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throws light on a much contested subject; namely, the nature of the
so-called "intertentacular organ" (i, Fig. 234, p. 469), described so
long ago as 1837 by Farre,[573] but looked for in vain by the majority
of later observers.

The failure to find this organ, even in species which possess it, in
certain individuals, according to Farre's statements, is now
satisfactorily explained by M. Prouho, who shows that while it is
absent in a large number of polypides, it is normally present in those
individuals which possess an ovary, and in those only; and that its
primary function is that of an oviduct.

The intertentacular organ is an unpaired ciliated tube, which is

situated between the two tentacles which are nearest to the
ganglion. In the retracted condition of the polypide, it opens from the
body-cavity into the tentacle-sheath; and in the expanded condition,
directly to the exterior.

In the remarkable case of Alcyonidium duplex, each zooecium

normally possesses two sexual polypides. The first of these
produces a testis and then becomes a "brown body." The second is
meanwhile developed, and produces an ovary and an intertentacular
organ, a structure which was not present in the male polypide. The
eggs pass through the intertentacular organ into the tentacle-sheath,
and attach themselves to the diaphragm (d, Fig. 234), where they
remain during their development.

Although the intertentacular organ has been found by Prouho in

female polypides only, it would perhaps be going too far to assert
that it is confined to polypides of that sex. Hincks[574] has observed
the passage of spermatozoa in enormous numbers through the
organ, although it may be noted that there is no sufficient proof that
eggs were not present as well in these zooecia. It further appears
that in some cases waste matters may be removed from the body-
cavity through the same passage.
It may be presumed that the egg is normally fertilised by a
spermatozoon, although this is at present largely a matter of
inference. It is believed by Joliet[575] that fertilisation is reciprocal,
although Prouho has come to the opposite conclusion. Joliet has,
however, very justly pointed out that the enormous number of
spermatozoa developed by a single individual would be
disproportionately large, if their function were merely to fertilise the
ovum in the same zooecium. According to his view, the egg is
fertilised by a spermatozoon after it has passed into the tentacle-
sheath or ovicell, or some other place where it is in free
communication with the outside water.

Development and Affinities.—Few parts of the history of the

Polyzoa are more fascinating than that which deals with their
development; and it is probable that no other is capable of giving so
much insight into the affinities of the several groups to one another
and to other groups of the animal kingdom.

Fig. 252.—Diagrams of larvae. A, Loxosoma, × 208; a, anus; b, brain,

with left eye and ciliated pit; c, ciliated ring; ep, epistome; m,
mouth; o, oesophagus; st, stomach; x, aboral adhesive organ: B,
Cyphonautes larva of Membranipora (Electra) pilosa, × about 90;
a, m, o, st as in A; c, anterior part, and c', posterior part of the
ciliated ring; e, epidermis; ms, adductor muscle of shells; p,
pyriform organ, of unknown function; sh, shell; v, vestibule; the
"internal sac" or sucker, by which fixation is effected, is seen
between a and ms. (B, after Prouho.)
The comparative study of the larvae of the Polyzoa may be said to
date from 1877, when J. Barrois published an elaborate
Monograph[576] on this subject. Although some of Barrois' earlier
opinions have been subsequently modified, this work still gives the
best figures of the external form of the beautiful larvae of many
genera. A detailed account of the larval forms of Polyzoa must be
omitted from want of space; and the general conclusions only can be
given.

