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CHAETOPODA

to be compared with similarly-placed cells in the nephridia of Amphioxus.

More usually, and indeed in nearly every other case among the Oligochaeta and Hirudinea, the coelomic aperture of the nephridium consists of several cells, ciliated like the nephridium itself for a greater or less extent, forming a funnel. The funnel varies greatly in size and number of its component cells. There are so many differences of detail that no line can be drawn between the one-celled funnel of Aeolosoma and the extraordinarily large and folded funnel of the posterior nephridia in the Oligochaete Thamnodrilus. In the last-mentioned worm the funnels of the anterior nephridia are small and but few celled; it is only the nephridia in and behind the 17th segment of the body which are particularly large and with a sinuous margin, which recall the funnels of the gonad ducts (i.e. coelomoducts).

Among the Polychaeta the nephridium of Nereis (see fig. 2) is like that of the Oligochaeta and Hirudinea in that the coiled glandular tube has an intracellular duct which is ciliated in the same way in parts. The Polychaeta, however, present us with another form of nephridium seen, for example, in Arenicola, where a large funnel leads into a short and wide excretory tube whose lumen is intercellular. In the young stages of this worm which have been investigated by W. B. Benham, the tube, though smaller, and with a but little pronounced funnel, has still an intercellular duct. That these organs in Polychaeta serve for the removal of the generative products to the exterior is proved not only by the correspondence in number to them of the gonads, but by actual observation of the generative products in transit. This form of nephridia leads to the shorter but essentially similar organs in the Polychaete Sternaspis, and to those of the Echiuroidea (q.v.) and of the Gephyrea (q.v.).

Though the paired arrangement of the nephridia is the prevalent one in the Chaetopoda, there are many examples, among the Oligochaeta, of species and genera in which there are several, even many, nephridia in each segment of the body, which may or may not be connected among themselves, but have in any case separate orifices on to the exterior.

2. Coelomoducis.—In this category are included (by Goodrich and Lankester) the gonad ducts of the Oligochaeta, certain funnels without any aperture to the exterior that have been detected in Nereis, &c., funnels with wide and short ducts attached to nephridia in other Polychaeta, gonad ducts in the Capitellidae, the gonad ducts of the leeches. In all these cases we have a duct which has a usually wide, always intercellular, lumen, generally, if not always, ciliated, which opens directly into the coelom on the one hand and on to the exterior of the body on the other. These characters are plain in all the cases cited, excepting only the leeches which will be considered separately.

