Topic 1: What determines the number

of species in a community?

Topic 1: Learning outcomes

1. Understand the different types of diversity (alpha, beta, gamma), and how
diversity varies across latitude
2. Explain the differences between evolutionary and ecological hypotheses for
latitudinal gradients in species richness, and why it is so difficult to determine
which of these actually generate the gradient
3. Describe the difference between the museum and cradle hypotheses, and
explain what evidence could be used to support each
4. Describe (verbally & mathematically) a species-area relationship
5. Explain which processes are important for within-region (nested) versus
between-region species-area relationships
6. Use Island Biogeography Theory to explain how species richness varies with
island size and isolation
7. Explain a dynamic equilibrium
8. Distinguish between processes that are neutral with respect to species identity,
and those that are based on species niches
9. Contrast dispersal limitation versus biotic limitation of local species richness
10. Explain the difference between a species fundamental and realized niche
11. Understand the potential influence of competition, facilitation, and predation
on local species richness
12. Design and interpret experiments to test for species interactions & dispersal
limitation.

Species-area curves: one of the oldest

statistical patterns in ecology

Log number of plant species

1859: First species-area curve (H.C. Watson)

Great Britain
S. England
Surrey

Village

Log area (square miles)

The relationship between diversity and area

Species richness increases with area, usually in a very
predictable way.
WHY????
Several factors that we now know influence diversity are
themselves affected by the amount of habitat (i.e. area):
-Resources
-Speciation
-Extinction
-Immigration

Why you should care about

species-area theory

Determining how many species

can live in a given sized patch is
increasingly crucial in our
fragmented habitats

Area and resources

Area -> Amount of resources - > Number of individuals->
Number of species
The number of individuals sets an
upper limit to how many species
can occur because:
1. A species cannot exist if its
population is not large enough
to be viable. (More individuals
hypothesis: REGIONAL SCALE)
2. Sampling more individuals
means a greater chance of
sampling any given species.
(Sampling effect: LOCAL SCALE)

Beetles in decaying wood

Area and immigration

What is the effect of area on immigration rate?
(At least for passive dispersers)
A. Larger areas are bigger targets for colonists Yes, for passive
dispersers
B. Larger areas have more resources Post-immigration process
C. Larger areas have more births Post-immigration process
D. Larger areas may be avoided by colonists
E. No effect

Area and emigration

-Its pretty unlikely that a species will be lost from an island or
habitat patch because they all just leave
-So for now, these effects can be ignored.

Speciation rate increase with

region area
But only on large islands
isolated for long periods of time
Anolis lizards on Caribbean islands:
speciation rate increases on islands
> 3000 km2, where lizards have
been present for the last 30 million
years.

Speciation rate increase with

region area
But only on large
islandseffects on
So area
isolated forspeciation
long periods ofare
timeNOT

relevant for smaller

Anolis lizardsareas
on Caribbean
and islands:
shorter
speciation rate increases
on islands
timeframes
> 3000 km2, where lizards have
been present for the last 30 million
years.

Area and extinction

Extinction rates decrease with island area
Reason 1: Total population size is smaller in small
regions, so declining populations reach zero faster
Reason 2: Declining or extinct populations on small
islands have less chance of being rescued by other
populations
Reason 3: Small islands may be subject to stronger or
more frequent disturbances

Area and extinction

Reason 1: Total population size is smaller in small regions, so
declining populations reach zero faster.

