Describing Communities

Basic Species Diversity Concepts

Species Richness
Diversity Indices
- Simpson's Index
- Shannon-Weiner Index
- Brillouin Index
Species Abundance Models

There are two important

descriptors of a community:
1) its physiognomy
(physical structure), as
described in the previous
lecture, and
2) the number of species
present and their relative
abundances (species
richness and diversity).

Species Diversity
By far, species diversity has received the greatest amount of
attention in community ecology. Diversity is an emergent
property of the community.
Current emphasis on biodiversity and conservation has
accelerated interest in this topic, both theoretically and
practically. Biodiversity often involves the inclusion of
genetic and ecosystem diversity as well. But, the focus for
many conservation efforts has remained on species.
Thus, we will devote considerable time to the development of
concepts in diversity.

Species Richness
The simplest way to describe a community is to list the
species in it.
Species richness (S) is the number of species on that list,
and is most often used as the first pass estimate of diversity
for a community. Term coined by McIntosh (1967).
How would one generate such a list? A simple and widely
used method is to define the boundaries of the community
and then walk through it seasonally, noting all the species
you encounter. This is what we call a flora.

Species Richness
Species Richness
Some communities are simple enough to permit a complete
count of the numbers of species presentthis is the oldest and
simplest measure of species richness.
Complete counts of species can often be done in very extreme
environments (certain deserts, polar regions, etc.); however,
this approach can rarely be taken in most mesic environments.
Estimating the total species richness of a community can be a
bit difficult and will be the subject of the subsequent lecture.
For now, lets continue to think about observed richness (Sobs)
from a community sample.

While many studies include S as a descriptive factor

associated with the community, it is largely
uninformative in as much as it does not reflect relative
abundance.
Example: suppose two communities (1 & 2) each contain
100 individuals distributed among five species (A-E):
A

Comm-1

20

20

20

20

20

Comm-2

96

Are these two communities equivalent?

Diversity Indices
In most instances, in order to have an effective measure of
diversity, we need to account for both species richness
and the evenness with which individuals are distributed
among species.
One way to do this is through the use of a proportional
abundance index. There are two major forms of these
indices: dominance indices and information indices.
While more than 60 indices have been described, we will
look at the three most widely used in the ecological
literature: Simpson's, Shannon-Weiner, and Brillouin .

Simpson's Index
Simpson's Index is considered a dominance index because
it weights towards the abundance of the most common
species.
Simpson's Index gives the probability of any two
individuals drawn at random from an infinitely large
community belonging to different species.
For example, the probability of two trees, picked at random
from a tropical rainforest being of the same species would
be relatively low, whereas in boreal forest in Canada it
would be relatively high.

Simpson's Index

Simpson's Index
The bias corrected form of Simpson's Index is:

( n ( n 1) )
=
( N( N 1) )
S

DS

i =1

where n i is the number of individuals in the ith species.

Since D s and diversity are negatively related, Simpson's
index is usually expressed as either reciprocal or
complementary forms (1/D or 1 -D) so that as the index
goes up, so does diversity.

Shannon-Weiner Index

A worked
example for
201 trees of 5
species
assessed in
several
quadrats:

Tree
spp.

No.
Individuals

100

50

30

20

Total

201

100 99 50 49
1 0
DS =
+
+ ...
= 0.338
201 200 201 200
201 200
1 / D = 1 / 0.338 = 2.96 or 1 D = 0.662

Shannon-Weiner Index

The Shannon- Weiner Indexbelongs to a subset of

indices that maintain that diversity can be measured much
like the information contained in a code or message
(hence the name information index).

The Shannon Index assumes that all species are

represented in a sample and that the sample was obtained
randomly:

The rationale is that if we know a letter in a message, we

can know the uncertainty of the next letter in a coded
message (i.e., the next species to be found in a
community).

where p i is the proportion of individuals found in the ith

species and ln is the natural logarithm.

H' = - pil n p i
i=1

The uncertainty is measured as H', the Shannon Index. A

message coded bbbbbb has low uncertainty (H' = 0).

Shannon-Weiner Index
A worked example from a community containing 100
trees distributed among 5 species:

Total

Species

Abund

pi

pi ln pi

50

0.5

-0.347

30

0.3

-0.361

10

0.1

-0.230

0.09

-0.217

0.01

-0.046

100

1.00

-1.201

Shannon-Weiner Index
The most important source of error in this index is
failing to include all species from the community in the
sample (important assumption, though rarely met).
Thus, a plant community ecologist must carefully
evaluate how well their community has been sampled.
We will look at various ways to do this later.
Values of the Shannon diversity index for real
communities typically fall between 1.5 and 3.5.

H' = 1.201

Shannon-Weiner Index
The Shannon index is affected by both the number of
species and their equitability, or evenness.

Evenness
We can compare the actual diversity value to the maximum
possible diversity by using a measure called evenness.
The evenness of the sample is obtained from the formula:

A greater number of species and a more even distribution

BOTH increase diversity as measured by H'.
The maximum diversity (H max) of a sample is found when
all species are equally abundant. H max = ln S, where S is the
total number of species.

Evenness = H'/H max = H'/lnS

By definition, E is constrained between 0 and 1.0. As with
H', evenness assumes that all species are represented within
the sample.

Brillouin Index

Brillouin Index

When the randomness of a sample cannot be guaranteed,

the Brillouin Index H B is preferable to the H':

HB =

lnN! -

ln n !
i

where N is the total number of individuals and n i is the

number of individuals in the ith species.
A worked example follows...

HB =

Species
1

No. Individuals
5

ln ni !
4.79

2
3

5
5

4.79
4.79

4.79

5
S=5

5
N = 25

4.79
= 23.95

lnN! -

ln n ! =
i

ln 25! - 23.95
= 1.362
25

Evenness
Evenness for the Brillouin Index is estimated as:

E=

HB
H Bmax

where H Bmax represents the maximum possible Brillouin

diversity, that is, a completely equitable distribution of
individuals between species.
In our example, we had complete equitability, therefore,
H Bmax = H B \ E = 1.0.

Species Abundance Models

One of the earliest observations made by plant ecologists
was that species are not equally common in a given
community. Some were very abundant, other were
uncommon.
A graphical way was sought to describe this pattern, and so
arose species abundance models.
These models are strongly advocated among some ecologists
because they emphasize abundance while utilizing species
richness information and therefore provide the most
complete mathematical description of the data.

Diversity Indices
As you have probably figured out, the choice of a
particular index is chosen with respect to the goals of the
study (emphasis on abundant vs. rare species) and to what
extent sampling can be assured to be random.
There are other factors that come in to play, but these are
the three most widely used measures of diversity that
incorporate both richness and evenness into the
determination.
Note: There is generally NO relationship between one
index and another.

Species Abundance Models

A species abundance model is generated by graphing the
abundance of each species against its rank order abundance
from 1 = highest to N = lowest.
One of four distributions usually arise:
Log normal distribution
Geometric series
Logarithmic series
McArthur's broken stick model

Species Abundance Models

Species Abundance Models

(Whittaker Plots)

(Changes through succession - Bazzaz 1975)

Species Abundance Models

There are a variety of mathematical methods used to fit
these models, but that will be deferred to a subsequent
lecture.
The ability to model the data permits one to determine
how close the abundance model fits the data and
whether one community differs from another in its
adherence to a particular distribution model.
For now, be able to construct a species abundance plot
and interpret the findings in a general fashion.