Intertidal Ecology

Outline
1. Zonation

upper limit set by physical environment
lower limit set by biological interactions
competition
predation

2. Species Diversity within a Rocky Intertidal Community

keystone predator (Pisaster)
set by periodic disturbance

3. Trophic Cascading Effects

Sea Otters and Kelp Forests Example

Vertical Zonation
the hallmark of the intertidal zone
Communities are divided into distinct bands, or zones, at characteristic
heights in the intertidal.

Species are not randomly distributed throughout the intertidal zone, but rather
are arranged within relatively narrow vertical ranges.

The zones look like sharply divided belts easily distinguished by the colors of the
assemblage (i.e. community) of organisms that live there

Physical Stresses: Often set Upper Limit to Species

Distributions:
Stress Factors
1. Desiccation

2. Temperature

3. Food Availability

4. Wave Energy

5. Salinity

6. Dissolved Oxygen

Biological Interactions: Often set Lower Limit to

Species Distributions:
Biological Factors
1. Competition for Space

note that space on a rock to attach
is the valuable resource that is in
short supply in the intertidal

2. Predation

The Cause of Zonation:

A Simple Barnacle Example

1. The Upper Limit of both species (grey and rock

barnacles) is determined by emersion; larvae that settle
too high in the intertidal dry out and die (physical
factor).

Little grey barnacles can tolerate drying better than
rock barnacles, so they settle higher in the intertidal.

Grey Barnacles

2. At Lower Levels, where the rock barnacles can

survive, the rock barnacle out competes the little grey
barnacle for space (biological factor) and this sets to
lower limit for the little grey barnacle.

3. The Lowest Limit of the adult rock barnacles is
determined by competition from mussels, and
predation by whelks or sea stars (biological factors).

Rock Barnacles

Summary:

General Rule of Zonation

upper limit at which a species can
occur in the intertidal is usually
determined by physical factors,

lower limit is usually determined by
biological factors

Species Diversity within a Rocky Intertidal

Community
1. Keystone Predators

2. Intermediate Disturbance

Starfish Predation:
Sets Lower Limits of Mussel Distributions in
Rocky Intertidal

Leads to Higher Species Diversity within a
Rocky Intertidal Community
(Paine, 1966)

1. Mussels can out compete most of the other

intertidal organisms for valuable space

2. Pisaster (Starfish) predation sets the lower
distributional limit to mussels and below this
distributional limit other species can settle in

3. The removal of Pisaster allows mussels to take
over and this leads to a decrease in the
communitys species diversity (just mussels
remain).

Keystone Predators: Predators that have effects on their communities

that are proportionately much greater than their abundance would
suggest are know as keystone predators

A rocky intertidal
starfish called Pisaster
is a classic type of
keystone predator

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Massive Die-Off of Starfish Along the US Westcoast

Required to Watch Before Next Wednesdays Lecture with Prof. Drew Harvell
h"ps://www.youtube.com/watch?v=2_I_B6U0GtI

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Physical Disturbance Can Also Regulate Species Diversity within

a Community
1. Physical disturbance (e.g., wave energy
from storms and log damage) can open
up gaps or patches in the rocky
intertidal.

2. Intermediate Disturbance Hypothesis disturbance maximizes species diversity
by periodically removing competitively
dominant species and allowing less
competitive species to reestablish
themselves

Note : Too much disturbance keeps the rock bare with few species. Too little
disturbance allows the dominant competitor for space to take over and form a
monoculture (single species)

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Trophic Cascade:
Sea Otter-Kelp Forest Example
1. Sea Otters eat sea urchins

2. Sea urchins are herbivores that
eat tiny young kelp (before they
grow large).

3. Removal of sea otters allows sea
urchins to grow to high
abundance

4. Low abundance of sea otters
leads to high abundance of sea
urchins and low abundance of
kelp forests

2004 KenAshman

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Summary
1. Zonation

upper limit set by physical environment

lower limit set by biological interactions

competition

predation

2. Species Diversity

Pisaster (starfish) - A Keystone Species

intermediate disturbance

3. Trophic Cascade Effects

Sea Otters and Kelp Forests

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Coral Reefs

15

Coral Anatomy

Calcium Carbonate Support Structure

NOTE: the process of building a calcium carbonate reef structure layer-by-layer is a very slow process:
<1mm per year to about 20 mm per year

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Coral Energetics
Zooxanthellae are
chlorophyll-containing algal
symbionts that live in the
tissue of the coral polyp
Corals receive 60- 90% of
their overall nutrition from
photosynthetic-derived
products!!!

