MOLLUSK
I
The
“Allow me the use of
Four species of oysters are grown commercially in
the
Each geographic area has traditional, even unique,
methods for cultivating, maintaining and harvesting its shellfish resources and
Chesapeake Ray is no exception. Generally colorful phrases and local jargon are
associated with these operations to the extent that the newcomer to this area
is often quite taken by them.
Unlike the European or Dutch flat oyster, Ostrea edulis, and
other members of the genus Osirea members of the genus Crassostrea have separate sexes. In the
summer during periods of spawning, both eggs and sperm. produced literally by the millions, are shed into the water
where fertilization takes place by chance contact. Fertilized eggs, or embryos,
develop into free-swimming, plankionic larvae called
veligers and are carried by currents. Development
continues as the bivalve shells form through the straight hinge and into the umbo stage. By the end of 10-25 days, depending on genetic
proclivities and environmental factors, baby oysters are ready to set into nonmotile stages for the rest of their lives.
By means of secretions from a “Cement gland”
oyster larvae attach themselves to various materials called clutch and the
newly settled oysters are called spat; a term both singular
or plural. A set of spat is called a strike, a heavy set is a “wrap up
set” or “wrap up strike.” Oysters in
It takes about 3 years for oysters to reach the
harvestable size of 3 inches in
Oysters that can be transplanted from one location
to another are called seed. Seed oysters can be of almost any size or age, but the are usually under 1½-inches long and 6 months to 1 or
even 1½-years old at the time of transplantation. Seed oysters are usually
transplanted along with the clutch material to which they have become attached.
Cultch, as used in
There are some privately owned and managed oyster
beds in Chesapeake Bay (more in
The
These methods largely involve a hunting approach
and ostensibly are carryovers from olden days. However, it should be noted that
each of these methods is technologically primitive, and fuel and other energy
expenditures are smaller than for other mechanized, more sophisticated fishing
practices. Recently, scuba diving has been introduced as a method of
harvesting in the Bay, and has created a considerable amount of concern (and
even animosity), among those engaged in the traditional pursuits.
Man, one of nature’s most aggressive and
persistent predators as well as polluters and despoilers, has managed to reduce
the harvest of oysters from
Infectious disease agents such as viruses,
bacteria, fungi, and protozoa contribute to fluctuations and declines in oyster
recruitment and productivity. Parasites and predators such as oyster drills,
flat worms, and crabs, as well as other plants and animals that compete for
space and food also contribute. Similarly, natural events such as storms,
hurricanes, drought, and abnormal temperatures can affect the health and even
accessibility of oysters. The 1973 summer hurricane Agnes is a good example of
a natural event causing catastrophic consequences. Large
amounts of fresh water run-off from the storm, carried chemicals and sediments
into the Bay and over shellfish beds, dropping shellfish populations
dramatically. Only recently has the Bay recovered from this natural
phenomenon.
Fishery management agencies involved with the Bay
try to keep abreast of new developments in oyster and shellfish culture.
Furthermore, using the best council available, they devise new strategies to
enhance recruitment, conserve, and if necessary, protect growing stages and
increase production of this economically important renewable resource.
Some of the oyster culture techniques used in
Probably foremost is the difficulty in overcoming
tradition in the multiple use of waterways in the
Off-bottom
culture systems, could probably be used effectively to
enhance recruitment of early life history stages of oysters. Suspended or
elevated off-bottom structures could be placed in broodstock
areas to capture seed for later transplant. Mechanized systems are used
currently in
Several
oyster hatcheries have been built in the
Pond culture of oysters along the edges of the Bay
and sea also hold promise even to the extent of using them in polyculture systems in which more than one species or type
of animal and plant can be grown simultaneously. Environmental conditions
(salinity, pH, dissolved oxygen. and temperature), type of food, and nutrients,
diseases, and predators can be more easily controlled or regulated in these
systems.
Culture of oysters in vertical systems such as
silos and in other systems such as tank farms and raceways are also being
attempted in some parts of the world all with possible future application to
To
return to our original question:
Would our Japanese
colleague still feel he could use
II THE MUSSEL
— A Bivalve for All Seasons
by JOHN
B. PEARCE
Of all the marine animals that the beach walker
might observe, the mussel and its colleague in the intertidal
environment, the barnacle, are by far the most common in those coastal habitats
that have pilings, rock outcroppings, and other hard surfaces where these
animals can attach. The common mussel, often referred to as the blue mussel,
is a medium size bivalve which can grow to three or four inches long. It has an
elongated foot which can be extended from between the two shells and can
actually be used to allow the animal to move about. This foot has a special
modification called the byssus, made up of several
threads which are produced by a gland in the foot. These threads are used to
attach the mussel to hard surfaces. The mussel is a highly social animal and
generally occurs in aggregations of hundreds or thousands where extensive rocky
coastlines occur, as in
The mussel is extremely adaptable, found
throughout the intertidal and sublittoral
zones. In areas which receive heavy wave spray, the mussel will occur high in
the intertidal environment. Moreover, because of its
adaptability, the blue mussel is circumboreal in its
distribution, i.e., it is found in temperate waters throughout the northern
hemisphere. The blue mussel occurs on the west and east coasts of the
Two other mussels are common to the east coast.
The horse mussel is a much larger species, most often found in deeper waters
and to the north of
The other common mussel on the East coast is the
ribbed mussel, a common denizen of marshes. It can be observed nested between
stalks of marsh grass in most of the common east coast wetlands. Neither the
horse mussel nor the ribbed mussel is regarded as suitable for human
consumption. They often have a bitter taste, and the ribbed mussel frequently
lives in environments which receive wastes or natural organic debris and is
therefore not considered fit to eat.
The blue mussel is, however, an important food
organism. It also functions within an important ecological niche in terms of
other marine animals. As previously mentioned, the blue mussel will often form
dense beds on rock surfaces or pilings, creating a habitat for many smaller
species of marine life. Polychaete worms, amphipods
or small shrimplike animals, snails, hydroids, small
brittle-stars, and other forms requiring protection from predation will often
live within and beneath the dense mussel beds. These small animals can move
about under the layer of mussels and thus receive considerable protection from
predators and from strong wave action and currents which might sweep them from
their preferred habitat. Because of the strong byssal
attachments, the mussels themselves are difficult to remove and are able to
withstand all but the strongest waves that pound exposed shorelines.
