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.