9 ISLAND BIOGEOGRAPHY

Objectives
• Explore the relationships of immigration and extinction
rates and species richness to island area and distance from
the mainland.
• Observe the accumulation of species on an island, and the
approach of immigration and extinction rates and species
richness values to equilibrium.
• Find equilibrium values of immigration and extinction rates
and species richness, both graphically and algebraically.
• Understand species-area curves and the underlying mathe-
matical relationships implied.
• Explore the interaction effects of area and distance.

INTRODUCTION
People have long known that larger islands, and islands closer to a mainland,
support a greater number of species than smaller or more distant islands. Most
ecology textbooks give examples of such species-area and species-distance rela-
tionships, not only for islands in the strict sense, but also for habitat islands
such as mountaintops and lakes. Few books explicitly state the mathematical rela-
tionship between number of species and area or distance, but most show them as
straight lines on log-log plots. This should indicate to you that the underlying
relationships are power functions. (See Exercise 1, “Mathematical Functions and
Graphs,” for definitions and examples of power functions and other kinds of func-
tions.) On linear axes, both relationships are curves, hence the term “species-area
curve” and what could be called the “species-distance curve.”
Having observed and quantified these relationships, ecologists proposed sev-
eral hypotheses to explain them. One of the best-known hypotheses is the equi-
librium theory of island biogeography developed by Robert MacArthur and
Edward O. Wilson.

The MacArthur-Wilson Model of Island Biogeography

MacArthur and Wilson (1967) modeled species richness (the number of species
present) on an island as the result of two processes: immigration and extinction.
In their model, species immigrate to an island randomly from a mainland pool.
The rate at which new species arrive at the island is determined by three factors:
126 Exercise 9

• The distance of the island from the mainland

• The number of species remaining in the mainland pool that have not already
established themselves on the island
• The probability that a given species will disperse from the mainland to the island

The rate at which species on the island go extinct is also determined by three different
factors:
• The area of the island
• The number of species present on the island
• The probability that a given species on the island will go extinct

In the simplest version of the model, all species have equal probability of reaching the
island and of going extinct once there. The model ignores interactions such as compe-
tition, predation, or mutualism between species on the island.
We will develop a spreadsheet model incorporating these ideas. Let us begin with
immigration. It seems reasonable to suppose that the farther an island lies from the main-
land, the lower the rate of immigration—in other words, immigration is inversely related
to distance. Since immigrants are drawn from a finite pool, as more species establish
themselves on the island, fewer species will remain in the pool that have not already
established themselves on the island. Based on these considerations, we can write a sim-
ple equation for the rate of immigration to an island. Let
I = immigration rate (Note: This is overall immigration rate of species to the
island, which is different from the probability that any one species will make
that journey)
P = total number of species in the mainland pool
S = species richness of the island
D = distance of the island from the mainland
c = colonization probability, or the probability that a given species will make it to
the island; here it is assumed to be equal for all species
f = a scaling factor for distance

Note that (P – S) is the number of species in the mainland pool that have not already
reached the island. Now we can write an equation for immigration:

I = c( P − S) Equation 1
fD
We must determine a values for c and f from actual data. Based on the work of
MacArthur and Wilson, we can begin with reasonable values of c = 0.10 and f = 0.01.
Note that Equation 1 is a power function, in which the variable D is raised to a constant
power, –1.
Turning our attention to extinction, we can write a simple equation for that as well.
Let
E = extinction rate
S = species richness of the island
A = area of the island
q = extinction probability for a given species (assumed to be equal for all species)
m = a power scaling factor for area

Now we can write an equation for extinction:

qS
E= Equation 2
Am

Values of q and m must be determined from actual data, and based on work by
MacArthur and Wilson, we can begin with a reasonable values of q = 0.20 and m = 0.25.
 Island Biogeography 127

