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Monogonopora, in which a single orifice serves for both male and
female organs, may be taken the common garden snail (Helix
aspersa), the accompanying figure of which is drawn from two
specimens found in the act of union (Fig. 53).
Fig. 53.—Genitalia of Helix aspersa Müller,
drawn from two individuals in the act of
union, from a dissection by F. B. Stead.
 A.G, albumen gland.
C, coecum.
Cr, crop.
D.S, dart sac.
E, eye (retracted).
Fl, flagellum.
H.D, hermaphrodite duct.
H.DF, ditto, female portion.
H.DM, ditto, male portion.
H.G, hermaphrodite gland.
L, liver.
M.G, M.G, mucous glands.
Ov, oviduct.
P.S, penis sac.
R.M, retractor muscle of penis.
Sp, spermatheca.
V, vagina.
V.D, vas deferens.

Beginning from the inside and proceeding outwards we have

firstly the hermaphrodite gland or ovo-testis (h.g.), a yellowish white
mass of irregular shape, embedded in the liver (l.) and forming part
of its spiral but not reaching quite to the apex. Within this gland are
developed the ova and spermatozoa. The former are rather large
round cells, produced within the outer wall of the gland, while the
spermatozoa, which are produced in the more central part, are
thread-like bodies, generally aggregated in small bundles. From the
hermaphrodite gland the ova and spermatozoa pass through the
upper part of the hermaphrodite duct (h.d.), which is always more or
less convoluted. Below the convoluted portion, the duct opens into
the albumen gland (a.g.), a large linguiform mass of tissue which
becomes dilated at the time of pairing, and secretes a thick viscid
fluid which probably serves to envelop the ova. Up to this point both
the male and female elements follow the same course, but on their
exit from the albumen gland they diverge. The hermaphrodite duct
becomes greatly enlarged, and is partially divided by a kind of
septum into a male and female portion. These run parallel to one
another, the larger or female portion (h.df.), through which the ova
pass (and which is sometimes termed the uterus) being dilated into a
number of puckered folds, while the smaller or male portion (h.dm.)
is comparatively narrow, and not dilated. At their anterior end, the
two portions of the duct separate completely from one another, the
female portion being then termed the oviduct (ov.) and the male
portion the vas deferens (v.d.).
Following first the oviduct, we find that it soon widens into the
vagina (v.), which is furnished with a pair of mucous glands (m.g.),
one on each side. These are much branched, and resemble little
bunches of whitish seaweed. A little above the mucous glands a long
tube diverges from the vagina, which is furnished with a produced
coecum (c.) and a pouch, the spermatheca (sp.) at the extreme end.
In this pouch, and in the duct leading to it, is stored the
spermatophore received in union with another snail. Just below the
mucous glands the vagina is joined by the dart sac (d.s.), which is
more fully described below. Finally, at its lower end the vagina unites
with the penis sac at a point just posterior to the common orifice.
Returning now to the male organs, we find that the vas deferens
is the continuation of the male portion of the hermaphrodite duct,
after its final separation from the female portion. It passes under the
retractor muscle of the upper right tentacle, which has been cut away
in the specimen figured, to dissect it out. Just before the vas
deferens widens into the penis sac, it branches off into a long and
tapering tube, the flagellum, in which the spermatozoa are stored
and become massed together in the long packet known as the
spermatophore. The penis sac (p.s.) is the continuation of the vas
deferens beyond the point at which the flagellum diverges. It joins
the vagina at its extreme anterior end, uniting with it to form the
common genital aperture, which cannot be exactly represented in
the figure. The penis itself lies in the interior of the penis sac, and is
a rather long muscular tube which is protruded during union, but at
other times remains retracted within the sac.
In the Helicidae generally, the form of the generative organs
varies with each separate species, sometimes merely as regards the
size of the different parts, at others in the direction of greater
simplicity or complication. The mucous glands may be absent, and
the flagellum greatly reduced in size, or absent altogether.
 The Dart Sac.—A remarkable part of the reproductive system in
many of the true Helicidae is the so-called dart, Liebespfeil, or telum
veneris. It consists of ‘a straight, or curved, sometimes slightly
twisted tubular shaft of carbonate of lime, tapering to a fine point
above, and enlarging gradually, more often somewhat abruptly, to
the base.’ The sides of the shaft are sometimes furnished with two or
more blades; these are apparently not for cutting purposes, but
simply to brace the stem. The dart is contained in a dart sac, which
is attached as a sort of pocket to the vagina, at no great distance
from its orifice. There are four different forms of sac. It may be single
or double, and each of these divisions may be bilobed, each lobe
containing one dart at a time. In Helix aspersa the dart is about 5/16
in. in length, and ⅛ in. in breadth at its base (see Fig. 54).
It appears most probable that the dart is employed as an adjunct
to the sexual act. Besides the fact of the position of the dart sac
anatomically, we find that the darts are extruded and become
embedded in the flesh just before or during the act of copulation. It
may be regarded, then, as an organ whose punctures induce
excitement preparatory to sexual union. It only occurs in well-grown
specimens. When once it begins to form, it grows very rapidly,
perhaps not more than a week being required for its entire formation.

