Composition of Head.

Fig. 21.—Embryo of Aphis. Copied from Mecznikow, Zeits. f. wiss. Zool., Bd. XVI., taf. xxx., fig. 30. References in text. × 220.

Though the jaws of the Cockroach are eminently primitive with respect to those of most other Insects, they are themselves derived from a far simpler arrangement, which is demonstrable in all embryonic Insects. Fig. 21 shows an Aphis within the egg. The rudiments of the antennæ (At), mandibles (Mn), and maxillæ (Mx1,Mx2) form simple blunt projections, similar to each other and to the future thoracic legs (L1,L2,L3). We see, therefore, that all the appendages of an Insect are similar in an early stage of growth; and we may add that a Centipede, a Scorpion, or a Spider would present very nearly the same appearance in the same stage. A Crustacean in the egg would not resemble an Insect or its own parent so closely.57Aquatic life favours metamorphosis, and most Crustacea do not begin life with their full quota of legs, but acquire them as they are wanted.

Fig. 22.—Head of larva of Goat Moth, seen from behind. Copied from Lyonnet.

Paired appendages of perfectly simple form are therefore the first stage through which all Insect-jaws must pass. Our second stage is a little more complex, and not nearly so universal as the first. A caterpillar (fig. 22) has its own special wants, and these are met by the unequal development of its jaws. The mandibles are already as complete as those of the Cockroach, which theyclosely resemble, but the maxillæ are stunted cylinders formed mainly of simple rings, and very like the antennæ. They show, however, the beginnings of three processes (palp, galea, and lacinia), which are usually conspicuous in well-developed maxillæ. The second pair of maxillæ (Lm) are coalesced, as usual, and form the spinneret. The mouth-parts of the Caterpillar do not therefore in all respects represent a universal stage of development, but show important adaptive modifications. The mandibles are rapidly pushed forward, and attain their full development in the larva; the first pair of maxillæ are temporarily arrested in their growth, and persist for a long time in a condition which Orthopterous embryos quickly pass through; the maxillæ of the second pair are not only arrested in their growth, but converted to a special use, which seems to stop all further progress. The labial palps, indeed, which are not at all developed in the caterpillar, survive, and become important parts in the moth; but the greater part of the labium disappears when the time for spinning the cocoon is over.

Fig. 23.—Mouth-parts of Honey Bee.

Fig. 23A.—Diagram of Mouth-parts of Honey Bee.

We come next to the Orthopterous mouth, which is well illustrated by the Cockroach. This is retained with little modification in all the biting Insects (Coleoptera and Neuroptera). The mandibles may become long and pointed, as inStaphylinusand other predatory forms; in some larvæ of strong carnivorous propensities (Ant-lion,Dytiscus,58Chrysopa) they are perforate atthe tip, and through them the juices of the prey are sucked into the mouth, which has no other opening. The labium undergoes marked adaptive change, without great deviation from the common plan, in the “mask” of the larva of the Dragon-fly. This well-known implement has a rough likeness, in the arrangement and use of its parts, to a man’s fore-limb. The submentum forms the arm, the mentum the fore-arm. Both these are simple, straight pieces, connected by an elbow-joint. The hand is wider, and carries a pair of opposable claws, the paraglossæ. In some Coleoptera the labium is reduced to a stiff spine, while in the Stag-beetle it is flexible and hairy, and foreshadows the licking tongue of the Bee. The maxillæ become long and hairy in flower-haunting Beetles, and even the mandibles are flexible and hairy in the Scarabæus-beetles. Fritz Müller has found a singular resemblance to the proboscis of a Moth in a species ofNemognatha, where the maxillæ are transformed into two sharp grooved bristles 12 mm. long, which, when opposed, form a tube, but are incapable of rolling up.59

In the Honey Bee (fig.23) nearly all the mouth-parts of the Cockroach are to be made out, though some are small and others extremely produced in length. The mandibles (Mn) are not much altered, and are still used for biting, as well as for kneading wax and other domestic work. The mandibular teeth have proved inconvenient, and are gone. The lacinia of the maxilla (Mx′) forms a broad and flexible blade, used for piercing succulent tissues, but the galea has disappeared, and there is only a vestige of the maxillary palp (Mxp). In the second pair of maxillæ the palp (Lp) is prominent; its base forms a blade, while the tip is still useful as an organ of touch. The paraglossæ (Pa) can be made out, but the laciniæ are fused to form the long, hairy tongue. This ends in a spoon-shaped lobe (not unlike the “finger” of an elephant’s trunk), which is used both for licking and for sucking honey.