The larvae of the Entoprocta (Fig. 252, A) resemble the so-called

"Trochosphere" of Polychaeta (see p. 274). The common characters
shared by the larvae of Chaetopoda, Echiuroid Gephyrea, Mollusca,
and Polyzoa, and by adult Rotifera, may well point to the derivation
of these groups from a common ancestor. On this assumption, it is
possible that the Polyzoa have been derived from forms which
existed long ages ago, which combined the common characters of
these groups, and the structure of which we can picture to ourselves
only so far as the "Trochosphere" larva can be taken to represent it
in a much simplified condition. Such a view harmonises well with the
great antiquity of the Polyzoa. Certain Ectoproct forms have a larva,
known as Cyphonautes (Fig. 252, B), which closely resembles the
larval form of the Entoprocta; and it is a fact which probably has
considerable significance that this type of larva is known to occur
only in those species of Membranipora (Electra), Alcyonidium, and
Hypophorella, which lay eggs.[577] This may perhaps be regarded as
a primitive form of development which has been lost in species in
which development takes place inside the parent. Cyphonautes
compressus (Fig. 252, B), one of the commonest objects taken in the
surface-net off our own coasts, is the larva of Membranipora
(Electra) pilosa. Whilst this larva is provided with a well-developed
alimentary canal, those of most other Ectoprocta possess a mere
rudiment of this structure, and depend for their nutrition either on
yolk present in the egg or on material supplied by the parent. In most
cases the mature larva has no recognisable trace of a digestive
system; and, although it has a free-swimming period, it does not
become truly pelagic.
The alimentary canal of the larva of Pedicellina is known to persist in
the primary individual of the colony. In all other known cases, even in
that of Cyphonautes, the larva at fixation loses practically all its
internal organs, and becomes a mere body-wall containing a mass of
degenerated larval tissues. It is in fact a zooecium containing a
"brown body." A polypide-bud is now developed, the body-cavity
appears as the result of the shrinkage of the "brown body," and the
primary individual of the colony is thereby established.

The larvae of the Ectoprocta form a tolerably complete series,

starting from Cyphonautes, itself allied to the larva of the Entoprocta,
and ending with the Phylactolaemata. Alcyonidium (Fig. 253, B)
possesses a rudimentary alimentary canal,[578] although the most
conspicuous structures are those connected with the fixation and
other phenomena of larval life. The larvae of many of the encrusting
Cheilostomes (Fig. 253, A) resemble that of Alcyonidium, while those
of Bugula, Scrupocellaria, etc., belong to a type easily derivable from
that of the encrusting forms. The branching Ctenostomes
(Bowerbankia, etc.) have a larva which may be regarded as derived,
along slightly different lines, from that of Alcyonidium. The
Cyclostomata and the Phylactolaemata have the most modified
forms of larva. That of the former group may owe some of its
peculiarities to the occurrence of a remarkable process of embryonic
fission, which takes place in the ovicell, and as the result of which
each egg gives rise to a large number of larvae.[579] The
Phylactolaemata have a larva which is not unlike that of
Bowerbankia.
 Fig. 253.—A, Aboral view of free larva of Lepralia foliacea Ell. and Sol.;
a, long cilia of pyriform organ; g, aboral groove: B, longitudinal
section of embryo of Alcyonidium, × 135; c, ciliated ring; g, aboral
groove; m, mouth; n, nervous system; p, "pyriform organ," of
unknown function; s, "internal sac" or "sucker," by which fixation is
effected; st, stomach.

We have seen that the larva at fixation becomes a zooecium, which

in the Gymnolaemata forms a polypide-bud after fixation. The
peculiarities of the Phylactolaematous larva may be explained by
assuming that it becomes a zooecium while it is still free-swimming.
Thus the larva of Plumatella develops one or sometimes two
polypides, which actually reach maturity before fixation takes place.
That of Cristatella develops from two to twenty[580] polypides or
polypide-buds at the corresponding period, and it is in fact a young
colony while still free-swimming.

Now in most colonial animals, such as Coelenterates and Ascidians,

the larva metamorphoses itself into a temporarily solitary animal,
which then gives rise to the remainder of the colony by budding. The
majority of the Gymnolaemata behave in this way; while the
Phylactolaemata may not only develop a multiplicity of polypides in
their larval stage, but the individuality of the zooecia is then just as
much obscured as in the adult state. These facts are more easily
explained if we assume that Cristatella is the end-point in a series
than if we suppose it to be a starting-point.
On the view maintained by many authorities, that the Polyzoa are
related, through Phoronis, with the Gephyrea and the Brachiopoda,
we should expect to find in those Polyzoa which most closely
resemble Phoronis in their adult state—that is to say in the
Phylactolaemata—some indications of affinity to that animal in their
development. This is emphatically not the case. The hypothesis that
the Phylactolaemata are related to Phoronis leads, moreover, to the
improbable conclusion that the similarities between the Entoproct-
larva and Cyphonautes, on the one hand, and the Trochosphere
larva of Polychaeta, on the other hand, is entirely superficial and
meaningless. In spite, therefore, of the similarity between Phoronis
and a single individual of the Phylactolaemata, and in spite of the
marked resemblance between its nephridia and structures which
have been described in Cristatella[581] and Pectinatella[582] the
comparative study of the development appears to indicate that the
resemblances between Phoronis and the Phylactolaemata are the
result of a coincidence rather than of any close relationship.

