BIO 160 LEC 2 Part 2 4.

Some individuals, because of their

physical or behavioral traits, have a
POPULATION GENETICS AND NATURAL higher chance of surviving and
SELECTION reproducing than other individuals in the
same population
Introduction to the theory of Evolution a. If an environment requires a
“Nothing in Biology makes sense except in the certain trait from an organism,
light of evolution” - Theodosius Dobhansky then the organism that possesses
(1973) this trait will survive and
reproduce more of the same
Charles Darwin (1809-1888) species. Thus, their genes
● Galapagos Islands, 1835, aboard become more predominant in
H.M.S. Beagle nature.
● He wondered at the sources of the
differences among clearly related
populations and attempted to explain
the origin of these sources
○ Different species of plants and
animals.
○ He wanted to know why these
slightly related animals were
different even though theory lived
in the same environment.

Darwin’s theory of natural selection

1. Organisms beget like organisms
a. Ex. Humans can only give birth to
humans.
2. There are chance variations between
individuals in a species. Some variations
are heritable
a. Chance variations came from the
parents passed down to them.
b. Traits can be transmitted to the
next generation.
3. More offspring are produced each
generation than can be supported by ● The environment favors the beetles that
the environment are orange because the evolutionary
a. He obtained this idea from forces (e.g birds) are selecting against
Thomas Malthus: Earth can the green-colored beetles.
support only a certain population Orange-colored beetles will become
level, thus organisms have to more predominant
compete with each other →
Survival of the fittest
Evolution & Adaptation
● Darwin (1859) proposed that differential
survival and reproduction of individuals
would produce changes in species
populations over time → Evolution
○ The changes are in terms of
genes or allele frequencies in a
population
● The environment acting on variation
among individuals in populations would
result in adaptation
● Adaptation: an evolutionary process
that changes anatomy, physiology, or
behavior, resulting in an increased
ability of a population to live in a
particular environment
○ Species would have a better
chance of surviving in the
environment.
Gregor Mendel (1822-1884)
● Darwin work nearly half a century to
uncover the loss of inheritance; but he
was not able to do so
● Gregor Mendel is the “Father of
Genetics”
○ Mathematics and
experimentation uncovered the
basic mechanisms of inheritance
(1866)
○ Choose to work with plants
(Garden pea plants which
showed great deal of variation)
● Mendel subdivided the organism into a
set of manageable characteristics (e.g.:
seed form, stem length)
● Characters were controlled by genes on
different chromosomes and found out
that these genes come in alternative
forms – alleles (could be dominant and
recessive)
Variation within Populations
● Phenotypic variations among individuals
in a population results from the
combined effects of the genes and the
environments.
● Phenotype is the combination of the
genes and the environment.
● Characters are what you inherited from
your parents and the environment you
are exposed to.
○ Because the phenotypic variation
among individuals is the raw
material or substrate upon which
the environment acts during the
process of natural selection,
determining the extent and
sources of variation within
populations is one of the most
fundamental considerations of
evolutionary studies.
Variation Within Populations: Plants
Phenotypic and genetic variation in Potentilla
glandulosa
● Jens Clausen, David Keck, and William
Hiesey (studied in 1940)
● Stanford University in California
● Populations of Potentilla at 30m
(low-land), 1,400m (mid elevation), and
3,050m (alpine elevation) were cloned
○ They grew this plant in 3 different
gardens
○ Exposed to different environment
conditions (e.g Temperature -
high elevation would have much
 colder temperatures compared to
lower elevation)
○ They were able to establish
experimental conditions that
could reveal potential genetic
differences among populations.
○ Their experiment also
demonstrated adaptation to local
environment conditions.
● The actual results did not support the
null hypothesis
What were tested?
● Mid-elevation and Alpine plants attained
1. Genetic differences – tested by using
their greatest heights in the
clones of different populations in
mid-elevation garden, the lowland plants
experimental gardens
grew the tallest in the low-land garden
2. Adaptation to environmental
● Variations we have seen in this plants
conditions – tested by transplanting
could be classified
plants from a population into all
○ Variations due to genetics
elevations
○ Variations due to environmental
adaptation
1. Due to Genetics: height and flowering
differed between three types of plants
a. Environmental effect on plant
morphology
b. Comparing the clones from
different elevations
2. Due to environmental adaptation:
response of a plant type differed at
different elevations
→ Phenotypic plasticity –
● Clones were transplanted into other
variation as a result of
gardens and compared.
environmental differences
Lowland plants differed in how they
grew in the low land garden, mid
elevation and alpine garden. Response
of a plant type (Form and Functions) in
different elevations

In summary, the plant demonstrated

genetic differences among populations
‘
and adaptations to their natural
● If there were no genetic differences
environments.
among populations, all plants would
3. Ecotypes – locally adapted and
grow equally well in gardens.
genetically distinctive population within a
 species. (Thus, the different elevation
plants were ecotypes)
Variation Within Populations: Fish
Variation in alpine fish populations
● Marlis Douglas & Patrick Brunner (2002)
● Lives in the headwaters and lakes of
Alps (South Central Europe) – refuges
for cold-water organisms (following the
last ice age), created geographically
isolated populations → genetic
divergence
● Coregonus (whitefish)
○ The most studied in terms of
morphological variation among
populations
● Ichthyologists have described 19 native
populations of Coregonus (whitefish)
from the central alps
○ There has been a significant
disagreement over the taxonomic
status of these 19 populations.
○ Classification of these
populations ranges from that of a
single variable species with 19
distinctive populations to dividing
the 19 populations into more than
a dozen separate species.
○ The taxonomic status of
Coregonus populations is made
even more difficult by the
hundred year old history of
intensive fisheries management.
○ This fisheries management
included raising them in
hatcheries and moving the fishes
between lakes.
● Do the populations belong in a single
variable species, or do they belong to a
dozen separate species? Are
introduced populations from the
hatcheries genetically similar to their
origin populations?
Genetic and morphological differences among
whitefish populations
● The researchers collected 907
specimens from 33 populations in 17
lakes in the central alpine region. They
used a mixture of anatomical and
genetic features to characterize the fish.
○ Anatomical features included fin
ray counts and pigmentation.
○ Genetic features included
Microsatellite DNA
● Anatomical features and Microsatellite
DNA (tandemly repetitive nuclear DNA
10 – 100 bp long) were studied in 33
populations
● The study demonstrated a moderate to
high level of genetic variation
● Genetic and morphological analyses
was able to distinguish the 19
historically recognized Coregonus
populations
○ Genotypic differences –
correctly assigned individual fish
to indigenous population with
~71% probability
■ If they were given an
individual fish, by getting
its microsatellite DNA
researchers will be 71%
able to assign what
population it came from.
○ Morphological differences – fin
ray counts (69%), pigmentation
(43%)
○ Combination of genetic and
phenotypic data – probability of
assigning a fish to a certain
population increased to 79%
● They found out that some:
○ Introduced populations have also
become genetically distinctive
from their source populations
○ Introduced populations were
genetically similar to the
 populations from which they
came (stock populations and
hatcheries)
● Coregonus of the central alps is made
up of a highly diverse set of populations
that show a high level of genetic
differentiation
● Populations are “evolutionarily
significant” → should be managed as
separate units, and not moved from
one basin to another
○ They suggest that these
populations should be considered
as evolutionarily significant units
(ESUs). These various units
should be managed separately
and they recommend that they
should not be removed from one
lake basin to another in order to
conserve the genetic diversity of
the fish.
Hardy-Weinberg Equilibrium Model
● Mathematical model that population
geneticists use to identify evolutionary
forces that can change gene
frequencies in population.
● Gene frequencies change when
populations evolve.
● Evolution is the change of gene
frequencies in a population over time.
Calculating gene frequencies
● For example:
○ Harmonia axyridis (Asian lady
beetles
■ Include variations in color
patterns on the wing
covers or Elytra colors
with over 200 variants
● Chia-Chen Tan & Ju-Chi Li (1934,
1946); Theodosius Dobzhansky (1937)
○ Determined that the variation in
color patterns shown by the asian
lady beetles is due to the effects
of more than a dozen of
alternative alleles for color
patterns.
 ● Phenotypic expression of those 2 alles
is shown in the figure.
○ 19-signata is a homozygous
genotype (SS) → 2 copies of the
same allele
○ Aulica is also a homozygous
genotype (AA)
○ Their product is a heterozygote
→ (SA)
allele frequencies due to chance or
random events
3. No immigration - entry and exit of
individuals can disrupt the equilibrium of
allele frequencies
4. All genotypes have equal fitness - no
matter what the appearance, there must
be equal chances in having a mate,
kids, and surviving.

Let’s consider a:
● Sample population: The frequencies in Hardy-Weinberg equilibrium
○ Signata SS = 0.81 (81%), can be represented by P and Q.
○ Heterozygote SA = 0.18 (18%),
○ Aulica AA = 0.01 (1%)
● Frequency of alleles
○ SS + ½ SA = 0.81 + ½ (0.18) =
0.9
○ AA + ½ SA = 0.01 + ½ (0.18) =
0.1
● Proportion of matings
○ S sperm x S egg = 0.81
○ S sperm x A egg = 0.09
○ A sperm x S egg = 0.09
○ A sperm x A egg = 0.01
● Proportion of genotypes produced by
this random mating
○ SS = 0.81,
○ SA = 0.09+0.09 = 0.18,
○ AA = 0.01
● The proportion of the genotypes is the
same as the proportion of the genotypes
of the sampled population
○ Allele frequencies in parents
are same in the offspring →
Hardy-Weinberg

Assumptions of the Hardy-Weinberg

Equilibrium
1. Random mating - no preference while
mating
1. No mutations - will introduce changes in
the frequencies of the alleles
2. Large population size – so that no
genetic drift could happen: change in
Process of Natural Selection

What is Natural Selection?

● Natural selection is a mechanism in
evolution that is a result of the
differences in survival and
reproduction among phenotypes
● Darwin pointed out that different
phenotypic characteristics will produce a
high rate of survival and reproduction.
However, this does not occur at all times
and will still vary depending on the
environment, under different
circumstances.
● Forms of Natural Selection: ● Here, cactus species vary on the
○ Stabilizing numbers of spines. Starting population
○ Directional is the light green curve will less than and
○ Disruptive more than the number of spine.
However the extreme phenotypes slowly
Stabilizing Selection disappeared in the population because
● Acts against extreme phenotypes of parasitism and predation.
● As a consequence, it favors the ○ Cacti with less spines were prone
average phenotype. to predation by peccaries (pig-like
animals)
○ Cacti with more spines were
prone to parasitism by insects
laying their eggs to the base of
the spine
○ Cacti with average number of
spines became more
predominant.

Directional Selection
● Happens when one extreme phenotypes
is being favored by selection
 ● In this example, the peccary is selecting
against the low-numbered spine cacti.
● After some time, the predominant
population for the cacti is the
higher-numbered spine.
Disruptive Selection
● Here, the average phenotypes have
lower reproduction and survival
● The predators of the cacti are humans.
Cacti with an average number of spines
produced prettier flowers, so people
used to pick at these cacti.
● Thus, more extreme phenotypes are
favored.
Evolution by Natural Selection and Genetic
Variation
General Postulate of Natural Selection:
● The environment determines evolution
of the anatomy, physiology and behavior
of organisms
● The only way that natural selection can
produce evolutionary change in a
population is if the phenotypic traits
upon which the natural selection acts
can be passed from generation to
generation.
○ Evolution by natural selection
depends upon the heritability of
traits.
Heritability
● Probability that a trait can be passed on
to the offspring
● Potential for evolution by natural
selection is determined by the extent to
which phenotypic variation is due to
genetic variation
● Heritability – proportion of phenotypic
variation in a trait that is attributable
to genetic variance
○ h2 = VG/VP (Heritability)
○ Vg = variance attributable to
genes
○ Vp = variance attributable to
phenotype
● VP = variation in phenotype
○ May be due to genetic effects or
environmental effects
○ VP = VG + VE
● Thus, h2 = VG/(VG + VE)
○ Heritability of a trait depends on
relative sizes of genetic versus
environmental variance →
increases with increased VG,
decreases with increased VE.
○ Scenario 1: If all phenotypic
variations is ALL due to genotypic
variance and none is due to
environment, then heritability
will be 1.
○ Scenario 2: None of the
phenotypic variations is due to
genetic variation and all is due to
the environment then, heritability
will be 0
Adaptive change in colonizing lizards
● Jonathan Losos, Kenneth Warheit &
Thomas Schoener (1997)
● Anolis lizards – approx, 150 spp. in
Caribbean Sea, 250 spp. in Central and
South America
● They measured the length of hind limbs
tradeoff in selection for:
○ Speed: Longer limbs – run faster
○ Efficient movement on narrow
branches: Shorter limbs – move
efficiently on narrow branch
surfaces
● They wanted to see how these lizards
would evolve over type based on their
environment.
● Observed 10 – 14 years after
colonization
● Lizards captured from Staniel Cay (had
diverse type of vegetation e.g. trees,
shrubs, grasses) and introduced into
small islands that differed in
vegetation (almost no trees and shorter
vegetation) → primary agent for
selecting for differences in hind limb size
 ● Anolis populations on experimental
islands and Staniel Cay show a
significant correlation between hind limb
length within populations and average
perch diameter used on the islands
● Increasing relative hindlimb length with
increasing perch diameter.
● Lizards tend to have longer hind limbs
when the diameter of the perch is wider.

