BIO1243 section 1: Evolution and Adaptation

Last updated 25/07/2023

Dr. Sanet Hugo
Office: Natural Sciences FF-049
Email: sanet.hugo@univen.ac.za

Content summary:
• What is evolution? How did the theory develop?
• Darwin’s tenets of evolution by natural selection
• How do we collect evidence?
▪ Fossils
▪ Comparative anatomy and embryology
▪ Biogeography
▪ Genetics
• Mendel’s laws of inheritance
• The four forces of evolution
▪ The Hardy Weinberg Equilibrium
▪ Genetic drift
▪ Mutation
▪ Gene flow
▪ Natural selection
• Macroevolution
▪ Adaptation
▪ Speciation
• Important events in the history of life on Earth

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Development of the Theory of Evolution
The scientific method
To help you understand how we came to
the Theory of Evolution, let’s begin by
revising the scientific method and relevant
terms.
As depicted in the diagram (Figure 1), the
process starts when we observe an
interesting pattern and we ask: “what
causes the pattern that we see?” To solve
the research question we formulate a
hypothesis.
A hypothesis is a proposed explanation for
the observation. It is an educated guess
about the cause of the observation, based
on what previous scientists have
discovered. Figure 1

We can further construct our hypothesis in a logical or mathematical form, or more

precisely, construct a null hypothesis and one or two alternative hypotheses to compare:
An alternative hypothesis HA is what we expect to observe (the results) if our proposed
explanation really cause the pattern that we are studying.
A null hypothesis H0 is what we expect to observe if our proposed explanation is wrong and
does not cause the pattern that we are studying.
We then conduct an experiment to collect observations under controlled conditions, with
which we can test our H0 and HA hypotheses. We analyse our experimental results to see if
it provides evidence in support of the alternative hypothesis or not. Based on this we accept
either the null hypothesis or the alternative hypothesis as our conclusion.
In some cases we are unable to conduct experiments, e.g. for large-scale systems like whole
ecosystems where it would be unethical or impossible to manipulate the system for an
experiment. In this case we test the hypothesis after collecting sufficient data through
further observation, while taking care that the observations are unbiased so that they are a
reasonable representation of reality.
Scientific studies should be repeated by other scientists – hyptheses should be tested
repeatedly to either strengthen the conclusions (if more supporting evidence is found), or to
discard hypotheses that turned out to be untrue (if new evidence supports the null
hypothesis or a completely different hypothesis) or inconclusive (the results do not provide
enough information to come to a conclusion).
A Scientific Theory is a widely accepted explanation that is based on a large body of
evidence. A hypothesis can become a theory if it has been repeatedly tested and we have
gathered a large amount of evidence in support of the theory. A scientific theory can still be
tested with new observations.

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This process is a cycle. The results and conclusions of our experiments often lead to even
more questions. Scientists always build on the work of those who came before them.

What is evolution?
A scientific theory that explains the origin and change in lifeforms from earlier ancestral
forms. Evolution is the central unifying principle of modern biology. Biologists do not explain
any patterns or do any studies without the background of evolution. It has been repeatedly
tested with new observations, and studies are still ongoing today. As a scientific theory it
has held up for more than 160 years, with an increasingly large mountain of evidence in
support and not one instance, so far, of evidence in contradiction to what is expected from
evolution.

What did we need to understand before we could understand evolution?

Understanding evolution was made possible by several major developments in human
thinking.
1. Separate supernatural and natural explanations
The Greek philosopher Anaximander, who about lived about 2600 years ago, proposed that
scholars leave behind the supernatural and search for natural explanations to explain the
natural world.

2. Hypothesis testing
A proposed explanation for an observed pattern, should be tested against more
observations. In other words, we must support our ideas with evidence.
The Greek philosopher Aristotle, who lived
about 2300 years ago, wrote that “we must
not accept a general principle from logic only,
but must prove its application to each fact; for
it is in facts that we must seek general
principles, and these must always accord with
the facts,”

3. Realising that the Earth is ever-changing

The Earth is constantly changing (Figure 2).
Climates, ecosystems, landscapes, geological
formations, even the chemistry of the
atmosphere, has changed much over geological Figure 2: Marine fossils like these snails are
sometimes found at the top of mountains.
and shorter timescales.
The tops of these mountains were once the
bottom of the ocean.

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4. Realising that the Earth is very old
James Hutton (in the 18th century) and Charles
Lyell (19th century) explained that massive
features of the earth like mountains and
canyons could be explained by simple
everyday processes, like soil erosion (Figure 3).
Early researchers after Hutton and Lyell argued
much about the age of Earth, but none of
them estimated that the Earth is more than
100 million years old. Today, we estimate the Figure 3: The Grand Canyon, the largest
age of the earth at about 4.54 billion years! canyon on Earth, was formed over a long
period by erosion. This process is still
5. The development of Natural History ongoing as the Colorado River is still
further eroding the canyon.
Natural History is a scientific field focusing on
the study and description of living organisms
and other natural objects like fossils. Natural History supplied many of the observations and
samples that inspired questions about the origin and development of life on Earth.
Aristotle can be thought of as one of the first biologists. He described hundreds of species
and classified them into groups, an early attempt at taxonomy.
Some more recent noteworthy Natural Historians, who asked questions about the origin of
the diversity of life, include: Erasmus Darwin (Charles Darwin’s grandfather) who proposed
that life was passed on through common ancestry, and Robert Chambers who hypothesized
that new species arise from old species in a process called progressive development.

The role of Natural Selection in driving the process of Evolution

Many of the central ideas of Evolution were already known before Charles Darwin. It was
already known that parents passed on their ancestry to their children and that species
developed from one another progressively. In other words, they evolved. So why did we
need Darwin and his idea? We needed to discover the mechanism or method by which the
process of evolution changes species over time.
Some useful terms
Population: Organisms of the same species in a certain time and area, which can breed with
each other to produce offspring.
Species: One or more populations of similar organisms that are able to breed and produce
offspring if they can meet. See more about species concepts later in the course.
Traits (or characteristics): Any aspect that describes an organism – appearance, anatomy,
behaviour, physiology, biochemistry, genetics, etc.

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Lineage: Direct descent from an ancestor. Examples in different contexts:
• A sequence of members of the same family, e.g. a child from a parent from a
grandparent from a great grandparent, etc.
• A sequence of species, each of which evolved from a previous ancestral species.
• A sequence of body cells which developed from an ancestral cell in the body.

