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296 Part Three Structure and Function of Marine Ecosystems
communities. As in the rocky intertidal (see Fig. 11.22),
they remove residents from the rocks, opening up space
for other organisms. Settling larvae and seaweed spores
colonize the patches cleared by grazers and carnivores.
(Laminaria longicruris)
Because these planktonic stages are often seasonal,
patches formed at different times may be colonized by
different species. This increases species diversity in a
given area. Carnivores may also act as a check on the
grazers, helping maintain a relatively stable situation.
This balance can be easily upset, however. For example,
explosive increases in sea urchin populations have been
reported in several subtidal communities (see p. 300),
as have mass mortalities. Like all biological communi-
ties, subtidal communities are constantly changing.
The species composition of rocky-bottom subtidal
communities is influenced by factors such as light,
competition for space, grazing, and predation.
Kelp Communities Among the most fascinating of
marine communities, kelp communities are also one FIGURE 13.21 Examples of North Atlantic kelps. The hollow-stemmed kelp reaches
of the most productive. Kelps are a group of large, fast- lengths of 12 m (40 ft) in deeper water.
growing brown seaweeds that live in relatively cold water
and are restricted to temperate and subpolar regions. (see
Brown Algae, p. 104). They are true giants compared with
other subtidal seaweeds or seagrasses, creating luxuriant forests
(see Fig. 6.8) that are home to a vast assortment of organisms.
There are several species of kelp. In the North Atlantic
and on the Asiatic coast of the North Pacific, various species
of Laminaria (Fig. 13.21) predominate. Their simple or cleft
blades may be 3 m (10 ft) long. The giant kelp (Macrocystis)
dominates kelp communities on the Pacific coasts of North
and South America and other parts of the Southern Hemisphere
(Fig. 13.22). Each kelp individual is attached to the rocky bot-
tom by a large holdfast (see Fig. 6.1). Several long stipes,
intertwined to form a trunk-like foundation, grow from a
single holdfast. The fronds, leaf-like blades, grow from the
stipes. A single stipe can reach 20 to 30 m (65 to 100 ft) or more.
In some Southern Hemisphere communities, Ecklonia rather than
Macrocystis is the main species (see photo on p. 283). Kelp
communities on the Pacific coast of North America are par-
ticularly diverse. A number of species, including the bull kelp
(Nereocystis), the elk kelp (Pelagophycus; Fig. 13.23), Alaria, and
Pterygophora, may be important or even replace Macrocystis.
The bull kelp consists of many blades, each up to 5 m (16 ft)
long, that hang below the surface suspended by a gas-filled
pneumatocyst located at the end of a long stipe. Alaria
( Fig. 13.21 ), which like many other species of kelp is edible,
is also important in the North Atlantic.
Large, dense patches of kelp are known as kelp beds. They
are called kelp forests when the fronds float on the surface in a
thick mat (Fig. 13.24). This floating canopy is characteristic of
Pacific kelp forests dominated by the giant and bull kelps. Ecklonia
and some species of Laminaria and Alaria also form a canopy.
Physical factors have a major influence on kelp communi-
ties. Temperature is of particular importance because kelps are
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Hollow-stemmed kelp
Edible kelp
(Alaria escuelenta)
Horsetail kelp
(Laminaria
digitata)
restricted to cold water. This is partially because kelps dont do
well in warm water and partially because warm waters tend to
lack the rich supply of nutrients that kelps need (see Patterns
of Production, p. 346). This dependence on cold water is reflected
in the geographic distribution of kelps (Fig. 13.22). The surface
waters of the oceans flow in great clockwise gyres in the Northern
Hemisphere and counterclockwise gyres in the Southern
Hemisphere (see Fig. 3.21). The currents that flow toward the
poles on the western sides of the oceans carry warm water from
equatorial regions. Because of this, kelps are restricted to high
latitudes on the western sides of the oceans. On the other hand,
kelps extend well down eastern shores, where cold, nutrient-rich
currents flow down from high latitudes. Ecklonia occurs in low
latitudes not far from the Equator along the southern coast of
the Arabian Peninsula (Fig. 13.22). It is found there only during
the summer, when the summer monsoons that blow northeast-
ward across the Arabian Sea cause strong upwelling that lowers
water temperature (see Fig. 15.31). The disappearance of the
monsoons when the winds reverse direction during the winter
causes an increase in water temperature, which kills the kelp.
All but a few species of kelp are limited to hard surfaces,
which may include substrates such as worm tubes and the hold-
fasts of other seaweeds. Given a suitable place to attach, kelps
will grow in water as deep as light allows. This can be quite deep,
up to 40 m (130 ft) in some species. Their fronds float at the
surface, basking in sunlight, while their stipes connect them to
the bottom far below.
Kelps are actually quite fragile for their giant size; in fact, they
are fragile largely because of their size. The fronds cause a lot of drag,
but the long, thin stipe breaks easily, and kelps are often torn from
the bottom. Drifting kelps cause more damage by entangling other
fronds. Kelps dont do well where there is heavy wave action, and
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CHAPTER 13 Life on the Continental Shelf 297