Rapid adaptation by soapberry bugs

● Originally, soapberry bugs that feed on

Sapindaceae hosts were native to the
US.

a. X-axis: Difference in height vegetation

with respect to Staniel Cay
b. Y-axis: Resulting morphological
difference of the generations of lizards
after the study period
c. Trend: lizards that were introduced to
the islands that differed from the original
also differed in morphological change
compared when they were placed to
lesser variation.

● Mid-20th century, different species of

Sapindaceae that were exoctic or
non-native were introduced.
Researchers found out that the bugs
shifted their preference from antive to
the introduced species.
● However, the Sapindaceae fruits have a
large variation in the diameter of their
fruits.
○ In Florida, the introduced host
plant has a smaller fruit
● Extent to which colonizing populations compared to the native host.
change morphologically from source ○ In the south central US, the
correlate with differences in vegetative introduced host plant produced
structure between experimental islands larger fruits than the native hosts.
and Staniel Cay
 ○ Beak length of a soapberry bug is
proportional to the diameter of
the fruits that it eats.
■ Larger diameter fruits will
have longer beaks.

However, how do we know that these new

characteristics were due to changes in their
genes and not only due to phenotypic
plasticity?
● Phenotypic Plasticity
○ Ability to change in response to
stimuli or inputs from the
environment
● The shift of bugs in preference from ● Using Transplantation Experiment
native to introduced species came with ○ They transplanted juvenile
evolutionary changes in the soapberry soapberry bugs from introduced
bugs. species to the native species and
● The current populations of the bugs in found out that length of the beaks
Florida have shorter beak lengths to were the same even if the hosts
match the introduced soapberries. were changed.
● While, in the South Central US, they ● Same length in different hosts, therefore
have longer beak lengths. genetic.

Soapberry bugs living on host plant species

with larger diameter fruits have longer beaks.
● Beak length is under selection for
appropriate length → the beak should
be long enough to reach inside the fruits
to get the seeds inside.
Change due to Chance
● Random processes such as genetic drift
can change gene frequencies in
populations, especially in small
populations.
Genetic drift in Chihuahua spruce
● Found in Mexico
● Genetic diversity and population size
○ Due to climatic warming and
fragmentation
● Researchers looked up for Allozymes -
various forms of an enzyme
● They compared the diversity of
allozymes which corresponded to the
genetic diversity.
● The diversity of different populations of
the Spruce were also compared.
● They were able to show that there us a
direct correlation between genetic
diversity and population size:
○ Genetic diversity as low when
population size was low
○ Genetic diversity is high when
population size is high.
Genetic variation in island populations
● Richard Frankham (1997)
● Studied in Australia
● Genetic variation of organisms were
compared between Islands vs. mainland
● Different organisms ranging from moose
and wolves to toads, insects and trees
● Hundreds of comparisons were done
between populations. Out of those, a
large chunk was able to demonstrate
that the relative genetic variation in the
mainland relatively greater than the
island.
● Only a few comparisons were able to
show that there is greater variation in
the island than in the mainland.
● Trend for higher genetic variation in the
mainland population is even greater
when the comparison is between
Endemics and closely related species in
the mainland.
○ There were more comparisons
between mainland and endemic
species that showed higher
genetic variation in mainland.
○ Less comparisons showed that
the genetic variation in the island
is greater than in the mainland.
● In general, there is a higher genetic
variation in the populations in the
mainland. More genetically diverse,
probably because it is larger and there
are more chances for variations to
evolve.
 ● Females with low levels of
heterozygosity – lower survival, lower
rate of hatching eggs, produced smaller
larvae, pupae spent greater time in
pupal stage which made the pupae
more susceptible to predators.

Genetic diversity and butterfly extinctions

● Melitaea cinxia (Glanville fritillary
butterfly)
● Study done in Southwestern Finland
● Researchers measure the genetic
diversity in different populations in terms
of heterozygosity
○ Small populations in small
meadows to Higher populations
in large meadows
○ Higher genetic diversity in larger
populations
○ Smaller populations had lower
genetic diversity due to
inbreeding.
● Inbreeding resulted from having low
heterozygosity) – contributed to higher
extinction rates
 POPULATION DISTRIBUTION, ○ Immigration & Emigration rates
ABUNDANCE, & POPULATION DYNAMICS ○ Growth rates

Gray Whale migration route from Alaska

(feeding ground during the summer) to Baja
(wintering grounds) = 18,000km

Population Distribution and Abundance

1. Distribution Limits - discusses how the
environment limits the population
distribution of species
2. Patterns on Small Scales - how
population in small scales can be
random, clamped or regular
3. Patterns on Large Scales - how
population at larger scales are clumped
4. Organism Size and Population
Density - relationship between this two

Population Dynamics
1. Dispersal - movement of individuals
from 1 population to another.
2. Metapopulations - shows ongoing ● The image above shows the annual
dispersal can join subpopulations to migration of the monarch butterflies from
form metapopulations the south to the north and back again.
3. Patterns of Survival - in terms of Every year, they do this over 1000 km.
survivorship curves and its 3 types In the spring, they mate, then females
4. Age Distribution move on and lay their eggs in host
plants (milkweeds). Eggs hatch, become
Population caterpillars and transform into adult
● a group of individuals of a single butterflies. Several generations are born
species inhabiting a specific area along the way until they reach their
● Characteristics: summer generation in the north. At
○ Distribution – size, shape, autumn time, when it starts to get cold,
location of area occupied they migrate southward in their
○ Spacing pattern wintering ground. Some of them do
■ Random not survive their journey, but the rest of
■ Clumped the population breed and restart their
■ Regular journey, again, the next spring.
○ Abundance – Number of
individual
○ Density – Number of individuals
per unit area
○ Age distribution
○ Birth & death rates
 Population Distribution and Abundance
Distribution Limits

Distribution Limits
● Environment limits the geographic
distribution of species
● Cases:

Kangaroo Distributions and Climates

● Marsupials found in Australia

● G. Caughley et al. (1987) found a close
relationship between climate and
distribution of the three largest
kangaroos in Australia.
● Eastern gray kangaroos (Macropus
giganteus) live in eastern Australia
where precipitation varies little
seasonally or falls mainly in summer
(dominance of summer precipitation →
temperate and tropical forest biomes
● Western gray kangaroos (Macropus
fuliginosus) live in southern Australia
where winter rainfall dominates →
Mediterranean woodland and shrubland
biomes
● Red Kangaroo (Macropus rufus) lives
in central and western Australia where
conditions are hot and dry → savanna
and desert biomes
● None of the 3 kangaroos seem to
occupy the northernmost parts of
Australia.
○ Probably because it is too hot for
the Eastern gray;
○ Too wet for Red kangaroos;
○ Too hot during the summer and
too dry during the winter for the
Western Gray Kangaroo
● It was carefully pointed out that it might
not just be the climate that is directly
affecting the limitations of the
distribution of the kangaroos.
● It can also be that the distribution is
indirectly affected by climate
because climate influences other
factors: food production, water supply
and habitat for the kangaroos.
● Climate can also affect the incidence
of parasites, pathogens and
competitors of the kangaroos.
 A tiger Beetle of Cold Climates

● Tiger Beetle (Cicindela longilabris)

● It lives at higher latitudes and higher
elevations than any other tiger beetle
species found in North America.
○ Its distribution in the North
extends from North Western to
Eastern part of Canada where
temperate and boreal forests
are found.
○ In the south, on the other hand,
these tiger beetles are found to
be at higher elevations like on top
of mountains wherein montane
coniferous forests, very similar
to boreal forests, are found.
● Despite being separated by as much
● During the last glacial period, this tiger as 3,000km and 10,000years of history,
beetle lived farther south compared to different populations of tiger beetles
its present range limits. However, when (tiger beetles in Maine & Wisconsin in
the glacial retreat happened, it also the North, and Colorado & Arizona in
followed the retreat of the glaciers. Now, the South) have similar physiology
its current range is restricted to the (e.g metabolic rates, water loss rates,
south boreal forest. preferred body temperature).
● The retreat of the tiger beetles together ○ Reason for the similarity in
with the retreat of the glaciers is physiology is the similar
supported by fossil records. environmental characteristics
of their habitats.
 Distributions of Plants Along
Moisture-Temperature Gradient
● Species of Genus Encelia
○ E. californica
○ E. actomi
○ E. farinosa
○ E. frutescens
● The four different species are distributed
in different areas in the Southwestern
North America.
● The distribution corresponds to variation
in the leaf pubescence of the plants.
Pubescence refers to how hairy the
leaves are.
● Plants that occur in the westernmost
side are the least pubescent or the
smoothest, least hairy leaves.
● As you go towards the East, the leaves
become more pubescent.
● The variation in pubescence can be
attributed to the climates in the
different areas.
● Western E. californica have smoother
leaves since the temperature there is
usually cool. It does not need hairs to
serve as protection against extreme
heat.
● On the other hand, the areas towards
the inner part of the continent are much
hotter. E. farinosa and E. frutescens
have more hair to protect them from
extreme heat and help regulate leaf
temperatures.
○ E. frutescens, however, is almost
smooth as th E. californica found
in the west. E. farinosa is very
much pubescent. Because of the
difference in pubescence in these
2 species, the amount of
photosynthetically-active
radiation absorbed by these 2
plants also differ greatly.
○ E. frutescens, being smoother,
absorbs a great amount of light,
while E. farinosa, because white
hairs, has less light absorbance
because it reflects a lot of the
light that hits it.
● How does E. frutescens avoid
overheating?
○ The leaves do not overheat
because they transpire at a
high rate. So, such high rates
have a cooling effect similar to
how seating can evaporatively
cool our skin.
● How does it get enough water to
evaporatively cool its leaves if it leaves
in the hottest and driest deserts?
○ Because of microclimates.
Although both species overlap
greatly in a geographic scale,
their microenvironments differ.
They have very distinct areas
where they occupy.
○ E. farinosa grows mainly on slope
habitats in shallow soils that store
limited water. Despite low
transpiration rates, pubescent
leaves of E. farinosa remain
relatively cool because they are
highly reflective
○ E. frutescens rate of transpiration
is very high which is made
possible by selecting areas with
great amounts of water,
specifically ephemeral stream
channels or desert washes.
○ Along the washes, the runoff
infiltrates into the deep soils. This
makes the water highly available
for E. frutescens available for its
high rate of transpiration.
 Microclimates affecting the distribution of
E. frutescens and E. farinosa

E. Californica is confined to a narrow zone along the

coast of California and Baja California, which is cool and
moist in the north and cool and dry in the south.

E. actoni lives gather inland, where

conditions are drier and warmer
than the areas inhabited by E.
californica.

E. farinosa and E. frutescens live still farther in areas

that are much hotter. The geographic distributions of
these 2 species overlap a great deal.

