BIOLS380

CH11: Between the tides

Done by: Yousif Adel
 Intertidal zone
-Intertidal zone: the area between the mean low tide & mean high tide
-Subtidal zone: the area that is always submerged
-The intertidal zone will be exposed during low tide

Intertidal substrates
-Substrate (type of bottom):
 Intertidal zone substrate can be rocky or soft bottom (sand or mud)
 Soft bottoms can vary from sand to silt or mixture (mud)
 Rocky substrate can vary as to the type of rock and the slope angle

Intertidal zone organisms

1. Epifauna: organisms that live on the surface of the substrate (ex: mud
snails on soft bottoms, barnacles on hard substrates)
2. Infauna: organisms that live in the substrate (ex: clams burrowing in
soft bottoms)
3. Meiofauna: organisms that are so small that they live between the
grains of soft substrate; these organisms can be sessile or motile

Challenges associated with the intertidal

(Study them from the book)
-Due to the exposure to air, organisms face a variety of challenges,
including:
 Desiccation (water loss) - Temperature & salinity changes
- Interrupted feeding - Wave action and tides - Limited space -
Oxygen availability & build-up of CO2 at low tide

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 1. Desiccation (water loss)
-When exposed, organisms must deal with potential water loss
-Water loss is more pronounced on hot, dry or windy days
-Organisms can deal with potential water loss by hiding or clamming up
-Hiding may involve moving to a tide pool or an area with more
moisture (motile organisms)
-Hiding may also mean that some organisms only live in areas where
moisture will remain when the tide is out (such as crevices in rocks or
shallow spots in soft bottoms)
-Clamming up: closing shells or walling itself off from the environment
to conserve moisture (such as an oyster closing its shell or a snail
walling itself off using its operculum)
-While moisture can be conserved in this way, no gas exchange or
feeding
-Periwinkles (Littorania) clamp themselves to rocks to seal in moisture
-They can seal off the opening of their shell by closing their operculum
-Some other organisms can tolerate drying out
-Some intertidal chitons can survive the loss of 75% of the water in their
tissues
-Some seaweeds, such as rockweeds (Fucus) can withstand a water loss
of as much as 90%. They quickly recover when the tide comes in and
wets their tissues

2. Temperature
-Tide pool fishes are much more tolerant of extreme temperatures than
related fishes that live below the tides
-Certain species of periwinkles from the high in the intertidal show
remarkable heat tolerance. One known to survive temperatures as high as 49˚C
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-Some snails (especially in the tropics) have ridges on their shells. These
ridges help the snail lose excess heat
-The shell ridges on the snail help it stay cool by radiating heat. The
white color helps by reflecting sunlight
-The color of the snail’s shell can help it tolerate high temperatures
-Snails subjected to high temperatures tend to be light in color

3. Salinity
-Salinity can change dramatically due to temperatures (evaporation in
tide pools) or weather (rain)
-Normal ocean salinity is around 33-35 0/00. Estuarine salinity normally
varies between 5-30 0/00
-Organisms in the intertidal are euryhaline (tolerate a wide variety of
salinities) & subtidal organisms are stenohaline (tolerate very low
variations in salinities)
-Salinity may begin at 20-25 0/00 and climb dramatically during the day
due to water loss when the intertidal is exposed
-Salinity may also drop during the day due to a sudden influx of fresh
water provided by a passing thunderstorm
-A salinity change of possibly 20 0/00 or more combined with a
temperature change of 40 degrees or more may occur

4. Interrupted feeding
 Feeding will stop:
1. If an organism clams up
2. For filter feeders when exposed to air
-A few organisms (such as snails) can still feed during low tide by
scraping algae off surfaces
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-For communities with semidiurnal tides, an animal could spend nearly
half of their day exposed and not feeding
-Some intertidal organisms feed on pieces of seaweed and other large
detritus that drift from deeper water
-Some are predators and move over the rocks in search for prey
-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