Note that Equation 2 is also a power function, in which the variable A is raised to a con-
stant power, m.
If you consider Equation 1, you can see that as species accumulate on an island
(i.e., as S increases), the immigration rate, I, will decrease. Inspection of Equation 2
shows that as S increases, the extinction rate, E, will increase. At some value of S, immi-
gration and extinction will become equal (i.e., I = E), and species richness will come to
an equilibrium. This is an equilibrium because every new species immigrating to the
island is balanced by one already-established species going extinct, and vice versa.
This is an important point of the model: Equilibrium species richness is determined by a
balance between immigration and extinction. Note that this is a statement about the model,
not about species richness on real islands, which is certainly affected by other factors
in addition to immigration and extinction. However, like other simple models, this
one has proven fruitful in stimulating thinking and research.
A second important point of the model is that the equilibrium in species richness is
a dynamic equilibrium. At equilibrium, immigration and extinction rates are equal,
but neither is zero. The rate of immigration or extinction at equilibrium species rich-
ness is called the turnover rate.
According to the model, then, the particular species inhabiting an island continue
to change, or turn over, indefinitely—even after species richness has reached equilib-
rium. That is, species continue to go extinct and are replaced by an equal number of
immigrating species. A biologist revisiting the same island at different times would,
according to the model, find different sets of species present, but (at least roughly) the
same total number of species.
This prediction of continuing turnover is an important feature of MacArthur and Wil-
son’s model. This model is often used in conservation biology to predict the number of
species that would be expected to persist or go extinct in nature reserves (which are often
habitat islands). However, it is not useful in planning for protecting specific species,
because of this prediction of continuing turnover.

PROCEDURES
This exercise is presented in four parts. In each part you will develop a spreadsheet
model and make graphs. Between parts, we return to a little mathematical exposition
to lay the groundwork for modeling.
First you will build a spreadsheet version of the MacArthur-Wilson model of island
biogeography. Using Equations 1 and 2, you will graphically estimate the species rich-
ness of an island. In the second part, you explore how the island’s area and distance
from the mainland affect its species richness. In the third part, you will examine the time-
course of species accumulation on an island. In the fourth part, we derive equilibrium
solutions for species richness and turnover rate.
As always, save your work frequently to disk.

INSTRUCTIONS ANNOTATION

A. The MacArthur-
Wilson island biogeog-
raphy model.

1. Open a new spreadsheet Enter the text items and values shown for “Parameters” and “Scaling factors.” These
and set up titles and col- are all literals, so just select the appropriate cells and type them in.
umn headings as shown in
Figure 1.
128 Exercise 9

A B C D E F G
1 Island Biogeography
2 Assumes all species have equal dispersal ability and risk of extinction.
3 Assumes no interaction between species on an island.
4
5 Parameters Scaling factors
6 Species pool on mainland (P) 1000 For distance (f ) 0.01
7 Area of island (A) 200 For area (m) 0.25
8 Distance from mainland (D) 300
9 Colonization probability (c) 0.10
10 Extinction probability (q) 0.20 Equilibium values
11 Species richness
12 Immigration, extinction, and species richness Immigration rate
13 Fraction of pool Species richness Immigration Extinction Extinction rate
14 0.0
15 0.1
16 0.2

Figure 1

2. Set up a series: 0.0, 0.1, In cell A14 enter the value 0.

0.2, . . . , 0.9, 1.0 in cells In cell A15 enter the formula =A14+0.1.
A14–A24. Copy the formula from cell A15 into cells A16–A24.
This series represents different fractions of the mainland pool present on the island,
from 0% to 10% and so on to 100%.

3. In cell B14 enter a for- In cell B14 enter the formula =A14*$C$6. This formula is based on the fraction of the
mula to calculate the actu- mainland pool in cell A14 and the total number of species in the mainland pool.
al number of species pres-
ent on an island.

4. In cell C14 enter a for- In cell C14 enter the formula =$C$9*($C$6-B14)/($G$6*$C$8). This corresponds to Equa-
mula to calculate the rate tion 1:
of immigration to an I = c( P − S)
fD
island already colonized
by the number of species
in cell B14.