Difference between E. frutescens and E. farinosa

 Distributions of Barnacles Along an
Intertidal Exposure Gradient
Balanus balanoides larvae settle throughout
intertidal zones but survive to adults mainly in middle to
lower intertidal zones.
Chthamalus stellatus larvae settle in the middle
and upper intertidal zones but survive to adults mainly in
the upper intertidal zone.
● The barnacles live in the marine
intertidal zone. There is a steep gradient
of physical conditions in the marine
intertidal zone from the shore up to the
sea.
● At certain times of the day, parts of the
intertidal zone are either submerged to
water or exposed to the air.
● The distribution of organisms along the
intertidal zone is expected to be related
to how much they can resist drying
or desiccation upon exposure to air.
● Joseph Connell (1960’s) - Intertidal
zones in Scotland
● He compared the distribution of Balanus
balanoides and Chthamalus stellatus.
Balanus is usually limited to the middle
and lower levels of the intertidal zone
whereas the Chthamalus is restricted to
the upper levels of the intertidal zone.
● It appears that Balanus is more
vulnerable to desiccation than
Chthamalus. A higher rate of
desiccation may exclude Balanus from
the upper intertidal zone.
● However, from the graph, the larvae of
Chthamalus can actually reach up to the
lower or middle part of the intertidal
zone, but the adults rarely occur there.
○ This can be explained by the
competition between the 2
species. Chthamalus is not a
strong competitor at these
lower levels of the intertidal
zone.
Population Distribution and Abundance
Patterns on Small Scales
● On small scales, individuals within
populations are distributed in patterns
that may be random, regular, or
clumped.
1. Random Pattern
a. An individual has an equal
probability of occurring anywhere
in an area.
b. Processes: Neutral interactions
between individuals, and
between individuals and the local
environment. There are no
interactions that may push or pull
individuals from each other.
2. Regular Pattern
a. Individuals are uniformly spaced
through the environment
b. Processes: Antagonistic
interactions between individuals
or local depletion of resources.
Interactions push different
individuals away such that they
end up having uniformly spaced
distribution patterns.
3. Clumped Pattern
a. Individuals live in areas of high
local abundance, separated by
areas of low abundance.
b. Processes: Attraction between
individuals or attraction of
individuals to a common source.
If individuals are attracted to a
particular resource where there
are other individuals, then a
clumped pattern can be obtained.
Note:
● Small scale patterns depend on the
individual itself.
● Small scale refers to small distances
over which there is little environmental
change significant to the organism
under study.
○ Small scale of a bacterium vs.
Small scale for a gray whale
Distribution of Tropical Bee Colonies
1. Regular and random distributions of
stingless bee colonies in the tropical dry
forest are related to levels of
aggression (Costa Rica - Hubbell &
Johnson, 1977)
a. Trigona, stingless bees can be
found in tropical and subtropical
environments where they gather
pollen and nectar from flowers.
They use trees as their nest and
their colonies are made up of
hundreds to thousands of
workers.
b. Colonies of the stingless bee,
Trigona fulviventris, which
interact aggressively, are
distributed regularly across the
 tract of forest. T. fulviventris from
rival colonies battle daily for 2
weeks for possession of this
potential nest tree.
i. They forage a lot and
group mainly on flowers
that occur in high density
clumps
c. The less interactive stingless
bee, T. dorsalis, is distributed
randomly across the same tract
of forest.
i. They feed singly or in
small groups and on more
widely distributed flowers.
2. They map out the distribution of trees
suitable for nesting. They found out that
potential nest trees are distributed
randomly throughout the study area,
and that the number of potential nest
sites was much greater than the number
of bee colonies.
3. Then, they map out the nest of the
stingless bees where they found out that
species that are REGULARLY
DISTRIBUTED are more aggressive
compared with species that are
RANDOMLY DISTRIBUTED.
a. Aggressive bees fight for
potential nest sites where battles
can occur for long periods of time
(2 weeks). The worker bees can
be 15-30 individuals where they
face and grapple each other.
They hit each other until one falls
down. It is a continuous aerial
display where they do not get
injured apparently. However, due
to an unoccupied nest up to a
thousand bees can die.
Distribution of Desert Shrubs
1. Phillips & MacMahon (1981) showed the
distribution of creosote bushes (Larrea
tridentata) change as they grow in the
Sonoran and Mojave Deserts.
2. They found out that as creosote
bushes enter different stages in life,
they may exhibit different patterns of
distribution.
a. Small shrubs establish in high
densities and produce clumped
distribution.
b. Mortality as the shrubs grow
reduces clumping and produces
a random distribution among
medium shrubs.
c. Competitions enforces a regular
distribution among large
shrubs
● The competition between the adults can
be seen when their root systems are
examined.
● The root systems of 32 creosote bushes
were excavated and mapped.
○ Hypothetical: If excavated
shrubs had a CIRCULAR ROOT
SYSTEMS, 20% of the area
would include extensive overlap
of 4 or more shrubs
○ Actual: The actual root systems
were not circular and overlapped
extensively in only 4% of the
area. The overlap of root systems
in the actual photo is minimized
compared when the actual root
systems are circular
(hypothetical).
○ This proves that competition
between plants produces root
systems such that competition is
minimized.
Population Distribution and Abundance
Patterns of Large Scales
● On large scales, individuals within a
population are clumped.
Bird Populations Across North America
● “Christmas Bird Counts” mapped (Terry
Root 1988); “Breeding Bird Survey” also
analyzed (Brown, Mehlman & Stevens
1995)
● These studies show that at the
continental scale, bird populations show
clumped distributions.
● There are hotspots with very dense
birds. As you move away from this,
densities become sparser and sparser.
● Christmas Bird counts - done every
winter
○ Distribution of Fish Crow and
American Crow
○ Although American crow is
widely distributed across the
continent, it is most abundant in
a limited number of hotspots.
○ On the other hand, Fish Crows
have a more restricted range
but still they show a pattern
wherein there are hotspots and
as you go outward it, the
densities of the crows become
more sparse → Within its
restricted range, fish crows live in
high density areas.
● Breeding Bird Survey - done during
June (breeding season of birds)
○ A relatively small proportion of
study sites yielded most of the
records of each bird species.
Observers counted a few
red-eyed vireos along most
census routes.
○ But, large numbers of red-eyed
vireos were encountered on just
a few census routes.
○ There are only a few areas where
red-eyed vireos occur at high
density. There are more areas
where red-eyed vireos occur at
low density.
Plant Distributions Along Moisture
Gradients
● Santa Catalina Mountains, Arizona (left)
and Great Smoky Mountains,
Tennessee (right). (Whittaker & Niering
1965; Whittaker 1956)
● Clumping patterns can also be observed
in wood plants and trees along
moisture-gradients in several mountain
ranges across North America.
● Environmental conditions in mountain
sides change substantially with
elevation.
● The higher the slopes are dirier whereas
the bottom of the canyon is more moist.
● Different trees and plants are distributed
with regards to the different amount of
moisture
○ Douglas firs are more abundant
in moist canyon bottom
○ Arizona madrones are more
abundant at mid slope.
○ On the mountainside, Mexican
pinyon pines are most abundant
on drier upper slope.
 Population Distribution and Abundance
Organism Size and Population Density

● Population density declines with

increasing organism size

● Animal Size and Population Density

● John Damuth, 1981
○ Body size and population density
of herbivorous mammals
○ Range: 307 spp. from rodents
(10g) to rhinoceros (106 g )
○ He collected the information on
the average densities ranging
from 1 individual per square km
to 10,000 individuals per square
km.
○ There is an inverse relationship
between the herbivory density
and body mass. Average
population density of herbivorous
mammals decreases with
increasing body size. → As their
body size increases, their density
decreases.
● Peters & Wassenber, 1983
○ Overall, average population
density decreases with increasing
body size across a wide
spectrum of animals groups
■ Aquatic Invertebrates
■ Terrestrial invertebrates
■ Mammals
■ Birds
■ Vertebrate poikilotherms
● Similarly, trees in Tennessee also exhibit ○ Many aquatic invertebrates live at
same pattern: higher population densities than
○ On mountainsides, table other animals of comparable size.
mountain pines are most ○ Mammals tend to live a higher
abundant on drier upper slopes. population densities than birds
○ Red maples are most abundant ○ From this, they concluded that it
at mid slope is best to study aquatic
○ Hemlocks are most abundant on invertebrates and birds
the moist valley bottom. separately from the other
mammalian groups

● Plant Size and Population density
● White, 1985
○ Plant population density
decreases with increasing plant
size across a range of plant
growth forms (smallest to largest)
○ Duckweed, Lemna, one of the
smallest flowering plants lives at
very high population densities.
○ As in animals, plant population
sanity decreased with increasing
plant size across a wide range of
plant growth forms
○ The coast redwood, Sequoia
sempervirens, one of the largest
trees, lives at one of the lowest
population densities.
Population Dynamics
● Populations are not frozen in time but
are in continuous flux.
○ Population distribution and
abundance may vary across time
● Patterns of distribution and abundance
result from a dynamic balance between
factors which are represented below:
● Nt = no. of individuals in a population at
some time t
● Nt-1 = no. of individuals in the population
at some previous time t – 1
● B = no. of births that have occurred
during time interval between t – 1 and t
● I = no. of immigrants to the population
during time interval
● D = no. of deaths
● E = no. of individuals that have
emigrated
Population Dynamics
● Area of ecology concerned with
factors influencing expansion,
decline, or maintenance of
populations.
● Population dynamics is a key to
understanding:
○ Decline and extinction of
endangered species
○ Control of noxious pest species
(e.g. parasites and pathogens)
○ Maintenance of economically or
culturally important species
● You may not noticed the dynamic nature
of populations but it become quite
apparent when a conspicuous species
expands its range:
Expansion of Africanized Honey Bees ■ Cold winter temperatures
● Africanized Honeybees will likely halt the northern
○ They live from tropical to cold spread of Africanized
environments honeybees (2005).
○ Across this extensive ■ Africanized honey bees
environmental range, the have not permanently
honeybees have differentiated colonized South America
into a number of locally adapted south of 340 S latitude
species.
○ Apis mellifera scutellata in Africa
○ They are adapted to the tropical
climate.
○ In 1956, Brazil wanted bees
adapted to tropical climates. By
this, they mated the regular
honey bees to african honey
bees producing the africanized
honey bees.
○ Unfortunately, Africanized honey
bees were found to be more
aggressive than the regular
European honeybees. → Swarms
of these Killer bees attack
humans, producing more stings.
○ Unfortunately, these honey bees
also do not produce a high
amount of honey expected of
them by scientists.
■ Reason: Warmer climate
and greater stability of
nectar sources eliminates
the need for storing large
amounts of honey and
maintaining large colonies
for survival through the
winter.
○ Produce swarms and disperse to
form new colonies at higher rates
(300-500km/year). → Fastest
dispersal rates.
■ Most of South America,
most of Mexico and all of
Central America has the
population of it.
 Population Dynamics
Dispersal
● Dispersal can increase or decrease
local population densities
● Different species have evolved different
ways for them to be dispersed away
from their parents.
○ Wind blown seeds of dandelions
can disperse long distances, and
then once they find their
destination they grow into a
sedentary stage the dandelion
plant
○ Waterborne larvae of barnacles
can also disperse and then they
become sedentary when they
become adults
○ Juvenile spiders that disperse by
spinning a silken thread that
catches the wind, and then they
become sedentary when they
build their web in the garden
● Dispersal is the movement of
individual/s away from the population
in which they were born
○ Immigration – dispersal into the
local population
○ Emigration – dispersal out of the
local population
Dispersal of Expanding Populations:
Eurasian Collared Doves
1. In less than 60 years Eurasian collared
doves (Streptopelia decaocto) expanded
their range to the farthest corners of
Europe
2. Eurasion collared doves began to
spread out of Turkey into Europe early
in the 20th century.
a. The expansion of the population
began suddenly and it was
relentless once it started.
b. By 1980’s most of Europe was
already colonized by this bird
species. The expansion also was
not initiated by humans.
c. The expansion took place in
small jumps
i. Only adult eurasian
collared doves are highly
sedentary while only the
young doves disperse
from the places where
they were born.
ii. Dispersing young doves
actually dispersed for only
a few kilometers away
from the nest where they
were born
iii. However, some of these
birds actually disperse
hundreds of kilometers
away from their nest, but
as seen from the graph,
most birds disperse only a
few kilometers.
iv. Once the birds have
chosen a mate, they nest
and become sedentary like
their parents.
d. These pulses of dispersal by the
young birds spread the Eurasian
collared dove population across
Europe at a rate of about 45
kilometers per year.
3. Most collared dove fledglings disperse a
few kilometers
4. But some disperse hundreds of
kilometers
 4. Hemlock spread north and west from
the southeast.
5. In contrast, hemlock did not reach its
present range limits until just 2,000
years ago.