5. Wave action
-Wave exposure can be a problem for intertidal organisms
-Rocky shores are often exposed to significant wave action
-The impact of waves varies along the shoreline
-Some areas are sheltered from the surf, others are fully exposed
-Exposure to waves strongly affects intertidal organisms
-Coping with wave shock:
-Some organisms can’t withstand wave shock and are found only in
sheltered locations
-Organisms exposed to the waves need some way to deal with wave
shock
-Sessile organisms anchor themselves firmly to the rocks to keep from
being washed away
-Seaweeds used holdfast
-Barnacles secure themselves with a glue
-Mussels attach themselves with their byssal threads
-Mobile animals cling strongly to the rocks
-Limpets & chitons use their muscular foot like a powerful suction cup

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-Some fishes have also suction cup, formed by modified pelvic fins
-Intertidal fishes tend to lack swim bladders, so they sink and stay on the
bottom
-Animals in exposed locations tend to have thicker shells
-A compact shape can help reduce the impact of the waves (e.g., sea
anemone)

6. Oxygen availability
 Oxygen can be exhausted:
1.If an organism clams up during low tide
2. In tide pools if there is a high density of organisms (particularly
algae) at night; no photosynthesis
-Carbon dioxide can also build up
-Some organisms have evolved the ability to exchange gases in both air
and water

7. Limited space
 In some intertidal communities, space may be limited. This is
particularly true in:
1. The rocky intertidal, where surface area is limited
2. In soft-bottomed communities, where many organisms select
hard substrates (such as oyster reefs) to prevent being washed
away
-Competition for space is a dominant ecological factor in the rocky
intertidal
-There are many ways to compete for space. One is to be the first to get
to open spots

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-For intertidal organisms, this means having an effective means of
dispersal
-Most rocky intertidal species disperse via larvae or spores, which settle
on the rocks to colonize open space
-Some other intertidal organisms take over space that is already
occupied
-Barnacles undercut their neighbors; loosening them from the rock
-Many intertidal organisms simply grow over their competitors, making
them vulnerable to waves
-Some species grow in colonies, gradually increasing the amount of
space they hold as the colony grows

Intertidal areas
1. Rocky
2. Soft bottom
2.1) Sandy shores
2.1) Mud flats

Vertical zonation of rocky shores

-The rocky intertidal community is usually divided into distinct bands or
zones at characteristic heights in the intertidal
-Zonation means that a given species is usually found only within a
particular vertical range and not throughout the intertidal (Vertical
zonation)
-Zonation is the result of complex interaction of physical & biological
factors

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  The upper limit of a zone is often set by physical factors
 The lower limit of a zone is often set by biological factors
-Intertidal area can be divided into upper, middle & lower zones
-The boundaries between zones are not absolute and some species occur
in more than one zone
 The upper intertidal:
-Rarely submerged, the inhabitants must be well adapted to withstand
exposure to air
-This zone lies mostly above the high tide mark (Splash zone)
-In many places, lichens form black tar-like spots on rocks in upper
intertidal
-Dark green mats of cyanobacteria (Calothrix) are abundant, they are
protected by a jelly-like coating and able to fix nitrogen
-Filamentous green algae (Ulothrix); resistant to drying out may bloom
here
-Other green, brown & red algae are occasionally found in the upper
intertidal (in moist spots)
-Large number of periwinkle graze on the algae
-The periwinkle (Littorina) is so abundant
-Limpets live in the high intertidal; They are grazers
-Shore crab can scrape algae off the rocks and eat animals and detritus
-Sea lice or sea roaches breathe air and live above the intertidal but
move down into the upper intertidal at low tide
 Middle intertidal:
-The middle intertidal is regularly submerged & uncovered by the tides
-Diurnal tide exposes organisms once a day & a semidiurnal tide twice

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-With the variation in emersion time, different heights within the middle
intertidal zone often support different organisms
-The middle intertidal is often made up of several separate vertical zones
-The upper boundary of the middle intertidal is almost always marked by
a band of acorn barnacles

Zonation in the intertidal

-Zonation due to varying
degrees of exposure;
upper intertidal (most
exposed) to middle and
lower intertidal (least
exposed)

 Why competition will be greater in the lower intertidal?

