Biogeography : Diversity of Species on Islands

The diversity o f species varies considerably from place to place, even when
the habitats appear to be the same. Conservationists have argued that the
size of a region is important for diversity, and so they often favor a few large
wilderness areas rather than many tiny ones. The subject is far from under
stood. We study one corner of it briefly.

The world is broken into patches of differing habitats. Often a habitat

a species finds acceptable is surrounded by a large expanse of unacceptable
territory. Examples are alpine meadows, farm woodlots, lakes, game
preserves, and islands. The following discussion is confined to islands ;
however, most of the ideas and results apply to other types of isolated habitats.
The material is adapted from R. H. MacArthur and E. O. Wilson ( 1 967,
Ch. 3) which treats the subject in much greater depth.

Studies have indicated that the size of an island is an important factor

in determining the number of species the island is likely to contain. Also,
islands closer to the mainland tend to contain a greater variety of species
than more isolated islands. It seems reasonable that the effects of migration
of species and extinction of species (on islands) can account for this. We
develop this idea and briefly consider some of its consequences.

A species can become established on an island only by migrating to it

and prospering there. An organism migrates by flying, being carried, drifting
on currents, and so on. Since a population on an island is relatively small,
it can die out because of random variations in the environment. As a result
we expect the list of specie-s present on an island to change much faster than
the list of species present on the mainland.