Dispersal in Response to Changing Food

Supply

1. Numerical response to changes in prey

availability
2. Kestrel and owl densities closely follow
variation in vole densities in western
Finland.
3. Dispersal in these predators (Kestrels
and owls) happens in response to
changing food supply.
a. Over several years, the
populations of kestrel and owls
follow the variation in the
population in voles (prey) in
western Finland. → Numerical
response to changes in prey
availability

Range Changes in Response to Climate

Change
1. It started from the time that the glaciers
retreated northward during the last ice
age around 16 000 years ago.
2. Maple spread north and east from the
southwestern part of its range.
3. Maple reached the northeastern part of
its present range about 6,000 years ago.
 Dispersal in Rivers and Streams upstream limit of the
1. Organisms that live in moving bodies of population.
water have to have characteristics that iv. During their migration back
allow them to maintain their positions up the river, some of
even if the water is moving. these snails in the
2. However, despite these means of migrating population may
staying in place, organisms still get join the populations that
washed downstream in large numbers are stationary along the
→ drift way.
a. Opposite of drift is the upstream v. Conversely some
movement individuals from the
b. Drift moves organisms stationary sub-populations
downstream, sometimes actively may join the population
as behavioral drift, sometimes that is migrating upstream
passively with floods.
c. Many organisms engage in 4. This proves that populations can be
upstream movements that appear very dynamic.
to compensate for downstream
drift.
3. The dynamic interplay between the
downstream and upstream dispersal is
called the colonization cycle.
a. In the colonization cycle,
upstream and down dispersal
and reproduction have major
influences on stream populations.
b. A wave of migrating snails,
Neretina latissima, in Rio Claro,
Costa Rica.
i. This species of snails lay
eggs and produce free
living planktonic larvae
that drift down to the
pacific ocean.
ii. When the larvae
metamorphose into small
snails they re-enter the Rio
Claro and begin moving
upstream in huge
migratory aggregations of
up to half a million
individuals.
iii. These aggregations move
very slowly and may take
up to one year to reach the
 Population Dynamics ○ It seems like butterflies were
Metapopulations dispersing – moving from one
population to another.
● Ongoing dispersal can join numerous ● Larger meadows tend to support larger
subpopulations to form a populations of these butterflies.
metapopulation.

Metapopulation
● A group of subpopulations living on
patches connected by exchange of
individuals among patches
● Ex. Individuals S, I, and R are members
of subpopulations which are exchanging
individuals through migration. These
different subpopulations are further
connected to other subpopulations
forming the metapopulations.

● Butterflies are more likely to leave

smaller populations than larger ones.
○ Butterflies in small populations
immigrate to populations that are
larger.
○ This indicates that alpine
A Metapopulation of an Alpine Butterfly meadows, because of the
● Rocky Mountain Parnassian Butterfly decrease in size, can eventually
(Parnassius smintheus) occurring in the lose the populations of butterflies.
alpine region. ○ Perhaps, the range of these
● They were studied in 20 Alpine butterflies will become more
meadows – ranging from small to large, contracted as more factors affect
and were also either adjacent or the decrease in size in the Alpine
separate. Meadows.
● They have been decreasing in size due
to fire suppression and global warming
● Researchers caught and marked the
butterflies. They were then released and
recaptured to estimate population size
and follow butterfly movements.
● They were able to recapture some
butterflies that were marked in one
subpopulation and they were recaptured
in a different subpopulation.
 Population Dynamics
Patterns of Survival
● A survivorship curve summarizes the
pattern of survival in a population.
Life Table
● Predicts whether a population grows or
sinks by identifying birth and death rates
for organsims at different ages, as well
as, the current age and sex makeup of
the current poulation.
● A life table summarizes birth and
death rates for organisms at different
stages of their lives.
● It shows the likelihood that organisms in
a population will live, die, and/or
reproduce at different stages in their
lives.
● Example:
● Life Table for the total population
of US in 2016 – Static Life Table
○ Takes a snapshot of a
population at a period of
time and does not follow
the different individuals
from birth to death.
● Parameters included:
○ Number of surviving
population
○ Number of dying
populations
○ Life expectancy at each
particular age
Types of Life Tables
1. Cohort Life Table – most reliable way
of estimating patterns of survival
within a population; More difficult to
come up with since one needs to
identify a large number of individuals
that are born at about the same time
and keep records on them from birth
to death
2. Static Life Table – record the age at
death of a large number of individuals;
shows a snapshot of survival within a
population within a short interval of
time.
Life Table to Survivorship Curve
1. Life table of the Dall sheep. Study done
in the Mt. Denali National Park in
Alaska.
2. The researcher collected the skulls of
608 sheeps that died from various
causes.
3. He identified at which age the sheep
died by counting the growth rings in their
horns and by studying toothware.
4. Table summarizes the survival patterns
of the sheeps based on the sample of
skulls.
a. First column -lists the ages
b. Second column - lists the number
of surviving sheep in each age
class
c. Third column - lists the number of
dying sheep at each age class
5. Although only 608 skulls were studied,
the numbers in the table were
expressed to a thousand. This
 adjustment is made to ease the
comparisons with other populations.
a. To allow comparisons to other
studies, number of Dall sheep
surviving and dying within each
year of life is converted to
numbers per 1,000 births
b. Subtracting the number of deaths
(ex. 199) from the number alive
at the beginning of each year (ex.
1000) gives the number alive at
the beginning of the next year
(801).
6. Plotting age on the x-axis and number of
survivors on the y-axis creates a
survivorship curve.
7. The shape of the curve (Type I) shows
that the Dall sheep surviving their first
year of life have a high probability of
surviving to about age 9 (relatively flat).
8. However, sheep 10 years old and older
are easier prey for wolves and die at a
high rate.
Rotifer
Survivorship Curve
1. In Type I, juvenile survival is high and
most mortality occurs among older
individuals.
a. At a certain age, the slope is
relatively gradual (flat)
b. At another certain age, mortality
is high, and the slope becomes
steeper.
2. In Type II, individuals in a population die
at equal rates, regardless of age.
a. Slope is a straight line and the
same.
3. In Type III, individuals showing
survivorship die at a high rate as
juveniles and then at a much lower rate
later in life.
a. Opposite of Type I
● Type I - Gradual decrease of organisms
in the beginning and then, a steep
decline towards the end.
● A similar pattern of survival by the
rotifer, Floscularia conifera, is complete
within 11 days.
White-crowned Sparrow and American
robin
● Type II - relatively straight line, slope
does not change that much as you go
from younger to older species.
● Like many other bird species,
white-crowned sparrows and American
robins show approximately constant
rates of mortality.
Perennial Plant - Cleome droserifolia
● Type III
● The vertical scale has been extended so
that survivors appear on the graph.
● Out of 1 million seeds, only 39 plants
survive to 1 year of age.
● Upon reaching 1 yer, the slope of the
graph became gradual – high
survivorship among individuals that
reach higher age (1yr and above).
Population Dynamics
Age Distribution
● It is similar to static and cohort life tables
such that they are all ways to estimate
patterns of survival within a population.
● The age distribution of a population
reflects its history of survival,
reproduction, and potential for future
growth.
● An age distribution consists of the
proportion of individuals of different
ages within a population.
● Philippines (2018) and Japan (2021)
Age Breakdown
● Age range that makes up the majority of
the population
○ Philippines - under 15
○ Japan - 45-50 & 60-74
Stable and Declining Tree Populations
Quercus alba, White oak in Illinois
Dynamic Population in a Variable Climate
Age distribution of a population of large
cactus finches Geospiza conirostris on the
island of Genovesa in the Galapagos
Islands

1. The age structure of this population of

white oaks shows that older trees are
being replaced by young trees.
2. This population of white oaks is
dominated by young individuals.
3. The population is declining.
● 1983 - A fairly even distribution
individuals among several age
classes suggests several years of
Populus deltoides, Rio Grande Cottonwood
successful reproduction
in New Mexico
● Six-year-old finches are absent because
the birds did not nest during the 1977
drought.

1. The age structure of this population

shows that older trees are not being
replaced by young ones
● Age distribution in 1983 contrasts with
a. The absence of young tree
1987 distribution even if they are only
suggests that this population will
separated by 4 years.
not persist
● 1987 - population is dominated by
b. 40- to 50-year-old trees dominate
4-year old birds. The population is
this population
dominated by birds hatched during the
2. Population is stable
abundant rains of 1983.
 ● Droughts in 1984 and 1985 prevented
nesting
● Droughts of 1984 and 1985 reduced
numbers of older birds
● The 1977 reproductive failure is still
evident in the 1987 distribution.

The study demonstrates the responsiveness of

the population age structure to environmental
variations.
 POPULATION GROWTH
1. Geometric and Exponential
Population Growth
a. growth experienced by a ● Annual plant (Phlox drummondii) –
population when the resources
are ABUNDANT
2. Logistics Population Growth
a. type of growth the population
experiences when resources
STARTS TO DEPLETE
3. Limits to Population Growth
a. shows that the environment can
limit the population growth by
changing the birth and death
rates
POPULATION GROWTH
● Definition: how the number of
individuals in a population increases or
decreases with time
● Growth is controlled by rates of
○ birth,
○ death,
○ immigration, and
○ emigration.
● Open vs. Closed population
● In a closed population,
○ growth is determined by birth
rate and death rate
○ no immigration and emigration,
○ immigration is equal
emigration
*Note: In the lectrue only closed populations will be
discussed. No interference of immigration and
emigration
GEOMETRIC & EXPONENTIAL
POPULATION GROWTH
● In the presence of abundant
resources, populations can grow at
geometric or exponential rates
Geometric growth in populations with
discrete generations
populations grow in discrete annual
pulses. (Discrete - every year, the
population is different from the previous
one)
● Every year, the current population will
develop, reproduce and leave behind
seed. Then, they will die. The seds they
left behind will start the new generation
of Phlox. Then this genertaion will die
but leaving behind a group of seeds to
start a new generation.
● The growth by any population with
PULSED reproduction can be modeled
by Geometric Population Growth.
○ This is where successive
generations differ in size by a
constant ratio.
○ The size of a population growing
gemotrically at any time t can be
modeled here:
● Nt = number at time t
● N0 = number at initial time 0
● λ = geometric rate of increase = Nt+1/Nt
○ Average number of offspring left
by an individual during a 1 time
to interval
t
● = number of time intervals in hours,
days, years, etc.
1. The starting population was 996.
2. After a year, the population increased to
2,408.
3. Dividing the starting population from the
current population, a λ value of 2.4177
will be obtained.
4. Growing geometrically, the number of
phlox growing at any point in time, can
 be generated by using the (1) equation, 1. r (per capita rate of increase) is a
or (2) multiplying the previous constant and N is a variable.
population size to the value of lambda. 2. As population size (N) increases, the
5. At the seventh year, the population grw rate of population growth increase
up to 480,924. (dN/dT) increases, because the
6. Multiplying it with the lambda value, the constant R is multiplied by an increasing
population will increase up to 1,162,730 size N.
by the 8th year. 3. The formula can also be expressed like
7. Hypothetical Situation: Could be this:
possible if the resources are abundant.

Exponential growth of a colonizing population

of Scots pine, Pinus sylvestris (Bennett 1983)
Exponential growth in continuously
reproducing populations with overlapping
generations

● Continuous population growth in an

unlimited environment can be modeled
as exponential growth:
dN/dt = rmaxN
Or
dN/dt = (b – d) N

● N = number of individuals
● t = time
● rmax = intrinsic rate of increase
○ (constant)
○ per capita rate of increase
● b = instantaneous birth rate
● d = instantaneous death rate
1. Over hundreds of years, the increase of
the population of Scots pine followed an
exponential growth.
2. They were actually able to count the
populations by using the records of
pollen in lake sediments. They carbon
dated these sediments and identified
what period the pines came from.
3. Their populatio started with a low
amount and increased quickly in an
exponential manner due to a very Exponential population growth
favorable environment. (Retreat of the
glaciers) Equations:
1. dN/dt=rmaxN (differential equation form)
Per capita rate of increase (r) may be 2. Nt =N0ermaxt (to calculate population size)
positive, zero, or negative
For a population to demonstrate exponential
● Graphing population size vs dN/dT: population growth, the population conditions
○ You will get a line with a slope of must be observed.
r Conditions:
■ r = b – d = realized or ● Start with low population density
actual per capita rate of ● Favorable environment (No food or
increase resource limitations)
■ Slope is r → determines
the shape of the growth. Aransas National Wildlife Refuge
○ r=0, no change in population size
(flat line; instantaneous brith rate
is equal to instantaneous death
rate)
○ r<0, decrease in population size;
r>0, increase in population size.

