Dynamic biological processes influence
population density, dispersion, and
• Population ecology is the study of populations in
relation to environment, including environmental
influences on density and distribution, age
structure, and population size
Density and Dispersion
• Density is the number of individuals per unit area
or volume
• Dispersion is the pattern of spacing among
individuals within the boundaries of the population
demographics
• A population is a group of individuals of a single
species living in the same general area
• Populations are described by their boundaries and
size
Density: A Dynamic Perspective
• In most cases, it is impractical or impossible to
count all individuals in a population
• Sampling techniques can be used to estimate
densities and total population sizes
• Population size can be estimated by either
extrapolation from small samples, an index of
population size (e.g., number of nests), or the
mark-recapture method
 APPLICATION

• Mark-recapture method
– Scientists capture, tag, and release a random
sample of individuals (s) in a population
– Marked individuals are given time to mix back into
the population
– Scientists capture a second sample of individuals
(n), and note how many of them are marked (x)
– Population size (N) is estimated by
sn
N=
x
Hector’s dolphins

Births Deaths
• Density is the result of an interplay between
processes that add individuals to a population and
those that remove individuals
• Immigration is the influx of new individuals from
other areas Births and immigration Deaths and emigration
add individuals to remove individuals
• Emigration is the movement of individuals out of a population. from a population.

a population

Immigration Emigration
 (a) Clumped

Patterns of Dispersion
• Environmental and social factors influence spacing
of individuals in a population
(b) Uniform
• In a clumped dispersion, individuals aggregate in
patches
• A clumped dispersion may be influenced by
resource availability and behavior
(c) Random

(a) Clumped

• A uniform dispersion is one in which individuals

are evenly distributed
• It may be influenced by social interactions such as
territoriality, the defense of a bounded space
against other individuals
(b) Uniform
• In a random dispersion, the position of each
individual is independent of other individuals
• It occurs in the absence of strong attractions or
repulsions

Demographics
(c) Random • Demography is the study of the vital statistics of a
population and how they change over time
• Death rates and birth rates are of particular
interest to demographers
Life Tables
• A life table is an age-specific summary of the
survival pattern of a population
• It is best made by following the fate of a cohort, a
group of individuals of the same age
• The life table of Belding’s ground squirrels reveals
many things about this population
– For example, it provides data on the proportions
of males and females alive at each age
Survivorship Curves 1,000

Number of survivors (log scale)

• A survivorship curve is a graphic way of
representing the data in a life table 100
• The survivorship curve for Belding’s ground
squirrels shows a relatively constant death rate
Females
10
Males

1
0 2 4 6 8 10
Age (years)

Figure 53.6

Number of survivors (log scale)

1,000
• Survivorship curves can be classified into three I
general types
– Type I: low death rates during early and middle 100
life, then an increase in death rates among older II
age groups
– Type II: the death rate is constant over the 10
organism’s life span
III
– Type III: high death rates for the young, then a
1
slower death rate for survivors 0 50 100
• Many species are intermediate to these curves Percentage of maximum life span
Reproductive Rates
• For species with sexual reproduction,
demographers often concentrate on females in a
population
• A reproductive table, or fertility schedule, is an
age-specific summary of the reproductive rates in
a population
• It describes reproductive patterns of a population

The exponential model describes population Per Capita Rate of Increase

growth in an idealized, unlimited
Change in Immigrants Emigrants
environment population = Births + entering − Deaths − leaving
size population population
• It is useful to study population growth in an
idealized situation • If immigration and emigration are ignored, a
• Idealized situations help us understand the population’s growth rate (per capita increase)
capacity of species to increase and the conditions equals birth rate minus death rate
that may facilitate this growth

• The population growth rate can be expressed
mathematically using differential calculus,
∆N = B − D
∆t
Where ∆N is change in population size, ∆t is time
interval, B is number of births, and D is number of
deaths
Exponential Growth
• Exponential population growth is population
increase under idealized conditions
• Under these conditions, the rate of increase is at
its maximum, denoted as rmax
• The equation of exponential population growth is
dN =
dt rmaxN

2,000
dN
= 1.0N
dt
• Exponential population growth results in a J-

Population size (N)

