Chapter 1: Why and How to Study Ecology

Road Map
1. Ecology is the study of that which limits the abundance of plants and animals. We
can study ecolollgy at the level of individual behavior, populations, communities,
or ecosystems.
2. Observing natural systems affords an insight into their workings, but field and
laboratory experiments provide the most rigorous tests of ecological ideas.
3. Ecological measurement must be made at the spatial and temporal scale
appropriate to the question being asked.
1. Outline
a. Basic definitions of terms and field of study
b. Understanding the difference between observational studies, and field and
laboratory experiments
c. Spatial and temporal implications in ecological interpretations
2. What Is Ecology?
a. Study of individuals, populations, communities, and ecosystems
b. Study of interactions among and between organisms, and their
environment
c. Importance to human activities
i. Example: Aswan High Dam (Figure 1.1)
1. Located on the Nile River in Egypt (Figure 1.2)
a. Potential benefits
i.
Several years of irrigation reserves
ii.
Add 526,000 ha in arable land
iii.
Produce 10 billion kilowatts of electrical
power annually
iv. Protect country from catastrophic flood
b. Benefits to date
i.
Saved rice and cotton crops from drought
damage in 1972 and 1973
ii.
Two to three crops annually, as opposed to
one
iii.
Increased productivity and annual income
from agriculture by 200%
iv. 380,000 ha of desert are being irrigated for
the first time
c. Unexpected ecological problems
i.
Increase in the incidence of schistosomiasis
(4780%)
ii.
Decreased phytoplankton blooms and fish
harvests in the Mediterranean (e.g., Sardine

annual catch decreased from 15,000 tons to

500)
iii.
Increased need for fertilization ($100
million annually). Fertilizer production uses
most of the energy generated by the Dam.
iv. Farmers overwater their land, resulting in
salt deposition (half of the irrigated acreage
is affected by salt)
d. Proper ecological studies could have predicted
resulting ecological problems of the dam
d. Ecologistsbest-equipped scientists to study natural ecosystems
i. Investigating environmental change on the local, regional, and
global scale
ii. Reductionist analyses and experimentation
iii.
Adapted concepts from agriculture, physiology, biochemistry,
genetics, chemistry, and mathematics
iv. Analytical and portable equipment
v. Challenges for ecologists
1. Acid deposition
2. Global climate change
3. Increasing use of fertilizer results in huge N inputs to
communities
4. Increasing use of pesticides
5. Species extinction
vi. Ecology and Environmental Sciences
vii. Four broad areas of ecology (Figure 1.3)
e. Behavioral Ecology
i. How behavior contributes to survivorship, reproduction, and
population growth
ii. Example: Forest tent caterpillars (Malacosoma; Figure 1.4)
1. Reside in silken tents and defoliate trees
2. Vulnerable to predators
3. Group living would appear to be counterproductive (i.e.,
assisting predators)
4. Benefits of group living
a. Multiple-layered large silken tents
b. Multiple and stronger silk trails to food sources
c. Pheromones to attract colony mates
d. Increased probability of propagating ones own
genes
f. Population Ecology
i. Populations: Groups of individuals from a single species which can
potentially interbreed
ii. What controls the abundance of a species, need to know how
populations
1. grow

2. are limited by food, competitors, and enemies

iii.
Knowledge of population ecology
1. Prevent extinctions
2. Lessen species endangerment
3. Maximize sustainable yields
iv. Example: Exotic SpeciesCentaurea diffusa
1. Invasive Eurasian plant in Montana
a. Success not due to the absence of natural enemies
b. Species secrete allelochemicals, which kill North
American species
c. Eurasian plants evolved with Centaurea
d. Experiments proved effects of allelochemicals
(Figure 1.5)
g. Community Ecology
i. Biodiversity on Earth
ii. Interest in species richness
1. Preservation of species-rich areas
2. Linkages between species richness and community function
iii.
Emergence of Earths biological diversity as a critical issue
1. Diversity-stability hypothesis (Figure 1.6)
2. Rivet hypothesis
3. Redundancy hypothesis
4. Idiosyncratic model
h. Ecosystems Ecology
i. Passage of energy and nutrients through communities
ii. Effects of energy and nutrients on communities
iii.
Human alterations of global nutrient cycles
iv. Example: Nitrogen cycle
1. Increased fertilizer use
2. Increased biomass
3. Reduced species richness
4. Minnesota prairie study (Figure 1.7)
3. Ecological Methods
a. Development of study plan
b. Example: Study of Locust Outbreaks
i. Interaction web (Figure 1.8)
ii. Interactions which may influence populations include
1. Predators
a. Birds or other vertebrates
b. Insect parasites
c. Bacterial parasites
2. Competitors
a. Other insects
b. Larger vertebrate grazers
3. Host Plants
a. Host plant quality

iii.

b. Host plant quantity

4. Physical factors
a. Temperature
b. Rainfall
Where to begin
1. Observations and Interpretations (Figure 1.9a, Figure 1.9b,
and Figure 1.9c)
a. Statistical tests to determine significance of
relationships
b. Causation vs. correlation
2. ExperimentationPredator study
a. Hypothesis: Increased predators will decrease locust
population
b. Two study groups
i.
Treatment Group: Locusts with predators
removed
ii.
Control Group: Locusts with nothing done
predators present
c. Measurements
d. Replications (Figure 1.10)
e. Statistical TestsANOVA and t-tests
f. Means and Standard Error
3. Types of Experimentation
a. Laboratory
b. Field
c. Natural
4. Laboratory Experiments
a. Most exact regulation of abiotic and biotic factors
b. Vary only the factor of interest
c. Disadvantage: Oversimplification of the biotic
community
d. Best uses: Physiological responses of individuals
5. Field Experiments
a. Conducted outdoors
b. Manipulation of abiotic or biotic factors (e.g.,
Elimination of a competitor)
c. Disadvantage: Methods of exclusion are unlikely to
be generated by nature
6. Natural Experiments
a. Use natural perturbations to disrupt biotic
community
b. Used to follow the trajectory of the perturbation
over time
c. Results can be extrapolated to other communities
7. Strengths and Weaknesses of the Types of Experiments
(Table 1.1)

4.

5.
6.

7.

8.

9.

8. Experimental Problems
a. Logistic problems lead to low replication
i.
Which leads to a Type I error; declaring that
a hypothesis is false when in fact it is true
ii.
Low replications lead to greater standard
error (SE) values
b. Difficulty in combining studies with differences in
statistical power
c. Meta-analysisa technique for combining studies
9. Mathematical Models
a. Experiments may not be possible
b. Give valuable signposts on how natural systems
might work
c. Indicate need for further data
d. Indicate need for further observations
Spatial ScaleContinuum
a. A space occupied by an individual (behavior ecology)
b. A local patch occupied by many individuals (a population)
c. A large enough space to comprise multiple populations (a community)
d. A biogeographic scale large enough to encompass a community, and its
nutrients and energy cycles (an ecosystem)
Determining the Correct Spatial Scale (Figure 1.12)
Temporal Scale
a. Proper choice of scale, depends on phenomenon and species studied
b. Short time scale studies for behavioral responses
c. Longer time scales studies for population dynamics and ecosystem
processes
Applied Ecology: The Value of the Worlds Natural Services
a. Direct and indirect services from ecosystems (Table 1)
b. Ecosystems provide more than one service (Figure 1)
c. Worth of ecosystems (Table 2)
Summary
a. Importance of ecology in addressing human perturbations
b. Four broad areas of ecology: behavioral ecology, population ecology,
community ecology, and ecosystems ecology
c. Understanding ecological processes through the use of different ecological
methods: laboratory, field and natural experiments, and modeling
d. Investigations must be conducted at the right spatial scale
Discussion Questions
a. What is the difference between ecology and environmental science?
b. What are ecological methods? How do we apply them to ecological
questions?
c. In a local park, forest, or even in your backyard think about five ecological
questions you could ask and the information you would need to answer
them. Do your questions relate to behavioral, population, community, or
ecosystems ecology or do they cross categories?

Chapter 2: Genetics and Ecology

1. Outline
a. Species occurrence due to evolutionary past
b. Mutations and chromosomal rearrangements result in a wide variety of
species on Earth
c. Genetic variability can be measured by allozymes or DNA sequencing
d. Mechanisms for reductions in genetic variability in populations
2. Evolutionary History
a. Importance of evolutionary ecology to the discipline
b. Example: Control of penguins in the Southern Hemisphere vs. their
absence in Northern Hemisphere
i. Penguins evolved in the Southern Hemisphere
ii. Unable to migrate to Northern Hemisphere
c. South America, Africa, and Australia
i. Similar climates (tropical to temperate)
ii. Characterized by different inhabitants
1. South America: Ex. Sloths, anteaters, armadillos, and
monkeys with prehensile tails
2. Africa: Ex. Antelopes, zebras, giraffes, lions, baboons,
okapi, and aardvark
3. Australia: Ex. No native placental mammals except bats,
variety of marsupials, egg-laying monotremes, duck-billed
platypus, and the echidna
iii.
Best explanation of differences: Evolution
3. Genetic Mutation
a. Increase in number of species is primarily due to mutation
b. Two types of mutation
i. Gene or point mutation
ii. Chromosome mutation
c. Point mutation
i. Results from a misprint in DNA copying
ii. Example (Figure 2.1)
iii.
Most changes are caused by frameshift mutations
iv. An addition or deletion in the amino acid sequence usually leads to
drastic and often fatal mutations
d. Chromosome mutation
i. Four types: deletion, duplication, inversion, and translocation
ii. Order of genes is affected (Figure 2.2)
iii.
Deletion
1. Simple loss of part of a chromosome
2. Most common source of new genes
3. Often lethal
iv. Duplication

4.

5.

6.

7.
8.

1. Arises from chromosomes not being perfectly aligned

during crossing over
2. Results in one chromosome being deficient and the other
one with duplication of genes
3. May have advantages due to increased enzyme production
v. Inversion
1. Occurs when a chromosome breaks in two places. When
the segment between the two breaks re-fuses, it does so in
reverse order.
2. Occurs during prophase
Measuring Genetic Variability
a. Genetic diversity is essential to the breeding success of most populations
b. Two individuals with the same form of enzyme are genetically identical at
that locus
c. Variations in gene loci are found through searching for variations in the
enzymes (allozymes)
d. Gel electrophoresis: Technique for determining differences in allozymes
e. Example of Gel electrophoresis: Figure 2.3
Gene Sequencing
a. Another method for assessing variations in the sequence of DNA
b. Made possible through the polymerase chain reaction (PCR) technique
i. Makes millions of copies of a particular region of DNA, thereby
amplifying even minute amounts of DNA
ii. Important uses in conservation biology, and rare and endangered
species
Mutations
a. Accelerated through human-made radiation, UV light, or other mutagens
b. Rate of occurrence: one per gene locus in every 100,000 sex cells
c. Only one out of 1,000 mutations may be beneficial
d. Estimated that only 500 mutations would be expected to transform one
species into another
e. Rate of mutation is not the chief factor limiting the supply of variability
f. Variability is mainly limited by gene recombination and the structural
patterns of chromosomes
Genetic Diversity and Population Size
a. Function of population size
b. Three factors: inbreeding, genetic drift, and neighborhoods
Inbreeding Depression
a. Mating among close relatives
b. Reduced survivorship (Figure 2.4)
c. Various types of inbreeding (Figure 2.5)
d. Effects of inbreeding on juvenile mortality (Figure 2.6)
e. Effects of inbreeding on small populations (Figure 2.7)
f. Example of inbreeding: Greater Prairie Chicken (Figures 2.8 and 2.9)
g. Example of inbreeding and relation to extinction: Glanville fritillary
butterfly (Figure 2.10)

9. Genetic Drift
a. Probability of the failure to mate
i. Loss of possible rare gene
ii. Loss of genetic information for subsequent generations resulting in
a loss of genetic diversity
iii.
Small populations more susceptible to drift
iv. The rate of loss of original diversity over time is approximately
equal to 1/2 N per generation
v. Example:
1. N = 500, then 1/2 N = 0.001 or 0.1% genetic diversity lost
per generation
2. N = 50, then 1/2 N = 0.01 or 1% genetic diversity lost per
generation
3. Over 20 generations, the population of 500 will still retain
98% of the original variation, but the population of 50 will
only retain 81.79%
4. 50/500 Rule: Need 50 individuals to prevent excess
inbreeding and 500 is the critical size to prevent genetic
drift
5. Effects of immigration on genetic drift (Figures 2.11 and
2.12). Often immigration of only one or two individuals
into a population can counteract genetic drift.
10. Neighborhoods and Effective Population Size
a. Effective population size is determined on mating range
b. Individuals may only mate within their neighborhood
c. Example: Deer mice. 70% of the males and 85% of the females breed
within 150 m of their birthplaces.
d. Harem Effects
i. Even within a neighborhood, some individuals may not reproduce
ii. In a harem structure, only a few dominant males breed
iii.
Effective Population Size
1. NE = (4 Nm Nf)/(Nm + Nf)
2. Where: NE = Effective Population Size; Nm = Number of
Breeding Males; Nf = Number of Breeding Females
iv. Example of Effective Breeding Size (Figure 2.13)
11. Applied Ecology: Can Cloning Help Save Endangered Species?
a. Dolly, the cloned sheepIan Williams, 1997 (Photo 1)
b. Can this technique be used to save endangered species?
i. Need knowledge of reproductive cycle
ii. Need for surrogate females
iii.
Expense associated with cloning
iv. Cannot address genetic diversity
12. Summary
a. New species arise from the accumulation of gene and chromosome
mutations

b. Genetic variation is reduced in populations due to inbreeding, genetic

drift, and neighborhoods. 50/500 Rule.
c. Humans can acquire more individuals of wild populations, which could
counteract genetic drift
d. Effective population size can be reduced by harem mating structures or
territoriality
13. Discussion Questions
a. Small population size is detrimental to genetic variability. Why is habitat
fragmentation detrimental to populations, and can linking conservation
areas by corridors or siting them close together help alleviate this
problem?
b. We can have inbreeding depression as well as outbreeding depression
(where local populations are highly adapted to their local environment,
and outbreeding reduces fitness). By what mechanisms do you think this
works, and what implications does it have for conservation biology?
c. In 1986, the California condor had declined to only 27 individuals. Since
then, over 150 condors have been bred and 88 released back into the wild.
What genetic problems do you think might be encountered in trying to
reestablish this population in nature?

Chapter 3: Extinction
1. Outline
a. Rate of extinction
b. Causes of extinction
c. Risks confronted by endangered species
d. Characteristics of species and their relationship to extinction
2. The Extinction Crisis
a. Extinction
i. All individuals die without producing progeny
b. Pseudoextinction
i. Species disappear over evolutionary time
ii. Lineage transformed into separate lineages
c. Fossil record
i. Extinct species to living species1,000:1
ii. Average life span of a species4 million years
iii.
Average extinction rate2.5 species per year
iv. Total number of species over time10 million
v. Biased fossil record
1. Favors successful, geographically wide-ranging species
2. Persist longer than the average
3. Biased toward vertebrates and mollusks
vi. Background extinction rates are probably higher than indicated in
fossil record

1. Example: Extinction rates 10 times higher than predicted

by fossil record
a. 1 every 100 years for mammals living today
b. 1 every 50 years for birds living today
d. Present extinction rate much higher than in the past or predicted
i. Effects due to humansDistant Past
1. Correlation between human population growth and the
number of extinctions (Figure 3.1)
2. Large-scale extinctions in North and South America
coinciding with the arrival of humans (11 thousand years
ago)
a. North America lost 73% of its genera of large
mammals
b. South America lost 80% of its genera of large
mammals
3. Large-scale extinctions in Australia coinciding with the
arrival of humans (13 thousand years ago)
a. Lost nearly all of its large mammals, giant snakes,
and reptiles
b. Nearly half of its large flightless birds
4. Probable causes of these extinctions
a. Hunting
b. Some climate change
ii. Effects due to humansRecent Past
1. Devastating effects on islands
a. Hawaii4th- and 5th-century Polynesians arrived
i.
Exterminated 50 out of 100 species of
endemic land birds
b. New Zealandend of 18th century
i.
Entire avian megafauna consisting of huge
land birds was exterminated
ii.
Accomplished through hunting and habitat
destruction
c. Madagascarlast 1,500 years
i.
Exterminated
1. Giant elephant bird, largest bird ever
recorded
2. 20 species of lemur, most larger than
any surviving species
3. 2 giant land tortoises
3. Patterns of Extinction
a. Islands vs. continental areas (Table 3.1)
i. Reasons for differences in extinction rate
1. Island species may consist of a single population
a. Single climatic event can lead to extinction

2. Island species may have evolved in the absence of

terrestrial predators
a. Characteristics contributing to extinction
i.
Flightlessness
ii.
Tameness
iii.
Reduced reproductive rates
b. Ex. Hawaii (Figure 3.2)
b. Causes of extinction (Figure 3.3)
i. Introduced species effects
1. Competition
a. Not been shown to eliminate an entire species
2. Predation
a. Rats, cats, and mongooses have accounted for at
least 112 of 258 extinctions of birds on islands
(43%)
3. Disease and parasitism
a. Avian malaria in Hawaii accounted for the loss of
50% of local Hawaiian bird species
ii. Habitat destruction
1. A prime cause of extinction
2. Ex. Deforestation
3. Subtle alterations (e.g., pollution) have not yet been shown
to cause extinction
iii.
Direct exploitationhunting
1. Caused numerous extinctions
2. Ex. Figure 3.4
4. Endangered Species
a. Definitiona species that is thought to be at risk of extinction in the
foreseeable future
b. Factors threatening species with extinction
i. Habitat loss or modification
ii. Hunting
iii.
Accidental or deliberate introduction of exotic species
iv. Deliberate eradication
v. Incidental
vi. Disease, both exotic and endemic
c. Characteristics of factors
i. Human in origin
ii. Species are threatened with several factors simultaneously
iii.
Relative importance as measured by frequency of occurrence
1. Ex. Threats facing terrestrial mammals in Australia and the
Americas
a. 119 species considered endangered
b. 75% threatened by more than one factor
c. 27 species face four or more threats

d. Major threat76% of the species are experiencing

habitat loss or modification
e. Figure 3.5
d. Significance of huntingvaluable fur and wood (Figure 3.6)
e. Overexploitation
i. Overharvesting for commercial interests
ii. Rare plants are threatened by collectors
f. Categorized threats to plants and animals in the U.S.
i. 1998, David Wilcove
ii. Five categories
1. Habitat destruction
2. Alien species
3. Overharvesting
4. Disease (both native and alien)
5. Pollution
iii.
Sample size: 1,880 species
iv. Results: (Figure 3.7) Habitat degradation again most important
g. Categorization of threats by class of species (Figure 3.8)
h. Categorization of threats by geographic areas (Table 3.2)
i. The majority of threatened mammals occur in tropical countries
1. Tropical countries have more species (therefore, should
have more endangered species)
2. Tropical countries have a higher percentage of endangered
species as well
ii. Bigger countries have more endangered species than smaller
countries (Figure 3.9)
iii.
U.S. has proportionally more endangered reptiles, amphibians, and
fishes
1. This could be because of better monitoring and
documenting activities
i. Correlations between human factors and extinction
i. 1995, Kerr and Currie
ii. Compared 90 countries
iii.
Six indices of human activities (Table 3.3)
iv. Human population explained the most variation in the proportion
of endangered species of birds
v. Per capita GNP explained the most variation in mammals
5. Species Characteristics and Extinction (Figure 3.10)
a. Rarity
i. Determined by
1. Geographic range
2. Breadth of habitat
3. Local population size
b. Ability to disperse
i. Rescuing a population through immigration
c. Degree of specialization

i.

Organisms that are specialized are more likely to become extinct

1. Limited food
2. Limited habitat
d. Population variability
i. Stable populations are less likely to go extinct
e. Trophic status
i. Applies to animals only
ii. Higher trophic levels more at risk
f. Life span
i. Shorter life span increases risk of extinction
g. Reproductive ability
i. More offspring decreases risk of extinction
6. Summary
a. Important causes of extinction
i. Introduced species (39%)
ii. Habitat destruction (36%)
iii.
Direct exploitation (23%)
b. Factors threatening species
i. Habitat destruction
1. Deforestation and ecosystem conversion
c. Species characteristics affecting the sensitivity to extinction
i. Rarity
ii. Ability to disperse
iii.
Degree of specialization
iv. Population variability
v. Trophic status
vi. Longevity
vii. Reproductive ability
7. Discussion Questions
a. Which type of organisms do you think deserve priority in conservation
efforts and why?
b. What ecological information would you need in order to list a species as
endangered?
c. If we are concerned with protecting rare species, should we also be
concerned with protecting subspecies (or races) or even individual
populations? Discuss the pros and cons of this issue.

Chapter 4: Group Selection and Individual Selection

1. Outline
a. Group selection vs. individual selfishness
b. Altruism
c. Benefits and trade-offs of group living
2. Group and Individual Selection

a. Regulation of populationsearly thoughts

i. Levels below which competition becomes important
ii. Avoid wastefulness
iii.
Development of Group Selection
1. Territoriality of birds
2. Increase in emigration correlated with increase in numbers
3. Mechanisms operate in the absence of limitations
4. High variation in reproductive rates
5. Examples of self-regulation
a. 1940, David Lack and Alexander Skutch
i.
Self-regulation of song birds
ii.
Tropics vs. temperate
iii.
Clutch size based on food
b. 1962, V. C. Wynne-Edwards
i.
Animal Dispersion in Relation to Social
Behavior
ii.
Groups of individuals control their numbers
to avoid extinction
iii.
Theory known as Group Selection
1. Successful groupsindividuals
would not act selfishly
2. Selfish groupsoverexploit their
environment and die out
iv. Development of Individual Selection
1. 1966, G. C. Williams
a. Adaptations and Natural Selection
b. Arguments against Group Selection
i.
Mutation
1. Cheater scenario
2. Clutch size based on maximizing the
number of surviving chicks (Figure
4.1)
ii.
Immigration
1. Selfish individuals can migrate to
new areas
iii.
Individual selection
1. Individuals die out more quickly
than groups
2. Individual selection a more powerful
evolutionary force
iv. Resource prediction
1. Group selection needs a reliable and
predictable source of food
2. No evidence
2. Self-Regulation
a. Intraspecific competition

b. Individuals strive to command as much resources as

they can
c. Act in self-interest
i.
Ex. Male lions that kill existing cubs when
they take over pride. Increase their own
offspring
ii.
Ex. Male langur monkeys kill infants
(Figure 4.2)
iii.
Ex. Female giant water bugs kill eggs in
masses being taken care of by males (Figure
4.3)
3. Altruism
a. Apparent cooperation
i. Grooming
ii. Hunting
iii.
Warning signals
b. Caring for copies of ones genes
i. Genes in offspring
ii. Genes in relatives
iii.
Coefficient of relatedness = r
1. Probability of sharing a copy of a particular gene
a. Parents to its offspring; r = 0.5
b. Brothers and sisters; r = 0.5
c. Grandparents to grandchildren; r = 0.25
d. Cousins to each other; r = 0.125
e. Figure 4.4
2. Implications of relatedness to altruism
a. 1964, W. D. Hamilton
b. Importance of passing on ones genes through
offspring as well as related individuals
c. Inclusive fitness
i.
Total copies of genes passed on to all
relatives
d. Kin selection
i.
Lowers individual chance of reproduction
ii.
Raises chances of relatives reproduction
iii.
Quantifying kin selection
1. rBC > 0
2. r = coefficient of relatedness
3. C = number of offspring sacrificed
by donor
4. B = number of offspring gained by
recipient
iv. Ex. Caterpillars
1. Aposematiccontain colors to warn
predators of bad taste or poison

2. Datana caterpillars (Figure 4.5)

3. Predator must kill one to learn
4. Advantage of animals to congregate
in groups (Figure 4.6)
v. Alarms from "sentries"
1. Increased risk of being attacked
2. Animals living near "sentry" most
likely relatives
3. Favors kin selection
4. Alternative to kin selection
a. "Sentries" that are forced to
live at the fringe
b. Alert for their own safety
c. If "sentry" is successful,
predator may seek new area
d. "Sentry" increases chances of
own survival
c. Altruism between unrelated individuals
i. "You scratch my back, Ill scratch yours"
ii. Reciprocal altruism
iii.
Evidence
1. Brooding success correlated to availability of helpers
2. Social hunting
a. Benefit: Bigger prey
b. Cost: Sharing meat
d. Altruism in social insects
i. Extreme example of altruismsterile castes in social insects
ii. Female workers
1. Rarely reproduce
2. Assist queen with her offspring (eusociality)
iii.
Soldier castes (Figure 4.7)
iv. Social insect reproduction (Table 4.1)
v. Relatedness
1. Females are diploid
2. Males are haploid
a. Formed without meiosis
b. Each sperm is identical
3. Sister relatedness
a. Each daughter receives an identical set of genes
from her father
b. Half of a females genes come from her diploid
mother
c. Total relatedness of sisters: 0.5 from father + 0.25
from mother = 0.75
d. Genetic system termed haplodiploidy
e. Relatedness and the queen

i.
ii.

vi.

Sons and daughters; r = 0.5

Maximize reproductive potential. 50:50 sex
ratio
iii.
Average relatedness for sterile workers
would be 0.5
iv. Better for female workers to have more
sisters
v. Colonies usually have more females than
males
4. Non-haplodiploid colonies
a. Termites
b. Mole rat from South Africa (Figure 4.8)
Lifestyles that promote eusociality in mammals
1. Individuals are confined to burrows or nests
2. Food is abundant enough to support high concentrations of
individuals
3. Adults exhibit parental care
4. Mothers can manipulate other individuals
5. Heroism is possible

4. Group Living
a. Dense living
b. Promote intense competition
c. Significant advantages to compensate
d. Guppies and the occurrence of predators (Figure 4.9)
e. "Many-eyes hypothesis"
i. Success of predator attacks
1. Prey alerted to attack (Figure 4.10)
2. Ex. Goshawks less successful attacking large flocks of
pigeons (Columba palumbus)
3. The bigger the flock (more eyes), the more likely the prey
will be alerted to the presence of a predator (Figure 4.11)
4. Cheating vs. the advantages of not cheating
f. Selfish-herd theory
i. Predators usually only take one prey per attack
ii. The bigger the herd, the lower the probability of an individual prey
being taken
iii.
Larger herds are attacked more, but probability of being taken
would still favor individual
iv. Geometry of the selfish herd
1. 1971, W. D. Hamilton
2. Prey prefer middle of herd to avoid predator
3. Predator difficulty in tracking large numbers of prey
4. Peripheral prey easier to visually isolate
5. More difficult for predator to reach the center of herd
v. Large herds are better able to defend themselves
vi. Conflicting variables

1.
2.
3.
4.

Competition for food

Presence of predator
Figure 4.12
Figure 4.13

5. Applied Ecology
a. Tragedy of the Commons
i. 1968, Garrett Hardin
ii. "Tragedy of the Commons"
iii.
Humans and cattle grazing
iv. Carrying capacity of land
1. Ex. Carrying capacity on a piece of land1,000 cattle
a. 10 ranchers share land, each with 100 cattle
b. One individual wants to add one cattle more than
his/her share
i.
Maximizes his/her profits at expense of
others
ii.
All of the cattle suffer very little
c. Tragedy
i.
What would happen if all ranchers did this?
ii.
Overgrazing
iii.
Not sustainable
2. Benefits of the environment often accrue to the individual
3. Cost of using the environment is usually borne by the entire
population
6. Summary
a. Group Selection
i. Past theory
ii. Population maintained at equilibrium based on group selection
1. Self-regulation of individuals
2. Prevent overexploitation of resources
iii.
Several flawsmutation, immigration, and resource prediction
b. Individual Selection
i. More likely
ii. Explanations for altruism
1. Kin selection
2. Caste systems of social insects
3. Haplodiploid mating systems
iii.
Occurrence of eusociality and cooperation
1. Haplodiploid organisms
2. Non-haplodiploid organisms
a. Confinement to burrows
b. High food concentrations
c. Parental care of offspring
d. Mothers can manipulate other individuals
e. Opportunity for heroism
iv. Group SizeTrade-offs

1. Competition for food

2. Protection from predators
7. Discussion Questions
a. If kin selection occurs in nature, how do you think animals recognize their
kin?
b. If it is equally valuable for a female to raise her own young or help raise
sisters (both having an r = 0.5), why do we see most females preferentially
raising their young?
c. Unrelated vampire bats that roost communally often rest next to the same
neighbors every night and sometimes regurgitate meals of blood to their
hungry neighbors. How can you explain this apparent act of altruism?
How, if at all, can animal behavior be important to how we design nature
reserves?

Chapter 5: Life History Strategies

1. Outline
a. Reproductive strategies
i. Species that reproduce throughout their lifetimes (iteroparous)
ii. Species that reproduce just once (semelparous)
b. Age structure
i. Growing populations
ii. Declining populations
c. Classification of mating systems
d. Continuum of life history strategies
i. r-selected
ii. K-selected
iii.
Carrying capacity
2. Reproductive Strategies
a. Semelparity
i. Organisms that produce all of their offspring in a single
reproductive event
ii. May live several years before reproducing or life span is one year
(ex. annual plants)
iii.
Ex. Figure 5.1
b. Iteroparity
i. Organisms that reproduce in successive years or breeding seasons
ii. Variation in the number of clutches and number of offspring per
clutch
iii.
Some species have distinct breeding seasons
1. Ex. Temperate birds and temperate forest trees
2. Lead to distinct generations
iv. Some species reproduce repeatedly and at any time during the year
(continuous iteroparity)

1. Ex. Some tropical species, many parasites, and humans

c. Environmental uncertainty
i. Favors iteroparity
ii. Survival of juveniles is poor and unpredictable
iii.
Selection favors
1. Repeated reproduction
2. Long reproductive life
3. Spread the risk over a longer period ("bet hedging")
d. Environmental stability
i. Favors semelparity
ii. More energy can be devoted to seed production rather than
maintenance
iii.
Annuals rely on seed storage during environmentally unstable
years
3. Age Structure
a. Semelparous organisms
i. Often produce groups of same-aged youngcohorts
ii. Cohorts grow at similar rates
b. Iteroparous organisms
i. Many young at different ages
c. Increasing populationslarge number of young
d. Decreasing populationsfew young
i. Loss of age classes
1. Influence on population
2. Ex. Overexploited fish populationsolder age classes
a. Reproductive age classes removed
b. Reproductive failure
c. Results in population collapse
3. Ex. Younger age classes, deer removing young trees
a. Figure 5.2
4. Mating Systems
a. Sex ratio
i. Applied ecology
1. Hunters prefer deer populations dominated by males
2. Effects of too many males on population growth
ii. Analysis of the ratio
1. Why is the sex ratio usually 1:1?
a. Arent males superfluous?
b. Answer: Selfish genes!
i.
Populationspredominately female
1. Selection would favor sons
ii.
Populationspredominately male
1. Selection would favor daughters
iii.
Over time, sex ratio would be kept at 1:1
iii.
Exception to 1:1
1. One male dominates in breeding

2. Occurs in species with

a. Low powers of dispersal
b. Inbreeding is frequent
3. Ex. The parasitic Hymenoptera
a. Females mate once and store sperm
b. Females control sex ratio
i.
Use sperm to create females
ii.
Without sperm to create males
c. Process termed haplodiploid
4. Ex. The mite Acarophenox (Figure 5.3)
b. Mating systems in animals
i. Monogamy
1. Exclusive mating
2. Common among birds (~90%) of species
ii. Polygamy
1. Individuals mate with multiple partners
2. Polygyny
a. One male mates with multiple females
b. Females mate with one male
3. Polyandry
a. One female mates with multiple males
b. Males mate with one female
iii.
Polygyny
1. Females must care for the young
2. Mammals tend to be polygynous
a. Ex. Figure 5.4
3. Influenced by spatial and temporal distribution of females
a. Monogamous relationships result from all females
becoming sexually receptive at the same time
b. Female receptiveness spread over weeks or months
polygyny can result
4. Resource-based polygyny
a. Critical resource is patchily distributed or in short
supply
b. Male can dominate resource and breed with more
than one visiting female
c. Disadvantages for the female
i.
Must share resources
ii.
More females means less success
iii.
Figure 5.5
5. Non-resource based polygyny
a. Harem-based
i.
Common in groups or herds
ii.
Protection from predators
iii.
Harem master does not remain for long
b. Communal courting areasleks

i.
iv.

