The economy of British hydroid zoophytes

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TivZ

Elonomy ot BRITISH
HYDROID ZOOPHYTES,/

Jan \e> A 4 \\e James

cMITHSONIAN
OCT 26 1993

LIBRARIES

xiv.

TABLE A.
Order I HYDROIDA.
Sub-order I. ATHECATA.

Polypites naked—without receptacles.

Polypites stalked, branched, and terminal ;
(CORYNIDZS).

Sub-order IT, THECAPHORA.

Polypites provided with horny receptacles.

a Polypites stalked and terminal ;

(CAMPANULARIID/).
6 Polypites disposed on both sides of axis;
(SERTULARIID).
e Polypites disposed of on one side only
of axis ;
(PLUMULARIID).

Pp. J. RB.

Page 6, line 11 from top,

”

”

”

‘CORRIGENDA.

for mesoglea read mesoglea.

. ” ” ” ? ” : 4
is 4 % ; “A Specialised read Specialized.
2 em 14 i $3 ,, sponges read Sponges.
- fe 10 . bottom, ,, coral polyps read Coral polyps.

NOTES ON THE WORK
OF Mr. Fis, RUPFORD. FOR THE
HASTINGS MUSEUM.

In putting these notes together I feel I am perform-
ing an urgent duty, the more pressing since during
Mr. Rufford’s life his innate modesty kept him ever in the
background, so that for many years the part which he
took in promoting the Hastings Museum was scarcely
understood, and his name in Hastings unknown outside a
small circle of fellow workers.

Mr. Rufford’s first connection with the Museum
was through the sale of the effects of the late
Mr. 8. H. Beckles, F.R.S., F.G.S., in April, 1891.

The Museum Committee was then fully organised, and
a grant had been obtained from the Committee for the
purchase of specimens. I was fortunate enough to
accidentally meet Mr. Rufford—not knowing him even by
name—and seeing him interested in the collection which he
was explaining to a boy by his side, I came up and spoke
to him. From this chance acquaintance a friendship
sprang up between myself and Mr. Rufford, which was
fruitful of so much during the last ten years for myself
and the Museum. I shall always look with pleasure upon
those days, at the same time regretting that they are
ended.

To return to the Beckles’ sale. I then mentioned the
position of the Museum Committee as purchaser; and
Mr. Rufford gladly consented to give me his advice as to
what to purchase.

The Beckles’ sale was an important step in the history
of the Museum, because a home had to be found for the
purchases. The Brassey Institute second floor was
granted by the Town Council, and a beginning was
made. In the minutes of April 2nd, 1892, I find a vote
of thanks was passed to Mr. Rufford for his
valuable aid in selecting fossils at the Beckles’ sale, and
again reference is made in the minutes of November 2nd,
1891, to expenses voted for removal of geological remains
from Cliff End, Fairlight, to the ae Institute.
This referred to the Iguanodon foot-print sand-cast now
at the Museum which he presented.

In October, 1891, the Rev. J. W. Tottenham gave his
munificent gift of his private Museum to the Museum,
and‘ after the removal which was carried out by me,
Mr. Rufford threw himself vigorously into the task of
arranging the specimens. Geology and Conchology and
kindred forms of life had his peculiar care. Con-
chology was well represented in the Tottenham collection.

At an early stage of the history of the Museum, the
Bradnam collection of local fossils from the Town Hall
formed part of the original nucleus, to this was added the
Beckles’ fossils, mostly from the Wealden strata. A
strong reinforcement was now to be added to our local
collection by the loan of Mr. Rufford’s private collection,
which being added to from time to time, has given a
marked geological character to the Museum, and caused
it to be respected by geologists and men of science who
visit the town.

At the opening of the Museum in the Brassey
Institute, on Tuesday, August 16th, 1892, I made a few
remarks from the platform in which after mentioning
donors and lenders I said, ‘‘ I must now turn to those who
_have given what is perhaps as valuable as money—that is
time and dearly-bought knowledge. I must in the first
place mention Mr. P. Rufford, our Hastings geologist, a
gentleman well known in the scientific world for his
researches amongst our Wealden flora; this gentleman
has given up nearly his entire time since the month
of May to arranging our specimens, both geological and
otherwise. As I have been intimately associated with
him during the last few months I can say that our
Museum could scarcely have taken shape without his
single-minded enthusiasm for science.”

Mr. Smith Woodward, representing the Geological
Department of the British Museum, referred to the
discoveries of Messrs. Charles Dawson and Philip Rufford
in the Wealden strata, and stated there was evidence that
very soon their work would surpass that of Gideon
Mantell, the great Sussex geologist.

On November 17th, 1893, Mr. Rufford was unani-
mously elected a member of the Museum Committee, from
which time he became one of its most useful and energetic
members, identifying himself thoroughly with its interests,
and sparing neither time or trouble in any work he might
set himself to accomplish.

XVil.

The collection, including a fine series of Mollusca,
was partly the cause that Mr. Rufford directed his atten-
tion to this section of Natural History, and to further
illustrate the section many shells were added by him.
The fishermen took to him specimens which were usually
thrown back into the sea after the day’s dredging, and by
this means a fine collection of Hydroids and Polyzoa
was formed, to which the energies of his later years were
specially directed. At one time an effort was made to
start Aquaria in the Museum. Mr. Rufford did
his share of the work with ability, but the difficulty
of keeping the water fresh marred his efforts in this
direction. It is not necessary to detail the quiet work
which Mr. Rufford carried out during the years between
1893 and 1899, making descriptive labels and displaying
specimens; the Museum tells its own tale. But mention
must be made of the pictorial work of illustration which
he carried out during this period with prints gathered
from the British Museum Catalogue of his Wealden flora
at the British Museum, and other kindred sources. This
was an improvement on the usual methods of labelling,
and one much encouraged by the Committee. A recogni-
tion of Mr. Rufford’s services was made by the Committee
during this period by the gift of a standard work on shells
which Mr. Rufford much appreciated.