The Theory of Evolution by Acquired Characteristics

Before Darwin, Jean-Baptiste Lamarck described the Theory of Evolution by Acquired
Characteristics in his book Philosophie Zoologique published in 1809. This theory contends
that organisms adapt to their environment through an “inner force” that compels them to
develop traits that they need to survive in the environment. The organism’s offspring then
inherit these acquired traits, so they start off being better adapted to their environment.
Further observation and experiment found evidence that contradicts this theory. For
example, August Weismann (1834-1914) discovered that only the germ cells (sperm and egg
cells) play a role in reproduction and that the germ cells and the rest of the body’s cells do
not share information. There is no way for acquired traits involving the body cells to be
inherited by the organism’s offspring. Therefore, we discarded Lamarck’s theory.

Figure 4: The diagram illustrates the evolution of tall giraffes over generations as suggested
by the Theory of Acquired Characteristics.

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The Theory of Evolution by Natural Selection
Both Charles Darwin and Alfred Wallace independently proposed that natural selection is
the mechanism driving the process of evolution. Both Darwin and Wallace separately
travelled the world and collected observations and samples of existing species and fossils of
extinct species. Darwin began his work earlier than Wallace and was the first to publish
about this theory in his book Origin of Species. Unlike Lamarck’s theory, Darwin and
Wallace’s theory contends that random variation is already present in all populations
instead of being acquired by some ‘inner need’. Some individual organisms in a population
could have traits that are better suited to survival and reproduction in the environment,
compared to other individuals in the same population that lack these traits. The well-
adapted organisms are more likely to survive and have offspring, which may inherit these
favourable traits from their parents. Therefore, a larger proportion of the next generation
will have these favourable traits. Natural selection selects certain individuals, but evolution
acts on populations, not on individuals!

Darwin’s Tenets of Natural Selection

Darwin proposed the following four principles that are needed for evolution to occur by
natural selection.

1. Variation
Individuals of the same species in a population
vary in appearance, behaviour, physiology, and
other traits (Figure 5). This variation is the
foundation upon which natural selection operates.
Without variation, there can be no natural
selection. Variation arises by chance. Different
traits can be advantageous, detrimental, or
neutral in the specific environment where the
organism lives.

2. Heritability
For natural selection to favour or oppose a trait,
the trait has to be heritable. That means the trait
must be consistently passed on from parent to
offspring. Darwin did not know about genetics,
but we now know that we inherit traits from our
parents through the variant of genes (alleles) that
they pass on to us.
The way that traits are expressed in an organism
(the phenotype) is always a combination of genes Figure 5
and environment. Strongly heritable traits tend to be more strongly influenced by the alleles
inherited from the parents, and these traits tend to remain unchanged by the environment
where the organism grows up. Some traits are only weakly heritable, usually when the
environment has a strong influence on the phenotype while genes play a smaller role. Some
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traits are not heritable at all, for
example, an injury that changes the
appearance or anatomy of an organism
(i.e. an acquired characteristic).

3. High rate of population growth

If there are enough resources and
optimal environmental conditions to
support all the individuals, then there
will be no competition (struggle)
between the individuals, and survival
will be easy for everyone regardless of
their traits. Figure 6: This dense population of crabs will
compete for space, food, and mates.
Most populations have more offspring
each year than local resources can
support leading to a struggle for
resources (Figure 6). Natural resources
that can become depleted by the
populations that depend on them are
called limiting resources. Each
generation may experience substantial
mortality as many organisms lose in the
competition. Similarly, there may be
competition among the surviving
individuals to find a mate and
reproduce. In each generation, many
individuals may fail to produce offspring.

4. Differential survival and reproduction

Not all individuals will survive the
struggle. Of those that survive, not all
will reproduce. Of those that reproduce,
not all will produce the same number of
offspring that survive.
If an individual possesses a heritable
trait that helps them survive in the local
environment or a trait that helps them
reproduce successfully, that individual
may contribute more offspring to the
next generation (Figure 7). These
offspring will then inherit traits that give
them higher evolutionary fitness, as the
offspring will also be better able to Figure 7: The brown beetles have a higher fitness
survive and reproduce. Evolutionary than the green beetles because they have a trait
fitness is determined by the number of (being brown) that help them survive.

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individuals that survive to produce offspring that also reproduce. A larger proportion of
individuals in the next generation will have these favourable traits – in other words, the
population has changed or evolved.

Some misconceptions about evolution

1. “Some species are more evolved than other species”
There is no ideal most-evolved species. Artificial
selection by humans (i.e. domestication of
livestock and cultivating of crops) is goal-
directed, but evolution by natural selection has
no goal. Some organisms can be better adapted
to a certain environment, but when the
environment changes, they may suddenly find
themselves poorly adapted to the new
conditions. Environmental change is constant on
Earth.

2. “If humans evolved from apes, why are there

still apes?”
Figure 8: Although humans are genetically
Also, other similar statements, like: “…why are
closely related to chimpanzees and bonobos,
there still single-celled organisms”
the last common ancestor we shared with
Humans did not evolve from the apes that live these apes lived about 7 million years ago.
today. Instead, we share a common ancestor with
them, which no longer exists (Figure 8).
Species do not become extinct just because they are less complex, and they do not survive
better just because they are more complex. There are still thousands of simple microbes
thriving today that are biologically similar to the earliest lifeforms on Earth. Any species,
even us, can become extinct if their environment changes so much that none/not enough of
the available individuals have traits that support survival in the environment.

3. “No-one has ever seen a species evolve, therefore, there is no evidence that it happens.”
On the contrary, we can observe change in populations over time. Some organisms have a
very short generation time and/or they are subjected to a rapid change in their
environment. Since humans are long-lived and we can keep written records for hundreds or
thousands of years, we have been able to gather data tracking change in populations, in
other words evolution. For example, the proportion of two different colour morphs (forms)
in populations of the peppered moth changed depending on how well each colour morph
could camouflage in the environment (Figure 9).

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 Figure 9: Populations of peppered moth Biston betularia can vary in colour. Before the industrial
revolution, the dark coloured moths were rare because they could not camouflage against the
pale lichens growing on the tree bark (left). As they could not hide from predators, they were
less likely to survive. Pollution from coal-burning factories during the industrial revolution killed
the lichens and turned the bark of trees dark with soot. During this time, the pale moths became
rare and the dark moths more abundant (right). Later, laws were passed to ensure less pollution.
When the air quality improved, the lichens grew back, meaning that the pale B. betularia moths
became more abundant again, while the dark moths became rare. These changes were observed
and studied between 1848 and 1973.

How do we collect evidence to test the theory of evolution?

Fossil record
A fossil is the remains of a once-living organism. These remains can include original material
from the dead body, petrified material where the original material has been replaced with
minerals, and casts, molds and imprints, like footprints (Figure 10).
Plant fossils and microscopic fossils like pollen and spores can tell us which kinds of plants
occurred in a location at a certain time in the geological past, and therefore, which kinds of
ecosystems and climates prevailed at that time and place.
Fossils need certain ideal conditions to form, e.g. being rapidly buried in sediments or soft
clay that hardens to become a protective rock layer. Therefore, organisms rarely become
fossilised and the fossil record preserves only a fraction of the diversity of life in past
geological ages, likely only the most abundant species.