 Because it is the least extreme of the zones (it is exposed to a lesser
degree than the upper intertidal)
-The lower intertidal always has more species for the same reason
(although species will vary greatly by location)

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 Zonation
-Biological interactions (competition for space, predation, grazing) are
often important factors in determining lower limit of rocky intertidal
organisms; physical factors (desiccation) in determining upper limit

-Larvae of both little gray barnacles and rock barnacles settle out of the
plankton over a wide 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 live higher on the shore
-Little gray barnacles that settle where the rock barnacles can survive;
outcompeted
-The lower edge of the rock barnacle zone occurs where predatory snails
can easily reach them
-In some places, competition from mussels helps determine the lower
limit

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 Middle intertidal
-Depending on many physical & biological factors, mussels, gooseneck
barnacles and brown seaweeds often dominate the middle intertidal
below the barnacles
-Many rockweeds have air bladders (pneumatocysts)
-Mussels are especially common on stormy, exposed shores
 Why mussels can’t live as high on the shore as barnacles?
 Because they dry out and at high levels, they don’t spend enough
time underwater to filter feed
-Sea stars are voracious mussel predators
-Sea stars can’t tolerate desiccation very well and need to be underwater
to search for food
-Mussels are the dominant competitors for space on many rocky shores
 Their upper limit is set by: desiccation & filter feeding time
 Their lower limit is set by: predatory sea stars

Competition
-Some species are better competitors than others and will exclude others
if the community is left un-disturbed
-If a habitat is disturbed, organisms move in and are excluded in a
predictable pattern (ecological succession)
-The climax community (end result) will eventually form when the
community is undisturbed for a long period of time

Ecological succession
-In the rocky intertidal areas, the first stage is usually a thin film of
bacteria and microscopic algae (such as diatoms) that cover the rock

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-The film may condition the rock surface; since the settling larvae of
some species prefer surfaces with the film over bare rock
-Seaweeds often move in next, followed by barnacles and finally
mussels (the dominant competitor)
-The final stage in an ecological succession is the climax community
-The highest diversity occurs when there is enough disturbance to
prevent the dominant competitors from taking over

-Other things being equal, the number of species depends on how often
disturbance occur
-In the context, predation on the dominant competitors counts as a
disturbance
 Why number of species is highest when disturbance is moderate?
 Because when disturbance is rare, the dominant competitors take
over and exclude other species. Intermediate levels of disturbance
prevent this and give other species a chance
-If disturbance happens too often, most species cannot get a foothold and
the number of species drops

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 Keystone predators
-Keystone predators whose effects on their communities much greater
than their abundance

Predation
-Predation by keystone predators such as sea stars strongly affect
occurrence and density of animals in the middle intertidal

-Removing predatory sea stars or keeping them out with cages shows
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 can monopolize the available space by overgrowing and
crowding out other species
-Sea stars thus maintain the diversity of the middle intertidal community
-Page 260 (read very well)

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 The lower intertidal
-The lower intertidal is immersed most of the time
-This makes it easy for predators like sea stars to feed, so mussels and
barnacles are rare
-The lower intertidal is dominated by seaweeds, including species of red,
green and brown algae
-They can’t tolerate drying out but in the lower intertidal they grow in
abundance
-Grazing and competition are important in the lower intertidal
-Light & space is an important resource
-Seaweed often compete by overgrowing each other, blocking the light
-The lower intertidal supports many seaweeds, besides Enteromorpha
and Irish moss
-These range from delicate reds and green to large tough kelps
-The kelps mark the lower limit of the intertidal and continue down into
the subtidal zone
-There is no precise boundary between the lower reaches of the intertidal
and the upper edge of the subtidal
-Coralline algae, red algae that deposit calcium carbonate in their tissues
can be abundant
-Surf grass, a flowering plant is common in the lower intertidal on the
Pacific coast of North America
-A host of small animals live among the seaweeds, hiding from
predators & staying moist during very low tides
-Sea urchins are common grazers on the seaweeds
-Also present in the lower intertidal sea anemones, polychaete worms,
snails, sea slugs and many others

