments. This serial arrangement appears to confirm Hagen’s suggestion that they are survivals of abdominal gills, which in Euphæa are completely evaginated.

Fig. 446.—Nymph of Euphæa, showing the lateral gills: a, one enlarged.—Folsom del.

In the Trichoptera, all of which, except Enoicyla, are apneustic, and most of which have tracheal gills, the latter are filamentous, and arise either from the dorsal and ventral sides of the abdominal segment, or they grow out from the sides; while in certain genera (Neuronia, Phryganea, etc.) the gills are represented by conical hooks on the sides of the 1st abdominal segment, which are evidently respiratory, as they contain numerous tracheæ. The tracheal gills are either single or more rarely form tufts (Figs. 447, 448).

In Hydropsyche (Fig. 448) the tracheal gills persist throughout life, while in other genera they only last through the pupal stage. When first hatched, the larva of Phryganea lacks gills. The larvæ of most of the Hydropsychidæ, Rhyacophilidæ, and Hydroptilidæ have no gills, though they appear well developed in the pupal stage. (Klapálek.)

Fig. 447.—A, an abdominal segment of Hydropsyche, with the tracheal gills (lbr): trl, longitudinal tracheal trunk; f, stigmatal branch. B, 5th abdominal segment of pupa of the same; l, the three lateral flaps of the tergite; br¹, br², branchiæ.

Fig. 448.—Imago, abdominal segments IV to VI, with the gills at a concealed in their natural condition; at b, drawn out with the needle; at c, projecting abnormally and dried.—This and Fig. 447 after Palmén.

Fig. 449.—Larva and pupa of Paraponyx stratiolata, enlarged; s, spiracle.—After De Geer (compare Hart’s figure of P. obscularis, living in the Illinois River).

The only lepidopterous larva known to be provided with tracheal gills is that of the pyralid genus Paraponyx. Its thread-like gills, arranged in tufts of three or four, arise from a common tubercle situated on the sides of nearly all the segments. Wood-Mason describes the East Indian P. oryzalis as “covered with a perfect forest of soft and delicate white filaments,” arranged in tufts disposed in four longitudinal rows. “The stigmata of the 2d, 3d, and 4th abdominal somites only are clearly discernible.” The caterpillar crawls “free and uncovered” over the submerged leaves of the rice plant “in the very midst of the water.” In a Brazilian species of Paraponyx described as Cataclysta pyropalis, by W. Müller, the tufts are reduced to simple unbranched filaments, and the case is more complex than in the European species (Fig. 449).

Fig. 450.—Anterior end of larva of P. stratiolata, showing the head and first two thoracic segments, with their gills: A, a tuft of gills, much enlarged.—After De Geer.

Fig. 451.—Larva (1) and pupa (2a) of Paraponyx pyropalis enlarged: st, stigmata.—After W. Müller.

Of coleopterous larvæ breathing by tracheal gills there are but few. The larva of Gyrinus (Fig. 454) respires by 10 pairs of slender, hairy abdominal gills similar to those of Corydalus, and the stigmata are entirely wanting. Somewhat similar are the tracheal gills of Hydrocharis caraboides. Hydrobius has shorter setose gills, our American species having seven pairs of short setose gills. It has two spiracles at the end of the body, through which the air is taken by thrusting the body out of the water. The larvæ of two other aquatic coleopterous genera, Pelobius and Cnemidotus, also have gills; those of the former situated at the base of the coxæ, and brush-like, but containing no tracheæ, though filled with blood, while those of Cnemidotus are very long, bristle-like, jointed, and arising from the dorsal side of the thoracic and abdominal segments. The stigmata are wanting. (Schiödte.)

The larva of the dipterous genus Tanypus respires by two caudal papilliform processes, in each of which a trachea ramifies.

Fig. 452.—Freshly hatched larva of Hydrobius: t, enlarged tracheæ, the heart between them; g¹-g⁷, the seven pairs of gills. A, end of body, enlarged, showing the two terminal stigmata.—Emerton del.

Certain larvæ with both stigmata and tracheal gills are enabled either to live in or out of water or on the surface, as in the case of certain beetles (Cyphonidæ, Elmidæ, Hydrophilidæ, Fig. 452), or the larval mosquito and Psychodes (Fig. 455); also the nymphs of dragon-flies.