The mussel is an important prey for many species
of marine fish and invertebrates. Mouths of the temperate water cunner and tautug are specially
equipped to tear smaller mussels from their substrate. Other fish, such as
dogfish and rays, also actively prey on mussels. In turn, as the mussels are
gradually removed through grazing, other prey species are exposed and different
forms of fish and invertebrates can then feed on them. In addition to fish,
sea-stars, lobsters, and crabs, other invertebrates may use mussels for food.
Thus, this important marine bivalve, which itself obtains food by filtering
water for tremendous quantities of particulate foods, is able to grow very
rapidly and, as a population, serves as an important element of intertidal and sublittoral food
chains.
In
many instances, mussels in some parts of the world have been avoided during the
summer months because of the possible toxic seafood poisoning which can
develop. Traditionally, it has been held that mussels, along with other
filter-feeding shellfish, should be collected or eaten only during a month
with an “r” in its name. Marine scientists now
understand that seafood poisoning develops when some species of
single-celled phytoplankton are present in the water. Toxic forms of these phytoplankters produce a poisonous substance which is distributed
throughout the tissues of bivalves when ingested. Scientists now are able to
detect the presence of such phytoplankton species and in many parts of the
world waters are routinely analyzed to determine if they are free of toxic
substances. This allows the mussels to be collected and prepared for
consumption without undo concern for
mussel poisoning.
The ecological requirements for mussel
recruitment and growth, as previously noted. are
extremely broad. For this reason many cultures throughout the world have for
more than a century had commercial operations in which floats, ropes, and other
devices are used to collect the early larval stages of the bloc mussel in
shallow coastal waters and embayments. By properly
placing such collecting devices, the mussel “farmer” is able to gather crops of
mussels numbered in the millions. Because the mussel tends to reproduce most of
the year, there is often a steady supply of larvae, and it is possible to have
a more or less continuous crop of mussels developing within a given area all
year long. Through the management of such extensive natural systems it becomes
possible to provide food for large segments of the world’s populations. As
previously noted, mussels are extremely nutritious and even one or two individual
animals eaten several times a week would provide the minimal protein
allocations necessary for normal body maintenance and growth. Because the
mussel can be grown in extensive numbers in compact areas, it becomes possible
to provide food to extensive populations near mussel growing areas, cutting
transportation and processing costs. It also ensures that the nutrients from
runoff from the land are picked up as they pass through estuaries and before
they are carried into the open oceans where they
are effectively lost from a food web which culminates in man.
The
mussel has not only provided food for other forms of marine life, but has also been a delicacy in the diet of
many human cultures. Romans regarded mussels as highly appetizing. Today,
people throughout much of
To ensure that mussels are suitable for use,
however, it is essential that their environment be free of sewage and industrial
wastes. As is true for many bivalves, mussels can take up sewage wastes,
including pathogenic microorganisms, and transfer them to the ultimate
consumer, man. Mussels are also capable of biomagnification
of industrial wastes. For instance, toxic levels of heavy metals and organic
substances such as DDT and PCB's can accumulate in bivalves.
Given
the foregoing, and the benefit of seafood easily grown in coastal waters, it
is obvious that we must preserve water quality in estuaries not yet polluted
and, further, we must upgrade the water quality of major urban estuaries that
are already burdened with pathogenic microorganisms and industrial wastes.
This can and should be done. It will require recognition that coastal waters
and estuaries are suitable for purposes other than waste disposal and
industrialization. It will take a concerted effort by scientific and federal
and state agencies to identify problems, the sources of contaminants, and
solutions. It will require massive amounts of money to upgrade the sewage
treatment and industrial waste treatment systems within such areas. The
alternative, however, is to allow continued degradation of coastal waters and
to allow areas that are already heavily polluted to remain relatively
unproductive. Since estuarine and coastal water areas are limited, it is
obvious that mankind must make the best use possible of habitats that are
available to it for the production of seafoods. The
mussel is one of the more productive organisms available, a
relatively inexpensive seafood which has not only luxury connotations,
but can also he a staple in the diet of mankind.
III Common Chitons of
by
JAMES DUGGAN in Naturalist
The second largest and most familiar group of
invertebrates is the mollusca (“soft-bodied”). Members of the group. which include
snails, slugs, clams, oysters, octopuses, and squids are widely used as food
for man. Some typical anatomical features of mollusca
are variously modified, or even lost in the more specialized forms, like the
clam. Less common than their more conspicuous and economically valuable relatives,
chitons display the molluscan
body plan in its most typical form.
Chitons
belong to the class Amphineura,, known to exist for about 570 million
years, long before the first fish swam the seas. There are about 600 species of
chitons existing around the world, sonic 140 of which
thrive along the coasts of
Chitons
are sluggish animals, and most browse on the algal growth of rocks near the
seashore. They move very slowly and may remain in one area for long periods.
The body is bilaterally symmetrical. At the anterior end is an inconspicuous
head. The bottom surface is mostly taken up by a broad flat, muscular foot,
abundantly supplied with a slimy secretion. The foot functions for adhesion as
well as locomotion. Over the foot lies the visceral mass (containing most
organs) covered by a heavy fold of tissue extending around over the foot. This
fold is the mantle, and its peripheral area is called the girdle. This girdle
may be smooth and marked or covered with scales, bristles, or calcareous spicules, depending on species.
On
its upper surface, the mantle secretes a shell, which in most chitons consists of eight separate plates, overlapping
from front to rear. The digestive system is simply a tube extending from a
mouth in the head to the anus. The mouth leads to a muscular chamber in which
is found the radula. This is a horny ribbon with 17 chitonous teeth, some of which are capped with magnetite
(iron) and is used to rasp off algal fragments. A subradula
organ in the mouth area is used to locate food. Not all molluscs
have radulas, but nothing like it exists anywhere
else in the animal kingdom.
A chiton’s nervous
system is primitive, a ring of nervous tissue around the esophagus, connected
to two pair of long nerve cords which go to muscles in the foot and mantle.
There are no sensory organs except for the subradula
organ and aesthetes.
