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Community Ecology – Species Abundance and Diversity
LSM2251 Ecology and Environments Li Daiqin National University of Singapore

Objectives - After studying this lecture you should be able to
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describe the properties that define the structure of community
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the concepts of species richness, relative abundance or evenness, and diversity

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know how to study species diversity start thinking about the processes that influence the structure and dynamics of communities

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Outline
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Introduction Species diversity Species abundance
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Lognormal distribution

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Quantitative index of species diversity Environmental complexity Disturbance and diversity
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Intermediate disturbance hypothesis
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Introduction
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Habitat: The place where an organism lives Community: A collection of all interacting species located in a particular geographical area (habitat). Guild: Group of organisms that all make their living in the same fashion.
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e.g. seed eating animals in the desert.

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Life form: Combination of structure and growth dynamics.
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e.g. tree, shrubs, forbs, grasses

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Introduction, cont’d
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The central issue in community ecology
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What factors are most important in structuring a community -- in determining its species composition and abundance of species present?

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Community structure includes attributes such as number of species, the relative abundance of species, and the kinds of species
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Introduction, cont’d
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The factors that shape community structure
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interactions between climate and topography the kinds and amount of resources over the time individuals’ adaptive traits species in the habitat interact overall patterns and actual history of abiotic an biotic environment

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Species Diversity
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Species diversity component
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Species richness (S): the total number of species present in a community Relative abundance or evenness: a measure of the apportionment or allotment of individuals among species
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not all species are equally abundant! Why?

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Species Diversity, cont’d
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Measurement of diversity has typically focused on:
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Number of species present Relative abundance of these species Some combination of number and abundance

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Species Diversity, cont’d
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Species richness (S)
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What is it?
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The total number of species present As simple as making a list…

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How to measure it?
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Global patterns of species richness (next lecture) Commonly used, but there is one major problem:
No. of individuals of species 1 99 50 No. of individuals of species 2 1 50 Species richness (S) 2 2 Total No. of individuals 100 100
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Example 1

Community A Community B

Species Diversity, cont’d
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Species richness is highly susceptible to sample size
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You will find more species if you sample a larger area
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implies that only counts of species from equal areas are directly comparable some (not all) of this effect is a consequences of having examined more individuals

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Species Diversity, cont’d
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When numbers don’t tell the entire story...
Consider two communities, each with same 5 species, compared with equal samples of 25 individuals -- which is more diverse? Example 2:
!! "! "! "! "! "! "! Community species 1 species 2 species 3 species 4 species 5 Total 5 A 21 1 1 1 1 25 B 5 5 5 5 5 25
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Species Diversity, cont’d – Species evenness

Species Diversity, cont’d – Species evenness
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When numbers don’t tell the entire story...
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Abundance patterns may complicate the determination of richness Note that a species-poor, but ”even” community may appear richer than a species-rich, but “uneven” community at small sample sizes Both species richness and abundance should thus be incorporated in measures of species diversity
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Species abundance
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Species Abundance

Commonness or the number of species and the number of individuals of each species in a community The proportion that a particular species contributes to the total abundance of a community
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Most concerned with relative abundance of species
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Question:
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e.g., for species A, the relative abundance is 0.25 (25 individuals/100 individuals for 5 species)

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What can you see?
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“What can I see (a pattern of the abundance of species) if I go out to study a community and quantify the abundance of species within a group of organisms in question such as spiders, butterflies, shrubs, or birds?” There are regularities in the relative abundance of species in communities that hold regardless of the ecosystem.
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Species Abundance, cont’d – The Lognormal Distribution
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Preston (1948, 1962a, b) graphed abundance of species in collections as frequency distributions.
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Lognormal distributions
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Bell-shaped, or normal curves.
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In most lognormal distributions, only portion of bell-shaped curve is apparent. #! Sample size has large effect. #! Significant effort is needed to discover rare species.

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Most species are moderately abundant and few are very abundant or extremely rare

Lognormal distributions (bell-shaped) of (a) desert plants, and (b) forest birds. Note that the x-axis is on a log2 scale.
Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5rd ed. McGraw-Hill.