60 90 120 150 180 150 120 90 60 30 0

60 60

Aleutian Is. Laminaria

Macrocystis
Macrocystis
30 30

Ecklonia
Arabian
Sea Pacific Atlantic
0 0
Ocean Ocean
Indian
Ocean Ecklonia
New Macrocystis
30 Zealand 30
Ecklonia
Macrocystis Macrocystis Tristan da
Cunha
Ecklonia
Chatham I.
Falkland Is.
Warm current
Kerguelen Auckland Is.
Macrocystis
Cold current
60 Macquarie I. 60
South Georgia I. Kelp distribution

60 90 120 150 180 150 120 90 60 30 0

FIGURE 13.22 The geographic distribution of kelps is greatly affected by surface temperatures, which are influenced by the surface circulation of the ocean.
Currents along the west sides of the continents transport cold water from polar regions; on the east sides warm water is transported away from the Equator. As a
result, kelps extend farther toward the Equator along the west sides of the continents than on the east sides. The opposite is true for reef-building corals, which
require warm water (also see Fig. 14.11).

floating fronds, also tend to restrict them

there.
Kelps can grow very fast, with the giant
kelp growing as fast as 50 cm/day (20 in/day).
Not surprisingly, kelp communities are
very productive. Primary production reaches
1,000 gC/m2/year in Ecklonia in Australia
and South Africa, around 1,500 gC/m2/year
in California giant kelp, and close to
2,000 gC/m2/year in Laminaria in the North
Atlantic (also see Table 10.1, p. 225).

Kelp communities are restricted to hard

substrates in cold, nutrient-rich water.

The large kelps we see are only one stage

in their life history. All kelps go through two
stages: a large, spore-producing sporophyte
and a microscopic gametophyte, which pro-
duces male and female gametes (Fig. 13.25).
The sporophyte is the organism we see.

FIGURE 13.23 The elk kelp (Pelagophycus porra) grows on the outer, deeper edges of some giant kelp Monsoons Winds in the northern Indian Ocean
beds (see Fig. 13.26). It has two impressive, antler-like branches from which large blades hang at the mercy that blow from the southwest in summer but
of the currents. from the northeast in the winter.
Chapter 15, p. 352; Figure 15.31
storms can be disastrous. Though they must extend their blades to
Upwelling The upward flow of cold, nutrient-rich
the surface, many species prefer to attach to the bottom in relatively deep water to the surface.
deep water, where wave action is reduced. Thus, the very adaptations Chapter 15, p. 351
that allow these species of kelp to live in deep water, large size and
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298 Part Three Structure and Function of Marine Ecosystems

