Species Diversity: ANon-Concept?
Species Diversity: A Non-Concept?
What determines the number and kinds
What determines the number and kinds
of species that occur in a particular
of species that occur in a particular
place?
place?
Why do number and kinds of species
Why do number and kinds of species
vary from place to place?
vary from place to place?
Species
Species Forest habitat
Foresthabitat Scrub Habitat
Scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity
Beta Diversity
Beta Diversity
Gamma Diversity
Gamma Diversity
8.
Species
Species Forest habitat
Foresthabitat scrub Habitat
scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity 10
10 7
7 3
3
Beta Diversity
Beta Diversity
Gamma Diversity
Gamma Diversity
9.
Species
Species Forest habitat
Foresthabitat Scrub Habitat
Scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity 10
10 7
7 3
3
Beta Diversity
Beta Diversity (F vs. S) = 7
(F vs. S) = 7
Gamma Diversity
Gamma Diversity
10.
Species
Species Forest habitat
Foresthabitat Scrub Habitat
Scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity 10
10 7
7 3
3
Beta Diversity
Beta Diversity (F vs. S) = 7
(F vs. S) = 7 (S vs. G) = 8
(S vs. G) = 8
Gamma Diversity
Gamma Diversity
11.
Species
Species Forest habitat
Foresthabitat Scrub Habitat
Scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity 10
10 7
7 3
3
Beta Diversity
Beta Diversity (F vs. H) = 7
(F vs. H) = 7 (H vs. F) = 8
(H vs. F) = 8 (F vs. G) = 13
(F vs. G) = 13
Gamma Diversity
Gamma Diversity
12.
Species
Species Forest habitat
Foresthabitat Scrub Habitat
Scrub Habitat Grassland habitat
Grassland habitat
A
A x
x
B
B x
x
C
C x
x
D
D x
x
E
E x
x
F
F x
x x
x
G
G x
x x
x
H
H x
x x
x
I
I x
x x
x
J
J x
x x
x
K
K x
x
L
L x
x x
x
M
M x
x
N
N x
x
Alpha Diversity
Alpha Diversity 10
10 7
7 3
3
Beta Diversity
Beta Diversity (W vs. H) = 7
(W vs. H) = 7 (H vs. F) = 8
(H vs. F) = 8 (F vs. W) = 13
(F vs. W) = 13
Gamma Diversity
Gamma Diversity 14
14
13.
Measuring Diversity
Measuring Diversity
Species Richness
Species Richness
Total number of species in an area
Total number of species in an area
can also be measured as biomass, basal area, % cover
can also be measured as biomass, basal area, % cover
Species Diversity
Species Diversity
Considers evenness and richness
Considers evenness and richness
Species Evenness
Species Evenness
Considers how abundance data are distributed among
Considers how abundance data are distributed among
the species
the species
A= 96
A= 96 F = 20
F = 20
B = 1
B = 1 G = 20
G = 20
C = 1
C = 1 H = 20
H = 20
D = 1
D = 1 I = 20
I = 20
E = 1
E = 1 J = 20
J = 20
14.
Measuring Species Diversity
MeasuringSpecies Diversity
Species Richness
Species Richness
The number of species in a given area (N0)
The number of species in a given area (N0)
Sample Size Issue!
Sample Size Issue!
Margalef Index Menhinick Index
Margalef Index Menhinick Index
R1 = S-1/ln(n)
R1 = S-1/ln(n) R2 = S/ n
√
R2 = S/ n
√
Where S = total number of species in area sampled
Where S = total number of species in area sampled
n = total number of individuals observed
n = total number of individuals observed
Interpretation:
Interpretation:
The higher the index the greater the richness
The higher the index the greater the richness
Example: S = 6 and n = 50
S = 6 and n = 20
R1 = 1.28
R1 = 1.66
15.
