Kuliah 12 & 13

Islands Biogeography
(Biogeografi Pulau)

Island Biogeography
In the science of biogeography, the
island is the first unit that the mind can
pick out and begin to comprehend. By
studying clusters of islands, biologists
view a simpler microcosm of the
seemingly infinite complexity of
continental and oceanic biogeography.
Robert H. Mac Arthur and Edward O.
Wilson (1967)

Darwin and Wallace

Galapagos and Malaysian
Archipelago
Observed unique character of island
biota
1880 Island Life - Wallace

Islands can serve almost as a laboratory for the

study of biogeography. The biota of an island is
simpler than that of a continental area, and the
interactions are easier to understand.

There are three types of islands:

a. Islands that were originally part of a nearby
continent, but were separated by rising sea
levels (land-bridge islands).
b. Islands that are part of a volcanic island arc.
c. Seamount chains which formed over
geological hotspots.

Continental/Landbridge Islands
Connected to mainland during
glacials
Oceanic Islands
Never connected to continent;
Usually formed by volcanic activity;
Usually separated from continent by
deep ocean.
Hotspot islands
Chain of islands

A.

Continental Islands: Formed on continent; may

have formerly been
connected to mainland by land bridge:

Island

Current Sea Level

Continent
Submerged Land Bridge
Continental Shelf

Former
Sea
Level

Examples of Continental Islands

1. British Isles
2. California Channel Islands
3. Tasmania
4. Penang

California Channel Islands: Group of eight islands

off the California coast; during last ice age, some were
connected to mainland by land bridge.

Continental Islands:
Two Unusual Cases
1. San Salvadors offshore cays: Rising sea level caused
erosion of San Salvador, leaving many small, erosionResistant islands, or cays (keys).

Cays on the horizon (arrow) were once

part of San Salvador.

B. Oceanic Islands: Never connected to

continent; usually formed by volcanic
activity and isolated from continent
by deep ocean.
Oceanic
Island
Current Sea Level

Continental
Shelf

Former Sea
Level

Sea Floor

Undersea
Volcano

Examples of Oceanic Islands

Iceland
Japan
Aleutians
Bermuda
Caribbean Islands
Hawaiian Islands
South Pacific Atolls
Et al.

Many Caribbean islands were formed

by volcanic activity at subduction
zone.

Volcanic activity at subduction zone along continent

can form oceanic islands (e.g., Japan; Aleutians).

Source: Wikipedia http://en.wikipedia.org/wiki/File:Japan_separation.png

Volcanic activity at mid-ocean ridge can form

ocean islands (e.g., Iceland).

Convergence of Two Oceanic Plates:

Denser plate sinks Under lighter plate
= subduction zone.

Source: Wikipedia
http://en.wikipedia.org/wiki/Plate_tectonics

Convergence of Oceanic and Continental Plates:

Subduction zone results in earthquake and volcanic
activity (e.g., Pacific Rim of Fire).

Source: Wikipedia http://en.wikipedia.org/wiki/Plate_tectonics

Volcanic Activity at Tectonic Plate Boundaries

Source: USGS http://pubs.usgs.gov/gip/hawaii/page10.html

Map of hot spots

http://www.math.montana.edu/~nmp/materials/ess/geosphere/advanced/activities/hotspots/index.html

The Hawaiian Island Archipelago is

actually
a very long and extended island chain

Northwest Movement of Pacific Plate Over

Fixed Hawaiian Hot Spot

Source: USGS http://pubs.usgs.gov/gip/hawaii/page12.html

What determines the number of

species on an island?

The types of islands have different

characteristic flora and fauna. Islands formed
by isolation from continents would have a
biota which would be a subset of that on the
continent. It would have changed, however,
as the result of independent evolution and
extinction. The biota of island arcs and
hotspot island chains originally arrived by
trans-ocean dispersal. In both cases, several
islands exist at one time, creating the
possibility for inter-island dispersal and a more
complex pattern of evolutionary change.

Dispersal to islands is typically by a

sweepstakes route,. The dispersing
organisms share adapations that allow
them to reach the island, rather than
adaptations allowing them to live there
once they reach it. This is one factor that
restricts the diversity of life on islands.

How do propagules disperse to

islands from the mainland?
1. Self propelled flying or swimming
(walking on land)
2. Blown by the wind
3. Float in or on water
4. Can be carried by another species

What limits dispersal?

Failure to find (land on) the island

Failure to survive over the dispersal
distance
Lack of food/nutrients/freshwater

Predation
Herbivory
Mortality due to disease/parasites
Short life span

Species Diversity
More
species on
bigger
islands

Fewer
species on
more
isolated
islands

Some flying
animals, such as
birds and bats, are
capable of reaching
even very distant
islands.