Aesthetes, unique to chitons, are mantle
sensory cells in the upper part of the calcareous plates. They provide dorsal sensory reception to
light. Most chitons are negatively phototactic and live under
rocks and ledges during the day and feed mostly at night or on cloudy days.
During
May of 1982, while working for the National Oceanic and Atmospheric
Administration (NOAA) aboard the Hydrographic Survey Ship, DAVIDSON, I was
able to collect, identify and observe four different species of chiton thriving in the cold waters of Southeast Alaska. The
locality was a rocky, fiord-like environment called the Bay of Pillars, located
near Kuiu Island about 110 miles SSW of the state
capitol, Juneau.
To
collect chitons for study one must remove the animal
from its substrate. This is not always easy but can be accomplished
successfully by inserting a thin bladed knife under the foot of the animal and
gently prying it oft the rock. A 10% buffered formalin solution is a good
preservative. To prevent the animal from curling up you can secure the chiton to a thin board with strong thread before immersion
in the fixative and leave it in the solution for about two weeks. The chiton can then be removed
from the board and left to dry in a shady spot.
The most abundant chiton
I observed was the Black Katy Chiton (Katharina tunicala).
This chiton has a thick, black girdle that covers
about two thirds of each plate. It lives on rocks exposed to heavy wave action
and full sunlight. Its range stretches from
I found a colorful, smaller chiton, the Lined Chiton (Tonicalla lineala), very abundant in tidal pools encrusted with a
pink corallinc algae, its primary food source. The
colors of the plates vary greatly, enabling it to camouflage itself against the
rocks surface. This chiton has a smooth brownish
girdle and ranges from
The Mossy
Chiton (Mopalia muscosa), tolerates
large changes in environmental conditions (i.e. salinity, temperature). This
animal has many stiff mossy hairs on its girdle, with dark brown to olive green
plates. It lives in protected areas where heavy wave action is absent. Its diet
consists of animal and plant matter
and can be found along the coast from
The
least common of the chitons collected was the Mottled
Red Chiton (Tonicella njarmorea), a chiton with very beautifully sculptured bright red-orange
and white lined plates. Its girdle is smooth with a leathery consistency. I
found this chiton in tidal pools along with the Lined
Chiton, feeding on coralline algae. This species
also ranges widely along the
The
chitons are of interest in the persistence of their
primitive body plan and their success in survival. The success of the molluscan plan, a compromise of a protective shell with
some degree of mobility, is proven by the existence of over 70,000 species of molluscs.
Sea shell collections have made the phylum
Mollusc the most familiar group of animals in the
sea. Collectors travel to all parts of
the world to find new molluscs. For example, there was a 32 year old man who
journeyed to a remote Indo-Pacific island to skin-dive and collect shells. Having picked up a uniquely decorated shell
to examine, he was stung lightly on the thumb before he could drop it. He did not feel the pain at first, but soon
discovered difficulty in breathing and although the wound was a little more
than a puncture mark, he died within three hours. What is the fascination with this group of
animals that would cause a man to travel halfway around the world and
eventually meet his death?
Over 100,000 living species of the phylum Mollusca have been described. Forty thousand of these are found in the sea
making this phylum the most abundant marine animal group. There is great diversity within the phylum to
which both clam and octopus belong. This large group is subdivided into six
classes, five of which are described below:
1. Amphineura (Polyplacophora) - the chitons
2. Scaphopoda - tusk shells
3. Pelecypoda (Bivalvia) -clams and oysters
4. Gastropoda - snails, slugs, nudibranchs,
limpets etc.
5. Cephalopoda - the octopuses and squids
6. Monoplacophora - Neopilina (Q1)
Octopus, clams, snails,
and squids certainly do not look very much alike. Why have biologists placed these different
animals in the same group? In placing
animals into groups, taxonomists look for similarities in development as well
as similarities in appearance. Animals
classified as mollusks go through a stage of larval development in which they
look the same. The larval forms of
marine mollusks are drifting zooplankton called trochophores. The trochophore
larva (see picture) is typical of marine mollusks. The hairlike cilia
propel the larva and also sweep food into their mouths. (Q2)
The body of an adult mollusk has a head, a
foot, and a visceral (VISS-uh-rul)hump. Inside the visceral hump are the digestive
organs, excretory organs, and the heart.
The visceral hump is covered by the mantle. This is a thin skin that in most species
secretes the calcium carbonate shell. The mantle hangs down over the sides and
back of the body, forming a space called the mantle cavity. The gills are in this cavity. The are the respiratory organs of the aquatic mollusks. Undigested materials go from the anus into
the mantle cavity before passing out of the animal. (Q3)
In most mollusks a current of water flows
through the mantle cavity. The water carries in oxygen and food, and carries
away carbon dioxide and other wastes.
You might think that mollusks with a shell have a problem disposing of
wastes. They do not. Water flows into the cavity through an
opening called the incurrent siphon. The water circulates over the gills, where
it picks up the carbon dioxide and wastes, and exits through the excurrent siphon. (Q4)
The type of shell or shells or the absence
of shells and the type of muscular foot are also characteristics used to divide
the mollusks into the classes mentioned above.
AMPHINEURA
(POLYPLACOPHORA)
The class Amphineura
includes the chitins with a "shell" made up of eight partially overlapping
plates lining the dorsal (back) surface of the body. The muscular foot of these mollusks is broad
and well adapted for clinging to rocks.
Most chitins are unable to keep their gills free of debris. The animals are therefore obligated to live in
clear water such as is common to rocky shores.
Food is obtained using a rasp-like radula to
remove algae, detritus etc. encrusted on the roks. Most species are light sensitive and are most
frequently found under rocks in tide pools.
Adult chitins seldom reach a length longer than two inches. The Gumboot chitin, found on the west coast
of the
SCAPHOPODA
The class scaphopoda
contains about 200 species of burrowing marine mollusks. These are the tooth or tusk shells. The shell is an enlogated
cylinder shaped
much like an elephant's tusk. The
attractive appearance of the shell and its relatively
scarcity are witnessed by the fact hat Northwest coast Indians use to use these
shells as money or wampum.
The shells, which are open at both ends,
average 25-50 mm (1-2 inches) in length.
