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Populations
42
Chapter 42 Populations
Key Concepts
42.1 Populations Are Patchy in Space and
Dynamic over Time
42.2 Births Increase and Deaths Decrease
Population Size
42.3 Life Histories Determine Population Growth
Rates
Chapter 42 Populations
Key Concepts
42.4 Populations Grow Multiplicatively, but the
Multiplier Can Change
42.5 Immigration and Emigration Affect
Population Dynamics
42.6 Ecology Provides Tools for Conserving and
Managing Populations
Chapter 42 Opening Question
How does understanding the population ecology of disease vectors help us combat infectious diseases?
Concept 42.1 Populations Are Patchy in Space and
Dynamic over Time
Populations: groups of individuals of the same
species
Humans have long been interested in
understanding species abundance:
• To increase populations of species that
provide resources and food
• To decrease abundance of crop pests,
pathogens, etc.
Concept 42.1 Populations Are Patchy in Space and
Dynamic over Time
Population density—number of individuals per
unit of area or volume
Population size—total number of individuals in
a population
Counting all individuals is usually not feasible;
ecologists often measure density, then multiply
by the area occupied by the population to get
population size.
Concept 42.1 Populations Are Patchy in Space and
Dynamic over Time
Abundance varies on several spatial scales.
Geographic range—region in which a species
is found
Within the range, species may be restricted to
specific environments or habitats.
Habitat patches are “islands” of suitable habitat
separated by areas of unsuitable habitat.
Figure 42.1 Species Are Patchily Distributed on Several Spatial Scales
Concept 42.1 Populations Are Patchy in Space and
Dynamic over Time
Population densities are dynamic—they change
over time.
Density of one species population may be
related to density of other species populations.
Figure 42.2 Population Densities Are Dynamic
Concept 42.2 Births Increase and Deaths Decrease
Population Size
Change in population size depends on the
number of births and deaths over a given
length of time.
“Birth–death” or BD model of population
change:
DBNN tt 1
Concept 42.2 Births Increase and Deaths Decrease
Population Size
Population growth rate (change in size over one
time interval):
Nt1 Nt N (Nt Nt) BD BD
DBDB
tt
DB
T
N
1)1(
Concept 42.2 Births Increase and Deaths Decrease
Population Size
Change in population size can be measured
only for very small populations that can be
counted, such as zoo animals.
To estimate growth rates, ecologists keep track
of a sample of individuals over time.
Concept 42.2 Births Increase and Deaths Decrease
Population Size
Per capita birth rate (b)—number of offspring
an average individual produces
Per capita death rate (d)—average individual’s
chance of dying
Per capita growth rate (r) = (b – d) = average
individual’s contribution to total population
growth rate
rNT
N
Concept 42.2 Births Increase and Deaths Decrease
Population Size
If b > d, then r > 0, and the population grows.
If b < d, then r < 0, and the population shrinks.
If b = d, then r = 0, and population size does not
change.
Concept 42.3 Life Histories Determine Population Growth Rates
Demography: study of processes influencing
birth, death, and population growth rates
Life history: timing of key events such as
growth and development, reproduction, and
death during an average individual’s life
• Example: Life cycle of the black-legged tick
Figure 42.3 Life History of the Black-Legged Tick
Concept 42.3 Life Histories Determine Population Growth Rates
A life history shows the ages at which
individuals make life cycle transitions and how
many individuals do so successfully:
• Survivorship—fraction of individuals that
survive from birth to different life stages or
ages
• Fecundity—average number of offspring
each individual produces at different life
stages or ages
Table 42.1
Concept 42.3 Life Histories Determine Population Growth Rates
Survivorship can also be expressed as
mortality: the fraction of individuals that do not
survive from birth to a given stage or age.
Mortality = 1 – survivorship
Concept 42.3 Life Histories Determine Population Growth Rates
Survivorship and fecundity affect r. The higher
the fecundity rate and survivorship, the higher r
will be.
If reproduction shifts to earlier ages, r will
increase as well.
Concept 42.3 Life Histories Determine Population Growth Rates
Life histories vary among species: how many
and what types of developmental stages, age
of first reproduction, frequency of reproduction,
how many offspring they produce, and how
long they live.
Life histories can vary within a species. For
example, different human populations have
different life expectancies and age of sexual
maturity.
Concept 42.3 Life Histories Determine Population Growth Rates
Individual organisms require resources
(materials and energy) and physical conditions
they can tolerate.
The rate at which an organism can acquire a
resource increases with the availability of the
resource.