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Limits to Coral Net Growth

1.
2.
3.
4.

Temperature

Sunlight

Space to Grow

Predation

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Temperature: Limits Coral Growth to Tropical Latitudes

Optimal 26-28C

Restricted 18-36C

!

Sunlight:

Limits Coral Growth to a depth range extending from the ocean
surface down to maximum of about 25 meters

Light required for zooxanthellae to photosynthesize

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Boundary for 20 C Isotherm: Most corals live within this boundary. Note the
cooler waters caused by upwelling on the southwest coast of Africa and off the
coast of Peru.

http://upload.wikimedia.org/wikipedia/commons/thumb/c/cb/20_Grad_Isotherme.png/1024px-20_Grad_Isotherme.png

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Location of Coral Reefs

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Depth Limit Where Corals Can Grow is Set by Light

Requirements
Because corals rely heavily
on the photosynthesis of
Zooxanthellae, they have a
depth limit, set by light
levels, below which they
cannot grow effectively this limit is typically about
25 meters

http://www.nature.com/scitable/knowledge/library/coral-reefs-15786954

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Charles Darwin first proposed the theory

Coral Reef Formation

1. A new island forms (e.g., mantle hot spot
or ocean-ocean plate collision) and a
fringing reef develops in shallow sunlit
waters close to shore of the island

2. The island slowly sinks with age (crust
cools and thins) and at the same time the
coral slowly upward by slowly secreting its
calcium carbonate support structure layerby-layer - this eventually forms a barrier
reef.

3. At time point the entire island is submerged
and all that is left is the reef - referred to
now as coral atoll.

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new island

fringing reef

barrier reef

coral atoll
http://www.nature.com/scitable/knowledge/library/coral-reefs-15786954

Coral Reef Formation

NOTE: If the island sinks too fast, or sea
level rises too fast, and the reef cannot keep
up with its upward growth, it eventually stops
growing because it becomes too submerged in
deep/dark waters

These dead reefs are referred to as drowned

reefs

http://www.nature.com/scitable/knowledge/library/coral-reefs-15786954

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Corals Compete for Space:

With Other Corals and With Macroalgae

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Corals Competing with Other Corals: Sweeper Tentacles

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Corals Competing with Other Corals: Sweeper Tentacles

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Competition with
Macroalgae
The competitive
advantage for taking
over space is shifted in
favor of macroalgae
when nutrients from
agricultural activities
run off the coast and
onto coral reefs

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Predation by Crown of Thorns Starfish (Acanthaster)

1. This starfish is an
important predator of
corals

2. One of the main sources of
coral mortality

3. Eutrophic conditions
increase phytoplankton
abundance that enhances
the growth of Acanthaster
larvae that eventually leads
to large increases in
abundance of adults and
subsequent high coral
mortality

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Coral Bleaching
1. It is the zooxanthellae algae that give
corals all of their beautiful natural
colors

2. Coral bleaching is the name given to an
event where corals expel their
symbiotic zooxanthellae algae due to
environmental stress such as unusually
warm water

3. Corals can recover and regain their
zooxanthellae if the stress is small or
short-lived (a couple of weeks)

4. Coral death follows if the stress is
extreme and/or prolonged.

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Corals are extremely sensitive to rises is sea temperature. Just 1 oC above normal
for a period of a few weeks is enough to result in coral beaching.

http://www.reefpix.org/albums/album62/DSCN1595.jpg

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32

Regions reporting bleaching

Number of Regions Reporting Coral Bleaching Worldwide

10
9
8
7
6
5
4
3
2
1
0
2004

2002

2000

1998

1996

1994

1992

1990

1988

1986

1984

1982

1980

1978

1976

1974

Year

Regions Were: Caribbean, Atlantic coast of South America, Western Pacific, Central Pacific, Eastern
Pacific, North Pacific, South East Asia, North East Asia, Western Indian Ocean and Eastern Indian Ocean.
Data collected from over 3800 bleaching records from nearly 100 countries in the ReefBase bleaching report
database (http://www.reefbase.org, Dec 23, 2002).

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The global Average Loss of Coral is Now at About 27% !!!

