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FIGURE 11.4 Mussels (Mytilus californianus) FIGURE 11.5 These seaweeds cant move to FIGURE 11.6 The shell ridges on this tropical
form dense clumps that retain moisture. This helps moist depressions at low tide, so they live there snail (Nerita plicata) help it stay cool by radiating
protect the mussels and provides habitat for smaller permanently. They dont grow on the surrounding heat. The snails white color probably helps by
organisms. rock because it dries out. reflecting sunlight.
be completely shut. These organisms typically clamp themselves Most intertidal organisms can tolerate a wide temperature
tightly to the rock to seal the opening. Some use mucus to make range. Tide-pool fishes, for example, are much more tolerant of
a better seal. They may also carve out shallow depressions, or extreme temperatures than related fishes that live below the tides.
home scars, in the rock that make the seal more effective. They Certain species of periwinkle from high in the intertidal show
do this by scraping the rock with their shell or their radula, remarkable heat tolerance. One is known to survive temperatures
slowly wearing the rock away. Every strategy has disadvantages, as high as 49C (120F) in the laboratory.
however: Clamming up can make it harder for animals to obtain Organisms deal with temperature extremes in ways besides
oxygen and food. simply being tough. Those that move to moist hiding places to
Some organisms use a combination of strategies. The avoid drying out, for example, also avoid high temperatures
periwinkles (Littorina) shown in Figure 11.2, for example, because such places tend to be cool as well as damp. Some snails,
clamp themselves to the rocks to seal in moisture. They can especially in the tropics, have pronounced ridges on their shells.
also seal off the opening of their shell by closing the opercu- Like the fins on a car radiator, these ridges help the snail lose
lum, a stiff plate that fits the opening like a door. Periwinkles excess heat (Fig. 11.6).
are still not immune to desiccation, however, so at low tide The color of a snails shell can also help it tolerate high
they congregate in moist, shady places, especially on hot, sunny temperatures. Snails that are regularly subjected to extreme high
days. Desiccation is a particular problem in hot, tropical cli- temperatures tend to be light in color. The light color helps
mates, where intertidal organisms are often confined to tide reflect sunlight to keep the snail cool. On Atlantic shores, for
pools or damp, shady spots. example, the dog whelk or dog winkle (Nucella lapillus) has two
Finally, some intertidal organisms use neither the run-and- color forms: white-shelled and brown-shelled. On shores
hide nor the clam-up approach. Instead, they simply tolerate exposed to heavy wave action, brown whelks predominate. On
drying out. Some intertidal chitons can survive the loss of 75% sheltered shores the white form is most common. The differ-
of the water in their tissues. Some intertidal seaweeds, such as ence appears to be related to temperature. Brown whelks absorb
rockweeds (Fucus; see Fig. 6.5), can withstand a water loss of as more heat than white ones and die in temperatures that dont
much as 90%, becoming almost completely dry and practically seem to harm the white snails. On the exposed coasts where
crunchy. They quickly recover when the tide comes in and wets brown snails live, there are dense mussel beds, which stay moist
their tissues. (see Fig. 11.4) and relatively cool. Wave spray also helps cool
the snails. Sheltered coasts have few mussels and little wave
spray so they are hotter, and white whelks have an advantage.
Some intertidal organisms avoid drying out by moving to or living
in wet spots. Others can close a shell to retain water. Still others
Salinity also fluctuates widely in the intertidal. When it
are able to dry out and then recover after the tide returns. rains, exposed intertidal organisms have to endure fresh water,
which is fatal to most marine organisms. Many simply keep
the fresh water out by closing their shellsanother benefit of
the clam-up strategy. Even so, rainstorms during low tide
Temperature and Salinity Emersion creates problems other than sometimes cause the mass mortality of intertidal organisms.
desiccation for marine organisms. Sea temperatures are relatively Tide-pool residents, too, face extreme variation in salinity. At
constant and mild because of the high heat capacity of water, but low tide, pools may be diluted by rain, lowering the salinity. On
air temperatures can be much more extreme. At low tide, intertidal hot, dry days, salinity rises because of evaporation. To cope, tide-
organisms are at the mercy of the suns heat and the freezing cold pool organisms can usually withstand a wide range of salinity as
of winter. Because tide pools are shallow, they too experience extreme well as temperature. They may also burrow or reduce their activ-
temperatures, though usually not as extreme as air temperatures. ity to ride out extreme conditions and wait for high tide.
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that scrape algae, bacteria, and other food from the rocks. Others
feed on pieces of seaweed and other large detritus that drift in from
deeper water. Some are predators and move over the rocks in search
of prey. When mobile animals seek shelter or clamp to the rocks to
avoid water loss at low tide they cant move around to find food.
Being unable to feed when the tide is out is not much of a
problem for animals low in the intertidal. They are immersed most
of the day and have plenty of time to feed. Higher in the intertidal,
however, animals may not be underwater long enough to get ade-
quate feeding time. This can cause them to grow more slowly or
even prevent them from living in the high intertidal at all.
FIGURE 11.7 Ocean waves expend tremendous energy when they break
on the shore.
Many intertidal animals are unable to feed when exposed at low tide.