1. Whooping crane, an endangered

species
2. Winters in the Texas Gulf Coast
3. Breeds in Wood Buffalo National Park,
Canada
4. Recovered from near extinction in 1941
5. From 22 to 220 individuals from 1940 to
2005 - Conservationists made it their
mission to conserve their population.
 Slowing of exponential growth LOGISTIC POPULATION GROWTH

● As resources are depleted, the

population growth rate slows and
eventually stops.
● Since everything on earth is finite, it
seems impossible that any population
on earth will continue to grow at an
exponential rate, indefinitely.
● At some point, the population will
encounter will encounter environmental
problem growth.
○ The population of Eurasian
collared doves does not follow
exponential growth at some point
○ It will eventually taper off and will
show a graph that is more of an
● Eurasian collared doves expanded S-Shaped curve or signmoidal.
beyond its historical range in the
Western Europe during the 20th century. Sigmoidal (S-shaped) growth curve
● As the birds spread to new territory, its
populations grew in exponential rates for Growth of yeast and Elephant seal
a decade or more.
● However, in less than 20 years,
population size was less than that
predicted by the exponential model,
suggesting that population has slowed.
● It seems that instead of following the
graph, the slope become a bit more
gradual and the population growth
has slowed down.
● Here, the environment slows down
populaition growth.

1. When the density of the yeast is very

low, population tends to increase at a
higher rate (slope is steep).
2. As it reaches a certain population level,
the population growth slows down and
levels off.
 1. At some point, the population plateus
and fluctuates around a population level.

Carrying capacity

● Represented by K
● Maximum sustainable population size
under prevailing environmental
conditions
○ It is the number o individuals from
a particular population that the
environment can support.
○ It is NOT a fixed number. It can
change depending on the
prevailing environmental
conditons that population is
exposed to.
● Determined by a complex interplay
among factors such as food,
parasitism, disease, and space

Carrying capacity is reached in these

populations at:
1. Days for the paramecium
2. Weeks for barnacles
3. Years for buffalos Logistic population growth
● The logistic model was proposed to
account for patterns of growth shown by
 populations as they begin to deplete
environmental resources.
● Populations ecologists build the logistic
population m,odel by modifying the
exponential growth model.
● The original model for exponential
growth by insertion an element, K
● Add an element that slows growth as
population size approaches K
● This formula shows how population size
can affect population growth.
● As N approaches K, growth decreases
until it reaches zero at N=K
● Growth is fastest when N=K/2
● N/K → “environmental resistance”
○ As the size of the population
growth gets closer to carrying
capacity, environmental factors
increasingly impedes further
population growth.
Relationship between population size, N, and
realized per capita rate of increase, r
● If N<K, r is positive and the population
grows.
○ If N = ½ K, the population growth
is highest.
○ Halfway the 0 and the carrying
capacity, the steepest point of the
curve is found here.
● If N = K, r = 0 and population growth
stops; growth rate is equal to 0 and will
plateau at the carrying capacity, K.
● If N > K, r is negative and the population
declines
● In the logistic model, the realized per
capita rate of increase or r will
decreases as N increases.
● r is maximum when the
population is very low.
● As the population increases, r
decreases until it reaches N = K
● Popualiton growth will stoip when
N = K where r = 0
● Beyond carrying capacity, r
becomes negative, and the
population growth enters a
decline.
Relationship of density to per capita rate of
increase in populations of Daphnia pulex
1. They increased the density of Daphnia.
2. They were able to show that as the
density increases, the per capita rate of
increase decreases.
3. At some point, around densities 24 and
32, r was negative which indicates a
decline in population growth.
LIMITS TO POPULATION
GROWTH
● The environment limits population
growth by changing birth and death
rates
Darwin’s Finches & Galapagos Rainfall
Medium-ground finch, which is the dominant
finch in the Daphne major island in Galapagos.
1. Galapagos islands experience a
variable rainfall year to year and plants
in the islands respond to this.
2. They were able to document a drought
at 1977.
a. It lead to a decline in the
poppulation growth of finches:
from 1200 individuals to 180
individuals – 85% decline in just 1
year.
b. Although it is possible that the
decline could be due to
emigration, the primary reason
for the decline is death due to
starvation. Death rates due to
starvation far exceeded birth
rates.
3. From 1977 to 1982, the population
averaged to about 300 individuals.
4. In 1982, abundant rains led to high rates
of finch population growth
a. It was due to increased birth rate
as a consequence to an
abundance of seeds that the
adult finches eat, and abundance
of caterpillar that the finches feed
to their young.
b. In the presence of abundant food,
birth rates greatly exceeded
death rates.
Environment and Birth and Death among
Galapagos finches
Availability of caterpillar and fledgling of young
medium ground finches on Daphne Major (data
from Gibbs and Grant 1987)
Populations of Darwins’ finches and their food
plans are instructuive of hwo the environment
can affect britha nd death rates.
Factors affecting carrying capacity
Population Limiting Factors
1. Limiting factors are environmental
factors that restrict the growth of
populations.
2. Types of Limting Factors:
a. Density-dependent factors -
limitations to population growth
that vary with population density.
i. The size of the population
itlself influences its
numbers.
ii. Ex. Availability of food and
water, Competition for
food, Disease, Predation
b. Density-Independent factors -
limitations to population growth
that are not dependent on
population density.
i. Ex. pollution, Extreme
weather, Seasonal
changes, Catastrophic
events
3. Major canges to any population can
create imbalance between individuals
and their environment.
4. Elimination of wolves in the Yellowstone
National Park,
a. Population of deer and elssoared
with no predator to reduce them.
b. Competition for food increased.
c. Both populations spread to new
areas totally changing the food
supply available to other species.
d. The imbalance was corrected by
reintroducing wolves into yellow
stone
1. Factors that INCREASE
a. Changing climate or other
environmental factors
b. Technologic efficiency advances
c. Decrease in body size and
resource demand
2. Factors that DECREASE
 a. Changing climate or other species within an
environmental factors ecosystem.
b. Environmental degradation ii. Some species of food for
c. Depletion of nonrenewable other species which is
resources known as predation
d. Introduction of new competitor iii. Predation occurs to both
animals and plants. For
Limiting factors can also be categorized into example, rabbits prey on
Abiotic Factors and Biotic factors the grasses and hawks
may prey on the rabbits.
1. Abiotic - nonliving When a resource is
a. Water, temperature, and scarce, competition occurs
minerals, that affects an and this can be direct or
ecosystem such as water and indirect.
climate iv. Direct competition -
i. Water is essential for life individuals fight each other
and organisms have for a single resource. Such
evolved different as when deer fight for
adaptations to ensure that mate.
they get the water they v. Indirect competition -
need. when one species
ii. Example: Animals either indirectly affects another
live in water or obtain species’ resources. If the
water through drinking or deer eat all the grasses
eating foods that contain leaving none for the
water. In desert, rabbits.
environments plants have b. Competition can lead to a
adapted to retain water decrease in the size of one
limit the loss of water or population and increase in the
reach water through size of another population or a
extensive root systems shift within a population.
iii. Climate is another abiotic c. Population can also be affected
factor that strongly by pathogens which are
influences plant and organisms that cause disease
animal populations as the i. The more dense the
space which affects shelter population, the greater the
avaibility and resources. chances the disease will
2. Biotic - living spread.
a. Prey, predators, pathogens,
injury, competions, disease, old
age
i. Food may be abundant or
scarce and the availability
of food may vary among
 LIFE HISTORIES
1. What is Life History?
2. Offspring Number vs. Size
a. The larger the offspring, tend to
produce lerss offspring and vice
versa
3. Adult Survival and Reproductive
Allocation
a. Where organisms tend to survive
lower, reproduce at an early
stage; tends to allocate more
energy to reproduction
4. Life History Classification
a. Life history can be classified on
the basis of a few characteristics
of the population like number of
offspring, survival, etc.
What is Life History?
Life History
● Describes the series of events over its
lifetime (birth to death) pattern of
growth and reproduction
○ It includes how resources are
allocated for growth,
maintenance, and reproduction.
○ It looks on how life history traits
affect the life table of an
organism.
● It is genetically determined and shaped
by the environment and natural
selection.
○ Darwin’s “natural selection” →
reproductive success of a
generation; the concept of fitness
(number of offspring or genes
contributed by an individual to
future generations).
○ Adaptation by natural selection is
based on variation in the
Darwinian Fitness Sense, and
since life history traits determine
the survival nd reproduction, the
two are the major components of
fitness.
○ An organisms is only
“evolutionary fit”, if it can survive,
reproduce and contribute to the
gene pool of the future
generation.
● Life history of organism involves:
○ Trade-offs among the different
characteristics of the population
such as
■ Fecundity - number of
offspring that certain
individual will produce
■ Relative offspring size
■ Survival
■ Age at reproductive
maturity
Adult mayflies generally live for one day
only
● Unlike mayflies, humans are born
helpless and needs to be taken care of
until they are old enough. At the start of
their teenage years, they become
reproductively mature and spend a
relatively long time in old years and
pass away.
● Adult mayflies can only live for 1 day.
Meanwhile, their larvae can live in water,
stream for about a year.
● Adult mayflies emerge after 1 year and
can only have 1 day to find a mate to
reproduce and lay eggs. However, they
get to produce hundreds to thousands of
eggs.
Environmental uncertainty dictates
reproductive strategies
The kind of life history reproductive
strategy that organism exhbites depends a lot
on environmental uncertainty.
1. Semelparity – all offspring produced in
a single reproductive event
a. (Ex: sockeye salmon)
2. Iteroparity – reproduction in
successive years or breeding
seasons
Sockeye Salmon
Sockeye salmon – swim as far as 6,000 km
from Pacific Ocean feeding grounds to
spawning streams, lay thousands of eggs, then
die from the exertion.
Semelparity vs. Iteroparity
1. Semelparous – Most insects and other
invertebrates, some fish (salmon), and
many plants (bamboo, ragweed) –
Some are small, short-lived, grow in
disturbed habitats
a. Century plant spends a relatively
long time growing and
developing. Once it reaches
reproductive maturity, it bears a
gigantic inflorescence that
flowers and roduces fruits. After
that, it dies leaving hundreds of
offsprings behind.
2. Iteroparous – Multiple cycles of
reproduction mean the organism must
balance growth, maintenance, escaping
predators, defending territory, etc
against reproduction – Most vertebrates,
perennial plants.
a. Mango trees are perennial and
undergoes a lto of reproductive
events throughout its lifetime
which can be possibly decades.
Degree of development at hatching
(*used in the field of ornithology)
1. Altricial – newly hatched or born are
relatively immobile, lack hair or
downfeather, and must be cared for
by adults; closed eyes are common
(ex: herons, hawks, woodpeckers, owls,
and most passerines; rodents,
marsupials, primates, cats, dogs, and
humans);
a. Altricial organisms are also called
“Nidicolous” – stays in the nest
longer (nidi = nest + colous
=stay)
2. Precocial – young are relatively
mature and mobile from the moment of
birth or hatching (ex: chicken, cow, deer,
turkey); “imprinting” is characteristic of
precocial birds; has some form of
downfeather.
a. Precocial organisms are also
called “Nidifugous” – leaves nest
early (nidi = nest + fugous = to
flee)
i. Once describe precocious,
you want to be described
as mature even when you
are still very young.
b. Imprinting - there is a critical
period of time in its life wherein it
forms attachments and develops
concepts of its own identity. Once
it sees a different species of bird
upon hatching, it will identify with
that species and not its own.
3. Two types are not sharply demarcated
→ range of precociality.
 Offspring Number vs. Size

Because all organisms have access to

limited energy and other resources, there is a
trade-off between the number and size of
offspring; those that produce larger
offspring are constrained to produce fewer,
whereas those that produce smaller
offspring may produce larger numbers.