Fig. 54.—Darts of British land snails: A,

Hyalinia excavata Bean; B, Helix
hortensis Müll.; C, Helix aspersa
Müll. (After Ashford.)
 The dart is almost confined to Helicidae, a certain number of
exceptions being known which border on Helix. Hyalinia nitida and
excavata are the only British species, not Helices, which are known
to possess it. It has not been noticed to occur in the slugs, except in
the N. American genus Tebennophorus. About one-third of the
British Helices are destitute of the dart.[260] H. rufescens possesses
a double bilobed sac, but only two darts, which lie in the lower lobes.
It does not use the darts, and could not do so, from the relative sizes
of dart and sac; it has often been watched when uniting, but the use
of the darts has never been observed. From this it has been inferred
that the darts are degenerate weapons of defence, and that they
were in fact at one time much stronger organs and more often used.
[261] This theory, however, does not seem consistent with the whole
circumstances of the occurrence, position, and present use of the
darts.
Hermaphrodite Mollusca.—(b) Digonopora.—As an example of
the Digonopora or hermaphrodite Mollusca with separate generative
apertures for the male and female organs, we may take the common
Limnaea stagnalis (Fig. 55). It will be seen from the figure that the
relative positions of the hermaphrodite gland and duct, and of the
albumen gland, are the same as in Helix. When the oviduct parts
company from the vas deferens, it becomes furnished with several
accessory glands, one of which (Gl.E.) probably serves as a
reservoir for the ova, and answers more or less to a uterus. The tube
leading to the spermatheca is short, and there is no divergent
caecum. The female orifice lies near to the external opening of the
branchial cavity. The vas deferens, which is very long, is furnished
with a large prostate gland. The penis sac is greatly dilated, and
there is no flagellum. The male orifice is behind the right tentacle,
slightly in advance of the female orifice (compare Fig. 102).
Fig. 55.—Genitalia of Limnaea stagnalis
L. (from a dissection by F. B. Stead),
× 2.
 A.G, albumen gland.
Ac.G, accessory gland.
F.O, female orifice.
Gl.E, glandular enlargement.
H.D, hermaphrodite duct.
H.G, hermaphrodite gland.
Li, liver.
M.O, male orifice.
P, penis sac.
Pr, prostate.
R.M, retractor muscle of penis.
Sp, spermatheca.
V.D, vas deferens.]

