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Population Ecology

Population defined
Population is a group of individuals of a single species living in the same general area.

Population Ecology
Study of populations in relation to their environment, including environmental influences on density and distribution, age structure, and population size

Population Ecology
Concepts: The physical environment limits the geographic distribution. On small scales, individuals within populations are distributed in patterns that may be random, regular, or clumped; on larger scales, individuals within a population are clumped. Many populations are subdivided into subpopulation called metapopulation. Population density declines with increasing organism size. Commonness and rarity of species are influenced by population size, geographic range, and habitat tolerance.

Population Characteristics
1. Natality total number of individuals added to the population through reproduction over a particular period of time.
Biotic communities i.e. Plants, fungi, bacteria sexual and asexual Animals usually sexual reproduction Human population natality is described in terms of birth rate number of individuals born per 1000 individuals per year

2. Mortality number of deaths in a population over a particular period of time. 3. Population Growth rate of increase subtracted by rate of decline; (immigration + birth rate) (death rate + emigration)



Births and immigration add individuals to a population.

Deaths and emigration remove individuals from a population.



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

Measuring density of populations is a difficult task.

We can count individuals; we can estimate population numbers.

Fig. 52.1
Copyright 2002 Pearson Education, Inc., publishing as Benjamin Cummings

It is the result of an interplay between processes that add individuals to a population (birth and immigration) and those that remove individuals (death and emigration)

Parameters that effect size or density of a population: Immigration


Population (N)



Figure 1. The size of a population is determined by a balance between births, immigration, deaths and emigration

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
Population bottlenecks occur when a populations size is reduced for at least one generation.

Patterns of Distribution and Dispersal Environmental and social factors influence the spacing of individuals in a population. Overall, dispersion depends on resource distribution.

Uniform Dispersion
Regular pattern 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. There is an antagonistic interaction between individuals.

Clumped Dispersion
Individuals in areas of high local abundance are separated by areas of low abundance Uneven distribution of resources Individuals are attracted to a common resource.

Random Dispersion
An individual has an equal probability of occurring anywhere in an area. The position of each individual is independent of other individuals. There is a neutral interaction between individuals and it occurs in the absence of strong attractions or repulsions.

Age Structure
The proportion of individuals in each age class of a population. Iteroparous species individuals that give birth to few offsprings at several reproductive periods; exhibit age structure; ex. mammals Semelparous species reproducing only once in a life time; no age structure; ex. mayflies, cicadas

Age Structure
Age structure has a critical influence on a populations growth rate Classification of ages based on reproductive stages:
1. Pre- reproductive stage 0 to 14 years 2. Reproductive stage 15 to 44 years 3. Post- reproductive stage 45 years and older

Life tables
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.

Survivorship Curves
A graphic way of representing the pattern of survival of individuals in a population from birth to the maximum age attained by each individuals. This is a plot of the number of individuals in a cohort still alive at each age.

Figure 53.5


Number of survivors (log scale)





4 6 Age (years)


Survivorship Curves
Three general types: Type I - low death rates during early and middle life and an increase in death rates among older age groups Type II - a constant death rate over the organisms life span Type III - high death rates for the young and a lower death rate for survivors

Figure 53.6


Number of survivors (log scale)


1 0 50 Percentage of maximum life span 100

Population Growth
A function of reproduction and immigration.
High biotic potential and high rate of immigration Biotic potential is the maximum reproductive potential of an organism.

The population growth rate expressed mathematically as:



where N is the change in population size, t is the time interval, B is the number of births, and D is the number of deaths

Births and deaths can be expressed as the average number of births and deaths per individual during the specified time interval B D bN mN

where b is the annual per capita birth rate, m (for mortality) is the per capita death rate, and N is population size

The population growth equation can be revised

The per capita rate of increase (r) is given by

Zero population growth (ZPG) occurs when the birth rate equals the death rate (r 0)

Population Growth
Principle: In the presence of abundant resources, populations can grow at geometric or exponential rates.

Exponential Growth
Population increase under idealized conditions Under these conditions, the rate of increase is at its maximum, denoted as rmax Equation: dN dt rmaxN Results in a J-shaped curve Rarely seen in nature.

Exponential growth of rabbits

dN dt


The J-shaped curve of exponential growth also characterizes some rebounding populations
For example, the elephant population in Kruger National Park, South Africa, grew exponentially after hunting was banned


Elephant population




0 1900




1940 Year




Kruger National Park, South Africa

Logistic Population Growth

Describes how a population grows more slowly as it nears its carrying capacity.
Carrying capacity (K) is the maximum population size the environment can support; varies with the abundance of limiting resources.

The per capita rate of increase declines as carrying capacity is reached.