growth takes place at the holdfast as well as

the end of the stipes.
Kelp forests along the Pacific coast of
North America are arranged in distinct depth
zones, each made up of species that grow at
a characteristic height above the bottom
(Fig. 13.26). This structure results from the
interaction of several physical and biological
factors. The floating canopy of the giant
kelp, for instance, develops only where the
water is deep enough to reduce wave action
but shallow enough for light to reach the
bottom, permitting growth from the hold-
fasts. Other kelps may contribute to the sur-
face canopy. These include the bull kelp,
found closer inshore, and the feather-boa
kelp (Egregia), which lives in shallower water
subject to wave action. The elk kelp forms a
midwater canopy in deeper water along the
outer edge of the giant kelp canopy.
The dense canopy of giant kelp forests
cuts down the amount of light underneath.
Diving under the canopy is like walking into
FIGURE 13.24 Giant kelp (Macrocystis pyrifera) forest at Cedros Island, Baja California, Mexico.
a dense forest on land: It takes a few moments
for your eyes to adjust to the dim light. The
kelps that live there also have to adapt to the
reduced light level, but life under the canopy
Male
gametophyte is rich and varied. Smaller kelps exploit
Germinating (n) the understory, the area below the canopy.
spores
They include Laminaria, Pterygophora, and
other kelps that have either erect fronds that
Spores
(n) stand above the bottom or fronds that lie
Female Sperm right on it.
gametophyte (n) A variety of shorter algae, mostly red
(n)
algae, live on the bottom under the two
Spores
overlying layers, even when light is greatly
reduced. Both branching and encrusting
Fertilization
Egg coralline algae are common. Some sea-
(n)
weeds are more abundant in shallow
water, where increased wave action reduces
the canopy.
Developing North Atlantic kelp beds are domi-
Sporophyte sporophyte nated by species of Laminaria that typi-
(2n) (2n) cally do not form a canopy. They are
similar to Pacific kelp forests, however, in
Immature
sporophyte that they include many species of sea-
Macroscopic (2n) Microscopic weeds and are arranged in depth zones.
stages stages
The complex, three-dimensional struc-
ture of kelp communities is exploited by
FIGURE 13.25 The life history of the giant kelp (Macrocystis) and other kelps includes a large, spore- many different animals. An assortment of
producing sporophyte. Spores settle on the bottom and develop into minute male or female gametophytes.
Each gametophyte releases male or female gametes, which, after fertilization, develop into the sporophyte, polychaetes, small crustaceans, brittle stars,
thus completing the cycle (also see Fig. 6.11a). and other small invertebrates live on the
kelp holdfasts, particularly those of the
In some kelps the sporophyte is an annual. In contrast, the giant giant kelp. Tube-dwelling polychaetes and other sessile organ-
kelp and others are perennial and can live for several years. These isms are common on the blades and stipes. Like the animals
long-lived kelps often lose fronds to grazers, storms, and waves. that live associated with the holdfast, they are mostly suspension
They are able to grow them back because, unlike many seaweeds, feeders. One conspicuous inhabitant of the blades is a bryozoan
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CHAPTER 13 Life on the Continental Shelf 299

Feather-boa kelp
(Egregia)

Canopy
Fronds on
surface or
in midwater Elk kelp
(Pelagophycus)

Understory
Fronds erect Laminaria
or close to
the bottom
Pterygophora

Algal turf Bull kelp

Short clumps, (Nereocystis)
filaments, and
encrusting algae

re
Giant kelp Insho
(Macrocystis)

py
cano
Kelp

ore
Offsh

FIGURE 13.26 Distribution of the major types of kelps and other seaweeds in a generalized giant kelp (Macrocystis) forest on the Pacific coast of North America.
The complex distribution of algae results from the effects of factors such as light, type of substrate, wave action, depth, number and type of grazers, and even time
of year because some of the seaweeds are annuals.