5. In cell D14 enter a for- In cell D14 enter the formula =$C$10*B14/$C$7^$G$7. This corresponds to Equation 2:
mula to calculate the rate
qS
of extinction on an island E=
already colonized by the Am
number of species in cell
B14.

6. Copy the formulae in Select cells B14–D14. Copy.

cells B14–D14 down their Select cells B15–D24. Paste.
columns to row 24. Save your work!

7. Graph immigration and Select cells B14–D24 and make an XY graph. Edit your graph for readability. It should
extinction rates against resemble the graph in Figure 2.
species richness.

8. Try changing the You should see that smaller or more distant islands have fewer species than larger or
parameter values in cells closer ones. We will examine these relationships more rigorously in the next part of the
C6–C10, one at a time, and exercise.
observe how equilibrium
species richnesss changes.
 Island Biogeography 129

Immigration and Extinction

60

50

40

30 Immigration
Extinction
20

10

0
0 200 400 600 800 1000
Species richness

Figure 2

Effects of Island Area and Distance from the Mainland

In Step 8 of the preceding section of the exercise, you experimented with different
parameter values to see the effects on species richness. In this section, we will examine
the effects of an island’s area and its distance from the mainland somewhat more rig-
orously.
To quantify these effects, let us compare three islands of the same area, but at three
distances from the mainland: 0.5, 1.0, and 2.0 times some distance that you specify (in
cell C8 of your spreadsheet). Looking at Equation 1, which models the immigration rate,
you can see that it includes distance but not area. Accordingly, we will compute immi-
gration rates on these three islands, and estimate the effects on species richness.
We will also compare three islands at the same distance from the mainland, but
having three different areas: 0.1, 1.0, and 10.0 times the area that you specified in cell C7
of your spreadsheet. Looking at Equation 2, which models extinction rate, you can see
that it includes area but not distance. Accordingly, we will compute extinction rates on
these three islands and estimate the effects on species richness.

INSTRUCTIONS ANNOTATION

B. The effects of dis-

tance and area on the
MacArthur-Wilson
model.

1. Add the column head- These are all literals, so just select the appropriate cells and type them in.
ings shown in Figure 3 to
cells I12–P13 of the
I J K L M N O P
spreadsheet you set up in
12 Fraction Species Effect of distance on immigration Effect of area on extinction
Part A (see Figure 1). 13 of pool richness Imm near Imm medium Imm far Ext small Ext medium Ext large
14 0.0
15 0.1

Figure 3
130 Exercise 9

2. Set up a series: 0.0, 0.1, Copy cells A14–A24 into cells I14–I24.
0.2, … , 0.9, 1.0 in cells This series represents different fractions of the mainland pool present on the island,
I14–I24. from 0% to 10% and so on to 100%.

3. In column J, calculate In cell J14, enter the formula = I14*$C$6. Copy this formula into cells J15–J24.
the actual numbers of
species present on islands,
based on the fraction of
the mainland pool in cell
I14 and the total number
of species in the mainland
pool.

4. In column L, calculate In cell L14, enter the formula =($C$6-$J14)*$C$9/($C$8*$G$6). This corresponds to
immigration rates to Equation 1:
islands at the distance I = c( P − S)
fD
specified in cell C8, using
the species richnesses cal- Note the use of an absolute column address for cell $J14.
culated in column J. Copy this formula into cells L14–L24.

5. In column K, calculate Copy the formula from cell L14 into cell K14, and edit it to multiply distance (cell C8)
immigration rates to by 0.5.
islands at half the distance The new formula should read =($C$6-$J14)*$C$9/($C$8*0.5*$G$6).
specified in cell C8, using Copy the formula from cell K14 into cells K15–K24.
the species richnesses cal-
culated in column J.

6. In column M, calculate Copy the formula from cell K14 into cell M14, and edit it to multiply distance (cell C8)
immigration rates to by 2.0.
islands at 2.0 times the The new formula should read =($C$6-$J14)*$C$9/($C$8*2.0*$G$6).
distance specified in cell Copy the formula from cell M14 into cells M15–M24.
C8, using the species rich-
nesses calculated in col-
umn J.

