Authorities.—Thucydides iv. 76; Diodorus xvi. 85-86; Plutarch, Alexander, ch. 9; Sulla, chs. 16–19; Appian, Mithradatica, chs. 42-45; W. M. Leake, Travels in Northern Greece (London, 1835), ii. 112–117, 192-201; B. V. Head, Historia Numorum (Oxford, 1887), p. 292; J. Kromayer, Antike Schlachtfelder in Griechenland (Berlin, 1903), pp. 127–195; G. Sotiriades in Athen. Mitteil. 1903, pp. 301 ff.; 1905, p. 120; 1906, p. 396; Έφημ. Άρχαιολ., 1908, p. 65.
CHAETOGNATHA, the name given by R. Leuckhart to a small group of transparent and for the most part pelagic organisms, whose position in the animal kingdom is a very isolated one. Only three genera, Sagitta, Spadella and Krohnia, are recognised, and the number of species is small. Nevertheless these animals exist in extraordinary quantities, so that at certain seasons and under certain conditions the surface of the sea seems almost stiff with the incredible multitude of organisms which pervade it. Rough seas, &c., cause them to seek safety in dropping into deeper water. Deep-sea forms also occur, but in spite of this the group is essentially pelagic.
As a rule the body is some 1 to 2 or 3 cm. in length, though some species are larger, by 4 or 5 mm. in breadth, and it is shaped something like a torpedo with side flanges and a slightly swollen, rounded head. It can be divided into three regions—(i.) head, (ii.) trunk, and (iii.) tail, separated from one another by two transverse septa. The almost spherical head is covered by a hood which can be retracted; it bears upon its side a number of sickle-shaped, chitinous hooks and one or more short rows of low spines—both of these features are used in characterizing the various species. A pair of eyes lie dorsally and behind them is a closed circlet, often pulled out into various shapes, of modified epidermis, to which an olfactory function has been attributed. The interior of the head is filled up with masses of muscle fibres which are mainly occupied with moving the sickle-shaped hooks. The trunk contains a spacious body-cavity filled during the breeding season by the swollen ovaries, and the same is true of the tail if we substitute testes for ovaries.
The skin consists of a transparent cuticle excreted by the underlying ectoderm, the cells of which though usually one-layered may be heaped up into several layers in the head; beneath this is a basement membrane, and then a layer of longitudinal muscle fibres which are limited inside by a layer of peritoneal cells. The muscles are striated and arranged in four quadrants, two dorso-lateral and two ventro-lateral, an arrangement which recalls that of the Nematoda, whilst in their histology they somewhat resemble the muscles of the Oligochaeta. Along each side of the body stretches a horizontal fin and a similar flange surrounds the tail. Into these fins, which are largely cuticular and strengthened by radiating bars, a single layer of ectoderm cells projects.
The mouth, a longitudinal slit, opens on to the ventral surface of the head. It leads into a straight alimentary canal whose walls consist of a layer of ciliated cells ensheathed in a thin layer of peritoneal cells. There is no armature, and no glands, and the whole tract can only be divided into an oesophagus and an intestine. The latter runs with no twists or coils straight to the anus, which is situated at the junction of the trunk with the tail. A median mesentery running dorso-ventrally supports the alimentary canal and is continued behind it into the tail, thus dividing the body cavity into two lateral halves.
There are no specialized circulatory, respiratory or excretory organs.
The nervous system consists of a cerebral ganglion in the head, a conspicuous ventral ganglion in the trunk, and of lateral commissures uniting these ganglia on each side. The whole of this system has retained its primitive connexion with the ectoderm. The cerebral ganglion also gives off a nerve on each side to a pair of small-ganglia, united by a median commissure, which have sunk into and control the muscles of the head. As in other animals there is a minute but extensive nervous plexus, which permeates the whole body and takes its origin from the chief ganglia. In addition to the eyes and the olfactory circle on the head scattered tactile papillae are found on the ectoderm.
