Order I.--POLYPLACOPHORA

[Illustration: 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.]

[Illustration: 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.

[Illustration: 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.]

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.

[Illustration: 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).]

_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.

[Illustration: FIG. 5.--Diagrams of the excretory and reproductive

organs of Amphineura (after Hubrecht).

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.

[Illustration: After Haller (_Arbeiten zool. Instit._), Vienna, 1882.

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