MBD 3

Evenness and Abundance
Models
James A. Danoff-Burg
Dept. Ecol., Evol., & Envir. Biol.
Columbia University
Today: Evenness and
Abundance Models
Evenness – a review
Evenness – new issues
Introduction to the Models
Geometric Series
Log Series
Log-Normal Series
Broken-Stick Model
All series / models will have some associated

worked examples and computer time
Lecture 3 – Evenness & Species Abundance Models © 2003 Dr. James A. Danoff-Burg, jd363@columbia.edu
Diversity of Diversities
Difference between the diversities is usually one
of relative emphasis of two main envir. aspects
Two key features
 Richness
 Abundance – our emphasis today
Each index differs in the mathematical method of
relating these features
 One is often given greater prominence than the other
 Formulae significantly differ between indices
Evenness
Definition of Evenness
 How equally abundant are each of the species?
 A simple way to combine abundance and richness
Rarely are all species equally abundant
 Some are better competitors, more fecund, more
abundant in general than others
Evenness increases diversity
Increasing evenness greater diversity
 True for all indices
S=4 S=4
N=8 N=8
Higher
Evenness,
Diversity
Site 1 Site 2
Evenness as an Indicator
For many ecosystems, high evenness is a sign of
ecosystem health
 Don’t have a single species dominating the ecosystem
 Often invasives dominate
 Paradox of enrichment
• E.g., polluted / enriched Lake Okeechobee, Florida
 Disturbed areas are mostly edge species
• Simple biodiversity
• Dominance of a few species ecologically, numerically
Evenness Across Locations
 Between ecosystem comparability is usually not possible
 Some areas have lower biodiversity naturally than others
• Tiaga is naturally much less even than the deciduous forest
• Tiaga is often dominated by a single species (e.g., Blue Spruce)
 Seasonality may confound the comparison as well
• Earlier in temperate growing season, less even than later
 This is a general principle for most all indices this term
 When would you want to compare across locations?
 Trying to prioritize areas for conservation
 Based largely on biodiversity (not ecol. uniqueness)
Today: Evenness and
Abundance Models
Geometric Series
Log Series
New Ideas on Evenness
Types of evenness
Consequences of those types of evenness (a.k.a.
species abundance models)
Methods of testing and evaluation
Introduction to each series
Types of Evenness
 Types of evenness patterns are called Species
abundance models
 Have four main types of abundance models
1. Geometric series
2. Log series
3. Log-normal series
4. Broken stick
 Decreasing dominance of a single species from
#1 to #4
 Possibly both numerical and ecological dominance
Today: Evenness and
Abundance Models
Geometric Series
Log Series
Hypothetical Model Curves
100
How would this
appear if:
Minimal Evenness (one species)
10
Maximal
1
Maximal Evenness evenness?
Per
Species Minimal
Abundance 0.1
evenness?
0.01
Next to Minimal Evenness (two species)
0.001
10 20 30 40
Species Addition Sequence
100
10 Broken Stick Model
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
Geometric Series
0.001
10 20 30 40
Abundance Models
Use all the available data
 Most thorough representation of the data
Observations
 Evenness increases from Geometric  Log  Log-
normal  Broken Stick models
 Dominance of any one species decreases from
Geometric  Log  Log-normal  Broken Stick
models
 Broken stick is the closest nature gets to maximal
evenness
Other Methods of Abundance
Curves
How else could these graphs be constructed?
 How would the data thereby be interpreted?
Possible ideas:
 Biomass
 Number of species per trophic level
• Trophic level ≅ Species
• Number of species ≅ Abundance
 Number of species per feeding guild
• Feeding Guild ≅ Species
• Number of species ≅ Abundance
 Others?
Testability
Simple visual inspection of a single curve is
insufficient
How to test each abundance model differs
between each model
 Geometric – simple rank / abundance plots with
abundance on log scale on Y axis (as already seen)
 Log – frequency distribution of # of spp vs. abundance
 Log-normal – similar to Log, but use a log scale on X
axis
 Broken stick – rank / abundance plot, using ranks,
rather than abundance (more in a moment)