Most of the Opisthobranchiata, but not all, have separate sexual

orifices. Numerous variations from the type just described will be
found to occur, particularly in the direction of the development of
accessory glands, which are sometimes very large, and whose
precise purpose has in many cases not been satisfactorily
determined.
Pelecypoda.—In the dioecious Pelecypoda, which form the great
majority, the reproductive system is simple, and closely parallel in
both sexes. It consists of a pair of gonads, which are either ovaries
or testes, and a pair of oviducts or sperm-ducts which lead to a
genital aperture. The gonads are usually placed symmetrically at the
sides or base of the visceral mass. The oviduct is short, and the
genital aperture is usually within the branchial chamber, thus
securing the fertilisation of the ova by the spermatozoa, which are
carried into the branchial chamber with the water which passes
through the afferent siphon.
Hermaphrodite Pelecypoda are rare, the sexes being usually
separate. The following are assured instances: Pecten glaber, P.
jacobaeus, P. maximus, Ostrea edulis, Cardium norvegicum,
Pisidium pusillum, Cyclas cornea, Pandora rostrata, Aspergillum
dichotomum, and perhaps Clavagella. The greater number of these
have only a single genital gland (gonad) on each side, with a single
efferent duct from each, but part of the gland is male and part
female, e.g. in the Pectens above mentioned. Pandora and
Aspergillum have two distinct glands, respectively male and female,
on each side, each of the two glands possessing its separate duct,
and the two ducts from each side eventually opening near one
another. It appears probable that the Septibranchiata (Cuspidaria,
Poromya, Lyonsiella, etc.) must also be added to the number of
hermaphrodite Pelecypoda which have separate male and female
glands.
It is worthy of remark that all the hermaphrodite Pelecypoda
belong to forms decidedly specialised, while forms distinctly
primitive, such as Nucula, Solenomya, Arca, and Trigonia are all
dioecious. In Gasteropoda similarly, the least specialised forms (the
Amphineura, with the exception of the Neomeniidae, and the
Rhipidoglossa) are dioecious. It is possible therefore that in the
ancestors of the Mollusca the separation of the sexes had already
become the normal type of things, and that hermaphroditism in the
group is, to a certain extent, a sign or accompaniment of
specialisation.[262]
Development of Fresh-water Bivalves.—The vast majority of
fresh-water bivalves either pass the larval stage entirely within the
mother, and do not quit her except in a perfectly developed form
(Cyclas, Pisidium), or assume a mode of development in which free
larvae indeed occur, but are specially modified for adaptation to
special circumstances (Unio). Cyclas and Pisidium, and no doubt all
the kindred genera, preserve their ova in a sort of brood-pouch
within the gills, in which the ova pass the earlier stages of their
development. But, even so, the larva of these genera retains some
traces of its original free-swimming habits, for a rudimentary velum,
which is quite useless for its present form of development, has been
detected in Cyclas.
The larva of Dreissensia (see Fig. 47, A), so far as is at present
known, stands alone among fresh-water bivalves in being free-
swimming, and to this property has been attributed, no doubt with
perfect justice, the fact of the extraordinarily rapid spread of
Dreissensia over the continent of Europe (chap. xvi.). In expelling the
ova, the parent slightly opens the shells and then quickly closes
them, shooting out a small point of white slime, which is in fact a little
ball of eggs. The general course of development is precisely parallel
to that of marine Pelecypoda, greatly resembling, so far as form is
concerned, certain stages in the growth of the larvae of Modiolaria
and Cardium, as figured by Lovén.[263]
In June and July the larvae appear in large numbers on the
surface of the water, when in spite of their exceedingly small size,
they can be captured with a fine hand-net. They pass about eight
days on the surface, feeding apparently on minute floating algae.
During this time, the principal change they undergo is in the
formation of the foot, which first appears as a small prominence
midway between the mouth and anus, and gradually increases in
length and flexibility. When the larva sinks to the bottom, the velum
soon disappears entirely, the foot becomes exceedingly long and
narrow, while the shell is circular, strongly resembling a very young
Cyclas.
Larvae of Unionidae.—The early stages of the development of
Unio and Anodonta (so far as the species of North America, Europe,
and Asia are concerned) is of extreme interest, from the remarkable
fact that the young live for some time parasitically attached to certain
species of fresh-water fishes. In order to secure this attachment, the
larva, which is generally known as Glochidium, develops a long
filament which perhaps renders it aware of the neighbourhood of a
fish, and also a larval shell furnished with strong hooks by which it
fastens itself to the body of its unconscious host (Fig. 56). According
to some interesting observations made by Mr. O. H. Latter,[264] the
ova pass into the external gill of the mother, in which is secreted a
nutritive mucus on which they are sustained until they arrive at
maturity and a suitable opportunity occurs for their ‘being born.’ If
this opportunity is deferred, and the Glochidia mature, their so-called
‘byssus’ becomes developed, and by being entangled in the gill
filaments of the parent, prevents their escaping. It is interesting to
notice that, when the nutritive mucus of the parent is used up, it
becomes, as it were, the turn of the children to provide for
themselves a secondary mode of attachment.
 Fig. 56.—A, Glochidium immediately after it is hatched: ad,
adductor muscle; by, ‘byssus’ cord; s, sense organs; sh,
shell. B, Glochidium after it has been on the fish for some
weeks: a.ad, p.ad, anterior and posterior adductors; al,
alimentary canal; au.v, auditory vesicle; br, branchiae; f,
foot; mt, mantle. (Balfour.)
The mother Anodonta does not always retain the Glochidium until
fish are in her neighbourhood. Gentle stirring of the water caused
them to emit Glochidium in large masses, if the movement was not
so violent as to cause alarm. The long slimy masses of Glochidium
were observed to be drawn back again within the shell of the mother,
even after they had been ejected to a distance of 2 or 3 inches.
It is a mistake to assert that the young Glochidium can swim.
When they finally quit the mother, they sink to the bottom, and there
remain resting on their dorsal side, with the valves gaping upwards
and the so-called byssus streaming up into the water above them.
There they remain, until a convenient ‘host’ comes within reach, and
if no ‘host’ comes within a certain time, they perish. They are
evidently peculiarly sensitive to the presence of fish, but whether
they perceive them by smell or some other sense is unknown. “The
tail of a recently killed stickleback thrust into a watch-glass
containing Glochidium throws them all into the wildest agitation for a
few seconds; the valves are violently closed and again opened with
astonishing rapidity for 15–25 seconds, and then the animals appear
exhausted and lie placid with widely gaping shells—unless they
chance to have closed upon any object in the water (e.g. another
Glochidium), in which case the valves remain firmly closed.”
In about four weeks after the Glochidium has quitted its host, and
the permanent shell has made its appearance within the two valves
of the Glochidium, the projecting teeth of the latter press upon the
ventral edge of the permanent shell, at a point about half way in its
lengthward measurement, retarding the growth of the shell at that
particular point, and indenting its otherwise uninterrupted curve with
an irregular notch or dent. As growth proceeds, this dent becomes
less and less perceptible on the ventral margin of the shell itself, but
its effects may be detected, in well-preserved specimens, by the
wavy turn in the lines of growth, especially near the umbones of the
young shell.
Mr. Latter found that all species of fish with which he
experimented had a strong dislike to Glochidium as an article of
food. Sometimes a fish would taste it “just to try,” but invariably spit it
out again in a very decided manner. The cause of unpleasantness
seemed not to be the irritation produced in the mouth of the fish by
the attempt of the Glochidium to attach itself, but was more probably
due to what the fish considered a nasty taste or odour in the object
of his attentions.