In June, 1899, Mr. Rufford finally presented to the
Committee the results of his labours. The letter was
brief which announced the gift, it contained this passage :

‘‘Dear Crake,

Thanks for your note, it may be well to specify
the collections which I have the pleasure to offer to the
Museum as a gift, viz. :

1. The Geological Collection from the Wealden
formation of Hastings and neighbourhood
at the Museum up to the present date.

Local recent Sponges.

Local Hydroids.

Local Echinoderms.

Local Polyzoa.

Local Mollusca.

Land and Freshwater Mollusca.

(Local and from other Districts British.) *

We now approach the last years of this life full of
study and joy in the search of the hidden secrets of the
earth. Since 1899, Mr. Rufford had been engaged in

SES OUP CO 20

XVill.

work for the Victoria History of the Counties of England,
work for a Continental Museum, and in writing and
illustrating the catalogues published in this volume, which
is elsewhere referred to, also he was busy in the illustra-
tion of the Polyzoa and Hydroids to place with his
specimens referred to in this volume. In the work of the
removal of the collection into its new home on the first
floor of the Brassey Institute in 1900, no member of the
Committee worked harder than Mr. Rufford.

In 1901 the idea was mooted in the Committee of
forming a Marine Biological Station in connection with
the Museum, and money was voted for the purchase of a
trawl net. This had Mr. Rufford’s hearty support and
during the winter of 1901, he paid a visit to the
Marine Biological Station at Naples, purchasing specimens
and himself studying on the spot, and visiting the fishing
grounds in the steam trawl of the Institution; this resulted
in an addition to the treasures of the Hastings Museum
of Mediterranean Meduse, etc.

Mr. Rufford heartily aided me with the work of
demonstrations to schools, and one of his last remarks to
me was, that he was very pleased with the boys and girls
of Tower Road Board School who were so interested in
his last lecture on Geology, as he feared that he had dealt
with matters rather above them.

The loss to the Museum in the coming years will be
great, as they will no longer have the willing aid of
Mr. Rufford, one of its best friends, with his large
scientific reading and experience which he was ever ready
to place at the service of all, and whose place it will be
difficult to fill, as such services can only be rendered by
one who has been long in sympathetic touch with the
Institution in all its aims and endeavours.

W. V. CRAKE,
Hon. Sec. of the
Hastings & St. Leonards Museum Association.

ST. LEONARDS-ON-SEA.
1902,

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INTRODUCTION.

The Hydroids are mostly minute creatures
and nearly all marine. The individual animal
is termed a_ Polypite. They rank in
organization just above the Sponges and
below the Sea-anemones and Coral-polyps,
to which they are closely allied.

Their structure is simple. A sack-like
stomach, the only external aperture being the
mouth, around which are arranged tentacles
armed with poisonous darts or stinging cells,
for the capture of prey.

The exterior of the body-wall is composed
of a cellular layer (Ectoderm), some of the
cells of which by extending and withdrawing
lobes, perform the office of muscles, of which
the Polypites proper are destitute. Within
this is a delicate non-cellular membrane
(Mesoglea), the inner lining being composed
of a layer of cells (Endoderm), which have the
property of throwing out pseudopodia and
flagella, by means of which circulation of the
food particles is kept up. Some of these cells
contain pigment and secrete a digestive fluid.

A very few Hydroids (ex. Hydra) lead a
solitary existence. The great majority,
however, by a plant-like process of continuous
budding form colonies, the members of which
are all organically connected, by reason of the

=x. INTRODUCTION.

Buds or newly-formed Polypites not being
detached. The connective parts are tubular,
and allow the circulation of nutriment to all
members of the colony.

In nearly all Hydroid colonies there is an
outer horny cuticle (the Polypary), which in
some cases does not clothe the Polypite itself,
but only the connecting parts (the Coenosarc).
These naked Polypites form the division
ATHECATA. Those in which the cuticle is
prolonged to form protective cups or calycles
constitute the division THECAPHORA.

The forms of the calycles vary, the rim
being either plain, dentate, or with denticles
which meet above the polypite and form a lid,
or operculum.

The arrangement of the calycles on the
stem and branches also differs. They may be
either terminal (ex. Campanularia), biserial—
opposite or alternate—(ex. Sertularia), or uni-
serial (exs. S. Plumularia, and Antennularia).

In one family of THECAPHORA there
are found certain Amaeeboid bodies called
Nematophores, more or less closely associated
with the Polypites, and provided with darts
and cuticular receptacles. The functions of
these bodies are not known. It is possible
that they are to be regarded as modified
Polypites.

The principle of Alternation of Generation
makes its first appearance in the Animal
kingdom in this group. The Polypite is not

INTRODUCTION. XXi.

endowed with the function of sexual
reproduction, but certain members produce
Special Buds which are so endowed, the
bearers of these buds becoming considerably
modified in consequence. They are devoid of
mouth or tentacles, and so do not concern
themselves with procuring food. They are
termed Blastostyles. The Special Reproductory
Buds, in the lower forms of Hydroids, are
liberated as Jelly-fishes, and disperse with
their contained ova, to other parts. In the
higher forms of Hydroids, these Reproductory
Buds remain attached, setting free the ova in
situ. Before liberation, the ovum undergoes
segmentation, and produces, by a process of
inversion, a central cavity—the future
stomach. The embryo is termed a Planule,
and is provided with cilia, by means of which
it swims away, shortly attaches itself to some
object, forms rootlets by the splitting up of
the expanded base of attachment, and also
develops a mouth and tentacles; it then
becomes an ordinary Polypite, which, as
growth proceeds, buds and forms a new
colony.

id ee

¥xiil.