Logistic Population Growth

The logistic model starts with the exponential model and adds an expression that reduces per capita rate of increase as N approaches K. dN dt rmax N



Produces a sigmoid (S-shaped) curve

Figure 53.10

Number of Paramecium/mL

Number of Daphnia/50 mL 0 5 10 Time (days) 15

1,000 800 600 400 200 0



60 30 0 0 20 40 60 80 100 120 140 160 Time (days)

(a) A Paramecium population in the lab

(b) A Daphnia population in the lab

Figure 52.11 Population growth predicted by the logistic model

Imposition of limits dN/dt = r N (K-N)/K

New or Changing Environment (no competition / limits)

Environmental Resistance
Sum of the total environmental limiting factor (both biotic and abiotic) that prevent the biotic potential (rmax) of a population from being realized

Some of the assumptions built into the logistic model do not apply to all populations.
It is a model which provides a basis from which we can compare real populations.

Severe Environmental Impact

Other populations have regular boom-andbust cycles. There are populations that fluctuate greatly.

Boom and then Bust

Water flee (Daphnia magna) is adapted to exploit new environment: high growth rate, resistant eggs produced before crash.

Boom and then really Bust

Reindeer introduced to Pribilov island. Initial exponential growth, crash, complete extinction.

Boom and sort of Bust

Predators were removed from Kaibab plateau. Mule deer population size increased from 4,000 to hundred thousand, then dropped and stabilzed at 10,000.

Boom but not much Bust

Sheep introduced to Tasmania: rapid initial growth, overshoot, drop, fluctuation around carrying capacity.

Boom & Bust & Boom & Bust & Boom & Bust

The familiar 10-11 year hare-lynx cycle might not be true. Biased data.

Factors influencing population growth:

1. Sex ratio and age distribution
Sex ratio relative number of males and females Number of females are very important since they determine the number of offsprings produced in a population. Polygamous number of males less important to population growth Monogamous both sexes important Age distribution number of individuals in the prereproductive period is a determinant factor in population growth rate.

Factors influencing population growth: 2. Carrying capacity

Population size that can be maintained in an area over time without harming the habitat. Combination of factors that sets the carrying capacity of an area is called environmental resistance. An environmental resistance can be any limiting factor (raw materials, energy supply, accumulation of waste products, interactions among organisms)

Factors influencing population growth: 3. Density-independent factor

population-limiting factor whose intensity in unrelated to population density severe storms and flooding sudden unpredictable severe cold spells earthquakes and volcanoes catastrophic meteorite impacts

Factors influencing population growth:

4. Density-dependent factors
population-limiting factor whose intensity is linked to population density. limiting resources (e.g., food & shelter) production of toxic wastes infectious diseases predation stress emigration As resources become limiting with increasing population size, biotic interaction intensifies. This decrease fitness of individuals decreasing growth

Biotic interactions whose effects decrease fitness of individuals:

1. 2. 3. 4.

Competition = intra / interspecific Amensalism = 0 Parasitism = + Predation = + -

*Fitness ability to survive and reproduce; relative number of offspring that survive

The environmental factor that influence the growth, survival, and reproduction of a species. Function / role of the organism Interspecific competition

Competitive Exclusion Principle
G. F. Gause (1934) Two species with identical niches cannot coexist indefinitely. The more effective competitor for limited resources will have higher fitness and will eventually exclude all individuals of the second species.

Fundamental niche
the physical conditions under which a species might live, in the absence of interactions with other species.

Realized niche
actual niche of a species whose distribution is limited by biotic interaction (competition, predation, disease, and parasitism) may be much smaller than the fundamental niche

Competition and Niches

Competition can have a significant ecological role and evolutionary influences on the niches of species.
Competition restricts the species to their realized niche but they still retain their capacity to inhabit the fuller range of environment, fundamental niche.

Biotic interactions whose effects increase fitness of individuals: 1. Mutualism - + + intimate relationship; one cannot do without the other; protocooperation can live with or without the relationship; ex. Plants and ants 2. Commensalism one organism benefits without affecting the other

Life history
How natural selection and other evolutionary forces shape organisms to optimize their survival and reproduction in the face of ecological challenges posed by the environment. Consist of the adaptations of an organism that influences aspects of its biology such as the number of offspring it produces, its survival, and its size and age at reproductive maturity.

Life history
Concepts: Because all organisms have access to limited energy and other resources, there is a trade-off between the number and size of offspring.
Darter species that produce larger eggs produce few eggs.

Organisms reproduce at an earlier stage when adult survival is lower; where adult survival is higher, organisms defer reproduction to a later stage.
Ex. The survival of adult snakes and lizards increases as their age at maturity also increases.