This is somewhat vague. Does a flock of migrating birds that stops on the
island for a day or a season become established and then die out ? Even if a
species " intends " to stay on the island, we are still faced with the problem
of what we mean by " become established "if the island is too small to
support a large population, the species will always be on the verge of
extinction. When is a species established in this case ? Since we are dealing
with a fairly crude model, we can afford to ignore these problems. A more
refined model would have to come to grips with them.
If we uderstood all the aspects of the situation (e.g., the
biological, geographical, and meteorological), we could determine the
probability of a particular species composition being present on the island
at a given future time. These would be tremendously complex calculations
involving vast quantities of data, and this approach would be hopeless.
completely50
G R A P H I C A L M ET H O D S
Let's combine practically all the endogenous variables into one measure :
the total number of species present on the island. It seems reasonable to
suppose that this should vary around some average number of species in a
steady state situation. We discuss this average. For a discussion of transient
behavior see R. H. MacArthur and E. O. Wilson ( 1 967, Ch. 3) or E. O. Wilson
and W. H. Bossert (197 1 , Ch. 4).
When the number of species present on the island is
that is, the rate of migration
of new species to the island equals the rate of extinction of species already on
the island. These rates depend in a complicated way on the species present,
the season, and many other factors. If we regard a year as a short period of
time, seasonality will present no problem. In this sort of crude averaging over
many species, which species are actually present probably doesn't matter
much. Therefore it makes sense to talk about rates in a crude way independent
of which species are actually present on the island.
In Figure 3 are plotted the number of species
on the island versus
the migration and extinction rates. The two smaller graphs illustrate the
effect of distance from the mainland and the effect of island size. We discuss
the reasons for the shapes and positions of the curves.
Let's consider the extinction rate curves. When more species are present
on the island, the chances that at least one species will become extinct in a
given time are greater. Hence the extinction rate curves have a positive slope.
Since extinction rates depend only on the island and the species present, the
extinction curve is not affected by the distance from the mainland. However,
we can expect that a species is more likely to die out on a small island because
the lack of space keeps the population lower. Thus the extinction rate curves
shift upward as the islands become smaller.
Why do the migration rate curves have a negative slope ? The migration
rate relates to species
The greater the diversity on
the island, the smaller the pool of potential migrating species on the mainland.
Hence the chances of migration decrease as the number of species on the
island increases. Migration rates depend on the distance of the island from the
mainland and on the size of the island. The rates decrease with distance,
because any given organism is less likely to reach the island. The rates increase
with island size because ( 1 ) an organism has a larger land area as a target
and (2) an organism is more likely to be able to establish itself on a larger
island.
It follows from Figure 3 that the number of species present increases
with island size and decreases with distance from the mainland. This is not
so surprising, since we practically put these results in as initial assumptions.
We can say something about species turnover by looking at the graphs
a bit more. Note that the equilibrium extinction rate (which equals the
in equilibrium,
migration and extinction cancel out numerically ;
N
not present on the island.C O M P A R AT I V E STAT I C S
51
(a)
Large -
Smai l - - Biogeography :
D ive r s i ty o f S pec i es on I s l a n d s
The diversity o f species varies considerably from place to place, even when
the habitats appear to be the same. Conservationists have argued that the
size of a region is important for diversity, and so they often favor a few large
wilderness areas rather than many tiny ones. The subject is far from under
stood. We study one corner of it briefly.
The world is broken into patches of differing habitats. Often a habitat
a species finds acceptable is surrounded by a large expanse of unacceptable
territory. Examples are alpine meadows, farm woodlots, lakes, game
preserves, and islands. The following discussion is confined to islands ;
however, most of the ideas and results apply to other types of isolated habitats.
The material is adapted from R. H. MacArthur and E. O. Wilson ( 1 967,
Ch. 3) which treats the subject in much greater depth.
Studies have indicated that the size of an island is an important factor
in determining the number of species the island is likely to contain. Also,
islands closer to the mainland tend to contain a greater variety of species
than more isolated islands. It seems reasonable that the effects of migration
of species and extinction of species (on islands) can account for this. We
develop this idea and briefly consider some of its consequences.
A species can become established on an island only by migrating to it
and prospering there. An organism migrates by flying, being carried, drifting
on currents, and so on. Since a population on an island is relatively small,
it can die out because of random variations in the environment. As a result
we expect the list of specie-s present on an island to change much faster than
the list of species present on the mainland.
This is somewhat vague. Does a flock of migrating birds that stops on the
island for a day or a season become established and then die out ? Even if a
species " intends " to stay on the island, we are still faced with the problem
of what we mean by " become established "if the island is too small to
support a large population, the species will always be on the verge of
extinction. When is a species established in this case ? Since we are dealing
with a fairly crude model, we can afford to ignore these problems. A more
refined model would have to come to grips with them.
If we
understood all the aspects of the situation (e.g., the
biological, geographical, and meteorological), we could determine the
probability of a particular species composition being present on the island
at a given future time. These would be tremendously complex calculations
involving vast quantities of data, and this approach would be hopeless.
completely50
G R A P H I C A L M ET H O D S
Let's combine practically all the endogenous variables into one measure :
the total number of species present on the island. It seems reasonable to
suppose that this should vary around some average number of species in a
steady state situation. We discuss this average. For a discussion of transient
behavior see R. H. MacArthur and E. O. Wilson ( 1 967, Ch. 3) or E. O. Wilson
and W. H. Bossert (197 1 , Ch. 4).
When the number of species present on the island is
that is, the rate of migration
of new species to the island equals the rate of extinction of species already on
the island. These rates depend in a complicated way on the species present,
