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Ecological communities

Daijiang Li

LSU

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Announcements

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Even though I have separate sections on Competition and Predation in the course, I don't have one on Mutualism. Mutualism is important ecologically, but often each mutualism is important for a different reason. Mutualisms are often very specific for the types of organisms that are involved. Because mutualism is caused by co-evolution, a lot of them have their own idiosyncrasies that don't generalize easily to a way of explaining the overall things that are common to mutualisms. This is a contrast to Competition and Predation where there are a lot of features common to the interactions between very different pairs of species.

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Levels of organization in Ecology

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What is an ecological community?

Community – collection of species that occur at the same place & time, circumscribed by natural (e.g., serpentine soil), arbitrary, or artificial (e.g., 1-m2 quadrat) boundaries

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An ecological community is a set of interacting species which utilize the same environment.

What is an ecological community?

Community – an association of interacting species inhabiting a defined area whose interactions can be + or - & direct or indirect.

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Community Ecology

Major focus

  • How are groupings of species distributed in nature? (e.g., diversity)

  • How are these groupings explained by the environment and the interactions between and among species?

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Biodiversity

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Calculating Biodiversity

There are multiple ways to calculate biodiversity:

  • Species diversity
  • Genetic diversity
  • Ecosystem diversity
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Species diversity

alpha, beta, and gamma diversity

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Measuring alpha species diversity

One of the most common measurement is species richness, the number of species in a community

What is the species richness for both communities?

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Species Richness

Impossible to count all species

Need to estimate based on sampling (quadrats, plots, transects, traps, etc.)

Richness depends on number of samples collected

More samples → more species

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Species Richness

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Species Richness

Use extrapolation to predict richness

  • Want to predict the asymptote
  • Try to collect as few samples as possible
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  • with a few samples, you collect mostly common species
  • with more samples, you start to collect some of the rarer species
  • only with intensive sampling do you start to collect all species
  • unless you’ve sampled enough to reach this plateau, beyond which no new species are added, you will miss species
  • biological inventories conducted by Nature Conservancy and others rely on extrapolation
  • try to predict the asymptote. Also try to collect as few samples as possible to achieve that asymptote

Rarefaction

Individual-based and sample-based
Procedures of individual-based rarefaction
Use rarefaction to compare diversity between two sites with different sampling effort

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Rarefaction Assumptions

  • underlying community is constant over time (no immigration or emigration of species)

  • well-mixed community (spatial distribution of each species is random)

  • individuals are sampled at random (brighter colored individuals aren't sampled more often than others)

  • sampling with identical methods (if used to compare two communities)

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Approximation

E(Sm)Si=1S(1pi)m

where S is the number of species, m is the total number of individuals in the sample, and pi is the proportion of species i in the original sample.

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Species richness estimators for extrapolation

Chao1 index

Sest=Sobs+((f1)22f2)

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If a sample of an ecological community results in the following abundances of species, what is the estimated overall richness of the community? Sp A (200 individuals), Sp B (140 individuals), Sp C (98 individuals), Sp D (30 individuals), Sp E (2 individuals), Sp F (2 individuals), Sp G (1 individual), Sp H (1 individual).

A. 8
B. 9
C. 10
D. 11

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Problem of Species Richness

Which community is more diverse??

Richness is an uninformative descriptor: equal weight given to rare and abundant species

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Commonness vs. Rarity in communities

The typical pattern of abundances of different species of a single group in the same area:

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Species abundance

  • Moth data from England
  • x-axis on log2 scale;
  • Partial bell-shaped curve -- log-normal distribution
  • Very common pattern in nature
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Species Diversity

Given the commonness of rarity, how to measure community diversity?

Two different things going on:

  • Species Richness (number of species)
  • Species Abundance (relative evenness)

Two potential solutions:

  • Shannon Index
  • Rank-abundance Curves
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Shannon Index

Balancing Richness and Abundance/Evenness

H=i=1Spilnpi Where: pi = proportion of the total sample represented by species i, ln = log base e, S = the number of species in the community

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Shannon Index

H=i=1Spilnpi

Species Abundance Pi ln(Pi) Pi x ln(Pi)
Yellow 17
Orange 1
Purple 1
Brown 1
Total 20 1

H' = ?

Species Abundance Pi ln(Pi) Pi x ln(Pi)
Yellow 5
Orange 5
Purple 5
Brown 5
Total 20 1

H' = ?