Sampling area andspecies
Sampling area and species
richness
richness
Relationship b/w sampling area and bird species richness in North America)
Measuring Species Diversity
MeasuringSpecies Diversity
Diversity Indices - Simpson’s Index
Diversity Indices - Simpson’s Index
= probability that 2 individuals selected
= probability that 2 individuals selected
at random will belong to the same species
at random will belong to the same species
=
=
i
i(n
(ni
i(n
(ni
i-1))/N(N-1)
-1))/N(N-1)
Where:
Where:
n
ni
i= total number of individuals in each species
= total number of individuals in each species
N = Total number of individuals in all species
N = Total number of individuals in all species
Interpretation:
Interpretation:
If probability is high, the diversity of sample is low
If probability is high, the diversity of sample is low
21.
Measuring Species Diversity
MeasuringSpecies Diversity
Diversity Indices - Shannon’s Index
Diversity Indices - Shannon’s Index H’
H’
H’= -
H’= -
i
i ((n
((ni
i/N)
/N) ln (n
ln (ni
i/N))
/N))
Where:
Where:
n
ni
i= total number of individuals in each species
= total number of individuals in each species
N = Total number of individuals in all species
N = Total number of individuals in all species
Interpretation:
Interpretation:
1.5 (low richness/eveness) to 3.5 (high richness
1.5 (low richness/eveness) to 3.5 (high richness
and eveness)
and eveness)
23.
Hill’s Family ofDiversity
Hill’s Family of Diversity
Numbers
Numbers
Units are given in numbers of species
Units are given in numbers of species
NO = total number of species in the sample
NO = total number of species in the sample
N1 = the number of abundant species
N1 = the number of abundant species
N2 = the number of
N2 = the number of very
very abundant species
abundant species
N1 = e
N1 = eH’
H’
(H’=Shannon’s index)
(H’=Shannon’s index)
the value of e 2.718
the value of e 2.718
N2 = 1/
N2 = 1/
(
(
=Simpson’s index)
=Simpson’s index)
25.
Measuring Species Diversity
MeasuringSpecies Diversity
Species Eveness
Species Eveness
How abundance data are distributed among species
How abundance data are distributed among species
A= 96
A= 96 F = 20
F = 20
B = 1
B = 1 G = 20
G = 20
C = 1
C = 1 H = 20
H = 20
D = 1
D = 1 I = 20
I = 20
E = 1
E = 1 J = 20
J = 20
E1
E1 Pielou’s J (1975)
Pielou’s J (1975) In (N1)/ln (N0)
In (N1)/ln (N0) Where: N1 = e
Where: N1 = eH’
H’
N2 = 1/
N2 = 1/
E2 Sheldon (1969)
E2 Sheldon (1969) N1/N0
N1/N0
E3 Heip (1974)
E3 Heip (1974) N1-1/N0-1
N1-1/N0-1
E4 Hill (1973)
E4 Hill (1973) N2/N1
N2/N1
E5 Modified Hill’s Ratio
E5 Modified Hill’s Ratio N2-1/N1-1
N2-1/N1-1
Interpretation:
Interpretation:
0 = less even, 1 = more even
0 = less even, 1 = more even
27.
Desert Lizard Diversity
DesertLizard Diversity
Lizard Species Number of
Individuals
Cnemidophorus tesselatus 3
Cnemidophorus tigris 15
Crotophytus wislizenii 1
Holbrookia maculata 1
Phrynosoma cornutum 10
Scleoporus magister 2
TOTAL Individuals 32
Number of individuals for each of 6 species
of lizards counted in a 1 hectare plot
Processes Explaining Diversity
ProcessesExplaining Diversity
Gradients
Gradients
•
• Historical Disturbance Hypothesis
Historical Disturbance Hypothesis
-
- landscape reflects historical events, not current
landscape reflects historical events, not current
environmental conditions (not in equilibrium)
environmental conditions (not in equilibrium)
Habitats catastrophically disturbed are “under saturated” in terms of
Habitats catastrophically disturbed are “under saturated” in terms of
species because there hasn’t been adequate time for adaptation and
species because there hasn’t been adequate time for adaptation and
speciation
speciation
Problems: evidence from tropics
Problems: evidence from tropics
39.
Extent of tropicsduring last glacial
Extent of tropics during last glacial
maximum
maximum
40.