EXPLANATION OF FIG. 1.

Fig. 1. Campanularia flexuosa, Bincks.
(after Hincks.)
A. Natural size of colony.
Bb. A shoot enlarged, bearing
B/ Male reproductive capsules.
B.’ Horny cup or receptacle.
B." Polypite extended.
B." Polypite retracted.
C. <A shoot enlarged, bearing
C./ C./ Female or egg-capsules,

D. Ccenosarc, or connecting tissue which

forms a tube.

XXxiv.

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PART I.

THE ECONOMY

—- OF ——

BRITISH
HYDROID ZOOPHYTES

SS

General Remarks. “Things of beauty
are generally small,” says Aristotle, and the
animals here represented, taken individually,
are generally so minute that they would
probably escape the notice of even the most
enthusiastic searcher, did they only occur
singly. They, however, have the habit in one
of the phases of their existence—the fixed
phase—of living collectively or in colonies,
and in this form they may be discovered
without much difficulty, in rock pools along
the sea-shore ; growing amidst the groves and
spinneys of seaweed, or upon rocks, seaweed,
and shells ; they are also carried about on the
coverings of living shell-fish, upon the backs

2 BRITISH HYDROID ZOOPHYTES.

of Crustacea, and, in fact, upon many other
creatures and objects. The shrimp trawlers’
nets and boats, and the “rubbish” from the
larger trawling vessels afford a rich source
whence many rare forms may be obtained.
Hydroids six-ft. high. If, however,
we were to look abroad in the Pacific, we
should find fixed hydroids of the stature of a
man, that is five or six feet high, but those
which we are likely to meet with here will
generally require the aid of a microscope, in
order that their beauty, structure, habits, and
remarkable life history may be observed.
They are, however, particularly convenient for
the microscopist, since they are of so
transparent a nature that their internal
structure and the operations of digestion and
circulation—not to mention the elaborate and
interesting stinging apparatus, which the
animals use for overcoming their prey—can be
readily observed. Some knowledge of each
step of the animal kingdom is so necessary as
affording the key to other parts, and for a
proper comprehension of the whole, that even
creatures so low in the scale of organization
as the present group, should not be despised.

GENERAL REMARKS. 3

If they require any testimonial to recommend
them to lovers of nature, it will be sufficient
to say that they are very closely allied to the
sea-anemones and the polyps, animals which
form those exquisite structures the Corals.

Hydroids allied to the Sponges
and Corals. In organization, the Hydroid
animals (or polypites) are a step below these ;
but, on the other hand, they have as
neighbours below them, though in a separate
order, the anomalous group of animals, the
Sponges.

‘‘Zoophytes,”’ so named by Linnzus.
The group with which we are dealing, in
common with others of like form and habit,
was styled by Linneus “ Zoophytes,” since,
in consequence of the peculiarity they evince
of forming tree-like growths and of the
resemblance of the animals themselves to
flowers, he regarded them as combining the
natures of both plants and animals.

The cause and nature of these structures
will be found a most interesting study. ‘The
polypites themselves can hardly be excelled
in chaste beauty, being usually of a crystalline
transparency sometimes picked out with

4 BRITISH HYDROID ZOOPHYTES,

Opaque white, or occasionally coloured pink,
red, or orange. In the free phase (Medusa)
the colouring is more vivid. In form the
polypites are particularly elegant, and have
only to be seen in their native element under
the microscope, when their graceful movements
will elicit the utmost admiration. Sometimes
they are suggestive of palm trees with the
crown of leaves fully expanded and gently
swaying in the air; but the illusion is often
quickly dispelled by the sudden closing of
the fancied leaves, and the animal nature is
revealed in the efforts of the polypite to
secure some minute prey. When the food has
passed into the stomach, the polypite expands
again to its original beauty.

Leaving the esthetic side of the subject
for a moment (in which there is ample scope
for gratification), we will turn to the scientific
and get some idea of the grade of organization
which the Hydroids hold, more especially
with reference to their near neighbours in
the animal kingdom.

Structure of Polypite. The Polypites,
in the typical and fixed phase of their
existence, are of very simple structure, and

STRUCTURE OF POLYPITE. 5

the organs may be very briefly enumerated,
riz.: a mouth, a stomach, and arms or tentacles
with which to grasp their prey.

Stinging apparatus. In addition, the
tentacles are provided with stinging darts, by
means of which the animals overcome the
struggles of their victims. These darts are
very intesting structures, consisting of sacks
more or less oval in form, containing long,
coiled-up tubes, which are shot out like
harpoons and penetrate or adhere to the
quarry, which may be a minute worm or
crustacean. These dart sacks are formed
within a single cell.

Darts poisonous. All experiments
and observations concerning these darts tend
to show that poison is infused into the victim.

Body and Stomach sack-like. The
body of the polypite may be compared with a
sack or bag, the tentacles being arranged
around the mouth or body.

Body wall consists of an inner and
outer layer of Cells, with intermediate
Membrane, Non-cellular. The whole
body wall, including tentacles, consists of an
outer layer of cells (“ectoderm”) and an

6 BRITISH HYDROID ZOOPHYTES.

inner layer of cells (“endoderm”). The latter
differ in character from the former, the
endoderm cells being capable of throwing out
and withdrawing lobular processes ( pseudo-
podia ”), and also others which are lash-like
(“flagella”). These two layers of cells are
separated by a fine membrane which differs
from the middle layer (“mesoderm”) of the
sponges, and other groups, in _ being
non-cellular and having no structure. It is
known as the (‘“ mesogloea ”).

