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Chiton

CHITON, the name [1] given to fairly common littoral animals of rather small size which belong to the phylum Mollusca, and, in the possession of a radula in the buccal cavity, resemble more especially the Gastropoda. Their most important characteristic in comparison with the latter is that they are, both in external and internal structure, bilaterally symmetrical. The dorsal integument or mantle bears, not a simple shell, but eight calcareous plates in longitudinal series articulating with each other. The ventral surface forms a flat creeping "foot," and between mantle and foot is a pallial groove in which there is on each side a series of gills. Originally the Chitons were placed with the limpets, Patella, in Cuvier's Cyclobranchia, an order of the Gastropoda. In 1876 H. von Jhering demonstrated the affinities of Neomenia and Chaetoderma, vermiform animals destitute of shell, with the Chitons, and placed them all in a division of worms which he named Amphineura. The discovery by A.A.W. Hubrecht in 1881 of a typical molluscan radula and odontophore in a new genus Proneomenia, allied to Neomenia, showed that the whole group belonged to the Mollusca. E. Ray Lankester (Ency. Brit., 9th ed., 1883) placed them under the name Isopleura as a subclass of Gastropoda. Paul Pelseneer (1906) raised the group to the rank of a class of Mollusca, under von Jhering's name Amphineura.

The Amphineura are divided into two orders: (1) the Polyplacophora, or Chitons; (2) the Aplacophora, or forms without shells, Neomenia, Chaetoderma and their allies.

Order I. - Polyplacophora

A. Dorsal view of Chiton Wosnessenksii, Midd., showing the eight shells. (After Middendorf.)

B. View from the pedal surface of a species of Chiton from the Indian Ocean, p, foot; o, mouth (at the other end of the foot is seen the anus raised on a papilla); kr, oral fringe; br, the numerous ctenidia (branchial plumes); spreading beyond these, and all round the animal, is the mantle-skirt. (After Cuvier.)

C. The same species of Chiton, with the shells removed and the dorsal integument reflected, b, buccal mass; m, retractor muscles of the buccal mass; ov, ovary; od, oviduct; i, coils of intestines; ao, aorta; c', left auricle; c, ventricle.

Each of the eight valves of the shell is made up of two distinct calcareous layers: (a) an outer or upper called the tegmentum, which is visible externally; (b) a deeper layer called articulamentum which is porcellaneous, quite compact, and entirely covered by the tegmentum. In the lower forms the two layers are coextensive and have smooth edges, but in the higher forms the articulamentum projects laterally beyond and beneath the tegmentum into the substance of the mantle. These projections are termed insertion plates; they are usually slit or notched to form teeth, the edges of which may be smooth and sharp, or may be crenulated. The anterior margin of each valve except the first is provided with two projections called sutural laminae which underlie the posterior margin of the preceding valve.

A. Lepidopleurus benthus.

B. Boreochiton cinereus.

C. Schizochiton incisus. a, anus; f, foot; g, gills; m, mouth; pa, mantle; pa', anal lobe of mantle; ps, pallial slit; te, pallial tentacles.

The tegmentum is formed by the fold of mantle covering the edge of the articulamentum, and extends over the latter from the sides. It is the first part of the shell formed in development. The tegmentum is much reduced in Acanthochiton, and absent in the adult Cryptochiton. The tegmentum is pierced by numerous vertical ramified canals which contain epithelial papillae of the epidermis. These papillae form pallial sense-organs, containing nerve-end bulbs, covered by a dome of cuticle, and innervated from the pallial nerve-cords. They are termed according to their size, micraesthetes and megalaesthetes. In the common species of Chiton and many others of the family Chitonidae the megalaesthetes are developed into definite eyes, the most complicated of which have retina, pigment within the eye, cornea and crystalline lens (intra-pigmental eyes) (fig. 2). The eyes are arranged in rows running diagonally from the median anterior beak of each valve to its lateral borders There may be only one such row on either side, or many rows. In some species the total number present amounts to thousands.