The following works will be found useful for further study of this
portion of the subject:—
F. M. Balfour, Comparative Embryology, vol. i. pp. 186–241.
F. Blochmann, Ueber die Entwickelung von Neritina fluviatilis Müll.:
Zeit. wiss. Zool. xxxvi. (1881), pp. 125–174.
L. Boutan, Recherches sur l’anatomie et le développement de la
Fissurelle: Arch. Zool. exp. gén. (2) iii. suppl. (1885), 173 pp.
W. K. Brooks, The development of the Squid (Loligo Pealii Les.):
Anniv. Mem. Bost. Soc. Nat. Hist. 1880.
„ „ The development of the oyster: Studies Biol. Lab.
Johns Hopk. Univ. i. (1880), 80 pp.
 R. von Erlanger, Zur Entwickelung von Paludina vivipara: Morph.
Jahrb. xvii. (1891), pp. 337–379, 636–680.
„ „ Zur Entwickelung von Bythinia tentaculata: Mitth. Zool.
Stat. Neap, x. (1892), pp. 376–406.
H. Fol, Sur le développement des Ptéropodes: Arch. Zool. exp.
gén. iv. (1875), pp. 1–214.
„ Etudes sur le développement des Mollusques.
Hétéropodes: ibid v. (1876), pp. 105–158.
„ Etudes sur le développement des Gastéropodes pulmonés:
ibid. viii. (1880), pp. 103–232.
H. Grenacher, Zur Entwickelungsgeschichte der Cephalopoden:
Zeit. wiss. Zool. xxiv. (1874), pp. 419–498.
B. Hatschek, Ueber Entwickelungsgeschichte von Teredo: Arb.
Zool. Inst. Univ. Wien, iii. (1881), pp. 1–44.
R. Horst, On the development of the European oyster: Quart.
Journ. Micr. Sc. xxii. (1882), pp. 339–346.
E. Korschelt and K. Heider, Lehrbuch der vergleichenden
Entwickelungsgeschichte der wirbellosen Thiere, Heft iii. (1893), pp.
909–1177 (the work is in process of translation into English).
A. Kowalewsky, Embryogénie du Chiton polii avec quelques
remarques sur le développement des autres Chitons: Ann. Mus. Hist.
Nat. Mars. Zool. i. (1883), v.
E. Ray Lankester, Contributions to the developmental history of
the Mollusca: Phil. Trans. Roy. Soc. vol. 165 (1875), pp. 1–31.
„ „ Observations on the development of the pond-snail
(Lymnaeus stagnalis), and on the early stages of other Mollusca:
Quart. Journ. Micr. Sc. xiv. (1874), pp. 365–391.
„ „ Observations on the development of the Cephalopoda:
ibid. xv. (1875), pp. 37–47.
W. Patten, The embryology of Patella: Arb. Zool. Inst. Univ. Wien,
vi. (1886), pp. 149–174.
M. Salensky, Études sur le développement du Vermet: Arch. Biol.
vi. (1885), pp. 655–759.
L. Vialleton, Recherches sur les premières phases du
développement de la Seiche (Sepia officinalis): Ann. Sc. Nat. Zool. (7)
vi. (1888), pp. 165–280.
 S. Watase, Observations on the development of Cephalopods:
Stud. Biol. Lab. Johns Hopk. Univ. iv. (1888), pp. 163–183.
„ „ Studies on Cephalopods: Journ. Morph. iv. (1891), pp.
247–294.
E. Ziegler, Die Entwickelung von Cyclas cornea Lam.: Zeit. wiss.
Zool. xli. (1885), pp. 525–569.
 CHAPTER VI
RESPIRATION AND CIRCULATION—THE MANTLE