7. In column O, calculate In cell O14, enter the formula =$J14*$C$10/$C$7^$G$7. This corresponds to Equation 2:
extinction rates for islands
qS
of the area specified in cell E=
C7, using the species rich- Am
nesses calculated in col- Again, note the use of an absolute column address for cell $J14.
umn J. Copy the formula from cell O14 into cells O15–O24.

8. In column N, calculate Copy the formula from cell O14 into cell N14, and edit it to multiply area by 0.1.
extinction rates for islands The new formula should read =$J14*$C$10/($C$7*0.1)^$G$7.
of 0.1 times the area speci- Copy the formula from cell N14 into cells N15–N24.
fied in cell C7, with the
species richnesses calculat-
ed in column J.

9. In column P, calculate Copy the formula from cell N14 into cell P14, and edit it to multiply area by 10.0.
extinction rates for islands The new formula should read =$J14*$C$10/($C$7*10.0)^$G$7.
of 10.0 times the area spec- Copy the formula from cell P14 into cells P15–P24.
ified in cell C7, with the Save your work!
species richnesses calculat-
ed in column J.
 Island Biogeography 131

10. Graph immigration Select cells J13–M24.

rates for near, medium- Hold down the control key or  while selecting cells O13–O24.
distance, and far islands Make an XY graph. Edit your graph for readability. It should resemble the one in Fig-
along with the extinction ure 4.
rate for a medium-sized
island against species rich-
ness. Immigration Rates and Island Distances

70

Immigration or extinction rate

60

50
Imm near
40
Imm medium
Imm far
30
Ext medium
20

10

0
0 200 400 600 800 1000
Species richness

Figure 4

11. Graph extinction rates Select cells J13–J24.

for small, medium-sized, Hold down the control key or  and select cells L13–L24.
and large islands, and the Hold down the control key or  and select cells N13–P24.
immigration rate for a Make an XY graph. Edit your graph for readability. It should resemble the one in Fig-
medium-distance island, ure 5.
against species richness.

Extinction Rates and Island Areas

100
90
Immigration or extinction rate

80
70
60 Imm medium
Ext small
50
Ext medium
40 Ext large
30
20
10
0
0 200 400 600 800 1000
Species richness

Figure 5
132 Exercise 9

*12. As an OPTIONAL exer- Select cells J13–P24, and make an XY graph. This will allow you to compare species
cise, graph three immigra- richness and turnover rates on islands of three different sizes, at three different dis-
tion rates and three extinc- tances from the mainland. However, your graph might be rather cluttered and hard
tion rates on a single to read.
graph.

The Time-Course of Species Accumulation on an Island

The graphical analyses above answer a variety of questions about species richness on
islands at equilibrium. However, they tell us nothing about how species richness
changes over time as it approaches equilibrium. To find out about that, we must model
the time-course of species accumulation.
We can follow the accumulation of species over time using a discrete-time model. The
number of species present on an island at time t + 1 will be the number present at time t
plus the number of new species that immigrated in the interval from time t to t + 1, minus
the number of species that went extinct in the interval from t to t + 1. In symbols,
St+1 = St + It – Et

Substituting the right-hand side of Equation 1 for It and the right-hand side of Equa-
tion 2 for Et, we derive

c( P − St ) qSt
St +1 = St + − m Equation 3
D A

INSTRUCTIONS ANNOTATION

C. Model the time-

course of species accu-
mulation.
1. Add the column head- These are all literals, so just select the appropriate cells and type them in.
ings shown in Figure 6 to
cells A26 and A27 though
D27 of the spreadsheet A B C D
you created in Part A (see 26 Time-course of species accumulation
Figure 1). 27 Time Species richness Immigration Extinction
28 0
29 1
30 2

Figure 6

2. Set up a linear time In cell A28 enter the value 0.

series from 0 to 50 in cells In cell A29 enter the formula =A28+1.
A28–A78. Copy the formula from cell A29 into cells A30–A78.

