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Ww York State Museum
, FREDERICK. iRee se MERRILL Tereetor
aay
: * Bulletin 71 .
ZOOLOGY 10 an =
FEEDING HABITS AND GROWTH a
OF
JAMES L. .KELLOGG Ph. D, Sat jae
PAGE PAGE
Dye SE A Se ey a 3 Growth under wire netting . ieee
Growth above the bottom....... 23
Soa Scr era eye OS SR EMES OS... yan pete eed
Thee TORN OTe NASON 55 hc ee pean yee a
SP Sep ee nae 18 | Description of figures........... 26
AB es TOF | PEioOunesw a8). . 2s. es bac a2
Peas 22 | Index.................. sees een 28
iqs6TT
ALBANY
UNIVERSITY OF THE STATE OF NEW YORK X
1903 .
Price 10 cents
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University of the State of New York
REGENTS
With years of election i
WILLIAM CROSWELL DOANE D.D. LY. D.
Chancellor, Albany
WHItELAW ReEip M.A. LL.D. Vice hans New York
CHAUNCEY M. DEPEW LL.D. - - - New York
CHARLES BE. Firce LL.B. M-A. l.o. D> Rochester
Wit11am H. Warson M.A. M.D. LL.D. - Utica
Henry KE. TurRNER LL.D. - : = ss Lowville
Si -Crarr- McKEL way MsAy LED. Tipe:
D.C.1,. Brooklyn
DANIEL BEACH Ph.D. LL.D. - - - Watkins
Puiny I. Sexton LL.D. er ee res aliens
T. GuiiForp SurrH M.A. C.E. LL.D. - Buffalo
Lewis A. Stimson B.A. L1,.D. M.D. (- - New York
ALBERT VANDER VEER M.A, Ph.D. M.D. - Albany
CHARLES R. SKINNER M.A. LL.D.
Superintendent of Public Instruction, ex officio
CHESTER S. Lord M.A. LL.D. - - - Brooklyn
THomas A. HenpRicK M.A. LL.D. - - Rochester
BENJAMIN B. ODELL JR LL.D. Governor, ex officio
ROBERT C. PRUYN M.A. - - - - Albany
WILLIAM NoTTINGHAM M.A. Ph.D. LL.D. - Syracuse
FRANK W. HIGGINS Lieutenant Governor, ex officio ei
Joun F. O'BRIEN Secretary of State, ex officio
CHARLES A. GARDINER LL.B. M. A. Ph.D. LL.D. New York
CHARLES S. FRANCIS B.S. - - - - Troy
One vacancy ;
SEHCRETARY
Elected by Regents
1900 JamES RUSSELL. PARSONS JR M.A. LL.D.
DIRECTORS OF DEPARTMENTS
1888 MrLvin DEwEY M.A. LL.D. State Library and Home Education
1890 JAMES RUSSELL, PARSONS JR M.A. LL.D.
Administrative, College and High School Dep’ |
1890 FREDERICK J. H. MERRILL Ph.D. State Museum iat
University of the State of New York
New York State Museum
FREDERICK J. H. Merritt Director
Bulletin 71
ZOOEOGY 10
FEEDING HABITS AND GROWTH
VENUS MERCENARIA |
Introduction
In a previous bulletin of the New York State Museum,! attention
was directed to the fact that both the hard clam, or little-neck, and
the common long-neck clam were rapidly diminishing in numbers,
not only in the waters of New York State, but also along the entire
Atlantic coast where these forms have previously been found. After
a careful examination of a large part of the coast of New England
and Long Island, it appeared that the apprehensions of many mar-
ket men and clammers concerning the growing scarcity of these
forms were well founded. It was not intended that the attitude of
an alarmist should be assumed. Clams still may be had at almost
any hotel or restaurant. Even if the natural beds alone are depended
-on, as heretofore, a certain supply may be had for some time. But
it is certainly true that, unless something is done to check or
modify the indiscriminate and unintelligent methods of taking these
forms now in vogue, the supply is finally to fail more or less
completely everywhere, as it has already failed in many localities.
That time is not remote. It is difficult for one not personally
familiar with the clam flats and beaches, and their histories, to
realize the truth of such a statement. While at any time one may
obtain fresh or canned lobsters in the market, it is difficult to
interest him by the statement that he may not long be able to
1Clam and Scallop Industries of New York State. N. Y. State Mus.
Bul. 43.
4 NEW YORK STATE MUSEUM
indulge his taste for them; yet even now lobsters are danger-
ously near extinction on our coast. But it is the consumer who
should be interested, if possible, because from him, through his
representatives in the Legislature, must come the action which
shall make possible new and intelligent methods of propagation
which may preserve the supply.
Unpleasant facts of this kind, in any case, should be eoneiiee
seriously by the public-spirited citizen; but his interest would be
enlisted, and his support obtained much more readily, if he could
_ be shown some practical way out of the difficulty.
It has been proved, I think beyond question, that, not only are
methods of cultivating the common clam, Mya arenaria, easy and
inexpensive, but the results of the labor involved are astonishingly
great. “Seed” clams may readily be obtained in many localities.
They may, when necessary, be transported from one place to
another without injury. The planting is a simple process. Small
individuals may even be sown broadcast on a soft bottom like so
much grain. Unlike the oyster, the salinity of the water makes
little difference with their growth. Most important of all, their
growth is extremely rapid.
This method of culture, the details of which have been carefully
worked out and tested in artificial beds, was developed after a study
of the life history, the habits, and the conditions of growth. Every-
thing of scientific interest concerning the form has not been
investigated. The early stages of development from the egg, for
example, are not yet known; but enough was known to devise an
entirely satisfactory and practical method of culture, and this
method has been thoroughly tested.
The question may be asked, why, if the demand is increasing
and prices are rising, if the supply has everywhere fallen off, and
if a cheap and eee method of culture has been devised, do
not those who are interested in supplying the market become clam
“farmers,” instead of remaining clam-diggers?
The answer is that ancient laws still leave beaches and flats to
the people. They are public grounds where all have equal rights.
On them any one may dig at any time. No man has a right to
plant and protect his clams, and clam culture is impossible. To
VENUS MERCENARIA 4 5
repeal a law of this character is extremely difficult, for it appeals
to the many as a cession of their rights to the privileged few. But
all would have equal rights to the property by lease or purchase.