• Examples: Photosynthetic rate increases
with sunlight intensity; an animal’s rate of
food intake increases with the density of
food
Figure 42.4 Resource Acquisition Increases with Resource Availability—Up to a Point
Concept 42.3 Life Histories Determine Population Growth Rates
Principle of allocation
Once an organism has acquired a unit of some
resource, it can be used for only one function
at a time, such as maintenance, growth,
defense, or reproduction.
In stressful conditions, more resources go to
maintaining homeostasis.
Once an organism has more resources than it
needs for maintenance, it can allocate the
excess to other functions.
Figure 42.5 The Principle of Allocation
Concept 42.3 Life Histories Determine Population Growth Rates
In general, as average individuals in a
population acquire more resources, the
average fecundity, survivorship, and per capita
growth rate increase.
Concept 42.3 Life Histories Determine Population Growth Rates
Life-history tradeoffs—negative relationships
among growth, reproduction, and survival
• Example: A species that invests heavily in
growth early in life cannot simultaneously
invest heavily in defense.
Environment is also a factor: if high mortality
rates are likely, it makes sense to invest in
early reproduction.
Concept 42.3 Life Histories Determine Population Growth Rates
Species’ distributions reflect the effects of
environment on per capita growth rates.
A study of temperature change in a lizard’s
environment, combined with knowledge of its
physiology and behavior, led to conclusions
about how climate change may affect
survivorship, fecundity, and distribution of
these lizards.
Figure 42.6 Climate Warming Stresses Spiny Lizards (Part 1)
Figure 42.6 Climate Warming Stresses Spiny Lizards (Part 2)
Figure 42.6 Climate Warming Stresses Spiny Lizards (Part 3)
Concept 42.3 Life Histories Determine Population Growth Rates
Laboratory experiments have also shown the
links between environmental conditions, life
histories, and species distributions.
• Example: Quantifying life history traits of
two species of grain beetles in different
temperature and humidity conditions
explained distributions of these species in
Australia.
Figure 42.7 Environmental Conditions Affect Per Capita Growth Rates and Species Distributions
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Population growth is multiplicative—an ever-
larger number of individuals is added in each
successive time period.
In additive growth, a constant number (rather
than a constant multiple) is added in each time
period.
In-Text Art, Chapter 42, p. 873 (2)
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Charles Darwin was aware of the power of
multiplicative growth:
“As more individuals are produced than can
possibly survive, there must in every case be a
struggle for existence.”
This ecological struggle for existence, fueled by
multiplicative growth, drives natural selection
and adaptation.
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Multiplicative growth with a constant r has a
constant doubling time.
The time it takes a population to double in size
can be calculated if r is known.
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Populations do not grow multiplicatively for very
long. Growth slows and reaches a more or less
steady size:
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
r decreases as the population becomes more
crowded; r is density dependent.
As the population grows and becomes more
crowded, birth rates tend to decrease and
death rates tend to increase.
When r = 0, the population size stops
changing—it reaches an equilibrium size
called carrying capacity, or K.
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
K can be thought of as the number of individuals
that a given environment can support
indefinitely.
When population density reaches K, an average
individual has just the amount of resources it
needs to exactly replace itself.
When density <K, an average individual can
more than replace itself; when density >K, the
average individual has fewer resources than it
needs to replace itself.
Figure 42.8 Per Capita Growth Rate Decreases with Population Density
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Spatial variation in environmental factors can
result in variation of carrying capacity.
Temporal variation in environmental conditions
may cause the population to fluctuate above
and below the current carrying capacity.
• Example: the rodents and ticks in Millbrook,
New York
Figure 42.2 Population Densities Are Dynamic
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Environmental changes affected fecundity of the
Galápagos cactus ground finches:
• When females were 7 and 8 years old, they
produced no surviving young, and
survivorship dropped.
• Low food availability during these years
resulted from a severe drought in 1985.
• When the females were 5, a wet year
produced abundant food and high fecundity.
Table 42.1
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
The human population is unique. It has grown at
an ever-faster per capita rate, as indicated by
steadily decreasing doubling times.
Technological advances have raised carrying
capacity by increasing food production and
improving health.
Figure 42.9 Human Population Growth
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
In 1798 Thomas Malthus pointed out that the
human population was growing multiplicatively,
but food supply was growing additively, and
predicted that food shortages would limit
human population growth.
His essay provided Charles Darwin with a
critical insight for the mechanism of natural
selection.
Malthus could not have predicted the effects of
technology such as medical advances and the
Green Revolution.