The proboscis of the Bee is therefore more like a case of instruments than a single organ. The mandibles form a strong pair of blunt scissors. The maxillæ are used for piercing, for stiffening and protecting the base of the tongue, and whenclosed they form an imperfect tube outside the tongue, which, according to Hermann Müller, is probably suctorial. The labial palps are protective and sensory. Lastly, the central part, or tongue, is a split tube used for suction; it is very long, so as topenetrate deep flower-cups, and hairy, so that pollen may stick to it. When the proboscis is not in use it can be slid into the mentum (M), while it and the mentum together can be drawn out of the way downwards and backwards.60

Fig. 24.—Mouth-parts of Burnet Moth.Fig. 24A.—Diagram of Mouth-parts of Moth.

Fig. 24.—Mouth-parts of Burnet Moth.

Fig. 24A.—Diagram of Mouth-parts of Moth.

In the singular suctorial mouth of Moths and Butterflies we observe, first of all, the great development of the maxillæ. Each forms a half-tube, which can be accurately applied to its fellow, so as to form an efficient siphon. In many species the two halves can be held together by a multitude of minute hooks.61At the base of each maxilla is a rudimentary palp (Mxp). The mandibles (Mn) are also rudimentary and perfectly useless. The labium, which was so important to the larva as a spinneret, has disappeared almost completely, but the labial palps (Lp) are large and evidently important.

Fig. 25.—Mouth-parts of Gad-fly (Tabanus).Fig. 25A.—Diagram of Mouth-parts of Gad-fly.

Fig. 25.—Mouth-parts of Gad-fly (Tabanus).

Fig. 25A.—Diagram of Mouth-parts of Gad-fly.

In Diptera both piercing and sucking parts are usually present. The Gad-fly (fig. 25) is typical. Here we recognise the labrum (Lbr), mandible (Mn), and maxilla (Mx′) of the Cockroach transformed into stylets. The maxillary palp (Mxp) is still sensory. A pointed process, stiffened by chitinous ribs, is developed from the back of the labrum. This is the epipharynx (Ep), a process undeveloped in the Cockroach, though conspicuous in some Coleoptera. All these parts are overtopped by the suctorial labium (Lm), which has a two-lobed expansion at the end. In the more specialised Diptera this becomes a kind of cupping-glass. The Gad-fly is intermediate between the Gnat, in which all the mouth-parts are converted into piercing organs of extraordinary length and sharpness, and such flies as the House-fly and Blow-fly, where the sucking labium forms an organ of the most elaborate kind, the piercing organs undergoing a marked reduction. Except where the labium is short, it is doubly or trebly hinged, so that it can be readily tucked away under the chin.

In Hemiptera the long four-jointed labium (Lm) forms a sheath for the stylets. When not in use the whole apparatus is drawn up beneath the head and prothorax. The mandibles (Mn) are sharp at the tip, and close like a pair of forceps, enclosing the maxillæ (Mx). These are of unequal length, only one reaching the end of the mandibular case. Both have saw teeth on the free edge. Palps are entirely wanting.

Fig. 26.—Mouth-parts of Bug. Copied from Landois, Zeits. f. wiss. Zool., Bd. XVIII., taf. xi., fig. 3.Fig. 26A.—Diagram of Mouth-parts of Bug.

Fig. 26.—Mouth-parts of Bug. Copied from Landois, Zeits. f. wiss. Zool., Bd. XVIII., taf. xi., fig. 3.

Fig. 26A.—Diagram of Mouth-parts of Bug.

