Chapter 8 Biodiversity theories

Any mechanisms that promote species

coexistence in a given community are
biodiversity maintenance mechanisms, or
community assembly rules.
There are two types of mechanisms:
stabilization and equalization or niche and
neutral theories.
Connell, J. H. 1978. Diversity in tropical rain forests and coral reefs. Science
199:1302-1310.
Chesson, P. 2000. Mechanisms of maintenance of species diversity. Annu. Rev. Ecol.
Syst. 2000. 31:343-66.
Siepielski, A.M. and McPeek, M.A. 2010. On the evidence for species coexistence: a
critique of the coexistence program. Ecology 91:3153-3164.

Overview of diversity theories

1.Competition theory: the Lotka-Volterra model
2.Limiting similarity/Niche specificity
3.Habitat niche diversification
hypothesis/Regeneration niche
4.Tilmans R* theory
5.Janzen-Connell spacing hypothesis
6.Intermediate disturbance hypothesis
7.Compensatory mortality hypothesis (rare species
advantage/common species disadvantage)
8.Competition-colonization tradeof
9.Source-sink efects/Mass efect (immigration)
10.Theory of island biogeography
11.Historical and regional efects
12.Fluctuating recruitment/Storage efects
13.Recruitment limitation
14.Neutral theory

Final project:
Review 4 of these mechanisms:
Historical note (origination),
Model description
How the mechanisms contribute to species
diversity
Comparison of the mechanisms
Theoretical/empirical evidence
Discussion
2nd project:
Compare the additive and multiplicative partition of
biodiversity (Lande 1996 vs Jost 2007 papers).
Or model BCI SAD and SAR: Fit logseries and
lognormal distribution to BCI SAD and fit random
placement, logarithmic, power and logistic model to
BCI SAR.

1. The Lotka-Volterra
competition model

Population
dynamics

Exponential growth

dx
rx
dt

Densitydependence

Accelerati
ng growth

Logistic growth

dx
Kx
rx
dt
K
In the above equation, r: intrinsic growth rate and K: carrying
capacity
The evolutionary strategies of r-K species:
r species: exploit empty niches, and produce many ofspring,
low probability to surviving to adulthood.
K species: strong competitors, and invest more heavily in much

Species
competition

1. Intraspecific competition: competition between individuals of the

same species.
Scramble competition: a finite resource shared equally amongst the
competitors so that the quantity of food per individual declines with
increasing population density. Another definition is competition for a
resource that is inadequate for the needs of all, but is partitioned
equally among contestants, so no competitor obtains the amount it
needs and in extreme cases all die.
Contest competition: Contest competition is a form of competition
where there is a winner and a loser and where resources can be
completely attained or not at all. Contest competition sets up a situation
where each successful competitor obtains all resources it requires for
survival or reproduction.
2. Interspecific Competition: competition between two species that
negatively afect each others population growth rates (or fitness).
Exploitation (scramble) competition: populations depress one
another through use of a shared resource.
Interference (contest) competition: populations behaves in a way

The Lotka-Volterra competition

model
dn1
K1 n1 12 n2

r1n1

K1

dt

dn2 r n K 2 n2 21n1
2 2

dt
K2

nt 1 nt (b i d e)

Lotka-Volterra
model
n1

Lotka-Volterra Competition Model

K 2 / 21

dn1
K1 n1 12 n2

rn1

K1

dt

K1

dn2 rn K 2 n2 21n1
2

dt
K2

nt 1 nt (b i d e)

Condition for stable equilibrium (species

coexistence): Intraspecific competition stronger
than interspecific competition:

K2
21
K1

K1
12
K2

K2

K1 / 12

n2

If this condition is violated,

competitive exclusion then
happens.

2. Limiting similarity/Gauses principle

The notion behind the niche hypothesis is limiting similarity, i.e.,
there must be a limit to the similarity of coexisting competitors in
utilizing a fixed amount of resources in a community, similar
species could not coexist Gauses principle.
The principle predicts that there must be as many
niches as the number of species. However, this
prediction is not observed in nature. In tropical rain
forest and aquatic systems, there are many more
species than niches.
The concept of niche can be perceived in diferent
ways. And there are fundamental niche and
realized niche (see next slide).

Hutchinson, G. E. 1959. Homage to Santa Rosalia, or why are there so many kinds of animals?