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Questions:
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So, why do smaller sample size result in only a part of the bell-shaped curve (see (a))? Why does the massive sampling efforts reveal only a portion of the bellshaped curve as well (see (b))? Is the lognormal distribution important? Why?

Effect of sample size on the lognormal distribution

Sample-based rarefaction curves of the butterfly communities of forest reserves (FR), forest fragments (FF), urban parks adjoining forests (PF), and isolated urban parks (IP) in Singapore using EstimateS version 6.0b (Data from Koh & Sodhi 2004)

Koh & Sodhi. 2004. Ecol. Appl. 16: 1696-1708

Lognormal Distribution
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How does this lognormal distribution occur? - does it reflect important biological processes? May (1975) proposed lognormal distribution is a statistical expectation - a product of various random environmental variables acting on many species within a community. Sugihara (1980) suggested that lognormal distribution is a consequence of the species within a community subdividing niche space.
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Quantitative Index of Species Diversity
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A variety of diversity indices in two broad categories:
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Dominance indices
Berger & Parker index: DBP = Nmax/N; where Nmax = the number of individuals in the most abundant species
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Community A in example 2: DBP =21/25 =0.84 Community B in example 2: DBP =5/25 =0.2 While community B is more diverse than community A, it has a lower value of DBP. Thus, to express greater diversity, we usually use D = 1/ DBP. Advantage: very easy to calculate Disadvantage: ???

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Simpson’s index: Ds = ! [(ni(ni - 1))/(N(N - 1))]; where ni = number of individuals in the ith species, N = total number of individuals in the community
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Like Berger & Parker index, the higher the value of Ds, the lower is the diversity. So, Simpson’s index is usually expressed as 1/ Ds or 1- Ds . Advantage: use information from a broader array of species, thus more accurate. Disadvantage: many rare species with one individual (very common) has no contribution to diversity index value. Limited use in conservation biology. Why?

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Common species influence estimates of diversity from dominance indices more heavily than do rare species.
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Quantitative Index of Species Diversity
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A variety of diversity indices in two broad categories:
Information-statistic indices
Brillouin index: HB = [ln (N!) - ! ln (ni!)/N] Shannon index: One of the most widely used indices

s H’ = - ! (pi)(log pi) i=1 Where: H’ = Shannon diversity index pi = the proportion of the ith species i = ith species S = number of species
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Quantitative Index of Species Diversity, cont’d
Example 1: Brillouin index
Species (i) 1 2 3 4 5 Total S= 5 Abundance 5 5 5 5 5 N = 25 ni! 120 120 120 120 120 ln(ni!) 4.79 4.79 4.79 4.79 4.79 ! ln(ni!) = 23.95

Therefore, HB = [ln(N!) - ! ln(ni!)]/N = [ln(25!) – 23.95]/25 = 1.362

Quantitative Index of Species Diversity, cont’d
Example 2: Shannon index
Species (i) 1 2 3 4 5 Total 5 Abundance 50 30 10 9 1 100 pi 0.5 0.3 0.1 0.09 0.01 1.00 pi ln pi -0.347 -0.361 -0.230 -0.217 -0.046 -1.201

Therefore, H’ = -! pi ln pi = - (-1.201) = 1.201. Note that even the rare species with one individual, species 5, contributes some value to the index. Think about what if an area has many rare species?

Quantitative Index of Species Diversity, cont’d
Example 3: Shannon index vs Brillouin index

Species 1 2 3 4 5 H’ HB

Individuals in sample 1 3 3 3 3 3 1.609 1.263

Individuals in sample 2 5 5 5 5 5 1.609 1.362

Quantitative Index of Species Diversity
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Information-statistic indices
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Take into account rare species Difference between Brillouin index and Shannon index:
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Shannon index does not change with species abundance (example above) while Brillouin does. #! Brillouin index discriminates between two communities with the same number of species and their proportional abundance while Shannon index does not, thus better discriminant ability. #! Brillouin index have some real advantage over Shannon index and may be preferably in conservation biology. Calculation: Shannon index is relatively easy.