A few points connected with the metamorphosis of the Polyzoa

deserve more special notice. There is generally great difficulty in
persuading larvae to fix themselves when kept in a small quantity of
water, which becomes over-heated in the air of a laboratory. The
difficulty may be surmounted by placing colonies containing
embryos, together with some clean pieces of the seaweed on which
the adults are habitually found, in a vessel closed by a piece of fine
muslin, and by leaving the vessel attached to a buoy or in a deep
tide-pool. The larvae being without an alimentary canal, fix
themselves, after a very short free life, on the seaweed.

It is probable that a great struggle for existence normally takes place

at the commencement of the metamorphosis. Any one who will
examine, in June or July, rocks covered by Fucus on which Flustrella
hispida is growing, will probably find numerous young fronds of
Fucus, from half an inch to an inch or two in length, growing under
the shelter of the older fronds. The bivalve larvae of Flustrella show
a marked preference for fixing on these young fronds—perhaps in
order that the duration of life of the colony may coincide with that of
the Fucus—and these young fronds are commonly covered by very
numerous recently-fixed larvae, and by young colonies of various
ages. Or, it is easy to observe, by placing pregnant colonies of
Bowerbankia in a vessel of water, that the larvae, which are hatched
out in thousands, fix themselves in dense masses on certain parts of
the wall of the vessel. It is clear that but a small proportion of these
larvae will find room for further development.

Next with regard to the mode of fixation. Attachment always takes

place by the surface on which the mouth or its rudiment is situated,
and the permanent alimentary canal opens on the opposite surface.
In Pedicellina, the one case in which the larval digestive organs are
known to become those of the first adult individual, this presupposes
a rotation of the alimentary canal, in order to bring it into its new
position.

It is well known that the larvae of other fixed animals may undergo a
somewhat similar change. Thus those of Ascidians and of Barnacles
fix themselves by their anterior end, and ultimately reach their adult
form by performing a kind of a somersault. The process may
perhaps be explained by supposing that some part of the anterior
end or of the oral surface is specially sensitive, and that the larva
fixes itself by that portion of its body which is best fitted for
ascertaining which is the proper substance on which to fix.

Budding.—The formation of a new individual may take place by the

outgrowth of part of the body-wall, as in Pedicellina (Fig. 243, p. 487)
and in Bowerbankia (Fig. 238, p. 480). In Pedicellina a young stalk is
formed by an outgrowth near one of the growing points, and the
upper part of this outgrowth becomes constricted off to form the
calyx. In other cases (cf. the growing ends of the branches in Fig.
237) a partition grows across the body-cavity at the growing edge of
the colony, and so cuts off a part destined to become a new
zooecium.
The zooecium formed in one of these ways acquires an alimentary
canal by the formation of a polypide-bud, some stages in the growth
of which are shown in Fig. 235 (p. 472). Contrary to what happens in
Coelenterates and Tunicates, in which the endoderm takes part in
the budding, there is good reason for believing that in Polyzoa the
polypide-bud is developed entirely from ectoderm and mesoderm.
[583] The bud is a two-layered vesicle, attached to the inner side of
the body-wall. Its inner layer is derived from the ectoderm, which at
first projects into the body-cavity in the form of a solid knob
surrounded by mesoderm-cells. A cavity appears in the inner,
ectodermic mass, and the upper part of the vesicle so developed
becomes excessively thin, forming the tentacle-sheath, which is
always developed in the condition of retraction. The lower part
becomes thicker; its inner layer gives rise to the lining of the
alimentary canal, to the nervous system, and to the outer epithelium
of the tentacles, which grow out into the tentacle-sheath (cf. Fig.
235). The outer layer gives rise to the mesodermic structures, such
as the muscles, connective tissue, and generative organs.