This may prevent them from living higher on the shore than they do.
FIGURE 11.8 Nearly all waves approach the coast at an angle. As a result,
they refract, or bend toward the coast. By the time the waves break they are
Radula A rasping ribbon of small teeth that is present in most molluscs.
almost parallel to shore.
Chapter 7, p. 129; Figure 7.21 inset
5
4 3 Incoming waves refract, or bend, to become
2 1 more parallel to the shore. This increases wave
impact at headlands and decreases it in bays.
erg
Wave en er
rgy
en
ne
er
ee
Wave
ve
Wav
Canyon
Exposed coast
(North Gavel, Scotland)
4
3
Absent
Meters
above
low 2 (a) (b)
tide
FIGURE 11.12 The giant green sea anemone (Anthopleura xanthogrammica;
see Fig. 11.26) lives low in the intertidal zone on the Pacific coast of North
1 America. (a) In sheltered locations the anemone grows relatively tall.
(b) Anemones from areas with heavy wave action are much shorter. This
reduces the body area that is exposed and allows more water to slide over the
top of the anemone. In other words, drag is reduced.
Spiral Sea Bladder
0 rockweed rockweed rockweed
Sheltered coast and can go with the flow (Fig. 11.13). There may also be safety
(North Haven, Scotland) in numbers (Fig. 11.14).
Meters 1
above Absent Organisms on exposed coasts deal with wave shock in different
low
tide ways, including strong attachments, thickened shells, low profiles,
0
and flexibility.
Spiral Sea Bladder
rockweed rockweed rockweed
Even the firmest grip on the rocks is of little help if the
FIGURE 11.11 Even closely related species may be affected differently by wave smashes a loose rock or drifting log into the rocks, crush-
wave exposure. On the Scottish coast the bladder rockweed (Fucus vesiculosus)
occurs only in sheltered locations, the sea rockweed (F. inflatus) only at exposed ing whatever is there. Waves can also turn rocks over, even
sites. The spiral rockweed (F. spiralis; see Fig. 6.5) occurs in both environments but large boulders. When this happens the organisms on top of the
lives higher up on the exposed coast. This is probably because wave splash reaches rock may be smashed or buried, and sunlight for the algae is
higher at exposed locations. Such differences may be caused by the effects of waves
on competitors or predators rather than the direct effects of wave shock. Diagrams blocked. The underside of the rock supports different organisms
like this are known as kite diagrams. A wider blue kite indicates that more
individuals of that species are at that elevation above low tide.
Carnivores
Herbivory
Sea stars, snails, Predation
fishes, crabs,
flatworms, birds Detritus feeding or export
Contribution to detritus pool
(a)
(b)
FIGURE 11.16 Most organisms in the intertidal zone live attached to the rocks, and most of the avail-
able space is occupied. This little patch of rock supports two species of mussels and two of barnacles, plus
limpets and sea anemones. What little open space there is doesnt last for long: A small bare patch (arrow)
has already been colonized by newly settled barnacles.
of space commonly limits rocky intertidal populations. Nearly Many intertidal organ-
all the space in the intertidal is occupied (Fig. 11.16), and isms simply grow over (c)
colonizers quickly take over what open space there is. Space is their competitors, mak-
so scarce that organisms may attach to each other instead of ing them vulnerable to FIGURE 11.17 The sea palm (Postelsia
palmaeformis; see Fig. 6.7) takes advantage
to the rocks. waves (Fig. 11.17), of wave action to compete for space.
smothering them, or in (a) Adults literally drip spores, which develop
the case of seaweeds, into young sporelings near the parent.
Rocky intertidal populations are often limited by space, not food blocking precious sun- (b) When the space below is occupied, the
sporelings grow on these competitors,
or nutrients. light. Some species grow which are eventually pulled from the rocks.
in colonies, gradually (c) This clears space for a new generation
increasing the amount of of young sea palms.
Not surprisingly, competition for space is a dominant bio- space they hold as the
logical factor in the rocky intertidal. There are many ways to colony grows.
compete for space. One is to be the first to get to open spots. Two species of mus-
For intertidal organisms, this means having an effective means sel provide an example of how competition and physical fac-
of dispersal; that is, organisms that depend on being the first to tors can interact to determine the distribution of organisms.
occupy new patches of open space must be good at getting them- On the west coast of North America the blue mussel (Mytilus
selves or their offspring from place to place. Most rocky intertidal galloprovincialis; also called the Mediterranean or bay mussel)
species disperse via larvae or spores, which settle on the rocks to
colonize open space. Having taken over the space, the organism
must either be good at holding on to it or be able to reproduce Photosynthesis
rapidly to disperse its young to the next opening. Both strategies CO2 H2O sun energy organic matter O2
are used in the intertidal. (glucose)
Rather than colonizing open patches, some intertidal organ- Chapter 4, p. 66
isms take over space that is already occupied. Barnacles, for
Limiting Resource A resource that is so scarce that it restricts the
instance, undercut their neighbors, loosening them from the
growth of a population.
rock. Owl limpets (Lottia gigantea) maintain a territory by bull-
Chapter 10, p. 213
dozing out intruders. Less forceful methods may also work.
