Chapter IX: Introduction: Habits–classification–structure–chilognatha–chilopoda (3)

_Definition_.—Insects are small animals, having the body divided into three regions placed in longitudinal succession—head, thorax, and abdomen: they take in air by means of tracheae, a system of tubes distributed throughout the body, and opening externally by means of orifices placed at the sides of the body. They have six legs, and a pair of antennae; these latter are placed on the head, while the legs are attached to the thorax, or second of the three great body divisions; the abdomen has no true legs, but not infrequently has terminal appendages and, on the under surface, protuberances which serve as feet. Very frequently there are two pairs of wings, sometimes only one pair, in other cases none: the wings are always placed on the thorax. Insects are transversely segmented—that is to say, the body has the form of a succession of {87}rings; but this condition is in many cases obscure; the number of these rings rarely, if ever, exceeds thirteen in addition to the head and to a terminal piece that sometimes exists. Insects usually change much in appearance in the course of their growth, the annulose or ringed condition being most evident in the early part of the individual's life. The legs are usually elongate and apparently jointed, but in the immature condition may be altogether absent, or very short; in the latter case the jointing is obscure. The number of jointed legs is always six.

EXTERNAL STRUCTURE.

The series of rings of which the external crust or skeleton of Insects is composed exhibits great modifications, not only in the various kinds of Insects but even in the different parts of the same individual, and at successive periods of its development; so that in the majority of mature Insects the separate rings are readily distinguished only in the hind body or abdomen. The total number of the visible rings, segments, somites, or arthromeres, as they are variously called by different writers, is frequently thirteen in addition to the head. This latter part is considered to be itself composed of the elements of several rings, but morphologists are not yet agreed as to their number, some thinking this is three while others place it as high as seven; three or four being, perhaps, the figures at present most in favour, though Viallanes, who has recently discussed[17] the subject, considers six, the number suggested by Huxley, as the most probable. Cholodkovsky is of a similar opinion. However this may be, the three rings behind the head constitute the thorax, which is always largely developed, though, like the head, its segmentation is usually very much obscured by unequal development of different parts, or by consolidation of some of them, or by both of these conditions. The third great division of the body, the abdomen, is also usually much modified by one or more of the terminal segments being changed in form, or even entirely withdrawn into the interior of the body. The existence of ten segments in the hind body can, however, be very frequently actually demonstrated, so that it is correct to speak of ten as the normal number.

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It is no reproach to morphologists that they have not yet agreed as to the number of segments that may be taken as typical for an Insect, for all the branches of evidence bearing on the point are still imperfect. It may be well, therefore, to state the most extreme views that appear to be at all admissible. Hagen[18] has recently stated the opinion that each thoracic segment consists really of three segments—an anterior or wing-bearer, a middle or leg-bearer, and a posterior or stigma-bearer. There seems to be no reason for treating the stigma as being at all of the nature of an appendage, and the theory of a triple origin for these segments may be dismissed. There are, however, several facts that indicate a duplicity in these somites, among which we may specially mention the remarkable constancy of two pleural pieces on each side of each thoracic segment. The hypothesis of these rings being each the representative of two segments cannot therefore be at present considered entirely untenable, and in that case the maximum and minimum numbers that can be suggested appear to be twenty-four and eleven, distributed as follows:—

Maximum. Minimum. {89}
Head 7 3
Thorax 6 3
Abdomen 11 5
-- --
Total 24 11

Although it is not probable that ultimately so great a difference as these figures indicate will be found to prevail, it is certainly at present premature to say that all Insects are made up of the same number of primary segments.

A brief account of the structure of the integument will be found in the chapter dealing with the post-embryonic development.

The three great regions of the Insect body are functionally as well as anatomically distinct. The head bears the most important of the sense organs, viz. the antennae and ocular organs; it includes the greater of the nerve-centres, and carries the mouth as well as the appendages, the trophi, connected therewith. The thorax is chiefly devoted to the organs of locomotion, bearing externally the wings and legs, and including considerable masses of muscles, as well as the nerve centres by which they are innervated; through the thorax there pass, however, in the longitudinal direction, those structures by which the unity of the organisation is completed, viz. the alimentary canal, the dorsal vessel or "heart" for distributing the nutritive fluid, and also the nerve cords. The abdomen includes the greater part of the organs for carrying on the life of the individual and of the species; it also frequently bears externally, at or near its termination, appendages that are doubtless usually organs of sense of a tactile nature.

In the lower forms of Insect life there is little or no actual internal triple division of the body; but in the higher forms such separation becomes wonderfully complete, so that the head may communicate with the thorax only by a narrow isthmus, and the thorax with the abdomen only by a very slender link. This arrangement is carried to its greatest extreme in the Hymenoptera Aculeata. It may be looked on as possibly a means for separating the nutrition of the parts included in the three great body divisions.

Along each side of the body extends a series of orifices for the admission of air, the stigmata or spiracles; there are none of these on the head, but on each side of most of the other segments {90}there is one of these spiracles. This, however, is a rule subject to many exceptions, and it is doubtful whether there is ever a spiracle on the last abdominal segment. Even in the young stage of the Insect the number of these stigmata is variable; while in the perfect Insect the positions of some of the stigmata may be much modified correlatively with the unequal development or consolidation of parts, especially of the thorax when it is highly modified for bearing the wings.