Most land animals must rely on

dispersal mechanisms like drifting on
masses of debris. Although this
process is likely rare, it certainly
happens and has been documented
for organisms like iguanas.

Long distance dispersal in plants is

much more likely. A great many plants
are adapted for such dispersal. In
addition, the long distance dispersal of
a plant species can typically be
accomplished by a single spore or
seed, where in animals it typically
requires a pair of organisms or a
pregnant female.

Some plants have developed

seeds or fruits that can be
carried in the sea without being
harmed.

There is no doubt that

the degree of isolation
of an island or island
group is a factor in
determining the biota
that it will support.

For conifers and flowering plants in the

Pacific, diversity is much lower in the more
isolated island groups of the central and

The ratio of observed species to the

expected number declines with distance
from New Guinea.

If we plot the number of genera vs. island

area, it becomes clear that the two are
related. The more isolated islands
(represented by ) have fewer genera
that less isolated islands of the same
size.

Also, when a species is lost by

extinction, it is more difficult to
replace it be immigration than in a
mainland situation.
For these, and other reasons, islands
tend to support fewer species than
mainland areas of similar size.

Species vs. Area

relationships
Sp

Area

Sp

Area

Species

Species vs. Area

relationships

S=c
Az
Area

explains 50% of variance in species richness of islands.

Species Area Relationship

S cA

S = species richness
c = constant, regionally specific
richness
A = sampling plot size
z = constant, shape of curve (slope)

Species

Species vs. Area example

S=cA

c=1
c=1 z=0.25
z=0.25
Area

Area
(km2)
Species

10

100

1000

10000

10

Species-Area Curve

S = cAz
C=
constant
A = area
z = 0.25

Species vs. Area

relationships

Log Species

S
S=
= cc
zz
A
A

Log
Log S
S=
= Log
Log cc +
+ zz
log
log A
A
Log Area

Reptiles on Islands

Pond Size and Fish

MacDonald 2001

Habitat and Mammals

Island populations are more likely to

go extinct than those on mainlands,
for several reasons:
1. Populations are typically smaller.
2. They have less genetic diversity.
3. They were not originally adapted to
the island habitat.

In 1963, Robert MacArthur and E.O.

Wilson presented a new hypothesis
to explain patterns of species
richness on islands. Their
equilibrium theory of island
biogeography proposed that the
lower number of species on islands
was not the result of insufficient
time, but rather the result of an
equilibrium process peculiar to all
islands.

The theory is based on the idea that, at

any given time, the number of species
on an island is the result of a balance
between two processes: extinction and
colonization.

MacArthur & Wilsons Equilibrium

theory
MacArthur R.H.

MacArthur R.H.
&Wilson
&WilsonE.O.
E.O.1967
1967
The
Thetheory
theoryof
ofisland
island
biogeography.
biogeography.
Princeton
PrincetonUniv.
Univ.Press.
Press.

Number of species on an
island

Theory of Island
Biogeography
MacArthur and Wilson 1963,
1967

Number of species
on islands is
determined via a
balance of
extinction and
immigration
(colonization)

Island Size
Richness = island size and distance
from mainland
Small islands
Less habitat
Smaller populations
Higher rates of extinction (intra,interspecific competition)

Island Distance
Richness = island size and distance
from mainland
Distant islands
Lower rates of colonization
However, this does depend on dispersal
mechanism of the species!

Theories, like islands, are often reached by

stepping stones
MacArthur and Wilson (1967)

Important Concepts

Equilibrium number of
species, but constant
turnover
Migration slows as
richness increases
Extinction increases
with richness
Equilibrium = rate of
extinction, rate of
colonization intersect
Colonization balances
extinction

Equilibrium no. of species

Theory of Island Biogeography

Immigration varies with the distance of the
island from the mainland (or the pool of
potentially colonizing species).

More distant islands are colonized less

frequently.
The rate of colonization declines as
species richness increases, because
there are fewer potential colonists and
fewer unexploited niches.

Theory of Island Biogeography

Extinction varies with the size of the island.

Big islands can support larger population

sizes, and large populations are less prone
to extinction. So, the smaller the island,
the greater the probability of extinction.
Extinction rates rise as colonization
increases; interspecific competition
displaces of some species and lowers
population sizes when species can coexist.

Theory of Island Biogeography

At the same time as colonisation is
occurring, some species will become
extinct.
As new species arrive they will compete
for limited resources and the
extinction rate will increase.
The rate at which one species is lost and
a replacement is gained is the
turnover rate.