These processes are fundamentally similar, whether in the

metamorphosed larva, in a young zooecium, in an old zooecium
after the formation of a "brown body," or in the germinating statoblast
of the Phylactolaemata.

CHAPTER XIX

POLYZOA (continued)

CLASSIFICATION—GEOGRAPHICAL DISTRIBUTION—PALAEONTOLOGY—
METHODS FOR THE EXAMINATION OF SPECIFIC CHARACTERS—
TERMINOLOGY—KEY FOR THE DETERMINATION OF THE GENERA OF
BRITISH MARINE POLYZOA
Our account of the Polyzoa would be manifestly incomplete without
some reference to the systematic arrangement of these animals. An
outline of the principal groups has been given on p. 475. So far, the
classification is easy, but it is otherwise when we attempt to
subdivide most of the groups any further.

Systems of classification which depend exclusively upon the external

characters of animals have been repeatedly shown to be
unsatisfactory. Now with regard to the Polyzoa, not only is it the case
that the great majority of forms are only known in their external
characteristics, but current systems of classification cannot be
regarded as final, because it is not yet certain which of the external
features have most systematic value. Two obvious points can be at
once selected—namely, the character of the zooecium and the
character of the entire colony. One or two instances will serve to
show what different results are obtained by depending exclusively on
either of these characters by itself.

According to the older writers, the habit of the colony was taken as
the most important generic character; and there can indeed be no
doubt that this feature has great importance within certain limits. Any
one who has examined different species of such genera as Flustra,
Cellaria, Bugula, Retepora, etc., must feel that the form of the colony
goes for a good deal. But a consideration of other cases shows that
there is great risk in the indiscriminate use of this method of
arranging the Polyzoa. The old genus Eschara, composed of forms
with an erect coral-like habit,[584] included species which are now
placed in such different genera as Lepralia, Porella, Microporella,
etc. The older works on Polyzoa include all encrusting forms of
Cheilostomata, with a completely calcareous front wall, in the genus
Lepralia, the members of which are now distributed in numerous
widely separated genera.

As an instance of the converse arrangement—essential similarity of

the zooecia with great differences of the general habit—may be
mentioned the common Membranipora (Electra) pilosa.[585]
Ordinarily growing in the form of close encrustations on seaweeds,
this species may take on entirely different habits of growth. The
zooecia are now dissociated, growing in single lines over the
substratum; now forming erect tufts, composed of single lines of
zooecia or of several rows. The erect, branching habit appears to be
induced in the first instance by the character of the seaweed on
which the colony begins life. Thus colonies which encrust the thin
branches of Corallina may have impressed on them something of the
mode of growth of the seaweed, so that when they extend beyond
the tips of the branches of the Corallina, they continue to grow in
delicate branches, which still retain more or less the same diameter
as those which form their base. An extreme variation results in the
beautiful form known as Electra verticillata, in which the zooecia are
arranged with great regularity in whorls, which together form erect
branches.[586] But with all these variations, the zooecia are so much
alike that it is hardly possible to regard the extreme forms as more
than varieties of a single species. A careful examination of this case
would convince most observers that the characters of the zooecium
are a more trustworthy guide to classification than those of the entire
colony, a result which was first clearly stated by Smitt, and amply
confirmed by Hincks.[587]

The avicularia of the Cheilostomata afford useful help in classifying

this group; but while certain genera are always provided with
avicularia, others include some species with these organs, and other
species without them. Again, while the species of some genera (e.g.
Cellepora) possess a great variety of forms of avicularia, the same
pattern of avicularium may characterise several widely different
genera. Further, the position of the avicularium may be very different
in species which are apparently closely related. Well-developed
vibracula, although constant in their occurrence in such forms as
Scrupocellaria (Fig. 254) and Caberea (Fig. 242), occur here and
there in species of encrusting forms which are ordinarily placed in
very different families.
Now although some of these discrepancies are perhaps due to
errors in classification, whereby species which are really allied have
been wrongly placed in distinct genera, this explanation would not
prove satisfactory in all cases. Thus in Bugula, a genus which is
specially characterised by the high development of its avicularia,
these organs are normally absent in B. neritina. The fact that this
species was rightly placed in the genus has been confirmed by the
discovery made by Waters[588] that avicularia occur in specimens
which are believed to be identical with that species.