The segments of the Insect are not separate parts connected with one another by joints and ligaments; the condition of the Insect crust is in fact that of a continuous long sac, in which there are slight constrictions giving rise to the segments, the interior of the sac being always traversed from end to end by a tube, or rather by the invaginated ends of the sac itself which connect with an included second sac, the stomach. The more prominent or exposed parts of the external sac are more or less hard, while the constricted parts remain delicate, and thus the continuous bag comes to consist of a series of more or less hard rings connected by more delicate membranes. This condition is readily seen in distended larvae, and is shown by our figure 48 which is taken from the same specimen, whose portrait, drawn during life, will be given when we come to the Coleoptera, family Cleridae. The nature of the concealed connexions between the apparently separate segments of Insects is shown at _m_, Fig. 47, p. 88.

As the number of segments in the adult Insect corresponds—except in the head—with the number of divisions that appear very early in the embryo, we conclude that the segmentation of the adult is, even in Insects which change their form very greatly during growth, due to the condition that existed in the embryo; but it must not be forgotten that important secondary changes occur in the somites during the growth and development of the individual. Hence in some cases there appear to be more than the usual number of segments, e.g. _Cardiophorus_ larva, and in others the number of somites is diminished by {91}amalgamation, or by the extreme reduction in size of some of the parts.

Besides the division of the body into consecutive segments, another feature is usually conspicuous; the upper part, in many segments, being differentiated from the lower and the two being connected together by intervening parts in somewhat the same sort of way as the segments themselves are connected. Such a differentiation is never visible on the head, but may frequently be seen in the thorax, and almost always in the abdomen. A dorsal and a ventral aspect are thus separated, while the connecting bond on either side forms a pleuron. By this differentiation a second form of symmetry is introduced, for whereas there is but one upper and one lower aspect, and the two do not correspond, there are two lateral and similar areas. This bilateral symmetry is conspicuous in nearly all the external parts of the body, and extends to most of the internal organs. The pleura, or lateral regions of the sac, frequently remain membranous when the dorsal and ventral aspects are hard. The dorsal parts of the Insect's rings are also called by writers terga, or nota, and the ventral parts sterna.

The appendages of the body are:—(1) a pair of antennae; (2) the trophi, constituted by three pairs of mouth-parts; (3) three pairs of legs; (4) the wings[19]; (5) abdominal appendages of various kinds, but usually jointed. Before considering these in detail we shall do well to make ourselves more fully acquainted with the elementary details of the structure of the trunk.

In the adult Insect the integument or crust of the body is more or less hard or shell-like, sometimes, indeed, very hard, and on examination it will be seen that besides the divisions into segments and into dorsal, ventral, and pleural regions, there are lines indicating the existence of other divisions, and it will be found that by dissection along these lines distinct pieces can be readily separated. Each hard piece that can be so separated is called a sclerite, and the individual sclerites of a segment have received names from entomotomists. The sclerites are not really {92}quite separate pieces, though we are in the habit of speaking of them as if such were the case. If an Insect be distended by pressure from the interior, many of the sclerites can be forced apart, and it is then seen that they are connected by delicate membrane. The structure is thus made up of hard parts meeting one another along certain lines of union—sutures—so that the original membranous continuity may be quite concealed. In many Insects, or in parts of them, the sclerites do not come into apposition by sutures, and are thus, as it were, islands of hard matter surrounded by membrane. A brief consideration of some of the more important sclerites is all that is necessary for our present purpose: we will begin with the head.

The head is most variable in size and form; as a part of its surface is occupied by the eyes and as these organs differ in shape, extent, and position to a surprising degree, it is not a matter for astonishment that it is almost impossible to agree as to terms for the areas of the head. Of the sclerites of the head itself there are only three that are sufficiently constant and definite to be worthy of description here. These are the clypeus, the epicranium, and the gula. The clypeus is situate on the upper surface of the head-capsule, in front; it bears the labrum which may be briefly described as a sort of flap forming an upper lip. The labrum is usually possessed of some amount of mobility. The clypeus itself is excessively variable in size and form, and sometimes cannot be delimited owing to the obliteration of the suture of connexion with the more posterior part of the head; it is rarely or never a paired piece. Occasionally there is a more or less distinct piece interposed between the clypeus and the labrum, and which is the source of considerable difficulty, as it may be taken for the clypeus. Some authors call the clypeus the epistome, but it is better to use this latter term for the purpose of indicating the part that is immediately behind the labrum, whether that part be the clypeus, or some other sclerite; the {93}term is very convenient in those cases where the structure cannot be, or has not been, satisfactorily determined morphologically.

In Figure 50 the parts usually visible on the anterior aspect of the head and its appendages are shown so far as these latter can be seen when the mouth is closed; in the case of the Insect here represented the bases of the mandibles are clearly seen (_g_), while their apical portions are entirely covered by the labrum, just below the lower margin of which the tips of the maxillae are seen, looking as if they were the continuations of the mandibles.