2. Definitions of niche
The Grinnellian niche embodies the idea that the niche of a species
is determined by the habitat in which it lives. In other words, the niche
is the sum of the habitat requirements that allow a species to persist
and produce ofspring.
The Eltonian niche encompasses the idea that the niche is the role
a species plays in a community, rather than a habitat.
The Hutchinsonian niche is an n-dimensional hypervolume, where
the dimensions are environmental conditions and the resources that
define the requirements of an individual or a species to practise "its"
way of life. Hutchinson further conceptualized fundamental niche
and realized niche.

Soberon, J. 2007. Grinnellian and Eltonian niches and geographic distributions of species.

3. Habitat niche diversification

hypothesis/Regeneration niche
Habitat niche diversification can be envisioned either
in a
spatial or temporal context (regeneration niche Grubb 1977).
Habitat niche theory predicts that species richness
increases if (1) the lengths of niche axes increase
(i.e., more niche types are included) or (2) the width
of the existing niches is narrowed (Hutchinson 1959)
or species have diferent life histories (ephemeral vs
late season species).
d

Species
1

Species
2

Species

Species
S

x1

Species
3

x2

Grubb, P. J. 1977. The maintenance of species-richness in plant communities:

the importance of the regeneration niche. Biological Review 52:107-145.
May, R. M. 1973. Stability and complexity in model ecosystems. Princeton

4. R*
theory
Diagram illustrating competition for potassium and
nitrogen between two plant species, using the
graphical approach of Tilman (1982). The solid curves
labeled A and B are resource dependent zero
net-growth isoclines for Species A and B. These show
the soil solution concentrations of nitrate and
potassium at which the growth of a species exactly balances all sources
of loss and mortality to that species. CA and CB are vectors that show the
amounts of N and K consumed by Species A and B. These define the
regions in which one or the other species is the competitive winner, or in
which the species stably coexist.
Two conditions: (1) On species is more limited by one resource, and the
other species is more limited by the other resource, and (2) each species
must consume more of the resource that limits its own growth.

Tilman, D. 1982. Resource competition and community structure. Princeton University Press.
Tilman, E. A, Tilman, D., Crawley, M. J. and Johnston, A. E. 1999. Biological weed control via
nutrient competition: potassium limitation of Dandelion. Ecological Applications 9: 103111.
Murray, M. G. and Baird, D. R. 2008. Resources-ratio theory applied to large herbivores.

5. Janzen-Connell spacing
hypothesis

Janzen-Connell model (Janzen 1970, Connell 1971). The x-axis

represents distance from the focal tree, and the y-axis represents the
density of seeds falling to the ground (seed shadow) or the proportion
of seeds surviving to maturity (escape curves).
Terborgh, J. et al. 2008. Tree recruitment in an empty forest. Ecology

6. Intermediate disturbance
hypothesis

IDH is a non-equilibrium process

Frequent disturbances prevents competitive
exclusion
Mid-domain hypothesis (my alternative
hypothesis)
Sousa, W. P. 1979. Experimental investigations of disturbance and ecological succession in a
rocky intertidal algal community. Ecological Monographs 49:227-254.
Sheil, D. and Burslem, D.F.R.P. 2003. Disturbing hypotheses in tropical Forests. Trends in Ecology

6. Intermediate disturbance
hypothesis

Species richness of the 2- to 10-cm dbh tree communities in the 99 Paracou

400 m2 terra rme
quadrats, as a function of the percentage of pioneer stems (%PS) or heliophilic
stems (%HS) in the same dbh class. Species richness per quadrat [E(S40)] is
calculated using Hurlberts rarefaction
method for a standard sample size n = 40. (A) E(S40) as a function of %PS. (B)
E(S40) as a function of %HS. Upper
and
curvesD..are
regression
lines
for rain
Molino,
J.-F.lower
and Sabatier,
2001.
Tree diversity
in tropical
A validation
of (C)
the intermediate
disturbance
hypothesis.
maxima and minima in steps offorests:
5% HS
(n 5 16).
Same as (B),
but quadrats