Figure 5.6

Polyandry
1. Practiced by a few species of birds
2. Ex. Spotted sandpiper in the Arctic tundra
a. Reproductive success not limited by food
b. Limited by the number of males needed to incubate
eggs
3. Ex. American jacana (Figure 5.7)
5. Life History Strategies
a. Success of populations
i. Reproductive strategies
ii. Survival strategies
iii.
Habitat usage
iv. Competition with other organisms
b. Continuum
i. r-selected to K-selected (Figure 5.8)
ii. r and K refer to population growth equation parameters
iii.
r-K classification (McArthur and Wilson, 1967)
1. Table 5.1
iv. r-selected
1. High rate of per capita population growth
2. Tend to disperse well
3. Semelparous
4. Poor competitive abilities
5. Ex. Weeds
a. Quickly colonize an area
b. Exist in disturbed habitats
c. Grow quickly
d. Reach reproductive age early
e. Devote large amounts of energy to seed production
f. Seeds disperse widely
g. Short life span
v. K-Selected
1. Populations increase slowly toward the carrying capacity
(K) of the environment
2. Low reproductive allocations
3. Iteroparous
4. High competitive abilities
5. Ex. Mature forest
a. Non-disturbed habitat
b. Grow slowly
c. Reach reproductive age late
d. Devote large amounts of energy to growth and
maintenance
e. Grow to large sizes and shade-out r-selected species

f. Long-lived and produce seeds repeatedly every year

while mature
g. Seeds are bigger than r-selected speciesprovide
food reserves to help them get started
c. Alternatives to the r and K continuum
i. Ruderals, competitors, and stress tolerators (Grime, 1977 and
1979)
1. Ruderals (botanical term for weed)
a. Adapted to cope with habitat disturbances
2. Competitors
a. Adapted to live in highly competitive but benign
environments (e.g., tropics)
3. Stress tolerators
a. Adapted to cope with severe environmental
conditions (e.g., salt marsh plants)
4. Stress, disturbance and competition triangle
a. Figure 5.9
5. Demographic interpretation (Silverton et al., 1992, 1993)
a. Growth-survival and fecundity triangle
b. Figure 5.10
6. Applied Ecology
a. Life history and the risk of extinction
b. K-selected species
i. All attributes set them at risk to extinction
ii. Tend to be biggerneed bigger habitat
iii.
Fewer offspringpopulations cannot recover as fast from
disturbance
iv. Breed later in lifegeneration time is long
v. Population size is smallhigh risk of inbreeding
vi. Examples
1. Florida panthers
2. Giant sequoia trees
3. Large terrestrial mammals (elephants, rhinoceros, and
grizzlies)
4. Large marine mammals (blue and sperm whales)
7. Summary
a. Life history concerns lifetime patterns in reproduction and growth patterns
i. Semelparous
ii. Iteroparous
b. Reproductive strategy strongly affects age structure
i. Low ratio of young to adultspopulation in decline
ii. High ratio of young to adultspopulation growing
c. Sex ratio
i. 1:1 ratio expected in most populations
ii. Polygynous
1. Males mate with more than one female

iii.

Polyandrous
1. Females mate with more than one male
iv. Monogamous
1. Each individual has one mate
v. Polygamy is often based on limited resources
d. Categorizing life history strategies
i. r-K continuum
1. r-selected
a. Poor competitors
b. High per capita population growth rate
c. Disperse well
d. Colonize new habitats
2. K-selected
a. Good competitors
b. Usually exist in mature habitats, close to the
carrying capacity
ii. Alternative life history strategies
1. Ruderals, competitors, and stress tolerators classification
2. Growth-longevity-fecundity triangle
8. Discussion Questions
a. What particular life history strategies are possessed by successful exotic
invaders like kudzu in the southeast or zebra mussels in the Great Lakes?
Can knowing their life histories help us in the war against exotics?
b. How could you test the idea that there is a trade-off between life history
strategies? What would happen if you plant salt marsh plants like Spartina
grass or mangroves in a terrestrial habitat vs. a freshwater habitat?
c. In some species, males are much bigger than females, a property called
sexual dimorphism. Speculate about the types of animals in which sexual
dimorphism would and would not occur.

Chapter 6: Population Growth

1. Outline
a. Tabulating changes in population age structure through time
i. Time-specific life tables
ii. Age-specific life tables
b. Fecundity schedules and female fecundity, and estimating future
population growth
c. Population growth models
i. Deterministic models
ii. Geometric models
iii.
Logistic models
iv. Stochastic models
2. Life Tables

a. Date on numbers of individuals at each age

i. Construct life tables
ii. Demonstrate the age structure of a population
b. Life table constructiondemography
i. How a population will grow
ii. Construction of data
1. Follow a cohort from births to deaths (age-specific life
table)
2. Snapshotage structure at a single point in time (timespecific life table)
iii.
Time-specific life table
1. Useful in examining long-lived animals
a. Ex. Dall Mountain Sheep (Figure 6.1 and Table 6.1)
2. Useful parameters in the life tables (Deevey, 1947)
a. x = age class or interval
b. nx = number of survivors at beginning of age
interval x
c. dx = number of organisms dying between age
intervals = nx - nx+1
d. lx = proportion of organisms surviving to the
beginning of age interval x = ns/n0
e. qx = rate of mortality between age intervals = dx/ns
f. ex = the mean expectation of life for organisms alive
at the beginning of age x
i.
Lx = average number alive during an age
class = (ns + ns + 1)/2
ii.
Tx = intermediate step in determining life
expectancy = Lx
iii.
Ex = Tx/nx
3. Assumptions that limit the accuracy of time-specific life
tables
a. Equal number of offspring are born each year
i.
Favorable climate for breeding?
b. A need for an independent method for estimating
birthrates of each age class
c. As a result, age-specific life tables are typically
reported
iv. Age-specific life tables
1. Needed for short-lived organisms
a. Time-specific life tables biased toward the stage
common at the moment
2. Follows one cohort or generation
3. Population censuses must be frequent and conducted over a
limited time
4. Ex. Table 6.2 and Figure 6.3
v. General types of survivorship curves (Figure 6.4)

1. Type I
a. Most individuals are lost when they are older
b. Vertebrates or organisms that exhibit parental care
and protect their young
c. Small dip at young age due to predators
2. Type II
a. Almost linear rate of loss
b. Many birds and some invertebrates
3. Type III
a. Large fraction are lost in the juvenile stages
b. Invertebrates, many plants, and marine invertebrates
that do not exhibit parental care
c. Large losses due to predators
vi. Comparison in the accuracy of life tables
3. Reproductive Rate
a. Fecundity
i. Age-specific birthrates
ii. Number of female offspring produced by each breeding female
b. Fecundity schedules
i. Fecundity information in life table
ii. Describe reproductive output and survivorship of breeding
individuals
iii.
Ex. Table 6.3
iv. Table components
1. lx = survivorship (number of females surviving in each age
class
2. mx = age-specific fecundity
3. Ro = populations net reproductive rate = lx mx
a. Ro = 1; population is stationary
b. Ro > 1; population is increasing
c. Ro < 1; population is decreasing
d. Table 6.3
4. Variation in formula for plants
a. Age-specific fecundity (mx) is calculated differently
b. Fx = total number of eggs, seeds, or young deposited
c. nx = total number of reproducing individuals
d. mx = Fx/nx
e. Figure 6.5
f. Table 6.4
g. Figure 6.6
4. Deterministic Models: Geometric Growth
a. Predicting population growth
b. Need to know
i. Ro
ii. Initial population size
iii.
Population size at time t

c. Population size of females at next generation = Nt + 1 = RoNt

i. Ro = net reproductive rate
ii. Nt = population size of females at this generation
d. Dependency of Ro
i. Ro < 1; population becomes extinct
ii. Ro = 1; population remains constant
1. Population is at equilibrium
2. No change in density
iii.
Ro > 1; population increases
1. Even a fraction above one, population will increase rapidly
2. Characteristic "J"-shaped curve
3. Geometric growth
4. Figure 6.7
5. Something (e.g., resources) will eventually limit growth
6. Population crash
7. Figure 6.8a
8. Figure 6.8b
9. Figure 6.8c
e. Human population growth
i. Prior to agriculture and domestication of animals (~10,000 B.C.)
1. Average annual rate of growth: ~0.0001%
ii. After the establishment of agriculture
1. 300 million people by 1 A.D.
2. 800 million by 1750
3. Average annual rate of growth: ~0.1%
iii.
Period of rapid population growth
1. Began 1750
2. From 1750 to 1900
a. Average annual rate of growth: ~0.5%
3. From 1900 to 1950
a. Average annual rate of growth: ~0.8%
4. From 1950 to 2000
a. Average annual rate of growth: ~1.7%
5. Reasons for rapid growth
a. Advances in medicine
b. Advances in nutrition
6. Trends in growth (Figure 6.9)
iv. Human population statistics
1. Population is increasing at a rate of 3 people every second
2. Current population: over 6 billion
3. UN predicts population will stabilize at 11.5 billion by
2150
4. Developed countries
a. Average annual rate of growth from 19601965:
1.19%

v.

vi.

b. Average annual rate of growth from 19901995:

0.48%
5. Developing countries
a. Average annual rate of growth from 19601965:
2.35%
b. Average annual rate of growth from 19901995:
2.38%
6. Developed vs. Developing (Table 6.5)
7. Fertility rates
a. Average number of live births typically borne by a
woman during her lifetime (Table 6.6)
b. Theoretic replacement rate: 2.0
c. Actual replacement rate: 2.1
d. Decline in fertility rate
i.
19601965: ~5.0
ii.
1990: 3.3
Overlapping generations
1. Many species in warm climates reproduce continually and
generations overlap
2. Rate of increase is described by a differential equation
a. dN/dt = rN = (b - d)N
b. N = population size
c. t = time
d. r = per capita rate of population growth
e. b = instantaneous birthrate
f. d = instantaneous death rate
g. dN = the rate of change in numbers
h. dt = the rate of change in time
i. dN/dt = the rate of population increase
j. Produces a "J"-shaped curve
k. Plot with the natural logarithm, produces a straight
line (Figure 6.8)
l. r is analogous to Ro
i.
In a stable population
ii.
R = (ln Ro)/Tc
1. Tc generation time
Logistic Growth
1. Occurs in populations where resources are or can be
limiting
2. Logistic growth equations
a. dN/dt = rN[(K - N)/K]; or
b. dN/dt = rN[1 - (N/K)]
i.
dN/dt = Rate of population change
ii.
r = per capita rate of population growth
iii.
N = population size
iv. K = carrying capacity

v.

3.
4.

5.

6.

7.

(K - N)/K = unused resources remaining

1. As N gets larger, the amount of
resources remaining gets smaller
2. When N = K, zero growth will occur
S-Shaped Curve: Figure 6.11
Logistic growth assumptions
a. Relation between density and rate of increase is
linear
b. Effect of density on rate of increase is instantaneous
c. Environment (and thus K) is constant
d. All individuals reproduce equally
e. No immigration and emigration
Testing assumptions
a. Early laboratory cultures
i.
Pearl, 1927
ii.
Figure 6.12
b. Complex studies and temporal effects
i.
Figure 6.13
Difficulty in meeting assumptions in nature
a. Each individual added to the population probably
does not cause an incremental decrease to r
b. Time lags, especially with species with complex life
cycles
c. K may vary seasonally and/or with climate
d. Often a few individuals command many matings
e. Few barriers to prevent dispersal
Effect of time lags
a. Robert May (1976)
b. Incorporated time lags into logistic equation
c. dN/dt = rN[1 - (Nt - /K)]
i.
dN/dt = Rate of population change
ii.
r = per capita rate of population growth
iii.
N = population size
iv. K = carrying capacity
v. = time lag between the change in
population size and its effect on population
growth, then the population growth at time t
is controlled by its size at some time in the
past, t -
vi.
Nt - = population size in the past
d. Ex. r = 1.1, K = 1000, and N = 900
i.
No time lag, new population size
1. dN/dt = 1.1 x 900 (1 - 900/1000) =
99
2. New population size = 900 + 99 =
999

3. Still below K
ii.
With time lag, where a population is 900,
although the effects of crowding are being
felt as though the population was 800
1. dN/dt = 1.1 x 900 (1 - 800/1000) =
198
2. New population size = 900 + 198 =
1098
3. Possible for a population to exceed K
e. Effect of response time
i.
Inversely proportional to r = (1/r)
ii.
Ratio of time lag ( ) to response time (1/r)
or r controls population growth
1. r is small (<0.368)
a. Population increases
smoothly to carrying capacity
b. Figure 6.14
2. r is large (>1.57)
a. Population enters into a
stable oscillation called a
limit cycle
b. Rising and falling around K
c. Never reaching equilibrium
3. r is intermediate (>0.368 and <1.57)
a. Populations undergo
oscillations that dampen with
time until K is reached
8. Species with discrete generations
a. Logistic equation
i.
Nt + 1 = Nt + rNt [1 - (Nt/K)]
1. In discrete generations, the time lag
is 1.0
ii.
r is small (2.0)
1. Population generally reaches K
smoothly
iii.
r is between 2.0 and 2.449
1. Population enters a stable two-point
limit cycle with sharp peaks and
valleys
iv. r is between 2.449 and 2.570
1. More complex limit cycles
2. Figure 6.15
v. r is larger than 2.57
1. Limit cycles break down

2. Population grows in a complex,

nonrepeating pattern, known as
"chaos"
5. Stochastic Models
a. Models of population growth which incorporate genetic variability and
extrinsic factors
b. Stochastic = variability
c. Models are based on probability theory
i. Figure 6.16
d. Stochastic models of geometric growth
i. dN/dt = rN = (b - d)N
ii. If b = 0.5, d = 0, and N0 = 10, integral form of equation
1. Nt = N0ert
2. So for the above example,
a. Nt = 10 x 1.649 = 16.49
iii.
Path of population growth
1. Figure 6.17
e. Probability of extinction = (d/b)N0
i. The larger the initial population size
ii. The greater the value of b - d
iii.
The more resistant a population is to extinction
f. Introduce biological variation into calculations of population growth
i. More representative of nature
ii. More complicated mathematics
iii.
Important for small populations
iv. For large populations, deterministic models will suffice
6. Applied Ecology
a. 1992 Johns Hopkins study
i. Developed countries
1. 70% of couples use contraceptives
ii. Developing countries
1. ~45% of couples use contraceptives
2. Africa, 14%
3. Asia, 50%
4. Latin America, 57%
iii.
China
1. 1950s and 1960s
a. Fertility was six children per woman
2. 1970s
a. Government planning and incentives to reduce
population growth
3. 1990
a. 75% use birth control
b. Fertility rate dropped to 2.2
iv. Other governments
1. 1976, only 97 governments supported family planning

2. 1988, 125 governments supported family planning

3. As of 1989, in 31 countries, couples have no access to
family planning
v. Women
1. Women in developing countries want fewer children
2. In virtually every country outside of Saharan Africa, the
desired number of children is below 3
b. Countries concerned about low growth rates
i. Some Western European countries and other developed countries
ii. Total fertility has dropped below the replacement level of 2.1
iii.
Reduced populations concerns
1. Affect political strength
2. Economic structure
7. Summary
a. Life tables provide information on how populations are structured. For
longer-lived organisms, time-specific tables provide a snapshot of the
populations age structure.
b. For shorter-lived organisms, age-specific life tables are used
c. Three types of survivorship
i. Type I: Most individuals die after they have reached their sexual
maturity
ii. Type II: An almost linear rate of loss
iii.
Type III: Most individuals are lost as juveniles
d. With information on survivorship and age-specific fecundity, net
reproductive rate, R0, can be calculated and population growth determined
e. Population growth can be examined using deterministic or stochastic
models
f. When environments are favorable, and resources are not limiting
i. Populations undergo geometric or exponential growth
ii. Results in J-shaped population growth curves
g. Limits on population growth
i. Exhaustion of space or resources
ii. Waste product accumulation
iii.
Results in an S-shaped growth curve
iv. Upper level is the carrying capacity, K
h. Population growth curves are not always smooth in trajectory
i. Curves may oscillate or appear to be random fluctuations
ii. Result of time lags
8. Discussion Questions
a. What limits the population growth of organisms most commonly
weather, resources, pollutants, natural enemies, disease, or social factors?
Pick some local plants and animals and make some educated guesses.
b. What do you think will limit human population growth? Do you think the
carrying capacity of the Earth for humans has still to be reached or have
we overshot it? Remember, time lags can cause carrying capacity to be
exceeded. If fossil fuels are used up, could the human population crash?

c. Compare the reproduction output of a family line, which has mothers

always having twin girls at age 20 years old, versus a family line which
has mothers having triplet girls at age 33. What conclusions can you draw?
(Hint: compare reproductive output after 100 and 200 years.) Should all
couples stop reproducing after two children?
d. In what circumstances might a population be expected to grow in a
geometric fashion, and what circumstances in logistic fashion?

Chapter 7: Physical Environment

1. Outline
a. Physical variables commonly limit the abundance of plants and
animals
i. Resources
ii. Variables critical to survival
b. Physical factors commonly limiting species
i. Extreme temperatures
ii. Wind
iii.
Salt
iv. Global climate change
c. Physical environment limits abundance and distribution
d. Physical environment can alter species composition
2. Physical Variables and Species Abundance
a. Liebigs Law of the Minimum (1840)
i. The distribution of a species will be controlled by that
environmental factor for which the organism has the
narrowest range of tolerance
ii. Optimum range (Figure 7.1)
iii.
Effects of competition (Figure 7.2)
3. Physical Variables
a. Temperature
i. Affects biological processes
ii. Organism's inability to regulate body temperature
1. Distribution of coral reefs (Figure 7.3)
2. Distribution of Larrea tridentata (Figure 7.4)
3. Distribution of vampire bats (Figure 7.5)
iii.
Mean temperature vs. Extreme temperatures
1. Frequency of extremes limits species
2. Ex. Agriculture and occurrence of freezing
temperatures
a. Distribution of oranges in Florida
b. Distribution of coffee in Brazil
iv. Correlations between temperature and species distribution

v.

vi.

1. Temperature maps may not coincide with what

organisms experience
a. Movement from sun to shade environments
2. Temperature at the local scale is much more
variable
a. Ex. Microclimates of a tree
i.
South-facing vs. north-facing canopy
ii.
Soil surface to top of canopy
b. Ex, Rufous grasshopper
i.
Restricted to steep sunny slopes
ii.
Combination of time and
temperature is important
1. Degree-days determine
development
High temperature
1. High temperatures denature proteins (temperatures
above 45 )
2. Organisms effectively cool themselves through
water loss
3. Life-history stages resistant to high temperatures
a. Resting spores of fungi
b. Cysts of nematodes
c. Seeds of plants
i.
Ex. Dry wheat grains (90)
ii.
Thermus aquaticus (67)
iii.
Thermophilic bacteria (100; Figure
7.6)
Fire
1. North America before the Arrival of Europeans
a. Fires started by lightning
b. Frequent and regular
c. Consumed leaf litter, branches, and
undergrowth before great quantities
accumulated
d. Large trees usually not damaged
e. Some species evolved to require fire
i.
Pinus banksiana
ii.
Pinus palustris
iii.
Serotinous cones
1. Cones sealed with resin
2. Require heat from fire to
open
2. North America after the Arrival of Europeans
a. Management practices
i.
Maintain "natural" environment
ii.
Preventing forest fires

iii.

vii.

Produced the opposite

1. Change in species
composition
2. Catastrophic fires (Figure
7.7)

Global Warming
1. Two Issues
a. Rate of global warming
b. Contribution by humans
2. Increased global warming = greenhouse effect
a. Atmosphere transmits short-wave solar
radiation
i.
50% passes through the atmosphere
unaltered to heat the Earth.
b. Energy absorbed by the Earth is radiated
back to the atmosphere as long-wave
radiation
c. Long-wave radiation, much is absorbed by
clouds
d. A large amount of energy absorbed in the
atmosphere is returned to the Earth, causing
the temperature to rise
3. Earth requires some "greenhouse effect"
a. Without any greenhouse effect
i.
global average temperature: -17
b. With greenhouse effect
i.
global average temperature: +15
c. Explains hot Venus (blanketed in CO2) and
cold Mars (which has little atmosphere)
4. Greenhouse gases
a. Table 7.1
b. Figure 7.8
5. Influence of natural sources
a. Nitrous oxide
i.
2/3 comes from natural soils and
oceans
b. Methane
i.
1/3 comes from bogs, swamps, and
termites
c. Dust and carbon
i.
Volcanoes
6. Human influences
a. 75% of increases in CO2 emisssions
b. 39% of methane output
c. 36% of nitrous oxide emissions
d. ~50% of all greenhouse emissions

e. Alterations in land use (~25%)

i.
Deforestation
ii.
Conversion to rice paddies
iii.
Increase in domestic animals
iv. Agricultural soils
f. Overall, humans account for 75% of the
increase in greenhouse gases
g. Is it possible to replace fossil fuels?
7. Evidence of temperature increases
a. Temperature record (Figure 7.9)
i.
Problems with record
ii.
Warming has occurred
8. Computer Models and Predictions
a. Too many variables to include in a single
computer model
b. Negative feedback mechanisms
c. Positive feedback mechanisms
d. UN Intergovernmental Panel on Climate
Change (IPCC)
i.
1996 Report
ii.
Lack of fit of models
1. the formation of other
pollutants
a. aerosols of sulfate
b. soot
c. reduce solar heating
e. Emphasizes the complexity of interactions
9. Environmental Impact
a. Speed and Extent of global warming
b. Focus on 2100
i.
Atmospheric CO2 will have doubled
ii.
Temperature will have risen 1 to
3.5C
10. Natural Ecosystems
a. Profound changes in natural ecosystems
b. Most species cannot evolve significantly or
rapidly enough to counter climate changes
c. Most species will not be able to disperse or
migrate fast enough to keep up with climate
change
i.
Figure 7.10
d. Rainfall patterns
i.
Increase in rainfall (Figure 7.11) in
most areas
1. Increased crop production

ii.

viii.

ix.

x.

xi.

a. Ex. Tropical countries

and rice production
Decrease in some areas already dry
1. Midcontinental America and
Asia
2. More droughts
3. More extinctions
4. Current grain-producing
areas would become drier

Wind
1. Can be caused by temperature gradients
2. Amplifies temperature effects on organisms
a. Increases heat loss through evaporation and
convection
b. Increases animal evaporation and plant
transpiration
3. Wind aids pollination
4. Wind disperses plant seeds
5. Affects mortality (Figure 7.12)
a. High winds
b. Severe storms
6. Modify wave action
Salt
1. Increases osmotic resistance to water uptake
a. Occurs in arid regions
b. Important to agriculture in arid regions
i.
Increases salt concentration
ii.
Decreases crop yield
c. Salt marshes
i.
Halophytes
1. adapted to high salt
concentrations
2. Ex. Spartina grasses (Figure
7.13)
pH
1. Few organisms can exist below pH 4.5
2. Ex. Lake trout in Eastern U.S. disappear when pH
drops below 5.2
3. Roots are damaged below pH 3 and above 9
a. Calciphobe: only grow on acidic soils
b. Calciphiles: only grow in basic soils
c. Neutrophiles: tolerant of either condition
Water
1. Protoplasm is 8590% water
2. Distribution of many plants limited by water
availability

3. Animal distribution affected by desiccation

4. Tolerance and Avoidance
4. Physical Factors and Species Abundance
a. Davidson, Andrewartha, and Birch
i. Thrips (Figure 7.14)
ii. Fed on rosebushes
iii.
Counted every 81 consecutive days
iv. 78% of variation in population maxima was accounted for
by weather
v. Predict the number of thrips using multiple regressions
1. Log y = -2.39 + 0.125a + 0.201b + 0.186c + 0.085d
a. Log y = log of thrip density
b. a = winter temperature
c. b = spring rainfall
d. c = spring temperature
e. d = size of overwinter population
vi. Figure 7.15
b. Africa Buffalo and environmental regulation
i. Rainfall and grass productivity in the Serengeti
ii. Buffalo density regulated by food availability
iii.
Figure 7.16
c. Woddell, Mooney, and Hill (1969)
i. Correlation between rainfall and creosote bush density
ii. Figure 7.17
5. Physical Factors and Numbers of Species
a. Importance of evapotranspiration
i. Figure 7.18
b. Robert Whittaker (1969)
i. Four hypotheses explaining distribution patterns
1. Figure 7.19
a. Competition causes sharp boundaries among
groups
b. Competition causes sharp boundaries
between species
c. Physical environment causes distinct
boundaries between groups
d. Physical environment causes distinct
boundaries between species
e. Evidence supports hypothesis (d) above
6. Applied Ecology
a. Diseases and Global Climate Change
b. Spread of tropical diseases poleward
i. Controlled by the range of their vectors
1. Ex. Mosquitoes and other insects
2. Insects are ectotherms

ii.

Increase in temperature = increase in range and activity of

vectors
1. Ex. Rwanda, 1987
a. 1C increase in temperature resulted in a
337% increase in malaria
c. Diseases likely to spread (Box Table)
d. Computer model prediction
i. Average global temperature increase of 3C
ii. 5080 million new cases of malaria per year
7. Summary
a. Local distribution patterns are limited by certain physical or abiotic
factors, such as temperature, moisture, light, pH, soil, salinity.
b. Temperature is one of the most important physical variables
affecting plant and animal distribution
i. Resistance to high temperature and fire
ii. Implication of climate change
c. Wind can amplify the effects of temperature, and can affect species
distribution directly.
d. Soil salt concentration and pH can affect water uptake in plants
e. Physical factors strongly influence population densities.
f. The physical environment can profoundly influence species
richness
8. Discussion Questions
a. Why do you think there is a "tree line" on the sides of many of the
Rocky Mountains beyond which no trees grow? Is it because of
low temperatures, low water availability, or high wind? How
would you devise an experiment to find out?
b. Do you think an estuary would be stressful habitat to live in?
Explain. Are there any disadvantages to living in an estuary?
(Think in terms of salinity, food availability, and competition.)
c. Are you interested in maintaining buffalo populations in the United
States? Why is fire so important to achieving this goal?
d. What is the value of laboratory experiments in attempting to
identifying limiting factors to explain the observed distribution
patterns of plants and animals in the field?

Chapter 8: Competition and Coexistence

1. Outline
a. Forms of competition: Interspecific and intraspecific
b. Intraspecific competition
i. Common in nature
ii. Described by the 3/2 thinning law
c. Interspecific competition

i.
ii.

Common in nature
Outcome affected by
1. Physical environment
2. Other species
d. Competition
i. Exists among 5575% of the species
ii. Mechanism: overuse of the same resource
e. Mathematical models, called Lotka-Volterra models, predict four
outcomes of competition
i. One species eliminated
ii. The other species is eliminated
iii.
Both species coexist
iv. Either species is eliminated, depending on starting conditions
f. Competing species can coexist through partitioning of resources
2. Species Interactions
a. Summary of biotic interactions (Table 8.1)
i. Herbivory, predation, parasitism
1. Positive for one population
2. Negative for the other population
ii. Batesian mimicry
1. Mimicry of a non-palatable species by a palatable one
2. Positive for one population
3. Negative for the other population
iii.
Amensalism
1. One-sided competition
2. One species has a negative effect on another, but the
reverse is not true.
iv. Neutralism
1. Coexistence of noninteracting species
2. Probably rare
v. Mutualism and commensalism
1. Less common
2. Symbiotic relationships
3. Species are intimately associated with one another
4. Both species may NOT benefit from relationship
5. Not harmful, as is the case with parasitism
vi. Competition
1. Negative effect for both species
b. Types of competition (Figure 8.2)
i. Interspecific
ii. Intraspecific
c. Characterizing competition
i. Resource competition
1. Organisms compete for a limiting resource
ii. Interference competition
1. Individuals harm one another directly by physical force

3. Intraspecific Competition
a. Quantifying competition in plants vs. animals
i. For plants, expressed as change in biomass
ii. For animals, expressed as change in numbers
iii.
Plants cannot escape competition
iv. Animals can move away from competition
v. Yoda (1963)
1. Quantify competition between plants
2. Yodas Law or self-thinning rule; 3/2 power rule
3. Describes the increase in biomass of individual plants as
the number of plant competitors decrease.
4. Log w = -3/2 (log N) + log c
a. w = mean plant weight
b. N = plant density
c. c = constant
5. w = cN3/2
6. Figure 8.3
4. Interspecific Competition: Laboratory Experiments
a. Field experiments
i. Organisms can interact with all other organisms
ii. Natural variations in the abiotic environment is factored in
b. Laboratory experiments
i. All important factors can be controlled
ii. Vary important factors systematically
c. Thomas Park competition experiments
i. Tribolium castaneum (Figure 8.4a) and Tribolium confusum
ii. Large colonies of beetles can be grown in small containers
iii.
Large number of replications
iv. Observed changes in population sizes over two to three years
v. Waited until one species became extinct
vi. Cultures were infested with a parasite Adelina
vii. T. confusum won 89% of the time
viii.
Without the parasite, no clear winner
ix. Microclimate effects (Figure 8.4)
1. T. confusum did better in dry environments
2. T. castaneum did better in moist environments
x.
Mechanism of competitionpredation of eggs
xi. Predatory tendencies varied with different strains
1. Figure 8.5
d. Interspecific competition: Natural systems
i. Assessing the importance of competition
1. Remove species A and measure the response of species B
2. Difficult to do outside of laboratory
a. Migration problems
b. Krebs or Cage effect
3. Examples in nature

a. Parasitic wasps
b. Figure 8.6
c. Used to control scale pest
d. Climate can alter competitive outcomes
5. The Frequency of Competition
a. Joe Connell (1983)
i. Competition was found in 55% of 215 species surveyed
ii. Figure 8.7
iii.
Effects of number of competing species
1. Single pairs: competition was almost always reported
(90%)
2. Multiple species: competition was reported in 50% of the
studies
iv. Differing opinionsSchoener (1983)
b. Common flaws of studies
i. Positive results tend to be more readily accepted than negative
results
ii. Scientists do not study systems at randommay work in systems
where competition is more likely to occur
c. Failure to reveal the true importance of competition in evolution and
ecological time
i. Most organisms have evolved to escape competition and lack of
fitness it may confer
ii. Competition may only occur infrequently and in years where
resources are scarce
d. Patterns of competition
i. Figure 8.8
e. Mechanisms of competition (Schoener, 1963)
i. Table 8.2
ii. Consumptive or exploitative
iii.
Preemptive
iv. Overgrowth
v. Chemical
1. Allelopathy
vi. Territorial
vii. Encounter
f. Differing views of competition
i. Gurevitch et al., 1992
1. Found no differences in competition between different
habitat types but did find filter feeders and herbivores
competed more than carnivores or plants.
ii. Grime, 1979
1. Competition unimportant for plants in unproductive
environments
iii.
Tilman, 1988
1. Competition occurs across all productivity gradients

6. Modeling Competition
a. Based on logistic equations for population growth
b. Growth equations for two populations coexisting independently
i. For species 1; dN1/dt = r1N1[(K1 - N1)/K1]
ii. For species 1; dN2/dt = r2N2[(K2 - N2)/K2]
1. r = per capita rate of population growth
2. N = population size
3. K = carrying capacity
4. Subscripts refer to species
c. Populations that compete
i. Conversion factor that quantifies the per capita competitive effect
of one species on another
ii. For species 1; dN1/dt = r1N1[(K1 - N1 - N2)/K1]
iii.
For species 1; dN2/dt = r2N2[(K2 - N2 - N1)/K2]
1. = per capita competitive effect of species 2 on species 1
2. = per capita competitive effect of species 1 on species 2
iv. dN1/dt = 0: zero-growth isocline
v. Four possible outcomes
1. Figure 8.12
d. Test of equations
i. Figure 8.13
e. Deficiencies
i. The maximal rate of increase, the competition coefficients, and the
carrying capacity are all assumed to be constant
ii. There are no time lags
iii.
Field tests of these equations have rarely been performed
iv. Laboratory tests have shown divergence
1. Figure 8.14
v. Mechanisms that drive competition are not specified
1. R* - Tilman (1982, 1987) alternative
a. Need to know the dependence of an organisms
growth on the availability of resources
b. Figure 8.15
7. Coexistence of Species
a. Niche
i. Grinnell (1918): a subdivision of a habitat that contains an
organisms dietary needs, its temperature, moisture, pH, and other
requirements
ii. Elton (1927) and Hutchinson (1958): an organisms role within the
community
b. Gause: two species with similar requirements could not live together in the
same place
c. Hardin (1960): Gauses principle, known as competitive exclusion
principle, where direct competitors cannot coexist
d. David Lack: Competition and coexistence in about 40 pairs of birds,
mediated by habitat segregation

i. Figure 8.16
e. Examples of coexistence
i. Darwins finches on the Galapagos
ii. Terns on Christmas Island (Ashmole, 1968)
f. Ranks for resource partitioning (Schoener, 1974)
i. Macrohabitat (55%)
ii. Food type (40%)
iii.
Time of day or year (5%)
g. Hutchinson (1959)
i. Seminal paper, "Homage to Santa Rosalia, or why are there so
many kinds of animals?"
ii. Examined size differences for
1. Sympatric species (species occurring together)
2. Allopatric species (occurring alone)
3. Table 8.3
4. Hutchinsons ratio, 1.3
iii.
Criticism of Hutchinson
1. Studies that supported Hutchinsoninappropriate statistics
2. Further tests showed no differences between species than
would occur by chance alone
3. Size-ratio differences could have evolved for other reasons
4. Biological significance cannot always be attached to ratios,
particularly to structures not used to gather food. Figure
8.17
iv. Support of Hutchinson
v. Figure 8.18 d/w analysis for separation on continuous resource sets
1. Figure 8.19
a. d = distance between maxima
b. w = measure of spread
2. Figure 8.20
vi. Discontinuous resource distribution
1. Figure 8.21
2. Figure 8.22
3. Niche overlap between two insect species that feed on a
shrub
a. Measured quantity
i.
PS = pi
ii.
PS = proportional similarity
iii.
= sum of all units, 1 to n, in resource set
iv. pi = proportion of least abundant member of
pair
v. PS < 0.70 indicates coexistence for single
resource
vi.
PS > 0.70 indicates competitive exclusion
for single resource

vii.