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Projected Rise in Sea Surface Temperature Relative to Coral Bleaching Threshold

Denoted by Solid Horizontal Lines

Hoegh-Guldberg 1999

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Ocean Acidification
1. Increases in CO2 in the atmosphere lLeads to Increases in oOcean
acidity

2. Projected Increases in ocean aAcidity lLead to slowing of calcium
carbonate precipitation and/or eventually coral dissolution
3. An atmosphere above
480 ppm is Expected to
Enhance Coral Loss We are now at 400ppm

4. Note that Twice PreIndustrial Atmospheric
CO2 = 550 ppm
Hoegh-Guldberg et al. (2007) Science Vol. 318. pp. 1737 - 1742

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Expected Scenario Of Reef Changes with Increasing Atmospheric CO2

375 ppm!
+1 oC

>500 ppm!
> +3 oC

450-500 ppm!
+2 oC
Hoegh-Guldberg et al. (2007) Science Vol. 318. pp. 1737 - 1742

NOTE: The atmospheric CO2 concentration is currently at 400 ppm and going up by about 3
ppm each year - so it will reach 450 in about 15 years

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Overall:

Coral Reefs Around The World Are In Rapid Decline...

1.Coral reefs around the world are being degraded at an accelerated
pace due to the combined effects of local-scale stressors caused by
land-based pollution and by global-scale stressors due to a
warming planet and increased ocean acidification.

2.It is now widely recognized that local-scale stressors on coral reefs
should be minimized as an immediate means of offsetting increases
in global-scale stressors.

3.Managing local water quality conditions to alleviate the pressure
from global-scale stresses is now a top reef management priority.

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Whales

39

Outline
1. Whale Evolution
2. Feeding Modes
3. Vocalizations
4. Migration

5. Sound Pollution
6. Whaling

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Pakicetus hoofed-mammal that is sometimes classified as the earliest whale. It lived

around 53 million years ago (after the age of dinosaurs). It looked rather like a
dog/wolf with hoofed feet and long, thick tail.

The shape of the ear region in the Pakicetus skull is highly unusual and only
resembles the skulls of whales. The special ear bone feature is diagnostic for
cetaceans and is found in no other species.

41

Ambulocetus ("walking whale") was an early cetacean that could

walk as well as swim. It lived during early Eocene some 50-49 million
years ago. It is a transitional fossil that shows how whales evolved from
land-living mammals.

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Rodhocetus lived in the Middle Eocene 46-47 million years

ago, and was still able to walk on land though not very well.

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Dorudon lived during the Eocene about 40 to 36 million years ago.

Dorudon reached about 5 meters in length and was a carnivore. The
overall look of Dorudon was very similar to the modern whale.
Dorudon lacked the ability to go on land. It had tiny hind legs that
barely protruded from the body.

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Basilosaurus lived at about the same time as Dorudon in the Eocene 40 to 37

million years ago. This creature had a pair of very small hind legs, about 2 feet
long. Its jaw was around 5 feet long and was equipped with cone shaped teeth
in the front and triangular shaped teeth in the back.

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46

Evolutionary Divergence of Baleen Whales From Toothed

Whales
1. All Baleen Whales (Mysticetes) are large filter-feeders

2. The exact means by which baleen are used differ among species:

gulp-feeding

skim-feeding

bottom plowing

3. The first members appeared about 35 million years ago. These
changes may have been a result of worldwide environmental change
and physical changes in the oceans.

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Odontocetes
(Toothed)

Mysticetes

(Baleen)

48

Feeding Modes

49

Feeding Mode of Baleen and Toothed Whales

50

Orca (Killer) Whale

Odontocete
(Toothed)

51

52

BBC Blue Planet

53

Baleen Whales

54

Baleen whales feed by

gulping large quantities of
seawater and then
squeezing the water
through the baleen filter
to retain krill and small
fish.

55

56

57

58

Swimming with a
Humpback (Baleen) Whale

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Whale Migrations

60

Pacific Humpback Migration

1. Summer
Feeding at high
latitudes with long
day length and lots
of biological
productivity

2. Winter
Calving at low
latitudes - usually
warm, but low or
no food

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Global Humpback Migration

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Pacific Grey Whale

Migration

63

North Atlantic Right Whale Migration

64

Vocalization In Whales
Odontocetes (toothed whales)

1. Produce rapid bursts of clicks and whistles

2. Do not make the long, low-frequency, sounds known as the whale song.

3. Single clicks are generally used for echolocation

4. Collections of clicks and whistles are used for communication.

5. The multiple sounds themselves are produced by passing air through a
structure in the head rather like the human nasal passage.

Mysticetes (baleen whales)

1. Often make the long, low-frequency, sounds known as the whale song.

2. Have a larynx that appears to play a role in sound production, but it lacks
vocal cords and scientists remain uncertain as to the exact mechanism.

