Echiuroidea

Echiuroidea (Gr. ἔχις, adder, and οὐρά, tail), the zoological name for a small group of marine animals which show in their larval life-history a certain degree of segmentation, and are therefore grouped by some authorities as Annelids. Formerly, together with the Sipunculoidea and Priapuloidea, they made up the class Gephyrea, but on the ground that they retain in the adult a large preoral lobe (the proboscis), that they have anal vesicles, that their anus is terminal, that setae are found, and finally that they are segmented in the larval stage, they have been removed from the class, which by the proposed further separation of the Priapuloidea on account of their unique renal and reproductive organs, has practically ceased to exist.  
  
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Fig. 1.—A, Bonellia viridis, Rol., ♀; B, B. fuliginosa. Both natural size. a, grooved proboscis; b, mouth; c, ventral hooks; d, anus.  
  
Echiuroids are animals of moderate size, varying roughly from one to six or seven centimetres in length, exclusive of the proboscis. This organ is capable of very considerable extension, and may attain a length in Bonellia viridis of about a metre and a half (fig. 1). It is grooved ventrally and ciliated. At its attachment to the body the groove sinks into the mouth. In Bonellia the proboscis is forked at its free end, but in the other genera it is short and unforked. The body is somewhat sausage-shaped, with the anus at the posterior extremity, surrounded in Echiurus by a single or double ring of setae. The skin is usually wrinkled, and in B. viridis, Thalassema lankesteri, Th. baronii, Hamingia arctica, and in the larva of many species, is of a lively green colour. A pair of curved bristles, formed in true setal sacs as in Chaetopoda, project from the body a short distance behind the mouth, and are moved by special muscles; they are of use in helping the animal to move slowly about, and they take a large share in the burrowing movements (C.B. Wilson, Biol. Bull., 1900), for some species tunnel in the mud and sand and form more or less permanent burrows, the walls of which are strengthened by mucus secreted from the skin. The openings of the burrows become silted up, leaving, however, a small aperture through which the proboscis is extruded. This organ carefully searches the neighbourhood for particles of food. When these are found the grooved proboscis folds its walls inwards, and the cilia pass the particles down the tube thus formed to the mouth. Echiuroids also move by extending the proboscis, which takes hold of some fixed object, and, then contracting, draws the body forwards. Recently it has been shown that Echiurus swims freely at night-time, using for locomotion both the proboscis and the contraction of the muscles of its body-wall. The motion is described as “gyratory,” and the anterior end is always carried foremost. Those species which do not burrow usually conceal themselves in crevices of the rocks or under stones, or at times in empty Mollusc or Echinid shells. They are occasionally used by fishermen for bait.  
  
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Fig. 2.—Female Bonellia viridis, Rol. Opened along the left side.  
  
a, Proboscis cut short.

b, Bristle passing through the mouth into the pharynx.

c, Coiled intestine.

d, Anal tufts or vesicles.

e, Ventral nerve cord.

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f, Ovary borne on ventral vessel running parallel with e.

g, Position of anus.

h, Position of external opening of nephridium.

i, Nephridium—the line points towards, but does not reach, the internal opening.  
  
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Fig. 3.—Adult male, Bonellia viridis, Rol. The original was 1.5 mm. long. The nervous system is not shown. (After Selenka.)  
  
a, Generative pore with spermatozoa coming out.

b, Anterior blind end of intestine attached to the parenchymatous tissue by muscular strands.

c, Green wandering cells containing chlorophyll.

d, Parenchymatous connective tissue.

e, Epidermis.

i, Intestine.

j, Vas deferens.

l, Internal opening of vas deferens.

m, The left anal vesicle.

n, Spermatozoa in the body-cavity.  
  