Principle of Allocation
1. Principle of allocation – if an organism
uses energy for one function such as
growth, it reduces the amount of energy
available for other functions Relationship between growth and
a. Because organisms have a reproduction
limited amount of energy and ● Growth is represented by relative wiidth
resources available for them, of the annual rings produced in woody
they have create a way to create plants while Reproduction is
trade-off between growth, represented by the number of cones
maintenance, and reproduction. produced.
2. Reproductive effort – time and energy ● There is an indirect relationship between
allocated to reproduction growth and production.
3. Trade-off between growth, ○ As the number of cones
maintenance, and reproduction. – produced increases, the number
There is a negative relationship between of the rings and size of the trees
annual plant growth and the allocation to decrease.
reproduction.
Fecundity and Clutch size
Percentage of annual production to 1. Fecundity – number of eggs or seeds
reproduction varies widely in different types of produced by an organism;
organisms: a. It is the physiological maximum
1. Perennials: 15-20% potential reproductive output of
2. Wild annuals: 15-30% an individual (usually female)
3. Crops: 25-30% over its lifetime → may depend
4. Corn and barley: 35-40% on body size and/or age.
5. Lizard: 7-9% 2. Clutch size – (applied to organisms that
6. Salamander: 48% lay eggs) the number of offspring per
brood (set of young hatched or born at
the saame time by the same parents)
a. It is defined as the number of
eggs released during a single
spawning event, and spawning
frequency is the number of
 spawning events in a time
interval
b. 1 or 2 large live young in mako
sharks, and 600,000,000 eggs
are layed by ocean sunfish.
● The size of the individual is directly
related to its fecundity.
Egg size & number in fish
● Fecundity of the Mako Shark and the
ocean sunfish differ greatly. Aside from
the clutch size difference, they also
differ in a lot of variable such as body
size and toher aspects of life history.
Organisms belonging in similar group of fish
family.
● Tom Turner & Joel Trexler (1998) →
study on darters, small freshwater fish in
the perch family, Percidae
● Even when they are belonging in the
same group of fish, there is still a
trade-off between offspring size and
number.
● The females of larger darter species
produce more eggs. There is direct
relationship between size and number of
darter eggs.
● Darter species that produce larger eggs
lay fewer. There is a negative
relationship between egg diameter and
number of eggs.
● Larger eggs also produce larger larvae.
● The larger larvae begin feeding on prey
that live on the stream bed at an earlier
age compared to smaller larvae. They
 spend more time in the area where they
are hatched compared to the smaller
larvae that drifts away from their
hatching place.
● Larvae hatching from larger eggs
disperse only in shorter distances and
therefore, carry genes in short distances
as well. This leads to evolutionary
consequences produced from offspring
size and number tradeoff.

Offspring size & number tradeoff has

evolutionary consequences
● Population of species producing fewer
larger eggs will be more isolated
genetically from other populations.
○ Ex. Darters producing larger eggs
have lower gene flow among
their populations.
● Because of their isolation, such
populations will differentiate genetically
more rapidly compared to populations of
species that produce many smaller
larvae that disperse longer distances.
Seed size and number in plants
● Plants that produce larger seeds
produce fewer.

● Darters producing more eggs

show higher rates of gene flow
among populations

Plant growth form and dispersal mode are

associated with differences in seed size
● The sizes of seeds of plants are while unassisted and wind-dispersed
influenced by plant growth form and are smallest.
dispersal mechanism
● This study classified the plants based on
their plant growth form:
○ Graminoid - grass-like
characteristics
○ Forb - herbaceous
○ Woody
○ Climbers
● On average, climbers produce the
largest seeds, while graminoid plants
produce the smallest seeds.

Large vs. Small Seeds

Large Small

Numbers Fewer More

Enivronment Relatively Higher

more stable frequency of
disturbances

Resistance Higher Less

● After that, they also classified the plants to hazards survival to resistant
based on their dispersal mechanism. the face of
○ Unassisted hazards
○ Assisted by wind
Ability to Lower Greater
○ Assisted by adhesion to animals colonize capacity to capacity to
○ Assisted by ants new areas colonize colonize
○ Assisted by vertebrates that eats newlyopened newlyopened
the fruits spaces spaces
○ Assisted by a scatterhoarding
animals like squirrels
● Vertebrate dispersed and
scatterhoarded seeds tend to be larger,
 Larger seeds survive better Larger seeds survive better
(Herbs & Grasses in grasslands) (Trees in temperate deciduous forests)

● Japan: Tree seedlings that grew from

● Grassland: Larger seedlings grew from larger seeds were taller. This is because
larger seeds → better chances of of the larger seeds providing larger
survival. energy reserves to boost seedling
growth.

● Seedlings growing from larger seeds

recruited (proportion of the seeds that
were recruited into seedlings) into the
population at a higher rate.
 Adult Survival & Reproductive Allocation
● Where adult survival is lower,
organisms begin reproducing at an
earlier age and invest a greater
proportion of their energy budget into
reproduction
● Where adult survival is higher,
organisms defer reproduction to a
later age and allocate a smaller
proportion of their resources to
reproduction.
Reproductive Effort
● Reproductive effort → allocation of
energy, time, and other resources to the
production and care of offspring
● Involves trade-offs with other needs:
Growth and Maintenance
● This graph generalizes that there is a
difference in the allocation of
reproductive effort in environments that
are unfavorable and favorable.
● Unfavorable environments have
organisms that allocate less of their
resources or energy to reproduction
● Favorable environments have
organisms that allocate more of their
resources or energy to reproduction.
Patterns of tradeoff are similar when looking at
species that belong to different groups or the
same groups.
Life history variation among different
species
● There is a high reproductive effort when
their is a high probability of dying before
reproduction.
● Most vertebrate species begin
reproducing before they reach
maximum body size (e.g. fish &
reptiles) → energy budgets differ before
and after reaching sexual maturity
○ Before sexual maturity: They
allocate their energy budget to
Maintenance & growth
○ After sexual maturity: They
allocate their energy budget to
Maintenance, growth, &
reproduction
● Those that delay reproduction will
grow faster & reach larger size →
larger size results to higher
reproductive rate
● But, delayed reproduction increases
probability of dying before
reproducing (being eaten by a
predator)
○ So, there should be a trade-off
between delayed reproduction
and mortality of organism.
Mortality rates play a pivotal role in
determining age at first reproduction
● Lizards and snakes that have higher
survival mature at a later age.
● When adult survival is high, age at
maturity is also high.
 Higher mortality in adults

● Higher mortality rates tend to be

correlated with higher reproductive
effort expressed in Gonadosomatic
Insex (GSI)
● Fishes with higher mortality rates
allocate a greater proportion of their
energy to reproduction.
● If species has high rates of mortality, not
only this organisms need to reproduce
at a younger age, but they also need to
allocate more of their energy into having
kids.

● Looking at the relationship from the

opposite perspective, fish with higher
mortality rates reach reproductive
maturity at an earlier age.
● When mortality is high, the age at
maturity is low.
○ This indicates that organisms
must reproduce as early as
possible if the environment they
are living in has high mortality
rate for adults.
● Higher adult mortality rate may also 1. Gonadosomatic Index (GSI) per fish
indicate that organisms tend to allocate species → ovary weight divided by the
more energy in reproducing. species body weight; adjusted for the
number of batches of offspring produced
by each species per year.
2. GSI for each of the 28 species included
in the analysis spanned more than a
30-fold difference
3. Species with higher mortality show
higher relative reproductive effort.
Life history variation within species
● Between different populations within
same species.

5 populations of Pumpkinseed Sunfish from

lakes in Ontario, Canada

● The relationship between adult:juvenile

survival and age at maturity was not
statistically significant.
○ It doesn’t matter whether more
juveniles or more adults die
because it will not affect the age
at which the population will
● Adult pumpkinseed survival was reproduce.
highest in Beloporine Lake.
● It is intermediate in Black and Vance
lakes.
● Lowest in lottle round and Warrens
Lakes.

There is a great variation in terms of survival of

adults and juveniles.

● However, it can be indicated that there

is a negative relationship between
adult:juvenile survival ratio and
reproductive effort. Pumpkinseed
sunfish populations with higher ratios of
adult to juvenile survival showed lower
reproductive effort.
○ Lower survival of adults means
that energy must be spent into
● The ratio of adult to juvenile survival reproducing or there must be a
varied widely across the five study higher reproductive effort for
lakes. adults that survive less.
 ● Where adult survival is lower relatiove to ● r selected species tends to have high
juvenile survival, natural selection will growth rates while K selected species
favor allocating greater resources to display traits that are associated to living
reproduction. densities close to carrying capacity.
● r and K selection does not apply to all
species.
Life History Classification ○ There are a multitude of
variations in life histories that
The great diversity of life histories can could not be accounted for simply
be classified on the basis of a few population by this classification.
characteristics.
Examples include fecundity or number *Other ecologist came up with ways to classify life
of offspring, survival, relative offspring history strategies.
size, and age at reproductive maturity.
Plant life history strategies
r and K selection ● J.P. Grime’s classification (1979)
● One of the first ways byu which diversity proposed that the variation in
of life histories was categorized was environmental conditions has led to the
through r and K selection. development of distinctive strategies or
● It was originally proposed by Robert life histories among plants. He used 3
McArthur and E.O. Wilson (1967); variables for this:
expounded on by Eric Pianka (1970, ○ Intensity of disturbance
1972) ■ Disturbance is defined as
● r - intrinsic rate of increase factors that destroy plant
● K - carrying capacity biomass
○ Intensity of stress
■ Stress is defined as
Population r selected (r K selected factors that limit biomass
attribute strategists) (K
strategists) production
○ Intensity of competition
Intrinsic rate High Low
of increase, ● He then classified the plants into 3
rmax types:
Competitive Not strongly Highly ○ Ruderals are dominate under
ability favored favored conditions of high disturbance
and low stress.
Development Rapid Slow ■ Ruderal are annual
Reproduction Early Late grasses or Weeds; Small
timing and rapid lifecycle to
invade new sites; Large
Body size Small Large dispersal area
Reproduction Single. Repeasted, ○ Stress-tolerant species
semelparity iteoparity predominate under conditions of
low intensity of competition,
Offspring Many, small Few, Large
 high competition and high lx, low fecundity, mx, and early
stress maturity, α.
■ Stress-tolerant live in ● Periodic life history
arease with limited ○ The periodic life history combines
resources but as high low juvenile survival, lx, high
ability to adapt fecundity, mx, and late maturity
○ Competitive species such as α.
birch predominate under ● Equilibrium life history
conditions of low disturbance ○ The equilibrium life history
and high intensity of combines high juvenile
competition. survival, lx, low fecundity, mx,
■ Competitive species live and late maturity α.
in stable environment,
slower life cycle, with more
resources for growth

Another way to classify life histories is by using

the different aspects of population dynamics.

Life history types can be defined by

trade-offs in juvenile survival, fecundity and
age at reproductive maturity

● Winemiller & Rose’s classification, 1992

● Animals are categorized into 3:
● Opportunistic life history
○ The opportunistic life history
combines low juvenile survival,
Using that scheme, different groups of animals
can be compared.
Fishes
● Compared to other vertebrate groups,
fgishes include a much greater range of
life history strategies.
Amphibians and reptiles
reproductive effort & Relative offspring size
Birds
Mammals
● Compared to other vertebrates,
mammas show the least variation.
Mammals are between opportunistic
and equilibrium life history.
● Humans, in particular, are in equilibrium
life history strategy. Because we have
high juvenile survival, low feucndity and
late maturity
Life history types based on Lifetime
1. One of the recent attempts to classify
life history strategies was done by Eric
Charnov et al. (2002, 2007)
2. They aimed to classify organisms based
on characteristics that are free of the
influences of size and time.
a. Size and time are responsible for
the many obvious life history
differences among species of
closely related taxa. Ex. small
and large mammals.
3. Facilitate the exploration of life history
variation within and among groups of
closely related taxa.
4. They took some key life hsiotryfeatures
and converted it to dimensionless
numbers. Ex. Relative offspring size and
Relative reproductive effort over life
span.
a. Relatifve Offspring Size (I/m) - ● Bird score higher in dimensionless
offspring mass at independence indicators of life histories, including
from parent/adult mass offspring size relative to adult mass
i. I stands for independence ● Fraction of adult biomass allocated to
because it is the mass of reproduction over the adult life span
the offspring at
independence from the
parent
ii. m is the adult mass at first
reprioduction
b. Relative Reproductive effort
over adult life span (C*E) -
proportion of adult body mass
allocated to reproduction per unit
time * adult life span.
i. C is the proportion of adult
body mass allocated tor
reproduction for unit time
ii. E is the adult life span.