The labrum is a somewhat perplexing piece, morphologists being not yet agreed as to its nature; it is usually placed quite on the front of the head, and varies extremely in form; it is nearly always a single or unpaired piece; the French morphologist Chatin considers that it is really a paired structure.

The gula (Fig. 49, B _d_, and Fig. 47, _z_) is a piece existing in the middle longitudinally of the under-surface of the head; in front it bears the mentum or the submentum, and extends backwards to the great occipital foramen, but in some Insects the gula is in front very distant from the edge of the buccal cavity. The epicranium forms the larger part of the head, and is consequently most inconstant in size and shape; it usually occupies the larger part of the upper-surface, and is reflected to the under-surface to meet the gula. Sometimes a transverse line exists (Fig. 49, A) dividing the epicranium into two parts, the posterior of which has been called the protocranium; which, however, is not a good term. The epicranium bears the antennae; these organs do not come out between the epicranium and the clypeus, the foramen for their insertion being seated entirely in the epicranium (see Fig. 50). In some Insects there are traces of the epicranium being divided longitudinally along the middle line. When this part is much modified the antennae may appear to be inserted on the lateral portions of the head, or even {94}on its under-side; this arises from extension of some part of the epicranium, as shown in Fig. 49, B, where _h_, the cavity of insertion of the antenna, appears to be situate on the under-surface of the epicranium, the appearance being due to an infolding of an angle of the part.

There is always a gap in the back of the head for the passage of the alimentary canal and other organs into the thorax; this opening is called the occipital foramen. Various terms, such as frons, vertex, occiput, temples, and cheeks, have been used for designating areas of the head. The only one of these which is of importance is the gena, and even this can only be defined as the anterior part of the lateral portion of the head-capsule. An extended study of the comparative anatomy of the head-capsule is still a desideratum in entomology. The appendages of the head that are engaged in the operations of feeding are frequently spoken of collectively as the trophi, a term which includes the labrum as well as the true buccal appendages.

The appendages forming the parts of the mouth are paired, and consist of the mandibles, the maxillae, and the labium, the pair in this latter part being combined to form a single body. The buccal appendages are frequently spoken of as gnathites. The gnathites are some, if not all, of them composed of apparently numerous parts, some of these being distinct sclerites, others membranous structures which may be either bare or pubescent—that is, covered with delicate short hair. In Insects the mouth functions in two quite different ways, by biting or by sucking. The Insects that bite are called Mandibulata, and those that suck Haustellata. In the mandibulate Insects the composition of the gnathites is readily comprehensible, so that in nearly the whole of the vast number of species of that type the corresponding parts can be recognised with something like certainty. This, however, is not the case with the sucking Insects; in them the parts of the mouth are very different indeed, so that in some cases morphologists are not agreed as to what parts really correspond with some of the structures of the Mandibulata. At present it will be sufficient for us to consider only the mandibulate mouth, leaving the various forms of sucking mouth to be discussed when we treat of the Orders of Haustellata in detail.

The upper or anterior pair of gnathites is the mandibles, (Fig. 50, _g_). There is no part of the body that varies more than {95}does the mandible, even in the mandibulate Insects. It can scarcely be detected in some, while in others, as in the male stag-beetle, it may attain the length of the whole of the rest of the body; its form, too, varies as much as its size; most usually, however, the pair of mandibles are somewhat of the form of callipers, and are used for biting, cutting, holding, or crushing purposes. The mandibles are frequently armed with processes spoken of as teeth, but which must not be in any way confounded with the teeth of Vertebrates. The only Insects that possess an articulated tooth are the Passalidae, beetles armed with a rather large mandible bearing a single mobile tooth among others that are not so. Wood Mason and Chatin consider the mandibles to be, morphologically, jointed appendages, and the latter authority states that in the mandible of _Embia_ he has been able to distinguish the same elements as exist in the maxillae. In aculeate Hymenoptera the mandibles are used to a considerable extent for industrial purposes.

The maxilla is a complex organ consisting of numerous pieces, viz. cardo, stipes, palpiger, galea, lacinia, palpus. The galea and lacinia are frequently called the lobes of the maxilla. The maxilla no doubt acts as a sense organ as well as a mechanical apparatus for holding; this latter function being subordinate to the other. In Fig. 68, p. 122, we have represented a complex maxillary sense-organ.

The labium or lower lip has as its basal portion the {96}undivided mentum, and closes the mouth beneath or behind, according as the position of the head varies. In most Insects the labium appears very different from the maxilla, but in many cases several of the parts corresponding to those of the maxilla can be clearly traced in the labium.

The mentum is an undivided, frequently very hard, piece, continuous with either the submentum or the gula, and anterior to this are placed the other parts, viz. the labial palpi and their supports, the palpigers; beyond and between these exists a central piece (Fig. 52, B, _e_), about whose name some difference of opinion prevails, but which may be called the ligula (languette of French authors), and on each side of this is a paraglossa. In the Orthoptera the single median piece—the ligula of Coleopterists—is represented by two divided parts. In some Insects (many Coleoptera) there is interposed between the mentum and the palpigers a piece called the hypoglottis (Fig. 52, B, _b_). It is not so well ascertained as it should be, that the pieces of the lower lip bearing the same names in different Orders are in all cases really homologous, and comparison suggests that the hypoglottis of Coleoptera may possibly represent the piece corresponding to the mentum of Orthopterists, the so-called mentum of beetles being in that case the submentum of Orthopterists.