The body is greatly elongated and the head and foot project from the
larger opening of the shell. These
animals burrow head down in sand or mud leaving only the posterior tip of the
shell above the sand. Water is drawn in
and expelled through the opening in the posterior tip of the shell. Oxygen and some food is
drawn in with the water. Using
thread-like tentacles, scaphopods capture microcopic food organisms from the sand surrounding them.
The majority of scaphopods
live burrowed in sand under water ranging in depth from 6 to 1830 meters
(20-5,500 feet). As a result the living
animals are not frequently encountered.
Judging from the number of shells washed up on beaches, however, scaphopods are not to be considered rare animals. (Q6)
PELECYPODA (Bivalvia)
Mollusks like clams, oysters, scallops,
and mussels are called bivalves because their shells have two halves or valves.
Bivalve mollusks belong to the class Pelecypoda or Bivalvia
(new one). A hinge connects the two
shells so they can be opened and closed by the animal. The shells are chalky structures made of
calcium carbonate (lime) and other materials secreted by the mantle. There are 3 layers: an inner, pearly layer next to the mantle; a
central chalky layer: and a thin outer layer made of the same material as the
hinge. (Q7)
Pelecypoda means
"hatchet footed". the hatchet shaped,
muscular foot helps the mollusks quickly dig into mud or sand. Many members of this class spend most of
their lives partly buried. The valves
are kept slightly apart and two siphons are extended into the water. Water flows into the mantle cavity through
the incurrent siphon. The water flows
through the gills before it is expelled through the excurrent
siphon. The gills remove dissolved
oxygen from the water and release carbon dioxide. They also serve another important
function. The gills are covered with a
thin layer of mucus which traps small food particles. Cilia on the surface of the gills carry the
food-bearing mucus to the mouth. These
mucus-feeders feed on dead and decaying organic matter and many
microscopic plankton found in the siphoned water. (Q8)
Many bivalve mollusks are important as
food. Clams, oysters, scallops and
mussels are all harvested in great numbers.
Many of these species are cultivated by man and serve as the basis for
thriving industries.
GASTROPODA
The most successful group of marine
mollusks is the gastropods;
sails, slugs, nudibranchs, limpets and
abalones. There are over 28,000 species
of marine gastropods. Most of these
animals have only one shell (univalves).
Some have no shells.
All gastropods possess a flat, muscular
foot used for creeping. Most gastropods
are slow travelers with a top speed of about 3 miles per hour. Gastropods also have a head, usually with
tentacles and eyes.
This group of mollusks
possess a rasping, tongue-like structure called a radula. The word radula
comes from a Latin word meaning scraper.
The ribbonlike oral structure contains rows of
serrations or teeth which the gastropod uses to file its food apart. Sometimes during the development of all
gastropods torsion occurs. Torsion is a
180 degree rotation of the internal organs.
This rotation results in the anus, gills, and mantle
cavity (all originally in the rear) all being located immediately behind the
head. The most widely accepted
theory to explain the survival importance of torsion was advanced many years
ago by the biologist Garstang. Garstang maintained
that torsion allows the head to be rapidly retracted into the mantle
cavity. This offers the gastropod
protection from attack. (Q9)
CEPHALOPODA
Octopus, squid, cuttlefish, and chambered
nautilus belong to the Cephalpoda, or
"head-footed" mollusks.
Cephalopods are he most highly developed mollusks and may even be the
most highly developed invertebrates.
The head of these headfooted
mollusks is well developed and contains a large, differentiated brain. This complex nervous system allows the
animals to be extremely active and provides for an effective control of the
activity. Well developed eyes, similar
to the human eye, are present. With the
notable exception of the chambered nautilus, most cephalopods have no external
shell. Instead, they have an internal
pen which supports the body.
Cephalopods possess a siphon through which
water can be expelled with great force.
This stream of water propels the animal forward or backward in a
jet-like manor. Some members of this
group have fins which enable the animal also to swim in a fish-like fashion.
The mouth is surrounded by eight or ten
sucker-bearing tentacles used for gathering food and for reproduction. A strong, parrot-like beak, frequently
covered with a poisonous slime, is present in the mouth for grasping food. Immediately behind the beak is a file-like radula which is used to grind through shells.
Effective protection is obtained via an
ink sac. The cephalopod can discharge a
cloud of black, inky fluid that hides its escape from enemies. The high degree of development of this group
is also seen in its behavior. Females
frequently watch over their young, which develop directly (no larval stage), as
they grow in "nest".
The giant squid is probably the largest
invertebrate in the world. It may grow
to be 18 meters long (58') and weigh 1,800 Kg (3,960 lbs). Sucker marks from these giants are often seen
on the skin of deep diving whales. In
spite of its reputation, the octopus is usually a timid animal. The largest octopus is found along the
Mollusks play a variety of ecological roles in
the life of the sea: from scavenger to
predator. Their diversity, abundance and
large range (where they are found) make mollusks one of the most important
groups of sea animals. Mollusks also
play a large role in man's interactions with the sea. This group provides the "shell
fish" which are widely harvested:
clams, oysters, mussels, abalone, squid, octopus, etc. It also provides the most realistic hopes for
"sea farming" or mariculture. The many members of the phylum Mollusca are both ecologically and economically important.
V Red
Gilled Nudibranch of the Old Sow by Robert Leahy UW v19 #3 p7
Fundy and
Along Fundy’s western
shoreline are several island communities known as the Quoddy
Loop. The area features rugged coasts that include deep water coves, inlets,
bays, and three islands (Moose, Deer, and Campobello) which collectively
border Old Sow.
The deep waters of the
The inhabitants of this underwater wilderness have
not been photographed extensively, primarily because of the cold water (average
temperature is 48 degrees F.) and the false assumptions that the marine life
here is not as colorful as in the warmer latitudes, nor is the water as clear.