Proportional similarity indices for two or

more resources can be combined
1. Multiply separate PS values to
determine overall PS value
2. Coexistence for two resources
a. 0.7 x 0.7 = 0.49 or less

8. Applied Ecology
a. Is the release of multiple species of biological control agents beneficial?
i. Control of pests in agriculture is of paramount importance
ii. Biological control is seen as a preferable alternative to chemical
control
iii.
Biological control viewed by some
1. Release a variety of enemies against a pest
2. Observe which enemy does the best job
3. Is this the best strategy?
a. Intensive competition for the prey leads to lower
effectiveness of the biological agents
b. Greater population establishment rate with fewer
enemy species (Figure 1 for Box 1)
c. Establishment rate of single-species releases were
significantly greater than the simultaneous release
of two or more species (76% vs. 50%)
9. Summary
a. Competition may be interspecific or intraspecific
b. Competition may be viewed as resource competition or interference
competition
c. Intraspecific competition between plants may be described by the 3/2 selfthinning rule
d. Outcome of competition can be influenced by
i. Environmental conditions
ii. The presence or absence of natural enemies
iii.
The genetic strain of the competitors involved
e. Experimental studies show that in nature competition occurs between
different types of organisms over a broad scale
i. Such studies focused on exotics, and generalizations to natural
ecosystems are questionable
ii. Competition between exotics and native species
1. Serious consequences for natural ecosystems
f. Frequency of competition
i. 55% to 75% of species involved
ii. Competition is often asymmetric
g. Six mechanisms of competition
i. Consumptive
ii. Preemptive
iii.
Overgrowth
iv. Chemical

v. Territorial
vi. Encounter
h. Lotka-Volterra model: early competition model
i. Two species interaction
ii. Four possible outcomes
1. Species 1 becomes extinct
2. Species 2 becomes extinct
3. Either species 1 or species 2 becomes extinct based on
starting conditions
4. Coexistence
iii.
Competition is minimized and species can coexist if they use
different resources
1. Hutchinsons 1:1.3 ratio
2. d/w values greater than unity
3. Proportional similarity values no greater than 70%
10. Discussion Questions
a. Which type of competition would you expect to be more important in
nature?
b. Much native vegetation in the Florida Everglades is being lost. Could this
be due to climate change or the influence of exotic invaders? Design an
experiment.
c. In the above question, how could you determine the mechanism of
competition? How could you differentiate among competition for light,
water, or nutrients?
d. In trying to understand how species compete, what advantages are there in
field observations, field experiments, laboratory experiments, and
mathematical models?
e. Using fully labeled graphs, explain the Lotka-Volterra approach to
competition theory. What predictive power does the Lotka-Volterra model
have? How is Tilmans R* concept an improvement? What other
improvements might you suggest?

Chapter 9: Mutualism

1. Outline
a. Mutualism: an association between two organisms that benefits both
b. Seed dispersal mutualism: disperser acquires a meal and the plant gets its
seed dispersed
c. Through mutualism, species are better able together to secure resources or
better able to defend themselves
d. Mutualism is difficult to model; models tend to result in runaway densities
e. Mutualism between two species can affect the entire community

f. Commensalism is an association between two species that benefits only

one, with the other species unaffected
2. PlantPollinator Mutualism
a. Most frequent type of mutualism (Figure 9.1)
i. 45% of all studies of mutualism
ii. Coevolved systems
b. Selective pressures for plants to develop intimate relationship with
pollinators
i. Figure 9.2
ii. More than 900 species of Ficus exist and virtually all must be
pollinated by its own species of agaonid wasp
iii.
Yucca plants and yucca mothshighly coevolved
iv. The distribution of each species is controlled by the availability of
the other species
c. Coevolution stable mutualism
i. Both species have mechanisms to prevent excessive exploitation
1. Ex. Yucca flower abortion if too many eggs are laid
d. Cheating in pollination
i. Bogs of Maine
1. Grass-pink orchid produces no nectar, but mimics the
nectar-producing rose pogonia
2. Some Bombus species cheat by biting through the base of
the flowers, taking the nectar without entering the plant nor
assisting with pollination
3. Seed Dispersal
a. Seed dispersal systems account for almost 30% of all mutualisms
i. In tropics some fruits are dispersed by birds that are frugivorous
1. Fruit provides balanced diet for birds
2. Birds disperse seeds
ii. Mechanisms for attraction
1. Birds and mammalsattractive colors, and odorless
(Birds)
2. Nocturnal batsgive off pungent odor
b. Seed dispersal mechanisms are not as obligatory as plantpollinator
systems
i. Performed by more generalist agents
ii. Wide array of adaptations
1. Ex. Parrot beaks to crack and peel fruits
2. Ex. Figure 9.3
4. A Variety of Mutualisms
a. Mutualisms and resources
i. Neotropical leaf-cutting ants and fungus
1. Figure 9.4
2. Typical ant colony: 9 million
3. Typical biomass of colony: Equivalent to the biomass of a
cow

4. Typically cuts the equivalent of a cows daily requirement

of fresh vegetation
5. Leaf-cutting ants harvest 17% of the total leaf production in
the forest
6. Ants take cut vegetation underground to grow special
fungal crops (natures farmers?)
7. Fungus grows specialized structures called gongylidia,
which serve as food for the ants
ii. Leaf-cutting ants and the parasitic fungus, Escovopsis
1. Escovopsis can infest the ants fungi gardens, destroying
the fungi inside
2. A mutualistic streptomyces
a. Occurs on the bodies of ants
b. Produces antibodies that keep Escovopsis in check
iii.
Nitrogen
1. Vital to plant and animal growth
2. Most species cannot fix atmospheric nitrogen
3. Fixation is carried out by soil bacteria and archaebacteria
a. Most live in the roots of plants
b. Mutualistic relationship with plant
c. Excess nitrogen is available to plant
d. Ex. Rhizobia bacteria in legumes
i.
Figure 9.5
iv. Mutualism under harsh environmental conditions
1. Strong facultative mutualism
2. Leguminous shrub, Retama, and an understory plant,
Marrubium vulgare
a. Grow in a semiarid region of Spain
b. Retama shades Marrubium, provides favorable
microclimate
c. Marrubium enhances the availability of water for
Retama
d. Figure 9.6
b. Mutualism and protection from natural enemies
i. Common example: ants and aphids
1. Figure 9.7
2. Aphids feed on plant sap and excrete honeydew
3. Ants drink the honeydew and in return protect the aphids
c. Mutualism and herbivory
i. Ants protect plants from herbivores
1. Common in the tropics
a. 377 myrmecophytic plants per hectare in a Brazilian
rain forest (Fonseca and Ganade, 1996)
b. 312 antplant associations at a single coastal site in
Mexico (Rico-Grey, 1993)
ii. Example: Ants and the Central American acacia trees

1. Acacia trees provide food and shelter for the ants inside
large thorns
2. Ants protect the acacia tree from other insects and
vertebrate herbivores
3. Ants also trim foliage away from competing plants and kill
neighboring plant shoots
4. Figure 9.8
iii.
Example: Fungi and plants
1. Fungi reduce vertebrate herbivory
2. Soay Sheep of Hirta Island (in the St. Kilda Archipelago)
Figure 9.9
3. Sheep overgraze native grasses
4. Periodically, sheep population crashes
5. Mutualism is the chief culprit
a. The main forage is the grass Festuca rubra
b. F. rubra contains an endophyte, the fungus
Acremonium, inside its blades
c. The fungus produces toxic alkaloids
d. These alkaloids function as an anti-herbivory
defense
e. In return, the fungus obtains food from the plant
f. Frequency of infection correlated to grazing
pressure. Heavy grazing causes higher infections
g. Fungi are in greatest concentration in basal regions.
Heavy grazing results in sheep reaching lower
blades.
d. Obligate mutualism
i. A mutualistic relationship, in which neither participant could
survive without the other
1. Ex. Lichen: a relationship between algae and fungi
a. Algae provides the photosynthate
b. Fungi provides a safe habitat
2. Ex. Many ruminants and symbiotic bacteria
a. Bacteria break down plant tissue to provide energy
for their hosts
3. Ex. The roots of most plants and fungi
a. Mutualistic association between the fungus and root
tissuemycorrhizae
b. Fungi obtain carbohydrates from their host
c. Fungi increase access to mineral nutrition and water
for the plant
5. Modeling Mutualism
a. Uses equations similar to Lotka-Volterra competition equations
b. For facultative mutualism
i. dN1/dt = r1N1[(X1 - N1 + N2)/X1]
ii. dN2/dt = r2N2[(X2 - N2 + N1)/X2]

1. dNi /dt = change in population size of species 1 or 2

2. ri = per capita growth rate for species 1 or 2
3. Ni = population size of species 1 or 2
4. Xi = Maximum population density of species 1 or 2
5. = Positive effect of species 2 on species 1
6. = Positive effect of species 1 on species 2
7. Figure 9.10ac
c. For obligate mutualism
i. Different equations are needed
ii. Models of facultative mutualism are, in general, more stable than
models of obligate mutualism
iii.
Figure (9.10df)
6. Mutualisms and Community Process
a. Mutualism can have strong indirect effects on the community
i. Ex. Mycorrhizal fungi and herbivory load
1. Pinyon pines and mycorrhizae: Density of needle scale
insect, Matsucoccus acalyptus:
a. Mycorrhizae can improve pine vigor and/or increase
plant investment in antiherbivory defenses
b. Density of needle scale insects will decrease
ii. Ex. Endophytes (fungi that live inside leaves) and vascular plant
hostsDefend host against herbivory
iii.
Mycorrhizal fungi and plant species diversity (Figure 9.11)
7. Commensalism
a. Commensal relationship: one member benefits and the other is unaffected
i. Ex. An orchid and a tropical tree: orchids gain a place to live and
the tree gains nothing
ii. Ex. Cattle egrets and cattle: Cattle stir up insect prey for egrets
(Figure 9.12)
b. Commensalism in the form of phoresy: association is passive and more
temporary transport of one organism by another
i. The transport of flower-inhabiting mites from bloom to bloom in
the nares of hummingbirds
c. Common commensalisms: plant mechanisms of seed dispersal
i. Ex. Seeds attached to animal fur
ii. Some mechanisms can become an antagonistic relationship
1. Pisonia (cabbage tree) produce a very sticky fruit
a. On some islands, birds and reptiles become so
entangled with the fruit that they die
iii.
Table 9.1
8. Applied Ecology
a. Humans in mutualistic relationship
i. Humans and agriculture (Box Photo 1)
ii. Origins of worlds crops (Box Table 1)
iii.
Crops are introduced around the world
iv. Livestock are domesticated

v.

vi.

Mutualism
1. Human population has increased
2. Crops and livestock populations have increased
3. Box Table 2
4. Box Table 3
Side-effects of mutualism
1. Pollution of bodies of water
2. Loss of topsoil
3. Depletion of water supplies
4. Salting of the land
5. Desertification
6. Severe loss of wildlife

9. Summary
a. In mutualism, both species benefit
b. In some mutualism, the partners either help each other obtain resources or
protect each other from enemies
c. Obligate mutualisms are mutualisms in which the species cannot live apart
d. Mathematical models are hard to construct because they often result in
runaway populations. Facultative mutualism models are generally more
stable than obligate mutualism models
e. In commensalisms, one member derives a benefit while the other is
unaffected
10. Discussion Questions
a. Where might mutualisms be more common: in temperate areas or in the
more stable tropics? Why?
b. In your perusal of the literature, which type of mutualism covered in the
lecture seems to be most common? Are mutualisms more common in one
particular kind of habitat? If so, why do you think it is so?
c. Boucher, James, and Kresler (1984) suggested that mutualisms are of four
varieties: nutritional, protective, energetic, and transport. Define and give
examples of each.
d. A flock of birds with different species or mixed herds of animals on the
African plains could represent facultative mutualisms. Discuss the
advantages and disadvantages of such a group.

Chapter 10: Predation

1. Outline
a. There are a variety of antipredator adaptations, which suggests that
predation is important in nature
b. Predatorprey models can explain many outcomes
c. Field data suggests that predators have a large impact on prey populations

d. Experiments involving the removal or introduction of exotic predators

provide good data on the effects of predators on their prey
e. Field experiments involving the manipulations of native populations show
predation to be a strong force
2. Introduction
a. Wolves in Yellowstone Park (Figure 10.1)
i. U.S. Fish and Wildlife Service, 1980s
ii. Reintroduce in Yellowstone Park and stabilize wolf populations in
Minnesota and Montana
iii.
Concerns
1. Cattle ranchers concerned: Decimate herd?
2. Are predators tied to the health of the main prey?
3. Can predators switch prey?
4. Ramifications to reestablishment
iv. Results: No major effects
b. Predation
i. Traditional view: carnivory
ii. Differences from herbivory
1. Herbivory is nonlethal
iii.
Differences from parasitism
1. In parasitism, one individual is utilized for the development
of more than one parasite
iv. Predatorprey associations
1. Figure 10.2
3. Antipredator Adaptations
a. Aposematic or warning coloration
i. Advertises an unpalatable taste
ii. Ex. Blue jays and monarch butterflies
1. Caterpillar obtains poison from milkweed
2. Blue jays suffer violent vomiting from ingesting caterpillar
iii.
Ex. Tropical frogs
1. Toxic skin poisons
2. Figure 10.3a
b. Camouflage
i. Blending of organism into background color
ii. Ex. Grasshoppers (Figure 10.3b)
iii.
Ex. Stick insects mimic twigs and branches
iv. Ex. Zebra stripes: blend into grassy background
c. Mimicry
i. Animals that mimic other animals
1. Ex. Some hoverflies mimic wasps
ii. Types of mimicry
1. Mllerian mimicry
a. Fritz Mller, 1879
b. Unpalatable species converge to look the same
i.
Reinforce basic distasteful design

ii.
iii.

d.

e.

f.

g.

h.

Ex. Wasps and some butterflies

Mimicry ring: a group of sympatric species,
often different taxa, share a common
warning pattern
2. Batesian mimicry
a. Henry Bates, 1862
b. Mimicry of unpalatable species by palatable species
c. Ex. hoverflies resemble stinging bees and wasps
(Figure 10.3d)
3. Difficulty distinguishing type of mimicry
a. Monarch butterflies and viceroy butterflies (Figures
10.3d,e)
Displays of intimidation
i. Ex. Toads swallow air to make themselves appear larger
ii. Ex. Frilled lizards extend their collars to produce the same effect
(Figure 10.3f)
Polymorphism
i. Two or more discrete forms in the same population
ii. Color polymorphism
1. Predator has a preference (usually the more abundant form)
2. Prey can proliferate in the rarer form
3. Ex. leafhopper nymphs (orange and black)
4. Ex. Pea aphids (red and green)
iii.
Reflexive selection
1. Every individual is slightly different
2. Examples: brittle stars, butterflies, moths, echinoderms, and
gastropods
3. Thwart predators learning processes
Prey phenologically separated from predator
i. Ex. Fruit bats
1. Either diurnal or nocturnal
2. Only nocturnal in the presence of predatory diurnal eagles
Chemical defense
i. Used to ward off predators
ii. Ex. Bombardier beetles
1. Possess a reservoir of hydroquinone and hydrogen peroxide
2. When threatened, eject chemicals into "explosion chamber"
3. Mix with peroxidase enzyme
4. Mixture is violently sprayed at attacker
Masting
i. Synchronous production of many progeny by all individuals in
population
ii. Satiate predators
iii.
Allows for some progeny to survive
iv. Common to seed herbivory
v. Ex. 17-year and 13-year periodical cicadas

i. Comparison of defense mechanisms

i. Table 10.1, chemical defense is most common
4. PredatorPrey Models
a. Effects of predators on prey
b. Depend on such things as prey and predator densities, and predator
efficiency
c. Graphical method to monitor relationship
i. Prey isoclines have characteristic hump shape
1. Figure 10.4
2. In the absence of predators, prey density would be equal to
the carrying capacity, K1
3. Lower limit, individuals become too rare to meet for
reproduction
4. Between these two values, prey populations can either
increase or decrease depending on predator density
5. Above the isocline, prey populations decline
6. Below the isocline, prey populations increase
ii. Predator isoclines
1. Threshold density, where predator population will increase
2. Predator population can increase to carrying capacity
3. Mutual interference or competition between predators
a. More prey required for a given density predator
b. Predator isocline slopes toward the right
iii.
Superimpose prey and predator isoclines
1. Figure 10.5
2. One stable point emerges: the intersection of the lines
3. Three general cases
a. Inefficient predators require high densities of prey
(Figure 10.5a)
b. A moderately efficient predator leads to stable
oscillations of predator and prey populations
(Figure 10.5b)
c. A highly efficient predator can exploit a prey nearly
down to its limiting rareness (Figure 10.5c)
4. All based on how efficient predator is
5. Shift in isoclines
a. Prey starvation (shift to left)
b. Food enrichment (shift to right) (Figure 10.5d)
i.
Carrying capacity changes
ii.
Predator isocline changes"paradox
enrichment": Increases in nutrients or food
destabilizes the system
d. Functional response
i. How an individual predator responds to prey density can affect
how predators interact with prey (Figure 10.6)
ii. Three types

1. Type I: Individuals consume more prey as prey density

increases
2. Type II: Predators can become satiated and stop feeding, or
limited by handling time.
3. Type III: Feeding rate is similar to logistic curve; low at
low prey densities, but increases quickly at high densities
iii.
Changes in prey consumption
1. Functional response changes (Figure 10.7)
a. Dictates how individual predators respond to prey
population
2. Numerical response changes
a. Governs how a predator population migrates into
and out of areas in response to prey densities
5. Field Studies of PredatorPrey Interactions
a. Field comparisons to models
b. Do predators control prey populations?
c. Importance of predators in controlling prey density
i. Kaibab deer herd
1. Kaibab Plateau (Northern Arizona)
2. Declared a national park around 1900
3. All big predators were removed and deer hunting was
prohibited
4. Estimates of tenfold increase in deer population
5. Reevaluated by Graham Caughley (1970)
a. Predator control had some impact; cessation of
hunting and removal of competing sheep and cattle
also had an impact
ii. Serengeti plains of eastern Africa
1. Large predators have little effect on large mammal prey
2. Most prey taken are either injured or senile
3. Contribute little to future generations
4. Prey are migratory
iii.
Moose population on Michigans Isle Royale
1. Wolf-free existence until 1949
2. Durwood Allen (1958) began to track wolf and moose
populations
3. Trends in populations (Figure 10.8)
a. Wolf population
i.
Peaked at 50 in 1980
ii.
Severe nosedive in 1981
iii.
Small recovery in the late 1990s
b. Moose population
i.
Increased steadily in the 1960s and 1970s
ii.
Declined as the wolf population increased
until 1981

iii.

A record population of 2500 was reached in

1995, when the wolf population was low
iv. Good evidence of prey population control by
predators
v. Confounded in 1996 when the moose
population crashedstarvation
iv. Canada lynx and snowshoe hare
1. Populations show dramatic cyclic oscillations every 9 to 11
years (Figure 10.9)
2. Cycle has existed as long as records have existed (over 200
years)
3. An example of intrinsically stable predatorprey
relationship
6. Introduced Predators
a. Method for determining the effects of predators
b. Dingo predations on kangaroos in Australia
i. Dingo
1. Introduced species
2. Largest Australian carnivore
3. Predator of imported sheep
4. Eliminated from certain areas
a. Spectacular increases in native species
i.
160-fold increase in red kangaroos
ii.
Over 20-fold increase in emus
5. Effects on feral pigs
a. Shortage of young pigs
b. Considerable impact on recruitment of pigs (Figure
10.10)
c. European foxes and feral cats in Australia
i. Damage domestic livestock
ii. Effects when removed (Figure 10.11)
d. Lamprey and the Great Lakes
i. Construction of Welland Canal allowed lamprey to enter the Great
Lakes
ii. Dramatic reduction in lake trout (Figure 10.12)
iii.
Trout recovered after lamprey population was reduced
7. Field Experiments with Natural Systems
a. Lions in South Africa
i. Kruger National Park, 1903
ii. Lions shot
iii.
Number of large prey increased
iv. Shooting of lions ends, 1960
v. Wildebeast increase so much that their numbers had to be culled
from 1965 to 1972
b. Gray partridge, European game bird
i. Figure 10.13

ii.
iii.

Over 20 million shot in Great Britain in the 1930s

Only 3.8 million shot in the mid-1980s
1. High chick mortality due to starvation
2. Reduced insects due to introduction of herbicides in the
1950s was suspected
3. However, smaller populations in areas where there was no
control of predators by gamekeepers
4. Predation control increased
a. The number of partridges that bred successfully
b. The average size of the broods
c. Partridge populations by 75%
c. Predators and rodents in Finland
i. Large-scale removal of predators, April 1992 and 1995 over 23
km2
ii. Large increase in rodent population by June (compared to control
plots) (Figure 10.14)
8. Applied Ecology
a. Humans as predatorswhaling
i. Exploitation necessary
ii. Is harvesting at any level sustainable?
b. History of Antarctic whaling
i. Figure 1
ii. 1930s, blue whales primarily harvested
iii.
1950s, blue whale population depleted, replaced with fin whale
iv. 1960s, fin whale population collapsed
v. 1960s, humpback whale population collapsed
vi. Prior to 1958, Sei whales hardly ever harvested
1. Reduction in other whales made Sei whale attractive
2. Peak harvest of about 20,000 by 196465
3. Catches declined thereafter due to limitations
vii. The relatively small minke whale
1. Was ignored in the southern oceans until 197172
2. Began to be taken, and is now the largest component of the
southern baleen whale catch
viii.
Whale ban proposed in 198586, took effect in 1988
ix. Iceland, Norway, and Japan, 1994
1. Argued for resumption of limited commercial whaling
c. Should we ban commercial whaling?
d. Whale populations are recovering
i. Ex. Blue whale populations have increased fourfold
ii. Ex. California grey whales have recovered to prewhaling levels
9. Summary
a. Predation is a strong selective force in nature
i. Aposematic coloration
ii. Camouflage
iii.
Batesian and Mllerian mimicry

iv. Intimidation displays

v. Polymorphisms
vi. Chemical defenses
b. Modeling predatorprey interactions
i. Even simple predatorprey models show
1. Decreased predatorprey oscillation
2. Stable cycles
3. Wildly increasing and unstable oscillations
ii. Difficulty in predicting or modeling how predators and prey
interact
1. Mutual interference between predators
2. Existence of specific predator territory sizes
3. Ability of predators to feed on more than one type of prey
c. Large-scale observations support
i. Predators only take weak and sickly individuals
ii. Prey populations influence predator numbers, not vice versa
d. Accidental or deliberate introductions of exotic predators
i. Profound effects on native prey populations
ii. Predators have important regulatory effects on prey
iii.
May not be indicative of "natural systems"
e. Evidence from natural systems
i. Most studies have concluded that predators have a significant
effect on prey
10. Discussion Questions
a. Should ranchers be concerned about the reintroduction into their vicinity
of large predators, like wolves and panthers?
b. Do sea lions, otters, or dolphins decrease the stock of fish available for
people that fish?
c. Would the number of deer available for hunters be the same in the
presence of large predators?
d. What data would you need to collect to answer the above three questions?
e. What can the effects of exotic predators tell us about the strength of
predation? What cant they tell us?
f. Which do you think more likely: that predators control prey populations or
that prey control predator populations? Would the answer vary according
to the particular system? Give an example.
g. What shortcomings do you think Rosenzweig and MacArthurs predator
and prey isoclines have? What would these shortcomings mean in terms of
determining how predators and prey interact?
h. A great many fish stocks seem to have been overfished. How do you think
we could prevent overfishing? What biological information do we need to
have, and how can we get it when we cant see the population in question?

Chapter 11: Herbivory

1. Outline
a. Plant chemical and mechanical defenses against herbivores
b. Mathematical models predict polyphagous herbivores have a greater effect
on plants than monophagous herbivores
c. Herbivores remove 1518% of terrestrial plant biomass; over 50% in
aquatic systems
d. Herbivore numbers are strongly influenced by chemical and mechanical
plant defenses, and the amount of nitrogen in the plant
e. Herbivores can change plant communities by preferentially eating
dominant plant species
2. Plant Defenses
a. Example of plant chemical defenses
i. Alkaloids
ii. Mustard oils
iii.
Terpenoids
iv. Phenylpropanes
v. Figure 11.1
b. Two general classes of compounds
i. Carbon nutrient balance theory
ii. Nitrogen compounds
1. Limited by carbon (due to shortages of light or water)
iii.
Carbon compounds
1. Limited by nitrogen
iv. Table 11.1
c. Secondary chemicals
i. Not part of primary metabolic pathways that plants use to obtain
energy
ii. Defense compounds
d. Strategies of plant defenses
i. Quantitative defenses
1. Substances that are ingested in large amounts by the
herbivore
2. Prevent digestion of food
3. Ex. Tannins and resins
4. Figure 11.2
ii. Qualitative defenses
1. Highly toxic substances
2. Very small doses can kill herbivore
3. Ex. Atropine
4. Figure 11.3
iii.
Apparency
1. Correlated to qualitative and quantitative defense strategies
2. Apparent plants
a. Long-lived

iv.

v.

vi.

vii.

b. Apparent to herbivores (mainly insects)

c. Defenses are mainly quantitative
i.
Effective against specialist and generalist
ii.
Long history of association with these Kselected plants
3. Unapparent plants
a. Weeds
b. Ephemeral and unavailable for long periods of the
year
c. Defenses are mainly qualitative
d. Herbivores are mainly generalist like vertebrates
4. Table 11.2
Mechanical defenses
1. Thorns and spines deter vertebrate herbivores
2. Generalizations (Peter Grubb)
a. In many open sites, plants are primarily close to the
ground and so are very spinose to protect them
b. Plants such as palms, with one or a few apical
meristems, are also likely to protect them with
spines
c. Evergreens, such as holly, in a deciduous forest are
likely to face severe herbivore pressure in the winter
and so are very spinose
Repellents
1. Thistles produce compounds that repel certain insect larvae
2. Potatoes synthesize a component of an alarm pheromone
released by aphids when they are attacked by predators
a. Alarm pheromone causes aphids to flee
b. As a result, the potatoes are free of the aphids
Reproductive inhibition
1. Plants (e.g., firs) contain insect hormone derivatives
a. If digested, prevent insect metamorphosis
b. Results in diminished reproductive output
2. Plants (e.g., floss flower) produce a chemical mimic of
insect molting hormone ecdysone
a. Insects digest plant material
b. Insects die when they molt prematurely
Masting
1. Occurs in a few tree species (e.g., some oaks)
2. Production of more seeds in some years
a. Satiates herbivores
b. Permits more seed to surviveEx. Beech
i.
In mast years, 3.1% of seeds destroyed by
boring moths
ii.
In non-mast years, 38% of seeds destroyed
c. Other benefits

i.
Enhanced pollination
ii.
Enhanced seed dispersal
viii.
Defensive associations
1. Protection from herbivores through association with
unpalatable neighbors
2. Ex. Chrysomelid beetle and the purple loosestrife
a. Loosestrife sometimes grows on its own, or in
thickets of an aromatic shrub, Myrica gale
b. Myrica secretes a volatile chemical that deters
insects from feeding on it
c. Chemical also interferes with beetle searching for
loosestrife
d. Figure 11.4
3. Opposite association: Associational susceptibility
a. The spilling over of herbivores from palatable
neighbors
b. Fall cankerworms prefer to feed on box elder trees
and rarely feed on isolated cottonwood trees.
c. When cottonwoods occur under box elder, the
cankerworms spill over and defoliate the
cottonwoods
d. Figure 11.5
ix. Mutualism
1. Plants that defend themselves through enlisting help from
other animals
e. Understanding plant defenses
i. Important to agriculture
1. Use knowledge to defend crops against insects
2. Problems
a. Long time needed to develop resistances
b. Resistance to one pest may increase susceptibility to
other pests
3. Benefits
a. Once resistance is developed, requires minimal cost
from farmer
b. Environmentally benign
4. Insect response to resistance
a. Certain chemicals that are toxic to generalist insects
actually increase the growth rates of adapted
specialist insects
b. Specialization of herbivores to supposedly toxic
plantsevolutionary "arms race" (Diagram)
c. Figure 11.6
3. Modeling Herbivory
a. Series of plantherbivore interactions models developed by Mick Crawley
(1997)

b. Models depend on the degree of polyphagy of the herbivores

i. Monophagous herbivores are highly specialized and feed on only
one species
1. Many insects tend to be specialized
ii. Polyphagous herbivores feed on many species
1. Many vertebrates tend to be generalists
2. Exceptions: Pandas and bamboo, and koala and eucalyptus
c. Models for monophagous herbivores
i. Simplest model assumes a carrying capacity, K, for the plants
ii. Rate of change in plant population
1. dV/dt = A - B
a. dV/dt = rate of plant increases (numbers or biomass)
b. A = gains by growth
c. B = losses by herbivory
2. dN/dt = C - D
a. dN/dt = rate of herbivore increase
b. C = gains by growth
c. D = losses by death
3. In the absence of herbivores, plant populations increase
logistically
a. dV/dt = rV[(K - V)/K]
b. r = plants intrinsic rate of increase
4. Losses for plants
a. B = bNV
i.
b = feeding rate of herbivores
ii.
bNV = functional response
iii.
Type I: accurately modeled by bNV
5. Herbivore population growth
a. C = cNV
i.
c = numerical response of herbivores; their
degree of attraction is affected by plant
density
ii.
cNV is essentially a measure of efficiency
with which herbivores turn food into
progeny
6. Losses for herbivores
a. D = dN
b. d = herbivore death rate
c. Assumption: in the absence of plants, the herbivores
starve and their numbers exponentially decline
7. At equilibrium:
a. DV/dt and dN/dt are zero
b. No population changes
c. A = B and C = D
d. V* = d/c

i.

Paradoxical results: plant equilibrium

abundance has nothing to do with plant
growth rate or carrying capacity
ii.
Figure 11.7a
d. Models for polyphagous vertebrates
i. Constant level of herbivoryh
1. Herbivore densities are not tied to any one plant
ii. Plant growth
1. dV/dt = rV[(K - V)/K] - h
2. For equilibrium plant biomass, V*
a. V = (aK r2K2 - 4rhK)/2r
b. Figure 11.7b
e. Caveats to models
i. Misleading: c and d are treated like independent parameters. Could
lead to unrealistic results
ii. There are at least 3 different types of functional response
4. Effects of Herbivory on Plants
a. Herbivore responses to plant defenses
i. Detoxify plant poisons
1. Oxidation
2. Conjugation
ii. Measured effects of herbivores on plants in the field
1. In 93 cases, 7% of leaf area was consumed
2. In forested systems, 515% defoliation by insects
3. May underestimate effects due to leaf turnover
4. Herbivory increases with plant productivity (Figure 11.8)
b. Removal experiments
i. Best way to estimate effects of herbivory on plants
1. Remove herbivores
2. Examine effects on plant growth and reproduction
3. Review of 246 experiments (Bigger and Marvier, 1998)
a. Meta-analysis statistical technique
i.
Mean effect size d was 0.47, which is a
medium-size effect
ii.
Meaning: the average size of plant protected
from herbivores was significantly larger
than those that were not
b. Stronger effects in aquatic systems
i.
Figure 11.9
c. Which type of plants are most affected?
i.
Effect size of herbivores was greatest on
algae (Figure 11.10), also on grasses and
shrubs
ii.
Effect size of herbivore was smallest on
woody plants
c. Biological control

i.
ii.

Evidence from biological control of weeds in agriculture

50% of the 190 major weeds in the U.S. are invaders from outside
the country
iii.
Interest in biological control due to expense of chemical control
iv. Biological control success stories
1. Klamath weed was controlled by two French beetles
2. A floating fern, Salvinia molesta, was controlled by the
weevil Cyrtobagus salvinae
3. The Florida alligator weed was controlled by the
alligatorweed flea beetle
4. Figure 11.11
v. Unsuccessful biological control stories
1. Control of Lantana camara in Hawaii
2. In a review of 701 examples of biological weed control,
only 26% were rated successful
d. Beneficial herbivory
i. Review of herbivory (Bigger and Marvier, 1998)
1. 60 comparisons demonstrated a reduction in plant size due
to natural levels of herbivory
2. 10 comparisons demonstrated an increase in plant size
a. Plants are stimulated to regrow after damage, and
may overcompensate
ii. Benefits of root herbivory (Figure 11.12)
iii.
Benefits of grazing and regrowth
1. More flowers and fruits on grazed plants (Gronemeyer et
al., 1997)
2. More likely to occur in perennials than annuals
5. Effects of Plants on Herbivores
a. Herbivores select those plants that have the most nutrition (in terms of
nitrogen)
b. Animal tissue contains 10 times more nitrogen than plant tissue
c. Red deer prefer to feed on grasses that were defecated upon by herring
gulls
i. As the amount of droppings increased, the nitrogen content of the
grasses also increased
d. Plant fertilization (i.e., nitrogen) had a positive effect on population size,
survivorship, growth, and fucundity of insects (Figure 11.13)
i. Response was greater in cultivated versus wild plants, and
broadleaf trees versus conifers (Figure 11.14)
e. Two variations of the nitrogen limitation theory
i. Stress hypothesis (White 1993)
1. Plant stresses tend to increase the availability of nitrogen
because many nitrogen-rich compounds are mobilized in
response to stress
a. Ex. Drought stressed plants accumulate high
numbers of herbivores

ii.