The larvæ of the Cyphonidæ (Helodes, Cyphon, Hydrocyphon) possess but a single pair of stigmata, situated in the penultimate abdominal segment, while at the end of the abdomen are delicate tracheal gills. The two main tracheal trunks are much swollen. When on the surface of the water the larva breathes through the stigmata situated near the end of the abdomen; when floating in the water, the larva, like that of Gyrinus, carries along at the end of its body a bubble of air. The gills are only of use, as Rolph thinks, when the insect is compelled to remain a long time under water.

The larva of our native Prionocyphon discoideus (Say) is described by Walsh as “vibrating vigorously up and down a pencil of hairs proceeding from a horizontal slit in the tail”; this pencil is composed “of three pairs of filaments, each beautifully bipectinate. I presume it is used to extract air from the water.” When the larva is at the surface the pencil of hairs touches the surface of the water, and occasionally a bubble of air is discharged from the tail. “The general habit is to crawl on decayed wood beneath the surface, occasionally swimming to the surface, probably for a fresh supply of air.” (Proc. Ent. Soc. Phil., i, p. 117.)

Fig. 453.—Larva of Psephenus, enlarged.

Fig. 454.—Larva of Gyrinus.—After Westwood.

The larvæ of the small water beetles of the family Elmidæ (Elmis, Potamophilus, Macronychus, and Psephenus) have similar habits. That of Elmis has ten dorsally situated pairs of spiracles, and on the end of the body bushy gills which are protruded at pleasure. The young larva is without spiracles, its tracheal system being closed. Macronychus and Potamophilus have similar habits. In the larva of the latter genus, which has nine pairs of spiracles, there are at the end of the body on each side three tufts of thread-like gills which are connected with the two main horizontal tracheæ, while the branches of the abdominal tracheæ are dilated into numerous (64) bladder-like sacs. The larva usually breathes through the caudal gills. When the water is low or dried up, the air is inhaled directly through the spiracles. (Kolbe.)

The larva of Psephenus lecontei, by its broad hemispherical body, is adapted to adhere to the smooth surface of rounded stones, in which situation we have found it. Although it is said by Rolph to have two pairs of spiracles, one pair on the mesothoracic and the other on the 1st abdominal segment, it probably rarely rises to the surface to breathe the air direct.

Fig. 455.—End of body of a Psychodes larva: A, end of body of a young, freshly moulted larva, side view: a, the three anal gills; b, the left air-cavity. B, older larva of the same species, with the open air-cavity seen from above. C, end of larva of another species as it goes down into the water with a bubble of air, b, between the crown of hairs of the air-cavity or tube: a, the two pairs of anal gills; b, the two main tracheæ.—After F. Müller.

It possesses five pairs of gills on the under side of the 2d to the 6th abdominal segments. Each gill has finger-shaped processes on its hinder edge, which are “from their constant motion evidently connected with respiration.” Tracheæ may be seen, according to H. J. Clark, entering the gills, and “the circulation of water among the branchiæ is kept up by the flapping of the tail-pieces.” The larva of Helichus fastigiatus is said by Leconte to be “very nearly allied, while the remotely allied Stenelmis crenatus has no external branchiæ.”[^[75]]

The larva of the mosquito also has two modes of respiration, breathing either at the surface of the water through the two spiracles situated on the projection (siphon) at the hinder end of the body which is thrust out into the air; or when at the bottom respiring by tracheal gills. The pupa also has a double mode of respiration, either taking in air at the surface by the two thoracic horns with stigmatic openings, or when submerged using its tracheal gills.

Besides its long caudal tracheal air-tubes, the larval Eristalis is said by Chun to thrust out from the anus a number (20) of short tracheal filaments which float about in the water and serve to absorb the air.

An aquatic Brazilian larva of the family Psychodidæ has been found by Fritz Müller to take down under the water a large bubble of air (Fig. 455, C), the main tracheal trunk ending each in an opening at the end of the body (A, B); besides this, while at the bottom it breathes by three digitiform tracheal gills; another species having two pairs (C, a).