There is not a great deal of difference between most of these structures and true nephridia. It is not clear, for example, to which category it is necessary to refer the excretory organs of Arenicola, or Polynoe. Both series of organs consist essentially of a ciliated tube leading from the coelom to the exterior. Both series of organs grow back centrifugally from the funnel. In both the cavity originally or immediately continuous with the coelom appears first in the funnel and grows backwards. In some cases, e.g. oviducts of Oligochaeta, sperm ducts of Phreoryctes, the coelomoducts occupy, like the nephridia, two segments, the funnel opening into that in front of the segment which carries the external pore. It is by no means certain that a hard and fast line can be drawn between intra- and intercellular lumina. Finally, in function there are some points of likeness. The gonad ducts of Lumbricus, &c., must perform one function of nephridia; they must convey to the exterior some of the coelomic fluid with its disintegrated products of waste. There is no possibility that sperm and ova can escape by these tubes not in company with coelomic fluid. In the case of many Oligochaeta where there is no vascular network surrounding the nephridium, this function must be the chief one of those glands, the more elaborate process of excretion taking place in the case of nephridia surrounded by a rich plexus of blood capillaries. A consideration of the mode of development and appearance of the coelomoducts that have thus far been enumerated (with the possible exception of those of the leeches) seems to show that there is a distinct though varying relation between them and the nephridia. It has been shown that in Tubifex, and some other aquatic Oligochaeta, the genital segments are at first provided with nephridia, and that these disappear on the appearance of the generative ducts, which are coelomoducts. In Lumbricus the connexion is a little closer; the funnel of the nephridium, in the segments in which the funnels of the gonad ducts are to be developed, persists and is continuous with the gonad duct funnels on their first appearance. In the development of the Acanthodrilid earthworm Octochaetus (F. E. Beddard) the funnels of the pronephridia disappear except in the genital segments, where they seem to be actually converted into the genital funnels. At the least there is no doubt that the genital funnels are developed precisely where the nephridial funnels formerly existed. If the genital funnels are not wholly or partly formed out of the nephridial funnels they have replaced them. In the genital segments of Eudrilus the nephridia are present, but the funnels have not been found though they are obvious in other segments. Here also the genital funnels have either replaced or been formed out of nephridial funnels. In Haplotaxis heterogyne (W. B. Benham) the sperm ducts are hardly to be distinguished from nephridia; they are sinuous tubes with an intra-cellular duct. But the funnel is large and thus differs from the funnels of the nephridia in adjoining segments. Here again the nephridial funnel seems to have been converted into or certainly replaced by a secondarily developed funnel. This example is similar to cases among the Polychaeta where a true nephridium is provided with a large funnel, coelomostome, according to the nomenclature of Lankester. The whole organ, having, as is thought but not known, this double origin, is termed a nephromixium. The various facts, however, seem to be susceptible of another interpretation. It may be pointed out that the several examples described recall a phenomenon which is not uncommon and is well known to anatomists. That is the replacement of an organ by, sometimes coupled with its partial conversion into, a similar or slightly different organ performing the same or an analogous function. Thus the postcaval vein of the higher vertebrata is partly a new structure altogether, and is partly formed out of the pre-existing posterior cardinals. The more complete replacements, such as the nephridia of the genital segment of Tubifex by a subsequently formed genital duct, may be compared with the succession of the nesonephros to the pronephros in vertebrates, and of the metanephros to the mesonephros in the higher vertebrates. It might be well to term these structures, mostly serving as gonad ducts, which have an undoubted resemblance to nephridia, and for the most part an undoubted connexion with nephridia, “Nephrodinia,” to distinguish them from another category of “ducts” which are communications between the coelom and the exterior, and which have no relation whatever to nephridia or to the organs just discussed. For these latter, the term coelomoducts might well be reserved. To this category belong certain sacs and pouches in many, perhaps most, genera of the Oligochaeta family, Eudrilidae, and possibly the gonad ducts in the Hirudinea. As an example of the former it has been shown (Beddard) that a large median sac in Lybiodrilus is at first freely open to the coelom, that it later becomes shut off from the same, that it then acquires an external orifice, and, finally, that it encloses the ovary or ovaries, between which and the exterior a passage is thus effected. To this category will belong the oviducts in Teleostean fishes and probably the gonad ducts in several groups of invertebrates.

Polychaeta.—This group may be thus defined and the definition contrasted and compared with those of the other divisions of the Chaetopoda. Setae always present and often very large, much varied in form and very numerous, borne by the dorsal and ventral parapodia (when present). The prostomium and the segments generally often bear processes sensory and branchial. Eyes often present and comparatively complicated in structure. Clitellum not present as a definite organ, as in Oligochaeta. The anus is mostly terminal, and there are no anterior and posterior suckers. Nervous system often imbedded in the epidermis. Vascular system generally present forming a closed system of tubes. Alimentary canal rarely coiled, occasionally with glands which are simple caeca and sometimes serve as air reservoirs; jaws often present and an eversible pharynx. Nephridia sometimes of the type of those of the Oligochaeta; in other cases short, wide tubes with a large funnel serving also entirely or in part as gonad ducts. Frequently reduced in number of pairs; rarely (Capitellidae) more than one pair per segment. Gonads not so restricted in position as in Oligochaets, and often more abundant; the individuals usually unisexual. No specialized system of spermathecae, sperm reservoirs, and copulatory apparatus, as in Oligochaeta; development generally through a larval form; reproduction by budding also occurs. Marine (rarely fresh-water) in habit.

The Polychaeta contrast with the Oligochaeta by the great variety of outward form and by the frequency of specialization of different regions of the body. The head is always recognizable and much more conspicuous than in other Chaetopoda. As in the Oligochaeta the peristomial segment is often without setae, but this character is not by any means so constant as in the Oligochaeta. The prostomium bears often processes, both dorsal and ventral, which in the Sabellids are split into the circle of branchial plumes, which surround or nearly surround the mouth in those tube-dwelling Annelids. Tomopteris is remarkable for the fact that the hammer-shaped prostomium has paired ventral processes each with a single seta. It is held, however, that these are a pair of parapodia which have shifted forwards. The presence of parapodia distinguish this from other groups of Chaetopoda. Typically, the parapodium consists of two processes of the body on each side, each of which bears a bundle of setae; these two divisions of the “limb” are termed