Plant vigor hypothesis (Price 1991)

1. Herbivores select fast growing parts of plants or fast
growing plants because these are higher in nitrogen
a. Ex. Many attacks by insects are on young trees
f. Herbivore density correlated with plant quality
g. Herbivore population patterns are dependent on other factors
i. Predation
ii. Parasitism
6. Herbivory Affects Community Structure
a. Herbivory can affect community richness, succession, and ecosystem
function
b. Darwin observed that competitively dominant grasses were kept in check
by grazing
c. United States and bison (Figure 11.15)
7. Applied Ecology
a. Pest control around the world
b. Huge monocultures are ideal targets for insect pests and diseases
c. Wide variety of control techniques (Table 1)
d. Pesticides in the U.S.
i. 50,000 pesticides are registered for use in the U.S.
ii. Five to six hundred million kg of pesticides used each year
iii.
70% for agriculture, 23% for forestry, 7% for home and garden
iv. Many are strong poisons
v. 60% of herbicides and 30% of insecticides are potentially
oncogenic
vi. 95% of human tissue samples contain pesticide residues
e. Effects on nontarget organisms
8. Summary
a. A variety of plant defenses demonstrate the strength and frequency of
herbivory in nature
i. Chemical defenses
ii. Mechanical defenses
iii.
Hormone mimics
iv. Mutualism
b. Chemical defenses
i. Quantitative
1. Build up in the gut of the herbivore preventing digestion
ii. Qualitative
1. Toxic compounds that can be lethal in small doses
c. Mathematical models
i. Effects of herbivores on plants depends whether they are
monophagous or polyphagous
d. Effects of herbivores
i. Herbivores remove 1518% of terrestrial plant tissue
ii. Herbivores remove 51% of aquatic plant tissue
iii.
Impact of biological control

iv. Impact in agriculture

e. Population densities of herbivores are strongly influenced by plant quality,
particularly plant nitrogen
f. Herbivory may have substantial effects on plant communities
9. Discussion Questions
a. Cannabin and cocaine are secondary chemicals produced by the hemp
plant (Cannabis sativa) and the coca plant, respectively. What do you
suppose are the roles of these substances in the lives of their plants?
b. From a theoretical standpoint, think about some differences that might
occur between monophagous and polyphagous herbivores. Why are the
majority of mammals polyphagous and yet many insects are
monophagous?
c. Given the advantages and disadvantages of different forms of pest control,
which do you think would be more suitable for annual crops? For
orchards? For forests? Why?
d. Do you think chemical defenses would be more likely to occur in lush
tropical leaves that have no other defenses or in tough desert plants that
can ill afford the losses to herbivores.
e. Do you think that if plants need to protect their valuable tissue from losses
to herbivores, flowers and fruits would have high levels of secondary
chemicals? Explain.

Chapter 12: Parasitism

1. Outline
a. Parasites feed on a host, but generally do not kill it
b. Hosts have evolved many defenses (e.g., immune responses) against
parasites
c. Models show that the rate of spread of diseases is governed by the density
of susceptibles in the population, the transmission rate of the disease, and
the length of life of the infected host
d. Parasites can substantially decrease host population size
e. Parasites can affect the structure of host communities
f. Parasitoids help in biological control by reducing the density of pests
2. Defining Parasites (Figure 12.1)
a. Parasite: a predatory organism that feeds off another but generally does
not kill it
b. Host: prey of a parasite
c. Parasitoid: Cases where the host does not survive but one host is
insufficient for the development of the parasitoid
d. Some parasites live with their host most of their lives (e.g., tapeworms)

e. Some parasites drop off after prolonged periods of feeding (e.g., ticks,
leeches)
f. Are mosquitoes and wildebeests parasites?
g. Some parasites are parasitic on other plants
i. Holoparasites: lack chlorophyll, and are totally dependent on
another plant for water and nutrients
1. Ex. Figure 12.2
ii. Hemiparasites: photosynthesize, but do not have a root system, so
they rely on the host for this function
1. Ex. Mistletoe
h. Monophagous: parasites that feed off one to three closely related species
i. Polyphagous: parasites that feed off many host species
i. Table 12.1
j. Ectoparasites: live on the outside of the hosts body (e.g., fleas and ticks)
k. Endoparasites: live inside the hosts body (e.g., tapeworms and bacteria)
l. Haustoria: plant parasite outgrowths that penetrate inside a host plant to
tap into its nutrient supply
m. Use of multiple hosts: fluke (Figure 12.3)
n. Parasites outnumber free-living species 4 to 1 (Figure 12.4)
3. Defense against Parasites
a. Cellular defense reactions
i. Eggs of parasatoids are rendered inviable by encapsulating them
b. Immune responses in vertebrates
i. Phagocytes may engulf and digest small alien bodies, and
encapsulate and isolate larger ones
ii. Hosts may develop a "memory," that may make them immune to
reinfection
c. Defensive displays or maneuvers
i. Actions intended to deter parasites
d. Grooming and preening behavior
i. Behavior intended to remove parasites
4. Modeling Parasitism
a. Differ from models of predation and herbivory
i. Life cycles of many parasites involve intermediate hosts
ii. Models of parasite population dynamics generally describe the
population growth rate by the average number of new disease cases
b. For microparasites, the number of infected hosts is the most important
factor
c. Rp = NBL
i. Rp = number of infected hosts, with p for parasite and R for net
reproductive rate
1. Transmission threshold; Rp = 1
2. For disease to spread; Rp > 1
3. For disease to die out; Rp < 1
4. Microparasites are transferred from host to host
ii. N = density of susceptible hosts in population

iii.
iv.
v.

B = transmission rate of disease

L = average period over which the infected host remains infectious
Generalizations
1. As L increases, so does Rp
2. If diseases are highly infectious, Rp increases
3. Large populations of susceptible hosts promotes the spread
of diseases
4. Critical threshold NT where Rp = 1
a. NT is an estimate of the number of susceptible hosts
needed to maintain the parasite population at a
constant size
b. NT = 1/BL
i.
If B is large, N is small
ii.
If B or L are small, the disease can persist
only in a large population
d. Many diseases undergo periodic cycles
i. Ex. Measles in England (Figure 12.5)
1. Peaks occur because host immunity is developed
2. New births lead to new susceptible hosts, and cycle repeats
e. Parasites spread by a vector
i. Life cycle of both parasite and vector become important in
controlling diseases
ii. Ex. Farmers use insecticides to kill aphids, which transmit viral
diseases to crops (rather than chemicals to kill the parasite)
iii.
Ex. Yellow fever was eradicated in the U.S. by inoculation rather
than eradication of all mosquitoes
5. Parasites Affect Host Populations
a. Using biological control to study the effects of parasites on hosts
i. Ex. Hawkins 1999: Biological control of pests, especially by
parasitoids, was greater in exotic, simplified, managed habitats
than in natural habitats
ii. Control is most often exerted by a single parasitoid species, in
contrast to natural systems, which require a suite of generalized
enemies
iii.
Thus, biological control projects cannot provide rigorous evidence
of the importance of parasites in natural systems
b. Effects of introduced parasites on natural systems
i. Chestnut blight in the Appalachian Mountains of North America
1. Virtually eliminated chestnut tree (Figure 12.6)
2. Introduced in New York in 1904
3. In Britain, 25 million elm trees (out of 30 million) were
wiped out by the disease between 1960s and the 1990s
(Figure 12.7)
ii. Rinderpest
1. A virus with at least 47 natural artiodactyl hosts, most of
which occur in Africa

2. The virus belongs to a class known as morbilliviruses,

which includes measles and distemper
3. Spread by food and water contaminated by dung of sick
animals
4. Can be fatal to certain animals (buffalo, eland, kudu, and
warthogs)
5. Major epidemic swept through Africa in the 1890s, leaving
vast areas uninhabited by certain species
6. 80% of hoofed stock died. Disease traveled 5000 km in
eight years
7. Brought under control in the 1960s, through the use of
cattle vaccinations
iii.
Endangered species
1. Many endangered species are threatened by diseases from
domestic animals (Table 12.2)
2. Ex. The demise of the marsupial wolf in Tasmania was
because of a distemper-like disease obtained from domestic
dogs
3. Some endangered species have been given vaccinations to
protect them from disease
a. Mountain gorillas were vaccinated for measles
c. Natural systems
i. Massive mortality of bighorn sheep from infection by lungworms
(Protostrongylus stilesi and P. rushi)
1. Predisposes animals to pathogens, which cause pneumonia
2. Infection rates of 91% and mortalities of 5075% have
been reported
ii. Colorado pine tree plantations and mistletoe. Mistletoe can cause
30% loss in extractable timber
iii.
Saline marshes in North America and the plant parasite, Cuscuta
salina (Figure 12.8a)
1. Infects the most common plant in California marshes,
Salicornia virginica, thus promoting the growth of two
other species, Limonium and Frankenia (Figure 12.8b)
iv. Parasite removal experiments
1. Fuller and Blaustein (1996) compared the survivorship of
parasite infected and uninfected free-living deer mice
a. Conducted in large outdoor enclosures
b. Decreased overwinter survivorship for those deer
mice infected with the protozoan Eimeria
arizonensis
c. Contamination spread through the digestion of
contaminated feces
2. Hurtrez-Bousss et al. (1997) reduced the number of
blowfly larvae parasites in young blue tits in Corsica

a. Blowfly larvae suck blood from chicks, causing

anemia and high mortality
b. Removal was accomplished by removing nests from
nest boxes, and microwaving the nests to kill the
parasites, and then returning the nests and chicks
c. Chicks from microwaved nests were found to have
greater body weight at fledging (Figure 12.9)
3. Stiling and Rossi (1997) manipulated parasitic infection
levels of a gall-making fly on a coastal plant, Borrichia
frutescens, on isolated islands off the coast of Florida
a. Low rates of parasitism treatment
i.
Allowed potted plants on one island to be
colonized by gallflies
ii.
Plants were removed before parasitoids
could find them
b. High rates of parasitism treatment
i.
They left plants on the island longer, to
allow the parasites to colonize the galls
c. Results: High degree of parasitism of gallflies
resulted in a significant reduction in the number of
new galls (Figure 12.10)
6. Parasites Affect Communities
a. Parasites affect the presence or absence of various species in a community
b. The meningeal brainworm Parelaphostrongylus tenuis
i. Usual host is the white-tailed deer, which is tolerant of the
infection
ii. All other cervids and the pronghorn antelope are potential hosts
1. Worm causes severe neurological damage
iii.
The worm makes the white-tailed deer a potential competitor with
other cervids, because they cannot survive in the same area as the
white-tailed deer. This phenomenon is known as apparent
competition
7. Parasites and Biological Control
a. Not all parasites are detrimental to humans
b. Many are used to protect crops from pests: Biological control
i. Only 16% of classical biological control would qualify as
economic successes
ii. Organisms used in biological control, are released in a "hit or
miss" technique: Just release a bunch of parasites and predators,
and hope that one of them does the job.
iii.
New techniques: Ex. novel parasitehost associations
iv. Review of 548 control projects: the more parasites that were
released, the lower the rate of establishment
c. Necessary attributes of a good agent of biological control (Huffaker and
Kennett, 1969)
i. General adaptability to the environment and host

ii. High search capacity

iii.
High rate of increase relative to the hosts
iv. General mobility adequate for dispersal
v. Minimal time lag effects in responding to changes in host numbers
d. Methods affecting the success of biological control (Stiling, 1990)
i. Factor of greatest importance: climatic match between the control
agents locality of origin and the region in which it will be released
ii. Importance of climatic variation was underscored by another
review of biological failures (Stiling, 1993)
1. Climate accounted for 34.5 % of the failures (Figure 12.11)
e. Risks of biological control
i. Reduction in native Hawaiian lepidopterans was partly due to wasp
species introduced for biological control of lepidopteran crop pests
1. Wasps attacked target exotic species but also nontarget
native species
2. Stresses the importance of more narrowly focused release,
rather than the traditional "hit and miss" technique
ii. Problems with crops
1. While it is of interest to ensure that the agent does not
adversely affect the crop, however, non-crop plants are not
as vigorously tested
8. Applied Ecology
a. Importance of parasites to plants and animals
b. Five main categories of disease-causing organisms that most plants and
animals are susceptible to (Table 1)
c. Leading causes of human death by disease, worldwide (Table 2)
9. Summary
a. The true definition of parasite is problematic. Parasites may include many
species that feed on plants, plus more traditional parasites such as
tapeworms, leeches, bacteria, viruses, and parasitoids that attack animals.
80% of all life-forms are considered parasitic
b. The presence of various parasite defense mechanisms is testament to the
importance of parasitism in nature
c. Mathematical models suggest that effective parasites will keep their hosts
alive as long as possible, thereby facilitating the transmission of parasites
to additional hosts.
d. The huge influence of introduced diseases, such as chestnut blight and
Dutch elm disease, provides evidence for the severe effects that parasites
can have on a hosts population and host density
e. Parasites of insects can often be used as control mechanisms against crop
and forest pests. This technique is termed biological control. Finding the
attributes of successful biological control agents is valuable.
Unfortunately, biological control agents can have a significant impact on
nontarget natural populations
10. Discussion Questions

a. Discuss some of the main differences between microparasites,

macroparasites, parasitoids, predators, and herbivores in terms of their life
history strategies
b. Why can we eradicate some diseases, such as yellow fever, through
vaccinations, while we have not been able to eradicate diseases such as
malaria?
c. Can we ever expect chemical pesticides to be replaced entirely by
biological control agents? Explain.
d. Parasites are usually very small organisms, capable of passing through
screen enclosures. With this in mind, how might you design an experiment
to remove parasites from a population and compare that populations
survival rates with those of an unmanipulated control population?
e. Do you think that biological control would be more successful on islands
or on mainland continental areas? Against native or exotic pests?

Chapter 13: Evaluating the Controls on Population Size

1. Outline
a. Mortalities that perturb population densities away from mean or
equilibrium values are known as key factors. Among the many key factors
are predation, competition, parasitism, and herbivory
b. Mortalities that act to return population densities to mean or equilibrium
levels are known as density-dependent factors. As with key factors, there
are many different types of density-dependent factors.
c. Groups of populations may exist with many individuals intermixing
between the groups. A collection of such groups is known as a
metapopulation, and the dispersal rate, not mortality, is a key to
understanding its group dynamics
d. Simple theoretical models that use only the productivity and number of
trophic levels in a system have been alleged to be able to predict when and
where various mortalities are likely to be important
2. Introduction
a. Migrating wildebeest (Figure 13.1)
i. The most common herbivore in Africa
ii. Wildebeest population in the Serengeti region of Tanzania
1. Decline in population from 1.2 million to 0.9 million.
Decrease due to:
a. Increased poaching
b. Change in climate
c. Renewed drought
d. Changes in the rate of predation
2. 40-year study (Mduma et al., 1999)

a. Predation played a minor role in decline (>3%)

b. Illegal harvesting accounted for 20,000 per year
c. Main cause of mortality: malnutrition brought on by
drought
b. Long-term detailed studies are needed to disentangle the effects of
multiple factors on populations
3. Comparing the Strengths of Mortality Factors
a. Number of factors can potentially affect populations away from the mean
or equilibrium
i. Common biotic forces
1. Ex. Competition, predation, parasitism, herbivory, and
mutualism
2. If biotic forces are of overriding importance, then
communities may be tightly knit entities
ii. Climate and weather
1. If abiotic forces are the most influential, then community
structure may be loose and ephemeral
b. Terms for population change
i. Extrinsic factors, such as weather, parasitism, and predation
ii. Intrinsic factors, such as endocrine and immune systems
c. Key factors
i. Definition: those factors which can disturb a population away from
the mean or equilibrium
ii. Key-factor analysis: technique for determining the importance of a
variety of factors affecting a population
1. First developed by R. F. Morris (1957)
2. Requires detailed information on the fate of a cohort of
individuals
a. Total mortality of a generation or cohort (K) is
divided into various causes, and the relative
importance of these causes are compared
iii.
Ex. Oak winter moth (Varley and Gradwell)
1. Table 13.1
2. Figure 13.2
3. Caterpillars were not actually counted
a. Passive-sampling techniques were used
b. Estimated caterpillars based on the number of silken
threads they put down to pupate on the soil
c. Used metal traps on the soil, to estimate the number
of caterpillars per m2
d. Determined number of healthy vs. sick caterpillars.
Some sick caterpillars were infected with a parasite
(Cyzenis). Number of sick caterpillars was usually
less than 10%
4. Some caterpillars died after they arrived in the soil to
pupate

a. Parasite (Craitichneumon) caused substantial

mortality, between 4050%
b. Predators (shrews and beetles) can eat between 60
70% of the pupae
c. Number of healthy pupae was determined by using
inverted metal traps to estimate the number of
emerging moths per m2
5. At each stage, the number of deaths and cause of death was
recorded
6. The importance of each mortality factor (k) was estimated
by calculating the amount that the factor reduced the
population
a. k = log Nt - log N(t + 1)
i.
Nt = density of population before it is
subjected to the mortality factor
ii.
N(t + 1) = population afterward
b. Ex. for pupal parasite of the oak winter moth
i.
k5 = log 28.4 - log 15.0 = 0.27
c. Ex. Mortality from overwintering
i.
k1 = log 658 - log 96.4 = 0.84
7. Total generational mortality, K is equal to the sum of ki
a. K = k1 + k2 + k3 + k4 + + kn
8. Overwintering loss is the key factor (Figures 13.3 and
13.3b)
iv. Other species and key factors
1. Table 13.2
2. No key factor of overriding importance
v. Criticisms of key factor
1. Key factors cannot always be precisely linked to specific
mortality agents
2. Intricate interactions between natural enemies, including
hyperparasitoids, and such factors fail to show up in a key
factor analysis
3. Populations can be influenced greatly by egg-bearing
females that disperse into a population and that also never
show up in key factor analysis
4. Various other ways to look at life table data (Table 13.3)
d. Real mortality
i. Definition: Mortality of a population compared with the population
size at the beginning of the generation
ii. Useful in comparing population factors within the same generation
iii.
Real mortality is generally greater for factors that operate early in
the organisms life cycle, because more individuals are actually
killed
e. Indispensable (or irreplaceable mortality)

i.

Definition: Part of the generational mortality which would not

occur if a mortality factor was removed from the life system (after
allowance is made for the action of subsequent mortality factors)
ii. Example: Conservation of sea turtles
1. 1000 eggs are laid
2. Predators (seagulls) eat 500 (50%) hatchlings
3. Predators (fish predators) eat 400 of the remaining 500
turtles (80%) before they become reproductive adults
4. 90 out of the remaining reproductive adults (90%) are lost
in fishing nets
5. 10 surviving adults (Figure 13.4)
6. Best method for conserving turtles
a. Protecting turtle eggs, allowing 1000 turtles to make
it to the sea would result in 40 surviving adults
b. Better to protect adults from fishing nets100
surviving adults
f. Mortalitysurvivor ratio
i. Definition: The increase in population that would have occurred if
the factor in question had been absent
ii. If the final population is multiplied by the mortalitysurvivor ratio,
then the resulting value represents, in individuals, the
indispensable mortality due to factor
4. Density Dependence
a. Predation, parasitism, competition, and abiotic factors can affect
population densities
b. Population densities can remain stable for long periods of time (Figure
10.12), therefore there must be factors that stabilize population density
i. Difficulty in determining these factors
1. It is necessary to compare different mortality factors
2. It is appropriate to determine which factors act in a density
dependent manner
c. Density dependence can be determined graphically (Figure 13.5)
i. Positive slope (population density vs. percent mortality)
1. Mortality increases with density
2. Mortality factor is acting in a density dependent manner
ii. Positive slope (k values vs. Log N)
1. Density dependence
2. Ex. Overwintering moth (Figure 13.6)
iii.
Factors that do not change with density are termed density
independent
1. Ex. Bacteria infect 10% of the population, regardless of
density
iv. Sources of mortality that decrease with increasing population size
are termed inversely density dependent
1. Ex. A lion will take the same number of prey, regardless of
density, because it is territorial

d. Determining which factors act in a density dependent manner

i. Ex. Examining 58 species of insects (Stiling, 1988)
1. The most important density-dependent mortality factor was
different for different species at different times
2. Generalizations are difficult to make
ii. Ex. Review of 51 populations of insects, 82 of large mammals, 36
of small mammals and birds (Sinclair, 1989) (Figure 13.7)
1. Insects showed a wide variety of causes of density
dependence
2. Larger taxonomic groups
a. Food was important for large mammals
b. Space and social interactions were important to
small mammals and birds
3. r-selected species with very high reproductive rates and
Type III survivorship curves (insects and fish) have early
juvenile density-dependent mortality
4. Species with intermediate reproductive rates and Type II
survivorship curves (birds and small mammals) have late
juvenile and prebreeding regulation
5. K-selected species with low reproductive rates and Type I
survivorship curves (large mammals) are at least partly
regulated through changes in fertility (Table 13.4)
iii.
Spreading the risk may be a good explanation for the apparent
random patterns relating mortality to density
iv. Animals may not always behave in a density-dependent manner
due to conflicting pressures
1. Ex. African hunting dogs may not prey on antelopes in a
density-dependent manner because of the presence of a lion
pride which could prey on the hunting dogs
v. No consensus on the frequency of density dependence in nature
1. When found, it offers control of population
2. No simple answer as to which factor will most likely affect
population densities
3. A complex of factors participates in the regulation of most
organisms
5. Metapopulations
a. A metapopulation is a series of small, separate populations that mutually
affect one another
b. Metapopulations are viewed as sets of populations persisting in a balance
between local extinction and colonization
c. Relevance to population biology: populations could be maintained by a
balance between local extinction and colonization. There is no mean or
equilibrium, local extinction could occur at any time, and density
dependence is irrelevant
d. Persistence depends on factors affecting extinction and colonization

i.

e.

f.

g.

h.

Ex. Interpatch distances, species dispersal abilities, and number of

patches in the metapopulation
Mathematical model of metapopulation dynamics (Levin, 1969)
i. dp/dt = mp (1 - p) - xp
1. dp/dt = rate of change in occupied patches
2. m = movement rate between patches
3. x = extinction rate
4. p = proportion of patches occupied
Harrison's (1991) review of empirical literature revealed few situations
that fit classical description of a metapopulation (Figure 13.8). More
common were:
i. Core-satellite, source-sink, or mainland-island populations, in
which persistence depended on the existence of one or more
extinction-resistant populations
ii. Patchy populations, in which dispersal between patches or
populations was so high, that colonists always "rescued"
populations from extinction
iii.
Nonequilibrium metapopulations, in which local extinctions
occurred in the course of species overall regional decline
Metapopulation study: Bay checkerspot butterfly (Harrison et al., 1988)
(Figure 13.9)
i. Metapopulation consisted of a population of 106 adult butterflies
on a 2000-ha habitat (Jasper Ridge near Stanford University), and
nine populations of 10 to 350 adult butterflies on patches of 1 to
250 ha
ii. Of 27 small patches that were suitable for populations to live in,
only those close to the large patch were occupied
iii.
Difference could not be explained by the quality of habitats
iv. Distance effect appeared to indicate that the butterflies capacity
for dispersal was limited
1. Large patch was dominant source of colonists to the small
patches
2. Persistence in this population was relatively unaffected by
turnover in small populations
3. Small populations acted as sink populations
4. In 1996, the population disappeared from Jasper Ridge
Metapopulation theory has had a long history
i. A study of a phytophagous mite as prey, and a predatory mite
(Huffaker, 1958; Huffaker et al., 1963)
1. Laboratory study, examining spatial heterogeneity using
oranges and Vaseline barriers
2. Prey was able to keep one step ahead of predators
3. At any one time, there was a mosaic of:
a. Unoccupied patches
b. Patches of prey and predators headed for extinction
c. Patches of thriving prey

4. The mosaic was capable of maintaining persistent

populations of predators and prey (Figure 13.10)
6. Conceptual models of population control
a. Bottom-up factors, or trophodynamics
i. Factors that act from the bottom of the food chain, for example
food (Figure 13.11)
ii. Tropic-level concept or trophodynamics (Lindeman, 1942)
1. Explained the height of the trophic pyramid by reference to
a progressive attenuation of energy passing up through
trophic levels
2. Based on the thermodynamic properties of energy transfer
b. Top-down factors
i. Factors which percolate down from the top of the food chain, for
example, natural enemies
ii. Hairston, Smith, and Slobodkins (1960) hypothesis (HSS) that
because the Earth appears green, herbivores must have little impact
on plant abundance
1. Herbivores must be limited by predators rather than food
supply
2. Plants are so common they endure severe competition
3. Natural enemies, by contrast are limited only by the
availability of prey
iii.
Ecosystem exploitation hypothesis (EEH; Oksanen et al., 1981)
1. Strength of various types of mortalities on different trophic
levels varied with the type of system involved (Figure
13.12)
2. As plant productivity increases, more herbivores are
supported
3. In the absence of carnivores, herbivores will be limited by
plant resources
4. Primary productivity will reach a point where there are
sufficient herbivores to support carnivores. This is the HSS
scenario
5. As productivity is increased further, secondary carnivores
are supported
6. Table 13.5. The importance of competition or predation on
a given trophic level alternates as the number of trophic
levels increases
iv. Little evidence to support HSS and EEH
1. Removal of higher trophic levels leaves lower trophic
levels present, although modified
2. Removal of primary producers leaves no system at all
3. Trophodynamics makes more sense
c. Environmental stress
i. Environmental stress hypothesis originated by Menge and
Sutherland (1987)

1. Postulates that the strength of various mortalities is

governed by environmental stress
2. In stressful habitats, higher trophic levels have little effect
because they are rare or absent, and plants are mainly
affected by environmental stress
3. In habitats of moderate stress, there would be a little
herbivory, but not enough to affect population densities.
Plant densities are higher and are affected by competition
4. In benign environments, there are many herbivores, and
herbivory, not competition or environmental stress, controls
plant abundance
7. Summary
a. Mortalities that perturb populations away from mean levels are termed
"key factors." Key factors can be identified through key factor analysis.
There are many key factors for plants and animals, and no generalizations
can be made as to which key factors are important.
b. Factors that return populations to equilibrium are called density-dependent
factors. Few generalizations can be made as to which factors commonly
act in a density-dependent fashion.
c. Sometimes density dependence does not occur because populations exist
in interdependent groups (metapopulations). In these groups, dispersal is
the key to understanding their dynamics.
d. Several different models have been proposed to describe the types of
mortality factors that should be most important in different systems.
i. Trophicdynamics, a bottom-up approach, suggests that populations
are severely limited by their food supplies. The attenuation of
energy severely limits the number of trophic levels.
ii. Top-down approaches, such as HSS and EEH
e. Environmental stress hypothesis as an alternative hypothesis. Hypothesis
postulates that biotic complexity decreases with increasing stress.
8. Discussion Questions
a. Contrast and compare a key factor with a density-dependent factor. Give
examples.
b. Think of some local plants, fish, birds, insects, or other animals in your
area. Do they tend to have the same densities from year to year? Do you
think you would have noticed if they didnt? Try to think of one example
in which the abundance changed from one year to the next. What was the
cause of this change? What do you think regulates the abundance of this
species?
c. Do you think that density dependence has to occur in most populations
because there are few examples of sigmoidal or geometric growth?
d. What do you think is the appropriate scale on which to look for density
dependence as a regulator of population size: temporal or spatial?
e. Are territorial animals likely to be more affected by different factors than
nonterritorial animals?

Chapter 14: The Main Types of Communities

1. Outline
a. In most instances, communities are less like real entities that act like
superorganisms than they are individualistic associations of species
adapted to the same conditions
b. Climate causes the distribution of the major communities, or biomes, on
Earth
c. Communities can be classified in many different ways, but the most
common is with the use of two descriptors: climate and type of vegetation
d. Some of the main differences between terrestrial communities concern the
diversity of species present in them
e. Aquatic communities are distinguished by the influence of different
physical variables
2. Are Communities More Than the Organisms They Comprise?
a. Communities: an ecological unit, consisting of assemblages of many
populations living in the same place and same time
b. Since many populations are dependent upon one another, studying the
community as a whole may provide more insight than studying the
individual populations
c. Communities have emergent properties, such as the ability to withstand
drought, or community resistance to invasion by exotic species. Emergent
properties may be related to the stability or functioning of the community
d. Early scientists equated the community to a superorganism
i. Frederic Clements (18741945) suggested that ecology was to the
study of communities as physiology was to the study of individual
organisms
ii. Clementss ideas are still shared by some todayGaia Hypothesis
(Lovelock, 1991)
iii.
Henry Allen Gleason (18821975) proposed an "individualistic"
concept of plant association, in place of Clementss hypothesis
1. Suggested that distinct ecological communities did not
exist
2. Instead, individual species of plants were viewed as being
distributed independently along gradients. Thus,
communities could not be assigned boundaries.
iv. Robert Whittakers studies (19531970) on the vegetation
communities along elevation gradients on mountains
1. Asserted the "principle of species individuality," which
stated that most communities intergrade continuously, and
that competition does not create distinct vegetational zones.
This supported Gleasons ideas.

e. Communities can occur on a wide range of scales and can be nested

i. Ex. The tropical forest community encompasses the community
living in the water-filled recesses of bromeliads, which in turn
encompasses the microfaunal communities digesting the cellulose
in insects guts
ii. Depends on frame of reference (Figure 14.1)
f. Richness: Number of species present
g. Diversity: Index of distribution of individuals between species
h. Classification of immense range of variation in communities into a
manageable system is of fundamental importance to the management and
conservation of the biosphere
i. Problem: Assumes that classification of natural environment can be
divided into discrete and discontinuous units, rather than different
parts of a highly variable natural continuum (the latter is probably
a more accurate description
ii. Thus, community structure is only loosely valid
3. Climate and Community Structure
a. Substantial differentials in temperature over the Earth
b. Variation is largely due to variations in incoming solar radiation
i. Angle at which solar radiation hits the Earthless solar energy
striking the Earths surface per m2 the closer you are to the poles
(Figure 14.2)
ii. Length of travel of solar radiation through the atmospherethe
closer you get to the poles, the greater distance through the
atmosphere solar radiation has to travel, resulting in greater
dispersion
iii.
Result: 40% less total annual insolation at the poles compared to
the equator
iv. Earths angle of rotation (23.5%) results in changes in day length
and difference in Northern Hemisphere vs. Southern Hemisphere
(Figure 14.3)
c. Understanding global climate circulation patterns
i. There is not a linear relationship between temperature at the
Earths surface and latitude (Figure 14.4)
ii. Global temperature differentials create winds and drive the
atmosphere
iii.
Classical model of atmospheric circulation (George Hadley, 1735)
1. Solar energy drives winds that influence the circulation of
the atmosphere
2. Temperature contrast between poles and equator create
thermal circulation
3. Warmth at the equator causes air to heat up and rise
4. As the warm air rises, it cools and becomes less buoyant.
However, it cannot sink, because of warm air behind it.
Instead it spreads north and south away from the equator,

iv.

v.

vi.

eventually returning to the surface at the poles (Figure

14.5)
5. When the Earths rotation is factored in, the surface flow is
deflected to the west, in the Northern hemisphere (Coriolis
effect)
New model needed
1. Empirical data showed that the one large cell that was
originally hypothesized was incorrect.
2. In the 1920s a 3-cell circulation in each hemisphere was
proposed (Figure 14.6)
a. Cell nearest the equator is called the Hadley cell
b. Subsidence zones (also called horse latitudes) are
areas of high pressure and are sites of the worlds
tropical deserts, because the subsiding air is
relatively dry, having released all of its moisture
over the equator
c. The trade winds from both hemispheres meet near
the equator in a region that has a weak pressure
gradient
i.
Intertropical convergence zone is called the
Doldrums
ii.
Here the light winds and humid conditions
provide monotonous weather
d. Circulation between 30 and 60 latitude, the net
surface flow is poleward, and because of the
Coriolis force, the winds have a strong westerly
component
i.
Secondary peak in precipitation from about
45 to 60
e. The distribution of major biomes are set by
temperature differentials and the wind patterns that
they generate
Ocean currents
1. Rotation of the Earth and winds create currents
2. Between continents, currents act as pinwheels
a. Running clockwise in the Northern Hemisphere and
counterclockwise in the Southern Hemisphere
b. Figure 14.7
Broad geographical temperature trends are affected by altitude and
landmass
1. Increasing altitude, temperature decreases in a process
termed adiabatic cooling
a. Increasing elevation means a decrease in air
pressure

b. At lower pressure, as air expands, it cools. Cooling

occurs at a rate of 10C for every 1000-m increase
in elevation
2. Landmass affects climate because land heats and cools
more quickly than the sea
a. Land reflects less heat, so it heats up quicker and
loses heat quicker than the sea
b. As warm air rises, cool air replaces: Pattern creates
onshore and offshore breezes
4. Classification of Communities
a. General habitat classifications
i. Based on physical characteristics and appearance of an area
ii. Independent of species living in area
iii.
Classifications cover a wide range of conditions and have little
heuristic use
iv. Ex. Forest, grasslands, wetlands, etc.
b. Life Zone Classification Scheme (Holdrige, 1967) (Figure 14.8)
i. Temperature and rainfall considered the most important
environmental factors
ii. Descriptive names given to communities
iii.
Differences between some communities are difficult to discern
c. Global community classification systems
i. Somewhere between complex definitions of communities and
oversimplified ones
ii. The best descriptions use a combination of a general definition of
the type of habitat and a climatic descriptor
iii.
Seven community descriptions
1. Tropical forest, always deciduous
2. Temperate forest, deciduous
3. Temperate forest, coniferous
4. Tropical grassland, called savanna
5. Temperate grassland, called prairie
6. Desert
7. Tundra
iv. The world can be divided into these types of communities or
biomes (Figure 14.9)
5. Terrestrial Communities
a. Tropical forests
i. Found in the equatorial regions where rainfall exceeds 240 cm a
year, and average temperature is more than 17C
ii. Neither scarcity of water nor low temperatures to limit tree growth
iii.
Soils are fairly poor yet support luxuriant vegetation (Figure 14.10)
1. Many nutrients are leached out by heavy rainfall
2. No rich organic layer, thus fallen leaves are quickly broken
down
3. Most of the nutrients are locked up in the vegetation

4. Areas do not support agricultural practices well for long

iv. Tropical forests occupy 23 % of the worlds total land area
v. Human population in tropical forests is about 20% of the total
world population
vi. High species diversity: sometimes reaching more than 50 tree
species per ha
vii. Rain forest trees are often smooth-barked with large, oval waxy
leaves, narrowing to drip tips at the apex
viii.
Many trees are shallow-rooted with large buttresses for support
ix. Tallest trees reach heights of 60 m
x.
Little light penetrates the canopy, and there is little ground cover
xi. Epiphytes are common
b. Temperate forests
i. Common forest in United States and Europe (Figure 14.11)
ii. Found in regions where temperature falls below freezing each
winter, but not usually below -12C, and average rainfall is
between 75 and 200 cm
iii.
Commonly, leaves are shed in the fall and reappear in the spring
iv. Species diversity is much lower in temperate forests than in
tropical forests
1. Several tree genera may dominant in a given locality
a. Ex. oaks, hickories, maples
v. Many herbaceous plants flower in spring before the trees leaf out
and block the light
vi. Canopy allows some light to penetrate to the ground, allowing
more ground cover than in the tropics
vii. Few epiphytes and lianas
viii.
Soils are richer because the annual leaf fall or detritus is not
quickly decomposed
ix. With proper agricultural practices, agriculture can flourish
x.
Temperate animals are adapted to the vagaries of the climate; for
example many mammals hibernate during the cold months
c. Deserts
i. Biomes that suffer from a water deficit
ii. Generally found around latitudes of 30 N and 30 S, between the
latitudes for tropical and temperate forests
iii.
About 1/3 of the Earths land area is occupied by hot, dry deserts
(Ex. Sahara, Kalahari, Atacama, Sonoran, Gobi, and Simpson;
Figure 14.12)
iv. Deserts are characterized by a paucity of water (< 30 cm per year)
and usually high daytime temperatures
1. Cold deserts are found west of the Rocky Mountains, in
eastern Argentina, and in central Asia
2. Lacking cloud cover, all deserts quickly radiate their heat at
night and become cold

v.

