(as it appears) of the epidermis, and that it performs the function of a spermatheca is shown by its containing spermatozoa, or, in Stuhlmannia, a spermatophore. In Polytoreutus, also, spermatophores have been found in these spermathecal sacs. We have thus the replacement of a spermatheca, corresponding to those of the remaining families of Oligochaeta, and derived, as is believed, from the epidermis, by a structure performing the same function, but derived from the mesoblastic tissues, and with a cavity which is coelom.
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| Fig. 13.—Female reproductive system of Hyperiodrilus.—XIII, XIV, thirteenth and fourteenth segments. | |
sp, Spermatheca. |
ov, Ovary. |
Alimentary Canal.—The alimentary canal is always a straight tube, and the anus, save in the genera Criodrilus and Dero, is completely terminal. A buccal cavity, a pharynx, an oesophagus and an intestine are always distinguishable. Commonly among the terrestrial forms there is a gizzard, or two gizzards, or a larger number, in the oesophageal region. There is no armed protrusible pharynx, such as exists in some other Chaetopods. This may be associated with mud-eating habits; but it is not wholly certain that this is the case; for in Chaetogaster and Agriodrilus, which are predaceous worms, there is no protrusible pharynx, though in the latter the oesophagus is thickened through its extent with muscular fibres. The oesophagus is often furnished with glandular diverticula, the “glands of Morren,” which are often of complex structure through the folding of their walls. Among the purely aquatic families such structures are very rare, and are represented by two caeca in the genus Limnodriloides. It is a remarkable fact, not yet understood, that in certain Enchytraeidae and Lumbriculidae the spermathecae open into the oesophagus as well as on to the exterior. The only comparable fact among other worms is the Laurer’s canal or genito-intestinal canal in the Trematoda. The intestine is usually in the higher forms provided with a typhlosole, in which, in Pontoscolex, runs a ciliated canal or canals communicating with the intestine. It is possible that this represents the syphon or supplementary intestine of Capitellidae, which has been shown to develop as a grooving of the intestine ultimately cut off from it. The intestine has a pair of caeca or two or three pairs (but all lie in one segment) in the genus Pheretima and in one species of Rhinodrilus. In Typhoeus and Megascolex there are complex glands appended to the intestine.
In Benhamia caecifera and at least one other earthworm there are numerous caeca, one pair to each segment.
Classification.—The classifications of Adolf Eduard, Grube and Claparède separated into two subdivisions the aquatic and the terrestrial forms. This scheme, opposed by many, has been reinstated by Sedgwick. The chief difficulty in this scheme is offered by the Moniligastridae, which in some degree combine the characters of both the suborders, into neither of which will they fit accurately. The following arrangement is a compromise:—
Group I. Aphaneura.—This group is referred by A. Sedgwick to the Archiannelida. It is, however, though doubtless near to the base of the Oligochaetous series, most nearly allied in the reproductive system to the Oligochaeta. It contains but one family, Aeolosomatidae. There are three pairs of spermathecae situated in segments III-V, a testis in V and an ovary in VI. There are a clitellum and sperm ducts which though like nephridia have a larger funnel and a less complexly wound duct. This family consists of only one well-known genus, Aeolosoma, which contains several species. They are minute worms with coloured oil drops (green, olive green or orange) contained in the epidermis. The nervous system is embedded in the epidermis, and the pairs of ganglia are separated as in Serpula, &c.; each pair has a longish commissure between its two ganglia. The intersegmental septa are absent save for the division of the first segment. The large prostomium is ciliated ventrally. The setae are either entirely capillary or there are in addition some sigmoid setae even with bifid free extremities. This genus also propagates asexually, like Ctenodrilus, which may possibly belong to the same family. Asexual reproduction universal.
Group II. Limicolae.—With a few exceptions the Limicolae are, as the name denotes, aquatic in habit. They are small to moderate-sized Oligochaeta, with a smaller number of segments than in the Terricolae. The alimentary canal is simple and a gizzard or oesophageal diverticula rarely developed. The vascular system is simple with as a rule direct communication between dorsal and ventral vessels in each segment. Nerve cord lies in coelom; brain in first segment or prostomium in many forms. Clitellum generally only two or three segments and more anterior in position than in Terricolae. Nephridia always paired and without plexus of blood capillaries. Spermatheca rarely with diverticula; sperm ducts as a rule occupying two segments only, usually opening by means of an atrium. Sperm sacs generally occupying a good many segments and with simple interior undivided by a network of trabeculae. Ova large and with much yolk. Asexual reproduction only in Naids. Egg sacs as large or nearly so as sperm sacs. Testes and ovaries always free. The following families constitute the group, viz. Naididae, Enchytraeidae, Tubificidae, Lumbriculidae, Phreoryctidae, Phreodrilidae, Alluroididae, the latter possibly not referable to this group.