Today: Evenness and
Abundance Models
Geometric Series
Log Series
100
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
0.001
Geometric Series
10 20 30 40
Geometric Series
Niche pre-emption is structuring the ecosystem
 Species 1 takes a certain percentage of the resources
and prevents others from using them
• Assumes competitive exclusion and resource exhaustion
 Species 2 takes a bit more
 Continues with other species until all resources are
used and all species are included
Minimal cooperation in ecosystem
Geometric Series
Assumes that the species abundance is roughly
proportional to total resource use
 Linear increase in abundance  linear increase in
resource use
 Interspecific per-individual resource use is comparable
Mostly commonly found in species poor
communities
 Early succession
 Degraded ecosystems (Enriched, Invaded)
 Harsh Ecosystems
Testing the Geometric Series
Model – An Exercise
Should have: straight line species plot & statistical
test 100
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
0.001
Geometric Series
10 20 30 40
Worked Example 2
Magurran p. 130-131
Use the plant feeder data from one of the gardens
Work individually
Create a Rank / Abundance graph as in Fig. 2.4 of
Magurran
 Only a gross approximation of whether it actually fits
Estimate k
 Use Excel and do so iteratively
Conduct Chi-square goodness of fit test (GOF)
 Use SPSS to do this
Today: Evenness and
Abundance Models
Geometric Series
Log Series
100
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
Geometric Series
0.001
10 20 30 40
Log Series Model
Closely related to geometric series
 Some studies have found both fitting the same data
 Similar to geometric in hypotheses about origin of
community
• Arrival of species to a novel environment
• Both say that a few factors predominantly structure the
community
• Both say that one (geometric) or a few (log) species dominate a
species
 Log differs from Geometric in assumptions about arrival
• Arrivals are randomly arranged
– Can get some clumped, some long intervals between arrivals
• In geometric series, the arrivals are regular and continual
Log Series Model Mathematics
Base equations
 Best fit to a GOF test between expected & obs data
 Base form of log series: α, (αx2 / 2), (αx3 / 3), … (αxn / n)
 Observed S = α[-ln(1-x)]
 x is calculated iteratively
• Using the following equation S / N =(1-x) / x[-ln(1-x)]
• Solve S / N, then plug numbers in for x to determine its value
• 0.9 > x > 1.0
• If N / S > 20, then x > 0.99
 α (a diversity index) = N(1-x) / x
• Plug in x once obtained and get α
Testing the Log Series Model –
Worked Example 3
Procedure:
 Construct a rank abundance plot as in the geometric
• Estimate rough estimate of how well it fits to theoretical
 Determine log2 based classes (octaves = doubling abd)
 Determine numbers of each class in observed data
 Estimate x
 Solve for α
 Calculate expected abundance for each abd level
 Group into classes
 Conduct GOF test
Today: Evenness and
Abundance Models
Geometric Series
Log Series
100
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
Geometric Series
0.001
10 20 30 40
Log Normal Series
Most communities fit the Log Normal Series
 Usually large, mature communities
 E.g., temperate forest trees
Ubiquity may be b/c of simple mathematics
 Normal distribution is often a consequence of large
numbers
 Central Limit Theorem
• Large # factors  random variation will result in normal
distribution
• Central assumption behind parametric statisticss
• ↑ probability with ↑ # of factors
Log Normal Series
Species are grouped into classes
 Octaves – most common (Log2)
 Any log base can be used
16
14 2
4
12
8
Number 10 16
of 8 32
6 64
Species 128
4
256
2 512
0
Number of Individuals
Log Normal Community
Assembly
Assumption about community formation
 Sequential breaking of empty niche space (Sugihara
1980)
• Each species that arrives splits the niche space
• Occupies a niche space proportional to its relative abundance
• Probability of niche space being subdivided is independent of its
sizes
• Breakages occur successively
 Mechanism can be through an ecological or
evolutionary process
Fit to model: not necessarily supports assumption
Other Explanations
Central Limit Theorem
 Not necessarily a biological explanation (May 1981)