Weeks to extinction

Daphnia

Maximum population size

Area and extinction

Reason 2: Extinct populations in small areas are less easily
rescued

Area and extinction

Reason 3: Small islands have higher disturbance rates

Maximum elevation (m)

Lizards absent
Lizards present

Before
hurricane

7 mo. after

Log10 Area

Clicker question
If you have a group of moss patches of different sizes, and you want
to maximize species richness in the small patches, which strategy
might you employ?
A)
B)
C)
D)
E)

Fence off the small patches to prevent emigration

Build bridges between patches to increase immigration
Reduce the area of the large patches
Increase the area of the large patches
Increase speciation rate on the small patches

How to make a species-area relationship (SAR)

Step 1: Pick an
ecological system
Here we will look at
mites living in the soil
beneath moss patches

How to make a species-area relationship (SAR)

Step 2:Choose what

type of SAR you
want to construct

Within region
(nested)

Between regions

How to make a species-area relationship (SAR)

Step 3:Count the

number of species in
each area

30 species

100 cm2

How to make a species-area relationship (SAR)

Step 4:Plot the

number of species
against the sampled
area
Species

Within-region (nested)
SAR: plot the cumulative
number of species
against the cumulative
sampled area!

Species = c(Area)z

between

within

Area

How to make a species-area relationship (SAR)

Q1. Mathematically,
how do the withinregion and betweenregion lines differ?
Species

Species = c(Area)z

Q2. Why do the withinregion and betweenregion lines differ?

between

within

Area

How to make a species-area relationship (SAR)

Log S = log c+ z*log A

between

within

Area

Log Species

Species

S = cAz

between
within

Log Area

SARs can be plotted in either linear or log-log space, but it

can be easier to see the patterns in log-log space

S = cAz

Log S = log c+ z*log A

If a SAR has c = 0.2 and z = 0.25 then:

A. Species increase proportionally with area
B. Species decrease with area
C. There are 0.2 species in a region size 0

Z = 1 : proportional
(linear in linear space)

Z= 0 : no relationship

D. Species increase with area, at a decelerating rate

E. Species increase exponentially with area

0 < Z < 1: decelerating

increase

How to make a species-area relationship (SAR)

Q1. Mathematically,
how do the withinregion and betweenregion lines differ?

Moss patches:
Between-region z = 0.38
Within-region z = 0.22

How to make a species-area relationship (SAR)

Q1. Mathematically,
how do the withinregion and betweenregion lines differ?
Between-region z > within-region z

Q2. Biologically, why do

the within- region and
between-region lines
differ?

Log Species

Log S = log c+ z*log A

between
within

Log Area

Within-region SAR
100 ha
60 ha
10 ha

Between-region SAR

60 ha

To keep the comparison fair, well

set the total amount of area to be
equal between the two regions
-In both cases, the sampling effect
and the increase in resources with
area are operating
-But in the between-region SAR,
there is also immigration at play!

30 ha
10 ha

100 ha
60 ha
10 ha

Log Species

Within-region SAR

Thats why the difference between

the between- and within-region
SARs is greatest for large areas.
(In the within-region SAR, large
areas cant get immigration from
anywhere!)

Between-region SAR

60 ha

between
within

Log Area

30 ha
10 ha

Discussion question
Youre biologists in charge of creating some nature reserves. Your
goal is to maximize gamma diversity across your reserves. Does it
make more sense to preserve a Single Large reserve or Several
Small reserves (the SLOSS debate)?

Why?

27

Topic 1: Learning outcomes

1. Understand the different types of diversity (alpha, beta, gamma), and how
diversity varies across latitude
2. Explain the differences between evolutionary and ecological hypotheses for
latitudinal gradients in species richness, and why it is so difficult to determine
which of these actually generate the gradient
3. Describe the difference between the museum and cradle hypotheses, and
explain what evidence could be used to support each
4. Describe (verbally & mathematically) a species-area relationship
5. Explain which processes are important for within-region (nested) versus
between-region species-area relationships
6. Use Island Biogeography Theory to explain how species richness varies with
island size and isolation
7. Explain a dynamic equilibrium
8. Distinguish between processes that are neutral with respect to species identity,
and those that are based on species niches
9. Contrast dispersal limitation versus biotic limitation of local species richness
10. Explain the difference between a species fundamental and realized niche
11. Understand the potential influence of competition, facilitation, and predation
on local species richness
12. Design and interpret experiments to test for species interactions & dispersal
limitation.