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-Most rocky intertidal fishes live in the lower intertidal or in tide pools
-Many fishes are present in the lower intertidal and are adapted to the
environmental extremes of the intertidal
-Most are carnivores

11.2: Substrate in the soft bottom intertidal

-Sediment size can vary between sand, silt, clay or mixture (mud)
-The smaller the sediment size, the less oxygen in the water filling
spaces

-The sediment size can be small so that all except the top few inches is
anoxic (without oxygen)
-Infauna must maintain a connection with the oxygenated zone via tubes,
siphons or extended burrows if they are located in the anoxic zone
-Anaerobic bacteria that break down organic matter thrive in the anoxic
zone
-Most animals are infauna, with adaptations to dig or burrow
-Poor epifauna
-The amount of organic matter is important to deposit feeders
-There are few primary producers
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-Detritus is the main source of food for intertidal soft bottom
communities
-The amount of detritus depends on grain size
-Coarse sand contain very little organic matter

Anoxic condition

-Muddy bottoms then a have double problem

-They have more organic material to decay and use up oxygen
-The flow of water that brings in new oxygen is reduced
-Sediments with absolutely no oxygen are said to be anoxic
-Anoxic condition is not a problem to anaerobic bacteria which can
break organic matter and hydrogen sulfide is produced
-Muddy sediments are often oxygen deficient
-Animals have adapted to low oxygen environment
-They have special hemoglobin and other adaptations to extract as much
as possible of what little oxygen there is in the interstitial water
-Some are sluggish, which reduced their oxygen consumption
-Some have symbiotic bacteria that help them live in low oxygen
sediments

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-Many animals pump oxygen rich water from the sediment surface with
siphons or through their burrows
-Their burrows and the effect of bioturbators (infauna that burrow
through) turn over or otherwise disturb the sediment (help oxygenate the
sediment)

Soft bottom intertidal communities

 The kind of sediment accumulates in an area depends on:
1. How much water motion there is
2. The source of the sediment
-The type of sediment strongly influences the community

The shifting sediments

-Soft bottoms are unstable & constantly shift in response to waves, tides
and currents
-Soft bottom organisms do not have a solid place for attachments
-Few seaweeds have adapted to this environment
-Seagrasses are the most common large primary producers on soft
bottoms and often form dense intertidal beds
-The animals that live on soft bottoms also lack solid attachments sites
-Few of them are epifauna (live on the sediment surface), most burrow
in the sediment for protection and to keep from being washed away
 Why the animals that burrow in the substrate are called infauna?
 Because they live in the sediment
-The kind of sediment on the bottom (especially the size of the grains) is
one of the most important factors affecting soft bottom communities
-Most people use the terms sand, silt, clay & mud

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-Those terms refer to sediments of particular grain sizes. Sand is
coarsest, followed by silt and then clay. Mud refers to silt and clay
together
-Most sediments are made up of a mixture of different particle sizes
-Sediment composition is directly related to the degree of water motion
 Why calm sheltered areas have muddy bottoms?
 Because fine sediment can settle out
-Organic particles sink at about the same rate as clay particles so the 2
tend to settle at about the same rate
-Clay sediments are normally organic rich
-Places with stronger waves and currents have coarser bottom sediments
-If the water motion is strong enough, it may carry away all the loose
materials, leaving bare rock or large rocks

Getting around
-Soft bottom animals have different methods to burrow through
sediments
-Clams and cockles can change the shape of their muscular foot
-Many soft body animals do something similar, using their body the way
clams use their foot

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-Burrowing in the lugworm. The worm expands the end of its body
(black arrows)
-The expanded end acts as an anchor and the rest of the body is pulled
along behind (White arrow)