Good beaches are very numerous, and there is little danger that
any would be excluded who might desire such property. The sale
and lease of bottoms to oystermen along the shores of Long Island,
- have apparently worked injustice to no person who is desirous of
entering that occupation. At a very few points on the coast, por-
tions of flats have been leased to clammers. These experiments
have failed because of a lack of adequate protection. Unless such
a system, with proper protection, is introduced by the repeal of
old, and the enactment of new laws, soft clam culture will be
impossible, and such laws can be had only when they are desired
by the people at large.
The little-neck clam, Venus mercenaria, grows most abundantly
below the low tide line, where it is taken by means of tongs. Much
of the shallow bottom about Long Island, in which clams were
formerly taken, has been leased to oystermen. The profit from
oyster culture is much greater, acre for acre, than that derived
irom the taking of hard clams, which are left to propagate by the
natural method. ‘The areas left to clammers are now limited, and
the greater part of the supply used in the canning industry comes
from the southern coast. At the same time, clams are rapidly
- diminishing in the available beds.
The little-neck is also found between tide lines. This fact
suggested experiments to determine whether they grow well in
such places. Beaches and flats are not now generally available
by lease. If this were given, these areas could be more easily
protected than those in deeper water, and the matter of planting
and digging would be greatly simplified. It is of the utmost
importance, however, that clams not continually submerged
should increase in size with some degree of rapidity, to insure the
success of culture methods under these conditions. An account
will be given of this growth in Venus.
Very little is known of the growth of lower organisms. Among
the Lamellibranchiata, the group of mollusks to which the clams
belong, much is known concerning the growth of the oyster,
which, for many years, has been artificially reared in Europe and
6 NEW YORK STATE MUSEUM
America. But, till very recently, no observations have been
made on the growth of any clam. In work for the United States
Fish Commission, the results of which have not yet been pub-
lished, Mya was reared in many places, the experiments being
carried out on a large scale. In many ways the results were
astonishing, particularly in regard to the rapidity of growth.
Not only was the actual amount of growth observed, but also
the conditions under which it was least and most rapid, or
altogether impossible. It was my desire to continue the same
line of work with Venus, as nothing was known concerning its
growth or the conditions governing it. Though from lack of
time and facilities, these experiments were not extensive, they
were most encouraging, and show that this form also increases
in size rapidly, even when exposed at low tide.
Feeding habits of Venus. Growth a matter of food
Within wide limits, rapidity of growth in clams seems to depend
directly on the amount of food. In order to make clear the con-
ditions under which rapid growth is possible, the feeding habits of
Venus should be described.
Before such a description is possible, some anatomical features
must be noticed. In a clam bed, the animal lies but a short dis-
tance below the surface of the bottom. Though the shell is entirely
hidden, the creature reaches up to the water above: by means of a
fleshy extension of the body, which has the form of a double
tube. These tubes are known as the siphons, and may quickly be
retracted within the valves of the shell. Ona smooth bottom, the
ends of the siphons may be seen, when the animal is undisturbed,
extending out to the level of the surface. A close inspection will
show that a steady stream of water is entering one tube [fig. 1,
m. S| and leaving the other [er. s]. The margin of the first tube
is crowned by short, tactile tentacles. When touched by foreign
bodies floating in the water, these sense organs cause a closing
of the incurrent siphon, or perhaps a retraction of the entire
structure. The microscopic diatoms, which form the food of
clams, are so small and so evenly diffused in the currents, that
they do not induce these movements.
| VENUS MERCENARIA 7
When the animal is removed from the bed, the tight fitting
valves of the shell are found to be firmly closed. It may be neces-
sary to break the shell in order to insert a knife blade by means of
which the two powerful muscles which connect the valves, and
by their strong contraction close them, may be cut. Removing
one half of the shell, it is seen that both shell valves are lined on
their inner surfaces by thin, fleshy flaps which grow out from the
sides of the body. These are known as the mantle folds [fig. 1, m],
and they inclose a large space, the mantle or branchial chamber,
in which is found the main part of the body. The body, however,
does not entirely fill the mantle chamber, but a large space remains
which is filled with water. The siphons are seen to be simply a
modified portion of the mantle. It is into this space that the
inflowing stream of water, bearing the microscopic food, must
enter. The manner in which the food is collected and passed into
the mouth will be described presently. While the mantle folds
are free at their margins, their edges are closely applied to each
other, and the mantle chamber is essentially a closed space, except-
ing for the siphonal openings.
If now one of these mantle folds be cut away, the body is
exposed from the side and appears as represented in figure I.
The mantle fold on the farther side is shown at m, lining the
entire inner surface of the shell valve, s.
Two large, conspicuous folds, ig and og, the gills, arising from
the side of the body, hang free in the mantle chamber. In this
position, they are continually bathed by the incoming stream of
water, and they perform a very important function in addition to
that of the aeration of the blood —that of food collection. Just
anterior to the gills, and behind the large anterior adductor muscle,
aa, are two small folds, ap and pp, the labial palps. The portion
of the palp seen in the figure, ap, is simply the lateral extension of
a fold which hangs in front of the mouth like a huge lip drawn out
to a point on the sides. The posterior palp is similarly placed
behind the mouth. The mouth opening is on the median line
behind the anterior adductor muscle, and is hidden from view by
the closely applied palps. It is a funnellike entrance to the
digestive tract, and, because the food of the clam is microscopi-
8 NEW YORK STATE MUSEUM
cally small, it is supplied with no special organs such as teeth or
rasping structures.
I would call particular attention to the relation in position
between these palps and the anterior edges of the gills; for I wish
presently to describe the manner in which food is transferred from
gills to palps, and by these into the mouth.
When the gills are removed, there is exposed the main mass of
the body [vm, fig. 2] which is made up chiefly of a large colored
gland, the function of which is the secretion of the digestive fluid,
and the greatly developed sexual glands. This body in anatom-
ical descriptions, is called the visceral ‘mass, to distinguish it
. from the muscular organ which is developed on its under or
ventral surface —the so called foot, f. The last named organ
is represented in the figure as being contracted within the
mantle chamber. It is capable of great distension and, in a
large clam, may be projected for a distance of two or three
inches from the edges of the shell. Though a fleshy structure,
it is, when protruded, quite tough and firm, being made rigid by a
large quantity of blood which is pumped into it by the heart, in
order to cause its distension. The foot is an organ of locomotion,
and is also used in burrowing. It is possible for Venus to creep
about by means of its thrusting and wormlike movements; but I
believe that the animal uses it in this way much less than is
generally supposed, and this is a point of much interest to the
clam culturist.