There is another part of the mouth to which we may call special attention, as it has recently attracted more attention than it formerly did; it is a membranous lobe in the interior of the mouth, very conspicuous in Orthoptera, and called the tongue, lingua, or hypopharynx; it reposes, in the interior of the mouth (Fig. 51, _o_), on the middle parts of the front of the labium; it is probably not entirely lost in Coleoptera, but enters into the composition of the {97}complex middle part of the lip by amalgamation with the paraglossae. It has recently been proposed to treat this lingua as the morphological equivalent of the labium or of the maxillae, giving it the name of the endolabium, but the propriety of this course remains to be proved;[20] the view is apparently suggested chiefly by the structure of the mouth of _Hemimerus_, a very rare and most peculiar Insect that has not as yet been sufficiently studied.

As the maxillae and labium are largely used by taxonomists in the systematic arrangement of the mandibulate Insects, we give a figure of them as seen in Coleoptera, where the parts, though closely amalgamated, can nevertheless be distinguished. This Fig. 52 should be compared with Fig. 51.

In speaking of the segments of the body we pointed out that they were not separate parts but constituted an uninterrupted whole, and it is well to remark here that this is also true of the gnathites. Although the mouth parts are spoken of as separate pieces, they really form only projections from the great body wall. Fig. 51, B, shows the intimate connexion that exists between the maxillae and labium; the continuity of the mandibles with the membrane of the buccal cavity is capable of very easy demonstration.

The head bears, besides the pieces we have considered, a pair of antennae. These organs, though varying excessively in form, are always present in the adult Insect, and exist even in the majority of young Insects. They are very mobile, highly sensitive organs, situate on or near the front part of the head. The antennae arise in the embryo from the procephalic lobes, the morphological import of which parts is one of the most difficult points connected with Insect embryology.

The eyes of Insects are of two sorts, simple and compound. The simple eyes, or ocelli, vary in number from one to as many as eighteen or twenty; when thus numerous they are situated in groups on each side of the head. In their most perfect form, as found in adult aculeate Hymenoptera, in Orthoptera and Diptera, ocelli are usually two or three in number, and present the appearance of small, perfectly transparent lenses inserted in the integument. In their simplest form they are said to consist of some masses of pigment in connexion with a nerve.

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The compound, or facetted, eyes are the most remarkable of all the structures of the Insect, and in the higher and more active forms, such as the Dragon-flies and hovering Diptera, attain a complexity and delicacy of organisation that elicit the highest admiration from every one who studies them. They are totally different in structure and very distinct in function from the eyes of Vertebrata, and are seated on very large special lobes of the brain (see Fig. 65), which indeed are so large and so complex in structure that Insects may be described as possessing special ocular brains brought into relation with the lights, shades, and movements of the external world by a remarkably complex optical apparatus. This instrumental part of the eye is called the dioptric part in contradistinction from the percipient portion, and consists of an outer corneal lens (_a_, Fig. 53), whose exposed surface forms one of the facets of the eye; under the lens is placed the crystalline cone (_b_), this latter being borne on a rod-like object (_c_), called the rhabdom. There are two layers of pigment, the outer (_e_), called the iris-pigment, the inner (_f_), the retinal-pigment; underneath, or rather we should say more central than, the rhabdoms is the fenestrate membrane (_d_), beyond which there is an extremely complex mass of nerve-fibres; nerves also penetrate the fenestrate membrane, and their distal extremities are connected with the delicate sheaths by one of which each rhabdom is surrounded, the combination of sheath and nerves forming a retinula. Each set of the parts above the fenestrate membrane constitutes an ommatidium, and there may be many of these ommatidia in an eye; indeed, it is said that the eye of a small beetle, _Mordella_, contains as many as 25,000 ommatidia. As a rule the larvae of Insects with a complete metamorphosis bear only simple eyes. In the young of Dragon-flies, as well as of some other Insects having a less perfect metamorphosis, the compound eyes exist in the early stages, but they {99}have then an obscure appearance, and are probably functionally imperfect.

In the interior of the head there exists a horny framework called the tentorium, whose chief office apparently is to protect the brain. It is different in kind according to the species. The head shows a remarkable and unique relation to the following segments. It is the rule in Insect structure that the back of a segment overlaps the front part of the one following it; in other words, each segment receives within it the front of the one behind it. Though this is one of the most constant features of Insect anatomy, it is departed from in the case of the head, which may be either received into, or overlapped by, the segment following it, but never itself overlaps the latter. There is perhaps but a single Insect (_Hypocephalus_, an anomalous beetle) in which the relation between the head and thorax can be considered to be at all similar to that which exists between each of the other segments of the body and that following it; and even in _Hypocephalus_ it is only the posterior angles of the head that overlap the thorax. Although the head usually appears to be very closely connected with the thorax, and is very frequently in repose received to a considerable extent within the latter, it nevertheless enjoys great freedom of motion; this is obtained by means of a large membrane, capable of much corrugation, and in which there are seated some sclerites, so arranged as to fold together and occupy little space when the head is retracted, but which help to prop and support it when extended for feeding or other purposes. These pieces are called the cervical sclerites or plates. They are very largely developed in Hymenoptera, in many Coleoptera, and in Blattidæ, and have not yet received from anatomists a sufficient amount of attention. Huxley suggested that they may be portions of head segments.

