PATTERNS OF SPECIES RICHNESS

• species richness is the number of species

present at a site

• a simple model of species richness:

-Each species utilizes a portion n of the

available resource dimension R,
overlapping with adjacent species by an
amount o.

- ñ is the average niche breadth within the

community

- õ is the average niche overlap within the

community
a) o
n

b)

-More species, ñ smaller

- species are more specialized in their use of
resources
c)

- More species because each overlaps more with its

neighbours (larger õ)

d)

- Contain fewer species when more of the resource

continuum is unexploited

- More species because resource axis is more fully

exploited (community more fully saturated)
• if a community is dominated by
interspecific competition, the resources
are fully exploited, species richness will
depend on:

iii. The range of available resources

iv. The extent to which species are
specialists
v. The permitted extent of niche overlap

• The role of predation in regulating

species richness:

• Keep species below their carrying

capacities much of the time
• Reduce the intensity & importance of
direct interspecific competition for
resources
• Permit much more niche overlap
• Permit a greater richness of species than
in a community dominated by
competition
Richness Relationships

3. Productivity

• Productivity increase from pole to the

tropics,
• decrease at higher altitude in terrestrial
environment, deeper water levels & arid
environment

a) Productivity Species richness

• productivity leads to an increased range

of available resources

• in a community dominated by competition,

an increase in the quantity of resources
would allow greater specialization without
the individual specialist species being
driven to very low densities
b) Productivity Species richness

• eg. cultural eutrophication in lakes, rivers

• competitive exclusion

• other factors varies in parallel with

productivity, eg. disturbance

• range of resource concentrations in

productive environment less than in
unproductive environment

2. Spatial heterogeneity

• species richness increase in spatially

heterogenous environment (a greater
variety of microhabitats, a greater range of
microclimates, etc)

• richness & the heterogeneity of the abiotic

environment
• animal richness & plant spatial
heterogeneity (plant species diversity &
structural diversity)

• spatial heterogeneity arises from the

abiotic environment or from the other
biological components of the community
capable of promoting an increase in
species richness

3. Climatic variation

• temporal niche differentiation in seasonal

environments

• in a seasonally changing environment,

different species may be suited to
conditions at different times of the year

• climatic instability may increase or

decrease richness
4. Environmental harshness

• environment dominated by an extreme

abiotic factor

• any organism tolerating the harsh

environment requires a morphological
structure or biochemical mechanism which
is not found in most related species

• characterized by other features associated

with low species richness

• unproductive, low spatial heterogeneity

Species Diversity
• An expression of community structure
• a measure of the variety of different
animal & plant species of a community
• Describe the number of species within
that community & their relative
abundances
• species-abundance relationships as a
single number
• diversity indices are functions of the
no. of species & the no. of individuals
for each species
• relationship between diversity &
environmental quality
• composed of 2 components:
ix. Species richness (total no. of species)
x. Evenness (how the species
abundances are distributed among the
species)
• evenness is derived from the spread of
individuals between the species

• the equitability/evenness is a function of

the species-abundance distribution
displayed by the community

•Species diversity as a measure of

community stability & as an index of the
maturity of a community

•Indices of community structure (diversity,

evenness, richness, similarity) have been
used in biomonitoring

• all measures of community structure

depend on sample size
Measures of Species Diversity

i. Margalef’s index:

Da = s-1

log N

ii. Menhinick’s index:

Db = s
√N

Da and Db are inadequate because they do

not differentiate between the same s and N
Various diversity indices computed for hypothetical
situations (random samples) of N individuals
distributed among s species, with ni individuals in
the ith species
 Hypothetical examples

Species Abundance A B C D E
1 n1 10 29 91 100 20
2 n2 10 19 1 100 20
3 n3 10 14 1 100 20
4 n4 10 11 1 100 20
5 n5 10 9 1 100 20
6 n6 10 7 1 100

7 n7 10 5 1 100

8 n8 10 3 1 100

9 n9 10 2 1 100

10 n10 10 1 1 100

s 10 10 10 10 5

N 100 100 100 1000 100

Da 4.50 4.50 4.50 3.00 2.00

Db 1.00 1.00 1.00 0.32 0.50

Ds 0.91 0.84 0.17 0.90 0.81

H’ 1.00 0.86 0.22 1.00 0.70

H’max 1.00 1.00 1.00 1.00 0.70

J’ 1.00 0.86 0.22 1.00 1.00

iii. Simpson’s index:

If 2 individuals are taken randomly from a

community, the probability that the 2 will
belong to the same species is:

L= ∑ni(ni – 1)

N(N – 1)
• The quantity L is a measure of dominance
(the concentration of N individuals among
s species)

• A collection of species with high diversity

will have low dominance

Ds = 1 - L

Ds = 1 - ∑ni(ni – 1)
N(N – 1)
A hypothetical set of species abundance
data

Species, i Abundance, ni Relative

abundance, pi
1 50 50/85 = 0.588
2 25 25/85 = 0.294
3 10 10/85 = 0.118
s=3 N = 85

Ds = 1 - 50(49) + 25(24) + 10(9)

85(84)

= 1 – 3140/7140

= 1 – 0.44

= 0.56
 Shannon & Weaver, 1963

H’ = - ∑ (Ni/N) log2 (Ni/N)

Ni/N : the proportion of the sample comprised by

the ith species

Ni/N = Pi

Log2 Pi = K

Pi = 2K
K= Log10 Pi

Log10 2

Evenness (J’) = H’ = H’
H’max log2S

J’ should be max when all species in a sample are

equally abundant
Species N P (= N /N) P log P % Rel.
i i i i 2 i
abundance
A 3 0.029 -0.148 2.9

B 4 0.038 -0.179 3.8

C 5 0.048 -0.210 4.8

D 2 0.019 -0.109 1.9

E 1 0.009 -0.061 0.9

F 23 0.221 -0.481 22.1

G 2 0.019 -0.109 1.9

H 17 0.163 -0.427 16.3

I 6 0.058 -0.238 5.8

J 33 0.317 -0.525 31.7

K 7 0.067 -0.261 6.7

L 1 0.009 -0.061 0.9

∑ N = 104 H’ = 2.809
i
Factors promoting species diversity:

c) The time hypothesis

• Diversity is a function of time; all

communities diversify with time
• Increases in diversity due to immigration
to an area, specialization of the existing
species, evolution of new species etc
• Diversity increase during succession

b) Spatial complexity

• Environmental complexity: a more

complex & heterogenous environment –
more niches are found – the fauna & flora
more complex & diverse
• An increase in species richness may
cause a further increase in spatial
complexity as animals & plants provide
habitats for others
c) Productivity hypothesis

• diversity increase or decrease with

productivity
• increase in abundance of all resources –
overall increase in production – diversity
increase
• increase in only a small part of the total
resource spectrum – diversity decrease
(community become dominated by those
species which are competitively superior in
exploiting these augmented resources)
• in a more severe environments – abiotic
considerations have the dominant influence
on community
• in benign/stable environments – biotic
interactions play a major role