1,500
shaped curve dN
= 0.5N
dt

1,000

500

0 5 10 15
Number of generations
 8,000

• The J-shaped curve of exponential growth

Elephant population
characterizes some rebounding populations 6,000

– For example, the elephant population in Kruger

National Park, South Africa, grew exponentially 4,000
after hunting was banned

2,000

0
1900 1910 1920 1930 1940 1950 1960 1970
Year

The logistic model describes how a The Logistic Growth Model

population grows more slowly as it nears • In the logistic population growth model, the per
its carrying capacity capita rate of increase declines as carrying
capacity is reached
• Exponential growth cannot be sustained for long in
• The logistic model starts with the exponential
any population
model and adds an expression that reduces per
• A more realistic population model limits growth by capita rate of increase as N approaches K
incorporating carrying capacity
• Carrying capacity (K) is the maximum population dN (K − N)
= rmax N
size the environment can support dt K
• Carrying capacity varies with the abundance of
limiting resources
 • The logistic model of population growth produces
a sigmoid (S-shaped) curve

Exponential

2,000
growth The Logistic Model and Real Populations
dN
= 1.0N
dt
• The growth of laboratory populations of paramecia
Population size (N)

1,500 fits an S-shaped curve

K = 1,500 Logistic growth • These organisms are grown in a constant
dN 1,500 – N
dt
= 1.0N( 1,500
) environment lacking predators and competitors
1,000

500 Population growth

begins slowing here.

0
0 5 10 15
Number of generations
 Number of Paramecium/mL
Number of Paramecium/mL

Number of Daphnia/50 mL
1,000 180 1,000

800 150 800

120
600 600
90
400 400
60
200 30 200
0 0 0
0 5 10 15 0 20 40 60 80 100 120 140 160 0 5 10 15
Time (days) Time (days) Time (days)
(a) A Paramecium population in (b) A Daphnia population in the lab (a) A Paramecium population in
the lab
the lab

• Some populations overshoot K before settling

Number of Daphnia/50 mL
180
down to a relatively stable density 150
120
90
60
30
0
0 20 40 60 80 100 120 140 160
Time (days)

(b) A Daphnia population in the lab

 • Some populations fluctuate greatly and make it
difficult to define K
• Some populations show an Allee effect, in which
individuals have a more difficult time surviving or
reproducing if the population size is too small
Life history traits are products of natural
selection
• An organism’s life history comprises the traits
that affect its schedule of reproduction and
survival
– The age at which reproduction begins
– How often the organism reproduces
– How many offspring are produced during each
reproductive cycle
• Life history traits are evolutionary outcomes
reflected in the development, physiology, and
behavior of an organism
• The logistic model fits few real populations but is
useful for estimating possible growth
• Conservation biologists can use the model to
estimate the critical size below which populations
may become extinct
Evolution and Life History Diversity
• Species that exhibit semelparity, or big-bang
reproduction, reproduce once and die
• Species that exhibit iteroparity, or repeated
reproduction, produce offspring repeatedly
• Highly variable or unpredictable environments
likely favor big-bang reproduction, while
dependable environments may favor repeated
reproduction
 “Trade-offs” and Life Histories
• Organisms have finite resources, which may lead
to trade-offs between survival and reproduction
– For example, there is a trade-off between survival
and paternal care in European kestrels

RESULTS

100
Male • Some plants produce a large number of small
Parents surviving the following

Female
seeds, ensuring that at least some of them will
80
grow and eventually reproduce
winter (%)

60

40

20

0
Reduced Normal Enlarged
brood size brood size brood size
 • Other types of plants produce a moderate number
of large seeds that provide a large store of energy
that will help seedlings become established

(a) Dandelion

• K-selection, or density-dependent selection, • The concepts of K-selection and r-selection are

selects for life history traits that are sensitive to oversimplifications but have stimulated alternative
population density hypotheses of life history evolution
• r-selection, or density-independent selection,
selects for life history traits that maximize
reproduction
Population Characteristics Population ecology is the study of interactions
within populations (i.e., intraspecific interactions)
Recall that populations are groups of interacting
conspecifics (e.g., inter-mating)
We can characterize individual populations in terms
of there…
Size (average vs. variation)
Density (& impacts on size; density dependence)
Patterns of Dispersion
Demographics (age structure, sex ratios)
Rates of growth (or decline)
Limits on population growth
Factors That Regulate Populations
• Resource availability affects life history
adaptations
• When resources are limited, the cost of
reproduction is high
– Selection will favor individuals that can
compete and utilize resources efficiently
– Can lower reproductive rates
– K-selected populations: adapted to thrive
when population is near its carrying
capacity