The principle of respiration is the same in the Mollusca as in all

other animals. The blood is purified by being brought, in successive
instalments, into contact with pure air or pure water, the effect of
which is to expel the carbonic acid produced by animal combustion,
and to take up fresh supplies of oxygen. Whether the medium in
which a mollusc lives be water or air, the effect of the respiratory
action is practically the same.
Broadly speaking, Mollusca whose usual habitat is the water
‘breathe’ water, while those whose usual habitat is the land ‘breathe’
air. But this rule has its exceptions on both sides. The great majority
of the fresh-water Mollusca which are not provided with an
operculum (e.g. Limnaea, Physa, Planorbis), breathe air, in spite of
living in the water. They make periodic visits to the surface, and take
down a bubble of air, returning again for another when it is
exhausted. On the other hand many marine Mollusca which live
between tide-marks (e.g. Patella, Littorina, Purpura, many species of
Cerithium, Planaxis, and Nerita) are left out of the water, through the
bi-diurnal recess of the tide, for many hours together. Such species
invariably retain several drops of water in their branchiae, and, aided
by the moisture of the air, contrive to support life until the water
returns to them. Some species of Littorina (e.g. our own L. rudis and
many tropical species) live so near high-water mark that at neap-
tides it must frequently happen that they are untouched by the sea
for several weeks together, while they are frequently exposed to a
burning sun, which beats upon the rocks to which they cling. In this
case it appears that the respiratory organs will perform their
functions if they can manage to retain an extremely small amount of
moisture.[265]
The important part which the respiratory organs play in the
economy of the Mollusca may be judged from the fact that the
primary subdivision of the Cephalopoda into Dibranchiata and
Tetrabranchiata is based upon the number of branchiae they
possess. Further, the three great divisions of the Gasteropoda have
been named from the position or character of the breathing
apparatus, viz. Prosobranchiata, Opisthobranchiata and Pulmonata,
while the name Pelecypoda has hardly yet dispossessed
Lamellibranchiata, the more familiar name of the bivalves.
Respiration may be conducted by means of—(a) Branchiae or
Gills, (b) a Lung or Lung-cavity, (c) the outer skin.
In the Pelecypoda, Cephalopoda, Scaphopoda, and the great
majority of the Gasteropoda, respiration is by means of branchiae,
also known as ctenidia[266], when they represent the primitive
Molluscan gill and are not ‘secondary’ branchiae (pp. 156, 159).
In all non-operculate land and fresh-water Mollusca, in the
Auriculidae, and in one aberrant operculate (Amphibola), respiration
is conducted by means of a lung-cavity, or rarely by a true lung,
whence the name Pulmonata. The land operculates (Cyclophoridae,
Cyclostomatidae, Aciculidae, and Helicinidae) also breathe air, but
are not classified as Pulmonata, since other points in their
organisation relate them more closely to the marine
Prosobranchiata. Both methods of respiration are united in
Ampullaria, which breathes indifferently air through a long siphon
which it can elevate above the surface of the water, and water
through a branchia (see p. 158). Siphonaria (Fig. 57) is also
furnished with a lung-cavity as well as a branchia. Both these genera
may be regarded as in process of change from an aqueous to a
terrestrial life, and in Siphonaria the branchia is to a great extent
atrophied, since the animal is out of the water, on the average,
twenty-two hours out of the twenty-four. In the allied genus Gadinia,
where there is no trace of a branchia, but only a lung-cavity, and in
Cerithidea obtusa, which has a pulmonary organisation exactly
analogous to that of Cyclophorus,[267] this process may be regarded
as practically completed.
 Fig. 57.—A, Siphonaria gigas
Sowb., Panama, the
animal contracted in
spirit: gr, siphonal groove
on right side. B, Gadinia
peruviana, Sowb., Chili,
shell only: gr, mark of
siphonal groove to right
of head.
Respiration by means of the skin, without the development of any
special organ, is the simplest method of breathing which occurs in
the Mollusca. In certain cases, e.g. Elysia, Limapontia, and Cenia
among the Nudibranchs, and the parasitic Entoconcha and
Entocolax, none of which possess breathing organs of any kind, the
whole outer surface of the body appears to perform respiratory
functions. In others, the dorsal surface is covered with papillae of
varied size and number, which communicate with the heart by an
elaborate system of veins. This is the case with the greater number
of the Aeolididae (Fig. 58, compare Fig. 5, C), but it is curious that
when the animal is entirely deprived of these papillae, respiration
appears to be carried on without interruption through the skin.
 Fig. 58.—Aeolis despecta
Johnst., British coasts.
(After Alder and Hancock.)
In the development of a distinct breathing organ, it would seem as
if progress had been made along two definite lines, each resulting in
the exposure of a larger length of veins, i.e. of a larger amount of
blood, to the simultaneous operation of fresh air or fresh water.
Either (a) the skin itself may have developed, at more or less regular
intervals, elevations, or folds, which gradually took the form of
papillae, or else (b) an inward folding, or ‘invagination,’ of the skin, or
such a modification of the mantle-fold as is described below (p. 172)
may have taken place, resulting in the formation of a cavity more or
less surrounded by walls, within which the breathing organs were
ultimately developed. Sometimes a combination of both processes
seems to have occurred, and after a papilliform organ has been
produced, an extension or prolongation of the skin has taken place,
in order to afford a protection to it. Respiration by means of a lung-
cavity is certainly subsequent, in point of time, to respiration by
means of branchiae.