Fig. 254.—A, Front view, and B, back view of part of a branch of

Scrupocellaria scabra, Van Ben., Durham Coast, × 43; a, lateral
avicularium; a', smaller median avicularium; ap, membranous
aperture; f, fornix; r, rootlet; s, seta of vibraculum; v.z, vibracular
zooecium.

1. The Cyclostomata appear to fall naturally into two main groups,

(A) the Articulata, including the Crisiidae (Fig. 237), distinguished
by their erect branches, divided at intervals by chitinous joints; and
(B) the Inarticulata, which include the remaining families, whether
erect or encrusting, agreeing in the negative character of being
unjointed.

2. The Cheilostomata consist of (A) the Cellularina, including the

flexible, erect forms, such as Bugula (Fig. 233) and Scrupocellaria
(Fig. 254); (B) the Flustrina, to which belong Flustra (Fig. 232),
Membranipora (Fig. 256, A, B), Micropora (Fig. 256, C), and other
forms in which the front wall of the zooecium is either membranous,
or depressed and marked off by a ridge-like margin; (C) the
Escharina, including the great majority of forms, in which no part of
the front wall remains membranous, the wall of the zooecium being
wholly calcified.

3. The Ctenostomata comprise (A) the Alcyonellea or encrusting

forms; and (B) the Vesicularina or branching forms. The zooecia in
the latter subdivision (Fig. 238) are given off from a tubular stem or
stolon, which is usually erect and branching.

We thus have the following arrangement of recent forms. The genera

mentioned are for the most part those which have already been
alluded to in the preceding account:—

Sub-class I. Entoprocta.
Loxosoma, Pedicellina, Urnatella.

Sub-class II. Ectoprocta.

Order 1. Gymnolaemata.
Sub-order 1. Cyclostomata.
A. Articulata. Crisia.
B. Inarticulata. Hornera, Idmonea, Tubulipora,
Stomatopora,
Diastopora, Entalophora, Lichenopora.
Sub-order 2. Cheilostomata.
A. Cellularina. Aetea, Eucratea,[589] Catenicella,
Cellularia, Gemellaria, Menipea, Scrupocellaria,
Caberea, Notamia (= Epistomia), Bicellaria, Bugula,
Beania.
B. Flustrina. Cellaria, Flustra, Membranipora, Electra,
Lunulites, Membraniporella, Cribrilina, Micropora,
Selenaria.
C. Escharina. Retepora, Microporella, Lepralia, Porella,
Smittia, Mucronella, Schizoporella, Schizotheca,
Mastigophora, Porina, Cellepora.
Sub-order 3. Ctenostomata.
A. Alcyonellea. Alcyonidium, Flustrella.
B. Vesicularina. Vesicularia, Amathia, Bowerbankia,
 Farrella, Hypophorella, Triticella, Mimosella,
Victorella, Paludicella.
Order 2. Phylactolaemata.
Fredericella, Plumatella (including Alcyonella), Lophopus,
Cristatella, Pectinatella.

Even this classification, which deals only with the larger groups,
must not be made use of without a word of warning. The division of
the Cheilostomata is a matter of great difficulty; and no scheme
which has yet been suggested can be regarded as more than
tentative. The great number of forms included in this group makes its
subdivision extremely desirable from the point of view of
convenience; but a further knowledge of the anatomy and of the
development of many of the forms of doubtful systematic position is
probably necessary before any scheme which is likely to be
permanent is put forward. Those who desire to make a further study
of the classification of the Polyzoa should refer to the works of
Hincks,[590] Busk,[591] MacGillivray,[592] and Gregory.[593]

The Polyzoa do not appear to lend any valuable assistance towards

settling the disputed problems of Geographical Distribution. They are
not in any case terrestrial, while the fresh-water species do not
always respect the limits between the great zoogeographical
regions. It has already been pointed out (p. 504) that Plumatella,
Fredericella, and Lophopus are believed to occur in Australia, and
the first-named genus is practically world-wide in its distribution.