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is found mainly in sheltered locations, whereas the California mainly by physical factors, whereas the lower limit is usually
mussel (M. californianus) is found on the wave-exposed open determined by biological factors, especially predation and
coast. As you might expect, the California mussel has a thicker competition.
shell and attaches more strongly to the rocks than the blue
mussel. On the other hand, the blue mussel does just fine on
Most rocky shores have a distinct pattern of vertical zonation.
open coasts as long as the California mussel is absent. The blue
The upper limit of a zone is often set by physical factors, the
mussel is apparently rare on open coasts not because it cant
lower limit by biological ones.
take the waves but because it is out-competed by the California
mussel. The California mussel, with its thicker shell, crushes
the blue mussel when the two grow together. On the other This rule is a useful starting point in understanding the
hand, the blue mussel, unlike the California mussel, tolerates a causes of vertical zonation. As with all generalizations, however,
lot of silt and can therefore survive in calm bays. there are exceptions. For example, zonation can be produced if
the larvae of a species settle only at a particular height on the
shore. One cannot just assume that the upper limit of a certain
Vertical Zonation of Rocky Shores species is set by physical factors. Instead, this hypothesis must
Rocky intertidal communities vary markedly from place to place, be tested by experiment (see Transplantation, Removal, and
but there is one feature that is remarkably constant. The rocky Caging Experiments, p. 252). Furthermore, different factors
intertidal community is usually divided into distinct bands, or zones, often interact to determine the limits, and sometimes the line
at characteristic heights in the intertidal (Figs. 11.18 and 11.19). between physical and biological factors is fuzzy. As noted previ-
Thus, a given species is usually not found throughout the intertidal, ously, for example, some filter feeders do not live high in the
but only within a particular vertical range. Biologists call this pattern intertidal because they are not submerged long enough to feed.
of banding vertical zonation. Is this upper limit due to a physical factoremersion by the
When the shore consists of an evenly sloping rock face, the tideor a biological onefeeding?
zones are often sharply defined belts that can be easily distin- Many rocky shores around the world show a pattern of
guished by the colors of the organisms (Fig. 11.18). Many shores vertical zonation that is quite similar in its general nature but
are much more uneven, with scattered boulders, channels, and highly variable in its particulars. Biologists studying a certain
gullies. Zonation may not be as obvious in such areas, but it area sometimes name the zones after the dominant organism,
usually exists nonetheless. as in the limpet zone or the mussel zone. These zones actu-
Marine biologists have spent a lot of time and energy studying ally contain a variety of organisms. The mussel zone, for exam-
the causes of vertical zonation. Though there is still much to be ple, is also home to worms, crabs, snails, seaweeds, and many
learned, we now know that zonation results from the complex other species. Also, the specific organisms found in the inter-
interaction of physical and biological factors. A general rule is that tidal vary from place to place. On the east coast of North
the upper limit at which a species occurs is usually determined America, for example, the mussel zone is often replaced by a
rockweed zone.
Instead of naming the zones after organisms, we
FIGURE 11.18 Intertidal organisms often form distinct bands, or zones, at different heights will simply divide the intertidal into the upper, middle,
on the shore.
and lower zones. While not especially informative or
creative, at least these names apply everywhere! Bear
in mind that the following sections are general descrip-
tions of typical intertidal communities and dont
apply exactly to any particular place. Furthermore, the
boundaries between zones are not absolute and some
species occur in more than one zone. The next time
you go to the shore, think about how these descrip-
tions apply. Perhaps more importantly, think about
how the place you visit is different. What might cause
the differences?
In many places, lichens (Verrucaria) form black, tar-like so abundant that the upper intertidal is often called the Littorina
blotches on rocks in the upper intertidal. The fungus part of the zone. Not all of the many species of periwinkles live high in
lichen soaks up water, storing it for long dry periods. Dark green the intertidal, but those that do are well adapted to it. They
mats of cyanobacteria (Calothrix) are abundant. They are pro- can breathe air like land snails, live out of water for months,
tected from drying out by a jelly-like coating and have the and tolerate extreme temperatures. They do have limits, how-
advantage of being able to fix nitrogen from the air. Small tufts ever, and may have to seek shelter on hot days (see Fig. 11.2).
of a filamentous green alga (Ulothrix) that is also resistant to Limpets (Acmaea, Lottia) may also be found in the high
drying out may spring up here and there. Other green (Prasiola, intertidal. Like periwinkles, limpets are hardy grazers. Shore
Enteromorpha), brown (Pelvetia), and red (Porphyra, Bangia) crabs occasionally venture into the upper intertidal, scurrying
algae are occasionally found in the upper intertidal, usually in over the rocks. They mostly scrape algae off the rocks with their
moist spots. claws, but they may also eat animal matter, dead or alive. Sea
Large numbers of periwinkles graze on the algae, scraping lice or sea roaches (Ligia; see Fig. 7.33) breathe air and live above
it from the rocks with their radulas. Periwinkles (Littorina) are the waters edge, moving into the upper intertidal at low tide.
Upper intertidal
Periwinkles dominant; limpets, Most of the upper intertidal zone lies above
lichens, encrusting algae also common the high tide line and is kept wet by wave
splash. The dominant primary producers are
lichens and cyanobacteria. Periwinkles are the
Barnacles dominant most common animals.