Theory of Island Biogeography

Number of Species

Theory of Island Biogeography

Number of Species

Theory of Island Biogeography

Extinctio
n rate

Immigration
rate

Number of Species

Theory of Island Biogeography

Extinctio
n rate

Immigration
rate

Near
mainland
Far

Number of Species

Theory of Island Biogeography

Small island

Immigration rate

Large

Far

Number of Species

Extinction rate

Near

Theory of Island Biogeography

Islands further from mainland have lower immigration rates

More distant islands have lower species diversity

Theory of Island Biogeography

Smaller islands have lower total populations

Probability of extinction increases with lower population

Smaller islands have lower

species diversity

Equilibrium Theory of Island Biogeography

Immigration rate decreases as

island diversity increases

Extinction increases as island

diversity increases

Species equilibrium on islands

is a balance of immigration and
local extinction

Island Size
Richness = island size and distance
from mainland
Small islands
Less habitat
Smaller populations
Higher rates of extinction (intra,interspecific competition)

Island Distance
Richness = island size and distance
from mainland
Distant islands
Lower rates of colonization

When a new
island forms,
species begin to
colonize. As
more and more
species
accumulate, the
colonization rate
begins to
decline. The
extinction rate,
on the other
hand, begins to
increase with
increasing

At some
point, the
two
processes
balance each
other, and
the number
of species on
the island
should
stabilize.
This
equilibrium
number is
known as S

The equilibrium theory can also be

used to explain the effect of size and
distance on the number of species
found on islands.
Consider two islands of similar sizes
but different distances from the
mainland pool. Since extinction rates
are a function of the available
resources and should be related to
the size of the island, we would
expect them to be similar on the two
islands. Colonization rates, however,
should be greater for the island near
the mainland than for the more

This should result in a difference in the

equilibrium number of species, with Nnear >
Nfar

A similar argument can be used to

explain the effect of island size. If two
islands are of relatively equal distance
from the mainland, we can expect
colonization rates to be similar.
Extinction rates, however, should be
greater on the smaller island.
Therefore, we expect a higher
equilibrium number of species on the
large island.

MacArthur and Wilson 1972

(mainland)

Some islands are of the land bridge

type while others arose at sea and
have never had a connection to the
mainland.
Oceanic islands confirm pretty
closely to the patterns predicted by
island biogeographic theory. Land
bridge islands are a different story.

Land bridge islands begin with the

species complement to be expected
of a mainland area. So, over time,
we expect the number of species to
diminish.

So we see a different pattern for the

number of species as a function of time
for a:
land-bridge
island..

or an
oceanic island.

Examples of Islands Studies

Island life is probably more hazardous

than that on the mainland. For one
thing, catastrophic events have more
severe effects. There is typically no
place to hide.

1883 - Eruption of Krakatau (Krakatoa), a

volcanic island in Indonesia (Aug. 26-27).

Half of Krakatau was blown away; remaining portion,

Rakata (a volcanic cone), plus neighboring islands,
left covered with
30-60 m of pumice and ash (= sterile landscape?).

Rakata and adjacent islands formed a

laboratory for study of island colonization
and tropical succession:

May 1884 - first researchers reach the

islands; find only a spider in a crevice
on the south side of Rakata.
October 1884 - grass shoots growing on
Rakata.

1886 - Botanists, and later zoologists, begin

monitoring colonization of Rakata:
nine species of flowering plants present on
beaches;
1897 - 23 species of flowing plants present;
development of coastal forest provided seeds
and fruits for colonizing bats and birds;
ferns (with spores that can be dispersed by
wind) were first colonizers away from the
coast;

1908 - 46 species of flowing plants

and 13 species of birds present;
1934 - 30 species of birds present; but,
at least 5 bird species present in early
20th century were now extinct on
Rakata;

 ~ 50% of inland plant species on Rakata

in 1897 have become extinct; however,
since 1934, 16 additional families of
higher plants have colonized.

 colonization by new plant species

was initially high, then dropped as
available space became occupied by
pioneer species;
immigration rate then increased as
developing forests created new
habitat (= potential new ecological
niches);
as forests replaced grasslands,
grasses, and insects and birds
dependent on grasses, became extinct
on island.

Biogeographical lessons from Krakatau:

Composition of plant and animal commun

at any given time reflect

colonization
local extinction
succession
disturbance

Recent studies* have re-evaluated

ecological succession and
extinctions on Rakata and adjacent
islands since 1883:
Most plant extinctions have been
species introduced by people, and
rare or ephemeral species;
Few naturally colonizing and
established species have become
extinct.
*e.g., Whittaker, R.J. et al. 1992 GeoJournal 28.2: 201-211.
Whittaker, R.J. et al. 2000, J. Biogeograpy 27(5):10491064

MacArthur and Wilson 1967

Theory of Island Biogeography

Why do islands have fewer species than same area on
continent?

Function of SIZE of island and DISTANCE from mainland

Small islands have higher extinction rates.

Farther islands have lower probability of immigration.