The great diversity of life histories may be classified on the basis of a few population characteristics.
r-selection and K-selection

Offspring number versus Size

Life history
Adult Survival and Reproductive Allocation Long-lived species delay reproduction
- Advantage: juveniles gain experience before high cost of reproduction

Short-lived species reproduce early

- Time is important; delay may mean no offspring

Classification of life history patterns

r selection
Refers to per capita rate of increase, r Species often colonizing new or disturbed habitats (pioneer species)

K selection
Refers to the carrying capacity, K Prominent in situations where species populations are near carrying capacity

Characteristics favored by r versus K selection

Population attribute Intrinsic rate of increase, rmax
Competitive ability Development Reproduction Body size

r selection High
Not strongly favored Rapid Early Small

K selection Low
Highly favored Slow Late Large


Single, semelparity
Many, small

Repeated, iteroparity
Few, large

Life history
r and K selection are end points in a continuum correlated with attributes of the environment and of populations.

Life history
r selection characteristics of variable or unpredictable environment.
Type III survivorship

K selection fairly constant or predictable environment

Type I survivorship

Evolution and Natural Selection

Evolution change over time; the study of interplay between heredity and environment.
Change in genetic composition of a population over periods of many generations. Genetic changes lead to changes in appearance, functioning or behavior over generations

Evolution Theory
Charles Darwin
In 1831 Darwin joined the H.M.S. Beagle as the naturalist for a circumnavigation of the world; the voyage lasted five years. It was his observations from that trip that lead to his proposal of evolution by natural selection. Galapagos Island Published the book Origin of Species (1859)

Alfred Wallace
South East Asia

Darwins Finches
Genetic studies show all arise from a single ancestral species.

Natural Selection
Natural selection process where there is differential reproduction and survival of individuals carrying alternative inherited traits
Results in differential representation of genotypes in the future generation Genotype: genetic constitution of an organism

Charles Darwin found out how selection leads to adaptive evolution:


1. Organisms beget like organisms. 2. There are chance variations between individuals in a species. Some variations are heritable. 3. More offspring are produced each generation than can be supported by the environment. 4. Some individuals are better suited to their environment and reproduce more effectively.

Industrial Melanism in Peppered Moth

Industrial melanism - adaptive melanism caused by anthropogenic alteration of the natural environment in terms of industrial pollution.

Peppered Moths

Types of Selection
1. Directional selection drives a feature in one direction. 2. Stabilizing selection favors intermediate traits; preserving the status quo 3. Disruptive selection traits diverge in two or more directions;

Directional Selection
Occurs where one extreme phenotype has an advantage over all other phenotypes. Population's trait distribution shifts toward the other extreme.

Stabilizing Selection
Acts against extreme phenotypes

Disruptive Selection
Favors two or more extreme phenotypes over the average phenotype in a population. Result is a bimodal, or two-peaked, curve in which the two extremes of the curve create their own smaller curves

Genetic variation and Natural Selection

Genetic variation - range (variance) of phenotypes; different chromosomal arrangements (cytogenetics); DNA sequence differences among individuals. Genetic variation within a population is absolutely necessary for natural selection to occur.
If all individuals are identical within a population then their fitness will all be the same. Same fitness Natural selection will not occur

Sources of Genetic Variation

Mutation: inheritable changes in a gene or a chromosome
Point mutation Chromosome mutation

deletion, duplication, inversion, translocation Genetic recombination

Sexual reproduction Two individuals produce haploid gametes (egg or sperm) that combine to form a diploid cell or zygote. Reassortment of genes provided by two parents in the offspring Increases dramatically the variation within a population by creating new combinations of existing genes.

Asexual reproduction: less variation (only mutation)

Evolution is a change in gene frequencies

Evolution is a change of gene frequencies within a population (or species) over time.
Gene frequency: allele frequency; the frequency of occurrence of an allele in relation to that of other alleles of the same gene in a population Hardy-Weinberg Principle: in a population mating at random in the absence of evolutionary forces, allele frequencies will remain constant

p = frequency of one allele (A) q = frequency of the alternative allele (a) p2 = frequency of genotype A q2 = frequency of genotype a 2pq = frequency of individual Aa

Hardy Weinberg Principle

Gene frequencies will remain the same in successive generations of a sexually reproducing population if the following five conditions hold:
1. Random mating
2. No mutations

3. Large population size

4. No immigration

5. No selection

Genetic drift
Changes in the gene frequencies in a small population due to chance or random events. Reduces genetic variation in a population over time by increasing the frequency of some alleles and reducing or eliminating the frequency of others.
One allele can become common in a population in the expense of the alternative allele

Usually caused by bottleneck events and founder effect.