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 A B

D
Figure 10: Different types of fossil (A) mold of a snail shell left in surrounding
material, (B) original material in amber, (C) petrified wood, (D) dinosaur footprints.

There are several methods to estimate the age of fossils so that they can be arranged in a
timeline.

1. Relative dating
Sedimentary rock is laid down in layers from old to new (Figure 11). The sequence of fossils
can be determined by the sequence of sedimentary layers. Relative dating on its own cannot
indicate the age of the fossil, but only give us the sequence of events from earlier to more
recent.

Figure 11: The sequence of layers in sedimentary rocks can be used to organise fossils from
older to newer. Certain layers can be identified even in different locations.

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2. Radiometric dating
Also known as absolute dating, or chronometric dating. The age of the fossil, or rock layer, is
determined by measuring radioactive decay of certain elements in the rock. The proportion
of the radioactive isotope in the fossil or surrounding rock is measured. Age is expressed by
the time needed for half of the
radioactive isotopes to decay to
stable isotopes (Figure 12). This
time period is known as a half-life.
Different elements have different
half-lives, and are used for different
time-scales. For example, Carbon
14 which is radioactive and
unstable is transformed into stable
Nitrogen 14. Carbon 14 has a half-
life of 5730. Other examples
include Uranium-Lead = 700 million
years, and Potassium-Argon = 1300
million years. Figure 12

Biogeography
The study of the distribution of
species on Earth. Species
geographic ranges change and
expand over time. Closely related
species (with a more recent
common ancestor) tend to live
geographically closer together. The
distribution of existing species and
extinct species (fossils) can tell us
where species evolved and how
they spread across the Earth.
Alfred Wallace’s biogeographical
observations from around the
world contributed to his ideas about Figure 13: Alfred Wallace described zoogeographic realms
evolution. He is known as the based on the geographical distributions of related animal
“Father of Biogeography”. species. A modern updated version is shown here.

Comparative anatomy
We can compare homologous traits from different species to study relatedness.
Homologous structures or traits, are traits found in different species that have the same
ancestral origin. In the different species the trait may evolve to have different shapes or
functions. For example, the molar teeth of a horse and a lion are homologous – their

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common mammal ancestor also had molars (Figure 14). However, in the lion the molars are
shaped to better cut meat, whereas the horse’s molars are shaped to grind grass. The large
difference in the function of horse and lion molars suggest that they are more distantly
related, i.e. that their common ancestor lived very long ago, compared to e.g. a lion and a
house cat, who are more closely related with similarly-shaped molars for cutting meat.

Figure 14: An African lion (Panthera leo) skull on the left and a horse (Equus caballus) skull on
the right.

Analogous traits are traits

in different species that
were not derived from
the same ancestral trait,
even though they may
have the same function
currently. For example,
the wing structure of
insects are outgrowths of
the exoskeleton
supported by sclerotised
(hardened with minerals)
veins, whereas the wings
of flying vertebrates are
supported by bones,
which insects do not have
(Figure 15).
Organisms from different
lineages that adapt to the
same conditions may
independently evolve
similar-looking traits with
the same function. This
process is called
convergent evolution,
when similar solutions are
used to solve the same
problem. For example, Figure 15

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the common ancestors of birds, bats and insects did not fly, but all three lineages
independently evolved wings to gain the advantages of flight.
Vestigial traits are traits that lost most, or all, of their original function over the course of
evolution. For example, kiwi birds are flightless birds that have greatly reduced vestigial
wings (Figure 16). Further, some snakes have spurs where the ancestral reptile species had
legs. The males currently use these spurs during reproduction, but the females do not seem
to have any use for their own spurs. Humans have many examples of vestigial traits in our
bodies, including wisdom teeth, appendix, tailbones, goosebumps, etc.
A
B

C D

Figure 16: Kiwi birds (Apteryx spp.) (A) have greatly reduced vestigial wings (B). Goosebumps on
humans (C) are one of our many vestigial traits. The muscles attached to our fine body hair still raise
the hair in response to cold and fright, even though we do not have the benefit of thick fur to keep
us warm or make us look bigger (D).

Comparative Embryology
Embryos at early stages of development may display some traits shared by common
ancestors. Ernst Haeckel (1834 – 1919) hypothesised that embryos, in the course of
development, repeat the evolutionary history of its ancestral species. He stated that
“ontogeny recapitulates phylogeny”. This hypothesis is no longer accepted today. However,
as in Comparative Anatomy, homologous traits of embryos of different species can be
compared to study relatedness and evolution. For example, all tetrapod vertebrates have
pharyngeal arches during early stages of embryonic development (Figure 17).

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 Figure 17: Different stages of development of various species’ embryos are shown here. The four
on the right are all mammals.

Genetics
The previous researchers like Darwin did not know about genes and DNA, but modern
evolutionary biology has been greatly enhanced by the use of genetics, especially now that
we have advanced DNA sequencing techniques and powerful computers that can determine
the nucleotide sequence of entire species genomes.
Here are a few genetics terms that will be useful for the rest of the course:
Nucleotide: The building blocks (monomers) of DNA. Four types exist – Adenines, Guanines,
Cytosines and Thymines.
DNA: A polymer (chain) of nucleotides, Deoxyribonucleic acid – the molecule that carries
the genes, i.e. code for inherited traits.
Gene: A length of DNA coding for a functional product, such as protein.
Alleles: Different versions of the same gene – each allele differs in its nucleotide sequence.
Genotype: The combination of two alleles that a diploid organism has for every gene.
Homozygotes of a particular gene have two identical alleles. Heterozygotes of a particular
gene have two different alleles.
Chromosome: A length of DNA packaged with proteins, with all or a part of the genes of an
organism. Diploid organisms like us have nearly identical pairs of chromosomes (they are

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homologous). We inherit one chromosome of each pair, from each parent. Humans have 23
pairs of chromosomes. Heterozygotes carry two different alleles on the two chromosomes.
Phenotype: The way a genotype is expressed in an organism.
Gene pool: All genes, including all the different alleles, in a population.
Genome: All DNA (including genes and regulatory or non-functional DNA) in an individual
organism.
All living organisms on Earth follow the same universal genetic code – this is the
fundamental homologous trait that all life shares with our earliest common ancestor! We
can compare DNA sequences among species to develop very detailed phylogenetic trees
depicting the relatedness between species and lineages. Homology is also important in
genetics – we compare homologous sections of chromosomes and DNA. More distantly
related species will have more differences in the genetic code in homologous DNA
segments. Genetic code differences arise from mutations, e.g. when a nucleotide is
inserted, deleted or replaced with another nucleotide, or when the structure of a
chromosome is changed. Mutation rate of different parts of the genome can be determined
and used as a “molecular clock” to estimate the time since divergence from common
ancestors.
Instead of comparative embryology, since the late 1970’s we have a new field of research
called evolutionary developmental biology (“evo devo”) which combines embryology with
genetics. Evo devo includes the study of the Hox genes – a set of genes shared by all
bilaterally symmetrical animals that work together to control the development of the
animal’s anatomy. Small changes in the Hox genes can lead to changes in an animal’s body
shape, e.g. length and position of the limbs and tail, number of fingers and toes, etc.
Comparative anatomy and biogeography are still very important to study the evolution of
organisms from Earth’s past. It is very difficult to extract real ancient DNA from fossils,
especially old fossils like the dinosaurs. Molecular clocks also need to be calibrated using
information from the fossil record and radiometric dating.