In true deserts, plants cover 10% or less of the soil surface. In

semiarid deserts, 1033%
vi. Only in rare circumstances do deserts consist of lifeless sand dunes
(Ex. Atacama Desert in western Chile, where no rainfall has been
recorded)
vii. Three forms of plant life are adapted to deserts:
1. Annuals
2. Succulents
3. Desert shrubs
viii.
Plant adaptations to low water availability
1. Spines or aromatic odors to ward off water-seeking
herbivores
2. Wider spaces between plants, because their roots are longer
and occupy greater areas to ensure maximum watergathering potential (Figure 14.13)
3. To conserve water, plants produce many small seeds
a. Animals that eat these seeds are common
ix. Because of the large amount of sunlight available, irrigated deserts
can be extremely productive for agriculture
1. However, high levels of water must flow through these
systems. This brings salts to the surface, which inhibit plant
growth
d. Grasslands
i. Occur in the range between deserts and temperate forests
ii. Rainfall ranges from 25 to 75 cm per year, which is too low to
support forests, but higher than what is necessary to support only
desert life-forms (Figure 14.14)
iii.
Grasslands can be divided into subcategories based on average
temperature
1. Prairie: temperate grasslands
2. Savanna: tropical grasslands
iv. Fire and grazing may also be important in preventing the
establishment of trees in grasslands
v. Grasslands show differentiation along moisture gradients
vi. Few original grasslands remain due to human activity. Grassland
generally supports agricultural practices well
e. Taiga
i. North of the temperate-zone forests and grasslands lies the biome
of coniferous forests commonly known as taiga (Figure 14.15).
Occurs only in the Northern Hemisphere
ii. Most of the trees are evergreen or conifers with tough needles.
Needles may persist for 3 to 5 years
iii.
Spruce (Picea), firs (Abies), and pines (Pinus) dominate the forests
iv. Some deciduous species occur along watercourses (e.g., aspens,
willow, and alders)

v.
vi.
vii.
viii.
ix.
x.

All species are tolerant to freezes and can withstand temperatures

of -60C
Conical shape of many conifers reduces the likelihood of their
branches being broken by snow
Understory is thin due to year-round canopy cover
Soils are poor because fallen needles decay slowly due to colder
temperatures
Organic matter can build up in the litter later. The layer of litter
also acidifies the soil
Snakes and amphibians are rare, and insects are strongly periodic.
Mammals are heavily furred

f. Tundra
i. Occupies roughly 17% of the Earths land surface (Figure 14.16)
ii. Tundra only exists in the Northern Hemisphere, north of taiga
iii.
Rainfall is less than 25 cm per year, often locked up as snow. Too
little water available for tree growth
iv. Summer temperatures are only 5C. Midwinter temperatures
average -32C
v. Vegetation grows close to the ground and consists of fragile, slowgrowing lichens, mosses, grasses, sedges, and occasional shrubs
vi. Permafrost occurs in the tundra, and because it is impenetrable,
water drainage is inhibited, and water lies on the surface during the
summer in shallow lakes and ponds
vii. Anaerobic conditions of the waterlogged soil and low temperatures
slow nutrient cycling. Organic matter cannot completely
decompose, accumulates in thick layers known as peat
viii.
Animals are adapted to the cold, through thick insulation. Many
birds migrate
g. Other biomes
i. Not all communities fit neatly into the six major biomes
ii. There are characteristic regions where one type of biome grades
into another
1. Some coniferous forests occur in temperate lowlands where
you would expect deciduous trees would dominate.
2. Ex. Most of New Jerseys coastal plain is sandy nutrientpoor soil that cannot support deciduous forest. Instead it
supports pine barrens
3. Chaparral, another distinct biome, is a Mediterranean scrub
habitat that is adapted to fire
a. Rainfall may be sufficient to support trees, but the
frequency of fires prevents the establishment of
trees
4. Mountain ranges need to be treated differently
a. Temperature decreases with altitude. Precipitation
also changes with altitude

b. Mountains can also cause "rain shadows."

Approaching clouds usually dump all their moisture
on the windward side of the mountain
i.
Ex. A biome may change from temperate
forest through taiga and into tundra on an
elevation gradient in the Rocky Mountains
6. Aquatic Communities
a. Marine habitats
i. Most extensive and uniform environment on Earth
ii. Marine habitats are found over more than 70% of the Earths
surface
iii.
Communities are affected by the depth that they occur
1. Intertidal zone: shallow water where land meets the water
2. Neritic zone: shallow regions over the continental shelves
3. Oceanic or pelagic zone: open ocean, which may reach
great depths
4. Benthic zone: sea floor
5. Photic zone: the stratum of water near the surface to which
light penetrates. Phytoplankton, zooplankton, and most fish
live in this zone
6. Aphotic zone: the dark zone below the photic zone.
Virtually no plant production, and is occupied by
invertebrates and luminescent fish
iv. Intertidal zone
1. Alternatively exposed and submerged by the daily cycle of
tides (Figure 14.17)
2. Resident organisms are subject to a great daily variation in
the variability of seawater and temperature.
a. They are also battered by waves
b. In temperate areas, they may be subject to freezing
in the winter and very hot temperatures in the
summer
c. At low tides, they may be dry and vulnerable to
predation
3. There is a vertical zonation consisting of three broad zones
a. Upper littoral: submerged only during the highest
tides
b. Midlittoral: submerged during the highest regular
tide and exposed during the lowest tide each day
i.
Life in this zone is quite rich (green algae,
sea anemones, snails, hermit crabs, and
small fish living in tide pools
ii.
Competition may be intense
c. Lower littoral: Exposed only during the lowest of
tides, and the diversity and richness of organisms is
great

v.

Coral reefs
1. Exist in warm tropical waters (Figure 14.18)
2. Currents and waves constantly renew nutrient supplies, and
sunlight penetrates to the ocean floor (allowing
photosynthesis)
3. Coral reefs are composed of organisms that secrete hard
external skeletons made of calcium carbonate
a. Skeletons form substrate for other coral and algae to
grow on
b. Reefs are one of the richest biomes on Earth
i.
3040% of all fish species on Earth are
found in the coral reefs
vi. Pelagic zone (open ocean)
1. Nutrient concentration is typically low, although
upwellings may periodically enrich the surface waters
2. Waters are typically cold, and are only warm near the
surface, where phytoplankton typically grow and reproduce
a. Their activity accounts for nearly half of the
photosynthetic activity on Earth
b. Increasing phytoplankton could help slow global
climate change
3. Zooplankton graze on the phytoplankton
4. Zone also contains free-swimming animals, called nekton,
which can swim against the currents to locate food (e.g.,
large squids, fish, sea turtles, and marine mammals).
Nekton organisms feed on plankton and on each other.
b. Freshwater
i. Divided into standing-water lentic habitats and running-water lotic
habitats
ii. Lentic habitats
1. Ecology governed largely by the unusual properties of
water
a. Water is least dense when frozen; ice floats, keeping
lakes from totally freezingadvantageous for fish
b. Water is densest at 4C, and as long as no water in
the lake is colder than 4C, the warmest water will
be at the surface, and temperature declines with
depth
2. Three layers are present (Figure 14.19)
a. Epilimnion: upper layer, which is warmed by the
sun and mixed well by wind
b. Hypolimnion: A cool layer too far below the surface
to be warmed or mixed
c. Thermocline: A transition zone between the two
other layers
3. Zonations in lakes

a. Autotrophic or photic zone: Upper layer to which

light penetrates
i.
The point at which photosynthate production
equals energy used in respiration is the
lower limit of the photic zone, and is known
as the compensation level or compensation
point
b. Profundal or aphotic zone: Zone of darkness, where
heterotrophs live and subsist on the rain of material
from above
4. Degree of productivity in lakes
a. Least productive lakes are termed oligotrophic
i.
Low nutrient content
ii.
Relatively clear and their compensation
point could be below the thermocline
iii.
Often contain fish such as trout
b. Eutrophic lakes
i.
Process of lake aging: eutrophication
1. Oligotrophic lakes eventually begin
to accumulate nutrients
2. Sediments begin to be deposited
3. Algae and rooted vegetation begin to
bloom
4. Organic matter begins to accumulate
on the bottom, and respiration
increases
5. The oxygen level decreases and
turbidity increases
6. Trout are excluded, and bass and
sunfish take their place
ii.
Humans speed up eutrophication
1. Increase nutrients through sewage
and fertilizers
2. Biochemical oxygen demand: a
measure of eutrophication.
Difference between the production of
oxygen by plants and the amount of
oxygen needed for respiration
5. Stratified nature of temperate lakes
a. Stratification in summer does not last all year long
(Figure 14.20)
b. In the fall, as the surface cools, it becomes more
dense and sinks, carrying oxygen to the bottom, and
forcing the bottom level upward
c. Lake is thoroughly mixed, and along with storms,
the thermocline disappears

d. In the spring, when the ice melts and sinks, and the
water temperature rises, another mixing occurs
(spring overturn)
6. Tropical lakes
a. Often isothermal or exhibit a weak temperature
gradient
b. Little mixing occurs
c. Deep lakes are generally unproductive, with
oxygen-poor, fishless lower depths
iii.
Lotic or running water habitats (Figure 14.21)
1. Flora and fauna are completely different from those in
lentic waters
2. Plants and animals are adapted to help them remain in place
despite an often strong current
3. Nutrients fail to accumulate and phytoplankton fail to
bloom due to flowing water
4. Current thoroughly mixes water, providing a well-aerated
regime. Therefore, animals are not well adapted to low
oxygen environments, and are particularly susceptible to
oxygen-reducing pollution (e.g., sewage)
5. In cool waters, trout may be present. In warmer, murkier
waters, catfish and carp may be abundant
7. Applied Ecology
a. The distribution of threatened species among biomes
i. It would be beneficial to know which habitats contained the most
threatened species so that we can focus on preserving these
habitats
ii. Data is only available for birds of the world and mammals of
Australia and the Americas (Figure 14.21 UN.01)
iii.
The habitat with the most threatened species is the tropical forest
iv. Frequency of endemic species in each biome. Of the top 25 "hot
spots" for endemic species, 15 are in tropical forests and 5 are in
Mediterranean scrub habitats
v. Threatened birds tend to occupy the same habitats as threatened
mammals
1. Noticeable differences
a. Far more threatened birds than mammals on oceanic
islands
b. Oceanic islands are also known to have many
species of threatened plants
vi. Tropical forests and oceanic islands appear to be good habitats for
maximizing the protection of endangered vertebrates
8. Summary
a. Community ecologists have long debated whether communities are
superorganisms or merely associations of species. The latter view is more
widely accepted

b. Differences in temperature between the poles and the tropics create three
cells of wind movement. Together with Earths rotation and the location of
landmasses, these cells govern the worlds temperature and rainfall
distribution, which in turn governs the types of communities
c. Global classification schemes for communities are based mainly on the
types of vegetation that prevail in different temperature and rainfall
regimes
d. Large-scale terrestrial communities are referred to as biomes and include
tropical forests, temperate deciduous forests, deserts, temperate
grasslands, tropical grasslands, temperate coniferous forests, and tundra
e. Aquatic communities include marine and freshwater communities
i. Vary in properties such as salinity, strength of current, availability
of light, and oxygen content
ii. Marine communities include the intertidal rocky shore, sandy
shore, coral reef, and open ocean
iii.
Freshwater communities are divided into lentic and lotic habitats
f. Tropical forests and oceanic islands have a high proportion of the worlds
endangered vertebrates, and tropical forest and Mediterranean scrub
communities are particularly rich in endemic species. These areas are of
particular interest to conservation biologists
9. Discussion Questions
a. What is the most meaningful scale on which to examine communities?
b. Are the broad divisions of biomes the most useful? What finer divisions
could be made? When would these categories be useful?
c. Which communities or biomes do you expect to be the most degraded by
humans? Explain.

Chapter 14: The Main Types of Communities

1. Outline
a. In most instances, communities are less like real entities that act like
superorganisms than they are individualistic associations of species
adapted to the same conditions
b. Climate causes the distribution of the major communities, or biomes, on
Earth
c. Communities can be classified in many different ways, but the most
common is with the use of two descriptors: climate and type of vegetation
d. Some of the main differences between terrestrial communities concern the
diversity of species present in them
e. Aquatic communities are distinguished by the influence of different
physical variables
2. Are Communities More Than the Organisms They Comprise?

a. Communities: an ecological unit, consisting of assemblages of many

populations living in the same place and same time
b. Since many populations are dependent upon one another, studying the
community as a whole may provide more insight than studying the
individual populations
c. Communities have emergent properties, such as the ability to withstand
drought, or community resistance to invasion by exotic species. Emergent
properties may be related to the stability or functioning of the community
d. Early scientists equated the community to a superorganism
i. Frederic Clements (18741945) suggested that ecology was to the
study of communities as physiology was to the study of individual
organisms
ii. Clementss ideas are still shared by some todayGaia Hypothesis
(Lovelock, 1991)
iii.
Henry Allen Gleason (18821975) proposed an "individualistic"
concept of plant association, in place of Clementss hypothesis
1. Suggested that distinct ecological communities did not
exist
2. Instead, individual species of plants were viewed as being
distributed independently along gradients. Thus,
communities could not be assigned boundaries.
iv. Robert Whittakers studies (19531970) on the vegetation
communities along elevation gradients on mountains
1. Asserted the "principle of species individuality," which
stated that most communities intergrade continuously, and
that competition does not create distinct vegetational zones.
This supported Gleasons ideas.
e. Communities can occur on a wide range of scales and can be nested
i. Ex. The tropical forest community encompasses the community
living in the water-filled recesses of bromeliads, which in turn
encompasses the microfaunal communities digesting the cellulose
in insects guts
ii. Depends on frame of reference (Figure 14.1)
f. Richness: Number of species present
g. Diversity: Index of distribution of individuals between species
h. Classification of immense range of variation in communities into a
manageable system is of fundamental importance to the management and
conservation of the biosphere
i. Problem: Assumes that classification of natural environment can be
divided into discrete and discontinuous units, rather than different
parts of a highly variable natural continuum (the latter is probably
a more accurate description
ii. Thus, community structure is only loosely valid
3. Climate and Community Structure
a. Substantial differentials in temperature over the Earth
b. Variation is largely due to variations in incoming solar radiation

i.

Angle at which solar radiation hits the Earthless solar energy

striking the Earths surface per m2 the closer you are to the poles
(Figure 14.2)
ii. Length of travel of solar radiation through the atmospherethe
closer you get to the poles, the greater distance through the
atmosphere solar radiation has to travel, resulting in greater
dispersion
iii.
Result: 40% less total annual insolation at the poles compared to
the equator
iv. Earths angle of rotation (23.5%) results in changes in day length
and difference in Northern Hemisphere vs. Southern Hemisphere
(Figure 14.3)
c. Understanding global climate circulation patterns
i. There is not a linear relationship between temperature at the
Earths surface and latitude (Figure 14.4)
ii. Global temperature differentials create winds and drive the
atmosphere
iii.
Classical model of atmospheric circulation (George Hadley, 1735)
1. Solar energy drives winds that influence the circulation of
the atmosphere
2. Temperature contrast between poles and equator create
thermal circulation
3. Warmth at the equator causes air to heat up and rise
4. As the warm air rises, it cools and becomes less buoyant.
However, it cannot sink, because of warm air behind it.
Instead it spreads north and south away from the equator,
eventually returning to the surface at the poles (Figure
14.5)
5. When the Earths rotation is factored in, the surface flow is
deflected to the west, in the Northern hemisphere (Coriolis
effect)
iv. New model needed
1. Empirical data showed that the one large cell that was
originally hypothesized was incorrect.
2. In the 1920s a 3-cell circulation in each hemisphere was
proposed (Figure 14.6)
a. Cell nearest the equator is called the Hadley cell
b. Subsidence zones (also called horse latitudes) are
areas of high pressure and are sites of the worlds
tropical deserts, because the subsiding air is
relatively dry, having released all of its moisture
over the equator
c. The trade winds from both hemispheres meet near
the equator in a region that has a weak pressure
gradient

i.

Intertropical convergence zone is called the

Doldrums
ii.
Here the light winds and humid conditions
provide monotonous weather
d. Circulation between 30 and 60 latitude, the net
surface flow is poleward, and because of the
Coriolis force, the winds have a strong westerly
component
i.
Secondary peak in precipitation from about
45 to 60
e. The distribution of major biomes are set by
temperature differentials and the wind patterns that
they generate
v. Ocean currents
1. Rotation of the Earth and winds create currents
2. Between continents, currents act as pinwheels
a. Running clockwise in the Northern Hemisphere and
counterclockwise in the Southern Hemisphere
b. Figure 14.7
vi. Broad geographical temperature trends are affected by altitude and
landmass
1. Increasing altitude, temperature decreases in a process
termed adiabatic cooling
a. Increasing elevation means a decrease in air
pressure
b. At lower pressure, as air expands, it cools. Cooling
occurs at a rate of 10C for every 1000-m increase
in elevation
2. Landmass affects climate because land heats and cools
more quickly than the sea
a. Land reflects less heat, so it heats up quicker and
loses heat quicker than the sea
b. As warm air rises, cool air replaces: Pattern creates
onshore and offshore breezes
4. Classification of Communities
a. General habitat classifications
i. Based on physical characteristics and appearance of an area
ii. Independent of species living in area
iii.
Classifications cover a wide range of conditions and have little
heuristic use
iv. Ex. Forest, grasslands, wetlands, etc.
b. Life Zone Classification Scheme (Holdrige, 1967) (Figure 14.8)
i. Temperature and rainfall considered the most important
environmental factors
ii. Descriptive names given to communities
iii.
Differences between some communities are difficult to discern

c. Global community classification systems

i. Somewhere between complex definitions of communities and
oversimplified ones
ii. The best descriptions use a combination of a general definition of
the type of habitat and a climatic descriptor
iii.
Seven community descriptions
1. Tropical forest, always deciduous
2. Temperate forest, deciduous
3. Temperate forest, coniferous
4. Tropical grassland, called savanna
5. Temperate grassland, called prairie
6. Desert
7. Tundra
iv. The world can be divided into these types of communities or
biomes (Figure 14.9)
5. Terrestrial Communities
a. Tropical forests
i. Found in the equatorial regions where rainfall exceeds 240 cm a
year, and average temperature is more than 17C
ii. Neither scarcity of water nor low temperatures to limit tree growth
iii.
Soils are fairly poor yet support luxuriant vegetation (Figure 14.10)
1. Many nutrients are leached out by heavy rainfall
2. No rich organic layer, thus fallen leaves are quickly broken
down
3. Most of the nutrients are locked up in the vegetation
4. Areas do not support agricultural practices well for long
iv. Tropical forests occupy 23 % of the worlds total land area
v. Human population in tropical forests is about 20% of the total
world population
vi. High species diversity: sometimes reaching more than 50 tree
species per ha
vii. Rain forest trees are often smooth-barked with large, oval waxy
leaves, narrowing to drip tips at the apex
viii.
Many trees are shallow-rooted with large buttresses for support
ix. Tallest trees reach heights of 60 m
x.
Little light penetrates the canopy, and there is little ground cover
xi. Epiphytes are common
b. Temperate forests
i. Common forest in United States and Europe (Figure 14.11)
ii. Found in regions where temperature falls below freezing each
winter, but not usually below -12C, and average rainfall is
between 75 and 200 cm
iii.
Commonly, leaves are shed in the fall and reappear in the spring
iv. Species diversity is much lower in temperate forests than in
tropical forests
1. Several tree genera may dominant in a given locality

v.
vi.
vii.
viii.
ix.
x.

a. Ex. oaks, hickories, maples

Many herbaceous plants flower in spring before the trees leaf out
and block the light
Canopy allows some light to penetrate to the ground, allowing
more ground cover than in the tropics
Few epiphytes and lianas
Soils are richer because the annual leaf fall or detritus is not
quickly decomposed
With proper agricultural practices, agriculture can flourish
Temperate animals are adapted to the vagaries of the climate; for
example many mammals hibernate during the cold months

c. Deserts
i. Biomes that suffer from a water deficit
ii. Generally found around latitudes of 30 N and 30 S, between the
latitudes for tropical and temperate forests
iii.
About 1/3 of the Earths land area is occupied by hot, dry deserts
(Ex. Sahara, Kalahari, Atacama, Sonoran, Gobi, and Simpson;
Figure 14.12)
iv. Deserts are characterized by a paucity of water (< 30 cm per year)
and usually high daytime temperatures
1. Cold deserts are found west of the Rocky Mountains, in
eastern Argentina, and in central Asia
2. Lacking cloud cover, all deserts quickly radiate their heat at
night and become cold
v. In true deserts, plants cover 10% or less of the soil surface. In
semiarid deserts, 1033%
vi. Only in rare circumstances do deserts consist of lifeless sand dunes
(Ex. Atacama Desert in western Chile, where no rainfall has been
recorded)
vii. Three forms of plant life are adapted to deserts:
1. Annuals
2. Succulents
3. Desert shrubs
viii.
Plant adaptations to low water availability
1. Spines or aromatic odors to ward off water-seeking
herbivores
2. Wider spaces between plants, because their roots are longer
and occupy greater areas to ensure maximum watergathering potential (Figure 14.13)
3. To conserve water, plants produce many small seeds
a. Animals that eat these seeds are common
ix. Because of the large amount of sunlight available, irrigated deserts
can be extremely productive for agriculture
1. However, high levels of water must flow through these
systems. This brings salts to the surface, which inhibit plant
growth

d. Grasslands
i. Occur in the range between deserts and temperate forests
ii. Rainfall ranges from 25 to 75 cm per year, which is too low to
support forests, but higher than what is necessary to support only
desert life-forms (Figure 14.14)
iii.
Grasslands can be divided into subcategories based on average
temperature
1. Prairie: temperate grasslands
2. Savanna: tropical grasslands
iv. Fire and grazing may also be important in preventing the
establishment of trees in grasslands
v. Grasslands show differentiation along moisture gradients
vi. Few original grasslands remain due to human activity. Grassland
generally supports agricultural practices well
e. Taiga
i. North of the temperate-zone forests and grasslands lies the biome
of coniferous forests commonly known as taiga (Figure 14.15).
Occurs only in the Northern Hemisphere
ii. Most of the trees are evergreen or conifers with tough needles.
Needles may persist for 3 to 5 years
iii.
Spruce (Picea), firs (Abies), and pines (Pinus) dominate the forests
iv. Some deciduous species occur along watercourses (e.g., aspens,
willow, and alders)
v. All species are tolerant to freezes and can withstand temperatures
of -60C
vi. Conical shape of many conifers reduces the likelihood of their
branches being broken by snow
vii. Understory is thin due to year-round canopy cover
viii.
Soils are poor because fallen needles decay slowly due to colder
temperatures
ix. Organic matter can build up in the litter later. The layer of litter
also acidifies the soil
x.
Snakes and amphibians are rare, and insects are strongly periodic.
Mammals are heavily furred
f. Tundra
i. Occupies roughly 17% of the Earths land surface (Figure 14.16)
ii. Tundra only exists in the Northern Hemisphere, north of taiga
iii.
Rainfall is less than 25 cm per year, often locked up as snow. Too
little water available for tree growth
iv. Summer temperatures are only 5C. Midwinter temperatures
average -32C
v. Vegetation grows close to the ground and consists of fragile, slowgrowing lichens, mosses, grasses, sedges, and occasional shrubs
vi. Permafrost occurs in the tundra, and because it is impenetrable,
water drainage is inhibited, and water lies on the surface during the
summer in shallow lakes and ponds

vii.

Anaerobic conditions of the waterlogged soil and low temperatures

slow nutrient cycling. Organic matter cannot completely
decompose, accumulates in thick layers known as peat
viii.
Animals are adapted to the cold, through thick insulation. Many
birds migrate
g. Other biomes
i. Not all communities fit neatly into the six major biomes
ii. There are characteristic regions where one type of biome grades
into another
1. Some coniferous forests occur in temperate lowlands where
you would expect deciduous trees would dominate.
2. Ex. Most of New Jerseys coastal plain is sandy nutrientpoor soil that cannot support deciduous forest. Instead it
supports pine barrens
3. Chaparral, another distinct biome, is a Mediterranean scrub
habitat that is adapted to fire
a. Rainfall may be sufficient to support trees, but the
frequency of fires prevents the establishment of
trees
4. Mountain ranges need to be treated differently
a. Temperature decreases with altitude. Precipitation
also changes with altitude
b. Mountains can also cause "rain shadows."
Approaching clouds usually dump all their moisture
on the windward side of the mountain
i.
Ex. A biome may change from temperate
forest through taiga and into tundra on an
elevation gradient in the Rocky Mountains
6. Aquatic Communities
a. Marine habitats
i. Most extensive and uniform environment on Earth
ii. Marine habitats are found over more than 70% of the Earths
surface
iii.
Communities are affected by the depth that they occur
1. Intertidal zone: shallow water where land meets the water
2. Neritic zone: shallow regions over the continental shelves
3. Oceanic or pelagic zone: open ocean, which may reach
great depths
4. Benthic zone: sea floor
5. Photic zone: the stratum of water near the surface to which
light penetrates. Phytoplankton, zooplankton, and most fish
live in this zone
6. Aphotic zone: the dark zone below the photic zone.
Virtually no plant production, and is occupied by
invertebrates and luminescent fish
iv. Intertidal zone

v.

vi.

1. Alternatively exposed and submerged by the daily cycle of

tides (Figure 14.17)
2. Resident organisms are subject to a great daily variation in
the variability of seawater and temperature.
a. They are also battered by waves
b. In temperate areas, they may be subject to freezing
in the winter and very hot temperatures in the
summer
c. At low tides, they may be dry and vulnerable to
predation
3. There is a vertical zonation consisting of three broad zones
a. Upper littoral: submerged only during the highest
tides
b. Midlittoral: submerged during the highest regular
tide and exposed during the lowest tide each day
i.
Life in this zone is quite rich (green algae,
sea anemones, snails, hermit crabs, and
small fish living in tide pools
ii.
Competition may be intense
c. Lower littoral: Exposed only during the lowest of
tides, and the diversity and richness of organisms is
great
Coral reefs
1. Exist in warm tropical waters (Figure 14.18)
2. Currents and waves constantly renew nutrient supplies, and
sunlight penetrates to the ocean floor (allowing
photosynthesis)
3. Coral reefs are composed of organisms that secrete hard
external skeletons made of calcium carbonate
a. Skeletons form substrate for other coral and algae to
grow on
b. Reefs are one of the richest biomes on Earth
i.
3040% of all fish species on Earth are
found in the coral reefs
Pelagic zone (open ocean)
1. Nutrient concentration is typically low, although
upwellings may periodically enrich the surface waters
2. Waters are typically cold, and are only warm near the
surface, where phytoplankton typically grow and reproduce
a. Their activity accounts for nearly half of the
photosynthetic activity on Earth
b. Increasing phytoplankton could help slow global
climate change
3. Zooplankton graze on the phytoplankton
4. Zone also contains free-swimming animals, called nekton,
which can swim against the currents to locate food (e.g.,

large squids, fish, sea turtles, and marine mammals).

Nekton organisms feed on plankton and on each other.
b. Freshwater
i. Divided into standing-water lentic habitats and running-water lotic
habitats
ii. Lentic habitats
1. Ecology governed largely by the unusual properties of
water
a. Water is least dense when frozen; ice floats, keeping
lakes from totally freezingadvantageous for fish
b. Water is densest at 4C, and as long as no water in
the lake is colder than 4C, the warmest water will
be at the surface, and temperature declines with
depth
2. Three layers are present (Figure 14.19)
a. Epilimnion: upper layer, which is warmed by the
sun and mixed well by wind
b. Hypolimnion: A cool layer too far below the surface
to be warmed or mixed
c. Thermocline: A transition zone between the two
other layers
3. Zonations in lakes
a. Autotrophic or photic zone: Upper layer to which
light penetrates
i.
The point at which photosynthate production
equals energy used in respiration is the
lower limit of the photic zone, and is known
as the compensation level or compensation
point
b. Profundal or aphotic zone: Zone of darkness, where
heterotrophs live and subsist on the rain of material
from above
4. Degree of productivity in lakes
a. Least productive lakes are termed oligotrophic
i.
Low nutrient content
ii.
Relatively clear and their compensation
point could be below the thermocline
iii.
Often contain fish such as trout
b. Eutrophic lakes
i.
Process of lake aging: eutrophication
1. Oligotrophic lakes eventually begin
to accumulate nutrients
2. Sediments begin to be deposited
3. Algae and rooted vegetation begin to
bloom

4. Organic matter begins to accumulate

on the bottom, and respiration
increases
5. The oxygen level decreases and
turbidity increases
6. Trout are excluded, and bass and
sunfish take their place
ii.
Humans speed up eutrophication
1. Increase nutrients through sewage
and fertilizers
2. Biochemical oxygen demand: a
measure of eutrophication.
Difference between the production of
oxygen by plants and the amount of
oxygen needed for respiration
5. Stratified nature of temperate lakes
a. Stratification in summer does not last all year long
(Figure 14.20)
b. In the fall, as the surface cools, it becomes more
dense and sinks, carrying oxygen to the bottom, and
forcing the bottom level upward
c. Lake is thoroughly mixed, and along with storms,
the thermocline disappears
d. In the spring, when the ice melts and sinks, and the
water temperature rises, another mixing occurs
(spring overturn)
6. Tropical lakes
a. Often isothermal or exhibit a weak temperature
gradient
b. Little mixing occurs
c. Deep lakes are generally unproductive, with
oxygen-poor, fishless lower depths
iii.
Lotic or running water habitats (Figure 14.21)
1. Flora and fauna are completely different from those in
lentic waters
2. Plants and animals are adapted to help them remain in place
despite an often strong current
3. Nutrients fail to accumulate and phytoplankton fail to
bloom due to flowing water
4. Current thoroughly mixes water, providing a well-aerated
regime. Therefore, animals are not well adapted to low
oxygen environments, and are particularly susceptible to
oxygen-reducing pollution (e.g., sewage)
5. In cool waters, trout may be present. In warmer, murkier
waters, catfish and carp may be abundant
7. Applied Ecology

a. The distribution of threatened species among biomes

i. It would be beneficial to know which habitats contained the most
threatened species so that we can focus on preserving these
habitats
ii. Data is only available for birds of the world and mammals of
Australia and the Americas (Figure 14.21 UN.01)
iii.
The habitat with the most threatened species is the tropical forest
iv. Frequency of endemic species in each biome. Of the top 25 "hot
spots" for endemic species, 15 are in tropical forests and 5 are in
Mediterranean scrub habitats
v. Threatened birds tend to occupy the same habitats as threatened
mammals
1. Noticeable differences
a. Far more threatened birds than mammals on oceanic
islands
b. Oceanic islands are also known to have many
species of threatened plants
vi. Tropical forests and oceanic islands appear to be good habitats for
maximizing the protection of endangered vertebrates
8. Summary
a. Community ecologists have long debated whether communities are
superorganisms or merely associations of species. The latter view is more
widely accepted
b. Differences in temperature between the poles and the tropics create three
cells of wind movement. Together with Earths rotation and the location of
landmasses, these cells govern the worlds temperature and rainfall
distribution, which in turn governs the types of communities
c. Global classification schemes for communities are based mainly on the
types of vegetation that prevail in different temperature and rainfall
regimes
d. Large-scale terrestrial communities are referred to as biomes and include
tropical forests, temperate deciduous forests, deserts, temperate
grasslands, tropical grasslands, temperate coniferous forests, and tundra
e. Aquatic communities include marine and freshwater communities
i. Vary in properties such as salinity, strength of current, availability
of light, and oxygen content
ii. Marine communities include the intertidal rocky shore, sandy
shore, coral reef, and open ocean
iii.
Freshwater communities are divided into lentic and lotic habitats
f. Tropical forests and oceanic islands have a high proportion of the worlds
endangered vertebrates, and tropical forest and Mediterranean scrub
communities are particularly rich in endemic species. These areas are of
particular interest to conservation biologists
9. Discussion Questions
a. What is the most meaningful scale on which to examine communities?

b. Are the broad divisions of biomes the most useful? What finer divisions
could be made? When would these categories be useful?
c. Which communities or biomes do you expect to be the most degraded by
humans? Explain.