Comparing the four kinds of suctorial mouths, of which the Bee, the Moth, the Fly, and the Bug furnish examples, we observe that the sucking-tube is formed in the Moth out of the two maxillæ, in the other three out of the labium. Of these last the Bee has the edges of the labium turneddown, so that the siphon becomesventral; in the Bug and Fly the edgesare turnedup, and the siphon becomesdorsal. The more specialised flies have the simple arrangement of the Bug complicated by a system of branching tubes, which are probably a special modification of the salivary duct. Similar as the mouth-parts of the four types may be in regard to their mode of working, they cannot be reduced to any common plan which differs materially from that presented by the jaws of the Cockroach.

In all Insects fusion of the primitive elements of the head begins so early and is carried so far, that it is extremely difficult to discover the precise way in which they are fitted together. The following facts have been ascertained respecting the development of the parts in question. At a very early stage of embryonic life the body of the Insect becomes divided into a series of segments, which are at fewest fourteen (in some Diptera), while they are not known to exceed seventeen.62Each segment is normally provided with a pair of appendages. The foremost segment soon enlarges beyond the rest, and becomes divided by a median groove into two “procephalic lobes.”63Of the appendages the first eight pairs are usually more prominent than the rest, and of different form; those of the eighth segment, which may be altogether inconspicuous, never attain any functional importance. The first four pairs of appendages are budded off from the future head, while the next three pairs form the walking legs, and are carried upon the thoracic segments. All the existing appendages of the fore part of the body are thus accounted for, but the exact mode of formation of the head has not yet been made out. The chief part of its walls, including the clypeus, the compound eyes, and the epicranial plates, arise from the procephalic lobes, and represent the much altered segment of which the antennæ are the appendages. The labrum is a secondary outgrowth from this segment, and, in some cases at least, it originates as a pair of processes which resemble trueappendages, though it is unlikely that such is their real character. No means at present exist for identifying the terga and sterna of the head, nor have the gena, the occipital frame, and the cervical sclerites (described below) been assigned to their segments.64It is worthy of notice that in the stalk-eyed Crustacea, the head, or what corresponds to the head of Insecta, consists of either five or six somites, taking into account a diversity of opinion with respect to the eyestalks, while only four pairs of appendages can be certainly traced in the head of the Insect. The mandibles and maxillæ exist to the same number in both groups, and are homologous organs, so far as is known; the numerical difference relates therefore to the antennæ, of which the Crustacean possesses two pairs, the Insect only one. Whether the pair deficient in the Insect is altogether undeveloped, or represented by the pair of prominences which give rise to the labrum,65is a question of much theoretical interest and of not a little difficulty.

The following table shows the appendages of the head and thorax in the two classes. The homologies indicated are, however, by no means established.66

Crayfish.Cockroach.Antennæ.————Eyestalks.Antennules.Antennæ.Mandibles.Mandibles.Maxillæ (1).Maxillæ (1).Maxillæ (2).Maxillæ (2).————Maxillipeds (1).Thoracic Legs (1).Maxillipeds (2).Thoracic Legs (2).Maxillipeds (3).Thoracic Legs (3).

The neck is a narrow cylindrical tube, with a flexible wall strengthened by eight plates, the cervical sclerites, two of which are dorsal, two ventral, and four lateral. The dorsal sclerites lie immediately behind the head (fig.14); they are triangular, and closely approximated to the middle line. The inferior plates (fig. 27) resemble segments of chitinous hoops set transversely, one behind the other, rather behind the dorsal sclerites, and close behind the submentum. There are two lateral sclerites on each side of the neck (fig. 27), a lower squarish one, which is set diagonally, nearly meeting its fellow across the ventral surface, and an oblong piece, closely adherent to the other, which extends forwards and upwards towards the dorsal side.

Fig. 27.—Ventral Plates of Neck and Thorax of Male Cockroach. I, prosternum; II, mesosternum; III, metasternum. × 6.