7. Compensatory mortality hypothesis

(Rare species advantage/common species
disadvantage)
The compensatory mortality hypothesis predicts that if more of the
common species experience higher mortality, or in other words if
mortality is frequency (or abundance)- dependent in a community, then
a high number of species can be maintained (Connell 1978). The
mechanism of frequency-dependent mortality prevents the dominance
of the common species, leading to a more even species-abundance
distribution, and thus allows species coexistence.
CCT (Webb & Peart 1999): A means of including rare species and
comparing their responses directly with those of more common species
was proposed by Connell et al. (1984). If species difer in abundance,
but all share a similar density dependent response to population
abundance, a community- level compensatory trend (Connell 1978)
will emerge: species population growth rates will decline as a function
of their abundances. Hereafter, we will refer to this as a communitylevel compensatory trend (CCT). We suggest that tests of population Connell, J.H.,
Tracey, J.G. and(which
L. J. Webb,
L.J. 1984.
growth, and
level density
dependence
are
alsoCompensatory
importantrecruitment,
for assessing
scale
mortality as factors maintaining rain forest tree diversity. Ecological Monographs 54:141-164.
efects)
augment,
but
do notofreplace,
community-level
analyses.
Chesson,
P. 2000.
Mechanisms
maintenance
of species diversity. Annual
Reviews in Ecology and
Systematics 31:343-366.
Webb, C.O. and Peart, D.R. 1999. Seedling density dependence promotes coexistence of Bornean

7. Compensatory mortality hypothesis

(Rare species advantage/common species
disadvantage)

Bagchi, R. 2010. Testing the Janzen-Connell mechanism: pathogens cause

overcompensating density dependence in a tropical tree. Ecology Letters

8. Competition-colonization
tradeofdynamic model:
Metapopulation

The superior competitor is not afected by the inferior species.

The inferior can colonize only sites in which both it and the
superior are absent (the term 1-p1-p2). However, species 1 can
invade into and displace species 2 (the term c1p1p2). If the
species have identical mortality rates, any two species
equilibrium is globally stable. This occurs because species 1
grows logistically and approaches its equilibrium. Once the 1 st
These
are the
necessary
and to its
species is at equilibrium, species
2 then
grows
logistically
sufficient conditions for the
equilibrium.
stable coexistence of the two
species. When m1=m2, c2>c1
Hastings, A. 1980. Disturbance,
becausecoexistence,
c1>m1.history, and competition for space.
Theoretical Population Biology 18:363-373.
Tilman, D. 1994. Competition and biodiversity in spatially structured habitats.

9. Source-sink efects/Mass efect

(immigration)
Many species, especially rare species, are thought to not be selfsustaining, and these species are locally maintained in a given
community only by constant immigration from source populations
located outside the community (Pulliam 1998). In discretely
distributed metapopulations, immigration plays a vital role in
rescuing rare populations.
Rescue effect: Populations
on islands that are less
isolated are less likely to go
extinct because individuals
from the source population
and other islands can
immigrate and rescue the
population from extinction.

Pulliam, H.R. 1988. Sources, sinks, and population regulation. Am Nat 132:652-661.
Shmida, A. and Wilson, W.V. 1985. Biological determinants of species diversity. J

The number of species

occurring on an island
(or a habitat) is
determined by
recurrent immigration
of new species onto the
island, and recurrent
extinction of resident
species.
When the immigration
and extinction rates are
equal, the number of
species is at an
equilibrium.

Near mainland

Far mainland

Small island

Large island

S1
S2
Number of species on island

Extinction rates

Immigration rates

10. Theory of island

biogeography

11. Historical and regional

efects
It stresses the importance of historical events and regional
species pool to the assembly of species at local
communities. Species diversity can difer between
comparable habitats in regions that have diferent histories
or geographical configurations.
For example, both regional and local diferences in tree
species diversity between eastern Asia, eastern North
America, and Europe clearly reflect the isolation of North
America from Tertiary centres of tree diversification in
Eurasia and rapid late Tertiary climate cooling in the
disappearance of species from the flora of Europe.

Ricklefs, R.E. 2004. A comprehensive framework for global patterns in biodiversity.

