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Ecology Ch 9

Learning Objec7ves
A"er studying this chapter you should be able to 1. Dis7nguish between Clements organismal concept of community organiza7on and Gleason's individualis7c concept. 2. Explain dierent methods of describing the rela7ve abundances of species in a community, such as the log-normal distribu7on and species diversity. 3. Dis7nguish between dominant species and keystone species, and understand their impacts on a community.

Learning Objec7ves
4. Explain how various processes contribute to structuring natural communi7es, such as compe77on, herbivory, preda7on and various forms of symbioses.

Group of organisms that live together at the same place and 7me and interact directly or indirectly.

Discrete vs. Con7nuous Organiza7on

OMen there is a gradual transi7on from one community to another. Abrupt transi7ons between communi7es oMen coincide with steep environmental gradients, or changes in land use by Transi7on between lake, humans. rocky, and forest communi7es. Such transi7on zones are called ecotones.

Organismal (Community Unit) Concept

Clements regarded a community as a superorganism. Rela7vely discrete borders. Succession has a predictable end-point, a climax community.

Environmental gradient

Vegeta7on recovery aMer a re usually shows a characteris7c succession of species towards the climax community.

Individualis7c (Con7nuum) Concept

Gleason proposed that a community is a coincidental assemblage of species that have similar environmental requirements. Species are distributed independently of others.

Con7nuum Along Gradients

Robert Whi[aker (1956) found that species show individualis7c distribu7on pa[erns along gradients of environmental factors, resul7ng in a con7nuous change of community composi7on.

Fig. 9.3, p. 245

Community Composi7on
Species abundance
The number of individuals of a specic species in a specic area.

Species richness
The number of species occurring in a specic area.

Species evenness
A measure of the similarity of the rela7ve abundances of species within a community.

Species diversity
An indicator of the number of species in an area taking into account their evenness.

Species Abundance
There are regulari7es in the rela7ve abundance of species in communi7es that hold regardless of the ecosystem. Preston (1948) developed concept of distribu7on of commonness and rarity.

Lognormal Distribu7on

Species Diversity
Species richness and species evenness. High species diversity
There are many equally or nearly equally abundant species.

Low species diversity

There are only a few species or only a few species are abundant.

Informa7on theory
A measure of uncertainty (rarity of species)


Community 1

Community 2

Community 3

2 2 2 2 2

1 2 1 5 1

8 1 1 0 0

Using informa7on theory

High species diversity
uncertainty, probability that the next individual selected randomly will be the same as the last.

Low species diversity

uncertainty, probability that the next individual selected randomly will be the same as the last.

Quan7ta7ve Index of Species Diversity

Shannon-Wiener Index:

H' =

pi ln pi

H' = value of SW diversity index. pi = propor7on of the ith species. ln = natural logarithm of pi (ln on your calculators). s = number of species in community.

ln pi

pi ln pi

pi ln pi =

ln pi

pi ln pi

pi ln pi =

Rank Abundance
Easy to compare diversity by using rank abundance curves. Rela7ve abundance of species plo[ed against their rank in abundance. Greater evenness indicated by lower slope. Line extending further to the right means higher species richness.

MacArthur and MacArthur (1961)

Keystone Species
Keystone species in communi7es are those that have a dispropor7onately large eect on the community. Beavers and starsh are good examples. Compare with Dominant species being the most abundant species in a community.
One can be assigned per trophic level.

Paine (1966) wanted to determine if Pisaster was the keystone species.

Pisaster removed
Removal experiment (control + experimental plots)
Experimentally removed starsh and showed that it led to change in community composi7on of the lowest trophic level. The starsh maintained a high diversity of species by suppressing the most compe77ve species (bivalves); when it was removed, the diversity of prey plummeted.

Beavers as Keystone Species

Beavers are considered keystone species because of their prominent role as ecosystem engineers. Such species maintain or create substan7al habitats for other species.

Interac7ons between organisms is the key subject area in community ecology. These interac7ons inuence the presence, and abundance of species and structures the community. Herbivory Preda7on Compe77on Disease Symbiosis o Mutualism o Parasi7sm o Commensalism

Summary of Interac7ons
Type of interacCon Preda7on/ Herbivory Compe77on Mutualism Commensalism Parasi7sm Species A Species B

+ - + + +

- - + 0 -

Preda7on and Herbivory

Exploita7on: Interac7on between popula7ons that
enhances tness of one individual while reducing tness of the exploited individual.

Kill and consume other organisms.

Consume live plant material but usually do not kill plants.

Plants usually remain alive, even though they are harmed. Hawkes & Sullivan (2001) conducted a meta-analysis of results from 81 studies of herbivory eects on plants. In general, herbivory reduced plant growth and reproduc7on
Results varied among species

Herbivorous Stream Insect and Its Algal Food

Lamber; and Resh studied inuence of caddisy (Helicopsyche borealis) on algal popula7ons on which it feeds.

On the 7les with no exclusion

H. borealis

An Alterna7ve Reac7on to Herbivory

Snow goose grazing in a salt marsh
Jeeries documents snow goose herbivory across a sub-arc7c salt marsh containing two main species: Puccinellia and Carex. What is the eect of snow goose grazing on these two plant species?

Standing biomass

Herbivory caused increased produc;vity, despite reduced standing biomass; this is overcompensa7on.

Overall biomass

What causes overcompensa7on?