Chapter 15: Global Patterns in Species Richness

1. Outline
a. From the tropics through temperate areas to the poles, there is a gradient in
species richness, with the highest richness in the tropics and the lowest in
the polar areas. Many biotic and abiotic factors explain the changes in
species richness over these large scales
b. Some communities in similar habitats from different parts of the world,
such as plants in deserts, converge in numbers of species. Others, such as
lizards in deserts, do not.
c. It is estimated that there are 12.5 million species on Earth
d. To preserve biodiversity, we can focus on saving countries with the largest
numbers of species or saving areas with the highest numbers of endemics
e. The preservation of biodiversity is important because recent experiments
show that communities perform best when they have a full complement of
species
2. Introduction
a. Canada has a system of 34 parks located in nearly all of the principal
biomes
b. The number of species they contain is commonly called species richness
c. Review of species of parks (Rivard, 1999)
i. Compared species list when the parks were established to a recent
survey
ii. Some species were lost from certain parks
1. Missing species were mainly those hunted by humans
(Figure 15.1)
iii.
Species richness increased in some parks
1. Influx of species also associated with humans (rats,
starlings, sparrows, and pigeons)
2. Other species have been deliberately introduced, including
game species
iv. Rivard and colleagues found that changes in species richness were
related to climate, particularly mean annual potential evaporation
v. Change in species richness in Canadian parks was highest in the
warmer, southern areas because these contain the most humans
d. Understanding species richness would improve management practices
3. Explanation of Species Richness Gradients

a. Background
i. Generally, the number of species in any habitat increases from
polar areas through temperate areas, and reaches a maximum in the
tropics (Figure 15.2a,b)
ii. Species richness is also increased by topographical variation.
Hence, the increase in birds and mammals in the West (Figure
15.2)
1. Mountains provide a wide range of habitats, thus increasing
species richness
2. Peninsular effect causes a decrease in species richness
iii.
Richness of trees in North America is not well linked to latitudinal
gradients (Figure 15.3)
1. Trees do not grow well in deserts, even though there is a
decrease in latitude and an increase in topographical
variation
2. Tree richness is linked to rainfall level
iv. Understanding theories that explain species richness is important in
order to conserve biodiversity on Earth
b. Biotic explanations
i. Spatial heterogeneity theory
1. There are more plant species in the tropics than any other
climatic zone
2. These plant species support higher numbers of herbivorous
animal species, and hence more carnivores
3. Increased plant richness increases richness in herbivores by
a. Increasing the numbers of monophagous herbivores
b. Creating a more diverse architectural complexity
4. Theory does not explain why there are more plant species
to begin with
ii. Competition theory
1. Advanced by Theodosius Dobzhansky (1950) (Figure on
top of page 253)
2. In temperate climates, natural selection operates through
harsh environmental extremes and species are r-selected
3. In the more constant tropical temperatures, species are
thought to be more K-selected, to compete more keenly and
interact more
a. Increased competition narrows the breadth of niches
available, allowing more species to pack along the
resource axes
4. Little data gathered to test this theory
iii.
Predation theory
1. Proposed by Robert Paine (1966) (Figure at top of page
254)
2. Contrary to the competition hypothesis

a. Argues that predators and parasites in the tropics

hold prey populations down to low levels such that
more resources remain and competition is reduced
b. Allows for more species to coexist
c. Increased richness promotes more predator species
3. Evidence from intertidal communities (Figure at bottom of
page 253)
a. Pisaster (starfish), top predator
b. Food web was fairly constant
c. Pisaster preyed on another predator Thais (whelk),
and on chitons, limpets, acorn barnacles, and
bivalves
d. If Pisaster was removed, diversity decreased from
15 to 8 species
i.
Mytilus increased in numbers, crowding out
other species
ii.
Pisaster prevented any species from
monopolizing the space
iii.
Pisaster was termed a keystone species
4. Darwin (1859) also noted where predation allowed the
coexistence of many prey, when he observed more grass
species in areas that were grazed
5. In order for this theory to explain tropical richness,
predation would have to be intense on the majority of
species at all trophic levels
a. No data to test such a theory
iv. Animal pollinators theory (Figure at bottom of page 254)
1. Most tropical plants are pollinated by animals
2. Close associations develop between plants and specific
pollinators, increasing the reproductive isolation between
plant populationsincreasing the rate of speciation
3. Coevolution of plants and pollinators ensures high rate of
speciation
4. Theory may apply to terrestrial systems, but cannot easily
explain the similar diversity gradients in aquatic
ecosystems
c. Abiotic theories
i. Time theory (Figure at bottom of page 255)
1. Species richness increases over time. Ancient, unglaciated
communities in the tropics contain more species. More
temperate areas were glaciated and species richness has not
yet returned to pre-glacial levels
2. Sanders (1968) provided evidence in support of the
evolutionary time theory

ii.

iii.

a. Compared bottom-dwelling invertebrate diversity in

glaciated and unglaciated Northern Hemisphere
lakes that occur at similar latitudes
b. Lake Baikal (an ancient unglaciated lake) in the
former Soviet Union had 580 species of benthic
invertebrates
c. Great Slave Lake (a glaciated lake) in north Canada
had only 4 species in the same zone
3. Southwood (1961)
a. Investigated insect herbivore diversity on British
trees
b. Found good correlation of insect richness with the
length of time the species of tree inhabited Britain
c. However, American ecologist D. Strong (1974)
found insect species diversity on each tree species
was better correlated with the area over which a tree
species could be found
Area theory
1. Based on the notion that larger areas contain more species
because larger areas can support larger populations
2. Bigger populations result in fewer extinctions
3. Large areas with climatic similarity will have greater
species richness
4. The tropical regions of both hemispheres are adjacent,
creating one large area with a similar climate. The result is
increased species richness
5. Area theory does not seem to account for the relatively few
species in the vast contiguous landmass in central Asia nor
in the deep sea (in comparison to shallow tropical surface
waters).
Productivity theory
1. Proposes that greater primary production by plants results
in greater overall richness (Wright 1983) (Figure at the
bottom of page 256)
2. Currie and Paquin (1987) showed that the diversity of tree
species in North America is best predicted by the
evapotranspiration rate (Figure 15.3). Annual
evapotranspiration is correlated with primary production
3. Gratehouse and Carey (1987) demonstrated a correlation
between richness of British butterflies, and sunshine and
temperature during the months they were on the wing
(suggesting a relationship between energy and species
richness)
4. Theory fails to explain
a. Why tropical seas have low productivity but high
richness

iv.

v.

b. Why eutrophic lakes and lakes polluted with

fertilizers have high productivity but low richness
c. While in North America, there is evidence for the
evapotranspiration theory, the pattern does not hold
for broad comparisons between continents
i.
The temperate areas of eastern Asia support
729 tree species, while climatically similar
North America supports 253 species, and
Europe supports 124 species
Evolutionary speed theory
1. Klaus Rohde (1995) proposed that effective evolutionary
time promotes high species richness. (Figure in the middle
of page 257)
2. High effective evolutionary time is the result of
evolutionary speed and geological time during which an
ecosystem has existed under more or less the same
conditions
3. Evolutionary speed is promoted by high temperatures,
which foster
a. Shorter generation times
b. Higher mutation rates
c. Increased selection
4. Generation time is accelerated by temperature, but no
evidence for higher mutation rates or increased selection
5. Evolutionary speed theory predicts
a. High richness in the tropics
b. High richness in the deep sea
c. Accounts for high richness in areas next to these
areas
Conclusion
1. Eight theories are neither exhaustive nor mutually
exclusive, and can be combined in many permutations
2. Some biotic explanations are insufficient: explanations that
invoke increased competition, predation, or disease are
secondary explanations (Table 15.1). A primary explanation
is still needed
3. There may be good correlations between abiotic variables
and species richness; however, it is not clear why increased
productivity promotes diversity and not simply higher
population densities of just a few species
4. The most realistic way to examine causes of richness
gradients was proposed by Ricklefs and Schluter (1993)
a. Different processes may act on different scales
i.
Biotic factors: local scale
ii.
Evolutionary processes: provincial or global
scale (Figure 15.4)

4. Community Similarity
a. Communities from different parts of the globe are similar
i. Similarity in vegetation in climatically similar areas around the
globe
1. Ex. Cacti-like plants occur in deserts around the globe
ii. If species converge in morphology, can communities converge in
species richness?
1. Similar species diversity in similar habitats (Table 15.2a)
2. Dissimilar species diversity in similar habitats (Table
15.2b)
b. Eric Pianka (1986): Most comprehensive field studies
i. Examined desert lizard species around the world
ii. 61 species in Australia, 22 in southern Africa, but only 14 species
in North America (Figure 15.5)
iii.
Australia had more lizard species than southern Africa, no matter
what the habitat was
iv. Suggests that there are indeed strong evolutionary constraints in
southern Africa
c. Robert Whittaker (1972)
i. Named the differences in richness
1. Within habitats, alpha () diversity
2. Between habitats, beta () diversity
3. Overall difference in diversity between two geographical
regions, gamma () diversity (which is the product of alpha
and beta diversity)
ii. Comparison of Australia and southern Africa (Figure 15.6)
d. John Lawton and colleagues (1993)
i. Examined convergence in diversity in guilds: the actual way
species utilize a common resource
ii. Ex. Bracken fern, Pteridium aquilinum
1. Widespread (Figure 15.7)
2. Over last 20 years, surveys of insects were conducted
3. Species assemblage varies remarkably, giving no evidence
of taxonomic similarity in the fauna (Figure 15.8)
4. Variation in total number of insects exploiting bracken fern
is partly a function of how common and widespread the
plant is in each geographical region (Figure 15.9)
iii.
Effects of biotic interactions
1. Distribution of species across resources on the plant is
idiosyncratic from locality to locality, with numerous
vacant niches (Figure 15.10)
2. Parts of plants go unutilized in certain areas of the world. It
does not look like there is convergence of feeding types
across regions nor does it look like competition is an
important factor

3. Only pattern: leaf seems to be more exploited than the rest

of the plant
iv. Summary of main rules governing species richness of insects on
bracken fern (Table 15.3). Different factors are important at
different scales
5. Global Species Richness
a. 2 million species have been classified, and best estimates suggest that
about 12.25 million exist, with a maximum estimate of 118 million (Table
15.4)
b. Insects represent 2/3 of the total number of species. Why?
c. In any given taxon, it is not the smallest or the largest organisms that are
the most abundant, but the species that are intermediate between smallest
and medium-sized (Figure 15.11)
d. Plot of number of species on Earth against body size, measured in length
(Figure 15.12). Insects fall right where the number of species is highest
e. In order to get a more accurate measure of the number of species on Earth,
we need to obtain a better estimate of the number of insect species
f. Based on observations by Terry Erwin (1982), an estimated 30 million
species of arthropods must be in the tropics alone (Figure 15.13)
i. Estimate is based on one canopy of one tree species in Panama:
1. He found 1100 species of beetles
2. 160 species specific to that species of tree
3. Beetles represent 40% of all arthropods, so he estimated
that there must be 400 arthropods specific to that tree
canopy, and a total of 600 insect species for the whole tree
4. 50,000 different species of tropical trees would result in 30
million arthropods on tropical trees all.
5. Add insects in the soil and insects in temperate forests and
grasslands, results in an estimate of 100 million
ii. Problems with estimate
1. Many tropical insects probably utilize more than one tree
species
2. Many insect species that were taken to be newly discovered
had actually been discovered earlier
3. Most probable estimatebetween 5 and 10 million
(Gaston, 1991)
6. Preserving Species Richness
a. Conservation of biodiversity differs from single-species focus because the
aim is to conserve the habitats that contain the most species
b. Which habitats contain the most species?
i. Generally, tropical habitats have the most species
ii. Ex. La Selva Forest Reserve in Costa Rica (13.7 km2) has almost
1500 plant species, which is more than is found in Britain (243,500
km2)
c. Methods for identifying areas for conservation

i.

Target countries with the greatest number of species

(megadiversity)
1. Mexico, Columbia, Ecuador, Peru, Brazil, Zaire,
Madagascar, China, India, Malaysia, Indonesia, and
Australia
2. Together, these countries hold up to 70% of the diversity in
those groups of organisms
3. Megadiversity approach works well because large countries
with large areas garner most of the available international
funds
4. Drawback: Although they contain the most species, they do
not necessarily contain the most unique species. What is
needed is a measure of uniqueness (number of endemics)
ii. Norman Myers and his colleagues (2000) identified 25 "hot spots"
for endemic forest plants (representing 1.4% of the worlds total
land area, that together contain
1. 133,149 endemic plant species (44% of the worlds total)
2. 9645 endemic vertebrate species (35% of the worlds total)
3. Figure 15.14 and Table 15.5
4. Tropical Andes has the most endemics (Figure 15.15)
5. Protecting these hot spots would prevent the extinction of a
larger number of endemics than would protecting areas of
similar size elsewhere
6. Areas that are enriched in endemics of one taxon are often
rich in endemics of another (Table 15.5)
7. Drawback to preserving species richness through endemism
is that most areas rich in species and endemics are in the
tropics. Thus, other major habitats would be ignored
iii.
Best strategy may be to take into account richness, endemism, and
type of habitat
7. Species Richness and Community Function
a. Concerns for loss of biodiversity (Ehrlich and Wilson, 1991)
i. Moral responsibility to protect what are our only known living
companions in the universe
ii. Humanity has already obtained enormous benefits from
biodiversity in the forms of foods, medicines, and industrial
products
iii.
The wide array of essential services provided by natural
communities
b. Natural communities provide essential services
i. Maintenance of the correct gaseous composition of the atmosphere
1. Prevents global warming and loss of soil biodiversity
ii. Maintenance of a reservoir of natural enemies to prevent pest
outbreaks
iii.
Maintenance of a reservoir of pollinators to pollinate crops and
other plants

c. More diverse communities perform better than less diverse ones

i. Shahid Naeem et al. (1994) experiment
1. A series of 14 environmental chambers, each 1 m2
2. Replicated terrestrial communities that differed only in
their biodiversity (Figure 15.16)
3. The communities consisted of 9, 15, and 31 species, spread
across four trophic levels, with species-poor communities
being subsets of the more diverse communities (Figure
15.17)
4. Experiment ran 6 months
5. Results: as richness increased two- to threefold, so did
community productivity (Figure 15.18)
ii. David Tilman and his colleagues (Tilman 1996; Tilman et al.,
1996) hypothesized that diversity increases both resistance and
resilience
1. Studies directly manipulated plant diversity and measured
plant production and nitrogen extraction rates from the soil
2. 140 plots, each 3 m x 3 m, with comparable soils
3. Sown with seeds of 1, 2, 4, 6, 8, 12, or 24 species of prairie
plants, and replicated 20 times
4. Results: More diverse plots used nutrients more efficiently
than less diverse ones and exhibited increased productivity
(Figure 15.19)
5. Same results were found in a sample of 30 unmanipulated
native prairies (Figure 15.20)
6. Reason: In more diverse communities, interspecific
differences in soil use among plant species allow fuller use
of nitrogen, the main limiting nutrient. So as species
richness increased, more nitrogen from the soil was used up
(Figures 15.19b and 15.20b)
iii.
Relationship between productivity and diversity is not linear
1. Strongest at lower levels of diversity and weaker at higher
levels of diversity
2. Productivity would probably increase dramatically as tree
richness increased from 0 to 40 species, but might increase
little after that (diversity matters, but only up to a point)
8. Applied Ecology
a. Loss in species richness weakens community resistance to invasion by
exotic species
i. Elton (1958) suggested that species-rich communities would be
more resistant to invasion from exotic species
1. Richer systems would more fully utilize limiting resources,
and few would be left over for invaders to use
ii. Elton also suggested that increased richness would reduce the
severity of plant disease

1. Transmission rates are proportional to the host species

population; therefore, in rich communities there are fewer
individuals of any one host
iii.
Increased plant richness should lead to increased diversity of insect
herbivores and hence increased predator and parasite richness
b. Field experiments to test Eltons ideas (Knops et al., 1999)
i. Small biodiversity experiment with 140 plots that were 3 m x 3 m
1. Each plot was weeded to control the species that inhabited
the plots
2. Small plots has either 1, 2, 4, 6, 8, 12, or 24 native
grassland species
ii. Large biodiversity experiment had 342 plots that were 13 m x 13 m
1. These plots were not weeded as carefully, but were large
enough to examine the effects of richness on disease and
insect herbivore attack rates
2. Large plots has either 1, 2, 4, 6, 8, or 16 native grassland
species
iii.
Results
1. The frequency of invaders decreased with increased species
richness (Figure 1) and the biomass of the invaders also
declined
2. The level of fungal disease also decreased with richness
(Figure 2)
3. Arthropod species richness increased with plant species
richness (Figure 3)
4. Overall, as species richness increases, fewer species invade
the area, plants are less susceptible to disease, and insects
are more diverse. It appears that Elton was right!
9. Summary
a. Species richness increases from poles to the equator. A variety of biotic
and abiotic factors have been proposed to explain this phenomenon. Most
explanations are incomplete or seem secondary. Over different scales, the
predictive power of the explanations varies considerably. For instance,
predation and competition may be important on the local scale,
evolutionary time may be important on a large scale
b. Convergence in species richness. Species richness between areas has been
examined with reference to the terms , , and diversity. diversity
exists within habitats, and diversity obtains between habitats. The total
diversity is the product of and .
c. Estimating the total richness of species on Earth is fraught with difficulty,
but a figure of about 12.5 million species has been proposed
d. A variety of methods have been proposed for managing richness and
biodiversity on Earth, including consideration of megadiversity and hot
spots

e. Evidence is emerging that species richness can affect ecosystem function:

In general, species-rich communities perform better than species-poor
communities
10. Discussion Questions
a. Summarize the evidence for and against abiotic and biotic factors as
drivers of the increase in species richness from the poles to the tropics
b. Although most groups of organisms become more species-rich toward the
tropics, sometimes the reverse pattern is found, as with aphids and
parasitic insects. Why do you think this might be?
c. To preserve biodiversity on Earth, should we focus primarily on speciesrich areas, on "hot spots" of rare or endemic species, on underrepresented
habitats, or on some combination thereof?
d. List emergent properties that communities might exhibit, such as stability,
resistance to invasion, and other such properties.

Chapter 16: Species Diversity

1. Outline
a. Diversity indices take into account numbers of individuals and numbers of
species, indicating how evenly the individuals are distributed among
species
b. By graphing the abundance of individuals of all species, rank abundance
plots provide us with a more accurate measure of diversity than do simple
diversity indices
c. Similarity indices indicate the proportion of species common to the
communities being compared
d. Cluster analysis permits the comparison of similarity among three or more
communities
2. Introduction
a. 1710, Dutch colonists on the Indonesian island of Bangku
i. Over the centuries, mining and deforestation on a massive scale
ii. Slash and burn agriculture
iii.
3% of the primary forest remained intact
b. Minimal restoration efforts and bird diversity
i. First glance: restored areas looked the same as unrestored sites
(Table 16.1)
ii. Further analysis: restored sites had more diversity (Figure 16.1)
3. Diversity Indices
a. Measures of diversity
i. Simplest measure: count number of species in an area (species
richness)

1. Major problem: does not take into account species

abundance
a. Community A has 99 individuals of species 1 and 1
individual of species 2
b. Community B has 50 individuals of species 1 and
50 individual of species 2
c. Both communities have the same species richness
2. Species richness is highly susceptible to sample size
a. The greater the number of individuals in the sample,
the higher the number of species recorded
ii. Therefore, we need diversity indices that incorporate both
abundance and richness
1. Two broad categories: dominance indices and informationstatistic indices
b. Dominance indices
i. These indices are weighted toward the abundance of the
commonest species
ii. Berger and Parker dominance index (1970)
1. DBP = Nmax/N; where
a. Nmax = total number of individuals in the most
common species
b. N = total number of individuals in the community
c. Usually expressed as the reciprocal = 1/DBP
2. Concept: if one species dominates a community, then the
community is not very diverse (Table 16.2)
a. Community 1: DBP = 1/0.1 = 10
b. Community 2: DBP = 1/0.55 = 1.82
iii.
Simpsons index (1949)
1. Gives the probability that any two individuals drawn at
random from an infinitely large community will belong to
different species
2. Ds = [(ni(n - 1))/(N(N - 1))]; where
a. S = number of species present
b. ni = number of individuals in the ith species
c. N = total number of individuals in the community
3. The higher the value of D, the lower the diversity, therefore
the index is usually expressed as the reciprocal = 1/Ds
4. Disadvantage: it is heavily weighted toward the most
abundant species
a. Common to all dominance indices
b. Rare species fail to change index
c. Regarded as more accurate than Berger-Parker
Index
c. Information-statistic indices
i. Can take into account rare species

ii.

Based on the rationale that diversity in a natural system can be

measured in a way that is similar to the way information contained
in a code or message is measured
1. A technique to measure uncertainty
2. The higher the value, the higher the uncertainty in
predicting a sequence
3. The more diverse a community
d. Shannon index (information-statistic index)
i. Hs = - pi ln pi; where
1. pi = the proportion of individuals found in the ith species
2. Example (Table on page 276)
ii. Even the rare species 5 contributes to the Shannon index, so if an
area has many rare species, their contributions would accumulate
iii.
Shannon index for real communities falls between 1.5 and 3.5
e. Brillouin index (information-statistic index)
i. Index is designed to reflect species abundance
ii. HB = [ln (N!) - ln (ni!)/N]; where
1. N = total number of individuals
2. ni = number of individuals in the ith species
3. N! = factorial of N
4. ni! = factorial of ni
x
iii.
Example (Table top of page 277)
f. Differences between Brillouin index and Shannon index
i. Shannon index does not change with abundance, as long as the
proportional abundance remains constant. Brillouin index does
change
ii. Brillouin index uses factorials, which quickly produces huge
numbers that are unwieldy. Shannon index is often chosen for its
computational simplicity
g. Comparison of Berger-Parker, Simpson, Shannon, and Brillouin
i. Generally values of diversity are correlated
ii. Example (Table 16.3)
1. Indices generally give the same results
2. 3 out of 4 indicate that bird diversity is greater in the
restored site
3. If rare species are valued, then information-statistics are the
most valuable
4. If total abundance of individuals is important, then
Brillouin is the best index
5. Summary of effectiveness (Table 16.4)
h. Evenness
i. Information-statistic indices are affected by both number of species
and their equitability or evenness

ii.

Compare a communitys actual diversity, HS, to the maximum

possible diversity, Hmax
1. Evenness = H/Hmax
a. E is constrained between 0 and 1.0
2. Generally, adding species and increasing evenness both
increase species diversity (Table 16.5)
4. Rank Abundance Diagrams
a. A more complete picture of the distribution of species abundance
b. Plotting proportional abundance (usually on a logarithmic scale) against
rank abundance
i. Figure 16.2
c. Rank abundance diagrams can be drawn for the number of individuals,
biomass, ground area covered (and other variables) plotted against rank
abundance
i. At least 18 different theoretical forms
ii. Three most well-known: geometric series, lognormal, and brokenstick
iii.
Species abundance is most equitable in the broken-stick, less
equitable in the lognormal, and less equitable still in the geometric
series
d. Geometric series
i. First or most dominant species to colonize a new area appropriates
a fraction of the available resources, and by competitive
interaction, preempts that fraction
ii. Second species then preempts a similar fraction, and so on
iii.
Figure 16.3
iv. Model fits
1. Plants from subalpine communities
2. Benthos is a polluted fjord
3. Best fit for communities of relatively few species, where a
single environmental factor predominates
e. Broken-stick
i. Resources (represented by a stick) are divided at one time into
segments over the length of the stick
1. Implying instantaneous colonization by all species
2. The segments are ranked in decreasing order of their
lengths
3. Abundance of species is assumed to be proportional to the
length of the segment
4. Figure 16.4
ii. Model fits
1. A few bird, fish, worm, and predatory gastropod
communities
2. Realistic on relatively few occasions
f. Lognormal
i. Developed by Frank Preston (1948)

ii.

More common than broken stick for most communities that are
rich in species
iii.
Graph of species number versus a logarithmic scale
1. Figure 16.5
iv. Lognormal distribution of a variety of communities
1. Figure 16.6
v. Existence of the truncated lognormal distribution
1. Plots of number of species (y-axis) against log of
individuals per species (x-axis), were truncated to the left
of the mode
2. Figure 16.7
3. Truncation due to species that were present in the habitat
but not in the sample
4. If larger samples were taken, more species would be
obtained
a. Mode would move to the right
5. Therefore, it is critical that communities are adequately
sampled
5. Community Similarity
a. Similarity coefficients: a method for directly comparing diversity of
different sites
i. Usually compare the number of species common to all areas
ii. Comparison will involve a simple presence-absence matrix for two
areas, A and B (Table at bottom of page 284)
1. a = number of species common to both sites
2. b = number of species in site B, but not in A
3. c = number of species in site A, but not in B
4. d = number of species absent in both samples
iii.
Is "d" meaningful?
1. d is a measure of the negative matches, potentially
biologically meaningful
2. In reality, it is almost impossible to know
3. Most similarity coefficients rely only on a, b, and c
b. Jaccard coefficient
i. Jaccard coefficient is calculated from
1. Cj = a/(a + b + c)
ii. Using previously presented data from Indonesia
1. Cj = 6/(6 + 10 + 3) = 0.32
c. Sorensen coefficient
i. Weights matches in species composition
ii. Sorensen coefficient is calculated from
1. Cs = 2a/(2a + b + c)
iii.
Using previously presented data from Indonesia
1. Cs = 12/(12 + 10 + 3) = 0.48
d. The simple matching coefficient Csm
i. Includes the number of species absent

ii.

Simple matching coefficient is calculated from

1. Csm = (a + d)/(a + b + c + d)
iii.
Using previously presented data from Indonesia
1. Csm = (6 + 21)/(6 + 10 + 3 + 21) = 27/40 = 0.675
6. Cluster Analysis
a. Used when research is being conducted on more than one site
b. Starts with a table or matrix giving the similarity between each pair of
sites (by using any similarity coefficient)
c. The two most similar sites are combined to form a single cluster
d. The analysis then proceeds by successfully combining similar sites until
all are combined into a single figure (dendrogram)
e. Single-linkage clustering: method for combining sites into clusters
i. Begin with a matrix of similarity coefficients (Table 16.6)
ii. Find most similar pairs (Conifer sites 2 and 3)
iii.
Second most similar sites (Oak sites 1 and 2)
iv. Third most similar sites (Conifer sites 1 and 2)
1. Therefore, conifer site is linked with cluster of conifer site
2 and 3, forming a community cluster
v. Last joining is between oak 1 and 2 cluster, and conifer 1, 2, and 3
cluster
vi. Figure 16.8
7. Applied Ecology
a. Weighting biodiversity indices: the use of ordinal indices
i. Most indices discussed so far treat species equally.
1. Is this right, or should weight be used?
2. Should more weight be given to a large, rare predator or a
small, rare nematode?
ii. Ordinal indices: indices that attempt to rank order species in
importance
iii.
Cardinal indices: indices that treat all species as equals
iv. Vane-Wright, Humphries, and Williams (1991)
1. Developed methods to weight "taxonomically rare" species
in diversity indices (Figure 1)
2. Species counts are multiplied by a weighting factor (Table
1)
3. Using a cladogram
a. Those species which have the most branches
between the stem and tips are set to unity
b. Sister groups are given a weight or score (W) equal
to the sum of all other branch scores
4. Methods may overweight the value of taxonomically
distinct species (Figure 2)
v. Information index (I)
1. Based on the number of branchings in the tree (Figure 1b)

2. Sum of the I values is then divided by the value for the

individual species, and the contribution is then expressed as
a percentage
8. Summary
a. Biodiversity may be expressed in a number of ways: species richness or by
various indices that take into account richness and abundance
b. Dominance indices include Simpsons index and the Berger-Parker index.
Both are easy to calculate and interpret, but both are biased toward
dominant or common species. Simpsons index may be more realistic
c. Information-statistic indices pay more attention to rare species. Shannon
index and Brillouin index are information-statistic indices. The Brillouin
index is more sensitive to species abundance, but is much more difficult to
calculate
d. Each of the various dominance and information-statistic indices may give
values slightly different than those obtained from other indices, but all of
the values are generally well correlated. Each has its own strength.
Choosing the appropriate index for the appropriate situation depends on
the question being asked.
e. Most diversity indices can be referred to as cardinal indices (they treat
species equally). Indices that attempt to weight rare species or any other
type of species are known as ordinal indices
f. Diversity indices attempt to describe whole communities with one
statistic. A more complete description can be obtained by plotting
proportional abundance of every species against the rank of its abundance.
Examples of abundance diagrams: geometric, lognormal, and broken-stick
g. To compare diversity among areas, compare diversity indices or use
similarity coefficients. Such indices include Jaccard, Sorensen, and Simple
Matching Coefficient indices.
h. Cluster analysis may be used to compare the similarity of two or more
sites
9. Discussion Questions
a. In developing ordinal indices of species diversity, what factors would you
use in weighting your indices: rarity, biomass, importance in nutrient
cycling, or others?
b. Calculate similarity coefficients between the unrestored and restored sites
and between the restored and unmined sites (Table 16.1). What can be
concluded?
c. Table in Discussion Questions
i. Calculate the species diversity and evenness of the two butterfly
communities. Are the results similar to those for birds in mined and
unmined areas in Indonesia discussed earlier? Explain.
Chapter 17: Stability, Equilibrium, and Nonequilibrium
1. Outline

a. Community stability includes resistance to change and resiliencethe

ability to rebound from change
b. More diverse communities are more stable
c. Laboratory experiments with simple communities have usually failed to
show a link between diversity and stability. Field experiments with
complex communities, by contrast, have shown such a link.
d. The intermediate-disturbance hypothesis suggests that the most diverse
communities, such as rain forests and coral reefs, exist at intermediate
levels of disturbance, due to storms or some other kind of turbulence
2. Introduction
a. Jan Bengtsson and his colleagues (1997)
i. Examined the stability of British woodland bird communities from
1971 to 1992 (Figure 17.1)
ii. Studied 18 different plots
iii.
Recorded number of bird species and number of birds
iv. Most species changed little in abundance
v. Concluded: density-dependent population regulation
3. Community Stability
a. Equilibrium: a community where no change in population size and number
of species occurs
i. Long-term data shows constancy over time (Figure 17.2)
b. Community predictability can decrease over time
i. Eastern Wood bird community
1. Adjacent years show more stability than between widely
separated ones
ii. Temporal scale of change should be accounted for
iii.
Environmental variability can cause increased variability over time
c. Importance of community stability
i. Changes in community richness may be indicative of something
wrong
1. Ex. Decline in species richness associated with DDT
contamination
ii. Instability can lead to the extinction of certain species
d. Type of species
i. Number of species
1. Simply counting the number of species may obscure
potential changes; i.e., some species go extinct or emigrate,
and others immigrate
2. May result in a turnover of species
ii. Thus, definition of stability presupposes a stable equilibrium for
each population (not just a constant number of species)
e. Defining stability
i. Resistance: how big a force is needed to change a community
ii. Resilience: the ability of the community to return to equilibrium
after a perturbation

1. Elasticity: how quickly a community can return to

equilibrium
2. Amplitude: how much disturbance the community can
return from
iii.
Figure 17.3
iv. Global stability: the ability to return from high-amplitude
disturbance
v. Local stability: the ability to return from low-amplitude
disturbance
vi. Resistance and resilience may or may not be correlated
1. Lakes are weakly resistant and weakly resilient
2. Deserts are highly resistant and highly resilient
3. Rivers are not particularly resistant, but may be resilient
(fast-flowing water often cleanses the rivers quickly)
vii. Can a community exist in multiple stable states?
1. If, following an oil spill an intertidal community recovers
and contains 95 of the original 100 species in the
community, is it the same community?
2. Most evidence suggests no multiple stable states.
Environment usually changes from one state to another.
4. Is There a Link between Diversity and Stability?
a. Linkage between diversity and stability first recognized by Charles Elton
(1958)
i. Disturbances in large communities would be cushioned by large
numbers of interacting species
b. Evidence for a link between diversity and stability (Elton)
i. Small, faunistically simple island communities are more vulnerable
to invading species than are species-rich continental communities
ii. Outbreaks of pests are often found on cultivated land or land
disturbed by humans. These areas have few naturally occurring
species
iii.
Tropical rain forests do not often have insect outbreaks like those
that are common in temperate forests
c. Evidence against a link between diversity and stability
i. Examples of invaders of continental regions that assume pest
proportions
ii. Agricultural systems may suffer from pest outbreaks not because
of their simple nature, but because their individual components
often have no coevolutionary history. Native monocultures (e.g.,
Spartina and Juncus) seem to be stable
iii.
Goodman (1975) argued that the stability of tropical ecosystems is
a myth
d. Mathematical argument
i. Increasing complexity actually decreases stability in models
ii. Food webs are likely to be stable only if the following inequality is
satisfied:

1. (SC)_ < 1 ; where

a. _ = interaction strength
b. S = number of species
c. C = connectance
2. Any increase in S or C would increase instability
5. Experimental Tests of Diversity-Stability Hypothesis
a. Nelson Hairston and colleagues' experiments with microorganisms (1968)
i. Various combinations of one, two, or three species of bacterial
prey; one, two, or three species of Paramecium predator; and
sometimes one or two species of protozoan that fed on
Paramecium
ii. Cultures were monitored for 20 days
iii.
Results: Two-trophic level systems with two species of
Paramecium, the least abundant species of Paramecium showed a
tendency to become extinct when only one species of prey was
present than with two or three prey species (Figure 17.4a)
1. Supported a link between diversity and stability
iv. Results: Number of bacteria species held constant at three, and the
number of Paramecium species was increased to two or three,
extinction rates increased (Figure 17.4b)
1. Does not support link between diversity and stability
v. Which of the species went extinct was dependent on which species
were together in the culture
1. Species were not interchangeable numerical units
2. Results argue against a simple link between diversity and
stability
3. When a third trophic level is added, in the form of the
predatory protozoan, there was a further decrease in
stability
b. Sharon Lawler and Peter Morin (mid-1990s)
i. Experiments with microorganisms, to examine changes in
population dynamics that occur as the food chain increases in
length
1. Bacteria were the prey at trophic level 1
2. Bacteriovorous protists occupied trophic level 2
3. Protozoans occupied trophic level 3, and could feed on
either trophic level 1 or 2 (Figure 17.5)
ii. Consensus focused on the variability of trophic level 2 as a
measure of stability
iii.
Protists (Colpidium and Tetrahymena) displayed constant
abundance in shorter food chains, where they occupied the top
trophic level
iv. Addition of predators significantly increased the variability of their
prey populations (Figure 17.6)
v. The degree of reduction in stability depended on the identity of
both prey and predator species

vi.