In order to understand the mechanism by means of which food
is collected, it is necessary to describe in more detail the structure
of gills and palps. The gills are the most complicated organs
in the bodies of lamellibranchs, and must be described here as
briefly and as simply as possible, without mentioning their won-
derful histological structure. Outer and inner gills are practically
the same. Suppose that one of these is carefully removed from
its line of attachment to the body, and studied by means of the
microscope from the surface and in section: such an examina-
tion shows the gill to be not a solid flap or fold, but an
exquisitely minute basketlike structure with an outer and
inner wall inclosing a space between. These walls are made
VENUS MERCENARIA ; ie)
of extremely fine rods placed side by side, as represented
in the most diagrammatic way possible in figure 3. In order that
these rods, r, may retain their position, they are in many forms,
irregularly fused with each other by secondary lateral growths of
tissue, ic. The outer and inner walls of the gill are also held
together by partitions which extend across the inner space between
‘them, p. The gill is thus seen to be basketlike, the walls being
made of rods between which are spaces, s, which put the interior
chamber in communication with the mantle space in which the
gills hang.
These rods, or filaments, of which the gill is made, contain an
interior space in which the blood flows. They were probably |
primarily developed in order that the blood of the body might be
brought in close contact with the water, that, by diffusion, the
carbon dioxid of the blood might pass outward through the thin
walls, while, by the same process, oxygen, carried by the water,
might pass into the blood. But, in addition to performing the
function of breathing, the gills have taken on that of collecting
minute organisms used as food. This is accomplished by a com-
plicated process.
We have seen that a constant stream of water entered the mantle
or branchial chamber. What becomes of it? And what is it that
causes the current? All of this water inthe mantle chamber streams
through the minute openings between the filaments of the gill and
enters its interior space. It now rises to the base of the gill, and
flows into a tube, the epibranchial chamber [fig. 1, ec], through
which it passes backward, leaving the body by the upper or exhalent
siphon, which is directly continuous with the epibranchial chambers
of the four gills. The currents which we first noticed, then, enter
the mantle chamber by the lower siphon, pass into the interiors of
the four gills, flow to their upper or attached edges, and are directed
backward and out through the upper siphon tubes of the mantle.
The cause of these rapid currents is revealed by a microscopic
examination of the rods or filaments of the gills. These are found
to be covered on their outer surfaces, which face the water on
both sides of the gill, with innumerable short, hairlike structures
which project perpendicularly from the surface. These cilia
IO NEW YORK STATE MUSEUM
are protrusions of the living protoplasm of the cells which form
the walls of the filaments. Each possesses the power of movement,
lashing in a definite direction, and recovering the original per-
pendicular position more slowly. This movement is so rapid that
it can not be seen till nearly stopped by inducing the gradual
death of the protoplasm. It is very effective in causing strong
currents in the surrounding water. ;
A microscopic examination, and direct experiment with minute,
floating particles, will show that other cilia are present on the
filaments than those which cause the water to enter the gills. The
diagrammatic figure of the gill [fig. 3] does not show why the
minute food particles may not be taken into the interior of the
gill by the entering stream of water, and finally out of the body
through the broad water channels. This is prevented by long
cilia arranged in bands which project out laterally between con-
tiguous filaments in such a way as to strain the water which enters
the gill, thus preventing all floating matter from entering. These
highly specialized cilia tracts of lamellibranch gills, I have called
the “straining lines.“t In some forms there is a single line, in
others there are two. In some cases the lines are formed by a
single row of cells; or a section across the line sometimes reveals
several closely crowded cells bearing the greatly elongated strain-
ing cilia.
That foreign matter is really excluded as the current of water
enters the gill, may be demonstrated by direct experiment on a
living gill. Carmine may be ground into a fine powder, and
suspended in water without becoming dissolved. If a small
amount of this is allowed to fall on the surface of a living gill, it
will be seen to lodge there. A wonderful thing now occurs. A
myriad of separate minute grains, which may represent the food
of the clam, are almost instantly cemented together by a sticky
mucus which is secreted by many special gland cells in the fila-
ments, and the whole mass, impelled by the oscillations of the
cilia, begins to move with some velocity toward the lower
or free edge of the gill. On this free margin is a groove
into which the material collected on the faces of the gill is turned.
1 Kellogg, J. L. Contribution to Our Knowledge of Morphology of Lamelli-
branchiate Mollusks. U.S. Fish Com. Bul. 1892.
VENUS MERCENARIA Il
This groove is also lined by ciliated cells, and the whole mass is
swept swiftly forward in it toward the palps. The natural food
of the clam, of course, is carried forward in the same way. It is
evident that a large proportion of the organisms floating in the
water which enters the mantle chamber must come in contact with
the sides of the gills, and be carried forward to the mouth folds,
to which they may be transferred.
These points may be made more clear by referring to the diagram
[fig. 4]. It represents a section made transversely across the fila-
ments of a typical lamellibranch gill. In a single gill there are
thousands of these rods. But five are shown here on each side,
standing in row to form the perforated walls of the gill. Each rod
is represented as being more or less oval, when its cut end is
viewed in this way. In three places are shown the lateral union
of filaments. The reference letters ig are supposed to be placed -
in the interior space of the gill, and » shows the nature of the
partition, or septum, which, at more or less regular intervals,
stretches across this space and holds the two walls of the gill
together.
The details of cellular structure have been drawn in two
filaments. The long, straining cilia, which stretch across the spaces
between rods, are shown at sc, and the arrow indicates the course
taken by the water current as it enters the interior of the gill. The
cilia which cause this entering current are the frontal cilia, fe.
Opening on the surface between them and the straining cilia are
the gland cells, gc, the secretion from which cements together the
food particles.
This figure is not intended to represent the details of structure
found in the gill of Venus, which is much more complicated in
‘aany ways. The general plan of structure and of function in that
form, however, is very much as represented, and this diagram is
used because it may be so much more easily described. :
If we now examine the palps with a hand lens, we may notice
that their inner surfaces—those nearest to the mouth — are
covered by a set of very fine parallel ridges. The lateral portions
of the palps are shown in figure 2, ap and pp. They are capable of
many movements. They may be bent and spirally twisted,
i2 NEW YORK STATE MUSEUM
lengthened or shortened, and, if their inner faces touch the edges
of the gills, any material which is being brought to this region is
transferred onto the ridges of the palp. This is accomplished by
strong cilia which are developed on the ridges. These same cilia
carry the foreign matter on across the ridges, and finally force it
into the mouth [arrow on pp].