No through alimentary Canal.
From the sack-like nature of the body, it will
be noted that there exists no through
alimentary canal such as we find foreshadowed
in the higher Coelenterates, and well-marked
in the Echinoderms. The base of the body of
the animal is used for attachment, or, as a
surface by which to crawl.

No Organs of Sight, etc. This
description applies to the polypite phase only
of the Hydroid career, in which stage sense
organs (organs of sight, etc.) such as are
found in another—the Medusa—phase are
wanting, as also certain “pores” which
subserve excretion.

THE NATURE OF BUDDING. 7

Nerves and Muscles. <As_ regards
nervous and muscular systems. In _ the
tentacles, branching nerve-cells are found at
the base of the ectoderm cells immediately in
connection with a layer of single unstriped
muscle filaments which lie between the
ectoderm and the fine membrane (mesoglcea),
and, according to Parker and Haswell, are a
derivative of the ectoderm, and may be
regarded as a Rudimentary Mesoderm.

Colony=budding. JReference has been
made to the peculiarity which specially
characterises these Hydroids of forming
groups or colonies, sometimes consisting of
millions of creatures all in organic connection.
These creatures, in common with many other
of the lower animals, have the faculty of
multiplying by means of budding, as it is
termed. Budding distinct from ova-
production. This method of reproduction
is entirely distinct from that which takes place
by means of ova, and may, perhaps, be tersely
described as being produced by a simple
inflation of the body wall (budding an
inflation of body wall); such inflation
when perfected by the formation of a mouth

8 BRITISH HYDROID ZOOPHYTES.

and tentacles, constitutes a new individual.
These buds, in some few species (Hydra, etc.)
become detached, and like their parents, lead
a solitary existence (solitary polypites), but,
they also, and far more frequently, remain
permanently connected with the parent
growth, and by continuous budding produce
branching plant-like structures. (colony
budding).

‘‘Coenosarc,” or connective parts,
Between the parent creature and the young
bud there is generally a new piece of stem
formed which serves as a connective part
joining the bud to the main stem. Probably
this arrangement enables the young bud the
better to obtain sufficient space for its develop-
ment, the expansion of the tentacles, and the
procurement of food. These intermediate
portions which connect the polypites, are
termed the Coenosare (meaning common flesh).

Protective resemblance to seaweeds.
So nearly do these growths resemble seaweeds
that the majority of observers mistake them
for seaweeds. This close resemblance may
therefore very probably serve as a protection
against their enemies which might show less

SKELETAL FRAMEWORK. 9

partiality for seaweed than for animal diet,
and so leave them undisturbed. Or it may
have this effect—that the unwary and desir-
able quarry upon which they feed can approach
them without fear or misgiving. However
this may be, the resemblance is so striking,
that even the student may now and then be
deceived.

Budding, common in lower animals.
The process of throwing out buds, which
become detached, is common to other low
forms of life besides the hydroids; but this
peculiar habit of continuous permanent
budding is nowhere else carried to so high a
degree, except in the nearly allied Coral
polyps, * and the somewhat distant Polyzoa.

The Skeleton. The polypary.
Hydroid animals, being of such an extremely
delicate and slender nature, demand, like
most other animals, some support and
protection in the form of a skeleton, either
internal or external. That which the hydroids
have developed, is in the form of an external

*For the purpose of distinction the hydroid animal is
termed a Polypite; the coral animal a Polyp; and the polyzoa
a Polypide. Polyp meaning ‘ many footed,” and the termina-
tions, ide and ite, “like.”

10 BRITISH HYDROID ZOOPHYTES.

tube or casing composed of a horny substance
called Chitine. This casing forms a somewhat
loose jacket, and clothes in the more simple
forms (ATHECATA) only those stem-like and
branching portions (Coenosarc) of the animal
structure. In a few exceptional cases the
skeleton is of carbonate of lime. These little
horny branching growths are frequently cast
upon the beach in tangled masses with
seaweed and other objects. On examination
with a lens, it will be generally found that the
tubes are empty, the animal part having

become decomposed. The entire horny
envelope of the colony is called the
polypary.

Polypite receptacles. In the higher
forms of hydroids (THECAPHORA) this covering
is more fully developed, and is expanded
so as to form receptacles for the polypites
themselves. These receptacles take the
form of chalices or elegant cups, often with
deeply scalloped margins and ringed stems.
Those with the ornamented margins bear a
close resemblance to the delicate little flower,
the Hare-bell, and they have consequently been
named ‘“ Campanulariide.”

‘“ GUARD-POLYPITES.” 11

Receptacle door, or lid. In some
cases, amongst the higher kinds, there are
devices for closing the top of the receptacle by
a lid of various forms, which is forced open
when the animal emerges, and closes when
it retires. The lid is either external or
internal, the latter form being the higher
development.

‘‘Nematophores.’’ Attention should
also be called to certain peculiar bodies called
‘‘Nematophores” or ‘‘Guard-polypites,” which
are found on the stem and branches of the
Plumulariidz, and often closely associated with
the polypites. They have been carefully studied
by Allman, and appear to be a prolongation of
the outer animal layer of the ccenosare, and
show the lobular movements peculiar to the
lowest form of animal life, eiz., the amoeba.
(Amoeba = like character of Nemato-
phore). They may be readily watched in
either Plumularia or Aglaophenia. In the
latter they are found exceptionally distributed
over the egg-case. Their functions are not
fully understood. In some instances dart sacks
are found in connection with them.

12 BRITISH HYDROID ZOOPHYTES.

Formation of Bud. It has already
been mentioned that each member of the
hydroid community is formed by what has
been described as an inflation of the body
wall, which consists only of an inner and an
outer layer of cells, and an intermediate
membrane.