Old
Sow drops to over 450 feet but the
underwater rim of this turbulent whirlpool appears to attract much of the
beautiful and prolific marine life which can be photographed during slack tide
in this “North Atlantic Caribbean.” Its near vertical
walls help provide a rich biosphere which has created an exceptional environment for growth and reproduction
of the red gilled nudibranch. Although their habitat
extends from the Arctic to
The red gilled nudibranch
is a member of the sea slug family — class Gastropoda,
phylum Mollusca. These gastropods don’t have true
gills, but breath through their skin or projections on
the dorsal surface. They vary in size, shape and color and are of two types:
the oval dorids, which have gills on their backs, one
pair of tentacles and feed on sponges and bryozoa;
and the aeolids, which have cerata
or fleshy protuberances from the main body which are not gills and two pairs
of tentacles. Gastropods are small animals. The adults rarely exceed two
inches in length and a half inch in width. They are translucent white, with an
opaque white strip down the middle of the back and a tapered tail.
The red gilled nudibranch is a member of the aeolid
group which feeds on hydroids and anemones. This shell-less mollusk appears
defenseless, but like most aeolids, utilizes ingested
nematocysts (stinging cells) of its hydroid prey to defend itself from
predators.
Up to 100 long
finger-like extensions protrude in rows on each side of its back. These
finger-like projections have a bright red (occasionally brown) core and an
opaque white ring near the tips.
These extensions are
not gills but extensions of the nudibranchs gut. The
ingested nematocysts are directed to the gut extensions and become a part of
its defense. The head has two pair of long
antennae,
and the front end of the foot has a sharp hook-like extension on each side.
These animals are
found mostly among seaweed and hydroids, their favorite dining entrees.
Reproduction takes place at any depth; however most egg ribbons observed and
photographed, were in 6 feet of water (at high tide) or deeper. Thr red gilled nudibranch spins a
spiral egg casing containing hundreds of tin embryos. The
developmental period of one to two weeks (depending on water temperature).
Few predators have been identified that seek out and feed on these eggs. Larva are free swimming and have small shells. The young
resemble adults and have large secretory glands which
may provide a defense similar to the distastefulness of butterflies. Their
color acts as a warning to shallow water fish that this morsal
does not taste good; thus increasing their safety on the run of Old Sow. Winter
flounder and cod appear to be the chief predators of this colorful aeolid.
Although prevalent
along the entire coast of Maine, the red gilled nudibranch
as well as the northern red tetia sea anemones appear
in larger numbers adjacent to the rim and in the bore of Old Sow. The rich
biosphere including a Field of hydroids created by the Fundy tidal flow
promotes astonishing growth an reproduction for this and many benthic animals
of the area. I also observed a low diversity of predators in this region. This
combined with the fact that nudibranch are brightly
colored and conspicuous during the day and may have a noxious defense to
compliment the ingested nematocysts, contributes to the large populations.
There are 32 species of nudibranchs
in the southern
VI Hermit Crabs--THE SHELL GAME by DAVID K. BULLOCH
Unlike other knights of the order Decapoda (prawns, shrimp, crabs, and lobsters), the hermit
crabs do not come equipped with full suits of armor. They lack any hard parts
aft of their cephalothorax, which leaves their soft
and defenseless hindquarters exposed to easy assault.
Their evolutionary response to nature which
provided only plated gloves and jackets but no pants has been to employ empty
snail shells as leggings. Among a few species in odd parts of the world, other
items have been pressed into service — bamboo joints, half a coconut shell,
and even a concavity in a bryozoan colony — but most
stick to gastropod attire.
Since hermits of various species come in different
size ranges, each must habituate the haunts of a similarly sized snail host to
find a custom fit. In the shallow waters of mid-Atlantic shores, Pagurus longicarpus (the
long-clawed hermit), the commonest of the small tideline
hermits, inhabits the shells of the periwinkle, Littorina littorea; the mud snail, Alectrion trivittata; and
the little moon snail, Natica pusilla.
A larger cousin, P. pollicaris (the flat-clawed hermit),
abundant in deeper waters from Maine to Florida, suit themselves in the
carbonate remains of the whelks, Busycon; the sand
collar snail, Natica; and moon snails,Polinices, or any other large shell it can find.
On the west coast, the tidepool
hermits, P. samuelis
and P. granosimanus,
have an overwhelming fondness for Tegula shells;
whereas P. hfrsuriusculus,
more plentiful in deeper water, prefer Acanthina or Olive/la.
Ten or more common species of hermits in the
tropics use the wide variety of shells found in warmer waters. The tropics are
also home to a land hermit, the soldier crab, Coenobita clypeatus, that leaves the
sea when just out of the larval stage, returns often when young, but wanders
further afield as an adult, returning only to
procreate. The adult has a fondness for the West Indian top snail, Cittariunz pica, and is the reason these shells
are often found deep in the rainforest and at relatively high altitudes.
Hermits
often share both the inside and the outside of their chosen mobile home with
others. The slipper shell, Crepidula, finds
the interior lip of a whelk a good place to live, and its presence doesn’t seem
to bother a hermit one bit. Deep inside, the polynoid
worm, Halo.sydna, or a polychaete,
Nenes, may set up housekeeping practically in
the bosom of the hermit. Whether these worms arrive before or during occupancy
by the hermit is unclear.
The outside of the shell can be cluttered with a
long list of life — anything from the lightweight fuzz of Htydractinia or a coat of bryozoa, to a substantial
load of barnacles or anemone that add serious wciglit
to the total load the hermit must haul.
Irrespective of
the burden, having an anemone on one’s shell is something of a status symbol in
the hermit’s world. A have-not, especially if it is bigger, will boldly
approach a have, grab the anemone around its midriff with its large claw, and
give it a squeeze, at which point the anemone will let go its hold on its
former home and, upon positioning, attach itself to the shell of its new conpanion.
Even fully
shelled, the hermit has enemies. The octopus, for one, and shccpshead, spotted kelpfish, and perch for others.
In
Sex has its
dangerous moments as well. Somehow the females growing receptiveness is sensed
by the male which will seize her shell and carry it (and her) with him
occasionally banging their two shells together. When both have reached the
necessary level of stimulation, they simultaneously pop out of their shells,
mate in a flash, and pop back into their respective
quarters.
Aside from the
daily grind of searching for food
and occasional sexual forays, the hermit finds house hunting a consuming
preoccupation. Although there have been a few reports to the contrary, most
species have the common decency to wait until the original occupant passes on
its legacy via natural causes. This is probably due more to the impervious
defense the snail can muster by
withdrawing its foot
and
blocking an attack than any altruism on the part of the hermit.