the season, and many other factors. If we regard a year as a short period of
time, seasonality will present no problem. In this sort of crude averaging over
many species, which species are actually present probably doesn't matter
much. Therefore it makes sense to talk about rates in a crude way independent
of which species are actually present on the island.
In Figure 3 are plotted the number of species
on the island versus
the migration and extinction rates. The two smaller graphs illustrate the
effect of distance from the mainland and the effect of island size. We discuss
the reasons for the shapes and positions of the curves.
Let's consider the extinction rate curves. When more species are present
on the island, the chances that at least one species will become extinct in a
given time are greater. Hence the extinction rate curves have a positive slope.
Since extinction rates depend only on the island and the species present, the
extinction curve is not affected by the distance from the mainland. However,
we can expect that a species is more likely to die out on a small island because
the lack of space keeps the population lower. Thus the extinction rate curves
shift upward as the islands become smaller.
Why do the migration rate curves have a negative slope ? The migration
rate relates to species
The greater the diversity on
the island, the smaller the pool of potential migrating species on the mainland.
Hence the chances of migration decrease as the number of species on the
island increases. Migration rates depend on the distance of the island from the
mainland and on the size of the island. The rates decrease with distance,
because any given organism is less likely to reach the island. The rates increase
with island size because ( 1 ) an organism has a larger land area as a target
and (2) an organism is more likely to be able to establish itself on a larger
island.
It follows from Figure 3 that the number of species present increases
with island size and decreases with distance from the mainland. This is not
so surprising, since we practically put these results in as initial assumptions.
We can say something about species turnover by looking at the graphs
a bit more. Note that the equilibrium extinction rate (which equals the
in equilibrium,
migration and extinction cancel out numerically ;
N
not present on the island.C O M P A R AT I V E STAT I C S
51
(a)
Large -
Smai l - -
(b)
(c)
F i g u re 3
Migration .and extinction curves for islands . (a) Typical curves. (b) Effect of
distance . ( c) Effect of size .
equilibrium migration rate) is greater for near islands than for far islands.
Hence the species composition for two islands of equal size should change
more rapidly on the island closer to shore. If the effect of island size on
migration rate is not too great, we can similarly conclude that the species
composition changes faster on small islands than on large islands. Since
small islands have fewer species at equilibrium than large islands, this effect
should be quite noticeable.
There is some data supporting the conclusions that species turnover is
relatively and absolutely more rapid on smaller islands. R. H. MacArthur
and E. o. Wilson ( 1 967, pp. 52-54) discuss the results of two botanical
surveys of some small islands off the Florida Keys. The first survey ( 1904)52
G R A P H I CA L M ET H O D S
was conducted quickly and s o may b e incomplete. Since the 1 9 1 6 survey
was quite complete, the species present in 1 904 and absent in 1 9 1 6 give some
measure of the turnover rate. Unfortunately the data involve only six islands,
two of which are very small . .
R. H. MacArthur and E. O. Wilson ( 1967, pp. 55-60) also report on
some results of R. Patrick. She suspended glass slides in a spring in
Pennsylvania and counted the number of diatoms of various species that
were present. The glass slides can be thought of as islands. Four experiments
were done two times each : A glass slide with an area of either 12 or 25 square
millimeters was placed in the water for either 1 or 2 weeks. The slides sub
merged for 1 week had more species present than those submerged for 2
weeks. We can explain this apparent contradiction by observing that as a
barren area becomes more populated the interaction between species may
cause extinction. This was not allowed for in our model. Clearly care must
be taken in modeling islands that are far from equilibrium. Because of this,
we do not consider the 1 week data further. We can check out two predictions
using the 2 week data :
1.
Larger area implies more species : The smaller slides had 24 and 2 1
species, and the larger had 2 9 and 28.
2. Smaller area implies a higher migration rate : The migration rate may be
reflected somewhat in the differences in the species composition of the
slides. (Why ?) Seven species appeared on one but not both of the smaller
slides. For the larger slides the number was one.
(b)
(c)
F i g u re 3
Migration .and extinction curves for islands . (a) Typical curves. (b) Effect of
distance . ( c) Effect of size .
equilibrium migration rate) is greater for near islands than for far islands.
Hence the species composition for two islands of equal size should change
more rapidly on the island closer to shore. If the effect of island size on
migration rate is not too great, we can similarly conclude that the species
composition changes faster on small islands than on large islands. Since
small islands have fewer species at equilibrium than large islands, this effect
should be quite noticeable.
There is some data supporting the conclusions that species turnover is
relatively and absolutely more rapid on smaller islands. R. H. MacArthur
and E. o. Wilson ( 1 967, pp. 52-54) discuss the results of two botanical
surveys of some small islands off the Florida Keys. The first survey ( 1904)52
G R A P H I CA L M ET H O D S
was conducted quickly and s o may b e incomplete. Since the 1 9 1 6 survey
was quite complete, the species present in 1 904 and absent in 1 9 1 6 give some
measure of the turnover rate. Unfortunately the data involve only six islands,
two of which are very small . .
R. H. MacArthur and E. O. Wilson ( 1967, pp. 55-60) also report on
some results of R. Patrick. She suspended glass slides in a spring in
Pennsylvania and counted the number of diatoms of various species that
were present. The glass slides can be thought of as islands. Four experiments
were done two times each : A glass slide with an area of either 12 or 25 square
millimeters was placed in the water for either 1 or 2 weeks. The slides sub
merged for 1 week had more species present than those submerged for 2
weeks. We can explain this apparent contradiction by observing that as a
barren area becomes more populated the interaction between species may
cause extinction. This was not allowed for in our model. Clearly care must
be taken in modeling islands that are far from equilibrium. Because of this,
we do not consider the 1 week data further. We can check out two predictions
using the 2 week data :
1.
Larger area implies more species : The smaller slides had 24 and 2 1
species, and the larger had 2 9 and 28.
2. Smaller area implies a higher migration rate : The migration rate may be
reflected somewhat in the differences in the species composition of the
slides. (Why ?) Seven species appeared on one but not both of the smaller
slides. For the larger slides the number was one.