Factors That Regulate Populations

• Populations far below carrying capacity,

resources abundant
– Costs of reproduction are low
– r-selected populations: selection favors
individuals with the highest reproductive
rates
• Most natural populations show life
history adaptations that exist along a
continuum of r- and K-selected traits
60
 Populations
Factors That Regulate Populations
• Populations
– Groups of individuals of the same species • Density-dependent
in one place
– Factors that affect the population and
• 3 characteristics of population ecology depend on population size
– Population range, area throughout which a • Density-independent
population occurs
– Other factors, such as natural disasters,
– Pattern of spacing of individuals affect populations regardless of size
– How population changes in size through
time

Factors That Regulate Populations

Factors That Regulate Populations
At high
populations, • Density-independent effects
locusts have – Rate of growth of a population at any
different instant is limited by something unrelated to
hormonal and the size of the population
physical – External environment aspects: cold
winters, droughts, storms, volcanic
characteristics Positive feedback: Allee effect
eruptions
and take off as Growth rates increase with – Populations display erratic growth patterns
a swarm population size because of these events
Many factors that regulate population
growth are density dependent
• There are two general questions about regulation
of population growth
– What environmental factors stop a population
from growing indefinitely?
– Why do some populations show radical
fluctuations in size over time, while others remain
stable?
Population Change and Population Density
• In density-independent populations, birth rate
and death rate do not change with population
density.
• In density-dependent populations, birth rates fall
and death rates rise with population density.

% of young sheep producing lambs

100
Mechanisms of Density-Dependent
Population Regulation
80

• Density-dependent birth and death rates are an

example of negative feedback that regulates 60
population growth
• Density-dependent birth and death rates are 40
affected by many factors, such as competition for
resources, territoriality, disease, predation, toxic 20
wastes, and intrinsic factors
0
200 300 400 500 600
Population size
Competition for Resources
• In crowded populations, increasing population
density intensifies competition for resources and
results in a lower birth rate

Toxic Wastes Predation

• Accumulation of toxic wastes can contribute to • As a prey population builds up, predators may
density-dependent regulation of population size feed preferentially on that species

© 2011 Pearson Education, Inc.

Intrinsic Factors
• For some populations, intrinsic (physiological)
factors appear to regulate population size

Territoriality
• In many vertebrates and some invertebrates,
competition for territory may limit density
Disease
• Population density can influence the health and
survival of organisms
• In dense populations, pathogens can spread more
rapidly
 Population Dynamics
• The study of population dynamics focuses on
the complex interactions between biotic and
abiotic factors that cause variation in population
size

Stability and Fluctuation

50 2,500
Wolves Moose
• Long-term population studies have challenged the

Number of wolves

Number of moose
40 2,000
hypothesis that populations of large mammals are
relatively stable over time 30 1,500
• Both weather and predator population can affect
population size over time 20 1,000
– For example, moose on Isle Royale collapsed
during a harsh winter, and when wolf numbers 10 500
peaked
0 0
1955 1965 1975 1985 1995 2005
Year
 Population ecology
Immigration, Emigration, and
Metapopulations • Distribution
– Metapopulations = network of populations that
• A group of Dictyostelium amoebas can emigrate interact occasionally by exchanging individuals
and forage better than individual amoebas • Occur when habitat is
patchily distributed and
separated by unsuitable
habitat
• Characterized by
suitable habitat patches
going extinct and getting
recolonized over time
• glanville fritillary
butterfly in Finland

• Metapopulations are groups of populations linked
by immigration and emigration
• High levels of immigration combined with higher
survival can result in greater stability in
populations
Populations
• Metapopulations
– Occur in areas in which suitable habitat is
patchily distributed and is separated by
intervening stretches of unsuitable habitat
– Dispersal
• Interaction may not be symmetrical
• Populations increase and send out many
dispersers
• Small populations have few dispersers
• Individual populations may become extinct
• Population bottlenecks may occur
Populations • Glanville fritillary
• Source–sink metapopulations butterfly
– Some areas are suitable for long-term (Melitaea cinxia)
habitat, others are not • None of the
– Populations in better areas (source) bolster populations is large
enough to survive
the population in poorer areas (sink)
for long on its own,
• Metapopulations can have two implications but continual
for the range of a species immigration of
– Continuous colonization of empty patches individuals from
prevents long-term extinction other populations
allows some
– In source–sink metapopulations, the
populations to
species occupies a larger area than it survive
otherwise might
90