Presented with a
cluster of empty shells, the hermit ritually repeats an elaborate inspection of
each one, turning the shell over and over, probing its passages, and if
eminently suitable, popping out of its old home into its new. Even a new shell
home does not stop it from going through the same routine with every encounter.
If it comes across a shell it has recently examined, the normal riganiarole will be
duly abbreviated as it somehow senses it is going over ancient history.
If a new shell is
already occupied by a hermit, neither is deterred from the shell game. If the
bigger of the two decides it wants to try out the shell of its smaller
acquaintance, it is sure to get its way. The vanquished, forced to withdraw
from the shell, is not molested by the victor and will usually pop into the victor’s
hand-me-down or, if that’s not possible, quickly burrow into the sand to get
out of harm’s way before some other predator comes along to take a snap at its
hindquarters.
The hermit’s house
hunting is entirely innate, unlearned, and hard-wired into its genotype. That
is not to say the hermit is incapable of adaptive behavior. It is, but the presence of an empty shell
automatically sets off a foreordained response that is present from the very
first time it sets eyes on a shell.
The ethologist, Konrad Lorenz, calls
this behavior a “species-characteristic drive-action” (the term in German “arteigene triebhandlung” doesn’t
translate into anything more succinct). It is a set of neurophysiological
sequences in which a selective response to a specific stimulus releases a fixed
motor pattern. Put another way, a pattern of movements is evoked by an impulse
internally produced but held in check until externally released by what has
come to be called an “innate releasing mechanism”
(IRM).
This pattern of
movement and its trigger, the IRM, is similar from one species of hermit to
another, indeed from one genus to another within the suborder Paguridea, with as much constancy as their common physical
characteristics.
These motor
sequences have a phylogenetic origin and are part and
parcel of the hermit’s inheritance. As such, hermits are highly resistant to
individual modification. However, all animals have a feedback circuit that
interprets the consequences of their behavior; specifically whether it is
useful or not under prevailing circumstances. If you were to partly fill some
shells with plaster of Paris or, more challenging, lightweight plastic foam,
the hermit will learn by examination that the shells are useless.
Because they are
ceaselessly active, hermits make good aquarium additions. They are easy to
maintain. Feed them brine shrimp and, occasionally, a
bit of seaweed. Forget hunks of meat that might entice other crabs. Hermits eat
only small fragments and will not tear big pieces apart.
If, by chance, you
want to entice a hermit crab out of its shell to temporarily get a better look
at it, give up all thoughts of pulling it out. It will hold fast with two
claw-like uropods affixed to its hindquarters and you
will only break it in two.
The
best way to get a hermit out of its shell is to tempt it with a new shell with
a hole drilled in an appropriate rear whorl. When the hermit takes up residence
in the shell with the back door open, you have only to prod into the hole with
something pointed to elicit its departure. In fact, one marine biologist so
acclimated a pet Pagurus that it immediately dropped out of its
shell without a poke whenever it was lifted out of the water. Put back in the
aquarium, it would await the return of its shell, and instantly resume
residence when the shell was dropped back into the water.
VII
Suspension Feeding by Bivalves: The Inside Story
The ocean is clouded with small particles of many
types, defying a careful and complete description. Large rivers empty specks of
degraded leaves, clay and silt particles, and detritus of many other types into
an ocean already filled with phytoplankton, smaller zooplankton, and a plethora
of animal and plant degradation products and exudates that collide in the water
column, to eventually form what is often termed marine snow. A typical coastal
water column has all this complexity, but even deep waters may be surprisingly
rich in suspended particles.
Suspension-feeding animals must face this
complexity and eek out a living by collecting particles, ingesting them, and
absorbing nutrients. Many of the particles in the water, however, have no
nutritive value, Of course one might imagine a suspension feeder ingesting all
particles indiscriminately, digesting only those that are palatable, but this
strategy would have two important disadvantages. First, the gut would be filled
with large amounts of indigestible material. This would waste opportunities to
digest perfectly good food. Second, materials might enter the gut that have
negative physiological effects and even are poisonous. For example, the saxitoxin-bearing dinoflagellate Alexandrium has strong negative effects on bivalve
function, so why would a bivalve want to ingest it? Why not reject the toxic
substance beforehand?
Long ago, a simple observation demonstrated that
bivalves could reject particles and even discriminate among different particle
types. If you placed an oyster in a dish and fed it a high concentration of
phytoplankton or a large amount of silt particles, you would soon see diffuse material
at the bottom of the dish, emanating usually from the incurrent siphon or from
between the shells. Flow did this happen? The internal space of a bivalve, its
mantle cavity, contains a series of structures that transport and potentially
reject particles before they cuter the mouth. Ibis process is known as preingestive selection. Particles enter a siphonlike entrance and then are drawn toward the surfaces
of the gills. The water movement within the mantle
is stimulated by a
complex set of ciliated tracts that heat continuously, creating a pressure
differential that moves water and particles through the incurrent siphon and toward
the gill surfaces. It appears that particles hit ciliated surfaces by inertial
impaction, whereupon they become subject to selectivity (see Box Figure 13.2b),
are moved by ciliary beating to more centralized ciliary tracts, and may then be transported to another main
structure known as the palp (Box Figure 13.2b). The
bivalve palp varies in structure depending upon
species, but it is typically highly ridged, and the ridges and valleys are also
ciliated. The palp moves particles along its own
ciliated systems and eventually transfers some to the mouth.
To investigate this type of feeding, particles
could be placed upon both gills and palps that have
been surgically removed and mounted on slide, If the
ciliated tracts were “smart,” they could reject bad particles before the palp had transferred them to the mouth. The rejected
material that is never ingested is known as pseudofeces;
this is the stuff mentioned earlier that was found in a dish with a bivalve, as
opposed to true feces, which is matter that is egested from the gut. Pseudofeces production may occur by the gill sloughing off
material directly or passing it to the palp, which
can also form and pass pseudofeces outside the mantle
cavity.