Mendelian inheritance theory

Although Darwin knew that traits that take part in evolution should be heritable, he could
not figure out the mechanism by which traits are inherited. This was solved by Johan Gregor
Mendel, who is known today as “The Father of Genetics” despite the fact that he did not
know about genes and DNA. Mendel was a supporter of Darwin’s Theory of Evolution by
Natural selection. He studied mathematics and physics at university. After university he
became a monk and worked at a monastery where he had access to libraries and
laboratories. Here Mendel studied the way traits were inherited across generations by
crossing thousands of common garden pea plants with certain traits like different flower
and seed colours, and differences in seed shape. Luckily for Mendel, he picked traits that
were highly heritable and determined by single genes.

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Example of Mendel’s experiments
Mendel crossed purple-flowered peas with white-flowered peas. Every plant in the next
generation (F1 generation) were purple-flowered.
Next, Mendel crossed plants from this F1 generation with each other. In the next generation
(F2 generation), a quarter of the plants had white flowers and ¾ plants had purple flowers.
How did this happen? Why did the white flowered phenotype disappear in the F1
generation and reappear in the F2 generation? Using his mathematical skills Mendel figured
it out.
Punnett square
A Punnett square is a diagram used to predict all possible genotypes of the offspring from a
particular combination of parent genotypes (Figure 18).
All organisms must have two alleles, one from each parent. In Mendel’s experiment the
colour of the pea flower petals are controlled by one gene that codes for proteins involved
in manufacturing the purple pigment. Let us name the alleles from Mendel’s experiment P
for purple and p for white. The P allele produces the pigment, while the p allele is a mutated
gene that fails to produce the pigment. In the first part of the experiment Mendel mated a
purple-flowered plant with the genotype PP with a white-flowered pp plant. Both genotypes
are homozygous.

Figure 18: We can use Punnett squares to help us see what happened to the pea plants
in Mendel’s experiment.

All plants in the F1 generation received one P allele from one parent and one p allele from
the other parent, giving them a heterozygous Pp genotype. Only one P allele is needed to
produce purple flowers. The P allele is dominant over the p allele as only the P allele’s
phenotype is expressed while the p allele’s presence is masked.
From the Punnett squares in Figure 18, we can see that, when both parents are
heterozygous, there is a ¼ chance for the F2 offspring to be homozyous dominant PP, ½

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chance to be heterozygous Pp ( ¼ + ¼ ), and ¼ chance to be homozygous recessive ¼. ¾ of
the offspring will have purple flowers ( ¼ + ½ ) while ¼ will have white flowers that is only
possible from a homozygous recessive genotype. This is why, when Mendel produced a
large number of plants in the second part of the experiment, 25% of all plants in this F2
generation produced white flowers.

Mendel’s Laws of Inheritance

Based on his experiments Mendel developed four laws of inheritance:
1. Law of particulate inheritance: Each particle (gene) is inherited whole.
2. Law of Dominance: One allele will be dominant over the other, which is recessive.
3. Law of Segregation: The two parental alleles of the gene segregate and the offspring
receives one allele from each parent.
4. Law of Independent assortment: Different traits are passed on to the offspring
independently. There is no mixing between traits.

The Modern Synthesis

Although Darwin’s natural selection theory is still valid, our modern understanding of
evolution and the drivers behind the process changed dramatically over the past 160 years,
and we are still discovering new aspects of evolution today. The “Modern Synthesis”
combine Darwin’s theory of evolution by natural selection with Mendel’s theory of
inheritance. We now define evolution as: The sum total of the genetically inherited changes
in the individuals who are members of a population's gene pool. Or more simply: Any
change in the frequencies of alleles in the gene pool of a population (Figure 19). This
approach is also known as microevolution.

Time
Figure 19: This population of squirrels has evolved because the frequencies of the two
alleles for coat colour have changed with each generation.

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Hardy-Weinberg Equilibrium
p2 + 2pq + q2 = 1
The Hardy-Weinberg equation gives us the null model for testing microevolution. It allows
us to predict the genotype frequencies of a population that is in Hardy-Weinberg
equilibrium, for any set of two allele frequencies. To test whether a gene in a diploid
organism is evolving we compare the observed genotype frequencies to the frequencies
predicted by the Hardy-Weinberg equation.
Null hypothesis (H0): The observed genotype frequencies of a certain trait in a population
will be equal to the genotype frequencies predicted by the Hardy-Weinberg equation. The
trait is in Hardy-Weinberg equilibrium and is not evolving.
Alternative hypothesis (HA): The observed frequencies will differ significantly from the
frequencies predicted by the Hardy-Weinberg equation. The population is not in Hardy-
Weinberg equilibrium; the population is evolving.
The Hardy-Weinberg equation was derived from Mendelian inheritance and is closely
related to the Punnett square. Columns and rows are multiplied in each cell.
For a trait with two alleles A and a, let p = frequency of A and
q = frequency of a.
p+q=1
(p x p) + (p x q) + (p x q) + (q x q) = 1
p2 + 2pq + q2 = 1
p2 = frequency of AA
2pq = frequency of Aa
q2 = frequency of aa
For example: For p = 0.7 and q = 0.3
p2 = 0.49
2pq = 0.42
q2 = 0.09
0.49 + 0.42 + 0.09 = 1

A trait in a population is in Hardy-Weinberg equilibrium only when all of the following are
true:
• Mutation never occurs.
• Natural selection never occurs.
• The population is infinitely large.
• All members of the population breed.
• All mating is totally random, no sexual selection.
• Everyone produces the same number of offspring.
• There is no migration in or out of the population.

