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795
CHAETOPODA

haemoglobin. (4) Terebelliformia. These worms are in some respects like the Sabellids (Cryptocephala). The parapodia, as in the Capitellidae, are hardly developed. The buccal region is unarmed and not eversible. The prostomium has many long filaments which recall the gills of the Sabellids, &c. The nephridia are specialized into two series, as in the last-mentioned worms. (5) Spioniformia (including Chaetopterus, Spio, &c.) and (6) Scoleciformia (Arenicola, Chloraema, Sternaspis) are the remaining groups. In both, the nephridia are all alike; there are no jaws; the prostomium rarely has processes. The body is often divisible into regions.

Literature.—W. B. Benham, “Polychaeta” in Cambridge Natural History; E. Claparède, Annélides chétopodes du golfe de Naples (1868 and 1870); E. Ehlers, Die Börstenwürmer (1868); H. Eisig, Die Capitelliden (Naples Monographs), and development of do. in Mitth. d. zool. Stat. Neapel (1898); W. C. M‘Intosh, ”Challenger” Reports (1885); E. R. Lankester, Introductory Chapter in A Treatise on Zoology; E. S. Goodrich, Quart. Journ. Mic. Sci. (1897–1900); E. Meyer, Mitth. d. zool. Stat. Neapel (1887, 1888), as well as numerous other memoirs by the above and by J. T. Cunningham, de St Joseph, A. Malaquin, A. Agassiz, A. T. Watson, Malmgren, Bobretsky and A. F. Marion, E. A. Andrews, L. C. Cosmovici, R. Horst, W. Michaelsen, G. Gilson, F. Buchanan, H. Levinsen, Joyeux-Laffuie, F. W. Gamble, &c.

EB1911 Chaetopoda Fig. 10.—Diagrams of various Earthworms, to illustrate external characters.jpg
Fig. 10.—Diagrams of various Earthworms, to illustrate external characters. A, B, C, anterior segments from the ventral surface; D, hinder end of body of Urochaeta.

A, Lumbricus: 9, 10, segments containing spermathecae, the orifices of which are indicated; 14, segment bearing oviducal pores; 15, segment bearing male pores; 32, 37, first and last segments of clitellum.

B, Acanthodrilus: cp, orifices of spermathecae; Female symbol.svg, oviducal pores; Male black symbol.svg, male pores; on 17th and 19th segments are the apertures of the atria.

C, Perichaeta: the spermathecal pores are between segments 6 and 7, 7 and 8, 8 and 9, the oviducal pores upon the 14th and the male pores upon the 18th segment.

In all the figures the nephridial pores are indicated by dots and the setae by strokes.

Oligochaeta.—As contrasted with the other subdivisions of the Chaetopoda, the Oligochaeta may be thus defined. Setae very rarely absent (genus Achaeta) and as a rule not so large or so numerous in each segment as in the Polychaeta, and different in shape. Eyes rarely present and then rudimentary. Prostomium generally small, sometimes prolonged, but never bearing tentacles or processes. Appendages of body reduced to branchiae, present only in four species, and to the ventral copulatory appendages of Alma and Criodrilus. Clitellum always present, extending over two (many limicolous forms) to forty-five segments (Alma). Segments of body numerous and not distinctive of species, being irregular and not fixed in numbers. In terrestrial forms dorsal pores are usually present; in aquatic forms a head pore only. Anus nearly always terminal, rarely dorsal, at a little distance from end of body. Suckers absent. Nervous system rarely (Aeolosoma) in continuity with epidermis. Vascular system always present, forming a closed system, more complicated in the larger forms than in the aquatic genera. Several specially large contractile trunks in the anterior segments uniting the dorsal and ventral vessels. Nephridia generally paired, often very numerous in each segment, in the form of long, much-coiled tubes with intracellular lumen. Gonads limited in number of pairs, testes and ovaries always present in the same individual. Special sacs developed from the intersegmental septa lodge the developing ova and sperm. Special gonad ducts always present. Male ducts often open on to exterior through a terminal chamber which is variously specialized, and sometimes with a penis.

Generative pores usually paired, sometimes single and median. Spermathecae nearly always present. Alimentary canal straight, often with appended glands of complicated or simpler structure; no jaws. Eggs deposited in a cocoon after copulation. Development direct. Reproduction by budding also occurs. Fresh-water (rarely marine) and terrestrial.

The Oligochaeta show a greater variety of size than any other group of the Chaetopoda. They range from a millimetre or so (smaller species of Aeolosoma) to 6 ft. or even rather more (Microchaeta rappi, &c.) in length.

EB1911 Chaetopoda Fig. 11.—Setae of Oligochaeta.jpg
Fig. 11.—Setae of Oligochaeta.
a, Penial seta of Perichaeta ceylonica.

b, Extremity of penial seta of Acanthodrilus (after Horst).

c, Seta of Urochaeta (Perier).

d, Seta of Lumbricus.

e, Seta of Criodrilus.

f, g, Setae of Bohemilla comata.

h, i, j, Setae of Psammoryctes barbatus (f to j after Vezhdovský).

Setae.—The setae, which are always absent from the peristomial segment, are also sometimes absent from a greater number of the anterior segments of the body, and have completely disappeared in Achaeta cameranoi. When present they are either arranged in four bundles of from one to ten or even more setae, or are disposed in continuous lines completely encircling each segment of the body. This latter arrangement characterizes many genera of the family Megascolicidae and one genus (Periscolex) of the Glossoscolicidae. It has been shown (Bourne) that the “perichaetous” condition is probably secondary, inasmuch as in worms which are, when adult, “perichaetous” the setae develop in pairs so that the embryo passes through a stage in which it has four bundles of setae, two to each bundle, the prevalent condition in the group. Rarely there is an irregular disposition of the setae which are not paired, though the total number is eight to a segment (fig. 10), e.g. Pontoscolex. The varying forms of the setae are illustrated in fig. 11.

Structure.—The body wall consists of an epidermis which secretes a delicate cuticle and is only ciliated in Aeolosoma, and in that genus only on the under surface of the prostomium. The epidermis contains numerous groups of sense cells; beneath the epidermis there is rarely (Kynotus) an extensive connective tissue dermis. Usually the epidermis is immediately followed by the circular layer of muscles, and this by the longitudinal coat. Beneath this again is a distinct peritoneum lining the coelom, which appears to be wanting as a special layer in some Polychaetes (Benham, Gilson). The muscular layers are thinner in the aquatic forms, which possess only a single row of longitudinal fibres, or (Enchytracidae) two layers. In the earthworms, on the other hand, this coat is thick and composed of many layers.

The clitellum consists of a thickening of the epidermis, and is of two forms among the Oligochaeta. In the aquatic genera the epidermis comes to consist entirely of glandular cells, which are, however, arranged in a single layer. In the earthworms, on the other hand, the epidermis becomes specialized into several layers of cells, all of which are glandular. It is therefore obviously much thicker than the clitellum in the limicolous forms. The position of the clitellum, which is universal in occurrence, varies much as does the number of component segments. As a rule—to which, however, there are exceptions—the clitellum consists of two or three segments only in the small aquatic Oligochaeta, while in the terrestrial forms it is as a general rule, to which again there are exceptions, a more extensive, sometimes much more extensive, region.

In the Oligochaeta there is a closer correspondence between external metamerism and the divisions of the coelom than is apparent in some Chaetopods. The external segments are usually definable by the setae; and if the setae are absent, as in the anterior segments