THORAX.

The thorax, being composed of the three consecutive rings behind the head, falls naturally into three divisions—pro-, meso-, {100}and metathorax. These three segments differ greatly in their relative proportions in different Insects, and in different stages of the same Insect's life. In their more highly developed conditions each of the three divisions is of complex structure, and the sclerites of which it is externally made up are sufficiently constant in their numbers and relative positions to permit of their identification in a vast number of cases; hence the sclerites have received names, and their nomenclature is of practical importance, because some, if not all, of these parts are made use of in the classification of Insects. Each division of the thorax has an upper region, called synonymically dorsum, notum, or tergum; an inferior or ventral region, called sternum; and on each side a lateral region, the pleuron. These regions of each of the three thoracic divisions are further distinguished by joining to their name an indication of the segment spoken of, in the form of the prefixes pro-, meso-, and meta-; thus the pronotum, prosternum, and propleura make up the prothorax. The thoracic regions are each made up of sclerites whose nomenclature is due to Audouin.[21] He considered that every thoracic ring is composed of the pieces shown in Fig. 55, viz. (1) the sternum (B', _a_), an unpaired ventral piece; (2) the notum (A), composed of four pieces placed in consecutive longitudinal order (A'), and named praescutum (_a_), scutum (_b_), scutellum (_c_), and post-scutellum (_d_); (3) lateral pieces, of which he distinguished on each side an episternum (B', _c_), epimeron (_e_), and parapteron (_d_), these together forming the pleuron. We give Audouin's Figure, but we cannot enter on a full discussion of his views as to the thorax; they have become widely known, though the constancy of the parts is not so great as he supposed it would prove to be. Sometimes it is impossible to find all the elements he thought should be present in a thoracic ring, while in other cases too many sclerites exist. As a rule the notum of the meso- and metathoraces is in greater part composed of two pieces, the scutum and the scutellum; while in the pronotum only one dorsal piece can be satisfactorily distinguished, though a study of the development may show that really two are frequently, if not usually, present. On the other hand, one, or more, of the notal sclerites in some cases shows evidence of longitudinal division along the middle. The sternum or ventral piece, though varying greatly in form, is {101}the most constant element of a thoracic segment, but it has sometimes the appearance of consisting of two parts, an anterior and a posterior. The pleuron nearly always consists quite evidently of two parts, the episternum, the more anterior and inferior, and the epimeron.[22] The relations between these two parts vary much; in some cases the episternum is conspicuously the more anterior, while in others the epimeron is placed much above it, and may extend nearly as far forwards as it. It may be said, as a rule, that when the sternum extends farther backwards than the notum, the epimeron is above the episternum, as in many Coleoptera; but if the sternum be anterior to the notum, then the episternum is superior to the epimeron, as in dragon-flies. We would here again reiterate the fact that these "pieces" are really not separate parts, but are more or less indurated portions of a continuous integument, which is frequently entirely occupied by them; hence a portion of a sclerite that in one species is hard, may in an allied form be wholly or partly membranous, and in such case its delimitation may be very evident on some of its sides, and quite obscure on another.

{102}The parapteron of Audouin does not appear to be really a distinct portion of the pleuron; in the case of _Dytiscus_ it is apparently merely a thickening of an edge. Audouin supposed this part to be specially connected with the wing-articulation, and the term has been subsequently used by other writers in connexion with several little pieces that exist in the pleural region of winged Insects.

The prothorax is even more subject to variation in its development than the other divisions of the thorax are. In the Hymenoptera the prosternum is disconnected from the pronotum and is capable, together with the first pair of legs, of movement independent of its corresponding dorsal part, the pronotum, which in this Order is always more or less completely united with the meso-thorax; in the Diptera the rule is that the three thoracic segments are closely consolidated into one mass. In the majority of Insects the prothorax is comparatively free, that is to say, it is not so closely united with the other two thoracic segments as they are with one another. The three thoracic rings are seen in a comparatively uniform state of development in a great number of larvae; also in the adult stages of some Aptera, and among winged insects in some Neuroptera such as the Embiidae, Termitidae, and Perlidae. In Lepidoptera the pronotum bears a pair of erectile processes called patagia; though frequently of moderately large size, they escape observation, being covered with scales and usually closely adpressed to the sides of the pronotum.

The two great divisions of the body—the mesothorax and the metathorax—are usually very intimately combined in winged Insects, and even when the prothorax is free, as in Coleoptera, these posterior two thoracic rings are very greatly amalgamated. In the higher forms of the Order just mentioned the mesosternum and mesopleuron become changed in direction, and form as it were a diaphragm closing the front of the metasternum. The meso- and meta-thorax frequently each bear a pair of wings.