Group III. Moniligastres.—Moderate-sized to very large Oligochaeta, terrestrial in habit, with the appearance of Terricolae. Generative organs anterior in position as in Limicolae. Sperm ducts and atria as in Limicolae; egg sacs large; body wall thick; vascular system and nephridia as in Terricolae. Only one family, Moniligastridae.
Group IV. Terricolae.—Earthworms, rarely aquatic in habit. Of small to very large size. Clitellum commonly extensive and more posterior in position than in other groups. Vascular system complicated without regular connexion between dorsal and ventral vessels, except in anterior segments. Nephridia as a rule with abundant vascular supply. Testes, and occasionally ovaries, enclosed in sacs. Sperm sacs generally limited to one or two segments with interior subdivided by trabeculae. Sperm ducts traverse several segments on their way to exterior. They open in common with, or near to, or, more rarely, into, glands which are not certainly comparable to the atria of the Limicolae. Egg sacs minute and functionless(?). Eggs minute with little yolk. Nephridia sometimes very numerous in each segment. Spermathecae often with diverticula.
Earthworms are divided into the following families, viz. Megascolicidae, Geoscolicidae, Eudrilidae, Lumbricidae.
As an appendix to the Oligochaeta, and possibly referable to that group, though their systematic position cannot at present be determined with certainty, are to be placed the Bdellodrilidae (Discodrilidae auct.), which are small parasites upon crayfish. These worms lay cocoons like the Oligochaeta and leeches, and where they depart from the structure of the Oligochaeta agree with that of leeches. The body is composed of a small and limited number of segments (not more than fourteen), and there is a sucker at each end of the body. There are no setae and apparently only two pairs of nephridia, of which the anterior pair open commonly by a common pore on the third segment after the head, whose segments have not been accurately enumerated. The intervening segments contain the genitalia, which are on the Oligochaeta plan in that the gonads are independent of their ducts and that there are special spermathecae, one pair. The male ducts are either one pair or two pairs, which open by a common and complicated efferent terminal apparatus furnished with a protrusible penis. The ganglia are crowded at the posterior end of the body as in leeches, and there is much tendency to the obliteration of the coelom as in that group. Pterodrilus and Cirrodrilus bear a few, or circles of, external processes which may be branchiae; Bdellodrilus and Astacobdella have none. The vascular system is as in the lower Oligochaeta. There are two chitinous jaws in the buccal cavity, a dorsal and a ventral, which are of specially complicated structure in Cirrodrilus.
Literature.—F. E. Beddard, A Monograph of the Oligochaeta (Oxford, 1895), also Quart. Journ. Micr. Sci., 1886–1895, and Proc. Zool. Soc., 1885–1906; W. B. Benham, Quart. Journ. Micr. Sci., 1886–1905; W. Michaelsen, “Oligochaeta” in Das Tierreich, 1900, and Mitth. Mus. (Hamburg, 1890–1906); A. G. Bourne, Quart. Journ. Micr. Sci., 1894; H. J. Moore, Journ. Morph., 1895; F. Vezhdovský, System d. Oligochaeten (Prague, 1884), and Entwicklungsgeschichtliche Untersuchungen; and numerous papers by the above and by G. Eisen, E. Perrier, D. Rosa, R. Horst, L. Cognetti, U. Pierantoni, W. Baldwin Spencer, H. Ude, &c., and embryological memoirs by R. S. Bergh, E. B. Wilson, N. Kleinenberg, &c.
Hirudinea.—The leeches are more particularly to be compared with the Oligochaeta, and the following definition embraces the main features in which they agree and disagree with that group. Setae are only present in the genus Acanthobdella. Eyes are present, but hardly so complex as in certain genera of Polychaetes. The appendages of the body are reduced to branchiae, present in certain forms. A clitellum is present. The segments of body are few (not more than thirty-four) and fixed in number. The anus is dorsal. One or two (anterior and posterior) suckers always present. Nervous system always in coelom. Coelom generally reduced to a system of tubes, sometimes communicating with vascular system; in Acanthobdella and Ozobranchus a series of metamerically arranged chambers as in Oligochaeta. Nephridia always paired, rarely (Pontobdella) forming a network communicating from segment to segment; lumen of nephridia always intracellular, funnels pervious or impervious. Alimentary