Middle
intertidal
Mussels,
barnacles, Few marine predators can reach the
seaweeds upper intertidal. Shore crabs occasionally eat
periwinkles and limpets. From time to time
Lower intertidal predatory snails, such as unicorn shells
Seaweeds,
surf grass (Acanthina), may also venture into the splash
zone. On the other hand, the upper intertidal
is often visited by predators from land. Birds
such as oyster-catchers (Haematopus) may eat
large numbers of limpets and snails. Raccoons,
(a) Pacific
rats, and other land animals are also known
to enjoy these tasty molluscs, as do people in
most of the world.
Lower intertidal
Irish moss Lichens Close, mutualistic associations between
fungi and microscopic algae; mutualism is a type of
Extreme lower symbiosis.
intertidal
Seaweeds Chapter 5, p. 99; and Chapter 10, p. 217
A
general principle in the intertidal zone If organisms can live lower in the intertidal, It may be time-consuming, difficult, or down-
is that physical factors, especially desic- why dont they? Experiments suggest that other right impossible to remove all the organisms of
cation, usually determine how high in organisms, either competitors or predators, interest from an area, or they may move back
the intertidal a particular organism can live. often set a species lower boundary. In removal into an area too soon after removal. Instead,
Biological factors such as competition and pre- experiments, one species is removed from an caging experiments can be used to keep them
dation set the lower limit. Where does this area, which is then compared to an untouched out. On the Channel Islands off the coast of
principle come from? How can you tell why a control area. A classic experiment examined Southern California, the intertidal is covered by
certain species of barnacle or seaweed is found competition between rock barnacles (Semibalanus a nearly unbroken carpet of seaweeds. When
where it is? balanoides) and little gray barnacles (Chthamalus marine biologists discovered that large numbers
Like many scientific questions, this one is stellatus), which normally live higher on the shore of spiny lobsters (Panulirus interruptus) moved into
best answered by experimentaltering natural (see Fig. 11.20). Rocks with newly settled juve- the intertidal during nighttime high tides, they
conditions to see what happens. In the inter- niles of both species were collected and placed wondered what effect the lobsters were having.
tidal, three general types of experiments have at different heights. The rock barnacles were They installed plastic cages to keep the lobsters
been particularly effective. In one of these, removed from some of the rocks, while other out. Just to make sure that the seaweed wasnt
organisms are transplanted from one place to rocks were not disturbed. affected by the shade of the cages, they also
another to see whether they can survive under High in the intertidal, little gray barnacles installed control cages that had roofs but no ends,
the new conditions. In one such experiment survived better than rock barnacles. They appar- allowing lobsters to enter unimpeded. Seaweeds
barnacles were grown on panels that could be ently tolerate drying better. Lower in the inter- continued to flourish in these control areas and
placed at various heights on intertidal pier pil- tidal, where the rock barnacles can survive, little in adjacent open areas, but within the fully
ings. The barnacles soon died if they were gray barnacles cant compete. They do quite well enclosed cages the seaweeds were soon replaced
moved higher than they normally lived. as long as the rock barnacles are removed, but by mussels. This demonstrated that seaweeds
Measurements of their body fluids showed that when the two species grow together the bigger, were able to dominate because the lobsters ate
they had lost a lot of water, so the cause of faster-growing rock barnacles crush, undercut, the mussels, which are superior competitors.
death was probably drying out at low tide. The or smother the little gray barnacles. Thus, the Caging experiments dont have to use actual
experiment showed that the barnacles cannot lower limit of little gray barnacles is determined cages, just some sort of barrier to keep the ani-
survive the conditions above their normal by a biological factor: competition. mals out. Plastic rings or strips of artificial turf, for
range. Thus, their upper limit is set by a physical Removal experiments have also shown the example, will keep out sea stars, limpets, and
factordesiccation. importance of grazing and predation. When lim- snails, which cant crawl over the turf. Cages can
Transplantation can also tell us when physi- pets are removed from test areas, for example, also be used to hold animals inside an area, rather
cal factors are not important. When barnacles a profusion of seaweeds usually results. While than keep them out, to determine their effects.
are transplanted lower in the intertidal than they limpets are present the algae cannot establish Transplantation, removal, and caging exper-
normally live, they do just fine. The same is true themselves because limpets scrape young algae iments have been extraordinarily effective in
of many other intertidal species, including certain off the rocks. If the seaweeds get a foothold, unraveling the web of physical and biological
mussels, snails, and seaweeds. In fact, some spe- however, they are often able to persist because interactions in the intertidal zoneso effective,
cies actually grow faster when moved below adult seaweeds can stand up to these grazers. in fact, that they have become an important
their normal range. Their lower limit, therefore, Another removal experiment, involving a sea tool for ecologists not only in the intertidal but
is not set by the physical environment. star, is summarized in Figure 11.22. in many other environments.