Species could not be regarded as simply interchangeable units,

casting doubts on mathematical models
c. Laboratory cultures involving many species are difficult to conduct and
maintain, because it is virtually impossible to duplicate the full array of
environmental conditions that exist in nature
d. Field tests of the diversity-stability hypothesis
i. Sam McNaughton working with large vertebrates in Africa for 20
years
1. Disturbed savanna communities in the Serengeti by
allowing buffalo to graze in certain areas
2. Experiments were repeated in species-rich and species-poor
communities
3. Results: Plant biomass changes more in the species-poor
community (Figure 17.7). In the species rich plots the noneaten species proliferate, maintaining overall plant biomass
4. McNaughton felt that this demonstrated a clear link
between diversity and stability
ii. McNaughton and Frank's (1991) work in Yellowstone Park
1. Examined the effect of severe drought on the composition
of species in the grassland plant community (Figure 17.8)
2. Calculated diversity with the Shannon index, then
calculated an index of resistance
a. R = 1 - (pq/2); where
i.
_pq/2 = change in abundance of the ith of n
species in community j, between 1988
(drought year) and 1989 (normal year)
b. R was greater in more diverse communities (Figure
17.9)
iii.
David Tilman (1996): biodiversity and stability in Minnesota
grasslands
1. Examined both population and ecosystem traits
2. Used many different grassland plots in a long-term study
3. Demonstrated that, in more diverse plots, biodiversity
stabilizes community and ecosystem processes, but not
population processes
a. Year to year variability in aboveground biomass
was significantly lower in plots with greater plant
species richness (Figure 17.10a)
b. Year to year variability in species abundance was
not stabilized by plant species richness (Figure
17.10b)
4. Differences between species and community biomass was
most likely due to interspecific competition
a. When climatic variables harm some species,
unharmed competitors will increase

b. Such an increase stabilizes the total community

biomass, but individual species biomass is more
variable
iv. Dan Doak suggested that the diversity-stability relationship was
caused by a statistical averaging effect. More species in a
community increases the likelihood of including a very productive
species. This is called the Portfolio Effect
6. Intermediate-Disturbance Hypothesis
a. Joe Connell (1978)
i. Highest local diversities are maintained in communities of
intermediate levels of disturbance (Figure 17.11)
ii. High levels of disturbance, only colonists (r-selected) will survive,
giving rise to low diversity
iii.
Low levels of disturbance, competitively dominant species will
outcompete other species, only a few K-selected species will
persist, giving rise to low diversity
iv. Examples
1. Richest tropical rain forests occur where disturbance by
storms is common
2. Coral reefs maintain their highest diversity in areas
disturbed by hurricanes
3. Richest plant communities in the southeastern United
States occur on Army bombing ranges
v. Many communities therefore exist in nonequilibrium states
b. Wayne Sousa (1979)
i. Intermediate-disturbance hypothesis experiment in a marine
intertidal situation
ii. Small boulders were easily disturbed and only had a mean 1.7
sessile plant and animal species (Figure 17.12)
iii.
Large boulders were rarely moved by waves, and had a mean of
2.5 species
iv. Intermediate-sized boulders had the most species, with a mean of
3.7 species (had a mixture of r- and K-selected species)
c. Hiura (1975) tested intermediate-disturbance hypothesis on a larger scale
i. Beech forest in Japan over 10 latitude
ii. Areas with intermediate disturbance maintained the highest species
diversity (Figure 17.13
iii.
Results are confounded by latitudinal diversity gradients
(temperature increases as one progresses south, which will increase
species diversity)
7. Applied Ecology
a. Can marine communities recover after oil spills?
i. Crude oil is a common contaminant of marine systems
ii. Water-soluble fractions can be lethal to fish and invertebrates and
may disrupt the body insulation of birds
b. Wreck of the Exxon Valdez near Valdez, Alaska (March 24, 1989)

i.
ii.
iii.
iv.
v.
vi.

vii.

Worst oil spill in U.S. waters: ~11 million gallons spilled (Figure
1)
By international standards, not that big
A week after the spill, the slick covered 900 square miles, and
hundreds of miles of shoreline. Officially, 27,000 birds, 872 sea
otters, and an untold number of fish died
Effects spread to the terrestrial environment, with 100 dead eagles
and most pairs failing to produce young that year
Exxon spent $3.2 billion on cleanup
Was cleanup a success?
1. State viewed recovery as successful if the affected
communities returned to their pre-spill condition
a. Probably unattainable because we cannot know
what the community was like before the spill
2. Exxon view was that recovery occurs by the
reestablishment of a healthy biological community in
which the plants and animals characteristic of that
community are present and functioning normally
a. This view pays little attention to densities and age
structure
3. If populations do not recover, it may not be possible to
determine if it is the blame of the oil spill, or other
environmental variables, such as a severe freeze
Examination of long-term consequences
1. Compared nine oiled areas that were set aside and not
cleaned, with (a) areas that were oiled and cleaned (Figure
2), and (b) areas that were not oiled
2. Percent cover of rockweed
a. 50% on unoiled areas
b. Returned to normal values by 1991 on oiled, noncleaned sites
c. Returned to normal values by 1992 on oiled,
cleaned sites
3. Conclusion: cleaning reduced diversity of species found in
soft sediment cores for two years after the spill. Oiled sites
that were not cleaned showed a reduction in abundance but
not diversity
4. Best cleaning: Leave the beach alone!

8. Summary
a. Stability can be thought of in different waysas resistance to change or as
resilience, which refers to the ability of the community to return to
equilibrium after perturbation. Resilience in turn can be divided into two
concepts: elasticity, which measures how quickly a community can return
to a former state, and amplitude, which measures how big a disturbance it
can return from.

b. A community could exist in more than one form or stable state, but the
evidence for this hypothesis is weak.
c. The anecdotal evidence for a link of diversity with stability, first suggested
by Elton (1958), has not held up well under scrutiny.
d. Experimental studies of communities of microorganisms have not shown a
strong link of diversity with stability. Large-scale field experiments with
plants have shown a link between diversity and stability.
e. In 1996, Tilman reconciled the opposing viewpoints in the diversitystability debate by showing how communities of prairie plants exhibit
stability in biomass and ecosystem function, but not in species abundance.
f. The diversity-stability link may be caused by a "portfolio," or statistical
averaging, effect.
g. The older, more conventional view called the equilibrium hypothesis
proposes that most communities are stable. The more modern
nonequilibrium viewpoint argues that disturbances are frequent and
species composition is constantly changing, so that stability is elusive.
h. The intermediate-disturbance theory suggests that the most diverse
communities, such as tropical forests and coral reefs, exist at intermediate
levels of disturbance.
9. Discussion Questions
a. Classify the biomes (temperate and tropical grasslands and forest, deserts,
tundra, rivers, lakes, estuaries, coral reefs, and the open sea) in terms of
their resistance and resilience.
b. Do you think that pesticides might reduce the stability of agricultural
systems? Explain how such an effect would work.
c. If the intermediate-disturbance hypothesis is correct, how does that
influence the conservation movement? Should conservationists promote
disturbance to maximize diversity?
d. What differences might you expect between equilibrium and
nonequilibrium communities in terms of numbers of species, stochastic
effects, and life history strategies of species?
e. How would you set about establishing a link between diversity and
stability in nature?
f. Do you believe in the idea of multiple stable states? Does this idea help or
hinder the restoration of natural habitats by polluters in terms of how far
they are expected to go to restore communities?
Chapter 18: Succession
1. Outline
a. Communities develop over time
b. In facilitation, each succeeding species makes its habitat more favorable
for subsequent species
c. In inhibition, each species inhibits the species that try to succeed it
d. In tolerance, a species neither facilitates nor inhibits its successors
e. Species richness usually increases during succession

f. Biotic processes, such as herbivory, competition, and mycorrhizae, can

deflect the path of succession
2. Introduction
a. Mt. St. Helens erupts; 8:32 a.m. on May 18, 1980
i. Blast zone felled trees over a 600-km2 area
ii. Avalanche of mud destroyed everything in its path
iii.
Succession of area
1. Using new information, found evidence of 400 prehistoric
avalanches
2. Scientist learned how important stochastic events are in
recovery
3. Traditional dogma: area should be slow to recovery
4. Truth: Some species were quick to colonize the area
a. Most seeds were blown in by wind
b. Dead wood was a bonanza for wood-boring insects,
and woodpeckers and finches
c. Most lakes were covered in ice, protecting the
inhabitants
i.
Algae in lakes bloom and more frogs
d. 120 new lakes were created
i.
Salamanders thrived especially in the new
lakes
e. Blast zone converted area into a meadow habitat
i.
Favorable for gophers, which then built new
tunnels
ii.
Salamanders used tunnels to get to new
lakes
b. Succession can be speeded up by chance events or indirect interactions
(Figure 18.1)
3. Development of Communities
a. Succession: gradual changes in a community that are predictable and
orderly
b. Primary succession: when plants invade an area in which no plants have
grown before
c. Secondary succession: a modification of longer-term primary succession.
It does not occur on bare ground, but on partially cleared land
d. Frank Egler, studies of secondary succession at Alton Forest (1950s)
i. Most species already existed in the ground (seed bank or old roots)
ii. Rate of root regeneration or seed germination governed order in
which species appeared
iii.
Eventually, larger, slower growing trees would outcompete smaller
pioneer species
iv. Egler called his theory the initial floristic composition model
e. Frederic Clements, father of successional theory (1916, 1936)
i. Emphasized succession as a deterministic phenomenon, with a
community proceeding to some distinct end point or climax

ii.
iii.
iv.
v.

vi.
vii.
viii.

Each unit in succession was called a "sere" or "seral stage"

Initial seral stage is termed the "pioneer seral stage" (Figure 18.2)
A disturbance could return a later stage to an earlier one
The community progresses toward a climax stage (where the most
K-selected species has occupied the site)
1. Climax community for any given region was dictated by
climate and soil conditions
Succession governed by abiotic disturbances is termed allogenic
Succession governed by biotic disturbances is termed autogenic
Key assumption: each invading species makes the environment a
little different, so that it becomes more suitable for K-selected
species, which invade and outcompete earlier residents. This
process is known as facilitation

4. Facilitation
a. Succession and the Alaskan glaciers
i. Succession following the retreat of the glaciers fit the Clementsian
pattern of facilitation
ii. Over 200 years, glaciers in the Northern Hemisphere have
undergone dramatic retreats (Figure 18.3)
iii.
Ecological sequence of succession in Glacier Bay (Figure 18.4)
1. As glaciers retreat, they leave tills and moraines, which are
deposits of stones and pulverized rock, respectively, and
serve as soil
2. Bare soil has low nitrogen content and organic matter
3. In the early stages, area is first colonized by a black crust of
blue-green algae, lichens, liverworts, horsetail, and the
occasional river beauty.
a. Blue-green algae is a nitrogen fixer, thus soil
nitrogen levels increase.
b. Organic matter is still minimal
4. A few seeds and seedlings of willow, Dryas, alders, and
spruce occur, but they are rare in the community
5. After about 40 years, the nitrogen-fixing Dryas drummondi
comes to dominate the landscape
a. Soil nitrogen has increased, and so has the soil
depth and litter fall
6. After about 60 years, the nitrogen-fixing alder forms dense,
close tickets.
a. Excess nitrogen produced by nitrogen-fixing
bacteria, and not used by the alder, accumulates in
the soil
b. Level of soil nitrogen increases dramatically, as
does litter fall
7. After about 75100 years, spruce trees begin to overtop the
alders, shading them out
a. Litter fall is still high

iv.

b. Large amount of needles turn the soils acidic

c. Shade causes competitive exclusion of many of the
original understory species
d. Western hemlock and mountain hemlock begin to
occur
8. After 200 years, a mixture of spruce and hemlock results
the climax community (Figure 18.5)
a. Spruce-hemlock is the climax community, if the soil
is well-drained
b. If soil is poorly drained, the forest is invaded by
Sphagnum mosses, which accumulate water and
further acidify the soil
i.
Most trees die due to lack of soil oxygen
ii.
Some lodgepole pine are found
iii.
This climax community is called a muskeg
bog
Evidence of facilitation
1. Ecologists suggested that each species facilitated the entry
into the community of the next species
2. Chapin et al. (1994) showed that facilitation in the Glacier
Bay region occurs only during part of the process
a. Dryas and alders dramatically increase soil nitrogen
levels. The increase facilitates the invasion of
spruce
b. On the other hand, alder shades out Dryas
c. Competition is evident after 50 years, when Sitka
spruce begins to shade out alder
d. Facilitation, originally thought to fuel the entire
process of succession, was only important in the
establishment stages. Competition was important in
the later phases of succession
3. Decomposition of plant material (e.g., logs)
a. Edwards and Heath (1963)
b. Oak leaves in nylon bags in soil; studied
decomposition rates
c. Nylon bags had different size meshes, which could
vary the size of decomposers entering the bags
d. The larger the mesh, the more complete the
decomposition
e. In small-meshed bags, microorganisms were unable
to decompose the leaves (Table 18.1)
f. In the soil, earthworms are most important in the
initial decay process
g. Thus, on a small scale, facilitation occurs in the
decomposition of plant material
4. Teresa Turner (1983)

a. Facilitation in the marine intertidal zone off the

Oregon coast
i.
Algae facilitated the succession of surfgrass,
because surfgrass seeds became attached to
the algal species and could then grow
5. Inhibition
a. Many types of succession do not show elements of facilitation
b. Possession of space is all important
i. Principal mechanism affecting succession is the inhibition of
subsequent colonists
c. Facelli and Facelli (1993)
i. Removed litter of Setaria faberii (an early successional species in
old fields)
ii. Removal increased the biomass of a later species, Erigeron annuis
iii.
Conclusion: release of a phytotoxin from decomposing Setaria
litter or physical obstruction by the litter may contribute to the
inhibition of Erigeron (Figure 18.6)
d. Inhibition: dominant method of succession in marine intertidal zones,
where space is limited
i. Early successional species are at a great advantage in maintaining
possession of valuable space
1. Wayne Sousa (1979)
a. Scraped rock faces clean and out put new rocks and
concrete blocks
b. First colonists: green algae Ulva and Enteromorpha
c. Later, replaced by large brown algae and finally by
red algae
d. By removing Ulva from the substrate, brown and
red algae were able to colonize more quickly
(Figure 18.7)
e. How does this occur in nature?
i.
Ulva is subject to herbivory by the crab
Pachygrapus and is also susceptible to
drying out.
ii.
The middle species, brown algae, is
commonly overgrown by epiphytes
iii.
Red algae is not susceptible to either of the
above
6. Tolerance and Other Patterns of Succession
a. Connell and Slatyer (1977); a continuum from inhibition and facilitation
i. Facilitation model: each species makes the environment more
suitable for the next
ii. Inhibition model: early colonists tend to prevent subsequent
colonization by other species
iii.
Tolerance model: any species can start succession, but the eventual
climax community is reached in a somewhat orderly fashion

1. Evidence: Eglers (1954) work on floral succession

a. Many floral communities, most species are present
at the outset. Whichever species germinated first or
grew from roots would start the successional
sequence
b. Works only for secondary succession
iv. Figure 18.8
b. John Lawton (1987): Fourth model of succession, random colonization
i. Succession proceeds by chance alone
ii. Governed by who arrives first, and happens to be present when
favorable conditions prevail
1. Ex. Mt. St. Helens succession
c. Secondary succession in the Piedmont Plateau, North Carolina
i. Different patterns could govern different seral stages
ii. Henry Oosting (1942) and Catherine Keever (1950)
1. Old growth forests were replaced by agriculture
2. Fields were abandoned, and trees returned
3. Continuum of ages in the fields of the region to study
succession
4. In most communities, there is a gradual overlapping of
species over relatively long periods of time. In the
Piedmont little overlapping occurred
5. One year after fields were abandoned, there were 35
species recorded; all annual and perennial species. Two
species dominated: crabgrass and horseweed
a. Early seres, rotting horseweed inhibits the growth of
asterinhibition model
6. In the second year, same species present, but now aster and
ragweed dominant. 26 new species were present
a. Aster stimulated the growth of broom hedge
facilitation model
7. In the third year, species richness declined. A third species,
broom hedge, became dominant, and remained so for
several years. During this time, seeds of pines and some
hardwoods arrived via wind dispersal
a. Arrival of hardwood seeds via wind is by chance
random-colonization model
8. Fifth year, pines became established, and a closed canopy
by year 10
9. Pine seedlings have a difficult time surviving under the
canopy, but hardwoods thrive
10. By 100 years, there are equal numbers of hardwoods as
pines
11. By 200 years, only scattered pines remain
12. Figure 18.9
7. Patterns in Species Richness During Succession

a. Eugene Odum (1969) and Fahriki Bazzaz (1979) summarized general

trends in succession (Table 18.2)
i. Species in early seral stages
1. Often wind dispersed, seeds can live for a long time,
maximizing their chances for successful colonization
2. Acquire nutrients quickly, grow fast, and reach maturity at
low biomass
ii. Plant species richness generally increases as succession proceeds
1. Ex. Old fields in Minnesota (Figure 18.10)
b. Animal and succession
i. Animals are simple followers of succession
ii. Johnston and Odum (1956) examined the richness of bird
communities (Figure 18.11)
1. Grasslands: ~2 bird species
2. Grass/shrub: 7 bird species
3. 35-year-old pine forest: 10 bird species
4. Old pine forest (60100 years): 1820 bird species
5. Climax oak-hickory forests: 19 bird species
c. Climax communities can experience a decrease in richness
i. Val Brown and G. Edwards-Jones: plant species richness increased
only in middle seres
1. Ex. At climax, a birch woodland, species richness drops
dramatically
2. Ex. Insect richness also decreased dramatically (Figure
18.12b)
d. Animals can affect succession
i. Crab herbivory in intertidal boulders
ii. In Africa, elephants are prolific grazers, maintaining open savannas
iii.
In marine systems, fish grazing can deflect succession
8. Biotic Interactions and Succession
a. Walker and Chapin (1987): biotic processes that can affect succession
(Figure 18.13)
i. Mechanisms promoting seed dispersal are more important to
primary succession, whereas buried seeds and surviving vegetative
propagules are more important to secondary succession
ii. Stochastic variation is more important in severe and low-resource
environments
iii.
Facilitation is more important in severe environments, in primary
succession, and in early stages of community development, where
low levels of nutrient, shade, and water may prevail
iv. Competition is probably more widespread than facilitation,
especially in more favorable environments
v. Maximum potential growth is particularly important in favorable
environments, in which resources are sufficient to promote rapid
growth

vi.

Differences in longevity among plant species are more important in

older communities
vii. Mycorrhizae may be especially important in severe environments,
where plants need extra help to survive
viii.
Insect herbivores and pathogens are more important on mature
vegetation
ix. Mammalian herbivores are more important in early and
midsuccessional communities
9. Applied Ecology
a. Restoration ecology
i. How to restore a community, after the habitat has been altered =
Restoration ecology
ii. Restoration ecology is in its infancy
1. Some cleanup methods appear to cause more damage than
good
a. Ex. Oil spill from tanker, Torrey Canyon. Cleanup
methods caused more damage to indigenous biota
than the oil did
iii.
Foundation of an economical and successful restoration program is
a clear understanding of the environment, plants, animals, and
people involved in it.
iv. Steps in a restoration program
1. The knowledge of why a species or community disappeared
in the first place
2. An understanding of the natural history of similar
ecosystems
3. Test plots
4. Soil preparation
5. Revegetation
6. Advanced techniques
7. The reintroduction of animal components
10. Summary
a. Primary succession occurs when species invade an area in which no
organisms are present, such as land unearthed after receding glaciers.
Secondary succession is the change in species composition following a
change in land usage, such as the reversion of old fields to forests after
agriculture has stopped.
b. Early theories of succession viewed the entire process as facilitative, in
which each species makes the environment more suitable for the next.
c. Later work on succession revealed the existence of inhibition, wherein
early colonists actually prevented colonization by other species.
d. In 1977, Connell and Slatyer recognized the existence of a third type of
succession, which they termed tolerance, which was intermediate between
the other two. In this model, any species can start the succession, but the
eventual climax community is reached in a somewhat orderly fashion.

e. In 1987, John Lawton proposed the random-colonization model of

succession, which posits that there is no facilitation or competition and
that succession proceeds by chance alone.
f. In succession, early communities are often species poor and later
communities are species rich. However, this is not always the case.
g. Recently, it has been recognized that the path of succession can be
modified by herbivory, disease, and other factors. Thus, there is no single
universal cause of succession, which is a multifaceted process.
11. Discussion Questions
a. If agriculture on once-virgin tropical forest areas were to stop, how could
we speed up the process of secondary succession so that the forests
returned?
b. Which types of communities (marine versus freshwater versus terrestrial,
forest versus prairie, temperate versus tropical, or primary versus
secondary) do you think would support which model of succession
(facilitation, tolerance, or inhibition)? For example, in marine intertidal
habitats, nitrogen levels of the soil are not important, but competition for
space on rocks is vital, so whoever arrives first might have an advantage
and inhibition might be common.
c. How does the existence of the climax stage in many communities affect
the diversity-stability debate? If diversity is not maximum at the climax,
does it mean that climax communities are less stable than others?
Chapter 19: Island Biogeography
1. Outline
a. Island biogeography theory suggests that the number of species on an
island is affected by the size of the island and by the proximity of the
island to a source pool of colonists. The theory also suggests that species
continually emigrate to islands and go extinct, so that species turnover
occurs.
b. There is much evidence to support the hypothesis that, as the size of an
oceanic island or a continental habitat island increases, so does the species
richness
c. There is also evidence to suggest that more distant islands have fewer
species
d. Although species turnover on islands occurs, it does not appear to be
common
2. Introduction
a. Island of Krakatau
i. Massive volcanic eruption in 1883
ii. Destroyed two-thirds of island. Also, eradicated life on
neighboring islands of Rakata, Sertung, and Panjang.
iii.
1930, a new island was formed from volcanic activity (Anak
Krakatau)
iv. Recolonization studies

1. Nine months after 1883 eruption: first colonist of Rakata

was a spider
2. 1896, 11 species of ferns and 15 species of flowering
plants. 16 species were dispersed by wind and another 8 by
sea
3. Recolonization of Rakata was greatly affected by how well
plants were able to disperse
4. Early plant communities were dominated by grasses
5. 25 years later, plant communities were dominated by
Cyrtandra bushes
6. In the 1920s, the plant communities were dominated by
Neonauclea trees
7. Initially, wind and sea dispersed plants were more easily
dispersed than those that required animals (Figure 19.1)
a. After 40 years, animal dispersed species became as
common as wind and sea dispersed
8. Recolonization of islands was based on the size of the
island and the distance of the island to source of colonists,
and the ability of an organism to disperse
3. Theory of Island Biogeography
a. Equilibrium theory of insular zoogeographyfirst comprehensive theory
of island biogeography: Robert MacArthur and E. O. Wilson (1963, 1967)
i. The number of species on an island, S, tends toward an equilibrium
number
ii. is the result of a balance between the rate of immigration and the
rate of extinction
1. Rate of immigration is highest when there are no species
present on the island
2. Rate of extinction is low at the time of first colonization
3. Eventually, rate of extinction will equal rate of immigration
(Figure 19.2)
4. Both immigration and extinction lines would be curved
(Figure 19.3)
a. Species arrive at an island at different rates
i.
Immigration curve starts out steep, but gets
progressively shallower
b. Extinctions rise at accelerating rates
i.
As more species arrive, competition
increases
ii.
r-selected species arrive first (poor
competitors followed by K-selected species
(better competitors)
iii.
is determined only by the islands area and position, which
influences the rate of immigration and extinction (Figure 19.4)
iv. Equilibrium is dynamic; hence following colonization of an island
1. Number of species remains constant

2. Extinction = immigration
3. Results in a turnover of species
b. Major modifications to MacArthur and Wilsons theory of island
biogeography
i. Target effect (Whitehead and Jones, 1969)
1. The rate of immigration depends on an islands size
ii. The rescue effect (Brown and Kodric-Brown, 1977)
1. The distance from an island to a source pool of potential
colonists affects both rate of extinction and rate of
immigration
iii.
Target and rescue effects complete MacArthur model (Figure 19.5)
iv. Concept of an island
1. Patches of particular habitat on continents are viewed as
islands in a sea of other unsuitable habitat (Figure 19.6)
c. Strength of MacArthur-Wilson model: generated falsifiable predictions
i. Prediction 1: the number of species should increase with increasing
island size
ii. Prediction 2: the number of species should decrease with
increasing distance of the island from the source pool
iii.
Prediction 3: the turnover of species should be considerable
4. Species-Area Effects
a. Oceanic islands
i. Studies of biogeography: Lesser Antilles
1. Islands enjoy a similar climate, surrounded by deep waters,
and no historical connections to the mainland (Figure 19.7)
2. Ricklefs and Lovette (1999) summarized species richness
for birds, bats, reptiles, amphibians, and lepidopterans
a. Over 19 islands, varied in area (131510 km2)
b. Significant relationship between area and richness
(Figure 19.8)
ii. Relationship of species richness to area
1. S = c Az; where
a. S = number of species
b. c = constant measuring the number of species per
unit area
c. A = area
d. z = slope
e. A z value of 0.301, means that, as we multiply the
area by 10, we double the richness of species
2. In logarithmic form: log S = log c + z log A
b. Habitat islands
i. James Browns (1978) studies of mountain ranges of the Great
Basin
1. Mountain ranges are essentially isolated from one another
(Figure 19.9)

2. Significant relationship between species and area for

mammals and birds
ii. Of particular interest is the value of z, where z is the slope of the
line
1. For continental areas, z is often between 0.15 and 0.25,
which is lower than insular situations, where z is often
between 0.20 to 0.40
a. As larger areas are sampled, fewer new species are
added on continents than on islands
b. Continents have more transient species
c. Browns mountaintops
i.
Birds; z = 0.165
ii.
Mammals; z = 0.326
iii.
Birds more transient, easier for them to
disperse between mountaintops
c. Species as islands
i. Species of host plants act as islands in a sea of other vegetation for
the herbivores that eat from the plants
ii. Elaborated by Donald Strong (1974)
1. Found a species relationship between geographical area of
distribution of British tree species and the number of insect
herbivore species (Figure 19.10)
2. Entire island of Great Britain was divided up into 10-km2
grids
3. Area the tree occupied in Britain was determined
4. Number of insect herbivores per species of tree was
determined
iii.
Hart and Horwitz (1991): Reasons for a species-area relationship
1. Extinction rates are greater on small islands
2. The passive effect of increased sampling effort in bigger
areas increases the number of rare species found
3. Speciation may be more likely in bigger areas, an
explanation also given for greater species richness in the
tropics
4. Larger areas contain more "core" areas, which are less
affected by disturbances. Also, perimeter areas contain
more species that are sensitive to these disturbances
5. The species-area relationship may be more likely the result
of an increased diversity of habitats on large islands than
just an increase in area relationships
iv. Larger areas often contain greater diversity of habitats
1. Barry Fox (1983): investigated the relationship between
species, area, and habitat diversity in Australian mammals
a. Classified habitats into seven broad types
b. Larger areas include more types of habitats (Figure
19.11a)

c. Number of mammalian species is well predicted by

area (Figure 19.11b)
d. However, species richness was better predicted
from the number of habitats than from area (Figure
19.11c)
2. Dan Simberloff (1976a,b) investigated the effect of area
alone on the richness of species
a. Chose habitats that do not change as you sample
bigger islands
b. Studied islands of pure mangroves of varying size
in the Florida Keys
c. Collected every species that fed on the islands
d. Reduction in area caused a reduction in the richness
of invertebrate species
e. Area of islands was reduced experimentally
i.
Seven months later, the insects became
reestablished at equilibrium
ii.
Insect densities dropped on all experimental
islands (Figure 19.12)
f. Results clearly indicate that area affects the number
of species
5. The Effect of Distance on Island Immigration
a. Jared Diamond (1972); relationship between distance and number of
species
i. Tabulated land birds on islands close to the source area (New
Guinea), and assumed these islands had 100% of the available
birds
ii. Documented drop-off in species with increasing distance from
New Guinea (Figure 19.15)
iii.
Degree of saturation: richness of bird species as a proportion of the
number found on New Guinea
1. Strong decline with increasing distance
iv. Supports MacArthur-Wilsons predictions
6. Species Turnover
a. Francis Gilbert (1980); investigations of turnover
i. Found 25 investigations to demonstrate turnover; determined that
most of them suffered from fatal flaws
1. Methodology, statistics, or quality of data
ii. Ex. Jared Diamond (1968) studied birds of Californias Channel
Islands National Park
1. Compared his list to that of A. B. Howell (1917)
2. Diamond reported that 510 species per island were no
longer present, but just as many species not listed by
Howell had apparently colonized the islandindicating
turnover

3. Results were challenged: Lynch and Johnson (1974)

pointed out that Howells list was not exhaustive and just a
summary of all known breeding records (some as old as
1860)
iii.
Comparing old list with new lists can be problematic
b. Simberloff and Wilson (1969, 1970); only study of turnover with any
merit
i. Censused small (11 to 25 m in diameter) red mangrove islands in
Florida Keys for all terrestrial arthropods
ii. Fumigated their experimental islands with methyl bromide to kill
all arthropods (Figure 19.16)
iii.
Periodically after fumigation, they censused all islands for several
years
iv. After 250 days, most islands had similar number of arthropod
species that they began with (Figure 19.17)
1. Supporting MacArthur-Wilson theory
v. Colonization and extinction rates were observed
1. Colonization rates during the first 150 days were higher on
nearer islands than far islands
a. Supporting MacArthur-Wilson theory
2. Calculated rates of turnover were very low (1.5 extinctions
per year
a. Data was weak support for MacArthur-Wilson
theory of turnover
3. Same species returned to island
a. Indicates the existence of biological processes that
shape the final community structure the same way
every time the island is recolonized
i.
Contrary to the theory of biogeography
1. Treats the dynamics of different
colonizing species as equivalents
2. Community properties unimportant
4. Conclusion
a. Turnover involves only a subset of transient or
unimportant species, with more important species
becoming permanent after colonization
vi. Take home message: turnover rates are low, which gives little
support to this part of MacArthur-Wilson theory
7. Applied Ecology
a. Theory of National Park Design
i. Shape, design, and management of nature reserves
ii. Centered on island biogeography theory, which suggests that large
parks hold more species than smaller ones
iii.
International Union for Conservation of Nature and Natural
Reserves (IUCN) stated that refuge criteria and management

iv.
v.

vi.