Hollow connection between parent
and bud. This statement implies that
there is a hollow connection between the
alimentary cavity of the parent and the new
bud, which allows the nutriment elaborated
by the parent to be conveyed to the young
bud for its nourishment. (Circulation of
nutriment.) This step is repeated with
every bud formed, so that it will be understood
that a regular circulation of nutriment
previously prepared in the stomach of each
polypite, can be continued throughout the
whole colony, and utilized by those members
requiring it.

Circulation maintained by ‘‘Cilia.”’
Each individual can either pass it at will into
the common channel, or can itself draw upon
it. The current of this material, passing up

CIRCULATION OF NUTRIMENT. 1s

and down the tubes, is maintained by means
of cilia—minute hair-like processes, which by
constant waving keep up the circulation.

Object of circulation. It may be
thought that this common supply is
unnecessary, that each member could find
ample food for itself. But there are the
immature buds; the intermediate parts of the
structure, that is, the ccenosarc, with the
Nematophores; and, more important than all,
certain other individuals yet to be described,
which in consequence of special and important
duties devolving upon them, are in many
cases rendered incapable of obtaining food
for themselves.

A brief description of these members of
the colony will follow.

Dispersal. It will be evident that if
multiplication of the individual be restricted to
the method already described, (viz., permanent
budding), dispersal of the species could not
take place. Nature has therefore allotted to
certain individuals of the colony the duty of
bearing other and special buds possessing the

14 BRITISH HYDROID ZOOPHYTES.

power of reproduction by means of ova.*
This order of reproduction is usually described
as “Alternation of generation,” and this is the
earliest instance of its occurrence in the animal
kingdom. It is very important to realize these
steps, more especially as they are to a great
extent obscured in many instances. We will
therefore enumerate them thus :

The ‘‘Alimentary”’ polypite. 1.—Ali-
mentary Polypite. The ordinary colonial
polypite, whose only duty is to obtain and
assimilate food for itself and the colony, but
which takes no part in the work of propaga-
tion.

The “Select”’ polypite. 14.t—‘‘Select”
Polypite. Certain of the above, which,
for reasons stated in the _ foot-note, we

*It is extremely interesting to note the parallelism with
this order in the plant world. For example: Amongst the
Ferns (Cryptogams), if one of the spores found on the back of a
frond be sown, the result will not be a fern, but a very small
and simple plant—a leafy expansion, ¢:lled a prothalium. It is
the function of this little plant to produce the male and female
elements ; and so soon as fertilization has taken place it dies
leaving the embryos (‘ oo-spheres”) to develop into proper
ferns.
+The ‘*Gonoblast” idea of Huxley and the ‘fertile
polypite” of Hincks. Some apology seems necessary for
presuming to suggest a more suitable term for these Buds, in

ORDER OF REPRODUCTION. 15

have called the “Select” Polypites, generally
more or less modified or atrophied, whose
special function is to produce—not eggs, but
special buds of either sex, which, in the case
of the females, do produce eggs.

‘‘ Reproductive” bud. 2.— Repro-
ductive Buds (“Gonophores ”). Those special
buds, whose main function is the reproduction
of the species by means of ova.

Modification of ‘‘ Select” polypite.
The important duties imposed upon the select
polypites have been instrumental in more or
less considerably modifying their original
character as alimentary polypites ; so much so,
that in many species they have lost the
tentacles, and even the mouth and stomach,
and have become mere stumps. (Loss of
tentacles, mouth and stomach.) Under
these circumstances, it will be seen how very
necessary for its maintenance, and for that of

the face of such authorities as the above-mentioned. These
terms appear, however, to mislead, since both expressions
unquestionably imply the seed or ova-bearing Buds. Now
these are not the seed or ova-bearing Buds, but polypites which
bear those Buds. The term “Select,” or Selected, is free from
this signification, and at the same time distinguishes these
Buds from the ordinary alimentary polypites.

16 BRITISH HYDROID ZOOPHYTES.

its future progeny, is the common supply of
nutriment which it can always obtain, and
which is kept circulating through the connect-
ing channels of the colony.

Division of labour. We have here, in
this ‘‘ select” polypite a primitive example of
the principle of division of labour; all its
energies, in many cases, being devoted to its
special function, just as the various cells of
which our bodies are composed have their
particular characteristics and special duties.

Specialised individuals. Among the
hydroid colony the specialisation of individuals
is not confined to the example previously stated.
There are others whose functions are not
so clearly understood (Hydractinia). Refer-
ence will be made to these in the description
of the specimens.

Sexes of ‘‘reproductive’’ buds. At
certain seasons of the year these ‘select ”
polypites throw out the reproductive buds.
These on a given colony, may be either all
male, or all female, or there may be some
male and some female, but the former
disposition is the more usual.

THE ‘‘ SWIMMING-BELL.” 17

Two methods of dispersal. When
these reproductive buds reach a certain stage
of maturity, it will be found that two different
methods of dispersal are adopted.

‘sReproductive’’ buds either fixed
or free. In one case, the buds (of either sex)
become detached and migrate, fertilization
taking place either before or after. In the
other they remain attached. The ova are set
free on the spot and disperse, and are termed
“ planulze.”

‘¢ Swimming-=bell.’”’ In the first case
(mainly obtaining amongst the lower forms),
the young bud is liberated; it then assumes
the condition known as the swimming-bell,
and propels itself through the water by means
of cilia, to which reference is made hereafter.
The bud itself is essentially a polypite, that is
to say, it possesses a typical mouth and
stomach.