The elements of the thoracic exoskeleton are simpler in the Cockroach than in Insects of powerful flight, where adaptive changes greatly obscure the primitive arrangement. There are three segments, each defended by a dorsal plate (tergum) and a ventral plate (sternum). The sterna are often divided into lateral halves. Of the three terga the first (pronotum) is the largest; it has a wide free edge on each side, projects forwards over the neck, and when the head is retracted, covers this also, its semi-circular fore-edge then forming the apparent head-end of the animal. The two succeeding terga are of nearly equal size, and each is much shorter than the pronotum, contrary to the rule in winged Insects.67

All the terga are dense and opaque in the female; in the male the middle one (mesonotum) and the hindmost (metanotum) are thin and semi-transparent, being ordinarily overlaid by the wing-covers. While the thoracic terga diminish backwards, the sterna increase in extent and firmness, proportionally to the size of the attached legs. The prosternum is small and coffin-shaped; the mesosternum partly divided into lateral halves in the male, and completely so in the female. The metasternum is completely divided in both sexes, while a median piece, carrying the post-furca, intervenes between its lateral halves in the male. Behind the sterna, especially in the case of the second and third, the flexible under-surface of the thorax is inclined, so as to form a nearly vertical step. In the two hinder of these steps a chitinous prop is fixed; each is Y-shaped, with long, curved arms for muscular attachment, and a central notch, which supports the nerve-cord. The hindmost of these, known as thepost-furca, lies immediately behindthe metasternum, and its short basal piece is attached between the lateral halves of that plate. Behind the mesosternum is a somewhat slighter prop, themedi-furca. A third piece of similar nature (theante-furca), which is well developed in some Insects—e.g., in Ants—is apparently wanting in the Cockroach, though there is a transverse oval plate behind the prosternum, which may be a rudimentary furca.

Fig.27shows two conical processes which lie in the middle line of the ventral surface of the thorax, one in front of the metasternum, the other in front of the mesosternum. These are the thoracic pits, tubular apodemata, serving for the insertion of muscles. The occurrence of stink-glands in the thorax of Hemiptera,68and of so-called poison-glands in the thorax ofSolpuga, led us to look for glands in connection with these processes, but we have found none.

Fig. 28.—The three Thoracic Legs of a Female Cockroach. I,s, sternum;cx, coxa;tr, trochanter;fe, femur;tb, tibia;ta, tarsus. In IIIAthe coxa is abducted, and the jointsa(episternum) andbslightly separated. × 4.

Three pairs of legs are attached to the thoracic segments; they regularly increase in size from the first to the third, but hardly differ except in size; the peculiar modifications which affect the fore pair in predatory and burrowing Orthoptera (Mantis,Gryllotalpa), and the third pair in leaping Orthoptera (Grasshoppers, &c.), being absent in the cursorial Blattina. Each leg is divided into the five segments usual in Insects (see fig. 28). The coxa is broad and flattened. The trochanter is a small piece obliquely and almost immovably attached to the proximal end of the femur, on its inner side. The femur is nearly straight and narrowed at both ends; along its inner border, in the position occupied by the stridulating apparatus of the hind leg of the Grasshoppers, is a shallow longitudinal groove, fringed by stiff bristles. The tibia is shorter than the femur in the fore leg, of nearly the same length in the middle leg, and longer in the hind leg; it is armed with numerous stiff spines directed towards the free end of the limb. There are usually reckoned five joints in the tarsus, which regularly diminish in length, except that the last joint is as long as the second. All thejoints bear numerous fine but stiff hairs upon the walking surface. The extremity of the fifth joint is segmented off, and carries a pair of equal and strongly curved claws.69