The mussels in this dark patch have taken over the space inside a closed cage,
These cages were used in California to test the effects of spiny lobsters which was removed for the photo. The surrounding rock is dominated by red
(Panulirus interruptus) in the intertidal zone. The lower cages have closed ends algae, as are areas inside open-ended cages, not visible in this photo. The con-
to keep lobsters out. The upper cages have open ends to allow lobsters free clusion of the experiment is that mussels would dominate the community if not
access and are controls for the effects of the cages themselves. eaten by lobsters.
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the tides on a regular basis. A diurnal tide exposes the organ- The lower limit of rock barnacles, like that of little gray
isms once a day, a semidiurnal tide twice. If the tide is mixed, barnacles, is determined by biological factors, in this case both
the lower of the two successive high tides may not submerge predation and competition. Whelks are major predators of bar-
the high part of the zone, and the higher of the two lows may nacles (Fig. 11.21). These snails use their radula to drill holes
not expose the bottom part. Thus, even in places with a semi- in barnacles shells. They also secrete chemicals that soften the
diurnal tide, parts of the middle intertidal zone may be sub- shells. If protected from the snails by wire cages, rock barnacles
merged or exposed only once a day. With so much variation in flourish below their normal limit. Unprotected barnacles, how-
emersion time, different heights within the middle intertidal ever, suffer heavy losses to snail predation. The only thing that
often support different organisms. In other words, the middle saves the rock barnacles is that the dog whelks do most of their
intertidal is often made up of several separate vertical zones. feeding during high tide. When emersed they lose a lot of
The upper boundary of the middle intertidal is almost water if they move around hunting barnacles, so to avoid des-
always marked by a band of acorn barnacles. In many places at iccation at low tide they cease feeding or move lower in the
least two species of barnacles, such as little gray (Chthamalus) intertidal. In the upper intertidal, the snails dont have enough
and rock barnacles (Balanus, Semibalanus), form distinct bands, time at high tide to eat the barnacles. Sea stars, which also eat
with little gray barnacles living higher up. barnacles, have the same problem as the snails. Rock barnacles
Experiments (see Transplantation, Removal, and Caging may also face competition from mussels. In some places, in
Experiments, p. 252) show that this zonation results from a fact, the lower limit of barnacles is set not by dog whelk
combination of the barnacles larval settlement pattern, desicca-
tion tolerance, competition, and predation (Fig. 11.20). The lar-
vae of both species settle over a wider range than is occupied by Semidiurnal Tide A tidal pattern with two high and two low tides each
adults. The upper limit of both species appears to result from day.
emersion, because larvae that settle too high in the intertidal die. Chapter 3, p. 61; Figure 3.34
Little gray barnacles tolerate drying better than rock barnacles
Diurnal Tide A tidal pattern with one high and one low tide each day.
and can therefore live higher up. At lower levels, where the rock
Mixed Tide A tidal pattern in which two successive high tides are of
barnacles can survive, they outcompete the little gray barnacles.
different heights.
Adult little gray barnacles, then, live perched in a narrow band
Chapter 3, p. 61; Figure 3.34
above their competitors where they can survive emersion.
Rock barnacles
Hig
wa h
ter
Desiccation
Larvae Competition
settle from rock
barnacles Hi
wa gh
ter
Little gray barnacles
Mortality of
juveniles
Hig
hw
Predation by dog ate
whelks and competition r
from mussels
Distribution
of adults
Rock barnacles
FIGURE 11.20 Larvae of both little gray barnacles (Chthamalus stellatus) and rock barnacles (Semibalanus balanoides) settle out of the plankton over a wider
range of the intertidal zone than is occupied by adults. At the upper edge, young barnacles dry out and die. Little gray barnacles are more resistant so they can
live higher on the shore. Little gray barnacles that settle where the rock barnacles can survive, however, are outcompeted. The lower edge of the rock barnacle
zone occurs where predatory snails, called dog whelks (Nucella lapillus), can easily reach them. In some places, competition from mussels also helps determine the
lower limit.
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Sea stars
Gooseneck
barnacles
All mussels Lower
eaten edge of Lower
mussel intertidal
zone
extended
Rockweeds
Natural state Sea stars removed Results
FIGURE 11.22 Removing predatory sea starsor keeping them out with cagesshows how they affect the middle intertidal community. Below a certain level
the sea stars can easily reach the mussels, their favorite food, and eat them all. Above this level the sea stars cannot eat all the mussels, but they eat enough to make
space available for other species. When there are no sea stars around, the mussels are able to live lower, and they monopolize the available space by overgrowing
and crowding out other species. Sea stars thus maintain the diversity of the middle intertidal community.
communities: Just because a species is relatively uncommon chitons may remove newly settled larvae and spores. If grazers
does not mean it is unimportant. Predators whose effects on move into a patch early on, the succession may never get past
their communities are proportionately much greater than their the bacterial and algal film stage and in some places grazers
abundance are known as keystone predators. limit even the film. Predators and other disturbances help deter-
Natural disturbances can have effects similar to those of mine whether the final stage is a solid mussel bed or a mixture
predation. When mussel beds get too dense, clumps of mussels of species.
