1. Chapter 53
Population Ecology

2. Overview: Counting Sheep
A small population of Soay sheep were introduced to Hirta Island in 1932
They provide an ideal opportunity to study changes in population size on an isolated island with abundant food and no predators; dramatic fluctuation

3. Population ecology is the study of populations in relation to environment, including environmental influences on density and distribution, age structure, and population size

4. Concept 53.1: Dynamic biological processes influence population density, dispersion, and demographics
A population is a group of individuals of a single species living in the same general area
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

5. 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, or the mark-recapture method

6. mixing > recapture (total; n, marked; x) x/n = m/N
Fig. 53-2
APPLICATION
capture > mark (m)> release > wait for
mixing > recapture (total; n, marked; x)
x/n = m/N
N = estimated population size
Figure 53.2 Determining population size using the mark-recapture method

7. 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
Emigration is the movement of individuals out of a population

8. Births and immigration add individuals to a population.
Fig. 53-3
Births
Deaths
Births and immigration
add individuals to
a population.
Deaths and emigration
remove individuals
from a population.
Figure 53.3 Population dynamics
Immigration
Emigration

9. Patterns of Dispersion
Environmental and social factors influence spacing of individuals in a population
(clumped, uniform, random)

10. In a clumped dispersion, individuals aggregate in patches
A clumped dispersion may be influenced by resource availability and behavior

11. A uniform dispersion is one in which individuals are evenly distributed
It may be influenced by social interactions such as territoriality

12. In a random dispersion, the position of each individual is independent of other individuals
It occurs in the absence of strong attractions or repulsions

13. Demographics
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

14. 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

15. Table 53-1

16. Survivorship Curves
A survivorship curve is a graphic way of representing the data in a life table
The survivorship curve for Belding’s ground squirrels shows a relatively constant death rate

17. Number of survivors (log scale)
Fig. 53-5
1,000
100
Number of survivors (log scale)
Females 10
Males
Figure 53.5 Survivorship curves for male and female Belding’s ground squirrels 1 2 4 6 8 10
Age (years)

18. Survivorship curves can be classified into three general types:
Type I: low death rates during early and middle life, then an increase among older age groups
Type II: the death rate is constant over the organism’s life span
Type III: high death rates for the young, then a slower death rate for survivors

19. Number of survivors (log scale)
Fig. 53-6
1,000 I
100 II
Number of survivors (log scale) 10
Figure 53.6 Idealized survivorship curves: Types I, II, and III
III 1 50
100
Percentage of maximum life span

20. 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

21. Table 53-2

22. Concept 53.2: 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

23. Evolution and Life History Diversity
Life histories are very diverse
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

24. “Trade-offs” and Life Histories
Organisms have finite resources, which may lead to trade-offs between survival and reproduction (cf. chicks transfer among nests)

25. RESULTS Male Female 100 80 60 40 20 Reduced brood size 3~4 Normal 5~6
Fig. 53-8
Male
Female
100
RESULTS 80 60 40 20
Reduced
brood size
3~4
Normal
5~6
Enlarged
7~8
Parents surviving the following winter (%)
Figure 53.8 How does caring for offspring affect parental survival in kestrels?

26. Some plants produce a large number of small seeds, ensuring that at least some of them will grow and eventually reproduce

27. Other types of plants produce a moderate number of large seeds that provide a large store of energy that will help seedlings become established

28. Concept 53.3: The exponential model describes population growth in an idealized, unlimited environment
It is useful to study population growth in an idealized situation
Idealized situations help us understand the capacity of species to increase and the conditions that may facilitate this growth

29. Per Capita Rate of Increase
If immigration and emigration are ignored, a population’s growth rate (per capita increase) equals birth rate minus death rate

30. Zero population growth occurs when the birth rate equals the death rate (r=0, when r=b-d)
Most ecologists use differential calculus to express population growth as growth rate at a particular instant in time: N t 
rN = bN – dN = (b-d)N
where N = population size, t = time, and r = per capita rate of increase = birth – death

31. Exponential Growth
Exponential population growth is population increase under idealized conditions
Under these conditions, the rate of reproduction (r) is at its maximum, called the intrinsic rate of increase (rmax)
Equation of exponential population growth: dN dt 
rmaxN
Exponential population growth results in a J-shaped curve

32. 2,000 = 1.0N 1,500 = 0.5N Population size (N) 1,000 500 5 10 15
Fig
2,000 dN =
1.0N dt
1,500 dN =
0.5N dt
Population size (N)
1,000
500
Figure Population growth predicted by the exponential model 5 10 15
Number of generations

33. The J-shaped curve of exponential growth characterizes some rebounding populations
8,000
6,000
4,000
2,000
1920
1940
1960
1980
Year
Elephant population
1900

34. Concept 53.4: The logistic model describes how a population grows more slowly as it nears its carrying capacity
Exponential growth cannot be sustained for long in any population
A more realistic population model limits growth by incorporating carrying capacity
Carrying capacity (K) is the maximum population size the environment can support

