head. Below each eye is a cheek area (gena), often divided into an
anterior and a posterior part, while a distinct chin-sclerite (gula) is
often developed behind the mouth.
Feelers.—Most conspicuous among the appendages of the head are the feelers or antennae, which correspond to the anterior feelers (antennules) of Crustacea. In their simpler condition they are long and many-jointed, the segments bearing numerous olfactory and tactile nerve-endings. Elaboration in the form of the feelers, often a secondary sexual character in male insects, may result from a distal broadening of the segments, so that the appendage becomes serrate, or from the development of processes bearing sensory organs, so that the structure is pinnate or feather-like. On the other hand, the number of segments may be reduced, certain of them often becoming highly modified in form.
Jaws.—The mandibles of the Hexapoda are usually strong jaws with one or more teeth at the apex (fig. 1, A, B, mn), articulating at their bases with the head-capsule by sub-globular condyles, and provided with abductor and adductor muscles by means of which they can be separated or drawn together so as to bite solid food, or seize objects which have to be carried about. They never bear segmented limbs (palps) and only exceptionally (as in the chafers) is the skeleton composed of more than one sclerite. The mandibles often furnish a good example of “secondary sexual characters,” being more strongly developed in the male than in the female of the same species. In most insects that feed by suction the mandibles are modified. In bugs (Heteroptera) and many flies, for example, they are changed into needle-like piercers (fig. 2, II), while in moths and caddis-flies they are reduced to mere vestiges or altogether suppressed.
As previously mentioned, a pair of minute jaws—the maxillulae—are present in the lowest order of insects, between the mandibles and the first maxillae. They usually consist of an inner and an outer lobe arising from a basal piece, which bears also in some genera a small palp (see Aptera).
In their typical state of development, the first maxillae offer a striking contrast to the mandibles, being composed of a two-segmented basal piece (cardo and stipes, fig. 1, C, ca, st) bearing a distinct inner and outer lobe (lacinia and galea, fig. 1, C, la, ga) and externally a jointed limb or palp (fig. 1, C, pa). Such maxillae are found in most biting insects. In insects whose mouths are adapted for sucking and piercing, remarkable modifications may occur. In many blood-sucking flies, for example, the galea is absent, while the lacinia becomes a strong knife-like piercer and the palp is well developed. In bugs and aphids the lacinia is a slender needle-like piercer (fig. 2, III), while the palp is wanting. In butterflies and moths the lacinia is absent while the galea becomes a flexible process, grooved on its inner face, so as to make with its fellow a hollow sucking-trunk, and the palp is usually very small.
The second pair of maxillae are more or less completely fused together to form what is known as the labium or “lower lip.” In generalized biting insects, such as cockroaches and locusts (Orthoptera), the parts of a typical maxilla can be easily recognized in the labium. The fused cardines form a broad basal plate (sub-mentum) and the stipites a smaller plate (mentum)—see fig. 1, C, sm, m—jointed on to the sub-mentum, while the galeae, laciniae and palps remain distinct. In specialized biting insects, such as beetles (Coleoptera), the labium tends to become a hard transverse plate bearing the pair of palps, a median structure—known as the ligula—formed of the conjoined laciniae, and a pair of small rounded processes—the reduced galeae—often called the “paraglossae,” a term better avoided since it has been applied also to the maxillulae of Aptera, entirely different structures. The long sucking “tongue” of bees is probably a modification of the ligula. In bugs and aphids (Hemiptera), the fused second maxillae form a jointed grooved beak or rostrum (fig. 2, IV) in which the slender piercers (mandibles and first maxillae) work to and fro.
This second pair of maxillae (or labium) form then the hinder or lower boundary of the mouth. In front or above the mouth is bounded by the labrum, while the mandibles and first maxillae lie on either side of it. A median process, known as the hypopharynx or tongue, arises from the floor of the mouth in front of the labium, and becomes most variously developed or specialized in different insects. The salivary duct opens on its hinder surface. It does not appear to represent a pair of appendages, but the maxillulae of the Aptera become closely associated with it. According to the view of R. Heymons, the hypopharynx represents the sterna of all the jaw-bearing somites, but other students consider that it belongs to the mandibular and first maxillary segments, or entirely to the segment of the first maxillae.
Neck.—The head is usually connected with the thorax by a distinct membranous neck, strengthened in the more generalized orders with small chitinous plates (cervical sclerites). These have been interpreted as indicating one or more primitive segments between the head and thorax. Probably, however, as suggested by T. H. Huxley (Anat. Invert. Animals, 1877), they really belong to the labial segment which has not become completely fused with the head-capsule. It has been shown by C. Janet (1889), from careful studies of the musculature, that the greater part of the head-capsule is built up of the four anterior head-segments, the hindmost of which has the mandibles for its appendages, and this conclusion is in the main supported by the recent work on the head skeleton of J. H. Comstock and C. Kochi (1902) and W. A. Riley (1904).
Thorax.—The three segments which make up the thorax or fore-trunk are known as the prothorax, mesothorax and metathorax (see fig. 3). The dorsal area of the prothorax is occupied by a single sclerite, the pronotum (fig. 3, d), which is large and conspicuous in those insects, such as cockroaches, bugs (Heteroptera) and beetles, which have the prothorax free—i.e. readily movable on the segment (mesothorax) immediately behind—smaller and of less importance where the prothorax is fixed to the mesothorax, as in bees and flies. The dorsal area of the mesothorax, and also of the metathorax, may be made up of a series of sclerites arranged one behind the other—prescutum, scutum, scutellum and post-scutellum (fig. 3, e, f, g, h), the scutellum of the mesothorax being often especially conspicuous. Ventrally, each segment of the thorax has a sternum with which a median pre-sternum and paired episterna and epimera are often associated (see figs. 3, 4). The recent suggestion of K. W. Verhoeff (1904) that the hexapodan thorax in reality contains six primitive segments is entirely without embryological support.
Legs.—Each segment of the thorax carries a pair of legs. In most insects the leg is built up of nine segments: (1) a broad triangular, sub-globular, conical or cylindrical haunch (coxa); (2) a small trochanter; (3) an elongate stout thigh (femur); (4) a more slender shin (tibia); and (5-9) a foot consisting of five tarsal segments. The fifth (distal) tarsal segment carries a median adhesive pad—the pulvillus—on either side of which is a claw. The pulvillus is