where this condition is usually not to be observed, there is embryological evidence that the existing state of affairs is derived from this. Commonly the nephridia are strictly paired a single pair to each segment, while the branches of the blood vascular system are similarly metameric. The alimentary canal is nearly always a straight tube running from the mouth, which is surrounded by the first segment of the body and overhung by the prostomium, to the anus, which is then either surrounded by the last segment of the body or opens dorsally a little way in front of this.
The Class as a Whole.—The Chaetopoda are with but few exceptions (Myzostomida in part, Sternaspis) elongated worms, flattened or, more usually, cylindrical, and bilaterally symmetrical. The body consists of a number of exactly similar or closely similar segments, which are never fused and metamorphosed, as in the Arthropoda, to form specialized regions of the body. It is, however, always possible to recognize a head, which consists at least of the peristomial segment with a forward projection of the same, the prostomium. A thorax also is sometimes to be distinguished from an abdomen. Where locomotive appendages (the parapodia of the Polychaeta) exist, they are never jointed, as always in the Arthropoda; nor are they modified anteriorly to form jaws, as in that group.
Fig. 1.—A, side view of the head region of Nereis cultrifera; | |
| E, Eye. M, Mouth. |
pl, Prostomial palp. pp, Parapodium. pr, Prostomium. |
The prostomium overhangs the mouth, and is often of considerable size and, as a rule, quite distinct from the segment following, being separated by an external groove, and containing, at least temporarily, the brain, which always arises there. Its cavity also is at first independent of the coelom though later invaded by the latter. In any case the cavity of the prostomium is single, and not formed, as is the cavity of the segments of the body, by paired coelomic chambers. It has, however, been alleged that this cavity is formed by a pair of mesoblastic somites (N. Kleinenberg), in which case there is more reason for favouring the view that would assign an equality between the prostomium and the (in that case) other segments of the body. The peculiar prostomium of Tomopteris is described below. The body wall of the Chaetopoda consists of a “dermo-muscular” tube which is separated from the gut by the coelom and its peritoneal walls, except in most leeches. A single layer of epidermic cells, some of which are glandular, forms the outer layer. Rarely are these ciliated, and then only in limited tracts. They secrete a cuticle which never approaches in thickness the often calcified cuticle of Arthropods. Below this is a circular, and below that again a longitudinal, layer of muscle fibres. These muscles are not striated, as they are in the Arthropoda.
Setae.—These chitinous, rod-like, rarely squat and then hook-like structures are found in the majority of the Chaetopoda, being absent only in certain Archiannelida, most leeches, and a very few Oligochaeta. They exist in the Brachiopoda (which are probably not unrelated to the Chaetopoda), but otherwise are absolutely distinctive of the Chaetopods. The setae are invariably formed each within an epidermic cell, and they are sheathed in involutions of the epidermis. Their shape and size varies greatly and is often of use in classification. The setae are organs of locomotion, though their large size and occasionally jagged edges in some of the Polychaeta suggest an aggressive function. They are disposed in two groups on either side, corresponding in the Polychaeta to the parapodia; the two bundles are commonly reduced among the earthworms to two pairs of setae or even to a single seta. On the other hand, in certain Polychaeta the bundles of setae are so extensive that they nearly form a complete circle surrounding the body; and in the Oligochaet genus Perichaeta (= Pheretima), and some allies, there is actually a complete circle of setae in each segment broken only by minute gaps, one dorsal, the other ventral.