Ugland & Gray 1982
 Species can be divided into three abundance classes
• Rare (65%), Intermediate (25%), Common (10%)
 Communities are composed of patches
 Abundance of species = sum of abd in all patches
  enough to result in Log Normal distribution
Log-Normal Miscellany
Missed species
 Very rare species will not be sampled
 Those less abundant than the critical number are beind
the veil line
 Need to estimate how many there should be there
 Smaller the sample  increased number behind veil
line
• Because have a higher veil line, relative to larger samples
Simplicity of calculations
 Would be there, but for the veil line
 Pielou (1975) created a fit to truncated log normal
Testing the Log-Normal Series
Model – Worked Example 4
 Use Pielou’s Truncated Log Normal process
 Estimating # of spp missed to the left of veil line
 Process (14 Steps!)
 Sort species, from most to least abundant, ln transform
 Calculate mean and variance of community
 Determine observed class abundance
 Calculate Gamma & Sobs
 Estimate Theta
 Calculate Mu, Vx, zo
 Lookup po of zo
 Estimate total S (including those behind veil line)
 Lookup po of zo of each class abundance
Today: Evenness and
Abundance Models
Geometric Series
Log Series
100
1
Per
Species
Abundance 0.1
Log-Normal Series
0.01
Log Series
Geometric Series
0.001
10 20 30 40
Broken Stick Model
Sometimes called random niche boundary
hypothesis
 “broken stick” – MacArthur (1957)
A stick randomly and simultaneously broken into S
pieces
No real relationship between earlier species
presence and niche size of subsequent arrivals
 Unlike all earlier models
Examples of fits to Broken
Stick
Successfully fit in past
 Passerine birds (MacArthur 1960)
 Minnows and Gastropods (King 1964)
In General:
 Best fit in narrowly defined communities of
taxonomically related organisms
No adequate diversity index needed if data fit
Broken Stick
 S is adequate measure of diversity
Broken Stick Model
Most equitable species abundance as ever
happens naturally
 Most biologically realistic “uniform” distribution
Theoretically, only looking at one resource
 E.g., space
Strongly subject to sample size
 Don’t have crowding limitations between species
Limitations of Broken Stick
Model
Not really applicable to a single sample
 Usually conceived of as the average spp abd.
Distribution
 Can be misleading to test fit of a single sample to theory
of equal resource partitioning
Fine to use as we will use it
 Adherence to a species abundance model
Testing the Broken Stick
Model – An Exercise
Procedure (5 steps)
 Calculate N and S
 Determine Observed species in each Log2 abundance
classes
 Calculate Expected species for each abundance level
(1-2000 or so)
 Determine Expected species in each Log2 abundance
class
Our Data This Term I
Relationship between plant biodiversity, pest
insect biodiversity, and beneficial insect
biodiversity
 Read website at
http://www.columbia.edu/itc/cerc/danoff-burg/web-
pages/gardens_main.htm
 Has a pretty good amount of background on the topic
 Field sites were in Manhattan and Brooklyn community
gardens
 Data collected during summer 2001
 I will also email you the data matrix
• Please begin looking it over so that you are comfortable with it
Our Data This Term II
Influence of Hemlock Woolly Adelgid on carrion
beetle biodiversity
 Separated by many (at least 3) trophic levels
 Adelgid is a phloem-feeding insect
 Carrion beetles are detritivores or predators on fly
larvae on carrion
 Field sites at Black Rock Brook, Black Rock Forest
 Data collected during summer 2001
 I will also email you these data
• Please begin looking over the data set
Next week:
Abundance, An Introduction
Read
 Magurran Ch 2
 Magurran Worked Examples 1-6
 Southwood & Henderson 2.1, 2.2, 13.1
We will conduct a few evenness and species
abundance models next week
Decide which of the two projects on which you are
interested in working collaboratively
 3 people per group

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Evenness and Abundance

All series / models will have some associated

10 Broken Stick Model

10 Broken Stick Model

10 Broken Stick Model

10 Broken Stick Model

10 Broken Stick Model

10 Broken Stick Model

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