Fig. 59.—Chiton squamosus

L., Bermuda: A, anus;
Br, branchiae; M, mouth.

Fig. 60.—Fissurella virescens

Sowb., Panama, showing
position of the branchiae: Br,
branchiae: E, E, eyes; F, foot;
M, mantle; T, T, tentacles.
The branchiae seem to have been originally paired, and arranged
symmetrically on opposite sides of the body. It is not easy to decide
whether the multiple form of branchia which occurs in Chiton (Fig.
59), or the simple form as in Fissurella (Fig. 60), is the more
primitive. Some authorities hold that the multiple branchia has
gradually coalesced into the simple, others that the simple form has
grown, by serial repetition, into the multiple. There appears to be no
trace of any intermediate forms, and, as a matter of fact, the multiple
branchia is found only in the Amphineura, while one or rarely two
(never more) pairs of branchiae, occur, with various important
modifications, in the vast majority of the Mollusca.
Amphineura.—In Chiton the branchiae are external, forming a
long row of short plumes, placed symmetrically along each side of
the foot. The number of plumes, at the base of each of which lies an
osphradial patch, varies from about 70 to as few as 6 or 7. When the
plumes are few, they are confined to the posterior end, and thus
approximate to the form and position of the branchiae in the other
Amphineura. In Chaetoderma, the branchiae consist of two small
feather-shaped bodies, placed symmetrically on either side of the
anus, which opens into a sort of cloaca within which the branchiae
are situated. In Neomenia the branchiae are still further degraded,
consisting of a single bunch of filaments lying within the cloaca, while
in Proneomenia there is no more than a few irregular folds on the
cloaca-wall (Fig. 61).

Fig. 61.—Terminal portions of the Amphineura, illustrating the gradual degradation

of the branchiae, and their grouping round the anus in that class. A, Chiton
(Hemiarthrum) setulosus Carp., Torres Str.; B, Chiton (Leptochiton) benthus
Hadd., Torres Str.; C, Chaetoderma; D, Neomenia; a, anus; br, br, branchiae;
k, k, kidneys; p, pericardium. (A and B after Haddon, C and D after Hubrecht.)