Evolutionary processes + local environmental filter

Biogeographical origin: (a) phylogenetic
clustering due to local speciation of
allopatric clades; (b) phylogenetic
overdispersion due to allopatric speciation
of two ancestral sympatric species caused
by the same biogeographical barrier.
Ecological origin: (c) phylogenetic
clustering due to habitat filtering of
phylogenetically conserved traits; (d)
phylogenetic overdispersion due to habitat
filtering of phylogenetically convergent
traits (when sister species arise from
ecological speciation); (e) phylogenetic
overdispersion due to competitive
exclusion of related species showing
phylogenetically conserved traits. The
communities compared occur in
contrasted habitats in cases (c) and (d),
and in similar habitats in Hardy,
caseO.J.
(e).
& Senterre, B. 2007. Characterizing the phylogenetic structure of
communities by an additive partitioning of phylogenetic diversity. JEcol

12. Fluctuating
recruitment/Storage efect
The mechanism hypothesizes that if there is an asynchronous
variability in relative recruitment rates between common and
rare species, then rare species are at advantage, thereby
promoting coexistence (Chesson and Werner 1981). The strength
of this efect depends on the degree of reproductive asynchrony
and the degree to which rare species are not limited by dispersal
and maximal fecundity. The test of this hypothesis clearly
requires long-term data on production of seeds and recruitments
of seedlings and growth of saplings.
Lottery model (Connells equal chance hypothesis): Species have
equal chance to colonize empty sites, hold them against
invaders and survive any environmental adversaries. Resources
are captured at random by recruits from a larger pool of
potential colonists. The species composition at any site would be
unpredictable, depending upon the history of chance
Sale, P.F. 1977. Maintenance of high diversity in coral reef fish communities. Am
colonization
Nat 111:337-359.
Chesson, P. L. and R. R. Werner. 1981. Environmental variability promotes

13. Recruitment
limitation
This hypothesis states that coexistence is promoted by
the failure of species to recruit seedlings in all
microsites favorable for their germination, growth and
survival, so that those vacant sites could be occupied
by other species (Hurtt and Pacala 1995).

Hurtt, G. C. and S. W. Pacala. 1995. The consequences of recruitment limitation:

reconciling chance, history, and competitive diferences between plants. Journal

14. Neutral
theory

Is nature symmetric?

No, but. . . .

Photo by Cristian Ziegler

How good an approximation is it?

What is symmetric neutral theory?

Symmetric neutral theory asks: To what extent are the
properties of ecological communities due to species
similarities rather than to species diferences?
Current theory is limited to communities of trophically
similar species, like tree species in a forest, not tigers and
trees.
Symmetry is the assumption that the population dynamics
of species (demographic parameters on the per capita basis)
in such communities are statistically identicalat least to a
first approximation
Symmetric neutral theory to ecology is what neutral theory
is to population genetics: it is a null model.
The basics of symmetric neutral theory are very easy to
understand. Its parameters are few and familiar from
population biology: birth and death rates, dispersal,
speciation, and density dependence (individuals of all species
experience the same rates).

Contrast with mainstream

theory
The mainstream theoretical paradigm asserts that
ecological nature is fundamentally asymmetric. It
emphasizes the inherent uniqueness of species in
ecological communities (i. e., niche-assembly theory).
According to this largely deterministic perspective,
competing species co-exist in closed, equilibrium, or
near-equilibrium assemblages by partitioning limiting
resources through niche diferentiation.
In contrast, in neutral theory, a stochastic perspective,
communities are open, non-equilibrium assemblages
whose stability arises only from the inertia of large
numbers.
Niche theory tends to be parameter-rich, complex, and
prediction-poor, whereas neutral theory is parameterpoor, simple, and prediction-rich.

Recipe for neutral theory:

Start simply, add complexity
reluctantly, kicking and screaming,
only as needed, stir vigorously

Parameters of symmetric neutral theory

J = local community size
Metacommunity
(Size JM )

One of many
local communities
(each of size J)

JM = metacommunity size

m = probability of immigration
(dispersal) that a death in the
local community is replaced by
immigrant from metacommunity
n

= probability of a
speciation
event
per birth
b, d =
per capita
birth, death
rates
c = density dependence

parameter
(units: individuals).

How communities and the biodiversity

therein are formed and maintained?
The traditional paradigm: Environmental
filtering because diferent species have
diferent response to environmental
conditions.
Adaptation, deterministic, directional

Hubbells Rules of Community

Assembly
Random walk (drift/density dependent
drift)
Dispersal limitation (immigration)
Speciation

Random Walk (Ecological Drift): What

happens if two competing species have
identical competition ability?
n1

Lotka-Volterra Competition Model

K 2 / 21

dn1
K1 n1 12 n2

rn1

K1

dt

K1

dn2 rn K 2 n2 21n1
2

dt
K
2

nt 1 nt (b i d e)

K2

K1 / 12

n2

Drift: Coexistence of
Neutral Species

Summary: How
biodiversity is
maintained in
nature?