We have described briefly and figured (Fig. 55) the sclerites of the mesothorax, and those of the metathorax correspond fairly well with them. In addition to the sclerites usually described as constituting these two thoracic divisions, there are some small pieces at the bases of the wings. Jurine discriminated and named no less than seven of these at the base of the anterior {103}wing of a Hymenopteron. One of them becomes of considerable size and importance in the Order just mentioned, and seems to be articulated so as to exert pressure on the base of the costa of the wing. This structure attains its maximum of development in a genus (? nondescript) of Scoliidae, as shown in Fig. 56. The best name for this sclerite seems to be that proposed by Kirby and Spence, tegula. Some writers call it paraptère, hypoptère, or squamule, and others have termed it patagium; this latter name is, however, inadmissible, as it is applied to a process of the prothorax we have already alluded to.

To complete our account of the structure of the thorax it is necessary to mention certain hard parts projecting into its interior, but of which there is usually little or no trace externally. A large process in many Insects projects upwards from the sternum in a forked manner. It was called by Audouin the entothorax; some modern authors prefer the term apophysis. Longitudinal partitions of very large size, descending from the dorsum into the interior, also exist; these are called phragmas, and are of great importance in some Insects with perfect flight, such as Hymenoptera, Lepidoptera, and Diptera. There is no phragma in connection with the pronotum, but behind this part there may be three. A phragma has the appearance of being a fold of the dorsum; it serves as an attachment for muscles, and may probably be of service in other ways. More insignificant projections into the interior are the little pieces called apodemes (Fig. 57, _e_); these are placed at the sides of the thorax near the wings. The apophyses are no doubt useful in preserving the delicate vital organs from shocks, or from derangement by the muscular movements and the changes of position of the body.

The appendages of the thorax are (_a_) inferior, the legs; (_b_) {104}superior, the wings. The legs are always six in number, and are usually present even in larvae, though there exist many apodal larvae, especially in Diptera. The three pairs of legs form one of the most constant of the characters of Insects. They are jointed appendages and consist of foot, otherwise tarsus; tibia, femur, trochanter, and coxa; another piece, called trochantin more or less distinctly separated from the coxa, exists in many Insects. The legs are prolongations of the body sac, and are in closer relation with the epimera and with the episterna than with other parts of the crust, though they have a close relation with the sternum. If we look at the body and leg of a neuropterous Insect (Fig. 58) we see that the basal part of the leg—the coxa—is apparently a continuation of one of the two pleural pieces or of both; in the latter case one of the prolonged pieces forms the coxa proper, and the tip of the other forms a supporting piece, which may possibly be the homologue of the trochantin of some Insects. In some Orthoptera, especially in Blattidae, and in Termitidae, there is a transverse chitinised fold interposed between the sternum and the coxa, and this has the appearance of being the same piece as the trochantin of the anterior legs of Coleoptera.

Beyond the coxa comes the trochanter; this in many Hymenoptera is a double piece, though in other Insects it is single; usually it is the most insignificant part of the leg. The femur is, on the whole, the least variable part of the leg; the tibia, which follows it, being frequently highly modified for industrial or other purposes. The joint between the femur and the tibia is usually bent, and is therefore the most conspicuous one in the leg; it is called the knee. The other joints have not corresponding names, though that between the tibia and the tarsus is of great importance. The spines at the tip of the tibia, projecting beyond it, are called spurs, or calcares. The tarsus or {105}foot is extremely variable; it is very rarely absent, but may consist of only one piece—joint, as it is frequently called[23]—or of any larger number up to five, which may be considered the characteristic number in the higher Insect forms. The terminal joint of the tarsus bears normally a pair of claws; between the claws there is frequently a lobe or process, according to circumstances very varied in different Insects, called empodium, arolium, palmula, plantula, pseudonychium, or pulvillus. This latter name should only be used in those cases in which the sole of the foot is covered with a dense pubescence. The form of the individual tarsal joints and the armature or vestiture of the lower surface are highly variable. The most remarkable tarsus is that found on the front foot of the male _Dytiscus_.

It has been suggested that the claws and the terminal appendage of the tarsus ought to be counted as forming a distinct joint; hence some authors state that the higher Insects have six joints to the feet. These parts, however, are never counted as separate joints by systematic entomologists, and it has recently been stated that they are not such originally.

The parts of the foot at the extremity of the last tarsal joint proper are of great importance to the creature, and vary greatly in different Insects. The most constant part of this apparatus is a pair of claws, or a single claw. Between the two claws there may exist the additional apparatus referred to above. This in some Insects—notably in the Diptera—reaches a very complex development. We figure these structures in _Pelopaeus spinolae_, a fossorial Hymenopteron, remarking that our figures exhibit the apparatus in a state of retraction (Fig. 59). According to the nomenclature of Dahl and Ockler[24] the plate (_b_) on the dorsal aspect is the pressure plate (_Druck-Platte_), and acts as an agent of pressure on the sole of the pad (C, _e_); _c_ and _d_ on the underside are considered to be extension-agents; _c_, extension-plate; _d_, extension-sole (_Streck-Platte_, _Streck-Sohle_). These agents are assisted in acting on the pad by means of an elastic bow placed in the interior of the latter. The pad (_e_) is a very remarkable structure, capable of much extension and retraction; {106}when extended it is seen that the pressure plate is bent twice at a right angle so as to form a step, the distal part of which runs along the upper face of the basal part of the pad; the apical portion of this latter consists of two large lobes, which in repose, as shown in our Figure (_f_), fall back on the pad, something in the fashion of the retracted claws of the cat, and conceal the pressure-plate.