Fig. 456.—Under side of body of larva of Blepharocera. showing the position of the tracheal gills: A, section of the body through a sucker, showing position of the gills. B, section of a sucker: br, gill with numerous tracheæ; gl, outlet of excretory gland; M, m, muscles.—After F. Müller.

The remarkable larvæ of the Blepharoceridæ (represented in the United States by Blepharocera capitata), which live permanently in swift streams, attached by median suckers to stones, are apneustic, and breathe solely by leaf-like tracheal gills (Fig. 456, br) attached to the under side of the second to sixth abdominal segments. Those of the European Liponeura are said by Wierzejski to be branched, tree-like. Also immediately in front of the anus and behind the last sucker are four membranous sacs provided with tracheæ, but which are not capable of being withdrawn. These are said by Müller to be the same as what Dewitz states to serve as gills, and by Wierzejski they are homologized with the four anal gills of Chironomus.

The double mode of respiration in the larva of the horse bot-fly has been described by Scheiber. On the hinder end of the body are the stigmatic plates, which contain two lateral gill-plates and the middle stigmatal leaf. Besides this there is a pair of slightly developed prothoracic spiracles. The embryo and also freshly hatched larva of Gastrophilus equi do not possess these gill-plates, but on the end of the body are, according to Joli, two long thread-like gills. The freshly hatched larva of the allied Cephenomyia rufibarbis bears two caudal projections. (Kolbe.) As in shrimps and other Crustacea the gills are kept in constant motion, the water being driven over them by the rapid movements of the telson, so in the larval may-flies, and in the case-worm (Macronema), the gills move more or less rapidly. In case-worms as well as larval Perlidæ, Sialidæ, Paraponyx, and Hydrophilidæ the abdominal region is constantly moved to promote respiration. (Kolbe.)

Blood-gills.—Fritz Müller describes in trichopterous larvæ certain delicate anal tubular processes into which the blood flows, and which do not as a rule contain tracheæ, though occasionally very fine tracheal branches. Müller compares them with the gills of crabs and of shrimps. They are eversible finger-like tubules. They are used when the tracheal gills are temporarily not available. Their number varies even in the same genus. There are six in certain Rhyacophilidæ; five in different Hydropsychidæ; in Macronema there are four, and they are green when filled with the green blood of that insect, the tracheal gills being whitish. In the freshly hatched larva, while the tracheal gills are present, no anal blood-gills are visible. Similar blood-gills also occur in the pupæ of certain caddis-flies. (Pictet.)

Similar anal gills filled with blood occur in the larvæ of the fireflies (Lampyris, etc.), and perhaps, Kolbe thinks, serve for respiration, though other authors believe them to be adhesive organs.

The larva of Pelobius has true blood-gills. (Schiödte. See p. 461.)

The eversible ventral segmental sacs of Scolopendrella, Campodea, and Machilis, as well as the ventral tube (collophore) of Podura, Smynthurus, etc., may, as Oudemans and Haase have suggested, serve a respiratory purpose, though they lack tracheæ, and differ from blood-gills in containing no gases; yet the blood is forced into them, causing their eversion. Oudemans observed that Machilis everted its sacs when the vessel in which it was put was filled with warm, damp air. The sacs are only thrust out when the creature is completely at rest.

Structures referable to blood-gills also occur temporarily in the embryo of Orthoptera; Rathke observed them in the mole-cricket; Ayres observed them in Œcanthus niveus, where they form two stalked broad oval appendages on the first abdominal appendages, which he regarded as gills. Patten observed them in Phyllodromia germanica, as pear-shaped structures occurring in the same situation, but regarded them as sense-organs, as did Cholodkowsky. Graber found these structures in the embryo of the May-beetle, which looked like the other embryonic limbs, but survived after the disappearance of the latter, being longer and broader and unjointed. These disappeared shortly before birth. In Hydrophilus they remain, Graber states, after birth. Nussbaum has seen them in Meloë.