-Heart urchin burrow through the sediment with their spines and tube
feet
-Many crustaceans such as amphipods use their jointed appendages to
dig
-Few deposit feeders eat their way through the sediment
-Sea cucumbers and certain worms can push the sediment through them.
They digest the organic matter and leave the rest of the sediment behind
-After digesting out the organic matter, the tropical cucumber has
deposited a pile of undigested sand pellets
-Some soft bottom animals are so small that instead of burrowing
through the sediment they live in the interstitial spaces between the
grains (meiofauna)

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 Feeding
-Detritus is the main source of food in the soft bottom intertidal
-Diatoms sometimes form highly productive mats on the sediment
surface, but most of the time they don’t account for much primary
production
-Plankton brought in by the tides also make a small contribution to the
food supply
-Sea cucumber / Sand dollar / Clams / Polychaete worms / Suspension
feeders / Olive shell / Moon snails / Predatory polychaete & worms /
Birds / fishes (Read from the book; how those organisms feed in soft
bottom Habitats, Page 267)

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 Zonation
 Why zonation on intertidal soft bottoms is not as obvious as in the
rocky intertidal?
 Because the organisms live in the sediments and can’t be seen

Soft bottom intertidal zonation

-The upper beach is inhabited by beach hoppers or sand fleas, which are
really amphipods and by isopods
 In warmer areas, these small crustaceans are replaced by ghost and
fiddler crabs that live by:
1. Catching smaller animals
2. Scavenging bits of dead meat
3. Gathering detritus
-Polychaetes, clams and others appear lower on the beach
-Zonation is even less obvious in muddy areas
-The bottom in such places is flat and fine sediments retain water
-The habitat therefore does not change very markedly between the high
and low water marks
-Intertidal communities on muddy bottoms are much like the subtidal ones
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Soft bottom intertidal food web

Rocky intertidal food web

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 Do it yourself summary
Read each sentence and identify the word/phrase that you believe
correctly completes the statement. Then check the answer key at the end
below to see if your answers are correct.
1. The …. is the material on which an organism lives.
2. On the …. coast of North America, rocky shores are common only
north of Cape Cod. They dominate on the …. coast and are practically
absent on the …. coast.
3. Animals that live on the substrate are called …. .Those that burrow
are called ….
4. Zonation in the rocky intertidal zone is related to the difference in ….
time at different heights.
5. Wave …. causes a wave to bend and become more parallel to the
shore. Wave energy is focused at ….
6. …. attach mussels to the rock. Molluscs such as …., use a muscular
foot.
7. …. usually is the most limiting resource for rocky intertidal
populations.
8. Typically, the most common inhabitants of the high intertidal zone are
….
9. Barnacles usually occupy the uppermost part of the …. intertidal.
Their upper limit is set by …. (Or ….); predation by …. (…., or ….)
largely determines their lower limit.
10. Sea stars are called …. because they play a central role in the
structure of rocky intertidal communities.
11. The occurrence of a regular pattern of recovery after a community is
disturbed is called …... Typical stages of this in the rocky intertidal are a
…. followed by seaweeds, then …., and finally …. (Or …., …. or ….).
The final stage in the process is called the ….

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12. The lower intertidal zone is dominated by…. . ….., as well as space,
is an important limiting resource here.
13. ….is a relatively coarse sediment and contains little organic matter.
It is found in …. (Or ….) areas. Silt and clay together are called ….,
which is fine and organic-rich. Muddy bottoms are found in …. (Or ….)
areas.
14. The most important food source for soft-bottom intertidal animals is
…. . As a result, most are …. feeders.

Answer key
1. Substrate
2. East / West / Gulf
3. Epifauna / Infauna
4. Emersion
5. Refraction / Headlands
6. Byssal threads / Limpets, chitons, etc.
7. Space
8. Lichens, cyanobacteria (blue-green algae) & periwinkles
9. Middle / Emersion / Drying out / Snails / Gastropods / Dog whelks
10. Keystone predators
11. Ecological succession / Bacterial or algal film / Barnacles / Mussels /
Mussels & Barnacles / Gooseneck barnacles / Climax community
12. Seaweeds / Light
13. Sand / Turbulent / Surf-swept / Mud / Calm / Protected
14. Detritus / Deposit

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