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The Four Forces of Evolution
The Hardy-Weinberg Equilibrium is disrupted by four forces: genetic drift, mutation, gene
flow, and natural selection. These forces change allele frequencies in a population from one
generation to the next by adding alleles to or removing alleles from the gene pool.
1. Genetic drift
This is the change in allele frequencies in
a population due to chance events. It
operates in every population in existence
because no population is infinitely large.
A parent only passes half of their alleles
to their offspring, and which allele is
passed on is a random event.
Further, due to random events in
individuals’ lives, not all individuals have
the opportunity to breed and produce
young and therefore not all can pass on
their genotype and their particular
alleles to the next generation.
Based on these random events that
happen to individuals and alleles, the
number of individuals carrying particular
alleles change across generations – the
frequencies of various alleles change
across generations (Figure 20).
Random events that happen to
individuals do not have a strong effect
on a large population with many
individuals – genetic drift in a large
population is small and the allele
frequencies don’t fluctuate very much
(Figure 21).
Genetic drift has a stronger effect in
small populations. Random events that
happen to the few individuals in small
populations have a great effect in the
population – allele frequencies can
fluctuate wildly (Figure 21). In a small
population it is more likely that an allele
can reach zero (no individuals carry that
allele) and become extinct, or that it can
become fixed (all individuals carry only Figure 20 Random reproduction pass alleles on to
that allele) (Figure 20, third generation). their offspring randomly.

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Figure 21 Allele frequencies fluctuate across generations.
Compared to large populations, allele frequencies in small
populations tend to fluctuate more than in large populations,
and are more likely to go extinct (hit zero) or become fixed
(become the only allele of that gene in a population).

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Let’s imagine a large population of beetles living on an island (Figure 22). The beetles are all
the same species, but they have a variety of alleles and genotypes, and this cause individuals
to have different phenotypes. There are two colour morphs in the population, and some
variation in the size of the adults. Even if we don’t know which alleles are present, we can
use the variation in appearance of the beetles to study various scenarios of microevolution.
If their colour and size do not affect their survival or reproduction (do not affect their
fitness), and no new beetles arrive on the island from elsewhere, then the number of blue
beetles and red beetles and different sizes will fluctuate randomly across generations =
genetic drift.

Figure 22 A population of
beetles living on an island.
There are two colour forms
and some size variation.

One day disaster strikes and most of the island burns down, killing most of the beetles
(Figure 23). By chance most of the surviving beetles are red. When a population becomes
small, only a few individuals can possibly pass their alleles on to the next generation. The
population may lose many alleles before it starts to recover in size – it has gone through a
population bottleneck and its genetic diversity is reduced.

Figure 23 A fire burns most of the

island and kills most of the beetles.
The survivors are mostly red
beetles.

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 Real world example 1: population bottleneck
Conservation biologists are concerned about endangered species like cheetahs Acinonyx jubatus
with small population sizes that have lower genetic diversity. Cheetahs had already been through a
population bottleneck in their evolutionary history and already had low genetic diversity before their
current problems even began.

Genetic drift has a stronger effect while the population is small. Although there was a
chance for the blue beetles to increase in number again and persist in the population, it is
more likely that they will reach zero, i.e. become extinct, because there were so few left
after the fire (Figure 24).

Figure 24 Genetic drift is strong in a small population. The frequency of the blue beetles
fluctuated in the population but ultimately drifted to extinction. The alleles producing blue beetles
were lost from the population.

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Only red beetles remain, which increase in number to form a large population with only red
beetles (Figure 25). This is called a founder effect – the few red beetles from which a large
population grew were the population’s founders, and no blue beetles are present.

Figure 25 After the fire, the

remaining red beetles reproduce
and the population becomes
large again. Now the whole
population is red beetles,
because all the founders were
red.

Real world example 2: founder effect

Many of our crop species like potatoes (left) and mielies (right) have more variety and much greater
genetic diversity in the regions where they were first farmed. Only a few individuals were introduced
to other places around the world. These individuals carried only a small number of all available
alleles, and they were the founders of all the other populations around the world. Their reduced
genetic diversity tend to make them more vulnerable to diseases, which affects food security.

Mutation and gene flow

There are two possible ways for genetic variation to increase in a population. One is through
mutation, which creates new alleles. It is very unlikely that specifically blue beetles can
return to the population through a new mutation. However, generally mutations are
common.

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Genetic variation in a population can also increase due to immigration of organisms into the
population from another population. This process can also reintroduce alleles that were
previously extinct from a population. In our imaginary example, if there was another island
close enough to the first island so that beetles can sometimes disperse by flying between
the islands, then alleles can be shared between the islands. If both colour forms exist on the
other island so some of the beetles coming from the other island are blue, the alleles
causing the blue phenotype can be reintroduced to the population (Figure 26).

Figure 26 Beetles disperse

between two neighbouring
populations. In this way blue
beetles are reintroduced to
the previously all-red
population.

This migration of individuals and alleles between populations is called gene flow. Gene flow
of individuals into a population (immigration), and mutations, increase genetic variation.
These two processes work in opposition to genetic drift, and gene flow out of a population
(emigration), which reduces genetic variation.

Natural Selection
Now, what if we had a large population of red and blue beetles, and one day a new bird
species arrives on the island (Figure 27).

Figure 27 A new beetle-

eating bird species arrives on
the island.

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These birds eat mostly red beetles because they are easier to see. More of the blue beetles
survive and they can therefore produce more young than the red beetles. Fewer of the
alleles for the red colour form is passed on to the next generation and the red beetles
become rare compared to the blue beetles (Figure 28). The blue beetles have a higher
evolutionary fitness, because they have a trait that helps them survive. This is a form of
directional selection, because the average beetle in the population is now a blue beetle,
whereas before the birds arrived the population was fairly evenly mixed. You may notice
that this imaginary example is similar to the real-world example of the peppered moth,
which you read about earlier in the guide.

Figure 28 If the birds

preferentially eat red beetles,
the red colour morph is
selected against and become
rare in the population relative
to the blue beetles.

If the red beetles continue to be selected against, the frequency of the red allele in the
population could decrease until it becomes extinct. Conversely the blue beetles are selected
for or favoured by natural selection and the frequency of blue beetles will increase until the
allele becomes fixed, meaning that there is no more fluctuation in allele frequency because
all of the beetles are blue (Figure 29).

Figure 29 If the birds continue to select against the alleles producing red beetles, these alleles
decrease in frequency and become extinct. Conversely, the alleles producing blue beetles (shown
here) increase in frequency and eventually become fixed.

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 Real world example 3: directional selection
Elephants (Loxodonta africana) in Mozambique’s Gorongosa National Park have a higher frequency
of tusklessness compared to other populations, because this population was heavily poached during
the civil war for their tusks and having tusks became a disadvantage. Elephants with a genetic
mutation that prevents the development of tusks were more likely to escape poaching and survive
the war. They then pass this trait on to their offspring.