Many marine forms also have a surprisingly wide distribution. Thus

among the British species which are described by Mr. Hincks as
occurring from Norway to New Zealand are Membranipora pilosa,
Scrupocellaria scruposa, Cellaria fistulosa, Microporella ciliata, and
M. malusii. Even if it should be proved that specific differences do
exist between the southern forms and our own, there can be no
doubt of the wide distribution of certain species. It was pointed out by
D'Orbigny that Bugula neritina has the habit of attaching itself to the
bottoms of ships, a fact which may possibly account for the wide
distribution of this species; although it would not be safe to assume
this explanation of the facts in all cases. Other Polyzoa, on the
contrary, have a more restricted range. Thus Catenicella is specially
characteristic of the Australian region.

It is perhaps surprising that marine Polyzoa should in so many cases

have so wide a range. Even though it is the rule for Polyzoa to have
free larvae, the period during which these larvae are free-swimming
is, so far as is known, a short one in most cases. Cyphonautes is a
common pelagic form (see p. 510), and probably remains for a
considerable period in the larval condition. Other Polyzoon-larvae
appear to fix themselves very soon after their birth; and this would
not appear to give much time for them to be carried to great
distances by ocean-currents. It may, however, be suggested that it
does not follow that because we know that a larva may, under
favourable conditions fix itself a few minutes after it becomes free,
we should be justified in assuming that that larva would not retain for
a long period the power of undergoing a normal metamorphosis
should it be drifted away from suitable fixing-grounds.

Palaeontology.[594]—The number of fossil Polyzoa is enormous.

D'Orbigny devoted two hundred plates and more than a thousand
octavo pages[595] to a Monograph on the Cretaceous Polyzoa of
France. Many of the fossil forms are extraordinarily well preserved,
and there is often no difficulty in recognising the identity between
certain fossil species belonging to the more recent formations and
living forms. It thus becomes necessary to consult Palaeontological
memoirs in working at recent Polyzoa.

While the great majority of fossil Polyzoa do not differ in any

essential particular from recent species, this is not altogether the
case with the Palaeozoic forms. Leaving out of account the
Stromatoporoids, which have been variously referred to the
Sponges, Hydrozoa, and Foraminifera, as well as to the Polyzoa, the
Palaeozoic strata contain large numbers of peculiar Cyclostomata,
together with members of the Trepostomata, a fourth Sub-order of
Gymnolaemata, allied to the Cyclostomata. The Trepostomata are
for the most part Palaeozoic, but a few survived as late as the
Jurassic period.[596] These, with the other Polyzoa from the same
formations, are considered by Dr. Gregory in his recently published
Catalogue of the Fossil Bryozoa in the British Museum (1896).

The number of Polyzoa recorded from the earlier secondary strata is

small. The majority of the known Jurassic forms belong to the
Cyclostomata; and one or two Cheilostomes are recorded from the
same period. Recent papers by Walford[597] on Jurassic Polyzoa
contain the description of genera which are believed to be
intermediate between the Cyclostomata and Cheilostomata,
particularly with regard to the characters of their ovicells. Although it
is not impossible there may be a connection between the ovicells of
these two groups, it has yet to be proved that the two sets of
structures are homologous.

The Cretaceous period marks the commencement of a large number

of Cheilostome genera, although the Cyclostomes still remain
numerous.

In the Tertiary formations the Cyclostomes gradually become less

numerous, and although in earlier geological periods they far
outnumbered the Cheilostomes, these relations are now reversed.
Certain Tertiary strata, and particularly the Coralline Crag (Pliocene),
are remarkable for the extremely large number of Polyzoa they
contain. It will be noticed that no mention has been made of the
Entoprocta, the Ctenostomata, and the Phylactolaemata. Their
absence in the fossil condition[598] need not, however, be a matter
for surprise, as none of these forms are so well suited for being
fossilised as are the calcareous Cyclostomata and Cheilostomata.
There is consequently no adequate reason for assuming that the
absence of a palaeontological record implies that these groups have
been recently evolved.
Determination of Genera of Marine Polyzoa.—The species to
which a Polyzoon belongs can only be determined, in most cases,
with the assistance of the low powers of a microscope. There are
very great advantages in the use of a binocular instrument, by
means of which a microscopic preparation appears with its parts
standing up in proper relief.