_ This, in brief, is the method by which clams and oysters wid
other lamellibranchs collect and ingest their food. The process,
till very recently, has not been closely studied, but this auto-
matic feeding process has been known in a general way for a
long time. It has sometimes been said that, if a lamellibranch
is to prevent suspended mud from being collected by the
gills, it must close its shell, thus entirely preventing all ingress
of water into the body. It has been found that these creatures
have no more control over the activities of the cilia which have
been described than a man has over the cilia in his trachea. As
long as the animals live, the cilia continue to lash in the same
definite directions, though their activities soon become lessened
after the shell is removed.
But I have found that the animal can prevent food or particles
of dirt from being taken to the mouth while the stream of water
is yet flowing. It seems never to have been suspected that
complicated mechanisms existed, by means of which collected
particles could at once be discharged from the body. They are
present, however, probably in all lamellibranchs, differing some-
what in different forms, and I shall describe the comparatively
simple one which is found in Venus. ©
If the mantle and gills are removed from one side of the body,
so as to expose the visceral mass and the foot, and the creature is
put into a dish of sea water, grains of carmine, which are allowed
to settle on the surface of the visceral mass, at once indicate the
presence of a ciliation there, as well as on palps and gills. These
experiments require care and patience, but they show with great
certainty that the most definite cilia currents exist in this region.
These are indicated by the arrows placed on the visceral mass in
figure 2. It will be seen that all the currents converge at a definite
point, x, just above the line of the base of the muscular foot on the
VENUS MERCENARIA ibe
posterior margin of the visceral mass. Any material, then, which
touches this surface, instead of being taken toward the mouth,
tends to be forced in the opposite direction. Immediately on
touching the wall of the visceral mass, the fine particles are
‘cemented together by an abundant mucus, as on the gills.’ When
much carmine or mud is used, a large ball of it is collected at x.
It will be noticed that this region lies directly in the path of the
incoming stream of water from the branchial or lower siphon; and
at first sight it would seem that from this position there could be
no means by which it could escape from the mantle chamber.
Clams undisturbed in the bottom, however, from time to time
may be seen to discharge a strong jet of water from both siphons.
This habit of many lamellibranchs is better shown in Mya. When
these clams are kept in a bucket of water over night, the floor will
be wet for many feet around it in the morning, and indeed one
may at any time when they are so kept, see them violently close
the shell by contracting the adductor muscles, thus emptying the
mantle chamber by throwing a strong jet out of both siphons.
This peculiar habit of all lamellibranchs which have been observed
is, without doubt, for the purpose of removing masses of material
which the animal can not use as food.
This is not the only means of discharging undesirable material
from the mantle chamber. If the entire body be removed, leaving
only the mantle lining the sheil on one side, it also will be found
to be ciliated. In this case, as illustrated in figure 5, everything
is swept downward toward the free edge of the mantle, and falls
into a line parallel with the edge, and is then directed backward.
Particles which may fall on the extreme edge are also passed into
this well marked stream. Everything is directed backward, but
_can not be carried out of the incurrent siphon against the stream
which is entering through it. Ina little bay beneath the base of
the siphon, where it is out of the current, the material is collected.
By the contraction of the adductor muscles, and the resulting
emptying of the mantle chamber, as described above, this col-
lected mass is expelled. !
But, in spite of the activities of these two surfaces, which tend.
to rid the body of material not fit for food, it is evident that, if
14 NEW YORK STATE MUSEUM
much mud is entering, large quantities of it must be collected on
the gills and be sent forward toward the mouth. I have spoken
of the fact that the palps are capable of extended movements. Ii
they are withdrawn so as not to touch the gills, material will
accumulate in the anterior parts of the gill grooves till masses are
formed so large that they fall off into the space of the mantle
chamber below — perhaps to be taken up by the currents on the
mantle. At any rate, they would be discharged when the mantle
space was emptied. I have no doubt, especially after what I have
observed in forms like Yoldia, that the palps of Venus are from
time to time withdrawn from contact with the gills, in order
that they may receive no material from them.
It is when we come to examine the palps that we find the most
complex arrangement for keeping material from entering the
mouth when that is desirable. A close examination of the inner
faces of the palp shows a narrow strip around its margin which
is without the ridges previously described. Both of these margins
are very densely ciliated. When suspended material falls on the
upper margin, it is carried up onto the surface of the ridges [fig.
2, um] and across them to the mouth. Anything which touches
the other margin, on the other hand, is swept with great rapidity
in the other direction— out to the end of the palp, where it
accumulates and is finally thrown off into the mantle chamber
below. It is true that this margin is narrow, and not much
material suspended in the water would strike it; but probably
when a large quantity is collected on other parts of the palp,
this edge is folded over so as to touch these heavily laden sur-
faces, and sweeps them clean.
It thus appears that there are extensive ciliary tracts for collect-
ing and conveying food to the mouth; but that, in addition to
these, there are other ciliated surfaces by means of which unde-
sirable material may be excluded without the necessity of closing
the shell. Because of the advantage of sustaining the
aeration of the blood, this must be of very great service when the
water is muddy.
In this description of the feeding habits of Venus many
important details have been omitted, particularly in regard to the
VENUS MERCENARIA 15
anatomy of the gill, which is much more complicated than is
indicated in the figures.
The question of food is an important one when we are searching
for means of rearing this clam by some culture method. In order
to force the growth of oysters in French claires, water is held in
reservoirs back of the beds till the contained diatoms may have
multiplied greatly, and is then allowed to run over the beds. Such
methods are expensive, and under proper natural conditions, Venus
will grow very much faster than either the European or American
oyster. Enough has been said of the food of Venus to make it
clear that, if it were raised on beaches or flats, we should not
expect to find so rapid a growth as if it were never exposed, for
feeding is impossible without water currents. I hope to show,
however, that growth seems to be very rapid even under these
circumstances.