1. Dierences in grazing intensity.
2. Defeca7on from grazers provides nutrients. Hik & Jeeries (1990) conducted an experiment on snow geese, nding that these factors could aect produc7vity under dierent circumstances.
Heavy grazing led to no recovery Removing feces (fun!) led to lower growth

Cycles of Abundance in Snowshoe Hares and Their Predators

Snowshoe Hares (Lepus americanus) and Lynx (Lynx canadensis).
Extensive trapping records.

Popula7on Fluctua7ons

Snowshoe Hares: Role of Food Supply

Live in boreal forests dominated by conifers. In winter, browse on buds and stems of shrubs and saplings such as aspen and spruce. Peaks in snowshoe hare popula7ons coincide with food shortages.

Snowshoe Hares: Role of Predators

Lynx (classic specialist predator) Preda7on can account for 6090% of mortality during peak densi7es. Complementary:
Hare popula7ons increase, causing food supplies to decrease. Starva7on and weight loss may lead to increased preda7on, all of which decrease hare popula7ons.


Both food and preda7on cause cycles in snowshoe hare popula7ons.

Predator Sa7a7on by Periodical Cicadas

Periodical cicadas Magicicada spp. emerge as adults every 1317 years.
Densi7es can approach 4,000,000 ind/ha

Homoptera Nymphs A marvel of nature, appears aimed at predator sa7a7on.


Modes of Compe77on
Direct aggressive interac7on between individuals.

Compe77on to secure resources rst.

Compe77on between individuals of the same species.

Compe77on between individuals of two dierent species.

Interference Compe77on Among Song Sparrows

Arcese studied how male song sparrows defend their territories on Mandarte Island (BC). There are more males than territories, so 20% of them are oaters among already-occupied territories. Floaters were more likely to challenge territorial males if they were young, old, diseased, or injured, but not males in their prime.

Interference Compe77on Among Song Sparrows

Lotka-Volterra Model
Eect of intraspecic compe77on on popula7on growth: dN / dt = rmaxN [(K N )/ K]
As N increases so does intraspecic compe77on and this slows popula7on growth.

Lotka-Volterra Model
Eect of interspecic compe77on on popula7on growth of each species:
dN1 / dt = rmax1N1 [(K1 - N1 - 12N2) / K1] dN2 / dt = rmax2N2 [(K2 - N2 - 21N1) / K2 ] 12: eect of individual of species 2 on rate of pop. growth of species 1. 21: eect of individual of species 1 on rate of pop. growth of species 2.
Assumes equal compe77ve ability within a species

Lotka-Volterra Model
In general, LV predicts coexistence of two species when, for both species, interspecic compe77on is weaker than intraspecic compe77on.

Compe77ve Exclusion Principle

Gause (1934) thesis
two species with similar ecology cannot co-exist in the same region complete compe7tors cannot coexist

Connell Mapping Study

Mapped all the barnacles growing in certain regions of the inter7dal zone Divided each study area in half and cleared all the Balanus o as they established themselves in half of the region Periodically re-mapped

Chthamalus has a large fundamental niche
Middle and upper inter7dal zones

In the presence of Balanus, Chthamalus has a realized niche that is more limited
Upper inter7dal zone

Resource ParCConing

Mechanism of co-existence
In nature, species compete but s7ll co-exist SpaCal heterogeneity facilitates co-existence
Patchy distribu7on of resources in landscape Not compe7ng everywhere

VariaCon in compeCCve ability within a species

Compe77ve ability can vary with genotype

CompeCCve equivalence between species:

Completely equal so that outcome of compe77on not always predictable

Intraspecic Compe77on Among Herbaceous Plants

Resource Compe77on:

As a stand of trees grows, it starts with many small individuals Compe77on for limi7ng resources Climaxes with few large individuals


Intraspecic Compe77on Among Herbaceous Plants

Logarithm of dry weight per plant

Symbiosis: Mutualism
Symbiosis is an in7mate rela7onship between two organisms. In mutualism, both partners benet. Faculta7ve mutualism Species that can live without their mutualis7c partner Obligate mutualism Species that can not live without their mutualis7c partner

Obligate Mutualism: Ants and Bullshorn Acacia

Herbivores a[emp7ng to forage on acacia plants occupied by acacia ants are met by a large number of fast, agile, aggressive defenders. Ant Benets:
Thorns Sugar and liquid Bel7an bodies

Plant Benets:

Sucker growth and survival

Janzen (1966)

Experimental Evidence For Mutualism

Plant Performance and Mycorrhizal Fungi The rela7onship

Mycorrhizal fungi give plants greater access to inorganic nutrients (P, water), mycorrhizae feed o the root exudates of the plant (sugars, aas)
Ectomycorrhizae (ECM)

Humans and Mutualism

Honey gatherers and Honeyguides African cultures Honeyguides
Indicatoridae Feed on waxes (beeswax)
Greater Honeyguide (Indicator indicator)

Guiding Behaviour and Myth

Dos Santos (1569) First no7ced they would eat the candles in the churches Noted that local people follow the birds to get honey Mutualis7c rela7onship
Birds lead humans to honey Humans give the birds access to wax

If you do not leave anything for the guide (honeyguide), it will not lead you at all in the future. If you do not leave anything for the guide, it will lead you to a dangerous animal the next 7me.

Watch HoneyGuide

Isack and Reyer (1989) Science

Isack and Reyer (1989) Science