Equilibrium Theories
EquilibriumTheories
Landscape is a reflection of current
Landscape is a reflection of current
environmental conditions (in equilibrium)
environmental conditions (in equilibrium)
• Productivity
Productivity
• Climate stability-Harsh habitat
Climate stability-Harsh habitat
• Habitat heterogeneity
Habitat heterogeneity
• Biotic interactions
Biotic interactions
• Large Area
Large Area
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
41.
Productivity
Productivity
What isthe link b/w productivity and
What is the link b/w productivity and
biodiversity?
biodiversity?
• Tropics 2200 g/m2/yr
Tropics 2200 g/m2/yr
• Temperate 1200 g/m2/yr
Temperate 1200 g/m2/yr
• Boreal 800 g/m2/yr
Boreal 800 g/m2/yr
Scale
Scale
• Estuaries, marshes are most productive
Estuaries, marshes are most productive
ecosystems on earth, with lowest diversity
ecosystems on earth, with lowest diversity
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
42.
Climate Stability(Harsh Habitat)
Climate Stability (Harsh Habitat)
Environments with low stability are harsher
Environments with low stability are harsher
and are less diverse
and are less diverse
Why?
Why?
Exceptions
Exceptions
• Areas with stable climate but low diversity
Areas with stable climate but low diversity
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
43.
Habitat Diversity(Heterogeneity)
Habitat Diversity (Heterogeneity)
What is the link?
What is the link?
Is it a direct relationship?
Is it a direct relationship?
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
44.
Biotic Interactions
BioticInteractions
Is speciation driven by competition in low
Is speciation driven by competition in low
lats and adaptation to physical stress in
lats and adaptation to physical stress in
high lats?
high lats?
• Exceptions: trees/plants
Exceptions: trees/plants
What about predation as a mechanism?
What about predation as a mechanism?
Circularity
Circularity
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
45.
Large LandArea
Large Land Area
Supports more individuals
Supports more individuals
Supports more species
Supports more species
Tropics? Boreal?
Tropics? Boreal?
Processes Explaining Diversity
Processes Explaining Diversity
Gradients
Gradients
46.
Diversity in TRFand Coral Reefs
Diversity in TRF and Coral Reefs
Equilibrium Viewpoint
Equilibrium Viewpoint
Stability is the major characteristic of a
Stability is the major characteristic of a
community. Following disturbance, it
community. Following disturbance, it
recovers and high diversity is maintained by a
recovers and high diversity is maintained by a
variety of mechanisms. Community reflects
variety of mechanisms. Community reflects
current conditions.
current conditions.
Non-Equilibrium Viewpoint
Non-Equilibrium Viewpoint
Communities rarely reach an equilibrium state
Communities rarely reach an equilibrium state
and high diversity results from changing
and high diversity results from changing
environmental conditions.
environmental conditions.
47.
Diversity in TRF
Diversityin TRF
•
• Janzen’s Hypothesis (1970): Biotic interactions
Janzen’s Hypothesis (1970): Biotic interactions
- host-specific herbivores
- host-specific herbivores
- seed predation
- seed predation
- canopy foliovores
- canopy foliovores
•
• Hubbell’s research (1979, 1980) to support Janzen
Hubbell’s research (1979, 1980) to support Janzen
•
• Non-equilbrium explanation (Connell 1978)
Non-equilbrium explanation (Connell 1978)
- coral reefs
- coral reefs
The Non-Equilibrium Hypothesis
TheNon-Equilibrium Hypothesis
(Connell 1978)
(Connell 1978)
Connell’s Conclusions
Connell’s Conclusions
TRF and Coral Reefs demonstrate Non-
TRF and Coral Reefs demonstrate Non-
Equilibrium Hypothesis
Equilibrium Hypothesis
Equilibrium and Non-Equilibrium are not
Equilibrium and Non-Equilibrium are not
mutually exclusive
mutually exclusive
Role of human disturbances
Role of human disturbances
53.
More Intermediate Disturbance
MoreIntermediate Disturbance
Hypothesis (Denslow 1980)
Hypothesis (Denslow 1980)
Intermediate levels of disturbance vary by
Intermediate levels of disturbance vary by
ecosystem
ecosystem
Ecosyste
m
Historic Rate of
Disturbance (years)
Prairie 2
Chaparral 30
Pine 50
Oak-HW 50-100
Spruce-Fir 1000