rank as distinct classes of Arthropods. Pocock, indeed, definitely associated the Chilopoda with the Hexapoda in a group, the Opisthogoneata (Opisthogonea), equivalent to a group, the Progoneata (Prosogonea), comprising the Diplopoda, Pauropoda and Symphyla. As the basis for this classification was taken the position of the generative orifices which open in the Opisthogonea at the posterior end and in the Prosogonea near the anterior end of the body. As a matter of fact, in the Chilopoda they are situated on the penultimate or pretelsonic somite; in the Hexapoda upon the antepenultimate somite (male) or a little farther forward (female). Moreover, the recent researches of Heymons into the embryology of Scolopendra, one of the Chilopods, has shown a close correspondence in the number of cephalic metameres between the Chilopoda and Hexapoda, a correspondence which has not yet been established in the case of the Diplopoda or Symphyla. This last discovery bears out the view of relationship between the centipedes and insects, to the exclusion of the Diplopoda, Symphyla and Pauropoda. But even if in the future it can be shown that all these groups can be brought into line with respect to the metamerism of the head, the position of the generative orifices will remain as a fundamental and constant character, distinguishing the Chilopoda from the other groups of so-called “Myriapods” and the Hexapoda from the Symphyla, which in many particulars they resemble.
Structure of the Chilopoda.—The exoskeletal elements of a typical somite consist of a dorsal plate or tergum, a ventral plate or sternum, a lateral or pleural membrane, often strengthened with chitinous sclerites, and a pair of appendages. At the anterior extremity there is a head-shield or cephalite, which bears eyes, when present, and a pair of antennae. In all centipedes, except the Scutigeridae, the preantennal portion of the cephalite is sharply reflexed, ventrally forming an area called the clypeus. The inferior edge of this bears the labrum, which is usually represented by a small median, and two large lateral plates. The appendages are modified as a single pair of antennae, four pairs of jaws or gnathites, a variable number of walking legs and a single pair of generative limbs or gonopods. The antennae, articulated to the forepart of the head and preoral in position, are long and flexible and consist of fourteen or more segments. The jaws of the first pair of mandibles are stout and bi-segmented, with a dentate cutting edge. Those of the second pair or maxillae vary considerably in structure in different groups. They are foliaceous and are usually regarded as biramous. In some genera (Scutigera, Lithobius) the inner branch consists of two distinct segments meeting those of the opposite side in the middle line. The outer branch, which is always larger, consists of three or four segments. Generally, however, the basal segments of the two branches are coalesced with each other and with the corresponding segments of the opposite side to form a single broad transverse plate. The above described condition seen in Scutigera suggests that two pairs of jaws may be involved in the formation of the maxillae in the Chilopoda. The jaws of the third pair, the palpognaths or second pair of maxillae, resemble dwarfed walking legs, and consist of five or six segments, of which the basal or coxa is united mesially to its fellow. The jaws of the fourth pair, the toxicognaths or poison-jaws, are long and powerful, and consist like the legs primarily of six segments, whereof the basal is large and usually fused with its fellow to form a large coxal plate, the second is small and generally suppressed by fusion with the third, the fourth and fifth are also small, while the sixth is transformed into a great piercing fang, at the tip of which opens the duct of a poison gland lodged within the appendage.
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Modified from Heymons, Bib. Zool., 1901, by permission of E. Nagele.