Anatomy (fig. 2).—A thin cuticle covers the epidermis, which contains mucus-secreting glands. Beneath the epidermis is a layer of circular muscles, then a layer of longitudinal, and finally in some cases a layer of oblique muscle-fibres. The inner face of this muscular skin is lined by a layer of epithelium. The coelomic body-cavity is spacious. It does not extend into the proboscis, which is a solid organ traversed by the nervous and vascular rings, but otherwise largely built up of muscle fibres and connective tissue. Many sense-cells lie in the epidermis. The ciliated ventral groove of the proboscis leads at its base into the simple mouth, which gives access to the thin-walled alimentary canal. This is longer than the body, and to tuck it away it is looped from side to side. The loops are supported by strands of connective tissue, which in some species are united so as to form a dorsal mesentery, whilst traces of a ventral mesentery are met with anteriorly and posteriorly (H.L. Jameson, Zool. Jahrb. Anat., 1899). The alimentary canal is divisible into fore-gut, mid-gut and hind-gut, and the first-named can be further divided into pharynx, oesophagus, gizzard and crop, mainly on histological grounds. The mid-gut is characterized by the presence of a ciliated groove, from which arises the collateral intestine or siphon, a second tube which rejoins the alimentary canal lower down. Similar collateral intestines are familiar in the Echinids and certain Polychaets (Capitellidae). The rectum receives the openings of a pair of very characteristic organs, the anal vesicles. Each consists of a branching tube, the tips of whose twigs terminate in minute ciliated funnels. The anal vesicles are thought to be excretory; whether this be so or not, they undoubtedly have some influence on the amount of fluid found in the coelom. The coelomic fluid contains as a rule both amoeboid and rounded corpuscles, and, when ripe, the products of the gonads. A closed system of vessels, usually called the vascular system, is present. There are, however, no capillaries connected with this, and it is confined to certain portions of the body. It can possess few of the functions usually associated with a vascular system, and its main use is probably to assist in the expansion of the proboscis. The system consists of the following parts:—A dorsal vessel applied to the alimentary canal is continued anteriorly into a median vessel, which traverses the proboscis to its tip. Here the vessel splits, and each half returns along the lateral edge of the proboscis; they reunite around the oesophagus and form a single ventral vessel, which lies above the ventral nerve-cord. The ventral vessel, which ends solidly behind, sends off a branch which forms a ring around the intestine and opens into the posterior extremity of the dorsal vessel. In Echiurus and Thalassema the same vessel forms a ring round a stout muscle, which connects the bases of the two ventral setae before passing to surround the intestine. Amoeboid corpuscles float in the fluid contents. The nephridia vary in number from a single one in Bonellia to three pairs in many species of Thalassema. Their external openings are ventral, and on the same level as the ciliated funnel-shaped nephrostomes. The posterior wall of the organ is produced into a long blind sac, which is lined by secretory cells. The nervous system is a single ventral cord, which starts from a circumoesophageal ring. This ring is involved in the growth of the proboscis, and is drawn out with it. Thus there is a lateral nerve near each edge of the proboscis which unites with its fellow dorsally above the oesophagus at the tip of the proboscis, and ventrally beneath the oesophagus, where they fuse to form the ventral nerve-cord. There are no specialized ganglia, but ganglion-cells are scattered uniformly along the nerve-cords. The ventral cord gives off rings, which run into the skin at regular intervals. The reproductive cells are modified coelomic cells, which lie on the ventral vessel. They escape into the coelomic fluid and there develop. When mature they leave the body through the nephridia. Bonellia and Hamingia are very interesting examples of sexual dimorphism. The female has the normal Echiuroid structure, but the male is reduced to a minute, flattened, planarian-like organism, which passes its life usually in the company of two or three others in a special recess of the nephridia of the female. Its structure may be gathered by a reference to fig. 3.

Larva.—The larva is a typical trochosphere, which, although of a temporary character, shows a distinct segmentation of the mesoblast, of the nervous system, and of the ciliated and pigmented structures in the skin, resembling that of Chaetopods. The preoral lobe persists as the proboscis. The sexes of the larvae are not determinable in the early stages, but when a certain growth has been reached in Bonellia the males seek the proboscis of the adult females, and passing into the mouth undergo there the transformation into the planarian-like parasite which is the fully-formed male. This now creeps along the body of the female and takes up its home in her nephridia.

Classification and Distribution.—The Echiuroidea consists of the following genera:—(1) Bonellia (Rol.), with four species, widely distributed, but inhabiting the temperate and warmer waters of each hemisphere. (2) Echiurus (Guérin-Méneville), with four species. This genus reaches from the Arctic waters of both hemispheres into the cooler temperate regions. (3) Hamingia (Kor. and Dan.), with one species, which has been taken in the Arctic Sea and the Hardanger Fjord. (4) Saccosoma (Kor. and Dan.) was described from a single specimen dredged about half-way between Iceland and Norway. (5) Thalassema (Gaertner, Lamarck), with twenty-one species. This genus is in the main a denizen of the warmer waters of the globe. Sixteen species are found only in tropical or subtropical seas, three species are Mediterranean (Mt. Stat. Neapel, 1899), whilst three species are from the eastern Atlantic, where the temperature is modified by the Gulf Stream (Shipley; see Willey’s Zoological Results, part iii. 1899; Proc. Zool. Soc. Lond., 1898, 1899; and Cambridge Natural History, ii.). The following are found in the British area:—E. pallasii (Guérin-Méneville), Th. neptuni (Gaertner), and Th. lankesteri (Herdman, Q.J.M.S., 1898).

Affinities.—The occurrence of trochosphere larva and the temporary segmentation of the body have led to the belief that the Echiuroids are more nearly allied to the Annelids than to any other phylum. This view is strengthened by certain anatomical and histological resemblances to the genus Sternaspis, which in one species, S. spinosa, is said to carry a bifid proboscis resembling that of the Echiuroids.

(A. E. S.)

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