Do bivalves manage to select against nutritionally
poor or even toxic particles before ingestion? A simple experiment suggests
that they do. One can make a mixture of silt and phytoplankton and feed the
particles to the bivalves. Presumably the phytoplankton are
suitable food and the silt is not. One can measure the organic content of the
food by sampling the experimental water and compare the results to measurements
made of pseudofeces. Such experiments generally show
that the pseudofeces are proportionally depleted in
organic material, suggesting that selective rejection of nonnutritious
material is occurring.
But where is the site of
selectivity? This has been a mystery for many years. It has been possible to
surgically removed structures such as gills and palps
to trace the directions of ciliary beating, but this
precludes observing the entire mantle cavity organ system as an integrated
unit and disrupts any special hydrodynamic conditions that might occur in the
intact mantle cavity.
Enter (literally) the surgical endoscope, which is
a tubular lens that may be made of a Fiber-optic bundle or may be a single
glass lens. For
bivalves, endoscopes of as little as I mn
in diameter are available, which allows for easy insertion into the mantle
cavity, especially of larger bivalves. The endoscope can be attached to a video
monitor and video tape recorder, allowing recordings of the movements of
particles within the mantle cavity.
Another instrument aids us greatly in studying
particle selectivity. The flow cytometer is an
instrument adapted
From medical uses to marine biology.
The principle is fairly simple. A small volume of water is drawn into a tube
and as particles pass through the tube, they are struck by a laser beam.
Particles reflect the laser light and can be counted and even sized, because
the larger the particle cross section, the greater the reflection.
Phytoplankton cells have a variety of photosynthetic pigments, many of which
fluoresce when bombarded by blue light. For example, when chlorophyll is
bombarded with ultraviolet Iight it fluoresces a
vivid red color.
The flow cytometer reads
the light reflected and fluoresced from each particle by means of a series of
detectors. The combined signal of each particle allows us to discriminate
among a variety of different particle types.
Flow cytometers are extremely useful in analyzing
marine water samples, but they also can be used to tell if a suspension feeder
is selecting among different particle types. For example, one could feed a
series of phytoplankton species and tell if some are being selectively
ingested merely by comparing counts in the water column with particles in the pseudofeces. If there are proportionally more particles in
the suspended food than in the pseudofeces, then the
bivalve must be selectively ingesting those particle types because they are
depleted in pseudofeces.
A simple experiment demonstrated an interesting
result. Sandra Shumway and colleagues’ pioneered
in using the flow cytometer to study selective
feeding by bivalves. They fed mixtures of algal cultures to see whether species
of algae were preferentially ingested, which could be told by comparing
available food particles in the water with the material rejected in the pseudofeces. On comparing samples of algal mixtures, it
was clear that a variety of bivalves were selectively rejecting certain
phytoplankton species and preferentially retaining others for ingestion.
But where did the selection occur? Roger Newell
and Stephen Jordan did
an experiment that set the stage for the problem. After feeding the eastern
American oyster Crassos!rei virguzica, a
mixture of silt and the alga Tetraselmis sp.,
they found that the pseudofeces were strongly
enriched with silt relative to the mixture in the water. This suggested that
oysters are able to ingest organic rich particles and reject nonliving
particles in the pseudofeces. Not only was the
selectivity clear, but it worked in favor of excluding indigestible material
(i.e., the silt) from ingestion. Because this study compared food in the water
and in pseudofeces, it was nut possible to
demonstrate where selection had occurred in the mantle cavity. Newell and
Jordan followed the popular wisdom that oysters selected particles on the palp, maintaining that the gills had no role.
The use of a surgical endoscope clarified this
problem definitively. J. Evan Ward and colleagues’ examined the
feeding of Crassslna virginica by
placing the tip of an endoscope between the open valves. The structure of
oyster gills is rather different from those of many other bivalves. The gill is
folded, or plicate, and the valleys are lined with cilia that beat dorsally.
The “hills” also are covered with cilia and can either transfer particles or
ventrally. If particles were transported dorsally, they arrived at a basal
ciliated tract that transferred particles in a slurry
toward the palp and were ingested. Particles
transported ventrally, however were moved to a so-called ventral groove and
were either rejected from the gill or transported to the palp
and rejected from there as pseudofeces. The
bidirectional option of particle transport immediately suggested the gill as
the possible site of particle selection, rather than the palp,
which was the standard belief of bivalve researchers for many decades.
A simple experiment
performed by J. Evan Ward and colleagues combined observations using the
endoscope with sampling directly from various ciliated tracts in the mantle
cavity. While the endoscope was focused on various parts of the gill, a fine
pipette was located to sample material from the dorsal and ventral ciliated
tracts. It was already known that the dorsal tract carries particles that will
be ingested and the ventral ciliated tract carries particles that are to be
rejected. Therefore, one could sample these two tracts and compare them with
the particles in the water, representing the available food. A flow cytometric analysis could then determine whether the gills
were sorting the good from the bad particles.
Ward and colleagues used an approach similar to
that of Newell and Jordan. They fed the oysters Crassostrea inginica and C. gigas
with aged cord-grass (Spartina
alterniflora) detritus, which they had ground to
match an alga (Rhodonzonas)
in particle size. The detritus was very poor in nutrients, much like the
silt used by Newell and Jordan. The results were immediately apparent,
especially because Rhodonzonas is a vivid red. The particles in the
dorsal tract were clearly red and the ventral tract was tan, the color of the
cord grass detritus. This demonstrate that it
was the gill doing the sorting, not the palp.
Not all bivalves work this way. Mussels mainly
transport particles ventrally and these particles are usually transported to
the palps. Here the structure of the gill (much
simpler) precludes sorting at this stage, and it is the palp
that does the job. Indeed, the palp sorts among the
cord grass detritus and Rhodonzone
way to get the job done, apparently.
This research opens tip of the “black box” and
allows us to mechanism is completely unknowm. No chemosensory cells have yet been found on
bivalve gills. Yet the impact of good and bad Particles is literally spread all
over the gill, so there must be a general system of detection. We also have no
idea why different bivalves have such different gill architectures. Mussels do
not select particles on the gills, yet oysters do this routinely. Both types of
bivalve often live in near shore estuaries with complex arrays of particle
types, nutritionally poor as well as appropriate. The results thus far
therefore just scratch the surface..