can be torn away by waves (see Fig. 11.14), exposing the bare Furthermore, steps in the succession may be skipped. For
rock. Drifting logs that batter the rocks and, in cold places, example, seaweeds may never be able to colonize a patch if
scouring ice and severe freezes, have the same effect of opening barnacle larvae get there first (see Fig. 11.16). The time of
up new space. This prevents mussels from monopolizing the year that the patch is created is important. If the patch opens
available space, allowing other organisms to persist. when barnacle larvae are abundant and seaweed spores are rare,
When a patch of space is cleared, new organisms often barnacles have the upper hand. There is thus an element of
move into the patch and get replaced by others in a regular chance in the development of the community. Stages may
sequence (Fig. 11.23). The term ecological succession is used also be skipped if the bare patch is small. A small patch cleared
to refer to such regular patterns of regrowth. In the rocky in the middle of a mussel bed may be taken over by other
intertidal the first stage is usually a thin film of bacteria and mussels that move in from the sides before the succession
microscopic algae, such as diatoms, that covers the rock. This gets started.
film may condition the rock surface, since the settling larvae of Without predation and other disturbances, the best com-
some species prefer surfaces with the film over bare rock. petitors take over and many species disappear from the area.
Seaweeds often move in next, followed by barnacles and finally Disturbance can prevent this. Mussels, for instance, are often
mussels, the dominant competitor. The final stage in an eco- unable to completely cover middle intertidal rocks because sea
logical succession is called the climax community. stars eat them or waves tear them away. This gives other spe-
cies a chance. Disturbance can thus increase the number of
species that live in an area, that is, the diversity, by interfering
The typical steps of ecological succession in the middle intertidal with competitive exclusion. The rocky intertidal in some
of many rocky shores are, first, a bacterial and algal film, then
seaweeds, barnacles, and finally the climax mussel community.
Bare rock
Seaweeds
PROGRESSION OF SUCCESSION
Barnacles
get there
first Barnacles
Climax
community
FIGURE 11.23 Ecological succession following the clearing of a patch in a mussel bed. The pathway taken and the end result depend on the size of the patch,
when it opens up, and just plain luckwhich organisms get there first.
places can be thought of as a mosaic of patches that were from taking over, but not so much that the community is
cleared at different times and are in different stages of succes- unable to develop (Fig. 11.24).
sion. Each patch supports a different set of organisms, so the
number of species in the area as a whole increases. On the
In the absence of disturbance a few dominant competitors,
other hand, if predation and other disturbances happen too especially mussels, take over the space in the middle intertidal
often, the community keeps getting knocked back to the start- zone. Moderate predation and other disturbance prevents this and
ing point. It never gets a chance to develop, and not many spe- allows other species to live there. Too much disturbance,
cies are able to live there. The highest diversity occurs when however, removes most species.
there is enough disturbance to prevent the dominant competitors
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The Lower Intertidal The lower inter- The story doesnt end there. If the
FIGURE 11.25 Biological interactions determine what type of seaweed predominates in New England tide pools. Unless something interferes, both types of pool
are self-sustaining.
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T
o understand intertidal ecosystems we
need to consider different scales. Intensive,
localized studies are needed to dissect out
the detail of how individual organisms interact,
while studies of an entire bay or a coastline
provide a broader perspective. Examining
links between the intertidal and coastal eco-
systems offshore provides an even bigger pic-
ture. Time scale is critical, too: A one- or
two-year study cant detect long-term changes
in a community, and what seems like a major
shift can just be natural variation. Similarly,
while studies of single factors, sea star preda-
tion for example (see Fig. 11.22), reveal much
about intertidal ecology, we also need to
understand how different factors interact
for example, how sea star predation, mussel
and algal growth rates, and larval settlement
might affect competition between mussels
and seaweeds.
Studies that combine large spatial and tem-
poral scales and multiple factors require time,
expertise, and funding that are usually beyond
the reach of individual investigators or
research teams, so marine ecologists are
increasingly joining forces. The Partnership for Robomussel is an electronic temperature recorder that not only looks but also absorbs heat like real mussels.
Interdisciplinary Studies of Coastal Oceans
(PISCO) is a consortium of marine scientists previously thought, and that coastal upwelling To learn how climate change might
doing integrated studies at many sites over (see Upwelling and Productivity, p. 351) directly affect intertidal communities, PISCO
many years along 2,000 km (1,250 mi) of the influences predation, grazing, and other pro- scientists placed Robomussels in mussel
Pacific coast of the United States. Research on cesses in the rocky intertidal. beds from Washington to Southern California
oceanography, molecular biology, genetics, In the summer of 2002 PISCO scientists to determine the effects of splashing waves,
physiology, and ecology provides the most discovered an unprecedented hypoxic (low- tide, and latitude on the mussels body tem-
comprehensive view ever of the nearshore oxygen) dead zone near the Oregon coast, peratures. They discovered hot spots along
ecology of this coastline. which has since recurred every summer, pro- the entire coast where mussel temperatures
In the rocky intertidal, PISCO scientists up gressively getting larger and lasting longer. were higher than would be expected based
and down the coast regularly measure the Unlike hypoxic zones caused by nutrient just on how far north they were. It was previ-
abundance, growth, and larval settlement rates overfertilization and decay of the resulting ously assumed that the impacts of global
of major species, study the intensity and effects algal growth this one forms when upwelling warming will mainly occur on the edges of
of predation and competition, and monitor currents carry water from the naturally occur- species ranges, as the southern end gets too
physical factors such as temperature and ring oxygen minimum zone (see The Oxygen hot and the northern end warms up, but this
salinity. All studies use the same sites and are Minimum Zone, p. 367) onto the shallow research indicates that mussels could die off
linked to research on the oceanography and nearshore shelf. Scientists suspect a large- in patches all along the coast, likely affecting
biology of adjacent nearshore waters. PISCO scale change in oceanographic conditions, the entire community.