35. The Logistic Growth Model
In the logistic population growth model, the per capita rate of increase declines as carrying capacity is reached
We construct the logistic model by starting with the exponential model and adding an expression that reduces per capita rate of increase as N approaches K dN dt 
(K  N) K
rmax N

36. Table 53-3
Table 53.3

37. The logistic model of population growth produces a sigmoid (S-shaped) curve

38. Fig
2,000
1,500
1,000
500 5 10 15
Number of generations
Population size (N)
Exponential
growth
1.0N = dN dt
K = 1,500
Logistic growth
1,500 – N
Figure Population growth predicted by the logistic model

39. The Logistic Model and Real Populations
The growth of laboratory populations of paramecia fits an S-shaped curve
These organisms are grown in a constant environments of limited resources but lacking predators and competitors

40. Number of Paramecium/mL
Fig a
1,000
800
Number of Paramecium/mL
600
400
200
Figure 53.13a How well do these populations fit the logistic growth model? 5 10 15
Time (days)
(a) A Paramecium population in the lab

41. Some populations overshoot K before settling down to a relatively stable density (females may use their energy reserves to continue reproducing for a short time)

42. (b) A Daphnia population in the lab
Fig b
180
150
120
Number of Daphnia/50 mL 90 60 30
Figure 53.13b How well do these populations fit the logistic growth model? 20 40 60 80
100
120
140
160
Time (days)
(b) A Daphnia population in the lab

43. 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 (positive interaction at low density while negative interaction at high population density)

44. The Logistic Model and Life Histories
Life history traits favored by natural selection may vary with population density and environmental conditions
K-selection, or density-dependent selection, selects for life history traits that are sensitive to population density
r-selection, or density-independent selection, selects for life history traits that maximize reproduction

45. There are two general questions about regulation of population growth:
Concept 53.5: 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?

46. 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

47. Birth or death rate per capita Birth or death rate per capita
Fig
Density-dependent
birth rate
Density-dependent
birth rate
Density-
independent
death rate
Density-
dependent
death rate
Birth or death rate
per capita
Equilibrium
density
Equilibrium
density
Population density
Population density
(a) Both birth rate and death rate vary.
(b) Birth rate varies; death rate is constant.
Density-dependent
death rate
Density-
independent
birth rate
Figure Determining equilibrium for population density
Birth or death rate
per capita
Equilibrium
density
Population density
(c) Death rate varies; birth rate is constant.

48. Density-Dependent Population Regulation
Density-dependent birth and death rates are an example of negative feedback that regulates population growth
They are affected by many factors, such as competition for resources, territoriality, disease, predation, toxic wastes, and intrinsic factors

49. 1. Competition for Resources
In crowded populations, increasing population density intensifies competition for resources and results in a lower birth rate
Population size
100 80 60 40 20
200
400
500
600
300
Percentage of juveniles producing lambs

50. 2. Territoriality
In many vertebrates and some invertebrates, competition for territory may limit density
Cheetahs are highly territorial, using chemical communication to warn other cheetahs of their boundaries

51. Oceanic birds exhibit territoriality in nesting behavior (nests a peck apart)

52. 3. Disease
Population density can influence the health and survival of organisms
In dense populations, pathogens can spread more rapidly (TB spread in densely populated cities)

53. 4. Predation
As a prey population builds up, predators may feed preferentially on that species (not energy-efficient at low prey densities)

54. 5. Toxic Wastes
Accumulation of toxic wastes can contribute to density-dependent regulation of population size (ethanol production by yeast; inhibition at a high concentration of ethanol)

55. 6. Intrinsic Factors
For some populations, intrinsic (physiological) factors appear to regulate population size (hormonal change and/or depressed immune system due to stress induced by high population density; even food and shelter are plenty)

56. Population Dynamics
The study of population dynamics focuses on the complex interactions between biotic and abiotic factors that cause variation in population size

57. Stability and Fluctuation
Long-term population studies have challenged the hypothesis that populations of large mammals are relatively stable over time
2,100
1,900
1,700
1,500
1,300
1,100
900
700
500
1955
1965
1975
1985
1995
2005
Year
Number of sheep

58. Changes in predation pressure can drive population fluctuations
Weather can affect population size over time
Wolves
Moose
2,500
2,000
1,500
1,000
500
Number of moose
Number of wolves 50 40 30 20 10
1955
1965
1975
1985
1995
2005
Year

59. Population Cycles: Scientific Inquiry
Some populations undergo regular boom-and-bust cycles
Lynx populations follow the 10 year boom-and-bust cycle of hare populations
Three hypotheses have been proposed to explain the hare’s 10-year interval

60. Number of hares (thousands) Number of lynx (thousands)
Fig
Snowshoe hare
160
120 9
Figure Population cycles in the snowshoe hare and lynx
Lynx
Number of hares
(thousands)
Number of lynx
(thousands) 80 6 40 3
1850
1875
1900
1925
Year

61. Hypothesis 1: The hare’s population cycle follows a cycle of winter food supply
If this hypothesis is correct, then the cycles should stop if the food supply is increased
Additional food was provided experimentally to a hare population, and the whole population increased in size but continued to cycle
No hares appeared to have died of starvation