Coelom.—The Chaetopoda are characterized by a spacious coelom, which is divided into a series of chambers in accordance with the general metamerism of the body. This is the typical arrangement, which is exhibited in the majority of the Polychaeta and Oligochaeta; in these the successive chambers of the coelom are separated by the intersegmental septa, sheets of muscle fibres extending from the body wall to the gut and thus forming partitions across the body. The successive cavities are not, however, completely closed from each other; there is some communication between adjoining segments, and the septa are sometimes deficient here and there. Thus in the Chaetopoda the perivisceral cavity is coelomic; in this respect the group contrasts with the Arthropoda and Molluscs, where the perivisceral cavity is, mainly at least, part of the vascular or haemal system, and agrees with the Vertebrata. The coelom is lined throughout by cells, which upon the intestine become large and loaded with excretory granules, and are known as chloragogen cells. Several forms of cells float freely in the fluid of the coelom. In another sense also the coelom is not a closed cavity, for it communicates in several ways with the external medium. Thus, among the Oligochaeta there are often a series of dorsal pores, or a single head pore, present also among the Polychaeta (in Ammochares). In these and other Chaetopods the coelom is also put into indirect relations with the outside world by the nephridia and by the gonad ducts. In these features, and in the fact that the gonads are local proliferations of the coelomic epithelium, which have undergone no further changes in the simpler forms, the coelom of this group shows in a particularly clear fashion the general characters of the coelom in the higher Metazoa. It has been indeed largely upon the conditions characterizing the Chaetopoda that the conception of the coelom in the Coelomocoela has been based.
Among the simpler Chaetopoda the coelom retains the character of a series of paired chambers, showing the above relations to the exterior and to the gonads. There are, however, further complications in some forms. Especially are these to be seen in the more modified Oligochaeta and in the much more modified Hirudinea. In the Polychaeta, which are to be regarded as structurally simpler forms than the two groups just referred to, there is but little subdivision of the coelom of the segments, indeed a tendency in the reverse direction, owing to the suppression of septa. Among the Oligochaeta the dorsal vessel in Dinodrilus and Megascolides is enclosed in a separate coelomic chamber which may or may not communicate with the main coelomic cavity. To this pericardial coelom is frequently added a gonocoel enclosing the gonads and the funnels of their ducts. This condition is more fully dealt with below in the description of the Oligochaeta. The division and, indeed, partial suppression of the coelom culminates in the leeches, which in this, as in some other respects, are the most modified of Annelids.
Nervous System.—In all Chaetopods this system consists of cerebral ganglia connected by a circumoesophageal commissure with a ventral ganglionated cord. The plan of the central nervous system is therefore that of the Arthropoda. Among the Archiannelida, in Aeolosoma and some Polychaetes, the whole central nervous system remains imbedded in the epidermis. In others, it lies in the coelom, often surrounded by a special and occasionally rather thick sheath. The cerebral ganglia constitute an archicerebrum for the most part, there being no evidence that, as in the Arthropoda, a movement forward of post-oral ganglia has taken place. In the leeches, however, there seems to be the commencement of the formation of a syncerebrum. In the latter, the segmentally arranged ganglia are more sharply marked off from the connectives than in other Chaetopods, where nerve cells exist along the whole ventral chain, though more numerous in segmentally disposed swellings.
Vascular System.—In addition to the coelom, another system of fluid-holding spaces lies between the body wall and the gut in the Chaetopoda. This is the vascular or haemal system (formerly and unnecessarily termed pseudhaemal). With a few exceptions among the Polychaeta the vascular system is always present among the Chaetopoda, and always consists of a system of vessels with definite walls, which rarely communicate with the coelom. It is in fact typically a closed system. The larger trunks open into each other either directly by cross branches, or a capillary system is formed. There are no lacunar blood spaces with ill-defined or absent walls except for a sinus surrounding the intestine, which is at least frequently present. The principal trunks consist of a dorsal vessel lying above the gut, and a ventral vessel below the gut but above the nervous cord. These two vessels in the Oligochaeta are united in the anterior region of the body by a smaller or greater number of branches which surround the oesophagus and are, some of them at least, contractile and in that case wider than the rest. The dorsal vessel also communicates with the ventral vessel indirectly by the intestinal sinus, which gives off branches to both the longitudinal trunks, and by tegementary vessels and capillaries which supply the skin and the nephridia. In the smaller and simpler forms the capillary networks are much reduced, but the dorsal and ventral vessels are usually present. The former, however, is frequently