(Membranipora) that forms thin, lace-like colonies. Its calcareous
encrustations weigh the blades down and cover photosynthetic
tissues, but their effect appears to be minimal. The rocky bottoms
around kelps are inhabited by sponges, sea squirts, lobsters, crabs,
hermit crabs, sea stars (see Fig. 7.45a), abalones (see Fig. 7.22b),
and octopuses, among others.
Fishes are very common in kelp communities. They use
the food resources and shelter provided by the kelp community
in many ways, thus occupying many different ecological
niches. For instance, fishes often use the available resources
by feeding and taking shelter in different areas within the for-
est. Some species feed close to the bottom. In kelp beds along
the Pacific coast of North America, bottom feeders include
many species of rockfi shes ( Sebastes ) and the kelp bass
(Paralabrax clathratus). The California sheephead (Semicossyphus
pulcher) uses its dog-like teeth to crush sea urchins, crabs, and
other bottom invertebrates. Surf-perches ( Rhacochilus,
Brachyistius ) and others may feed in different parts of the
canopy, around the holdfasts, or in the open water among
the kelp. Topsmelt ( Atherinops ) are plankton feeders that
take advantage of large swarms of opossum shrimps, or
mysids, and other planktonic animals found around kelps.
Fishes may define additional ecological niches by being active
at different times of the day or night.
Small algae are grazed by snails, crabs, sea urchins, and
fishes, but surprisingly few grazers eat the large kelps. One giant
kelp grazer, the Stellers sea cow, is now extinct (see Fig. 18.13).
Isopods (Phycolimnoria) are small crustaceans that burrow into
the holdfast, weakening it. A few fishes graze on kelps, but they
do not appear to cause much mortality. Instead of feeding on the
attached, actively growing individuals, animals use most of the
huge production of kelps in the form of drift kelp, pieces that
break loose and sink to the bottom or are washed ashore. As with
seagrasses, salt-marsh plants, and mangroves, much of this detri-
tus is exported to other communities.
Kelp beds form a multistoried, complex environment. Drift kelp
and understory seaweeds are a major food source but not the
live kelp themselves.
Sea urchins are by far the most important grazers in kelp
communities. Of special importance are the red (Strongylocentrotus
franciscanus) and purple (S. purpuratus) sea urchins on the Pacific
coast of North America and the green sea urchin (S. droebachien-
sis; see Fig. 7.47a) in both the North Atlantic and North Pacific.
Sea urchin populations sometimes explode (see Fig. 10.3). These
explosions have had devastating impacts on kelp communities in
Ecological Niche The combination of what a species eats, where it lives,
how it behaves, and all the other aspects of its lifestyle.
Chapter 10, p. 215
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300 Part Three Structure and Function of Marine Ecosystems

islands in the Aleutian chain, Alaska, kelp

100 forests were found to be healthy where
sea otters were common. In contrast,

Number in islands
75 there were many sea urchins and few

(100 in 1972)
Sea otter kelps on islands where sea otters were
50
abundance absent (Fig. 13.27). There is evidence
25 that otter populations have declined
because of predation by killer whales,
which normally prefer seals and sea lions.
1985 1989 1993 1997
Seals and sea lions have become increas-
ingly scarce since the late 1980s, perhaps
because overfishing has reduced their food
400
Sea urchin supply. Killer whales also appear to have
g per 0.25m2

300 biomass changed their feeding habits when an

important food source, the carasses of
200
harpooned whales tied to whaling ships,
100 dwindled when most commercial whaling
was banned. Overfishing thus may have
1985 1989 1993 1997 affected kelp forests by way of a chain of
events that includes seals and sea lions,
killer whales, sea otters, and sea urchins.
% loss of kelp 24 hrs.

Observations in the Aleutians, how-

60 Grazing
ever, do not help explain the situation in
50 intensity
40
Southern California, where sea otters
30 were wiped out almost 200 years ago, but
20 the destruction of kelps by sea urchins
10 was not observed until the 1950s. Heavy
fishing on other urchin predators, includ-
1985 1989 1993 1997
ing lobsters, crabs, and fishes, probably
plays a role in the increase in urchin pop-
ulations in Southern California and other
Number per 0.25m2