62. Hypothesis 2: The hare’s population cycle is driven by pressure from other predators
In a study conducted by field ecologists, 90% of the hares were killed by predators
These data support this second hypothesis

63. Hypothesis 3: The hare’s population cycle is linked to sunspot cycles
Sunspot activity affects light quality, which in turn affects the quality of the hares’ food (low sunspot activity > less O3 > more UV > plant produce more UV-blocking chemicals > fewer chemicals that deter herbivores > good quality food for hare)
There is good correlation between sunspot activity and hare population size

64. The results of all these experiments suggest that both predation and sunspot activity regulate hare numbers and that food availability plays a less important role

65. Immigration, Emigration, and Metapopulations
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

66. ˚ Aland Islands EUROPE Occupied patch 5 km Unoccupied patch Fig. 53-21
Figure The Glanville fritillary: a metapopulation
Occupied patch
5 km
Unoccupied patch

67. Concept 53.6: The human population is no longer growing exponentially but is still increasing rapidly
No population can grow indefinitely, and humans are no exception
The human population increased relatively slowly until about 1650 and then began to grow exponentially

68. 7 6 5 4 3 2 1 Human population (billions) The Plague 8000 B.C.E. 4000
Fig 7 6 5 4
Human population (billions) 3 2
The Plague
Figure Human population growth (data as of 2006) 1
8000
B.C.E.
4000
B.C.E.
3000
B.C.E.
2000
B.C.E.
1000
B.C.E.
1000
C.E.
2000
C.E.

69. Though the global population is still growing, the rate of growth began to slow during the 1960s
2005
Projected
data
Annual percent increase
Year
1950
1975
2000
2025
2050
2.2
2.0
1.8
1.6
1.4
1.2
1.0
0.8
0.6
0.4
0.2
Famine in China (60 million death)

70. Regional Patterns of Population Change
To maintain population stability, a regional human population can exist in one of two configurations:
Zero population growth = High birth rate – High death rate
Zero population growth = Low birth rate – Low death rate
The demographic transition is the move from the first state toward the second state

71. Birth or death rate per 1,000 people
Fig 50 40 30
Birth or death rate per 1,000 people 20
Figure Demographic transition in Sweden and Mexico, 1750–2025 (data as of 2005) 10
Sweden
Mexico
Birth rate
Birth rate
Death rate
Death rate
1750
1800
1850
1900
1950
2000
2050
Year

72. The demographic transition is associated with an increase in the quality of health care and improved access to education, especially for women
Most of the current global population growth is concentrated in developing countries (death rates decline rapidly; birth rates decline in a variable manner)

73. Age Structure
One important demographic factor in present and future growth trends is a country’s age structure
Age structure is the relative number of individuals at each age

74. Rapid growth Slow growth No growth Afghanistan United States Italy
Fig
Rapid growth
Slow growth
No growth
Afghanistan
United States
Italy
Male
Female
Age
Male
Female
Age
Male
Female
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
Figure Age-structure pyramids for the human population of three countries (data as of 2005)
15–19
15–19
10–14
10–14
5–9
5–9
0–4
0–4
10  8 6 4 2 2 4 6 8
10  8 6 4 2 2 4 6 8 8 6 4 2 2 4 6 8
Percent of population
Percent of population
Percent of population

75. Infant Mortality and Life Expectancy
Infant mortality and life expectancy at birth vary greatly among developed and developing countries but do not capture the wide range of the human condition

76. Life expectancy (years) Infant mortality (deaths per 1,000 births)
Fig 60 80 50 60 40
Life expectancy (years)
Infant mortality (deaths per 1,000 births) 30 40 20 20
Figure Infant mortality and life expectancy at birth in industrialized and less industrialized countries (data as of 2005) 10
Indus-
trialized
countries
Less indus-
trialized
countries
Indus-
trialized
countries
Less indus-
trialized
countries

77. Global Carrying Capacity
How many humans can the biosphere support?
The carrying capacity of Earth for humans is uncertain
The average estimate is 10–15 billion

78. 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

79. 13.4 9.8 5.8 Not analyzed Log (g carbon/year) Fig. 53-27
Figure The amount of photosynthetic products that humans use around the world
5.8
Not analyzed

80. Our carrying capacity could potentially be limited by food, space, nonrenewable resources, or buildup of wastes

81. You should now be able to:
Define and distinguish between the following sets of terms: density and dispersion; clumped dispersion, uniform dispersion, and random dispersion; life table and reproductive table; Type I, Type II, and Type III survivorship curves; semelparity and iteroparity; r-selected populations and K-selected populations
Explain how ecologists may estimate the density of a species

82. Explain how limited resources and trade-offs may affect life histories
Compare the exponential and logistic models of population growth
Explain how density-dependent and density-independent factors may affect population growth
Explain how biotic and abiotic factors may work together to control a population’s growth

83. Describe the problems associated with estimating Earth’s carrying capacity for the human species
Define the demographic transition