When we quantify the red and blue colour forms of the beetles, we can put them in two
categories, red and blue. This is a categorical trait.
Now let’s look at the variation in size among beetles. This is measured on a continuous
scale, for example we can measure the length of beetles with a caliper or ruler (Figure 30),
and we would find all kinds of sizes on the ruler and not just a clear difference between

Figure 30 The number of beetles in a population of 10 000, each with a certain length, is shown on
the y axis and the length (or any other continuous trait) is on the x axis. Here the mean length of
beetles is around 20 mm. Few beetles are smaller than 15 mm or larger than 25 mm.

Page 26 of 50
small and large categories. This is called a continuous trait and there are likely many genes,
each with various alleles, that work together to cause the continuous variation.
We can use a distribution curve to picture how the different sizes are distributed across all
individuals in a population (Figure 30). The trait, in this case it could be length in millimeters,
is on the horizontal x axis, and the number of beetles on the vertical y axis. The mean length
of individuals in the population is around 20 mm, and most of the beetles in the population
can be said to be between 17 mm and 23 mm in length, but a few individuals are very big
>23 mm and a few are very small <17 mm.
Back to our island, where we now find that the birds have all gone from the island, but all
beetles are blue with variation in adult size. One day, a ship visits the island, and some mice
that were on the ship jump off (Figure 31).

Figure 31 Mice are

introduced to an island with a
species of blue beetles with
variation in adult sizes.

The mice like to eat beetles, but because they are very small, they can eat only the small
beetles. Few of these small beetles survive long enough to reproduce, while the larger
beetles are not affected, and survive and have more young than the smaller beetles. The
alleles that cause the beetles to be larger are more likely to be passed on to the next
generation, and small beetles become very rare or extinct. After the mice have affected the
population (put selective pressure on the population), the population has mostly medium to
large sized beetles, and the average length of the beetles in the population will be bigger
(Figures 32 and 33). This is also a process of directional selection.

Page 27 of 50
 Figure 32 Due to predation by
mice the smallest beetles
were selected against and
became rare or absent in the
population.

Figure 33 Predation by mice on the smallest beetles caused the average size of the beetles to
increase. Few beetles are now smaller than 20 mm in length.

But what if the ship brought not only small mice that ate the small beetles, but also some
chickens escaped and lived on the island and ate only the biggest beetles (Figure 34)? There
are therefore selection pressures working at the smallest and the largest extremes of the
size variation, and the medium-sized beetles would have the better chance of survival to
produce young – alleles causing a medium size are more likely to pass on to the next
generation. The mean size of beetles in the population may not change, but the range of
sizes found on the island will be narrower, in other words there will be less variation in size
among the beetles (Figures 35 and 36). This is called stabilizing selection, and it tends to
decrease genetic variation in a population.

Page 28 of 50
 Figure 34 Mice that eat only
the smallest beetles, and
chickens that prefer to eat
the largest beetles, are
introduced to an island where
adult beetles vary in size.

Figure 35 Due to predation by

mice on the smallest beetles,
and chickens on the largest
beetles, medium sized
beetles were more likely to
pass on their alleles to the
next generation.

Figure 36 Selective pressure against the extremes of a trait will not change the population mean
(mean length) but will reduce variation in the trait.

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 Real world example 4: stabilizing selection

Bird clutch size tend to favour an

intermediate number. As many eggs as
possible are produced to ensure that as
many chicks as possible survive, but the
clutch size is limited by the amount of
energy and resources the parents can put
into the eggs and chicks.

But what if, instead of mice and chickens, a frog is introduced to the island (Figure 37),
which eats only beetles of a medium size?

Figure 37 Frogs are

introduced to an island with a
species of blue beetles with
variation in adult sizes.

Now the medium-sized beetles are selected against and become rare in the population
while the smallest and largest beetles have a greater evolutionary fitness and become more
abundant in the population (Figure 38). This is called disruptive selection or diversifying
selection. The size distribution of beetles is better described by a bimodal distribution,
which has two peaks, instead of a distribution with one mean size (Figure 39). Diversifying
selection tends to increase genetic variation in a population. It is an important mechanism
for speciation, our next topic.

Page 30 of 50
 Figure 38 Due to predation by
frogs on medium-sized
beetles, the smallest and the
largest beetles are more
likely to pass on their alleles
and become more abundant
in the population, while
medium-sized beetles
become rare.

Hjk

Figure 39 Selection against intermediate traits (e.g. medium length) will favour the more extreme
traits to become more abundant compared to the intermediate traits. Disruptive selection is not
well described by a mean size as few individuals have this size. It is better described by a bimodal
distribution with two peaks describing the two different sizes prevalent in the population.

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Real world example 5: diversifying selection
Black-bellied seedcracker finch (Pyrenestes ostrinus) have either large beaks and eat large seeds
(right) or small beaks and eat small seeds (left).

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Macroevolution and speciation
Macroevolution is evolutionary change on a large scale, referring not just to changes in a
population, but to changes of whole taxonomic groups above the species level, and leading
to speciation – the origin of new species.

What is adaptation?
Evolutionary adaptation is a characteristic or trait inherited from an organism’s parents
(heritable trait) that help an organism survive in its environment and reproduce to pass on
this trait to its own offspring. Adaptation is driven by natural selection acting on the
variation in traits in a population. Those individuals with traits that increase their fitness
(meaning how well they survive and reproduce) are better adapted to the conditions in
which they live.
Each species becomes adapted to a particular niche in its environment (Figure 40). An
ecological niche is:
• The place or habitat where a species lives, which has certain environmental
conditions – temperature, water availability, soil type, vegetation.
• The type of resources they use – type of food, water, environmental structures like
trees or rocks.
• The time of day or season when they are most active.
• The interactions they have with other organisms – do they eat animals or plants, do
other animals eat them, do they have a symbiotic relationship with another species
(depend on another species to live, like a parasite)?
• All of the above together!

Figure 40 Cape sundew

Drosera capensis lives in the
arid Cederberg Wilderness
Area. You will find this
species growing on the side
of the river to get enough
water. The soil and water
don’t have enough
nutrients, so sundews catch
and consume insects on
sticky leaves. This method
of living in and using the
environment is the
ecological niche of the
sundew.

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Types of adaptations can be divided into:
• Morphological adaptations (also known as anatomical adaptations) – adaptations of
the structure of the organism, like the feet, jaws, legs and head of animals, and the
leaves, stems, roots and flowers of plants (Figure 41).
• Physiological adaptations – adaptations that regulate a body function, like digestive
enzymes, fat and nutrient storage, blood clotting, waste filtering, water
conservation, etc. (Figure 42).
• Behavioural adaptations – the ways an organism responds to and interacts with its
environment, like defending itself from danger, building or finding shelter, migrating
to a new home range, or cooperating with others of the same species (Figure 44).