Growth experiments
Before speaking of these experiments, it will be well to make
it clear that the planting was done on a small scale, and was
pursued under the most adverse circumstances. I believe that the
results as we have them are perfectly certain — and they are most
satisfactory as they are; but I am also sure that under favorable
conditions growth would have been very much greater.
A trip was made to Riverhead, and the shore examined carefully
as far as Greenport. Many clams are found along this shore,
and several sites were located, which, so far as currents and char-
acter of bottom were concerned, seemed to be ideal. In every
case, however, | was assured that clams would not.be allowed to
remain unmolested for a week. So certain did this seem, that the
very much less favorable harbor at Cold Spring, on the sound,
was selected. Here also it appeared that no portion of any of the
beaches would be free from molestation by clam-diggers. The
only thing to be done was to ask the privilege of a small space on
an oyster bed which extended close to the low water mark. This
was granted by Captain Jones, who has my sincere thanks for
this favor, and also for the kindly interest which he showed in the
work.
The rights of the oystermen seem to be strictly respected. I
ventured to run some of my beds up on the narrow beach nearly
16 NEW YORK STATE MUSEUM
to the high tide line, marking them by labeled wires which were
run down out of sight. These I easily found in the winter, but some
of the beds had been raked clean. Others certainly escaped ob-
servation. Before planting, the ground was raked, that I might
be assured that no little-neck clams were presentinit. Jam very
positive that the beds and sealed wire cages on the oyster ground
had not been touched when they were examined after an interval
of six months.
But the unfavorable conditions were these. Everywhere above
and below these beds, oysters covered the bottoms as close as they
could lie. They take from the water the same floating organisms
which Venus uses for its food. Everywhere, too, above and below
low tide line, soft clams were burrowed almost as close as they
could be placed. They also use the same food. Now, we have
experimental evidence to show that the growth of all these forms
is, up to a certain point, directly proportionate to the amount of
food. They all grew here; for, on account of the conditions of the
upper harbor, where at high tide the shallow water, fed by fresh-
water streams, was warmed for hours by the sun, diatoms must
have multiplied with great rapidity, and, when carried out,
offered abundant food. But undoubtedly none of these lamelli-
branchs grew as they would if the life of the bottom had not been
so abundant.
As an example of the number of these organisms on the bot-
tom, this case may be cited. A flowerpot, 4 inches across the top,
filled with clean sand, was sunk nearly to the level of the ground
on June 19, 1901. In it was placed a little-neck clam. When
_ examined Dec. 28 of the same year—six months afterward—the
sand in this pot contained 11 soft-shelled clams ranging from half
to three quarters of an inch in length, besides the hard clam, which
had increased considerably in size. These soft clams had settled
in the pot from the swimming larval condition, as they settled
elsewhere on the bottom, and had begun to grow. It is most
reasonable to suppose that, if this hard clam had been growing on
almost any beach where less life was being supported, its growth
would have been more rapid, for diatoms are more or less abund-
ant all along the shore.
os
|
VENUS MERCENARIA M7
Another serious hindrance to the growth of clams is the presence
of the seaweeds, Ulva (sea lettuce) and Enteromorpha which, .
during the greater part of the year, grow profusely after their
attachment to large pebbles or other solid bodies on the bottom.
‘Not only the larger stones on these beds, but, especially, the wire
cages which were sunk into the bottom, were in December
more or less completely covered by them. In extended experi-
ments on the growth of the soft clam, Mya, the same difficulty
was met with in many localities. The masses of weed, flattened
out on the bottom by the tide currents, greatly hinder the clams
underneath from obtaining from the water their needed food.
My experiments with both forms show that this condition is
detrimental to the best results. If one were free to select
sandy ground which would afford no means of attachment, this
difficulty would not appear.
These matters are spoken of in detail because the results which
will be given should, without doubt, have been far greater. Any
one with rights to certain parts of a beach, who could watch his
beds at all times of the year, could, with very little labor, prevent
. these drawbacks. c
Still another difficulty attending the work at Cold Spring was
the fact that it was almost impossible to obtain clams small enough
for planting. None were to be had in this locality. A number
were sent from Jamesport, L. J., but most of them were of
marketable size, and hence too large for the most important part
of the experiment. The smaller ones came from New Bedford
Mass., and these had perhaps previously been received from Edgar-
town. It must however be said that the hard clam, like the oyster
and quite unlike the soft clam, Mya, will live for many days, and
even for weeks, after being removed from the water during the
hot summer time, without apparent injury. The soft clam may be
preserved in this way for a long time during the winter, and very
small individuals may safely stand much exposure in hot weather;
but the larger forms of this species succumb after a short time.
The tenacity of life in‘the small Venus may also be greater than
in the adult, but nothing is known in regard to it.
18 NEW YORK:STATE MUSEUM
Methods
Each clam was measured in sixteenths of an inch at the time of
planting, and also when taken from the bottom six months after-
ward. Merely to state the increase in length, however, gives no
adequate idea of the actual growth. It is much better to give the
increase in volume. To state that a clam increases from
1,2, to 14% inches in a certain time gives little idea of its actual
growth. If individuals of the two sizes are held in the hand and
compared by the eye, the bulk of one is seen to be much greater
than that of the other. It is really this increase in volume which
we wish to determine, so each clam was measured also by deter-
mining its displacement in water. A table was made showing the
displacement of clams of various sizes. For example, many indi-
viduals just 1 inch in length were measured in a graduated vessel.
There is some slight variation, because some are thicker than
others. The average of many measurements, however, show that
a clam of this length displaces 2.5 c.cm. ‘The average displace-
ment of other sizes was determined in the same way.
To illustrate the difference in the two ways of stating the increase,
we may compare clams I and 2 inches in length. One is 100%
longer than the other. One has a volume of 2.5 c.cm, the other a
volume of 22 c.cm; and, while a clam 1 inch long has increased in
length 100%, it has increased in bulk or volume 780%. This increase
‘in size or volume is what we wish to determine.
Suppose that in a certain bed are placed clams all of a size.
When these are dug, after a lapse of several months, some indi-
viduals will have increased in size more than others, though the
differences may not be great. In order to determine the increase
in such a bed, the arithmetical mean length of the whole series
has been calculated, and the volume of the mean has been com-
pared with the volume of the clams when planted.