research has already revealed that there is perhaps related to climate change, but
much more spatial variation in larval settle- research to understand this phenomenon For more information, explore the links provided on
ment and individual species distributions than continues. the Marine Biology Online Learning Center.
A host of small animals live among the seaweeds, hiding from worms (Spirorbis, Phragmatopoma), snails (Tegula, Nucella), sea
predators and staying moist during very low tides. Sea urchins slugs (Aplysia, Dendronotus), and many others.
(Strongylocentrotus, Echinometra; see Fig. 13.19) are common Most rocky intertidal fishes live in the lower intertidal or in
grazers on the seaweeds. Also present in the lower intertidal are tide pools. Gobies, clingfishes, sculpins (Oligocottus), pricklebacks
sea anemones (Metridium, Anthopleura; Fig. 11.26), polychaete (Cebidichthys), and gunnels (Xererpes) are among the most common.
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a wide range of particle sizes, from large pebbles Well-sorted (coarse) Well-sorted (fine) Poorly sorted
down to clay. Put it in a container with some
water and shake it up. As soon as you stop shak-
ing, the pebbles will sink to the bottom. As the
water gradually stops moving, first the coarse sand
and then finer particles will begin to sink. Very
fine material will remain suspended for a long
time, making the water look muddy. You might
have to let the container sit for weeks before the
finest clay settles out completely. If the container (a) Water drains quickly (b) Water drains slowly (c) Water blocked
is stirred even gently, the fine material will never
settle out. FIGURE 11.28 Sorting refers to the uniformity of grain size in the sediment. In well-sorted
sediments (a and b) the grains are all about the same size. There is a lot of space between the grains,
This demonstrates a general rule: Fine sedi- so water can flow through. The grain size, as well as sorting, affects how well water circulates; larger
ments remain suspended with even a small amount grains have larger spaces and are thus more porous. Poorly sorted sediments (c) have many different-
sized grains. Small grains fill in the spaces between the larger ones, making water flow difficult.
of water motion, whereas coarse sediments settle
out unless there is considerable flow. Bottom sed-
iments thus reflect prevailing conditions. Calm, sheltered areas Fig. 13.12). Coarse sands contain very little organic matter.
have muddy bottoms because fine sediments can settle out. That is why we tend to think of sand as clean. Silt and clay,
Particles of organic matter sink at about the same rate as clay on the other hand, are usually rich in detritus, which is why
particles, so the two tend to settle out together; clay sediments they are often smelly.
are therefore organic-rich. Places with stronger waves and cur- The grain size also affects the amount of available oxygen in
rents have coarser bottom sediments. If the water motion is the sediments. Oxygen in sediments is used up by the respiration
strong enough, it may carry away all the loose material, leaving of animals and, more importantly, decay bacteria. Below the
bare rock or large boulders. sediment surface there is no light and therefore no photosynthe-
sis, so the infauna depend on the circulation of water through
Fine sediments are found in calm areas such as bays and lagoons. the sediments to replenish the oxygen supply. Grain size and the
Coarser sediments are found in areas affected by waves and currents. degree of sorting strongly affect how porous a sediment is
(Fig. 11.28). Every backyard gardener knows that water flows
In our sediment experiment, all the particles would eventu- more easily through sand than clay. Water circulation through
ally settle out if the container were kept still. A constant flow of fine sediments is greatly restricted.
water through the container, however, Muddy bottoms, then, have a double
would carry away the fine material and only problem. First, they have more organic
certain particles, the coarse ones, would material to decay and use up oxygen.
remain. The coarse particles would be Second, the fl ow of water that brings in
sorted out from the fine ones. new oxygen is reduced. Except for the
upper few centimeters of mud the intersti-
tial water, or water between the grains, is
Living in the Sediment deficient in oxygen. If you go very far down
Living in sediment has advantages in the at all, in fact, the oxygen is completely used
intertidal. Soft bottoms stay wet after the up. Sediments with absolutely no oxygen
tide is out, so the problem of desiccation is are said to be anoxic.
not as critical as in the rocky intertidal. Anoxic conditions are no problem at
This depends on the grain size, however. all for bacteria that are capable of anaero-
Coarse sands drain, and therefore dry out, bic respiration, in which they break down
quite rapidly (Fig. 11.28). Partially because organic matter without oxygen. A noxious
of this, coarse sand beaches have relatively gas called hydrogen sulfi de (H2S) is pro-
little animal life. duced as a by-product (see Table 5.1). If
you have ever had to unclog the drain of
Oxygen Availability The amount of your kitchen sink, you have probably run
organic matter in bottom sediments is into hydrogen sulfide. It turns things black
particularly important to deposit feeders. and smells like rotten eggs. In the sedi-
Because there are relatively few primary ments, just as in your drain, hydrogen
producers, detritus is the main source of sulfide is produced when bacteria continue
food for intertidal soft-bottom communi- to decompose organic material after all
ties. Deposit feeders extract this organic FIGURE 11.29 The black layer in the sedi- the oxygen has been used up. A distinct
ments shows where the oxygen has been completely
matter from the sediments. The amount depleted and hydrogen sulfide (H2S) is produced by black layer appears where anoxic condi-
of detritus depends on grain size (see anaerobic bacteria. tions begin in the sediments (Fig. 11.29).