3. Begin with an uninhab- Enter the value 0 in cell B28.

ited island.
 Island Biogeography 133

4. Enter a formula to cal- Copy the formula from cell C14 into cell C28. This corresponds to Equation 1.
culate the number of
species immigrating to the
island in the interval from
time 0 to time 1.

5. Enter a formula to cal- Copy the formula from cell D14 into cell D28. This corresponds to Equation 2.
culate the number of
species going extinct on
the island from time 0 to
time 1.

6. Enter a formula to cal- In cell B29 enter the formula =B28+C28-D28.

culate the number of This corresponds to Equation 3:
species present on the
island at time 1. c( P − St ) qSt
St +1 = St + − m
D A

The formula calculates the number of species on the island as the number already there
plus the number immigrating to the island, minus the number going extinct, in the pre-
ceding time interval.

7. Copy the formulae in

cells C28 and D28 into
cells C29 and D29.

8. Copy the formulae in Save your work!

cells B29–D29 into cells
B30–D78.

9. Graph species richness, Select cells A27–D78 and make an XY graph.

immigration rate, and After you have made your graph, double-click on any data point in the species rich-
extinction rate against ness curve. In the Format Data Series dialog box, click on the Axis tab, and choose Sec-
time. ondary axis. Plot species richness on the secondary y-axis.

To label the second y-axis, open Chart|Chart Options|Titles.

Edit your graph for readability. Your graph should resemble the one in Figure 7.

Species Accumulation
Immigration or extinction rate

35 450
30 400
350
Species richness

25
300
Immigration
20 250
Extinction
15 200
Species richness
150
10
100
5 50
0 0
0 10 20 30 40 50
Time (t )

Figure 7
134 Exercise 9

Equilibrium Solutions
So far, you have estimated equilibrium species richness using graphs. In the next sec-
tion, we will calculate these quantities algebraically. We have two reasons for doing so.
First, calculations give us more precise results than estimating from a graph. Second,
these calculations will allow us to close the loop, metaphorically, with the original moti-
vation for MacArthur and Wilson’s model. As we said at the beginning of this exercise,
among the original observations from which this model sprang were the relationships
of species richness to island area and distance from the mainland–the species-area curve.
But nothing we have done so far explicitly shows a species-area curve. By finding
equilibrium solutions, we can develop these curves, and briefly indicate how they have
been used to test the model and to guide conservation decisions.
As we explained in the first section of this excercise, the MacArthur-Wilson model
tells us that species accumulate by immigration and are removed by extinction, and that
species richness reaches equilibrium when these two processes balance. Algebraically,
we can find the equilibrium species richness of an island by solving for Seq when I = E.
So, let’s do a little algrebra.
Let I = E

Substituting from Equations 1 and 2 above, we can derive the equation for Seq:

c( P − Seq ) qSeq
= m
fD A
Amc(P – Seq) = fDqSeq

AmcP – AmcSeq = fDqSeq

AmcP = fDqSeq + AmcSeq

AmcP = Seq(fDq + Amc)

A m cP = S
eq Equation 4
fDq + A m c

Equation 4 isn’t very pretty, but you can use it in your spreadsheet model to see how equi-
librium species richness relates to island area, to colonization and extinction probabilities,
and to the richness of the mainland species pool. In particular, we will see how the model
predicts species-area curves for islands at different distances from the mainland.

INSTRUCTIONS ANNOTATION

D. Calculate species
equilibrium.

1. Enter a spreadsheet for- In cell G11 enter the formula =C7^G7*C9*C6/(G6*C8*C10+C7^G7*C9). This corre-
mula for equilibrium sponds to Equation 4:
species richness into cell A m cP = S
eq
G11. fDq + A m c

2. Enter the spreadsheet In cell G12 enter the formula =C9*(C6-G11)/(G6*C8).

equivalents of Equations 1 In cell G13 enter the formula =C10*G11/C7^G7.
and 2 into cells G12 and These are the rates of immigration and extinction, respectively, on an island already
G13. colonized by the number of species in cell B14 (Equations 1 and 2). Use the values in
these cells to verify your graphical estimates in the previous parts of this exercise.
 Island Biogeography 135