With at least 5 indices (many others), so which index should we use? Take a look at example 4.
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Example 4: Abundance of bird species on unrestored tin-mining areas, in restored areas and in unmined pristine forest areas of Indonesia. Data from Sterling (2002)
Unrestored sites Yellow-vented bulbul Olive-backed sunbird Ashy tailorbird Spotted dove Orange-bellied flower-pecker Blue-throated bee-eater Cuckoo doves Magpie robin Less coucal Plaintive cuckoo Savanna nightjar White-headed munia Scaly-breasted munia Richard’s pipit Red-eyed bulbul Woodpecker Black-shouldered kite Striped tibabbler Large-tailed nightjar Eastern marsh harrier Scarlet-backed flower pecker Species richness (S) 7 5 2 1 1 0 2 0 0 0 3 3 0 2 0 0 0 0 0 0 0 9 Second-year restored sites 11 10 3 4 3 4 0 0 1 0 4 4 0 0 0 0 0 0 0 0 0 9 Unmined 44 12 20 8 8 5 4 13 5 7 0 0 0 0 6 4 4 4 0 1 1 16

Calculation of the Berger-Parker, Shannon, and Brillouin indices for the data in Example Data from Sterling (2002) Unrestored sites Second-year restored sites 4.0 7.76 2.02 1.88

Unmined 3.32 7.91 2.32 2.16

Berger-Parker (DBP) Simpson (Ds) Shannon (H’) Brillouin (HB)
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3.72 8.83 1.81 1.62

same general trend that restored sites are more diverse than unrestored sites. The value of Simpson’s index does not indicate this trend, showing that, on occasion, different indices can give different results. Results vary according to the particular distribution of individuals among species.

Quantitative Index of Species Diversity, cont’d - “so which index should I use?”
A comparison of the effectiveness of different diversity indices
Sensitivity to sample size High Moderate Moderate Low Low Biased toward richness (R) or dominance of a few species (D) R R R D D

Index Species richness (S) Shannon (H’) Brillouin Simpson Berger-Parker

Discriminant ability Good Moderate Moderate Moderate Poor

Calculation Simple Intermediate Complex Intermediate Simple

Widely used Yes Yes No Yes No

depends on the question asked!

•! There is no one single best diversity index. One’s choice of index often

•! Don’t mix diversity indices when comparing communities - you cannot compare community using a Shannon index for one and a Simpson’s index for another - must use one index throughout.

Quantitative Index of Species Diversity
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Diversity indices may be used to compare species diversity
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within a community ("-diversity) between communities (#-diversity) among communities over a wide geographical area ($-diversity)
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Rank Abundance Curves
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Measuring whole communities by just one statistic of diversity index may lose much valuable information. Plotting relative abundance of species (usually on a logarithmic scale) against their rank in abundance within a community - alternative way of measures. A rank abundance diagram can be drawn for the number of individuals, biomass, ground area covered (for plants), and other variables.
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Higher species richness and greater evenness indicated by lower slope.
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Rank Abundance Curves, Cont’d

Rank Abundance Curves of Caddisflies (Insecta: Trichoptera) from northern Portugal

Rank Abundance Curves of Caddisflies
Coastal ponds at Mira containing 18 species, while mountain stream at Relva having 79 species
Caddisflies are aquatic insects. More caddiflies are in a stream, the better quality of water is.

Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5th ed. McGraw-Hill.

Environmental Complexity
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In general, species diversity increases with environmental complexity or heterogeneity. MacArthur (1958) found warbler diversity increased as vegetation stature increased.
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Measured environmental complexity as foliage height.

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Many studies have shown positive relationship between environmental complexity and species diversity.
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Forest Complexity and Bird Species Diversity

Foliage Height Diversity and Bird Species Diversity

Heterogeneity and Diversity of Tropical Forests
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Jordan (1985) studied the relationship between vegetation and soils in Amazon forest and concluded that tropical forest diversity is organized in two ways:
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Large number of species live within most tropical forest communities. There are a large number of plant communities in a given area, each with a distinctive species composition.

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Soil and Topographic Heterogeneity and Diversity of Tropical Forest Trees

Molles, M.C. Jr. 2010. Ecology: concepts and applications, 5th ed. McGraw-Hill.

Plant Species Diversity and Increased Nutrient Availability - Negative Relationship

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Repeatedly observed negative relationship between nutrient availability and algal and plant species diversity. Adding nutrients to water or soils generally reduces diversity of plants and algae.
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Reduces number of limiting nutrients.