In one bed, for example, several clams 1 53, inches in length were
planted. In six months they were removed, and the length of each
individual carefully measured. There was some individual variation
in the length; so the mean length of the series was calculated. It
was found to be 14? inches. The average volume of clams 1,3,
inches long is 4.5 c.cm; that of individuals 143% inches long is
VENUS MERCENARIA 19
14.5 c.cm, or 3.22 times as great. The increase in volume in the
six months, therefore, was 2224.
Growth between tide lines
The most important point brought out in this experiment is the
fact that growth is considerable on bottoms exposed for several
hours at low tide. This is shown in the following cases. .
A line of flowerpots was run from below ordinary low water
mark up the steeply sloping beach to a point about two feet below
the ordinary high water line, the fall of the tide being about six
feet. The pots were sunk so that their tops were level with the
ground, and were separated by a space of about two feet. June 19,
1901, there was placed in each of these pots a clam 1.25 inches, or —
to give the measurements for convenience’s sake in sixteenths of
one inch — 1,4, inches in length. These were examined, after an
interval of six months, on Dec. 28. Some of the pots were empty
er contained dead shells. In the first or highest, the clam had
grown to a length of 144 inches, an increase of 148% in volume in
the half year. If we had no other example of growth, this would
be very suggestive, for the increase is great, the creature having
become in this short period almost two and a half times as large
as when planted.
We should expect to find still greater growth with longer
immersion.. In the second pot, the clam had increased 154%, and
in the third, still lower down, 172%.
The fourth pot was empty. In the fifth, the increase, instead of
being greater still, was only 87%. The explanation of this seems to
be perfectly clear, and is exemplified in several other cases. Around
‘the margin of this pot there had grown a large quantity of Ulva.
There was much of it at this level of the beach, while higher up
it was not abundant. Without doubt this seaweed was flattened
out over the top of the pot by the current, in such a way as to
prevent free access to the food-bearing stream, and for this reason
growth was not so rapid.
The presence of these weeds, which grow on so many bottoms,
should not seriously inconvenience the clam culturist. They may
be removed without difficulty with a rake, and do not grow
abundantly on a surface which is reasonably smooth. If it had
20 NEW YORK STATE MUSEUM
been possible to visit these beds a few times during the summer,
the results in the case of many lower beds would undoubtedly
have been different.
In pots still lower down, all of which were covered with Ulva,
the growth was much the same as in the fifth — from 80% to 100%
increase, : :
In this line of pots, then, the fact is demonstrated that between
tide lines, hard clams 1.25 inches long may increase 2.5 times or
more in volume in half a year. Localities more favorable for their
growth could easily be found. If experiments were made on a
large scale, I should expect to get a more rapid average growth
even where the forms were exposed at low tide, and a much greater
increase on bottoms which are never exposed. As it is, this growth
as compared with that of the oyster is marvelously rapid, just as
it is in the soft clam.
It should be noticed that we are not attempting to make extended
generalizations on the data given by four or five individual clams.
Two clams side by side will not increase at the same rate. It is
possible that one might grow twice as fast as another. But, if we -
had a single case in which we were certain of the amount of increase,
it would assuredly indicate the possibilities of growth, and the
chances are that it would not by any means be the limit of-
possibility.
On the other hand, when we compare the growth in pots 1, 2 and
3. and find a progressive increase from the higher to the lower pot
—an increase of 145%, 154% and 172%— our induction is founded
on insufficient data, and really means nothing. The result is as we
should expect it, but it may be entirely accidental. But it is sug-
gestive, and, if it were possible to observe many rows of clams
similarly placed, we might .reasonably expect to establish it.
Unfortunately it has not been possible to do this.
The simple case of the line of flowerpots has been spoken of
first because it was more or less typical of the results obtained in
many small beds planted under similar conditions. Many hundreds
of clams, after being carefully measured, were segregated into
groups according to length and planted together. Their growth
substantiates the results obtained in the flowerpots.
VENUS MERCENARIA 21
Very briefly the following results will be described. Several
small beds, each with an area of 16 square feet, were laid out on
the gravel between tide lines: A group of these was separated by
an interval of 20 or 30 yards from another group. Most of these
small plots were within the boundaries of the oyster bed already
mentioned, but some were above the line of the bed, and a few
of them were dug clean. Others were not discovered by clam
diggers, and apparently entirely escaped molestation.
In each of these small beds, clams all of a size were planted.
The number on a bed varied from 100 to 175. I would call particu-
lar attention to the fact that on the deeper beds, where the tide
currents were swiftest, larger stones were exposed, and there was
here an abundant growth of seaweed, which was not found farther
up on the beach. This always interfered seriously with the growth
of the clams.
For example, on these beds which were below the ordinary low
_ tide line, where we should expect to find the most rapid growth,
there was an increase in volume in clams 112 inches long, of 35%;
in those 144, inches long, of 41%; and in those 14% inches long of
42%. 1am allthe more certain that this low rate of growth is to be
explained by the presence of the seaweed, because I had previously
had the same experience in a much larger experiment in the soft
clam. F ortunately, as I have already stated, a little labor by one
who is able to be on the spot during the entire year would prevent
this result.
Some of the higher beds, however, which from the character
of the bottom were free from the weed, gave different results, and
show the possibilities of growth much better. On a bed only three
or four feet from ordinary high water line, there was placed on
July 6, 130 clams, 134 inches long. On Dec. 30, almost the entire
number was removed. Some had increased more than others.
The mean of the series was calculated, and showed an increase
of 255% in volume in a little less than six months.
On another bed, somewhat lower, 150 clams 1,8 inches long had
increased 157% in volume. One of the things to be expected is that
clams of smaller size would show a relatively greater growth. It
has not been possible to make comparisons to demonstrate this
22 NEW YORK STATE MUSEUM
because of the influence of the seaweed on so many beds. The
variation in the size of planted clams in this experiment was from
1°; inches to 14% inches in length, and this is not a very great
range.
On a third bed, also situated well up on the beach, clams 18
inches long when planted had increased 155% in volume in the six
months. Whether the amount of food in the summer is greater
than in the winter, I do not know. I have no doubt that the
increase goes on during the winter months, though, it may be,
with diminished rapidity. It would be extremely interesting to
carry out these experiments on a large scale through the entire
year. These facts certainly show that the possibilities of growth
in Venus are very great, and indicate that its artificial culture
between tide lines would be easy and inexpensive, and that it would
yield large results. Considering the place which the little-neck
has in the markets, it would seem that the artificial culture of the
form should yield a larger income than does the culture of the
oyster as carried on in Long Island sound. The latter is expensive
and laborious, and growth is very much slower than in the case of
either of the clams.
