canal sometimes with protrusible proboscis; never with gizzard or oesophageal glands; intestine with caeca as a rule. Jaws often present. Testes several pairs, rarely one pair, continuous with sperm ducts; ovaries, one pair, continuous with oviducts; generative pores single and median. No separate spermathecae or septal chambers for the development of the ova and sperm. Eggs deposited in a cocoon. Development direct. No asexual generation. Fresh-water, marine and terrestrial. Parasitic or carnivorous.
In external characters the Hirudinea are unmistakable and not to be confused with other Annelids, except perhaps with the Bdellodrilidae, which resemble them in certain particulars. The absence of setae—save in Acanthobdella, where five of the anterior segments possess each four pairs of setae with reserve setae placed close behind them (fig. 14), and the presence of an anterior and posterior sucker, produce a looping mode of progression similar to that of a Geometrid larva. The absence of setae and the great secondary annulation render the mapping of the segments a subject of some difficulty. The most reliable test appears to be the nerve ganglia, which are more distinct from the intervening connectives than in other Annelids.
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| Fig. 14.—Acanthobdella, from the ventral surface, showing the five sets of setae (S1 to S5) and the replacing setae (Sr) behind them. The three pairs of pigmented spots show the position of the eyes on the dorsal surface. (After Kovalevsky.) |
In the middle of the body, where the limits of the somites can be checked by a comparison with the arrangement of the nephridia and the gonads, and where the ganglia are quite distinct and separated by long connectives, each ganglion is seen to consist of six masses of cells enclosed by capsules and to give off three nerves on each side. This corresponds to the usual presence (in the Rhynchobdellidae) of three annuli to each segment. Anteriorly and posteriorly separate ganglia have fused. The brain consists not only of a group of six capsules corresponding to the archicerebrum of the Oligochaeta, but of a further mass of cells surrounding and existing below the alimentary canal, which can be analysed into five or six more separate ganglia. The whole mass lies in the seventh or eighth segment. At the posterior end of the body there are likewise seven separate ganglia partially fused to form a single ganglionic mass, which innervates the segments lying behind the anus and corresponding to the posterior sucker. So that a leech in which only twenty-seven segments are apparent by the enumeration of the annuli, separate ganglia, nephridia, lines of sensillae upon the body, really possesses an additional seven lying behind that which is apparently the last of the series and crowded together into a minute space. The annuli into which segments are externally divided are so deeply incised as to render it impossible to distinguish, as can be readily done in the Oligochaeta as a rule, the limits of an annulus from that of a true segment. As remarked, the prevalent number of annuli to a segment is three in the Rhynchobdellidae. But in that group (Cystobranchus) there may be as many as eight annuli. In the Gnathobdellidae the prevailing number of annuli to a segment is five; but here again the number is often increased, and Trocheta has no less than eleven. The reason for this excessive annulation has been seen in the limited number of segments (thirty-four) of which the body is composed, which are laid down early and do not increase. In the Oligochaeta, on the other hand, there is growth of new segments. It is important to notice that the metameric plan of growth of Chaetopods is still preserved.
The nephridia are like those of the Oligochaeta in general structure; that is to say, they consist of drain-pipe cells which are placed end to end and are perforated by their duct. The internal funnel varies in the same way as in the Oligochaeta in the number of cells which form it. In Clepsine (Glossiphonia) there are only three cells, and in Nephelis five to eight cells. In Hirudo the funnel is not pervious and is composed of a large number of cells. Externally, the nephridium opens by a vesicle, as in many Oligochaetes whose lumen is intercellular. In Pontobdella and Branchellion the nephridia form a network extending from segment to segment, but there is only one pair of funnels in each segment. Slight differences in form have been noted between nephridia of different segments; but the Hirudinea do not show the marked differentiation that is to be seen in some other Chaetopods; nor do the nephridia ever acquire any relations to the alimentary canal.
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Fig. 15.—Section of Acanthobdella (after Kovalevsky). | |
| c, Coelom. c.ch, Coelomic epithelium (yellow-cells). cg, Glandular cells. cl, Muscle cells of lateral line. cp, Pigment cells. ep, Ectoderm. |
g, Nerve cord. m, Intestine. mc, Circular muscle. ml, Longitudinal muscle. vd, Dorsal vessel. vv, Ventral vessel. |
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Fig. 16.—Section of Acanthobdella (after Kovalevsky). Identical letters as in fig. 2; in addition, cn, nerve cord; in, intestine; nf, parts of nephridium; on, external opening of nephridium; ov, ova; t, testis. |
Coelom.—The coelom of the Hirudinea differs in most genera from that of the Oligochaeta and Polychaeta. The difference is that it is broken up into a complex sinus system. The least modified type is shown by Acanthobdella, a leech, parasitic upon fishes, in which transverse sections (see figs. 15 and 16) show the gut, the nervous system, &c., lying in a spacious chamber which is the coelom. This coelom is lined by peritoneal cells and is divided into a series of metameres by septa which correspond to the segmentation of the body, the arrangement being thus precisely like that of typical Chaetopoda. Moreover, upon the intestine the coelomic cells are modified into chloragogen cells. In Acanthobdella the testes are, however, not contained in the general coelom, and the nephridia lie in the septa. It is remarkable, in view of the spaciousness of the coelom, that the funnels of the latter have not been seen. Ozobranchus possesses a coelom which is less typically chaetopodous than that of Acanthobdella, but more so than in other leeches. There is a spacious cavity surrounding the gut and containing also blood-vessels, and to some extent the generative organs, and the nervous cord. Furthermore, in the mid region of the body this coelom is broken up by metamerically arranged septa, as in Acanthobdella. These septa are, however, rather incomplete and are not fastened to the gut; and, as in Acanthobdella, the nephridia are embedded in them. In addition to the median lacuna there are two lateral lacunae, one upon each side. These regions of the coelom end at the ends of the body and communicate with each other by means of a branched system of coelomic sinuses, which are in places very fine tubes. Neither in this genus nor in the last is there any communication between coelom and vascular system. In Clepsine (Glossiphonia) there is a further breaking up of the coelom. The median lacuna no longer exists, but is represented by a dorsal and ventral sinus. The former lodges the dorsal, the latter the ventral, blood-vessel. The gut has no coelomic space surrounding it. A complex