Development of the ‘‘bell.” The
animal in the region of the tentacles becomes
greatly expanded laterally at the expense of
that portion of the body below the tentacles,
and now takes a cup or saucer-like form,
overhanging the free, oval end of the animal.

18 BRITISH HYDROID ZOOPHYTES.

A bell with a clapper will serve as an illustra-
tion. The clapper representing the polypite
with the mouth at the free end, the bell
corresponding exactly to the large expansion of
the base. Sometimes there is a_ thin
membrane partly closing the mouth of the bell
‘alled the * veil.”

Radiating canals. Opening out of the
base of the stomach and traversing the bell
radially, like the ribs of an umbrella, are four
or more channels which extend to the margin
of the bell and unite by running along it ; this
canal system serves to convey nutriment from
the stomach to parts of the bell.

Channels, the homologue of ten-
tacles. It has been conclusively shown that
these radiating canals represent the polypital
tentacles* which are formed after the same
manner as the buds, viz.: by the simple process
of inflation, thus producing long, tubular
processes, which, in most polypites are closed,
but in some are open.

The bud, therefore, in progressing through
the water, strongly contracts the bell, and
thereby expels the water, the re-action

*See Clavatella, Hincks’ Brit. Hyd. Zoophytes, p. 70, ef seq.

SECONDARY BELL-TENTACLES. 19

carrying it backwards.* The bell then
resumes its original form and the process is

repeated.
‘¢ Bell” and mouth tentacles armed
with dart-sacks. To complete the

description of the fully-developed free bud.
Tentacles are sometimes formed around the
mouth and also from the margin of the bell,
from which they hang as long streamers, and
are armed with powerful batteries of
dart-sacks.

Secondary tentacles. In a few species
which have not the bell fully developed and
therefore are not so well fitted for swimming,
the bell-tentacles throw out near their ends
secondary tentacles, and these they use with
which to walk as on stilts. Sometimes suckers
are formed at their ends.

Organs for seeing. At the base of the
bell-tentacles are little granular masses of
pigment, generally of an orange colour, in
some cases a crystalline body is embedded in

* Amongst the Molluscs, the Cuttle fish and others of its
kind, progress after the same manner, and in a backward
direction, by the sudden expulsion of water from a kind of
pouch. Some bivalves, also, propel themselves by suddenly
closing their valves and expelling the water.

20 BRITISH HYDROID ZOOPHYTES.

them. To these organs the power of sight is
attributed. In addition, along the margin
of the bell there occur other organs which
have more the appearance of eyes, very
prominent and staring in character.

Lithocysts, organs of direction.
These are little globular sacks containing a
calcareous spherule, and are termed lithocysts.
It was formerly thought that they might have
been organs of hearing, but they are now
regarded as organs of the sense of direction, in
steering a course.

Nerves and Muscles. With regard to
the nervous and muscular systems entailed by
the development of the “ bell” for swimming
purposes. Romanes, in his work on ‘ Jelly
Fish, Starfish and Sea Urchins,” gives an
interesting account of experiments’ in
dissection which he carried out on the
Hydromedusee, pointing conclusively to a
differentiation of tissue in the direction of
nerves and muscles.

Two rings of nerves. It is now,
however, established, that there exists around
the margin of the bell, two rings of nerves and
nerve cells, one ring occurring just above and
the other just below the “veil” process.

EXCRETORY ORGANS. 21

Primordial centralized nervous
system. These rings afford the earliest
example of a central nervous system in the
animal world.

As to muscles, the whole inner surface of
the bell is lined by fine cancellated muscular
fibres. Muscular tissue also occurs in the
yeu.”

Primordial specialised excretory
organs. In some species of these medusoid
free buds, “pores” occur leading out of the
marginal canal. These pores subserve excre-
tion of whole matter and mark the earliest
occurrence of special organs for this purpose
amongst animals.

In this free bud, which we _ have
cursorily described, with its “bell” and
trailing tentacles, we are introduced to
particular forms of creatures, which are well-
known as Jelly-fish, but whose life-history
is not so fully understood.*

*A distinction must, however, be made between the
naked-eyed and covered-eyed Jelly-fish ; the hydroid offspring
corresponding to the former only, the latter being the offspring
of the higher hydrozoans.

wv BRITISH HYDROID ZOOPHYTES.

Jelly-fish hatches her eggs, then
called ‘‘planulz.” The freed bud, or
jelly-fish, after seeing something of the world
of waters around it, may, in the case of a
female, settle down, attach itself to some
object, and ultimately give birth to a family,
the members of which sooner or later disperse.
They do not, however, in this, the larval
stage, resemble either the polypite or their
immediate parent the jelly-fish, but are little,
flat, conical bodies called planulee, which, in
their later stages, enclose a cavity and swim
by means of cilia.

‘Planula’’ is modified into a
polypite. Later on they attach themselves
to some object by their larger end, which
expands and divides into root-like filaments.
A mouth and tentacles are formed at the
upper end and the result is a polypite, similar
to that which originated the colony. These
polypites will then proceed to carry out the
principle of continuous budding, and thereby
form fresh colonies.

First method of dispersal. ‘*Repro-
ductive ” bud bodily transported. This
is one method by which the ova are

METHODS OF DISPERSAL. 23
transported, viz.: by the mother-bud freeing
herself from the colony and bearing the brood
away to another place, and thus establishing a
new centre of distribution.

Second method, Bud remains
attached, ova set free. The reproductive
buds in this case remain attached to the
parent, fertilization takes place, and the ova
(planulz) are set free, disperse, and go through
the same modifications as the planules of the
detached buds, preparatory to establishing
fresh colonies in the same manner.