Disturbance and Diversity
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Disturbance difficult to define as it involves departure from “average conditions.”
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Average conditions may involve substantial variation. Discrete, punctuated, killing, displacement, or damaging of one or more individuals that directly or indirectly creates an opportunity for new individuals to be established.
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Sousa (1979) defined disturbance:
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Disturbance and Diversity
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White and Pickett defined disturbance:
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Any relatively discrete event in time that disrupts ecosystem, community, or population structure and changes resources, substrate availability, or the physical environment. Two major characteristics:
Frequency !! Intensity
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Intermediate levels of disturbance maximize the number of organisms from different successional stages

Intermediate Disturbance Hypothesis

Intermediate Disturbance Hypothesis
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Connell (1975, 1978) proposed disturbance is a prevalent feature that significantly influences community diversity.
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Proposed both high and low levels of disturbance would reduce diversity.
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Intermediate levels promote higher diversity.
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Sufficient time between disturbances allows wide variety of species to colonize, but not long enough to allow competitive exclusion.

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Disturbance and Diversity in the Intertidal Zone
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Question: how does disturbance (ocean waves generated by winter storms) affect on diversity of algae and invertebrates growing on boulders in the intertidal zone.
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Sousa, W.P. (1979). Ecology 60: 1225-1239.

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Prediction: level of disturbance depends on boulder size.
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Large boulders require more force to move, thus less disturbance; smaller boulders are turned over more frequently, thus high frequent disturbance.

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Disturbance and diversity of marine algae and invertebrates on intertidal boulders

Boulders subject to intermediate levels of disturbance support greatest diversity of species.

Applications: Disturbance by Humans
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Human disturbance is an ancient feature of the biosphere. The effects of human disturbance fall within the framework of the intermediate disturbance hypothesis. Human disturbance threatens many species with extinction. Moderate human disturbance may also increase species diversity.

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Figure 16_22

Changes in number of plant species and coverage by the grass Brachypodium pinnatum following abandonment of chalk grassland in Netherland

Applications: Disturbance by Humans
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Whether could be declines in grassland diversity be reversed using traditional mowing?
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Bobbink and Willems (1987, 1991). Normal autumn mowing and summer mowing

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Results:
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Summer mowing: biomass of grass decreased 77% to 34%; number of plant species increased from 15.6 to 21.2/0.25m2 Autumn moving: grass biomass remained at 75-80%; number of plant species remained low. Four different mowing: (1) autumn; (2) later summer; (3) early summer; or (4) early summer and autumn. Results: two mowing greatly reduced grass biomass.

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How do summer and autumn mowing differ in their effects on grass?
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Why is autumn mowing not sufficient for restoring the diversity of plants?

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Take-home message
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Most species are moderately abundant; few are very abundant or extremely rare. A combination of the number of species (i.e., species richness) and their relative abundance (or species evenness) defines species diversity. Lower slope of rank-abundance curves indicates higher species evenness or richness Species diversity is high in complex environments. Intermediate level of disturbance promotes higher species diversity.
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Study Questions
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What are a community’s properties? How to describe a community? Why do many studies use specie richness to describe and compare communities while others not? Can you come up with a complete list of species for any particular area of substantial habitats anywhere? Why or why not? Why have so many studies of species diversity concerned on one or a few groups of relatively well-known organisms such as bird, butterflies, mammals, or plants? Distinguish between species richness and evenness. Discuss how each of these components of species diversity contributes to the values of Berger-Parker index, Simpson’s index, Shannon index and Brillouin index? Do they also influence the shape of rank-abundance curves? How? If you do a quick search at ecological journals (e-journals), you will find that in these days (say, after 2005), there are not many researchers who use species diversity indices for their community studies. Can you find out what methods are currently used to compare the diversity of communities? What are the differences between lognormal distribution of species and rankabundance curves? What can they tell you about a community? How sample efforts affect them? Do they have any implication in conservation biology? How does environmental structure affect species diversity? How does the intermediate-disturbance hypothesis influence the conservation movement if it is correct? Should conservationists promote disturbance to maximize diversity?
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