Branchiae. - The series of gills may extend the whole length of the body in the pallial groove, or may be confined to the posterior end. Each gill has the structure of a typical molluscan ctenidium, consisting of an axis bearing an anterior and posterior row of filaments or lamellae. The gills are thus metamerically repeated; there may be from four to eighty pairs, but there is often a numerical asymmetry on the two sides. The largest pair of branchiae is placed immediately behind the renal openings and corresponds to the single pair of other molluscs, the organs being repeated anteriorly only (Metamacrobranchs) or anteriorly and posteriorly (Mesomacrobranchs).

Intestine. - The digestive tube in the Polyplacophora, which are herbivorous, is longer than the body, and thrown into a few coils, the anus being median and posterior. The mouth leads into the buccal cavity, on the ventral side of which opens the radular caecum. Each transverse row of teeth of the radula contains 17 teeth, one of which is median, while the second and the fifth on each side are enlarged. Two pairs of glands open into the buccal cavity, and at the junction of pharynx and oesophagus is another pair called the sugar glands. The stomach is surrounded by the liver or digestive gland, consisting of two lobes which are symmetrical in the young animals, but in the adult the right lobe is anterior and smaller.

A, Chaetoderma.

B, Neomenia.

C, Proneomenia.

D, Chiton.

O, Ovary.

P, Pericardium.

N, Nephridium.

u, External aperture of nephridium.

g, External aperture of the genital duct of Chiton.

r, Rectum.

Cl, Cloacal or pallial chamber of Neomeniae and Chaetoderma.

Br, Ctenidia (branchial plumes).

Coelom, Gonads and Excretory Organs. - As in other molluscs the coelom is represented by a large pericardial cavity, situated above the intestine posteriorly, and a generative sac which is single and median and situated in front of the pericardium, except in the Nuttalochiton hyadesi, where the gonads are in a similar position, but are paired. The excretory organs are coelomoducts with an internal ciliated opening into the pericardium and an opening to the exterior. Both the openings are close together, the external opening being just in front of the principal gill near the posterior end of the body. The renal tube is doubled on itself, its middle part where the bend occurs being situated more or less anteriorly. The excretory surface is increased by numerous ramified caeca which extend beneath the body wall laterally and ventrally, and open into the tube (fig. 6). The sexes are distinct, and the ovary is frequently greenish in colour, the testis red. The gonad is transversely wrinkled and lies between the aorta and the intestine, extending from the pericardium to the anterior end of the body. A simple gonaduct on each side arises from the gonad near its posterior end and passes first forwards, then backwards, and lastly outwards to the external opening in the pallial groove, anterior to the renal aperture. There may be from one to nine gills between the genital and renal pores.

Heart and Vascular System. - The heart is enclosed in the pericardium, and consists of a median elongated ventricle and a pair of lateral auricles, so that the structure somewhat resembles that in the Lamellibranchiata. The openings of the auricles into the ventricle vary in different forms. In many of the lower forms (Lepidopleuridae, Mopalidae, Ischnochitonidae) the opening on each side is single and anterior. In the true Chitonidae there are generally two apertures on each side, and in two species three or four, another instance of the tendency to metameric repetition in the group. The auricles are connected with one another posteriorly behind the ventricle. The ventricle leads into a single anterior median aorta. As in other molluscs, the arteries do not extend far, but lead into inter-visceral blood-spaces. The venous blood is conducted from the tissues to a large sinus on either side above the pallial groove, and from this sinus passes to the gills by an afferent vessel in each gill on the internal or pedal margin of the axis. The oxygenated blood is carried from each gill by an efferent vessel on the external or pallial side of the axis to another longitudinal vessel which leads to the auricle on each side.

Nervous System. - There are no well-marked specialized ganglia in the central nervous system, nerve-cells being distributed uniformly along the cords. There are two pairs of longitudinal cords, a pedal pair situated ventrally and united beneath the intestine by numerous commissures, and a pallial pair situated laterally and continuous with one another above the rectum (fig. 7). The four cords are all connected anteriorly with the cerebral commissure which lies above the buccal mass anteriorly. From the points where the cords meet the cerebral commissure, arise on each an anterior labial commissure and a stomatogastric commissure. The letter bears two ganglion swellings, the buccal ganglia. The labial commissure gives off a subradular commissure which also bears two ganglia, these being in close relation to a special sense-organ called the subradular organ, an epithelial projection with nerve-endings, lying in front of the radula and probably gustatory in function. One osphradium or branchial olfactory organ is usually present on each side, on either side of the anus on the inner wall of the mantle, near the base of the last gill. In Lepidopleuridae an osphradium occurs at the base of each gill. The sense organs of the shell-valves have already been described.