At the base of each leg are several chitinous plates (fig. 28), upon which no small labour has been bestowed by different anatomists. They are arranged so as to form two joints intermediate between the coxa and the sternum, and these two joints admit of a hinge-like movement upon each other, while their other ends are firmly attached to the coxa and sternum respectively. (Compare III and IIIA, fig. 28.) These parts in the Cockroach may be taken for two basal leg-joints which have become adherent to the thorax. In other cases, however, they plainly belong to the thorax, and not to the leg. In the Mole-cricket, for instance, similar plates occur; but here they are firmly united, and form the lateral wall of the thorax. In the Locust they become vertical, and lie one in front of the other. Most authors have looked upon them as regular elements of a typical somite. They regard such a segment as including two pleural elements—viz., a dorsal plate (epimeron), and a ventral plate (episternum). We have already (p.34) given reasons for doubting the constancy of the pieces so named. It is not inconvenient, however, to denote by the termepisternumthe joint which abuts upon the sternum; for the joint which is applied to the coxa no convenient term exists, and its occurrence in Insects is so partial, that the want need not be supplied at present.70Both joints are incompletely subdivided. In the first thoracic segment of the Cockroach they are less firmly connected than in the other two.

Fig. 29.—Wings and Wing-covers of Male Cockroach. × 4.

Cockroaches of both sexes are provided with wings, which, however, are only functional in the male. The wing-covers (or anterior pair of wings) of the male are carried by the second thoracic segment. As in mostOrthoptera genuina, they are denser than the hind wings, and protect them when at rest. They reach to the fifth segment of the abdomen, and onewing-cover overlaps the other. Branching veins or nervures form a characteristic pattern upon the surface (figs.4, 29), and it is mainly by means of this pattern that many of the fossil species are identified and distinguished. The true or posterior wings are attached to the metathorax. They are membranous and flexible, but the fore-edge is stiffened, like that of the wing-covers, by additional chitinous deposit. When extended, each wing forms an irregular quadrant of a circle; when at rest, the radiating furrows of the hinder part close up fan-wise, and the inner half is folded beneath the outer.71The wing reaches back as far as the hinder end of the fourth abdominal segment. The wing-covers of the female are small, and though movable, seem never to be voluntarily extended; each covers about one-third of the width of the mesonotum, and extends backwards to the middle of the metanotum. A reticulated pattern on the outer fourth of the metanotum plainly represents the hind wing; it is clearly rather a degeneration or survival than an anticipation of an organ tending towards useful completeness.

The rudimentary wing of the female Cockroach illustrates the homology of the wings of Insects with the free edges of thoracic terga, and this correspondence is enforced by the study of the development of the more complete wings and wing-covers of the male. The hinder edges of the terga become produced at the later moults preceding the completely winged stage, and may even assume something of the shape and pattern of true wings; it is not, however, true, though more than once stated, that winged nymphs are common. Adults with imperfectly developed wings have been mistaken for such.

The structure of the wing testifies to its origin as a fold of the chitinous integument. It is a double lamina, which often encloses a visible space at its base. The nervures, with their vessels and tracheal tubes, lie between the two layers, which, except at the base, are in close contact. Oken termed the wings of an Insect “aerial gills,” and this rather fanciful designation is in some degree justified by their resemblance to the tracheal gills of such aquatic larvæ as those of Ephemeridæ, Perlidæ, Phryganidæ, &c. In the larva ofChloeon (Ephemera) dipterum(fig.30), for example, the second thoracic segment carries a pair of large expansions, which ultimately are replaced by organs of aerial flight. The abdominal segments carry similarly placed respiratory leaflets, the tracheal gills, which by their vigorous flapping movements bring a rush of water against their membranous and tracheated surfaces.

Gegenbaur72has argued from the resemblance of these appendages to wings, that the wing and the tracheal leaflet are homologous parts, and this view has been accepted as probable by so competent an observer as Sir John Lubbock.73

The leaflets placed most advantageously for propulsion seem to have become exclusively adapted to that end, while the abdominal gills have retained their respiratory character. At the time of change from aquatic to terrestrial life, which takes place in many common Insects when the adult condition isassumed, and which, according to Gegenbaur, was a normal event among primitive Insects, the tracheal gill is supposed to disappear, and in its place, at the next moult, an opening, the stigma, is formed by the rupture of an air-tube. Gegenbaur supposes that the primitive Insects were aquatic, and their tracheal system closed. The tracheal gill he takes to be the common structure which has yielded organs so unlike as the wing and the stigma.