● Birds, mammals. Lizards occupy

well-seprated regions in Charnovs’s life
history plane.
● Lifetime reproductive effort of birds is
high and their relative offspring size is
high as well which is why they re in the
upper right portion fo the plane.
● Meanwhile, lizards are low on both
relative offspring size and relative
reproductive effort, occupying the lower
left poriton fo the plane.
● Mammals tend to be similar in lizards in
terms of relative reproductive effort over
their adult life span. However, they have
a much higher relative offspring size.
 COMPETITION AND COEXISTENCE
1. Types of competition
2. The ecological niche - how it reflects the
environmental requirements of species
3. Mathematical model of competition -
provide theoretical foundation for
studying interspecific competition in
nature
4. Influence of environment on competition
5. Competition and niches - how
competition can have significant
ecological and evolutionary influence on
the niches of species
Types of Competition
Types of interactions between species
Competition
Competition happens when resources are
limited
● Exploitative competition: two
organisms are indirectly influencing
each other by consuming the same
resources (ex: different bird species
consuming fruits of the same trees).
○ They dont directly engage in
fighting.
● Interference competition: involved
organisms directly influencing each
other by preventing others to occupy
a habitat or access resources (ex:
competition between males of deer).
○ Direct engagement with each
other.

Competition interactions
Nature of Interaction Species Species
1 2 Specific ways by which competitors interact
with each other
Mutualism + + 1. Consumption – Utilization of a shared
resource by 2 species
Predation + 0
a. 2 species use the same shared
Predation, Herbivory, + - resources.
Parasitism 2. Preemption – Occupation of a site by
1st organism stops occupation by 2nd
Amensalism - 0
organism
Competition - - a. Usually sessile organisms - not
mobile; no legs for locomotion.
Neutralism 0 0 3. Overgrowth – Where organism covers
another preventing access to a
● In competition, both species involved resource.
are harmed – negatively affected. a. Trees shade other plants; large
canopies competing for sunlight.
Population growth is both limited by 4. Chemical interaction – Release of
density-dependent and density-independent toxin to inhibit or kill competing
factors. One of the density-dependent factors organisms
is competitions. When resources are limited a. Allelopathy in plants - one plant is
and there are multiple organisms involved, capable of releasing chemicals in
competition happens. the environment and other
organisms are negatively affected
(killed or slow down in growth).
 5. Territorial – Behavioral exclusion of 1st
organism by 2nd organism defending
territory
6. Encounter – Non-territorial encounters
cause a negative effect on one or
both species
a. Lion and wild dogs over an
antelope kill.
● Whent he density is very high, tadpoles
suffer, and it had to undergo a logner
amount of time to reach maximum body
size.
○ Aside from that, their maximum
body size is relatively lower the
the tadpoles growing in lower
densities.

Competition can be between organisms Relationship of body growth and density

belonging to different or the same species White Clove
1. Intraspecific – Competition between
members of own species
2. Interspecific – Competition between
individuals of two species (different)

Intraspecific competition affects growth

and development
● Intraspecific competition is usually
density-dependent. It increases
gradually, at first affects growth and
development, then individual survival
and reproduction.
● Intraspecific competition affects the
growth and development of plants and
animals.

Relationship of body growth and density

Tadpole ● As the density of the white clove
increases, the mean weight per plant
decreases.
● Growth is negatively effected by
intraspecific competition greatly shown
in higher densities.

Among plants, there is a phenomenon called

self-thinning.
Self-thinning in Plants
● It can happen in any species in plants.
● There were 4 set-ups that differ in
Individuals within a population somehow
density of the tadpole. Mean body mass
manage to choose among themselves
of the tadpole were compared along
who is the best competitor. Then, those
weeks.
are the ones which eventually survive
● When the density in the set-up is very
and grow larger. Others are left to die 1
low, the tadpoles tend to reach a higher
by 1.
body mass within a short period of time.
 even before reproducing (e.g being
eaten, starvation, etc.).
● Not only do females reproduce at a later
stage, but less of the females also
reproduce with increasing population
density

● Self thinning – progressive decline in

density and increase in biomass of
remaining individuals in a population.
● M. sativa planted at high density
initially.
● M. sativa population at the end of the
experiment consisted of larger plants
growing at lower density.
● As plantings of alfalfa, Medicago sativa,
grew, mortality thinned the stands as
surviving plants reached larger size.

Competition can also reduce reproduction.

Intraspecific competition can reduce

reproduction
● Intraspecific competition can function to
reduce fecundity.

Seals Corn and Shrub

● It shows how population density among
harp seals affect the mean age of
whelping.
● At very high population density, the age
at which the females reproduce is also
higher. This, however, is not good
because when reproduction is delayed,
there is a higher chance of mortality
● Same trend with harp seals: as you the
population density of corns is increased,
the reproduction becomes lesser.
● Yield per plant in corn and number of
seeds produced in shrubs gets lesser
as plant population density increases.
 Ecological Niche

Ecological Niche of a Species

● In ecological sense, Niche summarizes
the environmental factors that
influence the growth, survival, and
reproduction of a species
○ Everything that incfluences an
organisms life belongs to its
niche.
● Approximates when, where, and how a
species “makes its living”
● Niche literally means “a recessed place
in a wall where one could set or display
items”
○ Place dedicated for a specific
thing.
● Some of the earliest concepts of
ecological niche were proposed by:
○ Joseph Grinnell (1917, 1924) –
centered around influences of the
physical environment
○ Charles Elton (1927) – biological
interactions & abiotic factors

It took quite a long time for the concept of

niche to be fully developed. It was in the
context of interspecific competition that the
importance of niche was fully realized.

G.F. Gause’s Competitive Exclusion

Principle (1934)
● His work was focused on interspecific
competition which helped ensure a
prominent place for niche in modern
ecology, particularly the Exclusion
Principle.
● Exclusion Principle – “Two species
with identical niches cannot coexist
indefinitely”
 could be low to high in various
factors (e.g. low temperature,
high salt content) → where one
finds its position along the axis is
its niche.

● Hutchinson classified niche into 2 types:

● When two species of Paramecium (P.
aurelia and P. caudatum) were grown
separately, there is a distinct growth
curve for the 2.
● When they are grown together P. aurelia
outcompetes. P. caudatum which will
eventually disappear from the container.
Both of them have identical niches
(almost the same requirements for
survival) which prevents them from
coexisting.
○ Fundamental niche: range of
conditions and resources a
species can use to survive and
reproduce under no
interference by other species.
○ Realized niche: portion of
fundamental niches that a
species actually exploits as a
result of interactions with other
species (e.g., competition) →
usually becomes smaller.

G. Evelyn Hutchinson’s n-dimensional

hypervolume
● Her work crystallized the niche concept.
● Niche is an n-dimensional
hypervolume, where n equals the
number of environmental factors
important to the survival and
reproduction of a species
● Dimensions can refer to higher
amounts. In this example, there are only
3 dimensions (x,y,z) representing water,
temperature, and light.
○ But, obviously species will require
Niche dimensions
more factors to live. So, niche is
● Rarely do two or more species possess
a hypervolume because it cannot
exactly the same combination of
just be represented by limited
requirement.
number of axes. Other axis can
● Species may overlap on one dimension
be nutrients, space, etc.
of the niche, but not on another.
○ So niche, will be the space
occupied by the organism in this
hypervolume. Along the axis,
 Mathematical Model Of Competition

Interspecific competition can be analyzed

mathematically using the Lokta-Volterra
Model
● Lotka-Volterra model is the most
fundamental equation used to analyze
the competition between species

● Derived from logistic equation

Intrinsic rate of population * N * element

● There can be overlaps in dimension. that defines the carrying capacity
However, organisms cannot occupy the
same niche with all dimensions the ● Add influence of another species (a
same. competition component)

First equation - growth of the first

species
Second equation - groth of the second
species
● Where:

Aside from per capita competitive effect,

these are also referred to as
compettion coefficients

Niche in different dimensions (1,2,3). Involves the

different factors that define the niche of organisms
Lokta-Volterra Model *Neither one of the α is too big or too strong so
that it will drive the growth of the other to 0.

When population of species 1&2 stop

growing

Equate both equations to 0. Growth of either if

● α 2,1 and α 1,2N1 : referred to as effect the two species is 0. → Population stops
of intersecific competition growing
○ In terms of resource use, an
individual of species 2 is equal to
α individuals of species 1
○ α can work as a conversion
factor which can be used when
(Without species 2, it will just be a simple logistic
converting the number of
equation. With increasing species 2, the growth will be
individuals of species 2 to the slower and slower such that there will be no growth
number of species 1. anymore and eventually reach carrying capacity. With
● If there is noo interspecific competition, increasing species 2, the carrying capacity for species 1
then α 2,1 and α 1,2 are 0 and normal will be at lower and lower density → the environment
can support less and less of species 1 as species 2
growth reaches carrying capacity.
increases)
○ Formula will look like a logistic
And
equation and produce a sigmoid
shape curve.
● If there is an effect of Interspecific
competition (density dependent), then
the graph will look much different.
That is when
Predictions of Lokta-Volterra Model
1. L-V model predicts coexistence of 2
species when interspecific
competition is weaker than
intraspecific competition.
a. The competition between varying
species is not big enough for Or rearranging these equations
them to outcompete each other.
2. Otherwise, one species will exclude the
other.
● Once these equations are graphed,
isocline of zero population growth
can be obtained.
Isoclines of Zero Population Growth ● Note: All the arrows going left and right
● These plots are not plots of population will meet at the isocline of 0-population
with respect to time. Time has growth
disappeared in consideration.
● Isoclines of 0-population growth show Species 1
how the environment can be filled up or
in other words, the relative population
sizes of species 1 and 2 that will deplete
the critical resources.
○ For example (species 1), at one
extreme, we have the
environment completely filled up
by species 1 such that species 1
reaches carrying capacity (N1 =
K1)
○ At the other end of the extreme is
when the environment is
saturated by species 2 and
species 1 is completely absent
(0). This occurs when the
population or density of species 2
● This graph indicates when will species 1
is equal to K1/α1,2 (carrying
hit 0-growth point due to the competitive
capacity of species 1 divided by
effect of species 2.
the coefficient of competition).
○ In between, the 2 extremes, the
Species 2
environment is saturated by a
mixture of species 1 and species
2.
○ There is also the isocline
representing the dN1/dt = 0
because it is the isocline of 0
population growth.
■ Below this, there is the
dN1/dt > 0 which is
positive.
■ Above the line dN1/dt < 0
is negative.
○ If the population is below carrying
capacity, the population will keep
on increasing because the
dN1/dt is positive.
○ If somehow, the population
exceeds carrying capacity, the
dn1/dt > 0 and the population will
decrease.
 ● To get the isocline of species 2, use the
same steps for species 1 and replace
some of the variables.
● Interpretation for species 2 will be
similar to that of species 1. However,
the species 2 is on the y-axis.
Therefore, the point at which the
species 2 is 0 would be when the
population of species 1 is equal to K2/α
2,1 .
● The arrows will be going up and down.
Possible outcomes of Interspecific
competition
*upon combination of 2 isoclines
When two species compete, what are possible
outcomes?
1. Species 1 wins, species 2 loses
2. Species 1 loses, species 2 wins
3. Competition can go either way (unstable
equilibrium)
4. Coexistence (stable equilibrium)
Outcome 1: Species 1 inhibits growth of
species 2 and latter goes extinct
Outcome 2: Species 2 inhibits growth of
● Isocline of species 1 occurring above
the isocline of species 2.
○ Species 1 wins; population size
equals K1 or its carrying capacity.
● To understand how this happened, the
trajectories of the arrows - which
represent the direction of the growth of
the 2 species - should be analyzed.
○ Arrows to the left of species 1 are
going to the right.
○ Arrows above species 2 are
going down.
○ Sum of arrows will point towards
southeast. The way that the
trajectory is pointing is towards
the reaching of the carrying
capacity for species 1 → Species
1 will win.
● The overlap is the part where species 2
is declining → population is going down.
At the densities where species 2 is
declining, species 1 will grow. The area
where species 1 are going to the right
indicates the growth in species 1. The
down movement indicates that species
1 will exclude species 2.
○ Species 2 is above its carrying
capacity over the entire range of
possible combinations.
species 1 and latter goes extinct
● Same trajectory analysis can also be
used for species 2.
 ● Isocline for species 2 is above that of
species 1. Species 2 wins; population
size equals K2
● The trajectory of the arrows points
towards the reaching of the carrying
capacity for species 2 → species 2 will
win.
● Species 2 can grow at denisites where
species 1 population is declining. At the
overlap, th species 1 is going to the left
while species 2 is going up.
○ Species 1 is above its carrying
capacity at all possible
combinations → Its growth is
negative.
Outcome 3: Unstable situation, both inhibit
in a density dependent manner. Depending
on initial density, either can make other
extinct
Outcome 4: Each species inhibits its own
population growth more than competitor.
● Coexistence is possible only in the
situations where the isoclines cross.
However, only one of these situations
leads to stable coexistence. For
unstable situations, → Eventually
species 1 or 2 wins.
● In this situation K1 is greater than K2/α2,1
and K2 is greater than K1/α1,2.
● Most population growth trajectories lead
either to the points where N1 is equal to
K1 and N2 = 0. Or, N2 is equal to K2
and N1 = 0
● The populations of species 1 and
species 2 may arrive at the point where
the lines cross, but any environmental
variation that moves the population of
this point eventually leads to the
exclusion of one species by the other.
○ Either species 1 achieves
carrying capacity and excludes
species 2. Or, species 2 achieves
carrying capacity and excludes
species 1.
● If we take a closer look at the trajectory
of the arrows, above both isoclines,
both species are actually declining and
below the isoclines, both species are
increasing.
● However, in the little triangles where one
is increasing and the other is
decreasing,
○ Either species 1 is increasing and
species 2 is decreasing →
species 1 will reach its carrying
capacity or the other way around.
Neither can eliminate competitor
● The only situation where there is there is
coexistence. Species 1 and 2 coexist at
the crossover point of the isoclines.
 ● In this situation, K2/α2,1is greater than K1
and K1/α1,2. Is greater than K2.
● All growth trajectories lead to the point
where the isoclines of the 0 population
growth cross. → Stable equilibrium
Laboratory experiments support the
Lotka-Volterra Equations
● The two examples (Paramecium and
Diatom) shows that environment can
influence the dynamics of competition
between species.
Varying the Food supplies for 2
Paramecium Species
● However, they experimented in
changing the food supplies and with
different food supplies: Coexistence
was achieved.
Diatom experiment
● David Tilman, University of Minnesota
● Growth of Asterionella formosa (Af) and
Synedra ulna (Su) was compared.
● Both diatoms compete for silica for the
formation of cell walls.
● When grown apart from each other, they
show distinct curves from each other.
○ Amount of silica decreases as the
population of the diatom species
increases. → Silica is depleted.
● When the 2 species are grown together
and there is an insufficient silica,
○ Su drove Af to extinction.
● However, if adequate amount of silica
was prepared, Su and Af were shown to
coexist.