Development. - The eggs may be laid separately invested by a chitinous envelope, or as in Ischnochiton magdalenensis they may form strings containing nearly 200,000 eggs, or the ova may be retained in the pallial groove and undergo development there, as in Chiton polii and Hemiarthrum setulosum. One species Callistochiton viviparus is viviparous and its ova develop without a larval stage in the maternal oviduct. Segmentation is total and at first regular, and is followed by invagination, the blastopore passing to the position of the future mouth. By the development of a ciliated ring just in front of the mouth the embryo becomes a trochosphere. In the centre of the praeoral lobe is a tuft of cilia. Just behind the ciliated ring is a pair of larval eyes which disappear in the adult; these correspond to the cephalic eyes of Lamellibranchs. An ectodemic invagination forms a large mucous gland on the foot, which is more or less atrophied in adult life. The gonads originate by proliferation of the anterior wall of the pericardium. The shell-valves arise as transverse thickenings of the dorsal cuticle behind the ciliated ring, the tegmentum being the first part formed.

Classification.

Suborder I. Eoplacophora, Pilsbry. - Tegmentum coextensive with articulamentum, or the latter projecting in smooth unslit plates.

Fam. 1. Lepidopleuridae. - Terminal margins of end valves never elevated; form oval or oblong. Lepidopleurus cancellatus, Sow. North Atlantic and Mediterranean; various abyssal species. Hanleya hanleyi, Bean, north Atlantic. Hemiarthrum Microplax. The extinct Gryptochitonidae, Pilsbry, with other Palaeozoic genera, narrow and elongated in form with terminal margins of end valves elevated, belong to this group.

Suborder II. Mesoplacophora, Pilsbry. - Insertion plates well developed and slit.

Fam. 2. Ischnochitonidae. - All the valves with slits, and the inner layer well covered by the outer.

Subfam. 1. Ischnochitoninae. - No shell-eyes: sutural laminae separated; slits in the valves 1-7 do not correspond with the ribs of the tegmentum. Ischnochiton, Trachydermon, Chaetopleura, Stenoplax, Stenoradsia.

Subfam. 2. Callochitoninae. With shell-eyes and united sutural laminae. Callochiton laevis, North Atlantic and Mediterranean.

Subfam. 3. Callistoplacinae. No shell-eyes, slits in the valves 1-7 corresponding with the ribs of the tegmentum. Callistochiton (viviparous). Nuttalochiton.

Fam. 3. Mopaliidae. Each intermediate valve with a single slit; girdle hairy. Mopalia, Placiphorella, Plaxiphora, Placophoropsis.

Fam. 4. Acanthochitonidae. Valves immersed in the girdle, with small tegmentum. Acanthochiton (A. fascicularis, North Atlantic and Mediterranean). Spongiochiton, Katharina, Amicula, Cryptochiton (C. stelleri, arctic).

Fam. 5. Cryptoplacidae. Vermiform, with thick girdle and small valves; insertion and sutural plates strongly drawn forward, sharp and smooth. Cryptoplax, Choneplax.

Suborder III. Teleoplacophora, Pilsbry. - All the valves, or at least the seven anterior, with insertion plates cut into teeth by slits.

Fam. 6. Chitonidae. Characters of the suborder.

Subfam. 1. Chitoninae. No extra-pigmental eyes; insertion plates with pectinations between the fissures. Chiton, Eudoxochiton, Trachyodon, Radsia.

Subfam. 2. Toniciinae. Extra-pigmental shell-eyes. Tonicia, Acanthopleura, Enoplochiton, Onithochiton, Schizochiton, Lorica, Loricella, Liolophura.

