Conceptos Básicos en Ecología Cuantitativa
Transcript of Conceptos Básicos en Ecología Cuantitativa
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Basic Species Diversity Concepts
Species Richness
Diversity Indices- Simpson's Index
- Shannon-Weiner Index
- Brillouin Index
Species Abundance Models
Describing Communities
There are two important
descriptors of a community:
1) itsphysiognomy
(physical structure), as
described in the previous
lecture, and
2) the number of species
present and thei r 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
proper ty 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 speciesyou encounter. This is what we call a flora.
Species Richness
Some communities are simple enough to permit a complete
count of the numbers of species presentthis is the oldes t 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 totalspecies 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.
Species Richness
While many studies include Sas a descriptive factor
associated with the community, it is largely
uninformative in as much as it does not reflect relativeabundance.
Example: suppose two communities (1 & 2) each contain
100 individuals distributed among five species (A-E):
EDCBA
111196Comm-2
2020202020Comm-1
Are these two communities equivalent?
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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 distributedamong 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
( )( )
( )( )i
S
1
n 1D
N N 1
Si
i
n
=
=
The bias corrected form of Simpson's Index is:
where n i is the number of individuals in the ith species.
SinceDs and diversity are negatively related, Simpson'sindex is usually expressed as either reciprocal or
complementary forms (1/D or 1 -D) so that as the index
goes up, so does diversity.
Simpson's Index
A worked
example for
201 trees of 5
species
assessed in
several
quadrats:
201Total
1E
20D
30C
50B
100A
No.Individuals
Treespp.
662.0196.2338.0/1/1
338.020020101...
2002014950
20020199100
===
= + + =
DorD
DS
Shannon-Weiner Index
The Shannon-Weiner Indexbelongs to a subset ofindices that maintain that diversity can be measured much
like the information contained in a code or message
(hence the name information index).
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).
The uncertainty is measured asH', the Shannon Index. A
message coded bbbbbb has lo w uncertainty (H' = 0).
Shannon-Weiner Index
The Shannon Index assumes that all species arerepresented in a sample and that the sample was obtained
randomly:S
i i
i = 1
H' = - p ln p
wherep i is the proportion of individuals found in the ithspecies and ln is the natural logarithm.
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Shannon-Weiner Index
A worked example from a community containing 100
trees distributed among 5 species:
-1.2011.001005Total
-0.0460.011E
-0.2170.099D
-0.2300.110C
-0.3610.330B
-0.3470.550A
pilnpipiAbundSpecies
H'= 1.201
Shannon-Weiner Index
The most important source of errorin this index is
failing to include all species from the community in thesample (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.
Shannon-Weiner Index
The Shannon index is affected by both the number of
species and their equitability, or evenness.
A greater number of species and a more even distribution
BOTH increase diversity as measured byH'.
The maximum diversity (Hmax) of a sample is found when
all species are equally abundant.Hmax = lnS, where Sis the
total number of species.
Evenness
We can compare the actual diversity value to the maximum
possible divers ity by using a measure called evenness .
The evenness of the sample is obtained from the formula:
Evenness =H'/Hmax =H'/lnS
By definition,Eis constrained between 0 and 1.0. As with
H', evenness assumes that all species are represented within
the sample.
Brillouin Index
When the randomness of a sample cannot be guaranteed,
the Brillouin IndexHB is preferable to theH':
i
B
lnN! - ln n !H =
whereNis the total number of individuals and ni is the
number of individuals in the ith species.
A worked example follows...
Brillouin Index
= 23.95N= 25S = 54.7955
4.7954
4.7953
4.7952
4.7951
ln ni!No. IndividualsSpecies
i
B
lnN! - ln n ! ln 25! - 23.95H 1.362
25= = =
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Evenness
Evenness for the Brillouin Index is estimated as:
B
Bmax
HE =H
whereHBmax represents the maximum possible Brillouin
diversity, that is, a completely equitable distribution of
individuals between species.
In our example, we had complete equitability, therefore,HBmax =HB\ E = 1.0.
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 whatextent 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 relat ionship between one
index and another.
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 util izing speciesrichness information and therefore provide the most
complete mathematical description of the data.
Species Abundance Models
A species abundance model is genera ted by graphing the
abundance of each species against its rank order abundance
from 1 = highest toN= lowest.
One of four distributions usually arise:
Log normal distribution
Geometric series
Logarithmic series
McArthur's broken stick model
Species Abundance Models(Whittaker Plots)
Species Abundance Models(Changes through succession - Bazzaz 1975)
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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.