In the case of the calcareous forms, the external characters may be

more readily made out in a dry preparation than in any other way.
For this purpose, the colony should be washed with fresh water, in
order to remove the salts, which otherwise crystallise out on drying
and obscure the surface. Preparations of this kind must be looked at
with the aid of reflected light. Canada-balsam or glycerine
preparations are also valuable, whether stained or unstained; and
are essential for the examination of the softer forms. In the case of
erect species, both surfaces of the branch should be looked at. The
opercula, avicularia, and rosette-plates afford important systematic
characters in the case of the Cheilostomata.

It must not be forgotten to take account of the condition of the

zooecia at different ages. The old zooecia often become entirely
altered in form, by the deposition of additional calcareous matter, or
by the loss of certain parts present in the younger zooecia. Thus the
marginal spines may be entirely lost in the older individuals, while in
those forms which develop a "peristome" (see Fig. 255 and p. 524),
the characters of the orifice can often be determined in the young
zooecia only. It is thus essential to examine the growing ends of the
branches or the rim of the colony, as the case may be.

Fig. 255.—Illustrating the nature of a secondary orifice

(Cheilostomata). A, Mucronella coccinea Abildg., Scilly Is., × 40.
 The ovicell (o) overhangs the primary orifice, which is concealed
by the great development of the peristome, produced into the
mucro (mu); t, the three teeth (denticles) within the secondary
orifice; a, avicularium. B, Porella compressa Sowb., Norway, × 40;
p.o. primary orifice, above which is a concave lamina, the
beginning of the ovicell. In the lower zooecium the ovicell (o) is
further grown. The primary orifice is still visible, but it is partially
concealed by the growth of the peristome, which encloses a
minute avicularium; m, mandible of avicularium. C, Older part of
the same colony; pr, peristome; s.o, secondary orifice; o', adult
ovicell; p, pores.

In order to make preparations with the tentacles expanded,

hydrochlorate of cocaine, chloral hydrate or spirit should be added
gradually to the water. When the animals are completely
anaesthetised they may be killed by means of a 7-10 p.c. solution of
sulphate of copper (best made in distilled water or in rain water).
This method gives admirable results in the case of both fresh-water
and marine Polyzoa. The use of formaline (see p. 229) may be
strongly recommended for the Vesicularina.

The only recent work dealing with all the marine British forms is Mr.
Hincks' invaluable History of the British Marine Polyzoa.[599] As the
use of this book, unaided by any artificial help, is by no means easy
to the beginner, the following key has been compiled as an index to
the genera. The Entoproct forms, Loxosoma and Pedicellina (see pp.
488-491), are not included in the table.

Fig. 256.—Illustrating the terminology of the front surface of the

zooecium (Cheilostomata). A, Membranipora (Electra) pilosa L.,
Cromer, × 47; ap, the membranous "aperture;" o, orifice. B,
Membranipora flemingii Busk, Plymouth, × 60; ap, the aperture,
 enclosed in a calcareous "area" (a); av, avicularium; s, marginal
spines. C, Micropora coriacea Esper, Plymouth, × 43; a, area
(calcareous); o, operculum; ov, ovicell.

In order to facilitate the use of the table here given in conjunction

with Mr. Hincks' work, the nomenclature there adopted has been
followed throughout. References to other descriptions of the species
may be obtained by consulting Miss Jelly's admirable Synonymic
Catalogue of the Recent Marine Bryozoa.[600]

Terminology.—A few technical terms must of necessity be

employed. The colony is adherent when its zooecia are attached to
the object on which the colony is growing. The zooecium is the body-
wall of a single individual; and, except in transparent species, is the
only part which can be seen from the outside in the retracted
condition of the polypide or tentacles with the alimentary canal. The
outermost layer of the zooecium is known as the ectocyst; it may be
simply membranous, or calcified, or may be rendered opaque by
foreign bodies; its surface in calcareous forms is often marked by
pores (Fig. 239, C, p), which are vacuities in the calcareous wall,
closed externally by membrane. A special median pore (Fig. 241, A,
m.p) may occur, and is in some cases at least a complete perforation
through the body-wall.