Fig. 30. Chloeon (Chloeopsis) dipterum. Larva in eighth stage, with wings and respiratory leaflets. × 14. Copied from Vayssière (loc. cit.).Fig. 31.—Tricorythus. Adult larva, with three functional leaflets. The next leaflet in front is converted into a protective plate. × 7.A, protective plate of Tricorythus larva, seen from the outside. × 26.B, the same from within, showing the attached respiratory appendage.C, protective plate of Cœnis larva, without respiratory appendage.All the figures are copied from Vayssière.

Fig. 30. Chloeon (Chloeopsis) dipterum. Larva in eighth stage, with wings and respiratory leaflets. × 14. Copied from Vayssière (loc. cit.).

Fig. 31.—Tricorythus. Adult larva, with three functional leaflets. The next leaflet in front is converted into a protective plate. × 7.A, protective plate of Tricorythus larva, seen from the outside. × 26.B, the same from within, showing the attached respiratory appendage.C, protective plate of Cœnis larva, without respiratory appendage.All the figures are copied from Vayssière.

Fig. 31.—Tricorythus. Adult larva, with three functional leaflets. The next leaflet in front is converted into a protective plate. × 7.

A, protective plate of Tricorythus larva, seen from the outside. × 26.

B, the same from within, showing the attached respiratory appendage.

C, protective plate of Cœnis larva, without respiratory appendage.

All the figures are copied from Vayssière.

The zoological rank of the Insects (Ephemeridæ, Perlidæ, and Libellulidæ), in which tracheal gills are most usual, is not unfavourable to such an explanation. Lubbock has given reasons for regardingCampodeaand the Collembola (of the order Thysanura) as surviving and not very much altered representatives of the most primitive Insects, and he has shown that no great amount of modification would be required to convert the terrestrialCampodeainto the aquaticChloeon-nymph.74We must not forget, however, that tracheal gills are by no means restricted to these families of low grade. Trichoptera, a few Diptera, two Lepidoptera (NymphulaandAcentropus), and two Coleoptera (GyrinusandElmis),75have tracheal gills, and a closed tracheal system in the larval condition. We cannot suppose that these larvæ of higher orders represent an unbroken succession of aquatic forms, but if we refuse to adopt this alternative, we must admit that the closed tracheal system with tracheal gills may be an adaptive modification of the open system with stigmata.

It is wellknown76that in certain Ephemeridæ (e.g.,TricorythusandCœnis) a pair of anterior tracheal gills may become transformed into large plates, which partly protect the gills behind (fig.31). A similar modification of the second and third thoracic gills inProsopistomaandBætiscabrings all the functional respiratory organs under cover, and these enlarged plates resemble stiff and simple wings very closely.

Palmén77has subjected Gegenbaur’s hypothesis to a very searching examination. He observesthat:—

1. InCampodea, and presumably in other primitive Insects, the tracheal system is not closed and adapted for aquatic respiration, but open. Tracheal gills are not by any means confined to the lowest Insects. (See above, p.65.)

2. Tracheal gills are not always homodynamous or morphologically equivalent. In Ephemeridæ, some are dorsal in position, some ventral (first abdominal pair inOligoneuriaandRhithrogena); they may be cephalic, springing from the base of the maxilla, as inOligoneuriaandJolia;Joliahas a branchial tuft at the insertion of each of the fore legs.78In Perlidæ the tracheal gills may have a tergal, pleural, sternal, or anal insertion. In some Libellulidæ also, anal leaflets occur.79

3. Tracheal gills never perfectly agree in position and number with the stigmata throughout the body. Sometimes they occur on different rings, sometimes on different parts of the same ring. Gegenbaur’s statements on this point are incorrect.

4. Tracheal gills may co-exist with stigmata. In Perlidæ the tracheal gills persist in the imago, and may be found, dry and functionless, beneath the stigmata. In Trichoptera they gradually abort at successive moults, and in some cases remain after the stigmata have opened.