● Russian biologist G.F. Gause (known for

competitive exclusion principle)
● Competition between two species
● The two species of paramecium where
having a certain growth level when
grown separately.
● On the other hand, when grown together
or in mixed population, P. caudatum
actually shows a decrease in its groth
because P. aurelia has a high growth
rate and can tolerate a higher population
density
● Of the two Paramecium species
○ One with higher rate of growth:
Extinction of slower grower
 Influence Of Environment On Competition

Temporal variation in environment

influences competition interactions
● As environmental conditions vary, the
competition advantages change
● No one species can reach sufficient
density to displace its competitors;
● Thus leading to coexistence.

Shift in dominant grass species caused by

moisture

● At the beginning of the experiment,

Urochloa was dominant and
Heteropogon was inferior.
● Over the years, the amount of
rainfallthat fell during the rainy season
increased and that changed the
competition relationship between
Urochloa and Heteropogon and
Heteropogon became more dominant.
● Depending on how the future
environment will change will dictate how
the relationship will change.
Relative competition abilities change along
environmental gradients

● Effect of interspecific competition across

an environmental gradient
● Note changes in response when in
mixture
Silybum marianum
(milk thistle)

● Six species of thistle in mixed pastures

in southeast Australia were grown in
pots of sand with different concentration
of nutrients.
● They were either grown in monocultures
or as mixtures.
○ In monocultures or when thistle
are grown on their ow. → They
almost have an overlapping
growth pattern wherein the higher
the relative nutrient concentration
is, the higher is the total biomass
per plant.
○ In mixtures, there are competitive
abilities of each of the species.
■ Carduun nutans (red
graph) having the least
competitive ability.
■ At highest concentration of
nutrients, Carduus
pycnocephalus has the
highest competitive ability
or the one that can grow
the most biomass.
■ At low nutrient
concentrations, a different
species, Carthamus
lanatus is the one showing
the highest biomass per
plant.
○ The dominant species is different
with respect to the nutrient
 concentration at low and high
nutrients.

Altitudinal zonation of Chipmunks in Sierra

Nevada, Calif.

● Heller, 1971
● Four chipmunk species were observed.
The range that the species of
chipmunks roughly correspond to the 4
different altitudinal zones in this study
site.
● Upper range of minimus determined by
aggressive interactions with dominant
amoenus, which is at the limit of its
fundamental niche where the Piñon pine
belt grades into the arid, hot sagebrush
desert.
● Among the 4 chipmunks, Alpinus and
Amoenus are said to be the ones that
are aggressive.
○ Aggressive amoenus determines
lower range limit of speciosus,
(speciousus cannot go further
down because of the aggressive
amoenus) while at the upper
edge of its range, speciosus is
limited by the aggressively
dominant alpinus.
● Aggression is selected for in alpinus and
amoenus because of a seasonal,
potentially limiting food supply which is
economically defendable.
● Aggression not selected for in minimus
because it is not metabolically
feasible to engage in aggressive
interactions in the hot sagebrush
desert.
● Aggression not selected for in speciosus
because of predator pressure
(speciosus is extremely quiet and
secretive) and seasonal abundance of
food in its habitat.
 Competition & Niche
How competition shapes niches of organisms
Ecological & Evolutionary Influences of
Competition
1. Niche differentiation
2. Habitat shift
3. Character displacement
4. Competitive exclusion
5. Competitive release
Niche differentiation
Niche differentiation
● Process by which natural selection
drives competing species into
different patterns of resource use or
different niches.
● Separation of niches via resource
partitioning among related species.
Terns and the sizes of fish they consume
● Terns are closely related to each other
but they evolved to speciealize on
specific fish body lengths.
● There are still overlaps in terms of the
sizes of the fish that they consume, but
they exhibit different patterns regarding
the fish body length that they will
specialize in → So, that they can avoid
competing with each other.
● Niche separation via resource
partitioning
Desert Plants
● Desert plants have evolved different
root systems (lengths and widths) in
order to specialize in certain levels of
the soil → so that they will not compete
that much for the resources in the soil
such as water and nutrients.
 Species of Small cats that co-occur in Anolis Lizards
Isarael.

● They have evolved different sizes in

terms of diameter of their canine teeth
from smallest silvestris to the largest
caracal.
● Anolis lizards in the Caribbean all feed
● The size (diameter) of canine teeth for
on similar prey and are similar in size,
small cat that co-occur in Israel. Size is
but they make use of different foraging
correlated with size of prey selected by
perches → Means to avoid competition.
different species.
● Different species of Anolis occupy
● It appeared that these cats have
different parts of their habitats (trees and
evolved they have come to specialize on
shrubs)
the different species of prey and these
○ Highest – occupied by Anolis
specialization came with concurrent
ricordii
differentiation in their morphology as
○ Lowest – occupied by A
well → Morphological differentiation
etheridgei
○ This is actually an example of
● It can be expected that when other
“ghost of competition past” – this
species are removed from this scenario,
is the current citation now, but
A. ricordii can forage and perch on the
this was probably a result of all
lower branches of the trees.
the competition that happened in
○ The only reason why it is
the past. Species have evolved in
specializing on the highest tree
order to avoid competition.
is because of the presence of
other species
○ So, in other words, species can
expand its habitat when other
Habitat shift
species are absent.
● Change from one species population to
another in the habitats occupied.
● Use of habitat often changes
depending on presence or absence of
Character displacement
closely related species.
● Differences between similar species are
● Habitat tend to expand when other
accentuated in sympatry
species absent.
○ Two related species or
populations are considered
 sympatric when they exist in the
same geographic area and thus
frequently encounter one another.
● Species A and Species B live in
sympatry in the area where they
overlap.
○ In these areas, their characters
are displaced, so the population
of specia A and species B will
show a range in terms of the
character means.
● Moreover, they also live in areas where
they are completely separate from each
other.
○ In these areas where they don’t
overlap, theta have this particular
character mean which could be
any phenotypic characteristic (e.g
size of the leaf, size of the teeth,
etc. )
● Those individuals that live away from
the overlap will have a different average
character compared to those organisms
that live int he overlapping areas.
● Character displacement happens
because species wants to avoid
competition.
Geospiza (Galapagos Finches)
● These 2 species of Darwin’s finches
have populations in either Los
Hermanos or Daphne.
○ Daphne island only has G. fortis
while the Los Hermnaos island
only has G. fuliginosa.
● When they live allopatrically or separate
from each other, their beak depth tend
to overlap or the range of beak depeths
are not much different from each other.
● They have sympatric populations in
Santa Maria, and San Cristobal.
● When the two are living sympatrically, G.
fortis beak depth mean is shifted to the
right (towards the larger beak depth)
while the beak depth mean for G.
fuliginosa is shifted to the left (towards
smaller beak depth) to avoid
competition.
Morphological traits and food selection of
species shift depending on presence or
absence of closely related species.
Criteria for Character Displacement – (Mark
Taper & Ted Case, 1992)
*6 criteria to be fulfilled in order to consider
character displacement among organisms
1. Significant difference between sympatric
and allopatric
a. Morphological differences
between a pair of sympatric
species must be greater than the
differences between allopatric
populations of the same species.
2. Genetic basis of difference
a. Observed differences between
sympatric and allopatric
populations should have a
genetic basis → Differences that
are observed are not only due to
phenotypic plasticity.
3. Not due to founder effect
a. Differences between sympatric
and allopatric populations must
have evolved in place and must
not be due to the sympatric and
allopatric populations having
been derived from different
founder populations already
differing in the character
understudy.
4. Character variation affects resource use
5. Competition for resource under question
a. There must be a demonstrated
competition for the resource
under question and competitions
must be directly correlated with
similarity in the character.
6. Not due to differences in resources
available
a. Differences in the character
cannot be explained by
differences in resources available
to sympatric and allopatric
populations.
Competitive exclusion
1. Upper intertidal zone
a. Balanus dessicates in this zone,
allowing Chthamalus to thrive
2. Middle intertidal zone
a. Where it can survive, Balanus
outcompetes Chthamalus for
space.
3. Lower intertidal zone
a. Balanus is subject to predation to
the lower tidal zone.
Competitive exclusion principle
● Complete competitors can not coexist (2
species with identical niches). One
species must go extinct
● Complete competitions: two species
that live in the same place and
possess exactly the same ecological
requirements.
● Assumptions:
○ Exactly the same resource
requirement (no more, no less).
○ Environmental conditions remain
constant
● Obviously, most of the time, these
assumptions are hard to fulfill, which is
why most of the time species can
coexist
Competitive release

● Expansion in the food preferences

and foraging range of an animal that
follows a reduction in the intensity of
competition by other species.
● Densities of organisms often increase
when densities of competing species
are reduced.
Baleen whales
● There were about 1 million whales a
century ago and each can eat Antartic
Krill with amount of 4% of their body
weight.
● Now, they are less than 200,000 whales
and other krill-dependent predators
such as seals and penguins have been
found greatly increased in abundance.