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MILLIPEDE
469


as a single pair of antennae, two or three pairs of jaws and a variable number of walking-legs, of which one or more pairs may be transformed into gonopods. The antennae are short and very similar to the legs. They are preoral in position, and usually consist of seven segments, the seventh or distal segment being small, as a rule, and furnished with a sense organ which is probably olfactory or tactile in function. The mandibles or jaws of the first pair are the most anterior of the postoral appendages. They are large, powerful, and usually consist of three or two segments, a basal or cardo, which is sometimes absent, a second or stipes, and a third or mala, the latter being supplied with a strong tooth and pectinate lamellae. In all Diplopods, with the exception of the Pselaphognatha, there are only two pairs of jaws, those of the second pair forming a large plate, the gnathochilarium, which acts as a lower lip. It consists of several distinct sclerites, two external on each side, the proximal known as the cardo, the distal as the stipes, the latter being tipped with one or two lobes (malae) and far exceeding the cardo in size. Between the external plates there is a median proximal plate (mentum) generally of large size and often itself subdivided, and a pair of distal plates (linguae). Behind the base of the gnathochilarium there is a single large transverse plate, the hypostoma. In the Pselaphognatha, the jaws representing the gnathochilarium are differently constructed and an additional pair, the maxillulae, has been recently detected between the gnathochilarium and the mandibles. Behind the gnathochilarium, which from embryological data appears to result from the modification of a single pair of appendages, a legless somite has been detected in some embryos. Possibly the plate referred to above as the hypostoma is its sternal element.

   After Voges.
Fig. 3.—Inner view of ventral area of a single segment of Julus, much enlarged to show the structure and arrangement of the tracheal organs. The two pairs of tracheae are seen in situ, the posterior pair overlapping the anterior.
h, Posterior margin of the body-ring (tergum). t, Fine tracheae given off from it.
r, Anterior border. ms, Respiratory muscle attached to tracheal sac.
st, Tubular chamber of tracheae m, Ventral body muscle

The heart is a median dorsal vessel composed of a series of chambers each giving off a pair of arteries and furnished with a pair of orifices or ostia. According to Newport, the anterior chamber lying in the second segment is prolonged into an aortic trunk from which arise three pairs of lateral arteries dipping down on each side of the alimentary canal and uniting beneath it in a common ventral vessel. The heart is enveloped in a delicate pericardial membrane and is supported by lateral alary muscles. The alimentary canal is a simple tube extending usually straight through the body from mouth to anus. Only in the Oniscomorpha is it looped, thus suggesting the origin of this short-bodied group of millipedes from longer, more vermiform ancestors. A pair of so-called salivary glands opens into the fore-gut near its anterior extremity and one or two pairs of malpighian tubes communicate with the hind-gut at its junction with the broad mesenteric portion of the canal. Respiration is effected by means of tracheal tubes which communicate with the exterior by means of spiracles situated just above the bases of the walking limbs. Each spiracle leads into a longer or a shorter pouch whence the tracheae, which are of two kinds, arise. In the majority of the orders the tracheae are tufted, that is to say, they form two bundles of short simple tubules springing from the innermost corners of each pouch. In the Oniscomorpha, however, each pouch gives rise to a number of long tubes which extend through the body and somewhat resemble those of the Chilopoda except that they neither branch nor are extensive. As in the Chilopoda and Hexapoda the tracheae are strengthened and kept expanded by a slender spiral filament.

The ventral nerve cord consists of two strands so closely approximated as to be practically fused, with a small ganglionic enlargement for each pair of legs. Hence in the double segments there are two such ganglia, which in addition to the crural nerve give off on each side a large branching nerve to other organs in the segment. In the Opisthospermophora (Julus, Spirostreptus) and the Oniscomorpha (Glomeris, Sphaerotherium) the ganglia are spaced at equal distances on the cord, but in the Merochaeta (Polydesmus) they are grouped in pairs to correspond to the spacing of the legs. The apodous penultimate and anal segments are innervated from the last ganglion of the cord, as are also the gonopods of the males of the Oniscomorpha. The first (suboesophageal) ganglion of the cord supplies the mandibles and gnathochilarium and is connected by the oesophageal commissures with the bilobed cerebral nerve whence arises the nerves for the eyes, when present, and the antennae.

After G. C. Bourne, J. Linn. Soc. xix., Pl. 29, 1886.

Fig. 4.—Diagram of the nervous and circulatory system of Sphaerotherium obtusum, a South African species of Oniscomorpha.
c, Head.
oc, Eye-cluster.
ant, Antenna.
md, Basal segment of mandible.
tg2 and tg13, Part of the terga of the second and thirteenth segments.
cb, Cerebral ganglia supplying tr, Tracheal tubes with spiral the eyes and antennae.
oes, Oesophagus, cut through.
sb.gl, Suboesophageal or First ganglion of ventral chain.
gl2 and gl22, Second and twenty second ganglia of chain, the posterior nerve of each
 ganglion, lg.n, supplies the leg, the anterior, tr.n, the tracheal sac and other organs.
n.gon, Nerve to gonopods.
tr.s, Tracheal sac.

Eyes are sometimes absent, as in all the genera of Merochaeta and in many genera of other groups, as in Siphonophora, one of the Colobognatha, and several of the Juloidea (Typhloblaniulus). In other cases they are represented by one or two ocelli on each side (Stemmiuloidea); or by a vertical series of ocelli as in the Glomeroidea and Polyzonium amongst the Colobognatha. But in the majority of the orders they are represented by triangular or subspherical aggregations of ocelli recalling in a certain degree those of the Lithobiomorpha amongst the Chilopoda. They are simple in structure and consist externally of a cuticular corneal thickening or lens and internally of a retinular layer of enlarged epidermic cells, the