Chaetognatha are hermaphrodite. The ovaries are attached to the side walls of the trunk region; between them and the body wall lie the two oviducts whose inner and anterior end is described as closed, their outer ends opening one on each side of the anus, where the trunk joins the tail. According to Miss N. M. Stevens the so-called oviduct acts only as a “sperm-duct” or receptaculum seminis. The spermatozoa enter it and pass through its walls and traverse a minute duct formed of two accessory cells, and finally enter the ripe ovum. Temporary oviducts are formed between the “sperm-duct” and the germinal epithelium at each oviposition. A number of ova ripen simultaneously. The two testes lie in the tail and are formed by lateral proliferations of the living peritoneal cells. These break off and, lying in the coelomic fluid, break up into spermatozoa. They pass out through short vasa deferentia with internal ciliated funnels, sometimes an enlargement on their course—the seminal vesicles—and a minute external pore situated on the side of the tail.
With hardly an exception the transparent eggs are laid into the sea and float on its surface. The development is direct and there is no larval stage. The segmentation is complete; one side of the hollow blastosphere invaginates and forms a gastrula. The blastopore closes, a new mouth and a new anus subsequently arising. The archenteron gives off two lateral pounchs and thus becomes trilobed. The middle lobe forms the alimentary canal; it closes behind and opens to the exterior anteriorly and so makes the mouth. The two lateral lobes contain the coelom; each separates off in front a segment which forms the head and presumably then divides again to form anteriorly the trunk, and posteriorly the tail regions. An interesting feature of the development of Chaetognaths is that, as in some insects, the cells destined to form the reproductive organs are differentiated at a very early period, being apparent even in the gastrula stage.
The great bulk of the group is pelagic, as the transparent nature of all their tissues indicates. They move by flexing their bodies. Spadella cephaloptera is, however, littoral and oviposits on seaweed, and the “Valdivia” brought home a deep-sea species.
The three genera are differentiated as follows:—
Sagitta M. Slabber, with two pairs of lateral fins. This genus was named as long ago as 1775.
Krohnia P. Langerhans, with one lateral fin on each side, extending on to the tail.
Spadella P. Langerhans, with a pair of lateral fins on the tail and a thickened ectodermic ridge running back on each side from the head to the anterior end of the fin.
The group is an isolated one and should probably be regarded as a separate phylum. It has certain histological resemblances with the Nematoda and certain primitive Annelids, but little stress must be laid on these. The most that can be said is that the Chaetognaths begin life with three segments, a feature they share with such widely-differing groups as the Brachiopoda, the Echinoderma and the Enteropneusta, and probably Vertebrata generally.
See O. Hertwig, Die Chaetognathen, eine Monographie (Jena, 1880); B. J. Grassi, Chetognathi: Flora u. Fauna d. Golfes von Neapel (1883); S. Strodtman, Arch. Naturg. lviii., 1892; N. M. Stevens, Zool. Jahrb. Anat. xviii., 1903, and xxi., 1905.
(A. E. S.)
CHAETOPODA (Gr. χαίτη, hair, πούς, foot), a zoological class, including the majority of the Annelida (q.v.), and indeed, save for the Echiuroidea (q.v.), co-extensive with that group as usually accepted. They are divisible into the Haplodrili (q.v.) or Archiannelida, the Polychaeta containing the marine worms, the Oligochaeta or terrestrial and fresh-water annelids (see Earthworm), the Hirudinea or leeches (see Leech), and a small group of parasitic worms, the Myzostomida (q.v.).
The distinctive characters of the class Chaetopoda as a whole are partly embodied in the name. They possess (save for certain Archiannelida, most Hirudinea, and other very rare exceptions) setae or chaetae implanted in epidermal pits. The setae are implanted metamerically in accordance with the metamerism of the body, which consists of a prostomium followed by a number of segments. The number of segments in an individual is frequently more or less definite. The anterior end of body always shows some “cephalization.” The internal organs are largely repeated metamerically, in correspondence with the external metamerism. Thus the body cavity is divided into a sequence of chambers by transverse septa; and even among the Hirudinea,