3. Enter the row and col- We will use this part of the spreadsheet to calculate species area curves for islands at
umn labels shown in different distances from the mainland.
Figure 8 into cells
R11–X13.
R S T U V W X
11 Species-Area Relationships at Different Distances
12 Equilibrium species richness Turnover rates
13 Area Near Medium Far Near Medium Far
14 10
15 50
16 100

Figure 8

4. To represent a wide Enter the values10 and 50 into cells R14 and R15, respectively.
range of island areas, set In cell R16, enter the formula =R14*10. Copy this formula into cells R17–R24.
up a series 10, 50, 100, 500,
1000 … , 500,000, 1,000,000
in cells R14–R24.

5. In column T, calculate In cell T14 enter the formula =$R14^$G$7*$C$9*$C$6/($G$6*$C$8*$C$10+$R14^$G$7*

the equilibrium species $C$9), which again corresponds to Equation 4. Copy this formula into cells T15–T24.
richnesses of islands at the Note that the address $R14 has an absolute column reference but a relative row refer-
distance specified in cell ence.
C8, with the areas given in
column R.

6. In column S, calculate Copy the formula from cell T14 into cell S14. Edit the formula to multiply distance by 0.1.
the equilibrium species The edited formula should read =$R14^$G$7*$C$9*$C$6/($G$6*0.1*$C$8*$C$10+
richnesses of islands at 0.1 $R14^$G$7*$C$9).
times the distance speci- Copy the edited formula from cell S14 into cells S15–S24.
fied in cell C8, with the
areas given in column R.

7. In column U, calculate Copy the formula from cell S14 into cell U14. Edit the formula to multiply distance by 10.
the equilibrium species The edited formula should read =$R14^$G$7*$C$9*$C$6/($G$6*10*$C$8*$C$10+
richnesses of islands at 10 $R14^$G$7*$C$9).
times the distance speci- Copy the edited formula from cell U14 into cells U15–U24.
fied in cell C8, with the
areas given in column R.

8. Graph equilibrium Select cells R13 though U24, and create an XY graph. Edit your graph for readability;
species richness against It should resemble Figure 9. The three species-area curves will rise very quickly, almost
island area for islands at following the vertical axis on the left, and then abruptly level out.
near, medium, and far dis-
tances from the mainland.
136 Exercise 9

Species-Area Curves

1,200

Equlibrium species richness

1,000

800
Near
600 Medium
Far
400

200

0
0 200,000 400,000 600,000 800,000 1,000,000

Island area

Figure 9

9. Change both vertical As in Figure 10, the species-area curves should become almost straight lines on the log-
and horizontal axes to log- log plot.
arithmic scales.

Species-Area Curves

1,000
Equlibrium species richness

100
Near
Medium
Far
10

1
1 100 10,000 1,000,000

Island area

Figure 10
 Island Biogeography 137

QUESTIONS
1. How can you estimate the equilibrium species richness of an island from Figure 2?
2. Is the equilibrium of species richness stable or unstable?
3. Is the equilibrium of species richness static or dynamic?
4. How does greater distance from the mainland affect species richness on an
island?
5. How does greater distance from the mainland affect the turnover rate on an
island?
6. How does larger area affect species richness on an island?
7. How does larger area affect the turnover rate on an island?
8. (OPTIONAL) How do area and distance from the mainland interact to determine
species richness and turnover rate on an island?
9. How do species accumulate on an island over time? That is, does species rich-
ness increase linearly, exponentially, logarithmically, or otherwise?
10. What does Figure 7 tell us about the changing state of species equilibrium?
11. How is species richness related to island area?
12. How do the species-area curves differ for islands at different distances from the
mainland?

LITERATURE CITED
MacArthur, R. H. and E. O. Wilson. 1967. The Theory of Island Biogeography.
Princeton University Press, Princeton, NJ.