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248
CHITON

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

EB1911 Chiton Fig. 1.jpg
Fig. 1.—Three views of Chiton.

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.

EB1911 Chiton Fig. 2.jpg
Fig. 2.—Pallial eye and aesthetes of Acanthopleura spiniger (Moseley).


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.

EB1911 Chiton Fig. 3.jpg
From Lankester, Treatise on Zoology.
Fig. 3.—Ventral aspect of three species of Polyplacophora showing position of gills.

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.

EB1911 Chiton Fig. 4.jpg

Fig. 4.—Diagrams of the alimentary canal of Amphineura
(from Hubrecht).
 A. Neomenia and Proneomenia.
 B. Chaetoderma.
 C. Chiton.
o, Mouth.
a, Anus.
d, Alimentary canal.
l,  Liver (digestive gland).

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