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Mysticete Vocalization:
Sexual Selection or Navigation

The complex and haunting sounds of the Humpback Whale (and some Blue
Whales) are believed to be primarily used in sexual selection during mating
season, but the simpler sounds of other whales have a year-round use.
1. While many toothed whales are capable of using echolocation to detect
the size and nature of objects (e.g., prey), this capability has never been
demonstrated in baleen whales.

2. Given the poor visibility of aquatic environments, and the fact that sound
travels so well in water, simple sounds may play a role in navigation. For
instance, the depth of water or the existence of a large obstruction ahead
may be detected by loud low frequency noises made by baleen whales.

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Spectrograms are used to visualize the whale vocalization

Bowhead Whale - Shallow, Coastal Arctic

Song (two voiced) normal speed
Slide- Kip Reddington

Frequency

1. Time on the x-axis

2. Frequency on the y-axis

3. Loudness is denoted by
Brighter Colors

Christopher W Clark

Time
Christopher Clark (Cornell Lab of Ornithology)

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Spectrograms are used to visualize the whale vocalization

Frequency

1. Time on the x-axis

2. Frequency on the y-axis

3. Loudness is denoted by
Brighter Colors

Time
Christopher Clark (Cornell Lab of Ornithology)

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Sources of Anthropogenic Sound in the Ocean

1. Commercial Shipping

Engine and Propeller Noise

2. Naval Operations

Low Frequency Active Sonar

3. Oil Exploration

Seismic Surveys with Explosive Air Guns/Cannons

Repeated Every Ten Seconds, 24 hours a day, For Days and
Weeks at a Time

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Spectrograms are used to visualize the whale vocalization

Bowhead Whale - Shallow, Coastal Arctic
Song (two voiced) normal speed
Slide- Kip Reddington

Frequency

1. Time on the x-axis

2. Frequency on the y-axis

3. Loudness is denoted by
Brighter Colors

Christopher W Clark

Time
Christopher Clark (Cornell Lab of Ornithology)

70

The Tides of Ambient Noise (Payne & Webb 1971)

he Tides of
Noise
(Payne & Webb 1971)
NoAmbient
shipping =
Gulf of
CA, Baja Mex

Ambient Ship Noise

No shipping = Gulf of CA, Baja Mex

Low Boat
Traffic
Range (km)

Range (km)

Shipping = Mediterranean
Shipping = Mediterranean

High Boat
Traffic

Distance (km)
Distance (km)

Christopher W Clark

Time (1 day)
Time (1 day)

Christopher Clark (Cornell Lab of Ornithology)

Christopher W Clark

71

Noise Pollution
!

1. Acoustic Habitat is no different than spatial habitat and must be

preserved.

2. Noise Pollution is every bit as destructive as other forms of more
familiar marine pollution (oil, nitrogen, plastic)
Dr. Christopher Clark Cornell Lab of Ornithology...

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Whaling...

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Whaling
An international convention on whaling culminated in the signing of the
International Convention for the Regulation of Whaling (ICRW) in 1946

The aim is to "provide for the proper conservation of whale stocks and thus make
possible the commercial whaling and the orderly development of the whaling
industry".

The International Whaling Commission (IWC) was set up under the terms of the
ICRW to make decisions on quota levels based on the findings from the Scientific
Committee of the IWC. Countries which are not members of IWC are not bound
by its regulations and conduct their own management programs.

The members of the IWC voted on 23 July 1982 to apply a moratorium to all
commercial whaling beginning in 1985...

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Japan continues to whale by claiming its whaling operation is now for scientific
purposes. The IWC allows lethal scientific whaling, but only when it addresses questions
vital to management. Numbers vary each year, but on average it is close to 1,000 Minke
(not-endangered), 50 Fin (endangered), 50 Humpback (endangered) and 5 Sperm
(endangered) whales each year. !
NOTE: The IWC found the Japanese research to be unnecessary, and that the same ends could be
accomplished by non-lethal methods, but Japan continues this practice

Norway registered an objection/reservation when the 1982 IWC whaling

moratorium was signed, and was thus not bound by it. In 1993 Norway decided to
exercise its reservation against the moratorium and resumed domestic commercial whaling.
They now take about 600 Minke (not-endangered) whales each year

Iceland issued licenses in October 2006 for a commercial whale hunt in addition to its
continuing hunt for scientific purposes. Iceland has an exemption to the moratorium
through the reservation it made in 2002. They take about 150 Fin (endangered) and
200 Minke (not-endangered) whales each year.

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