The tentacles are protruded through the orifice, which in

Cheilostomata is usually guarded by a movable chitinous lid, or
operculum (Fig. 256, A, o). Should the ectocyst be thickened or
raised into a ridge surrounding the orifice, a tubular passage results,
known as the secondary orifice (Fig. 255), at the deeper end of
which is the true orifice. The peristome (Fig. 255, C, pr) is the raised
or thickened part which gives rise to the secondary orifice. Should
the zooecium be outlined by a raised ridge, the part so enclosed is
known as the area (Fig. 256, C, a), if calcareous. The aperture or
opesia (Fig. 256, A, B, ap) is a membranous part of the front surface;
and may consist of the whole or part of the area. The orifice or the
aperture is commonly provided with spines (Fig. 256, B, s).
 Fig. 257.—A, Cribrilina annulata Fabr., Norway, × 33; c, calcareous
bars concealing the membranous aperture: B, Membraniporella
nitida Johnst., Plymouth, × 45; a, calcareous bars growing up
round the margin of the aperture; b, the same, further developed;
c, the same, completely formed (as in A); av, avicularium; o,
immature, and o', mature, ovicell; s, marginal spines.

The avicularium and the vibraculum are specially modified zooecia

(see p. 482), which occur in a great variety of forms, in certain
Cheilostomata only. The operculum of the ordinary zooecium is
represented by the mandible (Fig. 239, B, m) in the avicularium, and
by the seta (Fig. 242, s) in the vibraculum. The representative of the
zooecium itself is known as the avicularian (Fig. 239, A, a.z) or
vibracular zooecium (Fig. 242, v.z).

An ovicell is a swelling in which the embryo develops, in certain

Cyclostomata (Fig. 237) and Cheilostomata (Fig. 241, A, o). A stolon
(Fig. 238, B, st) is a stem, not formed of fused zooecia, from which
new individuals originate. An internode, in a jointed colony, is the
part between any two joints. The fornix or scutum (Fig. 254, A, f) is a
modified spine which in some Cheilostomata overhangs the
aperture. A mucro (Fig. 255, A, mu) is a spike or protuberance
developed just below the orifice. A sinus (Fig. 239, B, s) is a slight
bay on the lower margin of the orifice.

The orifice opens at the upper end of the zooecium, on its front
surface. The length of the zooecium is the distance from the upper to
the lower ends, and the width the distance between its sides.

1. One or more of the following characters: orifice provided with

 an operculum; avicularia or vibracula present; a globular
ovicell above the orifice of certain zooecia (Cheilostomata)
7
Opercula, avicularia, vibracula, and ovicells completely
absent, or inconspicuous. Calcareous or non-calcareous. If
calcareous, the orifice is not at the end of a free cylindrical
portion
3
Calcareous; zooecia cylindrical, often united for the greater
part of their length, but usually ending in a free cylindrical
portion, which bears the terminal orifice. The zooecia may
be much obscured by calcifications surrounding their basal
parts
2

2. Zooecia long, tubular, with a lateral membranous region at

the upper end, given off quite separately from a creeping
stolon
Aetea
Zooecia more or less united to one another, orifice without
chitinous operculum (Cyclostomata[601])
63

3. Zooecia without marginal spines; arising from a branching

axis, which is not formed of zooecia
74
Colony adherent; or erect, fleshy and slightly branched; or
erect, encrusted with earthy matter and repeatedly branched
72
Characters not as above 4

4. Zooecia minute, boat-shaped, united by a delicate tube.

Aperture large, with marginal spines
Beania mirabilis
Colony delicate, erect; zooecia wider above than below 5

5. No marginal spines 6