Order 2. - Aplacophora, von Jhering.

Chaetoderma was first described by S. LovOøΩn, in 1841, and was for a long time believed to be a Gephyrean worm. Neomenia, mentioned first by Michael Sars in 1868 under the name Solenopus, was afterwards included among the Opisthobranchs by J. Koren and D.C. Danielssen. C. Gegenbaur placed the two genera in a division of Vermes which he called Solenogastres.

The chief points in which the Aplacophora differ from the Polyplacophora are: (1) they are worm-like in shape; (2) there is no distinct foot, and the mantle bears no shell-valves, but only numerous calcareous spicules; (3) the digestive tube is straight.

Neomenia and its allies are marine animals living at depths of 15 to 800 fathoms on soft muddy ground; they are found crawling on corals and hydrozoa, on which they feed. The British genera are: Neomenia, Rhopalomenia and Myzomenia. They have been taken in nearly all seas except the South Atlantic and S.E. and N.W. Pacific. About forty species are known. Chaetoderma, of which nine species have been described, has similar habits and distribution, but feeds chiefly on Protozoa. The order Aplacophora is divided into two suborders.

Suborder I. Neomeniomorpha. - Aplacophora with a distinct longitudinal ventral groove; bisexual with paired genital glands and no distinct liver. The whole of the skin except the ventral groove corresponds to the mantle of Chiton. The cuticle, in some species very thick, contains numerous spicules which are long, hollow and calcified; they are secreted by epithelial papillae. In some species there are also sensory papillae comparable to the aesthetes of Chitons. A small longitudinal projection in the ventral groove represents the foot. Into the groove open mucous glands, a large one anteriorly and another opening into a posteriorly cloacal, branchial cavity.

A, Lateral view.

B, Ventral view.

C, Dorsal view.

D, Ventral view of a more extended specimen.

a, Anterior.

b, Posterior extremity.

c, Furrow, in which the narrow foot is concealed.

Branchiae. - In Neomeniidae and most of the Parameniidae there is a circlet of gills on the inner walls of the cloacal chamber. These gills are simple folds or laminae of the body wall. In other species they are absent.

Intestine. - -The mouth opens into a muscular pharynx lined by a thick cuticle. Into the pharyngeal cavity open salivary glands and radular sac. The former are paired and ventral, and open on a subradular prominence. In some species there is a second dorsal pair. Neomenia and other genera have no salivary glands.

The radula when present comprises several transverse rows of teeth, and each transverse row may have several teeth (polystichous), two teeth (distichous), or one tooth (monostichous). It is a curious fact that in the original type Neomenia the radula is entirely absent, as it likewise is in several genera of Proneomeniidae. The oesophagus is short and leads into a long, straight stomach, provided with numerous symmetrical lateral caeca. The stomach opens into a short straight rectum which opens into the branchial chamber.

Coelom, Gonads and Excretory Organs. - The coelom differs from that of the Chitons in the fact that the cavities of the genital organs are continuous with it, and in the fact that there is only one pair of coelomoducts resembling the renal organs of Chitons, but serving also as genital ducts. The gonads are paired and hermaphrodite, they form a pair of anterior prolongations of the pericardium, extending nearly to the anterior end of the body. Ova are developed on the median, spermatozoa on the outer wall of each genital tube. The pericardium is ciliated internally on its dorsal and lateral walls. The urino-genital tubes arise from the posterior angles of the pericardium, pass first forwards, then backwards, and unite to open by a common opening into the cloaca below the anus except in Strophomenia, where the openings are separate. Usually each tube is provided with caecal appendages on its proximal portion, and these serve as vesiculae seminales, while the distal portion is enlarged and glandular and secretes the egg-shell.

Heart and Vascular System. - There is a heart in the pericardium consisting of a median ventricle attached, except in Neomenia, to the dorsal wall of the pericardium, and in Neomenia a pair of auricular ducts returning blood from the gills to the ventricle. The aorta is not independent as in Chitons, but is a sinus like the other channels of the circulation. A single median ventral sinus passes backwards to the gills or cloaca. The blood is coloured red by haemoglobin in blood corpuscles.