5. Stigmata do not form by the breaking off of tracheal appendages, but by the enlargement of rudimentary trachealbranches, which open into the main longitudinal trunks. In larvæ with aquatic respiration these branches exist, though they are not functional.

Palmén’s objections must be satisfactorily disposed of before Gegenbaur’s explanation, interesting as it is, can be fully accepted. Palmén has proved, what is on other grounds clear enough, that stigmata are more ancient than tracheal gills, aerial tracheate respiration than aquatic. But there is nothing as yet to contradict the view that the first Insect-wings were adapted for propulsion in water, and that they were respiratory organs before they became motor. It is Gegenbaur’s explanation of the origin of stigmata, and not his explanation of the origin of wings, which is refuted by Palmén.

In the abdomen of the female Cockroach eight terga (1–7; 10) are externally visible. Two more (8, 9) are readily displayed by extending the abdomen; they are ordinarily concealed beneath the seventh tergum. The tenth tergum is notched in the middle of its posterior margin. A pair of triangular “podical plates,” which lie on either side of the anus, and towards the dorsal surface, have been provisionally regarded by Prof. Huxley as the terga of an eleventh segment. Seven abdominal sterna (1–7) are externally visible. The first is quite rudimentary, and consists of a transversely oval plate; the second is irregular and imperfectly chitinised in front; the seventh is large, and its hinder part, which is boat-shaped, is divided into lateral halves, for facilitating the discharge of the large egg-capsule.

Fig. 32.—Under side of Abdomen of Male and Female Cockroach. × 4.

In the male Cockroach ten abdominal terga are visible without dissection (fig.33, p. 70), though the eighth and ninth are greatly overlapped by the seventh. The tenth tergum is hardly notched. Nine abdominal sterna are readily made out, the first being rudimentary, as in the female. The eighth is narrower than the seventh, the ninth still narrower, and largely concealed by the eighth; its covered anterior part is thin and transparent, the exposed part denser. This forms the extreme end of the body, except that the small sub-anal styles project beyond it. The podical plates resemble those of the female.

Pleural elements are developed in the form of narrow stigmatic plates, with the free edge directed backwards. These lie between the terga and sterna, and defend the spiracle.80

The modifications of the hindmost abdominal segments will be more fully considered in connection with the reproductive organs.

The high number of abdominal segments found in the Cockroach (ten or eleven) is characteristic of the lower orders of Insecta. It is never exceeded; though in the more specialised orders, such as Lepidoptera and Diptera, it may be reduced to nine, eight, or even seven. The sessile abdomen of the Cockroach is primitive with respect to the pedunculate abdomen found in such insects as Hymenoptera, where the constricted and flexible waist stands in obvious relation to the operations of stinging and boring, or to peculiar modes of oviposition. The first abdominal segment, which is especially liable to dislocation and alteration in Insects, occupies its theoretical position in the Cockroach, though both tergum and sternumare reduced in size. The sternum is often altogether wanting, while the tergum may unite with the metathorax.

Fig. 33.—Profile of Male and Female Cockroach. × 4.

The externally visible appendages of the abdomen are the cerci and the styles of the male Cockroach. The cerci are found in both sexes; they are composed of sixteen rings each, and project beneath the edge of the tenth tergum. They are capable of erection by special muscles, and are supplied by large nerves.81The sub-anal styles are peculiar in their insertion, being carried upon the sternum of their segment (the ninth).

The abdominal segments are never furnished with functional legs in adult Insects, but representatives of the lost appendages are often met with in larvæ. According to Bütschli,82all the abdominal segments are provided with appendages in the embryo of the Bee, though they disappear completely before hatching. Some Hymenopterous larvæ have as many as eight pairs of abdominal appendages, Lepidopterous larvæ at most five (3–6; 10).83

The Muscles; the Fat-Body and Cœlom.