˚
Aland
Islands The human population is no longer growing
EUROPE exponentially but is still increasing rapidly

• No population can grow indefinitely, and humans

are no exception

Occupied patch
5 km Unoccupied patch
The Global Human Population 7

Human population (billions)

6
• The human population increased relatively slowly
until about 1650 and then began to grow 5
exponentially
4

2
The Plague
1

0
8000 4000 3000 2000 1000 0 1000 2000
BCE BCE BCE BCE BCE CE CE

2.2

2.0

1.8
• The global population is more than 6.8 billion

Annual percent increase

1.6
people
1.4
• Though the global population is still growing, the 2009
1.2
rate of growth began to slow during the 1960s Projected
1.0 data
0.8

0.6

0.4

0.2

0
1950 1975 2000 2025 2050
Year
Regional Patterns of Population Change
• To maintain population stability, a regional human • The demographic transition is associated with an
population can exist in one of two configurations increase in the quality of health care and improved
– Zero population growth = access to education, especially for women
High birth rate – High death rate • Most of the current global population growth is
– Zero population growth = concentrated in developing countries
Low birth rate – Low death rate
• The demographic transition is the move from the
first state to the second state

Age Structure
Rapid growth Slow growth No growth
Afghanistan United States Italy
• One important demographic factor in present and Male Female Age Male Female Age Male Female
85+ 85+
future growth trends is a country’s age structure 80–84
75–79
80–84
75–79
70–74 70–74
• Age structure is the relative number of individuals 65–69 65–69
60–64 60–64
at each age 55–59
50–54
55–59
50–54
45–49 45–49
40–44 40–44
35–39 35–39
30–34 30–34
25–29 25–29
20–24 20–24
15–19 15–19
10–14 10–14
5–9 5–9
0–4 0–4
10 8 6 4 2 0 2 4 6 8 10 8 6 4 2 0 2 4 6 8 8 6 4 2 0 2 4 6 8
Percent of population Percent of population Percent of population
 Rapid growth Slow growth
Afghanistan United States
Male Female Age Male Female Age
85+ 85+
80–84 80–84
75–79 75–79
70–74 70–74
65–69 65–69
60–64 60–64
55–59 55–59
50–54 50–54
45–49 45–49
40–44 40–44
35–39 35–39
30–34 30–34
25–29 25–29
20–24 20–24
15–19 15–19
10–14 10–14
5–9 5–9
0–4 0–4
10 8 6 4 2 0 2 4 6 8 10 8 6 4 2 0 2 4 6 8
Percent of population Percent of population

No growth
Italy
Age Male Female
85+
80–84 • Age structure diagrams can predict a population’s
75–79
70–74 growth trends
65–69
60–64 • They can illuminate social conditions and help us
55–59
50–54 plan for the future
45–49
40–44
35–39
30–34
25–29
20–24
15–19
10–14
5–9
0–4
8 6 4 2 0 2 4 6 8
Percent of population
 Figure 53.25

Infant mortality (deaths per 1,000 births)

60 80
Infant Mortality and Life Expectancy
50

Life expectancy (years)

• Infant mortality and life expectancy at birth vary 60
greatly among developed and developing 40
countries but do not capture the wide range of the
human condition 30 40

20
20
10

0 0
Indus- Less indus- Indus- Less indus-
trialized trialized trialized trialized
countries countries countries countries

Global Carrying Capacity Estimates of Carrying Capacity

• How many humans can the biosphere support? • The carrying capacity of Earth for humans is
• Population ecologists predict a global population uncertain
of 7.8−10.8 billion people in 2050 • The average estimate is 10–15 billion
Limits on Human Population Size
• The ecological footprint concept summarizes the
aggregate land and water area needed to sustain
the people of a nation
• It is one measure of how close we are to the
carrying capacity of Earth
• Countries vary greatly in footprint size and
available ecological capacity Gigajoules
> 300
150–300
50–150
10–50
< 10

Human Population Growth

• Our carrying capacity could potentially be limited Ecological
by food, space, nonrenewable resources, or Footprint: amount
buildup of wastes of productive land
• Unlike other organisms, we can regulate our required to support
population growth through social changes an individual at the
standard of living
of a particular
population through
the course of
his/her life
112