A B

C D

Figure 41 The limbs of these four mammals are adapted to different kinds of movement:
A. climbing chimpanzee; B. running antelope; C. swimming seal; and D. flying bat.

Page 34 of 50
A B

Figure 42 Some examples of physiological adaptations: A. penguins have a layer of fat under their
skin to keep them warm; B. desert kangaroo rats get water from their metabolic processes and
have special kidneys to save water; C. mangrove trees can grow in salt water by filtering out the
salt before water enters the root; D. many plants, like this thornapple, is poisonous to prevent
herbivores from eating them.

Of course, an adaptation can be

classified as more than one of these
types, for example the desert
rodents’ physiological adaptations to
conserve water often depends on
structural adaptations to their
kidneys, and animals with
morphological adaptations for
camouflage (cryptic colouring) often
combine this with behaviours to
ensure that the camouflage is used
to its full effect, like choosing the
right background and staying very
still (Figure 43).

Figure 43 Can you see the owl?

Page 35 of 50
A B

C
D

Figure 44 Some examples of behavioural adaptations: A. phototropism, meaning plants bending

in the direction of light; B. parental care – here an alligator mother guards her babies on their
nest; C. tool use – here an octopus carries a seashell and a coconut shell, closing the two shells
whenever a danger is sensed; D. migration – here birds fly to a distant region where more food
will be available, the V formation improves energy use during the long flight.

What is a species?
In general, species are organisms that appear similar and that can possibly breed with each
other, but not with organisms of a different species. Species comprise one or more
populations, each population with many individuals of that species. You may think it is easy
to tell the difference between species when thinking of familiar species, like a cow or a
chicken or a pig. However, many species are so similar that it becomes difficult to decide
whether they are separate species or whether they are two populations of the same species
that look different because the populations have different alleles giving them different
outward appearances (Figure 45).
Biologists use three different species concepts to understand whether a group of organisms
are a species:
The morphological species concept depends on similarity in a group of individuals’ outward
appearance to define and describe species. It was the first method used long before
genetics and evolution were known or understood. Linnaeus, who first designed the
binomial classification method for naming species, classified similar-looking individuals as
species, with similar species grouped together in a genus. This species concept is considered

Page 36 of 50
deficient today to definitively describe species, but it is still used to recognise when a
species may be newly discovered and have not yet been described by taxonomists.

Figure 45 These two parrots (A)

A are a breeding pair of one sexually
dimorphic species, Eclectus
roratus – the male is green and
the female is red. Conversely, the
two similar looking plovers (B and
C) are two distinct species.
Genetic analysis showed Kentish
plover Charadrius alexandrinus (B)
and white-faced plover Charadrius
dealbatus (C) had evolved
separately for half a million years.

B C

The most widely used species concept currently is the biological species concept, which
depends on whether a group of individuals is reproductively isolated, meaning the
individuals of a species can only breed and produce young with each other and not with
individuals from another species, and there is no exchange of alleles between species. There
are several ways in which reproductive isolation can occur, including differences in
behaviour that prevents mating, or physiological or genetic incompatibility that prevents
them from producing surviving young that are fertile even if they do mate. However,
occasionally this concept fails, for example for species that reproduce asexually, or when
different species produce hybrid offspring that are fertile (Figure 46).
To supplement the biological species concept, the evolutionary species concept can be used
to define species by their genetic differences and evolutionary history (Figure 45 B and C).
This can be used to separate species that had a long history of evolving separately without
interbreeding, even if they could interbreed and produce fertile hybrid young under unusual
circumstances like in captivity (Figure 46), or when people move species between
continents.
A

Page 37 of 50
 A B

C Figure 46 The savannah cat breed (C)

resulted from several generations of
hybridisation between servals Leptailurus
serval (A) and housecats Felis catus (B).
However, this is only possible in captivity.
Servals and housecats would not naturally
choose to breed with each other (behavioural
barrier), and the first generations of
savannah cats have problems with fertility
and survival of the embryos (chromosomal
and physiological incompatibility).

Speciation
Since environmental conditions are always changing, natural selection constantly work on
populations. Populations with enough genetic variation have individuals with alleles better
suited to the new conditions, which become more abundant in the next generation.
Eventually, the population had so many changes in allele frequencies at so many different
genes that it has a unique genotype and become a new reproductively isolated species.
Directional selection and diversifying selection are particularly important mechanisms for
such changes, but genetic drift and founder effects also contribute, while gene flow can
slow the process.
This speciation by accumulation of changes can happen in two ways:
• Anagenesis – one species is transformed to a new species over time in response to a
changing environment and directional or stabilizing selection (Figure 47a). Also
called phyletic speciation.
• Cladogenesis – two or more species are derived from an ancestral species, usually as
a result of diversifying selection of populations in the same place or directional
selection of populations living in different habitats with very different environmental
conditions (Figure 47b). Also called divergent or branching speciation.

Page 38 of 50
 Figure 47 Anagenesis (a) or phyletic
speciation involves a species
changing over time until it is so
different from the ancestral species
that they cannot be considered the
same species. Cladogenesis (b)
involves different populations of an
ancestral species evolving separately
until the two populations are
different species.

Four different modes of speciation are proposed (Figure 48):

• Allopatric speciation – a population of a single species splits in two due to the
formation of a physical barrier and the two populations evolve separately until they
have accumulated so many changes that they are different species, for example
squirrel species separated by the Grand Canyon (Figure 49). Since gene flow is
absent genetic drift may play an important role to change the separate populations.
• Peripatric speciation involves part of the population splitting off from the main
population to expand into a new isolated niche with different environmental
pressures (similar to allopatric speciation).
• Sympatric speciation is speciation in the same time and place, driven by disruptive
selection (Figure 50) or by polyploidy. Polyploidy is a mutation where an organism
has more than two sets of chromosomes which can make it impossible to reproduce
with the original population (common in plants).
• Parapatric speciation is speciation by directional selection in different habitats, but
instead of being geographically isolated like with allopatric speciation, there is a
contact zone between the populations where some gene flow occurs and where the
two species may hybridize later (hybrid zone) (Figure 51).
• Adaptive radiation is when multiple species evolve from one ancestral species in a
relatively short time, typically when many new adaptations evolved to exploit
multiple available ecological niches (Figure 52). This could happen in response to a
large change in the environment that opens many new environmental niches, or if
the ancestral species arrives in a new environment with many available niches
(Figure 53). As the ancestral species become abundant and individuals begin to
compete for resources, multiple sub-populations evolve different adaptations to
occupy many different niches (Figure 52).