SPECIAL REFERENCES.Viallanes.Histologie et Développement des Insectes. Ann. Sci. Nat., Zool., Tom. XIV. (1882).Kühnein Stricker’s Histology, Vol. I., chap. v.Plateau.Various Memoirs in Bull. Acad. Roy. de Belgique (1865, 1866, 1883, 1884). [Relative and Absolute Muscular Force.]Leydig.Zum feineren Bau der Arthropoden. Müller’s Archiv., 1855.Weismann.Ueber zwei Typen contractilen Gewebes, &c. Zeits. für ration. Medicin. Bd. XV. (1862).

SPECIAL REFERENCES.

Viallanes.Histologie et Développement des Insectes. Ann. Sci. Nat., Zool., Tom. XIV. (1882).

Kühnein Stricker’s Histology, Vol. I., chap. v.

Plateau.Various Memoirs in Bull. Acad. Roy. de Belgique (1865, 1866, 1883, 1884). [Relative and Absolute Muscular Force.]

Leydig.Zum feineren Bau der Arthropoden. Müller’s Archiv., 1855.

Weismann.Ueber zwei Typen contractilen Gewebes, &c. Zeits. für ration. Medicin. Bd. XV. (1862).

The muscles of the Cockroach, when quite fresh, appear semi-transparent and colourless. If subjected to pressure or strain they are found to be extremely tender. Alcohol hardens and contracts them, while it renders them opaque and brittle.

The minute structure of the voluntary or striped muscular fibres of Vertebrates is described in common text-books.84Each fibre is invested by a transparent elastic sheath, the sarcolemma, and the space within the sarcolemma is subdivided by transverse membranes into a series of compartments. The compartments are nearly filled by as many contractile discs, broad, doubly refractive plates, which are further divisible into prismatic columns, the sarcous elements, each being as long as the contractile disc. Successive sarcous elements, continued from one compartment to another, form the primitive fibrils of the muscle. In cross-section the fibrils appear as polygonal areas bounded by bright lines. Outside the fibres, but within the sarcolemma, are nuclei, imbedded in the protoplasm, or living and formative element of the tissue.

The muscular fibres of Insects present some important differences from the fibres just described. The nuclei are often found in the centre, and not on the surface of the fibres in both Insects and Crustacea. In both classes the fibrils are frequently subdivided into longitudinal strands, which have not been distinguished in Vertebrate muscles (Viallanes). The sarcolemma is often undeveloped. Lastly, Insects, like other Arthropoda, exhibit the remarkable peculiarity that not only their voluntary muscles, but all, or nearly all, the muscles of the body, even those of the digestive tube, are striated.85

The arrangement of the muscles in an Insect varies greatly according to situation and mode of action. Some of the abdominal muscles consist solely of straight parallel bundles, while the muscles of the limbs usually converge to tendinous insertions. In certain larvæ, where the segments show hardly any differentiation, the muscles form a sheet which covers the whole body, and is regularly segmented in correspondence with the exo-skeleton. As the movements of the body and limbs become more varied and more energetic, the muscles become grouped in a more complicated fashion, and the legs and wings of a flying Insect may be set in motion by a muscular apparatus almost as elaborate as that of a bird.

The following short notes on the muscles of the Cockroach, aided by reference to the figures, will render the more noteworthy features intelligible. A very lengthy description, far beyond our space or the reader’s patience, would be required to explain in detail the musculature of the head, limbs, and other specialised regions.

Sternal Muscles of Abdomen.—Thelongitudinal sternal muscles(fig. 34) form a nearly continuous transversely segmented sheet, covering the ventral surface between the fore-edge of the second abdominal sternum and the fore-edge of the seventh. These muscles, in conjunction with the longitudinal tergal muscles, tend to telescope the segments.

Theoblique sternal muscles(fig. 34), which are very short, connect the adjacent edges of the sterna (2–3, 3–4, 4–5, 5–6, 6–7). They extend inwards nearly to the middle line, but, like the longitudinal sternal muscles, they are not developed beneath the nerve-cord. Acting together, the oblique sternal muscles would antagonise the longitudinal, but it is probable that they are chiefly used to effect lateral flexion of the abdomen, and that only the muscles of one side of the abdomen contract at once.

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