10 Kelp density places.

8 Another possibility is that a decrease in
6 the amount of drift algae in Southern
4 California might have caused the urchins to
switch to feeding on live kelps. Sewage pol-
2
lution and temperature increases are factors
HEALTHY OVERGRAZED that may have caused a drop in the avail-
KELP FORESTS 1985 1989 1993 1997 KELP FORESTS
ability of drift kelp. Some of these factors
FIGURE 13.27 The effect of predation of sea otters (Enhydra lutris) by killer whales (Orcinus orca) on may also stimulate the growth and survival
the populations of sea urchins and kelps in kelp forests in the Aleutian Islands, Alaska. The effect is an of sea urchins. Some organic compounds
example of a trophic cascade, so called because the effects of a predator extend throughout the food web.
In the graphs, years without bars indicate years without data and not zero values. released in sewage, for example, are used as
nutrients by juvenile sea urchins.
The harvesting of Pacific abalones for
different parts of the world. Sea urchins normally feed on drift food in Southern California has also been proposed as a possible
kelp. During population explosions, or plagues, however, the cause of sea urchin plagues. These large molluscs compete with
urchins eat live kelps and other seaweeds. When the sea urchins sea urchins for shelter in rock crevices. Removal of the abalones
eat the holdfasts or stipes, the kelps break loose, float away, and may provide more space for urchins.
die. The sea urchins may even climb up the kelp, weighing down It is also possible that fluctuations in the number of sea urchins
the fronds and allowing other urchins to reach them. The urchins are caused by the higher survival of their planktonic larvae, perhaps
may completely clear large areas, which are then known as urchin the result of more favorable temperatures or more abundant food.
barrens or urchin deserts. Encrusting coralline algae are virtu- It is likely that a combination of factors causes urchin plagues,
ally the only seaweeds left on these barrens. which probably are natural fluctuations in population size.
The reasons for such outbreaks of urchins remain unclear. In addition to grazing by sea urchins, another factor affecting
In the North Pacific a possible cause is the decline in sea otters the health of kelp forests is climate. A strong El Nio in 1983
(Enhydra lutris; see Fig. 9.12), an urchin predator that has dis- produced severe storms and unusually warm currents, which
appeared from most of its former range. In a study of several caused high kelp mortality. The 19971998 El Nio was
cas28193_ch13_283-302.indd Page 301 8/1/07 5:29:56 PM luxminarayan
similarly destructive. A La Nia that followed the 19971998
El Nio, however, brought cold, nutrient-rich currents to the
California coast, stimulating the recovery of kelp forests.
Kelp communities may be severely disrupted by strong wave
action, grazing by sea urchins, warm currents, and pollution.
The recovery of Southern California kelp forests has pro-
gressed well in a few areas. It has been aided in part by transplant-
ing healthy kelps tied to blocks into depleted areas (see Restoration
of Habitats, p. 417). To appease fishers, who were angry at the
possibility of sea otters feeding on valuable shellfish, it was agreed
that sea otters were to be kept from naturally migrating from
central to Southern California. A 2007 survey showed 3,026 sea
otters along the California coast, a slight increase since a 2004
peak of 2,825 and continuing a recovery after decreases in numbers
from 1995 to 2002. Sea otters are unfortunately being infected by
protozoan parasites of humans. One such parasite is Giardia,
which is found in the small intestines of humans, dogs, and cats.
It has also been found in whales. Toxoplasma, a parasite carried
by cats, has been linked to lethal brain infections in sea otters,
seals, and dolphins. Marine animals get these parasites from sew-
age and urban-runoff pollution.
Will measures to stimulate the recovery of kelp beds have an
effect on kelp beds? Nobody really knows. One thing is for sure:
Some Southern California kelp forests have not recovered and
perhaps never will.
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CHAPTER 13 Life on the Continental Shelf 301
A similar situation with a different cast of characters has
been described in the Gulf of Maine on the Atlantic coast of the
United States. Overfishing of cod (Gadus) and other bottom-
feeding fishes reduced pressure on young lobsters (Homarus),
crabs, and sea urchins. The number of sea urchins steadily
increased and they grazed on kelp, which is used by lobsters as
shelter. Heavy harvesting of sea urchins for export to Japan led
to a reduction in their numbers in the early 1990s. This develop-
ment and continued overfishing of bottom fishes have resulted in
a sharp increase in the number of lobsters. A total of 28.5 million
kg (63 million lb) of lobsters worth around $290 million were
caught during the 2005 season, more than 2.5 times the 1945
1985 average.
Evidence from other parts of the world, however, seems to
indicate that catastrophic disturbances of kelp communities are
usually followed by a recovery, all in recurring cycles. Sea urchins
and kelps are apparently kept in a delicate balance that can be
tipped one way or the other by factors such as climate fluctuation,
the effects of nutrient pollution on the survival of urchin larvae,
and the removal of urchin predators.
El Nio A warming of the surface water in the Eastern Pacific, part of
large-scale changes in atmospheric and ocean current patterns, or ENSO.
Chapter 15, p. 353
La Nia A cooling trend of the surface water in the Eastern Pacific.
Chapter 15, p. 355