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Index Higher when species more evenly distributed

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What is the Shannon Index of Diversity for the following community?
A. 0
B. 1
C. 2.83
D. 17

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Prop = 1, and ln1 = 0. Therefore, it’s 0.

Shannon Index

Weakness

  • Can’t tell if differences in measurements are due to richness or evenness (or can you?)

  • Difference between 0.58 & 1.39. What does it mean?

When all species are equally abundant: H=ln(4)=1.39: Community B has max diversity

Evenness E = H'/H'max

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Rank-Abundance Curve

Slope ≈ evenness; Length ≈ richness

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Rank-Abundance Curve

Slope ≈ evenness; Length ≈ richness.
Most species are moderately abundant; few are very abundant or extremely rare

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Beta diversity

To compare diversity between communities

Jaccard similarity:

J(A,B)=|AB||AB|

|AB| is the shared species between communities A and B, while |AB| is the total number of shared and non-shared species.

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Beta diversity

There are many other beta diversity indices

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Distance-decay in community similarity

A common observation: communities which are farther apart tend to be more dissimilar than two communities which are close together

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How have alpha and beta diversity changed over time?

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How have alpha and beta diversity changed over time?

Changes in taxonomic and phylogenetic diversity in the Anthropocene

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Community composition

Community composition describes the diversity and identity of species within a given community

keystone species is a species that has a large impact on community structure relative to it's abundance / biomass in the community (e.g., top predator)

foundation species is a species that can play a substantial role in determining what species can exist within the community (e.g., beavers, earthworms). Usually, this is a species that defines much of the structure of a community through e.g., habitat-forming

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Community assembly

Community assembly is the set of processes by which communities are formed

Whereas the figure is drawn as if these processes occur sequentially, in reality they occur more-or-less simultaneously or in any order

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Succession, Disturbance & Stress

Succession – directional change in community composition at a site (as opposed to simple fluctuations), initiated by natural or anthropogenic disturbance, stress, or the creation of a new site

Some biologists restrict the definition to directional replacement of species after disturbance

Disturbance – a discrete event that damages or kills residents on a site (and potentially creates opportunities for other individuals to grow or reproduce); either catastrophic or non-catastrophic (Platt & Connell 2003 Ecological Monographs)

Stress – factor that reduces growth, reproduction, or survival of individuals (and potentially creates opportunities for other individuals)

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Scales of Disturbance

Intensity & Severity (how much damage & death?); Frequency (how often?); Extent (how much area was affected?)

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Disturbance & Succession

Primary Succession – succession that occurs after the creation of a “blank slate,” either through catastrophic disturbance or de novo creation of a new site

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Disturbance & Succession

Secondary Succession – succession that occurs after non-catastrophic disturbance (including “old fields”)

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Primary successsion example

Primary succession along the Glacier Bay chronosequence

One of the world’s most rapid and extensive glacial retreats in modern times (so far); eliminated ~2500 km2 of ice in ~200 yr, exposing large expanses of nutrient-poor boulder till to biotic colonization

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Primary succession along the Glacier Bay chronosequence

Reconstructed patterns of stand development at several sites within the chronosequence;

intensively analyzed tree-rings

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Primary succession along the Glacier Bay chronosequence

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Primary succession along the Glacier Bay chronosequence

Species richness generally increased with successional age (a common pattern in succession studies, even though at times a mid-succession peak is observed)

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Primary succession along the Glacier Bay chronosequence

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Secondary succession

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Disturbance & Succession

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Disturbance & Succession

Henry Chandler Cowles (1869-1939)

Lake Michigan sand dunes – late 1800s

Concluded that sites on the dunes were older further inland, i.e., formed a “chronosequence” (also space-for-time substitution) from which temporal change could be inferred

Referred to distinct plant assemblages as “societies”

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Succession: superorganism or individualism?

Frederic E. Clements (1874-1945)

Radical, “superorganism” view of communities; species interact and act en masse to promote a directed pattern of community development through “seral” (intermediate) stages, ending in a “climax” community

Henry A. Gleason (1882-1975)

individualistic view of succession” in which “every species is a law unto itself”

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Our modern population-biology view derives primarily from Gleason’s conceptual model, even though Clementsian ideas of deterministic progression through seral to climax stages dominated ecological theory well into the 20th century

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Why aren't all communities predictable?

Historical contingency or priority effects

Stochasticity

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Announcements

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