Wandering habits of Venus
The soft or long-neck clam, Mya, is capable of locomotion only
when very small. As the body increases in size, the foot, or loco-
motor organ, becomes relatively smaller. An individual 2 inches
long, while it can not move along the stirface of the bottom, is
still able to use the foot as a burrowing organ. When it has
attained a length of 3 or more inches, however, it seems to be
incapable even of covering itself in the bottom.
In the case of the hard clam, Venus, on the contrary, the foot
remains throughout life a very well developed locomotor organ.
Though no definite experiments have been made to demonstrate
what it is able to do, one might assume, from the size of the
organ and its power of extension as demonstrated in aquaria,
that the animal is able at all times in its life, not only to burrow
but also to move from one locality to another, as the fresh-water
clams, with a similar foot, are known to do.
VENUS MERCENARIA 23
The beds in this experiment were planted with the fear that the
clams would wander. The result, however, showed conclusively
that they do not have this habit — or that they did not exhibit it
in this particular case. The clams were found where they were
placed within the limits of the original beds. Careful digging
around the margins of the beds failed in every instance to show
any wandering tendencies.
Growth under wire netting
In order to be perfectly certain that clams should have no means
of escape, three cages of wire netting were constructed, bounding
the margins of the area containing clams in each case to a depth
of 5 inches and covering the top. These forms never burrow to
a greater depth than this, and there was no possibility of escape.
In each case the netting remained intact, and certainly was not
disturbed. These beds were exposed only during the full moon
tides. Here also the seaweed seemed to play an important part
in the results. In one case the netting was sunk so deep as to be
covered with sand, and consequently no seaweed attached, as it
did on the other cages. Growth was much more rapid here,
though the clams in this bed were smaller when planted, and, asa
consequence, a more rapid growth should have been expected.
The results were as follows:
Cage 1 Clams planted July 6, 1,5 inches long. Some seaweed
was attached to the wire of the cage. The clams were removed
Dec. 30. The increase in volume was 145%.
Cage 2 Planted July 6, 1;% inches in length. Removed Dec. 30.
A very large quantity of weed over the cage. Increase in vol-
ume, 78%.
Cage 3 Planted July 6, 14 inches long. This cage was sunk
so deep that no weed was attached on the surface. The increase
Dec. 30 was 222% in volume in the six months.
Growth above the bottom
In methods of oyster culture as developed in France, the forms
are placed in racks above the bottom, and from the tide which
sweeps over them, they are enabled to obtain nourishment enough
for comparatively rapid growth. It would be an interesting
Go Nelpretateneon eh gee pet
24 NEW YORK STATE MUSEUM
thing to show that clams could be made to erow in this way.
The clam culturist could then make himself independent of
beach rights, and perhaps more easily obtain a lease of ground
for such a purpose below low water mark. .
But one or two very small experiments on the soft clam have
indicated that the creatures do not do well under these conditions.
At Cold Spring a wire rack was constructed, and anchored above
the bottom in a swift current. Into it were put several hard clams
ranging from 1,4, to 24? inches in length. Every one of these
seemed to be in a healthy condition at the end of six months, but
not one had increased a particle in size. Not being able to cover
the body in sand, they seem to have remained most of the time
with valves closed. They may possibly have moved about at °
times, for their shells were worn, but more likely this was due to
the fact that they were rolled about in the cage by the currents.
On their smooth, clean surfaces numbers of Anomias, or silver
shells, had attached and grown, as shown in figure 6. ;
Though this small attempt to induce growth above the bottom
ended in failure, it should, on account of its importance, be
repeated on a large scale under as many different conditions as
possible, in the hope that some combination of circumstances
might prove to be the right one.
Enemies
Neither of the clams is molested by the starfish after it has
become large enough to burrow, though the very small soft clam,
‘and perhaps the hard clam also, is destroyed in great numbers by _
small starfish, before itis able to cover itself. So far as I have
been able to discover, there is but one natural enemy of Venus
which might possibly be destructive. It is the gastropod mollusk,
Lunatia [fig. 7], which is abundant in some localities. It is found
in numbers at Cold Spring. On several occasions I have observed
it digging below the surface and attacking both hard and soft clams
in their burrows. By long continued labor, it files a smooth, clean
hole through the shell of its victim by means of a rasping organ in
its mouth cavity, and then destroys the soft parts of the body
within. Figure 8 illustrates the character of the borings on shells
VENUS MERCENARIA 25
taken from the beds at Cold Spring. In every case the perforation
is near the prominence of the shell called the umbo, directly over
the pulpy visceral mass, which might most easily be sucked up
through the opening. It is a curious fact that this region of
the shell is selected by Lunatia for boring in any lamellibranch
which it attacks. It may not invariably be so, but I have many
shells of different species which have been drilled in this region,
and have happened to notice no exceptions to it.
No matter how numerous it might be, this enemy would prob-
ably not be as troublesome to clam culture as the starfish is to the
oyster industry. In several | places I have seen it collected by
fishermen for bait, simply by pegging a bit of fish, or even a dead
starfish on the bottom. In a short time numbers of them will be
found collected on the bait. By some such simple means, if it
were desirable, a clam bed probably could easily be rid of the
creatures.
Conclusion
- This experiment on the growth of Venus from lack of means
and time and favorable locality has been a limited one. In order
fully to demonstrate the feasibility of the artificial culture of the
form, it should be carried out on a very much larger scale, and
should be extended through a longer period of time. There can
be no doubt about the accuracy of the results in the case of the
wire cages, the growth in which has been described; and, from
their position, I have no reason to think that the clams were dis-
turbed on the other beds which have been cited as examples of
growth. Some of the higher beds seem to have been discovered
by clammers, and these were raked clean.
The figures giving the percentages of growth, though not numer-
ous, at least indicate the fact that the most essential feature of the
culture of the little-neck clam — rapidity of growth —is all that
could be desired. Neither has anything appeared which would
suggest a natural difficulty in the way of artificial culture.