Nervous System. - Ganglionic enlargements are more conspicuous than in the Chitons. In front of the buccal mass is a median cerebral ganglion. From this pass off two pairs of cords, the pleural and pedal, in Proneomenia separate from their origin, in Neomenia united at first and diverging at a pleural ganglion. The pedal cords anteriorly form a pair of pedal ganglia united by a thick commissure. The supra-rectal commissure may be present and bear an ovoid ganglion; or may be wanting. With regard to sense organs the epithelial papillae of the mantle have been mentioned. There is also in some genera a median retractile sensory papilla on the dorsal posterior surface above the rectum, not covered by the cuticle.

Development has only been described in Myzomenia banyulensis, by G. Pruvot. It closely resembles in the early stages that of Chitons. The external surface of the trochosphere is formed of a number of ciliated test-cells. The ectoderm behind the ciliated ring develops spicules, and the post-oral region of the larva elongates. Later the ciliated ring or velum disappears and seven imbricated calcareous plates, made up of flattened spicules, are formed on the dorsal surface. This appears to indicate that the Neomeniomorpha are descended from Chiton-like ancestors, and that they have lost their shell valves.

Classification of the Neomeniomorpha. - Fam. 1. Lepidomeniidae. Slender, tapering behind, with subventral cloacal orifice; thin cuticle without papillae; flattened spicules; no gills. Lepidomenia, Ismenia, Ichthyodes, Stylomenia, Dondersia, Nematomenia, Myzomenia, M. banyulensis, Mediterranean and Plymouth.

Fam. 2. Neomeniidae. Short, truncate in front and behind; cloacal orifice transverse; gills present; rather thin cuticle; no radula. Neomenia (N. carinata, N. Atlantic and N. and N.W. Scotland), Hemimenia.

Fam. 3. Proneomeniidae. Elongated, cylindrical, rounded at both ends; thick cuticle with acicular spicules; radula polystichous or wanting. Proneomenia, Amphimenia, Echinomenia, Rhopalomenia (R. aglaopheniae, Mediterranean and Plymouth), Notomenia, Pruvotia, Strophomenia.

Fam. 4. Parameniidae. Short and truncated in front; thick cuticle, often without papillae; gills and radula present. Paramenia, Macellomenia, Pararhopalia, Dinomenia, Cyclomenia, Proparamenia, Uncimenia, Kruppomenia.

Suborder II. Chaetodermomorpha. - Aplacophora without distinct ventral groove, with single median unisexual gonad, with differentiated hepatic sac, and with cloacal chamber furnished with two bipectinate gills. There are only two genera in this suborder: Chaetoderma, and Limifossor from Alaska. The characters therefore are very uniform. The body is worm-like and cylindrical, the posterior half a little thicker than the anterior; the posterior extremity forms the enlarged funnel-like branchial or cloacal chamber. The anterior extremity is also somewhat enlarged. The whole surface is uniformly covered with short compressed calcareous spicula embedded in the cuticle.

Branchiae. - The single pair of branchiae are placed symmetrically right and left of the anus, and each has the structure of a ctenidium bearing a row of lamellae on each side as in the Polyplacophora.

Intestine. - The mouth is anterior, terminal and crescentic, and beneath it is a rounded ventral shield. On the floor of the pharynx or buccal mass is a rudimentary radula, which in many species consists of a single large tooth, bearing two small teeth or a row of teeth. In other species the radula is more of the usual type consisting of several transverse rows of two or three teeth each. Two pairs of salivary glands open into the buccal cavity. The digestive tube is straight and simple, wider in its anterior part, into which opens the duct of the hepatic caecum (fig. 4, B). The latter extends backwards on the ventral side of the intestine.

Coelom, Gonads and Excretory Organs. - These are closely similar in their relations to those of the Neomeniomorpha. The chief difference is that the gonad or generative portion of the coelom is single and median, opening into the pericardium by a single posterior aperture. The excretory organs or coelomoducts arise from the posterior corners of the pericardium, run forwards and then backwards to open by separate apertures lateral to the gills (fig. 5, A). There are no accessory generative organs.