The mode in which Insects are able to walk on smooth perpendicular surfaces has been much discussed, and it appears highly probable that the method by which this is accomplished is the exudation of moisture from the foot; there is still, however, much to be ascertained before the process can be satisfactorily comprehended. The theory to the effect that the method is the pressure of the atmosphere acting on the foot when the sole is in perfect apposition with the object walked on, or when a slight vacuum is created between the two, has apparently less to support it.

The legs of the young Insect are usually more simple than those of the adult, and in caterpillars they are short appendages, and only imperfectly jointed. If a young larva, with feet, of a beetle, such as _Crioceris asparagi_ be examined, it may be seen that the leg is formed by protuberance of the integument, which becomes divided into parts by simple creases; an observation suggesting that the more highly developed jointed leg is formed in a similar manner. This appears to be really the case, {107}for the actual continuity of the limb at the chief joint—the knee—can be demonstrated in many Insects by splitting the outer integument longitudinally and then pulling the pieces a little apart; while in other cases even this is not necessary, the knee along its inner face being membranous to a considerable extent, and the membrane continuous from femur to tibia.

Turning to the wings, we remark that there may be one or two pairs of these appendages. When there is but one pair it is nearly always mesothoracic, when there are two pairs one is invariably mesothoracic, the other metathoracic. The situation of the wing is always at the edge of the notum, but the attachment varies in other respects. It may be limited to a small spot, and this is usually the case with the anterior wing; or the attachment may extend for a considerable distance along the edge of the notum, a condition which frequently occurs, especially in the case of the posterior wings. The actual connexion of the wings with the thorax takes place by means of strong horny lines in them which come into very close relation with the little pieces in the thorax which we have already described, and which were styled by Audouin articulatory epidemes. There is extreme variety in the size, form, texture, and clothing of the wings, but there is so much resemblance in general characters amongst the members of each one of the Orders, that it is usually possible for an expert, seeing only a wing, to say with certainty what Order of Insects its possessor belonged to. We shall allude to these characters in treating of the Orders of Insects.

Each wing consists of two layers, an upper and a lower, and between them there may be tracheae and other structures, especially obvious when the wings are newly developed. It has been shown by Hagen that the two layers can be separated when the wings are recently formed, and it is then seen that each layer is traversed by lines of harder matter, the nervures. These ribs are frequently called wing-veins, or nerves, but as they have no relation to the anatomical structures bearing those names, it is better to make use of the term nervures. The strength, number, form and inter-relations of these nervures vary exceedingly; they are thus most important aids in the classification of Insects. Hence various efforts have been made to establish a system of nomenclature that shall be uniform throughout the different Orders, but at present success has not {108}attended these efforts, and it is probable that no real homology exists between the nervures of the different Orders of Insects. We shall not therefore discuss the question here. We may, however, mention that German savants have recently distinguished two forms of nervures which they consider essentially distinct, viz. convex and concave. These, to some extent, alternate with one another, but a fork given off by a convex one is not considered to be a concave one. The terms convex and concave are not happily chosen; they do not refer to the shape of the nervures, but appear to have been suggested by the fact that the surface of the wing being somewhat undulating the convex veins more usually run along the ridges, the concave veins along the depressions. The convex are the more important of the two, being the stronger, and more closely connected with the articulation of the wing.

The wings, broadly speaking, may be said to be three-margined: the margin that is anterior when the wings are extended is called the costa, and the edge that is then most distant from the body is the outer margin, while the limit that lies along the body when the wings are closed is the inner margin.

The only great Order of Insects provided with a single pair of wings is the Diptera, and in these the metathorax possesses, instead of wings, a pair of little capitate bodies called halteres or poisers. In the abnormal Strepsiptera, where a large pair of wings is placed on the metathorax, there are on the mesothorax some small appendages that are considered to represent the anterior wings. In the great Order Coleoptera, or beetles, the anterior wings are replaced by a pair of horny sheaths that close together over the back of the Insect, concealing the hind-wings, so that the beetle looks like a wingless Insect: in other four-winged Insects it is usually the front wings that are most useful in flight, but the elytra, as these parts are called in Coleoptera, take no active part in flight, and it has been recently suggested by Hoffbauer[25] that they are not the homologues of the front wings, but of the tegulae (see Fig. 56), of other Insects. In the Orthoptera the front wings also differ in consistence from the other pair over which they lie in repose, and are called tegmina. There are many Insects in which the wings {109}exist in a more or less rudimentary or vestigial condition, though they are never used for purposes of flight.