26 NEW YORK STATE MUSEUM
DESCRIPTION OF FIGURES
Figure 1
Side view of large Venus mercenaria. Mantle fold on right side
of the body has been removed. The edge of the left fold of the
mantle is shown at m. The exhalent, ex. s, and inhalent, in. s,
siphons are modified parts of the mantle. |
Water bearing food and other floating substances enters the
space between the mantle folds — the mantle chamber — through
the inhalent siphon. Hanging in this chamber are the foot, f, and
gills, og and ig. -Cilia on the gills cause water to enter them,
forcing it to their bases, into the epibranchial chambers, ec, and
then backward and out of the body through the excurrent siphon.
This is indicated by fine, dotted arrows. The two large transverse
muscles —the anterior and posterior adductors— which, by their
contraction, close the valves of the shell, are shown at aa and pa.
Reference letters: aa, anterior adductor muscle; pa, posterior
adductor muscle; ec, epibranchial chamber ; og and 1g, outer and
inner gills; ap and pp, anterior and posterior palps; ex. s and m. s,
exhalent and inhalent siphons; f, foot; m, edge of left mantle fold;
Ss, ventral margin of shell.
Figure 2
Drawn to show that floating particles which touch the surface
of the visceral mass are taken posteriorly and thrown off into the
mantle chamber at x. From this region, they are removed from
the body by the contraction of the adductor muscles, which dis-
charges a large part of the water in the mantle chamber.
At pp is shown the striation of the inner side of the posterior
palp, over which food is taken to the mouth. The unstriated
margin is also shown.
Other reference letters as in figure 1.
Figure 3
Paper model of lamellibranch gill. A diagrammatic figure to
show the basketlike structure of the gill.
Reference letters: ic, interfilamentar connections; , partition
or septum holding the two halves of the gill together; r, a rod
or filament; s, space between filaments. ;
VENUS MERCENARIA ¥ 27
Figure 4
Diagrammatic section across the filaments of a typical gill.
Arrows represent the course taken by water which enters the
gill. Reference letters: 1g, interior of gill; p, septum between
sides; gc, gland cells, the secretion from which cements floating
particles into a mass on the outer surfaces of the gill; fc, fine
frontal cilia causing water to enter gill; sc, straining cilia pre-
venting solid matter from entering the gill and moving it to the
ventral margin.
Figure 5
View of inner surface of left mantle fold of Venus, showing
course taken by particles which touch it. These are discharged
from the body when the stream entering the mantle chamber
through the lower siphon is reversed by contraction of adductor
muscles.
Figure 6
Hard clams kept in wire cage above the bottom for six months.
All shells were covered by attached Anomia, or silver shells.
: Figure 7
Lunatia, a gastropod mollusk, which bores shells and destroys
clams.
Figure 8
Venus shells bored by Lunatia.
INDEX
Anatomical features of Venus mer-
cenaria, 6-12; cilia, 9-10; foot, 8;
gills, 7, 8-11; mantle folds, 7; palps,
7-8, 11-12, 14; siphons, 6 7, 13.
Beaches and flats, public grounds, 4.
Cilia of Venus mercenaria, 9-10.
Cultivation of Mya arenaria, 4, 6; of
Venus mercenaria, 15-17.
Enemies of Venus mercenaria, 24-25.
Enteromorpha interferes with clam
culture, 17.
Experiments to determine growth of
Venus mercenaria, 15-17.
Feeding habits of Venus mercenaria,
6-15.
Figures, description of, 26-27.
Foot of Venus mercenaria, 8.
Gills of Venus mercenaria, 7, 8-11.
Growth of Mya arenaria, 6.
Growth of Venus mercenaria, depen-
dent on food, 6; experiments to
determine, 15-17; method of meas-
uring, 18-19; between tide lines,
19-22; under wire netting, 23.
Hard clam, see Venus mercenaria.
Jones, Capt., acknowledgments to, 15.
Kellogg, James L., cited, 3, 10.
Little-neck clam, see Venus merce-
naria. ©
Lobsters, near extinction, 4.
Locomotion, of Mya arenaria, 22; of
Venus mercenaria, 22-23.
Long-neck clam, see Mya arenaria.
Lunatia destructive to Venus merce-
naria, 24—25.
Mantle chamber, 7, 9, 13, 14.
Mantle folds of Venus mercenaria, 7.
Mya arenaria, cultivation of, 4, 6; ob-
stacles to cultivation of, 17; enemies,
24; rapidity of growth, 6; growth,
above the bottom, 24; locomotion,
22; power of emptying mantle cham-
ber, 13.
Oysters, cultivation, 5, 16; growth, 15,
growth above the bottom, 23.
Palps of Venus mercenaria, 7-8, 11—
12, 14. '
Planting, difficulty of obtaining clams
for, 17.
Racks to hold oysters, 23—24.
Seaweeds interfere with clam culture,
W7pe10=205.29,
Seed clams, 4.
Siphons of Venus mercenaria, 6,7, 13.
Soft-clam, see Mya arenaria.
Starfish destructive to clams, 24.
Straining lines, 10.
Tide lines, growth of Venus merce-
naria between, 19-22.
Ulva, interferes with clam culture,
17, 19-20:
Venus mercenaria, supply in danger
of failing, 3-4; where found, 5;
anatomical features, 6-12 ; feeding —
habits, 6-15; growth experiments,
15-17; methods of measuring
growth, 18-19; growth between tide .
lines, 19-22; wandering habits,
22-23; growth under wire netting,
23; growth above the bottom, 23—
24; enemies, 24—25.
Wandering habits of Venus, 22-23.
Wire netting, growth of Venus merce-_.
naria under, 23.
Wire rack to hold clams above bot-
tom, 24.
Fa.
eC.
A gg
comes in Ss. AU
Figure 3
Figure 7
Figure 8
University of the State of New York
New York State Museum
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York; issued as part of Museum bulletin 15 and the 48th Museum Report, —
v.1. 59x67 em. 1894. Seale 14 miles to 1 inch. Separate edition out of
print. a ae <4
Geologie Map of New York. 1901. Scale 5 miles to 1 inch. In atlas
form $3: mounted on rollers $5. Lower Hudson sheet 60c. =
The lower Hudson sheet, geologically colored, comprises Rockland, Orange, Dutchess, Put-
nam, Westchester, New York, Richmond, Kings, Queens and Nassau counties, and parts of
Sullivan, Ulster and Suffolk counties; also northeastern New Jer sey and part of western Con-
necticut.
Map of New York showing the Surface Configuration and- Water Sheds.
1901. Boa 12 miles to 1 inch. I5C.
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