The heart and vascular system are similar to those of the Neomeniomorpha, the only important differences being that the ventricle is nearly free in the pericardial cavity, and that the latter is traversed by the retractor muscles of the gills.

Nervous System. - There are two closely connected cerebral ganglia, from which arise the usual two pairs of nerve cords. Pallial and pedal on each side are closer together than in the other groups, and posteriorly they unite into a supra-rectal cord provided with a median ganglionic enlargement (fig. 7, C). A small stomatogastric commissure bearing two small ganglia arises from the cerebral ganglia and surrounds the oesophagus.

The development is at present entirely unknown.

General Remarks on the Amphineura.

The most important theoretical question concerning the Amphineura is how far do they represent the original condition of the ancestral mollusc? That is to say, we have to inquire which of their structural features is primitive and which modified. Their bilateral symmetry is obviously to be regarded as primitive, and the nervous system shows an original condition from which that of the asymmetrical twisted Gastropods can be derived. But in many other features both external and internal the three principal divisions differ so much from one another that we have to consider in the case of each organ-system which condition is the more primitive. According to Paul Pelseneer the Polyplacophora are the most archaic, the Aplacophora being specialized in (1) the great reduction of the foot, (2) the disappearance of the shell (Cryploplax among the Polyplacophora showing both reductions in progress), (3) the disappearance of the radula. But it is a widely recognized principle of morphology that a much modified animal is by no means modified to the same degree in all its organs. A form which is primitive on the whole may show a more advanced stage of evolution in some particular system of organs than another animal which is on the whole more highly developed and specialized. Thus the independent metamerism of certain organs in the Chitons is not primitive but acquired within the group: e.g. the shell valves and the ctenidia. And although embryology seems to prove that the Neomeniomorphs are derived from forms with a series of shell-valves, nevertheless it seems probable that the calcareous spicules which alone are present in adult Aplacophora preceded the solid shell in evolution.

It is held by some morphologists that the mollusc body is unsegmented, and therefore is to be compared to a single segment of a Chaetopod or Arthropod. In this case there should be only one pair of coelomoducts in the adult, the pair of true nephridia which should also occur being represented by the larval nephridia. There should also be only a single coelom, or a pair of lateral coelomic cavities. On this view then the Aplacophora are more primitive than the Polyplacophora in the relations of coelom, gonad and coelomoducts; and the genital ducts of the Chitons have arisen either by metameric repetition within the group, or by the gradual loss of an original connexion between the generative sac and the renal tube, as in Lamellibranchs and Gastropods, the generative sac acquiring a separate duct and opening to the exterior on each side.

Literature. - A. Sedgwick, "On certain Points in the Anatomy of Chiton," Proc. R. Soc. Lond. xxxiii., 1881; J. Blumrich, "Das Integument der Chitonen," Zeitsch. f. wiss. Zool. lii., 1891; A.C. Haddon, "Report on the Polyplacophora," Challenger Reports. Zool. pt. xliii., 1886; H.N. Moseley, "On the presence of Eyes in the Shells of certain Chitonidae, and on the structure of these Organs," Quart. Journ. Mic. Sci. new ser. xxv., 1885; A.A.W. Hubrecht, "Proneomenia Sluiteri," Nied. Arch. f. Zool. Suppl. 1., 1881; A. Kowalewsky and A.F. Marion, "Contr. OøΩ l'histoire des Solenogastres ou Aplacophores," Ann. Mus. Marseille, Zool. iii., 1887; A. Kowalewsky, "Sur le genre Chaetoderma," Arch. de zool. expOøΩr. (3) ix., 1901; P. Pelseneer, "Mollusca," Treatise on Zoology, edited by E. Ray Lankester, pt. v., 1906; E. Ray Lankester, "Mollusca," in the 9th ed. of this Encyclopaedia, to which this article is much indebted.

(J. T. C.)

[1] The Gr. was a garment in the shape of a loose tunic, varying at different periods: see Costume: Greek.

Note - this article incorporates content from Encyclopaedia Britannica, Eleventh Edition, (1910-1911)

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