‘¢Select’’ and ‘‘ Reproductive ’”’
buds nearly obliterated, The buds styled
“ select” and “reproductive,” demand some
further remarks. In many cases, their real
identity as originating in distinct buds is
almost wholly lost, and it is difficult to recog-
nize any trace of bud unless it is known that
a solution of the difficulty is to be found
in certain links or intermediate forms. We
will, therefore, set those interested in such
matters at once on their guard. Hincks calls
attention to these gradations, and his work on
the Hydroids should be consulted by all who
desire to study this group of animals.

24 BRITISH HYDROID ZOOPHYTES.

Modification and atrophy of ‘‘select”’
bud. Beginning with the “select” polypite,
that is to say, with the individual which
displays the first step in the series of
phenomena connected with reproduction. In
the lower division (ATHECATA or naked
polypites) some members which bear the
gonophores are not modified at all, but in
many species the tentacles become more or
less aborted and the polypite stunted, and
when the higher division (THECAPHORA or
sheathed polypites) is reached the “ select ”
polypites do not obtain nourishment from
external sources, in fact, they have no mouths
and are wholly unrecognizable as polypites.

There is, however, one exception in this
division, amongst our British species, and that
exception is the Genus HALECcIUM, Oken, where
the “select” polypites are fully developed, in
other cases they are represented by a mere
stump (‘Blastostyle”) bearing the reproductive
buds In Tuecapnora, that which repre-
sents the “select” polypite is provided with a
receptacle, fairly strong and modified to the
special circumstances of the case. This modi-
fied receptacle is more in the nature of a case

ATROPHY OF BUDS. 25

or vessel because it contains not only the
aborted “select” polypite, but also the repro-
ductive buds bearing the ova. It is therefore
termed a ‘ capsule.”

Modification and atrophy of
‘sReproductive” bud. In the ATHECATA
and simpler THECAPHORES, the “ reproductive ”
bud is generally a Medusa, but in some species
of ATHECATA the bud remains attached.

In the THECAPHORA, however, we soon
reach a certain point—the turning point—
where, as in such cases as Gonothyrea Lovéni,
Allman, liberation fails to take place and
thenceforward the jixed bud is the distinguish-
ing feature. The bud having now become a
fixture, loses its individuality and the bell and
tentacles dwindle away and the creature wastes
down to a mere sack, which contains in its
walls the ova. The bud, in both ATHECATA
and THECAPHORA, is nearly always enclosed
in a fine envelope, which eventually ruptures.

We have seen, therefore, in the final
stages, the “Select” bud and the “repro-
ductive” bud, both becoming obsolete, and
almost all that remains ostensibly in their
place, are the ova within the Capsule. Under

26 BRITISH HYDROID ZOOPHYTES.

these circumstances, it is difficult to
discriminate between these two buds, and it
will therefore be convenient when this is the
case, to suppress their individuality and to
refer to the whole body as the egg-capsule, or
simply, the capsule. Where, however, the
“select” polypite is recognizable, as in
ATHECATA, and is distinct from the reproductive
bud, the latter with its envelope is also called
the “ gonophore.”

Process of Budding, and Budding
of Medusz. The process of budding, as
previously stated, is by inflations of the body
wall, which is composed of an inner and an
outer layer. This process takes place not
only in the polypites, but also in the Meduse,
in which latter case the buds are Meduse.

Ovaries are situated in body wall,
also in radiating canals of Meduse.
Ova are formed between the outer and inner
layers of the alimentary cavity. This is the
case also in the Medusz, but with some of
these ova also occur in sacks (inflations of the
layers) in the canals which radiate from the
stomach cavity. The important distinction
between budding and ova-production should
therefore be clearly understood.

COMPARISON WITH SPONGES. 27

Larva is a ‘‘planule,’” a polypite,
an Amoeba, or a Medusa. Hincks
states that in nearly all species the ova
develop into planules, and thence become
modified into polypites; but in the fresh water
Hydra and a few other species, the ova at
once take the form of polypites. One instance
he cites in which the egg gives place to an
amoeboid form. There are some cases also,
where the ovum of the hydroid-medusa does
not revert to the fixed hydroid state but is
hatched out a Medusa.

Hydroids compared with the
sponges. Having briefly considered the
nature of the Hydroids, a few remarks
may not be out of place to state in what
respect they differ from their neighbours on
either side.

The group of organisms immediately
below the Hydroids, is the Sponges. They
are a peculiar and somewhat anomalous group
of animals. It is difficult to define their
actual affinities..

Suppose, however, we take a densely-
branching hydroid, such as Hudendrium
rameum, Pallas and deprive the animal colony

\

28 BRITISH HYDROID ZOOPHYTES.

of the horny covering, and also of the
polypites, leaving only the stem and branches,
i.e., a system of tubes or channels (ccenosarc),
with their ends open. Let there now be
developed between the inner and outer layer
(of which the ccenosarc, like the polypite, is
composed) a middle and a much thicker layer
of simple flesh substance called protoplasm,
containing, however, numerous “flesh particles”
or cells. If we then picture the cilia through-
out this canal system maintaining by their
movements a constant circulation, not of partly
assimilated nutriment as in the hydroid colony,
but water containing food which is supplied
it to all parts of the system; and furthermore,
that the larger end of the main channel or
central stem, into which all other channels
eventually lead, be open forming an exit for
the impoverisbed water, we shall gain an
approximate idea of the comparative systems
of a hydroid colony and a simple sponge.

As we go higher in the sponges, instead
of the channel being lined with cilia
and exercising stomachic functions, the cilia
become restricted to little chambers lined by
cells of peculiar form (collar cells), where also

COMPARISON WITH CORAL POLYPS. 29

probably food assimilation is localized, thus
suggesting a comparison with the stomach of
the polypite.