Finally, Wheeler has discussed at length these embryonic organs, which he regards as glandular structures, and calls pleuropodia, their primitive function having been that of limbs. He has detected them in the embryo of Periplaneta orientalis, Mantis carolina, Xiphidium ensiferum (Fig. 387); also in the Hemiptera (Cicada septemdecim, Zaitha fluminea), and in Sialis infumata. He discards the view that they were once gills or sense-organs, and concludes that they were glands. But, as we have suggested, their function once that of gills, and still respiratory in Synaptera, has perhaps become in the winged insects glandular and repugnatorial. Instead, then, of being modified abdominal limbs afterwards serving as glands, as Wheeler claims, we are inclined to believe that they functioned as blood-gills.

k. Tracheal gills of adult insects

Tracheal gills are known to be retained by a few insects in the imago stage, the nymphs in all stages breathing by them. The most notable example is the perlid genus Pteronarcys, in which, as Newport states, there are eight sets, comprising 13 pairs of branchial tufts distributed over the under surface of the thoracic and first two abdominal segments.

The first set, consisting of three pairs of tufts, partly encircling the neck like a ruff, arises from the soft membrane connecting the head and prosternum. The thoracic tufts originate between and behind the coxæ, as well as on the front margin of the meso- and metathoracic segments. The number of filaments in each tuft varies from about 20 to 50 or more, the densest tufts being those of the two hinder thoracic segments. Each filament is usually simple, though in a few cases they are branched (Fig. 457, A).

The adult Pteronarcys is nocturnal, flying only at dewfall or in the night, and Mr. Barnston observed it when on the wing, “constantly dipping on the surface of the water”; by day it hides “in crevices of rocks which are constantly wetted by the spray of falling water, under stones and in other damp places.” It may thus be compared with the Amphibians, Necturus and Proteus, whose gills are retained in adult life. A similar large Chilian Perlid (Diamphipnoa lichenalis Gerst.) differs in completely lacking the thoracic gills, though there are four pairs on the abdomen, i.e. a pair on each of the first four segments. In this form the number of individual filaments in the largest tufts may amount to about 200.

Another Perlid (Dictyopteryx signata) is said by Hagen to have two pairs of gill-tufts on the under side of the head; the first pair situated on the base of the submentum, the second on the membrane connecting the head and prosternum.

Kolbe states that in the imagines of Perla marginata and P. cephalotes on the hinder edge of the thoracic stigmata arise three very small chitinous plates, which, on their under side and on the edges are beset with numerous short white filaments. These completely correspond to the filaments of the tuft-like larval gills. Persistent anal gills also occur in the imagines of Perla.

Fig. 457.—Under side of Pteronarcys regalis, showing the situation of the gills (g, b, f) and the sternal orifices: A, a branchial filament showing the direction of the current of blood; c, d, tracheæ. B, end of the abdomen enlarged.—After Newport.

In Nemoura lateralis and cinerea the tracheal gills are differently disposed. On each side of the anterior edge of the prosternum arise delicate tightly twisted filaments, like those of the larva. (Einführung, p. 536.)

Hagen also states that in the dragon-fly, Euphæa, the gills of the nymphs are retained in the imago, and Palmén remarks that in Æschna the rectal gills of the nymph persist in the imago, though not used for respiration.

Palmén gives an instance of a caddis-fly (Hydropsyche, Fig. 448) retaining its gills through the imago stage, but they are unfit for respiration, as they are minute and shrunken.

A walking-stick (Prisopus flabelliformis) found in the mountains of Brazil has the remarkable habit, according to Murray, of spending “the whole of the day under water, in a stream or rivulet, fixed firmly to a stone in the rapid part of the stream,” with its head turned up stream; but leaving the water at dark. The under side of the body, including the head, is hollowed so that the creature may adhere, sucker-like, to smooth stones; the claws, claspers, and flaps on the legs aid in retaining its hold, while the outer margin of the legs is dentate and thickly fringed with hair to repel the water.

Another form, closely related to Prisopus, from Borneo (Cotylosoma dipneusticum) is said by Wood-Mason to be even more profoundly modified for an aquatic life, since it has not only spiracles, but also, as he claims, tracheal gills. From each side of the body, in fact along the lower margins of the sides of the metathorax, there stand straight out five equal, small, but conspicuous ciliated oval plates, “which, when the insect is submerged and its stigmata are closed, doubtless serve for respiration.” The author did not note the actual presence of tracheæ in these plates.