VII Pearls
A pearl is unique. It is the only gem created by a
living creature! A pearl is formed when a foreign particle enters the body of
an oyster (or other mollusk) along with some of the special cells which secrete
nacre - the pearly substance of the pearl. For centuries, divers have searched
the seas hoping to find an oyster with a pearl!
Today man has formed a partnership with the
mollusk which has resulted in the fabulous CULTURED PEARL INDUSTRY. The center
of this industry is in
The mollusks are
collected and opened and the pearls are separated from the bodys.
The lovely harvest is sorted according to size, weight and color. The pearl
industry recognizes 7 colors as acceptable: white, cream, pink, green, blue and
black. The Japanese Ministry of International Trade has enacted very strict
regulations which govern the growing, sorting and selling of the pearls. This
guarantees that all Japanese cultured pearls will be of highest quality.
IX
About Octopi ....
Octopi are
the phylum Mollusca which includes chitons, snails and clams. Their closest relatives are the
chambered nautilus, cuttlefish and squids.
The
largest and smallest octopi are found off the
The most
often encountered on the west cost of
Octopi
have the most complex brain of the invertebrates (animals with out backbones).
They have long term and short-term memories as do vertebrates. The octopus
learns to solve problems by trial-and-error and experience. Once the problem is
solved, the octopus remembers and is able to solve it and similar problems
repeatedly.
Sense of
touch is acute in the suckers. The rim of the cups are
particularly sensitive. A blindfolded octopus can differentiate between objects
of various shapes and sizes as well as a sighted octopus.
The
octopus has highly complex eyes which compare to human visual acuity. Focusing
is done by moving the lens in and out rather than by changing its shape as the
human eye dose.
When
threatened, the octopus will often try to escape by releasing a cloud of
purple-black ink to confuse the enemy. It will change color and jet away to
safety. Several blotches of ink can be released before the ink sac is empty.
The ink is toxic to the octopus who will become ill or
perhaps die if released in a confined area where the octopus can't escape the
ink.
Color change
is initiated by the eyes. If the octopus is disturbed, cells in the skin (chromatophores) will be active in an attempt to blend in
with the surroundings. The chromatophores consist of
three bags containing different colors which are adjusted individually until
the back ground is matched. Coloration reflects mood, white for fear, red for
anger, brown is the usual color.
Many
octopi produce venomous secretions. In members of the Octopus genre, this venom
is fatal to crabs and lobsters, their main food source.
Octopi have separate sexes (male and female)
and fertilization is internal. In some species, the male can be distinguished
by greatly enlarged sucker discs on the tentacles, which is used to remove a
mass of spermatophore from within his mantle cavity
and insert it into the mantle cavity of the female. Within two months after
mating the female attaches strands of clustered eggs to the ceiling of her
lair. More than 200,000 eggs may be laid. She shields the eggs, oxygenates them
with streams of water from her syphon and cleans them
with her suction cups. Most females will not eat after laying eggs and die soon
after they hatch. Young octopi are carried about in water currents for about a
month before they settle to the bottom. Only one or two out of 200,000 eggs
will survive to become adult.
1
The
1. What species
of oyster is grown in the
2. Why are exotic
species not grown there?
3. What happens
to eggs from fertilization until attachment?
4. What do
oysters use to attach themselves?
5. What sex are
most oysters during their first 3 years of life..What happens later in life?
6. What is a
clutch? a seed? a spat??
7. What do the
watermen do in the
8. How much has
the harvest declined....what has caused it?
9. How did
hurricane Agnes affect the population?
10. Why is
off-bottom culture not useful in the
11. How does off
bottom culture system work?
12. What type of
culturing is being tried?
13. Will oyster cultivation in the
1. What is a byssus
2. Where,
geographically (range) are mussels found?
3. What are the
three types of mussels listed....What use is each for food for man?
4. How do mussels
benefit other organisms in a mussel bed?
5. What organisms
feed on mussels?
6. What can cause
poisoning in mussels?
7. What is the
nutrition value of mussels?
8. What is the
main drawback of raising mussels in coastal waters?
9. What is the main idea of the last paragraph?
3
"Common Chitons of
1. To what class do chitons
belong?
2. How many
species are there and how old is this group?
A) B)
3. What percentage of the known chitons lives along the coasts of
4. How big is the largest chiton
and what is it called?
A) B)
5. What kind of symmetry do
these animals exhibits?
6. What is the function of the muscular foot?
7. What layer secretes the shell?
8. Describe the chiton's
digestive system.
9. Describe the chiton's
nervous system.
10. What do they
eat?
11. Who first
discovered the chitons?
12. Why is the molluscan
body plan so successful?
1. Which one of the five classes mentioned above
do you think has the greatest number of species?
2. Which is one observation taxonomists use to
place animals in the phylum Mollusca?
3. What are two functions that the mantle
performs?
4. What
substances of importance to the mollusk does the incoming water carry?
5. Since chitons can
only live in clear water, how might we use them as an indicator of
environmental pollution?
6. Considering the way scaphopods
capture their food, do you think more scaphopod
species would be found living in the mud or sand? Why?
7. Shells play a number of different roles in
the life of marine mollusks. What are
two ways in which shells are
used to help marine animals survive.
8. What is the path of water flowing through a pelecypod mollusk?
9. How has torsion increased the survival
chances of gastropods?
10. List three structures and behaviors that make
marine scientists think that cephalopods are the most highly developed molluscs:
1....................................
2....................................
3.....................................
11. Describe the
BEAK in Cephalopods.
12. Which
organisms best provide hope in mariculture?
5
"The Red Gilled Nudibranch of the Old Sow"
1. Where does the
red-gilled nudibranch live?
2. Describe the
place called "Old Sow".
3. How high is the tidal range and what is the
speed of water movement in and out?
A) B)
4. What is the
family, class, and phylum for the nudibranch" A)
B) C)
5. What does the red-gilled nudibranch eat?
6. What does it use to protect itself7
7. What
animals are the chief predators?
6. Hermit Crabs
1. What can be found outside the shell?
2. How would you get a hermit crab out of its
shell?
3. What does the presence of an empty shell do to a
hermit crab?
4. What is meant by the analogy "provided only
plated gloves and jackets but no pants"?
5. To what Order do hermit crabs belong?