To complete the sponge simile. Instead
of the horny tubing suited to the hydroid
form, there is an intricate network of horny
fibres, serving as a skeleton to support the
otherwise somewhat flaccid body-substance,
and in addition, the structure is rendered more
solid by the secretion of minute needles of
carbonate of lime or flint, in the flesh and fibre.
These needles may be the homologue of the
calcareous skeleton of some hydroids, ¢.¢., a
foreign species of Hyvracrinta and the fossil
Parkeria and others.

Hydroids compared with the Sea-
anemones and the coral polyps. Higher
in the scale of development, above the
Hydrozoa, are placed the Sea-anemones and
the Coral polyps. In the hydroid animals the
stomach is a simple sack, but in the anemones
and coral polyps a slight though important
transformation has taken place. It is first
indicated in the higher medusz. Instead of
the simple sack-like stomach, the mouth
portion is turned inwards. (e.g. A certain kind

30 BRITISH HYDROID ZOOPHYTES.

of non-spilling inkstand). The preparation of
food is carried on by this portion. The
interior is called the body cavity, as distinct
from the stomach.

Arising out of the body wall and directed
towards the pendant stomach or centre of the
animal, are certain little vertical fleshy plates
(Mesenteries). The anemones do not possess
these modifications, but the coral polyps do,
and in addition secrete carbonate of lime, or
horny coral, in the deposition of which the
mesenteries take part, by secreting what are
known as the “ septa,” or rays of the coral.

Hydroids and Polyzoa compared.
There is so much outward similarity between
these hydroid-zoophytes and certain other
creatures of much higher standing, viz., the
Polyzoa and Bryozoa (Molluscoida) that it
may be desirable to point out the wide gap
which separates these two groups.

In appearance, they have much in com-
mon. They are very minute; they have the
habit of permanent budding (thereby produc-
ing plant-like growths); they secrete a poly-
pary-like covering; the animals themselves
are transparent, and have a circle of tentacles

COMPARISON WITH POLYZOA. 31

arranged round the mouth. On examination
under the microscope, however, a considerable
advance upon the structure of the hydroid will
be observed. There isa well-formed through
alimentary canal, entirely cut off from the body
cavity, there is also a gizzard, and well-defined,
though simple, muscular and nervous systems.
The polypary, as it may be called, partakes to
some extent, of a horny substance, but
carbonate of lime also enters largely into the
composition, more especially in the higher
forms. The receptacle also, or cell of the
polyzoan is more a part of the animal than
is the case in the hydroids, where the whole
polypary hangs like a loose-fitting garment
on the compound animal, the former condition
resembling more the relation of the shell to
the shell-fish in the ordinary mollusc, to which
the polyzoan is nearly allied.

32 BRITISH HYDROID ZOOPHYTES.

TABLE C.

Systematic Table to show the position of the
HYDROMEDUSZE (CRASPEDOTA)
(Hyprois, Hyprozoa, and Hyprorpa, of various Authors)
in the classification of the COALENTERATA.

PHYLUM CCELENTERATA.

Radially symmetrical animals with only one cavity in the body
—the gastrovascular space—which serves alike for digestion and
circulation. The generative cells are always either ectodermal
or endodermal.

Sub-phylum I. CNIDARIA.
Coelenterata with thread-cells.

Cuiass I. HYDROMEDUSAS (CRASPEDOTA).
Cnidaria in which the medusa has a velum and the polyp is
without gastral ridges or filaments.

Order 1. HYDRIDA.
Solitary polyps without medusoid buds. Both generative
products are developed in the ectoderm of the polyp.

Order 2. HYDROCORALLIN A.
Colonial Hydromeduse, consisting of a meshwork of coenosarcal
canals, the ectoderm of which secretes a hard calcareous matter,
filing up the spaces of the meshwork. Polyps of two forms,
gastrozooids and dactylozooids. 2 Families.

Order 38. TUBULARLAT (@YMNOBLASTEA).
Without hydothecz and gonangia. Polyps, when more than
one, forming permanent colonies. Generative individuals,
when set free, are Anthomeduse. 4 Sections. 14 Families.

ANTHOMEDUSA. The Medusee of this Order.

Craspedota without otocysts, with ocelli at the base of the
tentacles, and with manubrial gonads; radial canals, usually
4, rarely 6 or 8; budded from polyps of the Tubularia.

4 Families. 18 Sub-families.

TABLE OF CCELENTERATA 33

Order 4. CAMPANULARIAS (CALYPTOBLASTEA).

With hydrothece and gonangia. Colonial. Generative in-
dividuals, when set free, are Leptomeduse.
4 Sections. 7 Families.

LEPTOMEDUS2. The Medusz of this Order,
Craspedota partly with, partly without otocysts; ocelli present
or absent, gonads on radial canals ; budded from polyps of the
Campanularie. 4 Families. 13 Sub-families,

Order 5. TRACHOMEDUS®.

Hydromedusze without hydrosome (polyp stage); with mar-
ginal sense-tentacles in pits or vesicles, with endodermal otoliths.
Ocelli usually absent. Gonads radial. Radial canals, 4, 6, or 8,
often with centri-petal canals. With thread-cell thickening of
ectoderm round the edge of the umbrella.

4 Families. 8 Sub-families.

Order 6. NARCOMEDUSZE.

Craspedota with free auditory tentacles. Tentacles inserted
dorsally on the ex-umbrella, and connected with its edge by
peroniums. Radial canals, when present, in the form of flat,
radial, gastric pouches. 4 Families.

Order 7. SIPHONOPHORA.

Freeswimming polymorphic colonies of Hydromeduse, pro-
duced by budding from an original, probably medusoid,
individual. Gonads in gonophores, which, as a rule, are not
set free.

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