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| Fig. 1. |
| A, Diagram of anterior extremity of an early embryo of Scolopendra, ventral view; cl, clypeus; lb, labrum; m, mouth; p.a, preantennal appendage; a, antenna; int, premandibular rudiment; mdl, mandible; mx, maxilla; p.g, palpognath; t.g, toxicognath; lg. 1, first pair of walking legs. B, Posterior end of a later embryo of Scolopendra, ventral view, showing the anal segment or telson (t); the legs of the last pair in the adult (lg. 21) and the two rudimentary pairs of legs (lg. 22, lg. 23). |
The tergal elements of the somites bearing the antennae, mandibles and maxillae appear to be represented by the head-shield or cephalite. The tergal element of the somite bearing the palpognath is usually suppressed; that of the toxicognath is sometimes of large size as in some Geophilomorpha (Himantarium), sometimes small as in Scutigera, Lithobius, Craterostigmus, sometimes suppressed probably by fusion with the tergum of the first leg-bearing somite as in the Scolopendromorpha. The sternal plates of all the jaw-bearing somites have disappeared, except in the case of the somite of the toxicognath, where it may be vestigial. In the case of the somites bearing the walking legs the tergal and sternal elements are preserved without fusion with the corresponding plates of the preceding or succeeding somites, so that great flexibility of the body is retained. The only exception to this is presented by Scutigera, where the terga corresponding to the somites bearing the fifteen pairs of legs are reduced by fusion and suppression to seven. The walking legs are articulated to the inferior portion of the pleural or lateral area of the somites close to the external margins of the sterna, which widely separate those of the left from those of the right side. Generally speaking the legs resemble each other, although as a rule they progressively increase in length towards the posterior end of the body. They consist typically of six segments, of which the basal is termed the coxa and the apical the tarsus. The tarsus is armed with a single terminal claw, and, except in the Geophilomorpha and a few genera of other orders, is divided by a mesial transverse joint into two segments, as is the case in Scolopendra and Lithobius for example. But in some of the longer-legged, swift-footed centipedes of the order Lithobiomorpha (e.g. Henicops, Cermatobius) the tarsi are further subdivided. The multiplication of sub-segments reaches its maximum in Scutigera, where the tarsi are extremely long, slender, flexible and annulated. The legs of the last pair are directed backwards in a line parallel with the long axis of the body, so that their coxae, fused in some cases with the pleural sclerites (Scolopendra, Geophilus), or free and of large size (Scutigera, Lithobius), serve to protect the small genital and anal somites. They are often greatly modified. In the males of some species of Lithobius one or more of the segments is inflated or furnished with tubercle-bearing, tactile bristles; in some Geophilomorpha the whole limb is thickened in the male sex. In most Scolopendromorpha the basal segment is armed beneath with spines or spikes (Dacetum, Scolopocryptops); sometimes the whole appendage is thickened and terminated by a sharp and serrate claw (Theatops, Plutonium). In these cases the legs act as weapons of defence and offence. In other cases (Newportia) the tarsi lose the claw, become many-jointed and act as feelers, while in Alipes the terminal segments are flattened, leaf-like and furnished with a peculiar stridulating organ. The genital somite is always small and sometimes retractile within the somite bearing the last pair of legs. Its tergal plate is usually retained, but its sternal plate is generally suppressed. In females of the Lithobiomorpha and Scutigeromorpha the appendages of this somite—the gonopods—are jointed, forcipate and relatively well developed although small. In the females of the other orders they are greatly reduced or absent. In the males their development varies considerably. They are well developed in Scutigera, where they form two pairs of digitiform sclerites, whereas in the Geophilomorpha they are reduced to a pair of very short, two-jointed limbs. The anal somite is always small and limbless. In Craterostigmus the genital and anal somites are represented by a pair of elongate valves projecting between the legs of the last pair. The structure of the gonopods is unknown, and the homology between the two valves and the skeletal elements of the somites in question not clearly understood.
A study of the development of Scolopendra has shown that the antennae of the adult are the appendages of the second postoral metamere and the mandibles those of the fourth, the first postoral metamere, which has a pair of transient preantennal appendages, and the third, which has no appendages, being excalated at an early stage of embryonic growth. Furthermore, behind the legs of the last pair two pairs of appendages are present. The second of these persists as the gonopods of the adult, but the first is suppressed. Possibly, however, it is represented in the male of Scutigera by the anterior branches of the gonopods. The cerebral or cephalic portion of the nervous system consists of a quadrilobate mass. From the two upper lobes, which are set transversely, arise the ocular nerves; from the two lower lobes, which are united by a transverse commissure, spring the antennal nerves in front and the chords which form the oesophageal collar behind. These chords unite below the oesophagus to form the compound suboesophageal ganglion, whence the nerves for the four pairs of jaws arise. The ventral system consists of a double chord uniting in each of the leg-bearing segments in a ganglionic swelling which gives off four pairs of nerves to the limbs and tissues of the somite. There is a single ganglion in the genital segment.
Eyes are frequently absent. When present they may be either simple or compound, i.e. consisting externally of a single lens (monomeniscous) of or an aggregation of lenses (polymeniscous). Simple eyes vary in number on each side of the head from one, as in Henicops, to many as forty, as in some species of Lithobius. In