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FIGURE 11.30 Burrowing in clams and cockles. The foot does all the work. The clam first pushes its foot down through the sediment. Then the clam expands
the end of the foot, which anchors the clam as it pulls its body down.
meiofauna, but most have independently The bent-nosed clam ( Macoma; see
evolved a worm-like shape (see Life in Fig. 12.11) also concentrates on the upper
Mud and Sand, p. 291). layer of sediment. It uses a long siphon
to suck up food particles from the sur-
Feeding As already noted, detritus is the face. Other animals catch particles as
main source of food in the soft-bottom they settle out and are thus on the line
intertidal (Fig. 11.33). Diatoms sometimes between suspension feeding and deposit
form highly productive mats on the sedi- feeding. Some polychaete worms
ment surface, but most of the time they (Terebella) have long, sticky tentacles that
dont account for much primary produc- they spread out on the bottom to gather
tion. In any case, most animals dont dis- FIGURE 11.32 After digesting out the organic food (see Fig. 13.11). Other polychaetes
tinguish between the diatoms and detritus. matter, this tropical sea cucumber (Holothuria edulis) produce a mucus net that they use instead
Plankton brought in by the tides also make has deposited a pile of undigested sand pellets. of tentacles.
a small contribution to the food supply. There are also suspension feeders,
Many different methods of deposit feeding have evolved. which dont wait for the detritus to settle. An olive shell
One of the more common methods has already been described: (Olivella) from Central America makes a mucus net, but instead
Animals such as sea cucumbers and various worms take in of spreading the net on the bottom the snail holds the net up
sediment as they burrow, digest the detritus and small organ- in the water to filter food. Sand, or mole, crabs hold a pair of
isms, and leave the rest of the sediment in their wake. This large, bushy antennae up into the water to trap food. By quickly
technique is more common on mud bottoms than sand, prob- burying and uncovering themselves, they combine feeding and
ably because mud contains more organic matter. Sand is also locomotion. When the tide is coming in and they want to
abrasive and hard on the digestive system. move up the beach, they uncover just before a wave hits, ride
Sand dollars (Mellita, Dendraster; see Figs. 7.47 and 13.8) the wave up, and raise their antennae. Then they burrow to
are more selective. They use their tube feet to pick up organic keep from being washed back down. As the tide recedes, they
particles. Sand dollars burrow near the sediment surface, where do the opposite, burrowing before the wave arrives, then pop-
detritus is still accumulating and is therefore more abundant. ping out to ride the backwash down the beach.
Soft bottoms also have their share of
predators. Moon snails (Polinices) burrow
through the upper sediments looking for
Carnivores Herbivory
clams. When a snail finds a clam, it drills
Fishes, Predation
a hole in the clams shell and eats it.
birds, Detritus feeding
moon snails or export Several polychaetes and other worms are
Contribution to also important predators. Birds can be
detritus pool major predators during low tide (see
Fig. 12.12). At high tide, fishes can come
in. Fishes often dont eat an entire animal.
Instead, they just nip off clam siphons or
Suspension feeders Deposit feeders
other bits that stick out.
Consumers Clams, Polychaetes,
sand crabs, clams
Zonation Because the organisms live in
polychaetes the sediments and cant be seen, zonation
on intertidal soft bottoms is not as obvi-
ous as in the rocky intertidal. Zonation
does exist, however, especially on sandy
beaches (Fig. 11.34). Water drains rap-
From other idly from the sand, and because the beach
communities slopes, the upper part is drier than the
Detritus (rocky intertidal,
estuaries, lower part.
subtidal) The upper beach is inhabited by
beach hoppers, or sand fleas, which are
really amphipods, and by isopods. In
Producers warmer areas, these small crustaceans are
Plankton
(from other communities)
Bottom diatoms replaced by ghost (Ocypode) and fiddler
crabs (Uca; see Fiddler on the Mud,
p. 273) that scurry about catching smaller
FIGURE 11.33 A generalized food web on a sandy beach. animals, scavenging bits of dead meat,
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e
h tid
Hig
Ghost crab
Amphipod Inte
rtid
al
Soft-shelled
clam
ide
wt
Lo
Lugworm Sub
tida
Isopod l
Bristleworm
Sand dollar
Moon snail
Coquina
shell
Blue crab
Sea cucumber
FIGURE 11.34 Generalized zonation pattern on sandy beaches of the Atlantic coast of North America. Sheltered and exposed beaches have different com-
munities, and the animals shown here dont usually all occur on the same beach.