practice should be based on the equilibrium theory of island

biogeography
1. Recommendations are on shaky ground
Large areas cost a lot of money
Which is better, single large areas or several small ones?
1. Single large preserves may buffer populations against
extinction
2. Many studies show that multiple small sites contain more
species (broader range of habitats)
Fauna were shown to be richer in collections of small national
parks than in large parks
Smaller parks are better for maintaining diversity
Implications for future land purchases

vii.
viii.
8. Summary
a. Island biogeography theory predicts that the equilibrium number for
species on an island is determined by a balance between immigration of
species onto that island and extinction of species already there.
b. The theory suggests that the number of species, , is determined by an
island's size and position relative to a source pool of colonists. Extinction
should increase on small islands, because of their smaller populations, and
immigration should decrease on far islands, because colonists have a
difficult time reaching distant places.
c. Island biogeography theory also suggests that there is much turnover on
islands as new species arrive and old ones become extinct. There is little
evidence, however, to support this prediction. Most turnover that has been
documented suggests that rates of turnover are low and center mainly on
transient species.
d. Island biogeography theory may be applied to "habitat islands" as well as
real islands. In the relationship between species richness and area, the
slope of the line may be steeper for true islands than habitat islands and
steeper for poor dispersers like mammals than for good dispersers like
birds.
9. Discussion Questions
a. How can island biogeography be important for conservation biologists?
Should we place reserves close together to facilitate colonization among
them? Should we connect reserves via corridors?
b. How does logging national forests, particularly in the western United
States and Canada, affect species richness in the fragments of forests that
remain? How would you design a logging policy if you were a forester?
c. The MacArthur-Wilson theory suggests a linear relationship between area
and species richness. Can you see any limit to the relationship?
Chapter 20: Trophic Structure
1. Outline
a. Food webs connect all species that feed on one another in an ecosystem

b. Generally, the number of links per species and the connectance remain
constant between different food webs
c. Problems such as imperfectly known links between species, the taxonomy
of species, and the amount of prey eaten by each species can profoundly
affect food web patterns
d. Ecological guilds are groups of species that feed on the same resources in
the same way, and the species involved are likely to be competing for
resources
e. Keystone species have an effect on an ecosystem out of all proportion to
their biomass
2. Introduction
a. A. G. Tansley (1935) coined the term ecosystem, which not only included
the biotic community of organisms in the area, but also the abiotic
environment around the community
b. Ecosystem ecology is concerned with the movement of energy and
material through communities
i. Applied at any scale
c. Most ecosystems do not have clearly defined boundaries (Figure 20.1)
d. Advantage of ecosystem ecology is the common currency of energy or
nutrients, which allows communities and populations to be compared
between and within trophic levels
3. Food Web Complexity
a. Few ecosystems can be characterized by a single unbranched food chain
b. More correct to draw relationships as an elaborate interwoven food web
(Figure 20.2)
c. Measures of food web complexity
i. Chain length = the average number of links between trophic levels
ii. Connectance = actual number of links/potential number of links
iii.
Linkage density = number of links per species
4. Food Web Patterns
a. Generalizations about food webs
i. Connectance remains constant as the number of species in the food
web increases (Figure 20.3)
ii. Mean chain length increases as the number of species in the web
increases (Figure 20.4)
iii.
Top predators tend to be rather large and sparsely distributed,
whereas herbivores are smaller and more common (termed
pyramid of numbers)
1. Charles Elton (1927) described pattern
a. Small pond
b. Numbers of protozoa may run into the millions and
those of Daphnia and Cyclops (their predators)
number in the hundreds of thousands, possibly
10,000 beetle larvae, and 100 fish (Figure 20.5a)
2. Exceptions to the pyramid

a. Oak tree (one producer) supports thousands of

herbivorous beetles, caterpillars, and other primary
consumers, which support tens of thousands of
predators and parasites (Figure 20.5b)
b. Reconcile this exception
i.
Weigh the organisms in each trophic level
ii.
Oak tree weighs 30,000 kg, all herbivores in
tree weigh 5 kg, and the predators sum about
100 g
iii.
Figure 20.5c
c. Inverted pyramids can still occur even when
biomass is used
i.
Biomass of phytoplankton supports a higher
biomass of zooplankton in the English
Channel (Figure 20.5d)
1. May be because the production rate
of phytoplankton is much higher
than zooplankton, and the small
standing crop of phytoplankton
processes large amounts of energy
(Figure 20.5e)
3. The most realistic pyramid is the energy pyramid, which is
never inverted
5. Problems with Food Web Patterns
a. Numerous problems with data that may invalidate the observed patterns in
food webs
i. Predation on "minor" species is frequently omitted. Food webs are
far more complex than is reported
1. Gary Polis (1991)
a. Conducted an intensive 10-year study of the local
desert ecosystem
b. Found 174 species of plants, 138 vertebrate species,
55 species of spiders and scorpions, an estimated
20003000 species of insects, and an unknown
number of microorganisms and nematodes
c. Compared to literature, chains were longer,
omnivory was common, and linkage density was
much greater (Table 20.1)
ii. Data on quantities of food consumed, indicating the thickness of
connecting links, are usually absent from published work
1. Robert Paine (1992)
a. First example in which interaction strengths were
calculated in the food web
i.
Most connections were found to be weak, so
the web was essentially very simple
2. Blake and Wallace (1997)

iii.

iv.
v.
vi.

a. Showed how misleading a normal food web for a

riverine system in Alabama can be, because it
implied the equivalence of all food resources
b. Great variation in the strengths of linkages (Figure
20.6)
In food web theory, species are often aggregated into "trophic
species"
1. Ex. Various types of insects are often lumped together
2. This grouping disguises important biology
Data on the importance of chemical nutrients are sparse
Hard to define web boundaries
Food web links may obscure positive effects of higher trophic
levels on their food production.
1. Ex. Pollinators have a net positive effect on their hosts

6. Guilds
a. Guilds: a group of functionally similar organisms within a tropic level
b. Term was coined by Dick Root (1967)
i. In his studies of birds, he meant to describe a group of species that
fed on the same resources and in the same way
1. Ex. In an insect community feeding on plants, we may have
leaf chewing guild, sap sucking guild, leaf miners, stem
borers, stem gallers, root feeders, and flower feeders
(Figure 20.7)
c. Interest in Guilds
i. Guilds represent arenas of the most intense competition
ii. Guilds represent the basic building block of communities
d. Main problem with guild theory
i. How much overlap in diet does there have to be for species to
belong to the same guild?
ii. Answer may depend on which resources one selects for analysis
1. Ex. The fox is an insectivore if you base diet overlap on
species richness, but not if you base it on biomass of prey
e. Patterns found in guild analysis
i. Density compensation within guilds could maintain overall guild
abundance at or near carrying capacity
1. Fortunes of different species within the guild vary
individually in response to factors other than resource
variability
ii. Evidence: John Lawton and colleagues (1984, 1993)
1. Herbivore community of bracken fern in England, Brazil,
South Africa, New Mexico, Borneo, and Hawaii
2. Insects were arranged into four guilds (chewers, suckers,
miners, and gallers) and further divided based on where
they fed on the plant (main stem, leaves, and leaf veins)
3. Results failed to support the guild theory (Figure 15.10)
iii.
Evidence: Ashbourne and Putnam (1987)

1. Herbivores feeding on red oak

2. Insects arranged into four basic guilds (chewers, suckers,
miners, and gallers), and included three more (species that
folded one leaf in half and fed inside; species that tied two
leaves together and fed inside; and species that rolled
leaves up and fed inside
3. Their result was opposite to Lawton. Striking similarities
were seen between the guild composition of the two
regions (Figure 20.9)
7. Keystone Species
a. Keystone species: a species that has an effect on an ecosystem out of
proportion to its abundance or biomass
i. Ex. The beaver can completely alter an ecosystem by building
dams
b. Difference between a keystone species and a dominant species
i. A dominant species has a large effect in a community because it is
very common
1. Ex. Spartina is a dominant species in a salt marsh due to its
large biomass and role in energy flow
2. Figure 20.10
c. Types of keystone species
i. Keystone enemies
1. Ex. Pisaster starfish and predatory whelks in the rocky
intertidal zone
a. Removal of either results in the community being
dominated by mussels
ii. Keystone prey
1. Ex. Palm nuts, figs, and nectar
a. They are critical to tropical forest fruit-eating guilds
b. Without the fruit trees, wholesale extinction of
frugivores would occur
iii.
Keystone habitat modifiers
1. Ex. Gopher tortoises
a. Burrows provide homes for an array of mice,
possums, frogs, snakes, and insects
b. Without the burrows, many of these creatures would
be unable to survive
c. Figure 20.10
d. Both gopher tortoises and beavers are considered
ecosystem engineers, because they modify the
habitat and cause ecological changes (Table 20.2)
8. Summary
a. Many food webs can be compared by simple properties such as
connectance and linkage density
b. The analysis of food webs has revealed some common patterns. Two are
that, as the number of species in a food web increases, chain length tends

to increase, but connectance remains constant. Another is that there is a

pyramid of numbers, with fewer species on higher trophic levels than on
lower trophic levels.
c. Because food webs are usually imperfectly known, the generalizations that
have resulted from analyzing them may be incorrect. The inadvertent
omission of minor or mobile species, lack of knowledge about the
strength, importance, or beneficial effects of connecting links between
species, and the taxonomic lumping of species into groups or guilds can
all obscure true food web patterns.
d. It is possible to split up trophic levels into units called guilds. Guilds are a
valuable analytical tool because they focus attention on groups of species
most likely to be competing for resources.
e. Keystone species have a large influence on ecosystems out of all
proportion to their abundance.
9. Discussion Questions
a. How might food web properties change if we included all the
decomposers in food webs?
b. Do you think food webs would be different in complexity between twodimensional habitats, such as grasslands, versus three-dimensional
habitats, like forests?
c. Would you expect differences in chain lengths between disturbed and
undisturbed areas? How would disturbance affect connectance or linkage
density?
d. Would fertilizing an ecosystem increase the length of a food chain?
Explain.
e. Do you expect the position of a species in a food web to change as it ages
and increases in size? If so, does this affect how we construct food webs?

Chapter 21: Energy Flow

1. Outline
a. The energy content of plants can be measured by calorimetry, harvesting,
CO2 production, O2 production, or chlorophyll concentration
b. The production of plants is limited by temperature, moisture, and nutrients
such as nitrogen and phosphorus
c. Primary production is highest in the tropics and decreases toward the poles
d. The efficiency of the transfer of energy from plants to animals can be
measured by three indices: production efficiency, consumption efficiency,
and trophic-level transfer efficiency
e. The production of animals is limited mainly by the production of plants
2. Measuring Production
a. Plants make up 99.9% of Earths living mantle
i. Thus, when measuring energy of an ecosystem, we are mainly
interested in plants

ii.

Gross primary production: we measure plant production, because

plants represent the first and primary trophic level. This is
equivalent to energy fixed through photosynthesis
iii.
Net primary production: this is equal to gross primary production
minus the energy plants lose through respiration. It is a measure of
how much biomass is accrued by plants
b. Measurement methods
i. Calorimetry
1. Sample of dry material is burned in a small chamber
2. The greater the energy in a sample, the more heat that it
will generate
ii. Harvesting
1. Samples of plant biomass are obtained at the beginning of
the growing season, after which it is dried and weighed
2. Repeated at the end of the growing season
3. Difference between two measurements if plant production
in the study in a known unit of time
iii.
CO2 uptake method
1. Plants are sealed in chambers, and CO2 uptake due to
photosynthesis is measured
2. The higher the CO2 uptake, the greater is the production of
plant biomass
iv. O2 output
1. Production in many aquatic plants is measured by O2 output
instead of CO2 intake
2. As photosynthesis occurs, oxygen is released. The amount
of photosynthesis that occurs can be calculated from the
amount of oxygen released
v. Chlorophyll concentration
1. We know how much carbon is assimilated per gram of
chlorophyll
2. By measuring chlorophyll concentration, we can estimate
carbon production
c. Scaling up from individual estimates to ecosystems
i. Monumental task
ii. John Teal, examined energy flow in a Georgia salt marsh (Figure
21.1)
1. Most incident solar energy went to two types of plants:
Spartina cordgrass and marine algae
2. Cordgrass and algae are the producers in the system
3. Energetics
a. Most energy is used by the plant in the energetically
costly process of photosynthesis
b. The remainder of the energy is mostly lost in
respiration

c. The energy that is accumulated in biomass is

relatively small (about 6% of the incident solar
energy)
d. Herbivores take very little of this production, eating
only 0.8%
e. The remaining 99.2% of the plant biomass dies in
place and rots on the muddy ground, to be
consumed by bacteria, the major decomposers in the
system
f. Bacteria are eaten by crabs and nematodes. Very
little energy is removed (via tides) from the system
4. Where the energy goes
a. Plants use 98.6% in plant respiration and
photosynthesis
b. Decomposers degrade 20% of the energy that plants
use
c. Animal consumers degrade only 3.8% of the energy
that decomposers use
3. Limits to Primary Production
a. Terrestrial systems
i. Water is a major determinant
1. Production shows an almost linear increase with annual
precipitation, at least in arid regions (Figure 21.2)
ii. Temperature is important
1. Temperature affects production by affecting metabolic rates
iii.
Combined effects of temperature and moisture
1. Actual evapotranspiration rate
2. Could predict the aboveground production with good
accuracy in North America (Figure 21.3)
3. High evapotranspiration is ideal for plant growth, and
primary production is then maximized
iv. Length of growing season in forests
1. The more days available for forests to grow, the more wood
they can put on
v. Nutrient deficiency
1. Particularly N and P
2. Can limit primary production
3. Fertilizers are commonly added to boost production in
annual crops
4. Cargill and Jeffries (1984)
a. Demonstrated that salt marshes and grasses in
subarctic conditions in Hudson Bay were limited by
N and P (Figure 21.4)
b. Once N was added, then P became limiting
c. Addition of N and P together increased production
the most

5. Leibigs (1842) Law of the Minimum

a. The abundance of species is controlled by the most
limiting factor
b. Aquatic systems
i. Light and nutrients
1. Most important limiting factors
2. Light is likely to be in short supply due to the fact that
water readily absorbs light
a. At one meter in depth, half the infrared energy has
been absorbed. At 20 meters, only 5 to 10% of the
radiation is left
b. Too much can also inhibit growth, by overheating
the plants
3. N and P: most important nutrients
a. Can limit production over large areas of the ocean
b. Occur in low concentrations
i.
Few nutrients are tied up in standing crops
1. Soil contains 0.5% N
2. Seawater contains 0.00005% N
c. Enrichment of seawater with N and P can result in
algal blooms
d. P is particularly important in limiting freshwater
lakes
i.
Lakes polluted with runoff enriched with P
form fertilizers or sewage
ii.
These inputs can cause huge algal blooms,
which can clog the lake and increase
turbidity; a process called eutrophication
iii.
David Schindlers (1974, 1977) study of
lake eutrophication
1. Added P to lakes in northwestern
Ontario
2. Phytoplankton biomass increased
3. Composition of algal community
changed from green algae to bluegreen algae (Figure 21.5)
a. Blue-green algae can produce
some secondary chemicals,
so zooplankton prefer green
algae
b. Result is the lakes become
clogged with the scum of
blue-green algae
e. Marine systems
i.
N and P limit production
ii.
Iron can also be limiting

1. Experiment in Sargasso Sea showed

that the addition of iron boosted
production (Figure 21.6)
4. Patterns in Primary Production
a. Efficiency of primary production
i. Efficiency of gross primary production =
1. Energy fixed by gross primary production/energy in
incident sunlight
ii. Phytoplankton communities have very low efficiencies: <0.5%
1. Severely limited by nutrients (Figure 21.7)
iii.
Highest efficiencies occur in temperate forests
iv. Net primary production
1. 50 to 70% of the energy fixed by photosynthesis is lost in
respiration (less than 1% of solar energy is actually
converted into net primary production)
2. In temperate zones, this 1% is equivalent to 300 to 600
calories of primary production per cm2 of land
b. Global distribution of primary production
i. Primary production is highest in the tropical rain forest and
decreases progressively toward the poles (Table 21.1)
1. Gradient is due to temperature decreasing toward poles
ii. Production in the open ocean is very low
iii.
Marine production is highest in upwelling areas of the continental
shelves
5. Secondary Production
a. Biomass of plants
i. Accumulates in a community as a result of photosynthesis
ii. Eventually, goes in one of two directions
1. Detritus feeders
a. Most biomass dies in place and is available to
detritus feeders. This material is known as dead
organic matter
b. Carry out 8090% of the consumption of matter
2. Herbivores
a. Feed on living plant biomass
b. Constitute the greatest selective pressure on plants
b. Measuring energy intake
i. Herbivores are generally confined to an area, and the mass of
vegetation per unit time can be recorded
ii. Carnivores intake of prey items of known biomass and energy
content can be recorded
iii.
Of the energy assimilated, some goes to production (growth) but
much goes to maintenance (respiration)
iv. To measure production

1. Weigh organism before and after a meal, allowing

sufficient time for the meal to have been absorbed by the
body
c. Efficiency of secondary production
i. Three main types of efficiency can be used to compare species at
different trophic levels
1. Production efficiency
a. Defined: how well an organism uses the energy it
assimilates from its food
b. Product efficiency =
i.
[net production/assimilation] x 100
c. Product efficiency in vertebrates is generally less
than invertebrates
i.
Vertebrates use more energy in respiration
d. Endotherms may use over 98% of assimilated
energy for respiration, whereas ectotherms may
only use 90%
i.
Energy cost of homeothermy
e. Consequences of differences
i.
Ex. Sparsely vegetated deserts can support
healthy populations of snakes and lizards,
but mammals might easily starve
1. Komodo dragon (Figure 21.8) eats
its own weight every two months
2. Cheetah eats four times its own
weight every two months
2. Consumption efficiency
a. Defined: measures the amount of one trophic level
eaten by the trophic level above that is incorporated
into the latters biomass
b. Consumption efficiency =
i.
[intake at trophic level n/net production at
trophic level n - 1] x 100
c. Values
i.
015% for terrestrial herbivores
ii.
3040% for zooplankton
iii.
50100% for carnivores
d. Animals are much better equipped to digest meat
than to detoxify and digest chemically protected
plants
3. Trophic-level transfer efficiency
a. Defined: a measure of the energy available at one
trophic level that is acquired by the trophic level
b. Trophic-level transfer efficiency =
i.
[assimilation at trophic level n/assimilation
at tropic level n - 1] x 100

c. Appears to average around 10%, although there is

much variation (Figure 21.9)
d. Trophic level efficiency is no higher than this for
two main reasons
i.
Many organisms cannot digest all parts of
their prey
ii.
Much of the energy assimilated by animals
is used in maintenance, so much of their
energy is lost as heat
e. 10% transfer of energy from one trophic level to the
next necessitates short food chains, most with no
more than four or five links
i.
There simply is little energy to pass on to
higher trophic levels
6. The Limits to Secondary Production
a. Strong relationship between primary production and biomass/consumption
of herbivores (Figure 21.10)
b. Implications
i. Secondary chemicals which make plants poisonous or distasteful to
herbivores are a much less important influence on consumption at
the ecosystem level than they are at the population level
c. Overall, ecosystems are transformers of energy, thus:
i. Plants are the most important organism on the planet. Limits to
plant production control energy flow
ii. The second most important flow of energy is that utilized in the
breakdown of plants by bacteria
iii.
Birds, animals, and insects contribute almost nothing to energy
flow
1. Cannot ignore them, because:
a. They could be keystone species or ecosystem
engineers
b. Herbivores provide selective pressure on plants
c. Herbivores and carnivores play key roles in the
ecosystem
7. Applied Ecology
a. Biomass fuels
i. Energy content of plants can be recovered as heat in the
combustion of biomass materials
ii. Crops can be planted to produce biomass fuels as a source of
renewable energy
iii.
Some areas would produce a far greater yield than others (Figure
1)
iv. Fuel crops would be in competition with food crops in densely
populated countries at the high latitudes
1. Greatest potential for producing biomass is in the humid,
densely vegetated, less densely populated tropical countries

v.
vi.

vii.

Rather than chopping down existing tropical forests, use

harvestable crops such as sugarcane for fuel
Mechanism for generating biomass fuels
1. Anaerobic glycolysis (fermentation) to break down organic
polymers into liquid alcohols (ethanol)
2. 1:3 ratio of ethanol to gas for some cars and trucks, with
only small modifications to the engine
Ethanol fuel
1. Disadvantages: caloric value is only half that of petroleum
and more expensive to produce
2. Advantages: burns cleaner

8. Summary
a. In ecosystems, energy is lost with each transfer up the food chain. In
contrast, chemicals are not dissipated; they remain in the ecosystem and
often concentrate at higher trophic levels.
b. In most ecosystems, plant material goes not to herbivores, but to
decomposers after the plant dies. Most primary production is used in plant
respiration, so plants are the most important consumers. Bacteria are next
in importance as decomposers, degrading about 20% of the energy that
plants use. Herbivores degrade less than 10 percent of the energy the
bacteria use.
c. In general, primary production is highest in tropical forests and decreases
progressively toward the poles. There are exceptions to this trend,
however: Temperate salt marshes are as productive as many tropical areas,
and tropical deserts are not as productive as temperate grasslands. Thus,
temperature and rainfall both limit primary production.
d. Nutrient deficiency, particularly of nitrogen and phosphorus, can limit
primary production. In aquatic systems, the availability of light can also be
important. In freshwater lakes, excess phosphorus can cause huge algal
blooms and turbid water, a process known as eutrophication.
e. The limit to secondary production is available primary production. This
means that, at a large scale, plant defenses do not effectively reduce
consumption by herbivores.
9. Discussion Questions
a. Eutrophication is a phenomenon most often observed in lakes, causing
huge algal blooms. However, eutrophication can occur in marine
environments, too, where it causes "red tide," a dinoflagellate bloom that
is toxic to fish. What do you think might cause "red tide," and how could
you test your ideas?
b. Based on your knowledge of what limits primary production, how could
we improve crop yields? Are there any drawbacks to your suggestions?
c. What limits the efficiency of secondary production? Are native herbivorerangeland systems such as bison-prairie likely to be more efficient than
exotic systems like cattle-grassland?
Chapter 22: Nutrients

1. Outline
a. Soils exhibit a characteristic series of layers or horizons: O, A, B, C, and
E. The availability of nutrients is dependent on soil characteristics
b. In most systems, nutrient availability is limited because the most limiting
nutrients, such as nitrogen and phosphorus, are bound up in living plants,
in the soil, or in lake sediments
c. Light is an important limiting resource for plants, and different
photosynthetic pathwaysthe so-called C3 and C4 pathwayshave
evolved to take advantage of low light levels and high light levels,
respectively
d. Through their feeding activities, herbivores can affect the degree of soil
nutrients. There is a close relationship between nutrient availability and
the abundance of organisms
2. Introduction
a. Nutrients, such as nitrogen, carbon dioxide, and water, play an important
role in the distribution of organisms
b. In addition to the basic building blocks of hydrogen, carbon, and oxygen,
organisms require a wide range of elements (Table 22.1)
c. Macronutrients: elements required in the greatest amount
i. Nitrogen, phosphorus, sulfur, potassium, calcium, magnesium, and
iron
ii. Other nutrients are needed in trace amounts
3. Soils
a. A comparison of elements in the soil with those that accumulate in
vegetation (Table 22.2)
b. Certain elements are likely to be limiting
i. Ex. Nitrogen
c. Soil: complex loose terrestrial surface material in which plants grow. Soil
is composed of
i. Weathered fragments of parent material in organic matter in
various stages of breakdown
ii. Water
iii.
Minerals and organic compounds dissolved in water
iv. Teeming life, mainly microscopic
1. When organisms die, they decay in soil through the
processes of other organisms, especially bacteria and fungi
2. Humus is produced: finely ground organic matter
d. Effect of soil on plant distribution: Serpentine soils
i. Serpentine rock is basically magnesium iron silicate
ii. Plants must be tolerant of low nitrogen, low phosphorus, and high
magnesium
iii.
Conditions are lethal to many plants
iv. Some species have adapted to these soils, forming stunted endemic
communities (Figure 22.1)
e. Soil-forming processes
i. Fall of litter and weathering of rocks tend to act from the top down

ii.

Soils develop a vertical structure referred to as the soil profile

(Figure 22.2)
iii.
Five main layers or horizons
1. O horizon: topmost layer, consisting of organic debris
a. Oi: true litter layer
b. Oa: humus layer, where decomposition of leaves
occurs
2. A horizon: where most plant roots are located, dead organic
matter from the O horizon mixes in with topsoil in this
layer
3. E horizon: zone of eluviation
4. B horizon: organic material is largely absent, mineral
parent material is less thoroughly weathered. Material from
the A horizon may be leached here
5. C horizon: generally consists of weathered parent bedrock
material
6. Not all horizons may exist in a given soil
f. Soil texture
i. Based on the sizes of mineral particles making up the soil
1. Sand (<2 mm)
2. Silt (<0.05 mm)
3. Clay (<0.002 mm)
4. Figure 22.3
ii. Heavy soils
1. Consists mainly of clay
a. Water soaks in slowly and is retained well
b. Potential for being highly fertile
iii.
Light soils
1. Sandy soils
2. Well aerated
3. Free movement of roots and water
iv. Relatively infertile (water drains readily and nutrients are leached)
g. Fertility
i. Based on how minerals are retained
ii. Cation minerals (positive charge when dissolved) are stored on the
surface of soil particles
iii.
Anions are dissolved in soil water
iv. Cations and anions are more prevalent in heavy soils
v. Plants remove cations and replace them with hydrogen
1. Cation exchange capacity:
a. A measure of the number of sites per unit of soil on
which hydrogen can be exchanged for mineral
cations
b. A measure of potential soil fertility

2. Percentage base saturation: percentage of the exchange

capacity filled by calcium, magnesium, and similar
elements
a. 60% means that 60% of the sites are filled by basic
ions and 40% are filled with hydrogen ions
h. Soil development
i. Strongly influenced by
1. Parent material
2. Rainfall
3. Evaporation
4. Temperature
ii. Vegetation is also important
1. Conifers produce strongly acidic soils with a heavy layer of
undecomposed litter and hardpan of leached clay in the B
horizon
2. Deciduous forests produce a thinner, less acidic litter that
grades into the mineral soil below
3. Grasslands are even less acidic, nutrients near the surface,
dark soils due to organic matter added to soil each year
from aboveground biomass as well as at various soil depths
by death each year of many fine roots
a. These soils are termed mollisols
i. Soil types
i. Based on vegetation and other features
ii. 11 different types of soils, known as orders
1. Podzols: acid soils of coniferous forests
a. Clay particles have broken down in A horizon
b. Soluble ions have leached down to lower horizons
c. Lower fertility
d. Soils are known as spodosols
2. Laterization
a. A process brought on by heavy rainfall in the
tropics, where clay is broken down and silica
leaches into lower layers.
b. Only iron oxides and aluminum hydroxide are left
in the A horizon
c. Characteristic red color
d. Poor quality
e. Soils are known as oxisols
iii.
Influence of soil nutrient limitations and influence on animals
(Figure 22.2)
4. Nutrient Availability
a. Biogeochemical cycles: path of nutrients through an ecosystem
i. Nutrients can accumulate in individuals or populations. These can
then act as "pools" of nutrients

ii.

iii.

iv.

v.

Nutrients cycle between pools through meteorological, geological,

or biological transport
1. Meteorological inputs
a. Dissolved matter in rain or snow
b. Atmospheric gases
c. Dust blown in the wind
2. Geological inputs
a. Elements transported by surface and subsurface
drainage
3. Biological inputs
a. Movements of animals or parts of animals from one
ecosystem to another
Types of nutrient cycles
1. Local cycles
a. Involve no long distance transfer of elements (e.g.,
phosphorus)
2. Global cycles
a. Involve an interchange between the atmosphere and
the ecosystem
b. Most elements are involved in global cycles
c. Biosphere: the area of Earth where all living
creatures exist (one large ecosystem), united by
global nutrient cycles
Nutrient turnover time
1. Time taken for a nutrient to pass through a complete cycle
2. Generally 1050 years
3. Inputs and outputs are small in comparison to what is held
in biomass and held within the system, producing slow
turnover times
4. Demonstrated at Hubbard Brook Experimental Forest
(Figure 22.6)
a. Nitrogen was only added to the system in
precipitation (6.5 x 104 kg m-2 yr-1) and nitrogen
fixation (14 x 1034 kg m-2 yr-1)
b. Export to streams: (4 x 10-4 kg m-2 yr-1)
c. Low movement of nitrogen emphasizes how
securely it is held and cycled within the ecosystem
d. Observation confirmed in large-scale experiments at
Hubbard Brook (Figures 22.7 and 22.8)
Nutrient use efficiency
1. Efficiency can vary according to species and location
2. Evergreen species are more efficient in nutrient use than
deciduous species
3. Plants in the Southern Hemisphere are more efficient than
plants in the Northern Hemisphere

4. In contrast to terrestrial systems, freshwater streams and

lakes only have a small fraction of nutrients locked up in
living biomass
b. The nutrient recovery hypothesis
i. Limiting nutrients and population abundance
1. Ex. Lemmings of the tundra in North America and Eurasia
(Figure 22.10)
a. Studied in detail near Point Barrow, Alaska
b. As lemming numbers increase, their feces and urine
stimulate plant growth
c. Increase in plant growth results in an increase in
lemming population
d. Eventually the number of lemmings becomes so
great that their foraging thins out the plant cover
e. More sunlight reaches the ground, thawing out the
soil, so nutrients leach further down the soil and are
not available for plants
f. Intense foraging and loss of nutrients reduces the
quality of forage, resulting in a drastic reduction in
lemmings
2. Nutrient recovery hypothesis (Frank Pitelka, 1964; Arnold
Schultz, 1964)
a. Description of the process in which nutrients
control populations (i.e., the lemmings)
5. Light
a. Light can be a limiting resource for photosynthesis
b. Photosynthetic pathways
i. One reason for the variation in photosynthetic rates among plant
species
ii. 3 different biochemical pathways
1. C3 pathway (Calvin-Benson cycle)
a. CO2 from the air is first converted to 3phosphoglyceric acid (PGA), which is a threecarbon molecule
2. C4 pathway (Hatch-Slack)
a. CO2 is first fixed by producing malic and aspartic
acids (four-carbon molecule)
b. Typically, C4 plants do not reach light saturation
even under the brightest sunlight
c. Always produce more photosynthate per unit area
of leaf than do C3plants
d. Most species are grasses and sedges and more
common in the tropics (Figure 22.11)
e. C4 plants can absorb CO2 more effectively than C3
plants

i.

Do not have to have their stomates open as

much
ii.
More water use efficient
f. The concentration of rubisco is lower in C4 plants
than in C3 plant
i.
Total nitrogen content is lower in C4 plants
ii.
Nutritionally less attractive to some
herbivores than C3 plants
g. Why haven't C4 plants penetrated into the temperate
regions?
i.
C4 plants do not photosynthesize as
efficiently in the shady wet environments of
the northern latitudes
3. Crassulacean acid metabolism (CAM)
a. Open stomates at night to take up CO2opposite of
the typical plant
b. Minimizes water loss during the day
c. CO2 is stored as malic acid and used during
photosynthesis during the day
d. CAM plants have very low rates of photosynthesis
e. Can switch to C3 mode in the daytime, if the
weather is cool
f. Adapted to very dry desert environments
6. Organismal Effects on Nutrient Availability
a. Shade tolerance
i. Sun-loving species soon overtop shade-loving species
ii. However, their shade is too dark for their own seedlings and
inhibits their growth
iii.
Light levels below the crowns of sun-loving species are still
sufficient for those that are intermediate in their shade tolerance,
and below these species are those species most tolerant to shade
iv. Result is a stratified canopy
b. Nitrogen
i. Mark Ritchie and colleagues (1998)
ii. Excluded herbivores from an oak savanna
1. Herbivores preferentially eat nitrogen-fixing legumes and
nitrogen-rich woody plants, leaving prairie grasses that use
nitrogen highly effectively
2. Figure 22.12
iii.
The effect of preference for high-nitrogen plants would be the
removal of nitrogen from the local system
c. Exotic species effect on ecosystem function
i. Northern end of the Great Plains in Canada
ii. Agropyron cristatum, an introduced C3 perennial grass
1. Planted widely since the 1930s
2. Dominates millions of acres of grasslands

3. Outcompeted native prairie grasses over large areas

iii.
Nitrogen levels of normal undisturbed prairie were compared to
disturbed prairie (Figure 22.13)
1. Disturbance alone slightly decreased soil nitrogen
2. Agropyron depleted soil nitrogen greatly
a. Low soil nitrogen in turn reduced plant diversity
7. Applied Ecology
a. Food webs and the passage of pesticides
i. Dichloro-diphenyl-trichloroethane (DDT)
1. First synthesized in 1874
2. Insecticide properties first recognized by Swiss scientist
a. Won Nobel prize in chemistry in 1948
ii. Applications
1. First applications were human health issues before and after
World War II
2. Use in agriculture also began
3. Global production peaked in 1970: 175 million kilograms
were produced
4. Peak production in the United States was 90 million
kilograms in 1964
5. Most industrial countries banned DDT after the early
1970s. A few less developed countries still use it today
iii.
Chemical and physical properties of DDT
1. Persistent in the environment
2. Does not easily degrade to less toxic forms by microbial
actions or physical agents
3. Persistence in soil is typically 10 years
4. Amount of DDT in U.S. soils is negligible
iv. Biological concerns
1. Low water solubility and high solubility in fats or lipids
a. Great affinity for organisms and tends to
concentrate in biological tissue
2. Organisms at the top of a food web are effective in
accumulating DDT from their food
a. Tend to have large concentrations of DDT in their
lipids
b. Figure 1
3. DDT was used to control mosquitoes (and therefore malaria
in less developed countries)
4. Ex. Lake Michigan
a. Largest concentration in gulls, which fed on fish,
which fed on insects
b. Effects on birds
i.
Interference with the metabolic process of
egg formation

ii.
iii.

Thin-shelled eggs that often broke under the

weight of incubating birds (Figure 2)
DDT was responsible for the decrease in the
populations of many bird species

8. Summary
a. The most critical limiting elements are nitrogen and phosphorus, which
exist at relatively low levels in soils. Soil structure can greatly affect the
availability of nutrients to organisms.
b. Nutrients cycle between biological, geological, and meteorological pools.
Most nutrients are bound up in living things or in soils. The nutrient
recovery hypothesis states that populations of some species can recover to
high levels following the release of nutrients into the system.
c. Light can be limiting for plant photosynthesis. The C3 and C4 metabolic
pathways in plants have evolved to take advantage of low-light and highlight environments, respectively.
d. Although the abundance of many organisms is affected by nutrient
availability, organisms themselves can affect nutrient cycles. Herbivores
can increase the availability of nutrients in the soil by feeding or decrease
nutrient availability by eating nitrogen-rich plants.
9. Discussion Questions
a. What sort of organisms live in soil? (You may need to do some research in
the library to find out.) Where are their densities greatestin the O, A, B,
E, or C horizon? What characteristics do these species possess that enable
them to live in the soil? What might govern their densities?
b. It is thought that human alterations have approximately doubled the rate of
nitrogen input into the terrestrial nitrogen cycle. How has this happened?
What effects will it have?
c. Slash-and-burn agriculture is common in tropical countries. Trees are cut
down and burned in place, and crops are planted on the newly cleared
fields. How does this practice affect nutrient cycles in general and soil
nutrient levels in particular?
d. How would you determine whether herbivores increase or decrease soil
nutrients?