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
Many believe that the human population has
now overshot its carrying capacity for two
reasons:
• Technological advances and agriculture
have depended on fossil fuels—a finite
resource.
• Climate change and ecosystem degradation
have been a consequence of 20th century
population expansion.
Concept 42.4 Populations Grow Multiplicatively, but the Multiplier
Can Change
If the human population has indeed exceeded
carrying capacity, ultimately it will decrease.
We can bring this about voluntarily if we
continue to reduce per capita birth rate.
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
Many species occupy habitat patches separated
from other patches by unsuitable
environments.
Each patch is occupied by a subpopulation,
the set of subpopulations in a region is a
metapopulation.
Individuals may move in or out of
subpopulations.
Figure 42.10 A Metapopulation Has Many Subpopulations
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
The BIDE model of population growth adds the
number of immigrants (I) and emigrants (E) to
the BD growth model.
EDIBNN tt 1
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
In the BD model, populations are considered
closed systems—no immigration or
emigration.
In the BIDE model, subpopulations are
considered open systems—individuals can
move among them.
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
Small subpopulations in habitat patches are
vulnerable to environmental disturbances and
chance events and may go extinct.
If dispersal is possible, individuals from other
subpopulations can recolonize the patch and
“rescue” the subpopulation from extinction.
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
Immigrants also contribute to genetic diversity
within subpopulations.
This gene flow combats the genetic drift that can
occur in a small population that reduces a
species’ evolutionary potential.
Concept 42.5 Immigration and Emigration Affect
Population Dynamics
In the metapopulation of Edith’s checkerspot
butterfly, all but the largest subpopulation went
extinct during a severe drought between 1975
and 1977.
In 1986, nine habitat patches were recolonized
from the Morgan Hill subpopulation.
Patches closest to Morgan Hill were most likely
to be recolonized because adult butterflies do
not fly very far.
Figure 42.1 Species Are Patchily Distributed on Several Spatial Scales
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
Understanding life history strategies can be
useful in managing other species.
Conserving endangered species
• Larvae of the endangered Edith’s
checkerspot butterfly feed on two plant
species found only on serpentine soils.
• The two plant species are being suppressed
by invasive non-native grasses. Grazing by
cattle can control the invasive grasses.
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
Fisheries
• Black rockfish grow throughout their life.
• Older, larger females produce more eggs,
and the eggs have larger oil droplets, which
give the larvae a head start on growth.
• Because fishermen prefer to catch big fish,
intense fishing reduced the average age of
female rockfish from 9.5 to 6.5 years.
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
• These younger females were smaller,
produced fewer eggs, and the larvae did not
survive as well.
• Population density rapidly declined.
• Management may require no-fishing zones
where some females can mature and
reproduce.
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
Reducing disease risk
• The black-legged tick’s life history indicates
that success of larvae in getting a blood
meal has greatest impact on the abundance
of nymphs.
• Thus, controlling the abundance of rodents
that are hosts for the larvae is more
effective in reducing tick populations than
controlling the abundance of deer, the hosts
for adults.
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
Conservation plans begin with inventories of
habitat and potential risks to the habitat.
Largest patches can potentially have the largest
populations and genetic diversity and are given
priority.
Quality of the patches is evaluated; ways to
restore or maintain quality are developed.
Ability of the organism to disperse between
patches is evaluated.
Figure 42.11 Habitat Corridors Can “Rescue” Subpopulations from Extinction (Part 1)
Figure 42.11 Habitat Corridors Can “Rescue” Subpopulations from Extinction (Part 2)
Figure 42.11 Habitat Corridors Can “Rescue” Subpopulations from Extinction (Part 3)
Concept 42.6 Ecology Provides Tools for Conserving and
Managing Populations
For some species, a continuous corridor of
habitat is needed to connect subpopulations
and allow dispersal.
Dispersal corridors can be created by
maintaining vegetation along roadsides, fence
lines, or streams, or building bridges or
underpasses that allow individuals to avoid
roads or other barriers.
Figure 42.12 A Corridor for Large Mammals
Answer to Opening Question
By understanding the factors that control
abundance and distribution of pathogens and
their vectors, we can devise ways to control
their abundance or avoid contact.
Black-legged ticks are vectors for the bacterium
that causes Lyme disease.
For these ticks, abundance of hosts for larvae
(rodents) determines tick abundance.
Answer to Opening Question
Rodent abundance depends on acorn
availability.
Acorn production can be used to predict areas
that are likely to become infested with ticks,
and measures can be taken to minimize
human contact.