The abdomen, or hind body, is the least modified part of the body, though some of the numerous rings of which it is composed may be extremely altered from the usual simple form. Such change takes place at its two extremities, but usually to a much greater extent at the distal extremity than at the base. This latter part is attached to the thorax, and it is a curious fact that in many Insects the base of the abdomen is so closely connected with the thorax that it has all the appearance of being a portion of this latter division of the body; indeed it is sometimes difficult to trace the real division between the two parts. In such cases a further differentiation may occur, and the part of the abdomen that on its anterior aspect is intimately attached to the thorax may on its posterior aspect be very slightly connected with the rest of the abdomen. Under such circumstances it is difficult at first sight to recognise the real state of the case. When a segment is thus transferred from the abdomen to the metathorax, the part is called a median segment. The most remarkable median segment exists in those Hymenoptera which have a stalked abdomen, but a similar though less perfect condition exists in many Insects. When such a union occurs, it is usually most complete on the dorsal surface, and the first ventral plate may almost totally disappear: such an alteration may involve a certain amount of change in the sclerites of the next segment, so that the morphological determination of the parts at the back of the thorax and front of the abdomen is by no means a simple matter. A highly modified hind-body exists in the higher ants, Myrmicidae. In Fig. 60 we contrast the simple abdomen of _Japyx_ with the highly modified state of the same part in an ant.

Unlike the head and thorax, the abdomen is so loosely knitted together that it can undergo much expansion and contraction. {110}This is facilitated by an imbricated arrangement of the plates, and by their being connected by means of membranes admitting of much movement (Fig. 47, _m_, p. 88). In order to understand the structure of the abdomen it should be studied in its most distended state; it is then seen that there is a dorsal and a ventral hard plate to each ring, and there is also usually a stigma; there may be foldings or plications near the line of junction of the dorsal and ventral plates, but these margins are not really distinct pieces. The pleura, in fact, remain membranous in the abdominal region, contrasting strongly with the condition of these parts in the thorax. The proportions of the plates vary greatly; sometimes the ventral are very large in proportion to the dorsal, as is usually the case in Coleoptera, while in the Orthoptera the reverse condition prevails.

Cerci or other appendages frequently exist at the extremity of the abdomen (Fig. 47, _n_, p. 88); the former are sometimes like antennae, while in other cases they may be short compressed processes consisting of very few joints. The females of many Insects possess saws or piercing instruments concealed within the apical part of the abdomen; in other cases an elongate exserted organ, called ovipositor, used for placing the eggs in suitable positions, is present. Such organs consist, it is thought, either of modified appendages, called gonapophyses, or of dorsal, ventral, or pleural plates. The males frequently bear within the extremity of the body a more or less complicated apparatus called the genital armour. The term gonapophysis is at present a vague one, including stings, some ovipositors, portions of male copulatory apparatus, or other structures, of which the origin is more or less obscure.

The caterpillar, or larva, of the Lepidoptera and some other Insects, bears a greater number of legs than the three pairs we have mentioned as being the normal number in Insects, but the posterior feet are in this case very different from the anterior, and are called false legs or prolegs. These prolegs, which are placed on the hind body, bear a series of hooks in Lepidopterous larvae, but the analogous structures of Sawfly larvae are destitute of such hooks.

Placed along the sides of the body, usually quite visible in the larva, but more or less concealed in the perfect Insect, are little apertures for the admittance of air to the respiratory {111}system. They are called spiracles or stigmata. There is extreme variety in their structure and size; the largest and most remarkable are found on the prothorax of Coleoptera, especially in the groups Copridae and Cerambycidae.

The exact position of the stigmata varies greatly, as does also their number. In the Order Aptera there may be none, while the maximum number of eleven pairs is said by Grassi[26] to be attained in _Japyx solifugus_: in no other Insect have more than ten pairs been recorded, and this number is comparatively rare. Both position and number frequently differ in the early and later stages of the same Insect. The structure of the stigmata is quite as inconstant as the other points we have mentioned are.

The admission of air to the tracheal system and its confinement there, as well as the exclusion of foreign bodies, have to be provided for. The control of the air within the system is, according to Landois[27] and Krancher,[28] usually accomplished by means of an occluding apparatus placed on the tracheal trunk a little inside of the stigma, and in such case this latter orifice serves chiefly as a means for preventing the intrusion of foreign bodies. The occluding apparatus consists of muscular and mechanical parts, which differ much in their details in different Insects. Lowne supposes that the air is maintained in the tracheal system in a compressed condition, and if this be so, this apparatus must be of great importance in the Insect economy. Miall and Denny[29] state that in the anterior stigmata of the cockroach the valves act as the occluding agents, muscles being attached directly to the inner face of the valves, and in some other Insects the spiracular valves appear to act partially by muscular agency, but there are many stigmata having valves destitute of muscles. According to Lowne[30] there exist valves in the blowfly at the entrance to the trachea proper, and he gives the following as the arrangement of parts for the admission of air:—there is a spiracle {112}leading into a chamber, the atrium, which is limited inwardly by the occluding apparatus; and beyond this there is a second chamber, the vestibule, separated from the tracheae proper by a valvular arrangement. He considers that the vestibule acts as a pump to force the air into the tracheae.

SYSTEMATIC ORIENTATION.