Bottleneck event
Severe reduction in a population size Northern elephant seals
reduced genetic variation probably because of a population bottleneck humans inflicted on them in the 1890s. hunting reduced their population size to as few as 20 individuals at the end of the 19th century. Their population has since rebounded to over 30,000 - but their genes still carry the marks of this bottleneck: they have much less genetic variation than a population of southern elephant seals that was not so intensely hunted.

Founder effects
Occurs when a small number of individuals, representing only a small fraction of the total genetic variation in a species, starts a new population. Small population size means that the colony may have:
reduced genetic variation from the original population a non-random sample of the genes in the original population

Afrikaner population of Dutch settlers in South Africa

descended mainly from a few colonists today, the Afrikaner population has an unusually high frequency of the gene that causes Huntingtons disease, because those original Dutch colonists just happened to carry that gene with unusually high frequency

Selective pressures influence adaptation

Related species in different environments
Experience different pressures Evolve different traits

Convergent evolution unrelated species may evolve similar traits.

Because they live in similar environments

The process of generating new species from a single species. Concept of species
Morphological species concept
A species is defined as a morphologically consistent group of organisms than can be distinguished from all other species
Can fail. So called cryptic species

Biological species concept

A group of populations whose individuals can interbreed and produce fertile offspring and cannot interbreed with other species Reproductive isolation
Still fails. If you cannot tell the individuals apart morphologically, how can you tell if they are interbreeding or not Also, some species can interbreed and produce viable offspring Bontebok and Blesbok in South Africa

Genetic species concept

A group of populations whose individuals have a distinct genetic makeup and who do not interbreed with others groups of populations
Bontebok and Blesbok are genetically distinct as well as being morphologically different.

Mechanisms of Speciation
Allopatric speciation: geographic speciation; species formation due to physical separation of populations; allopatric species occupy area separated by time and space; probably most vertebrates.
Vicariance: separation of an individual taxon or biota due to the formation of a physical barrier to gene flow or dispersal

Sympatric speciation: species form from populations that become reproductively isolated within the same area; sympatric species occupy the same place at the same time; plants and insects.

Mechanisms of Speciation

Genetic isolation mechanisms (reproductive barriers)

Pre-mating mechanisms
- Factors

which prevent individuals from mating

Post-mating mechanisms
- Genomic

incompatibility, hybrid inviability or sterility

Pre-mating mechanisms
1. 2. Geographic isolation: Species occur in different areas, and are often separated by barriers. Temporal isolation: Individuals do not mate because they are reproductively active at different times. This may be different times of the day or different seasons. The species mating periods may not match up. Individuals do not encounter one another during either their mating periods, or at all. Ecological isolation: Individuals only mate in their preferred habitat. They do not encounter individuals of other species with different ecological preferences. Behavioral isolation: Individuals of different species may meet, but one does not recognize any sexual cues that may be given. An individual chooses a member of its own species in most cases. Mechanical isolation: Copulation may be attempted but transfer of sperm does not take place. The individuals may be incompatible due to size or morphology. Gametic incompatibility: Sperm transfer takes place, but the egg is not fertilized.



5. 6.

Post-mating mechanisms
1. Zygotic mortality: The egg is fertilized, but the zygote does not develop. 2. Hybrid inviability: Hybrid embryo forms, but is not viable. 3. Hybrid sterility: Hybrid is viable, but the resulting adult is sterile. 4. Hybrid breakdown: First generation (F1) hybrids are viable and fertile, but further hybrid generations (F2 and backcrosses) are inviable or sterile.

Allopatric speciation
1. 2. Geographically isolated The separated populations diverge (through changes in mating tactics or use of their habitat) Reproductively isolated (such that they cannot interbreed and exchange genes)


An ancestral fish population was split into two by the formation of the Isthmus of Panama about 3.5 millions years ago. Since that time, different genetic changes have occurred in the two populations because of their geographic isolation. These changes eventually lead to the formation of different species. The porkfish (Anisotremus virginicus) is found in the Carribean Sea and the Panamic prokfish (Anisotremus taeniatus) is found in the Pacific Ocean.

Speciation via geographic isolation and divergence

Allopatric speciation
Ring species - population of a single species encircling an area of unsuitable habitat. As a result, the species becomes geographically distributed in a circular, or ring, pattern over a large geographic area.

200 years ago, the ancestors of apple maggot flies laid their eggs only on hawthorns, which are native to America. But today, these flies lay eggs on hawthorns and domestic apples that were introduced by immigrants and bred there. Females generally choose to lay their eggs on the type of fruit they grew up in, and males tend to look for mates on the type of fruit they grew up in. So hawthorn flies generally end up mating with other hawthorn flies and apple flies generally end up mating with other apple flies. This means that gene flow between parts of the population that mate on different types of fruit is reduced.