Page 39 of 50
Figure 48 Diagrams of four modes of speciation. Eventually, as shown in the last step, the two
populations have become so different that they are reproductively isolated species – even if
any geographic barriers disappear and the species meet again, they cannot interbreed.

Figure 49 Allopatric speciation: Harris’s ground squirrel Ammospermophilus

harrisi (left) and white tailed ground squirrel A. leucurus (right) have a common
ancestor of which the population became split by the Grand Canyon.

Page 40 of 50
 Figure 50 Sympatric speciation:
Soapberry bugs have specialised
proboscises to feed on the fruit of the
balloon vine. When the golden rain
tree fruit was brought into the area by
people, mutant soapberry bugs with
short proboscises (who would
otherwise have starved) began to
exploit the flatter fruits. Two different
soapberry populations formed that live
in the same area but tend not to breed
together; they are in the process of
becoming different species.

Figure 51 Parapatric speciation: Between the central African tropical rainforest and savanna
biomes is a transition area (ecotone) with characteristics of both biomes. The little greenbul
Andropadus virens populations living in the ecotone tend to differ from those in the forest, for
example, they have longer wings to fly faster and better escape predators where there is less
tree cover. There is a large amount of gene flow between the forest and ecotone populations
which would tend to mix gene pools and prevent or slow speciation, but these two habitats
are so different that natural selection continues to change the morphology of the populations
to become dissimilar. From Smith et al. 2008. Science 276 1855-1857

Page 41 of 50
 Figure 52 The finches that Darwin encountered on the Galapagos islands is a famous example
of adaptive radiation. Many different finch species, each with distinct adaptations to different
food sources and different ways to exploit the resources, have evolved in a short amount of
time by evolutionary time-scales, from a population of one species that arrived on the islands
from the South American mainland.

Gradualism and punctuated equilibrium are two hypotheses about how quickly evolution
occurs (Figure 54). Neither of these two hypotheses are currently considered to be more
accurate, and both are probably contributing to the evolutionary history of species to
different degrees and at different times.
Gradualism refers to the idea that evolution proceeds gradually over a long time, with
consistent small changes in the population average. This is the original idea adopted by
Darwin, and is sometimes observed in the fossil record, when a series of transitional fossils
show similar organisms with gradual morphological changes over time (Figure 55).
Punctuated equilibrium is the idea that species have long periods of very little evolutionary
change, but this is occasionally interrupted by periods of rapid change. This is observed in
the fossil record of some species that appear in the fossil record abruptly, after which they
seem to change very little over a long time (Figure 56). Massive ecological change can
happen rapidly (see the section on extinction later), and species can also face drastically
different environmental conditions when they disperse to new environments (e.g., the
Galapagos finches). These periods of drastic change can place strong selective pressures, on
populations, that may also go through bottlenecks, forcing strong genetic drift in a short
amount of time.

Page 42 of 50
 Figure 53 The diagram depicts the
mainland and islands over time, with
different species indicated by different
capital letters. The adaptive radiation of
finches on the Galapagos islands began
with a few individuals of one species (A)
arriving on an island where there were
many available ecological niches. The
finches most likely arrived first on the
island closest to the mainland, and then
spread to other islands. Speciation
occurred on each island as they adapt to
the variety of available niches at each
island.

Figure 54 Gradualism refers to the hypothesis that evolution happens gradually and
constantly, whereas punctuated equilibrium refers to the hypothesis that periods of slow
change are sometimes punctuated by periods of rapid evolution.

Page 43 of 50
Figure 55 The fossils of prehistoric horses map out a gradual change from a dog-sized three-
toed animal appearing first in the late Eocene around 30 million years ago, to the large one-
hooved modern horse we know today.

Figure 56 A cockroach
preserved in amber that is
between 40 million and 50
million years old (Eocene).
This ancient species doesn’t
look much different from
cockroaches that are alive
today.

Page 44 of 50
Important events in the history of life
Where did life originate and how?
Various scientific fields, like astronomy, astrophysics, geology, geochemistry and geophysics
consistently estimate the age of the earth as around 4.54 billion years, with the oceans
forming around 4.41 billion years ago. The earliest undisputed evidence of life on Earth
(microbial fossils) dates from at least 3.5 billion years ago, but there is some evidence
suggesting that life began as early as 4.28 billion years ago.
The earliest lifeforms were microbes and likely lived at hydrothermal vents on the ocean
floor (Figure 57), where they used the energy and minerals in the water to drive their
metabolism. They didn’t use oxygen (anaerobic) as there was very little free oxygen
available on Earth, and oxygen was toxic to them. They were likely similar to the
archaebacteria (Figure 57) that still live in extreme environments like these very hot
hydrothermal vents today. Life was prokaryotic microbes for most of the history of life on
Earth.

Figure 57 The earliest microbial lifeforms likely lived at hydrothermal vents in the ocean (left)
and were similar to the extremophile archaebacteria (right).

Later some microbes produced their own food

through photosynthesis (they were autotrophic), and
the waste product of photosynthesis is oxygen (Figure
58). They were very successful and became very
abundant, filling the ocean with dissolved oxygen.
Between 2.4 and 2 billion years ago The Great
Oxygenation Event occurred when this oxygen
became so much that it began filling the atmosphere.
After this time, anaerobic microbes were limited to
habitats where oxygen is absent, because oxygen is
toxic to them. However, an oxygen atmosphere
paved the way for the diversification of eukaryotes.

Figure 58 Photosynthetic bacteria like these Cyanobacteria

produce free oxygen on Earth through photosynthesis.

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The origin of eukaryotes
Genetic studies estimate that eukaryotes first appeared at least 2.7 billion years ago,
possibly evolving from archaebacteria. Eukaryotes are characterised by a variety of
organelles with different functions, and sexual reproduction. Sexual reproduction allowed
eukaryotes to evolve faster than prokaryotes, because it randomises the alleles that are
passed on, creating more variation among individuals. The next great development for
eukaryotes came when some formed symbiotic relationships with aerobic bacteria living
inside them (Figure 59), most notably the mitochondria that produce energy in eukaryote
cells, including our own. Eukaryotes that had mitochondria could develop many adaptations
that depend on rapid access to energy. Some eukaryotes also formed relationships with
photosynthetic bacteria, which allowed them to benefit from photosynthesis – these
bacteria became the chloroplasts that are found in plant cells today (Figure 59).

Figure 59 Eukaryotes evolved many organelles with different functions over time, notably the
nucleus. They further acquired a symbiotic relationship with aerobic bacteria that produced
energy for the cell (mitochondria), while some eukaryotes also acquired a symbiotic relationship
with photosynthetic bacteria that produced food for the cells (chloroplasts).

The origin of animals

Complex multicellular organisms, that have cells differentiated into several functions,
evolved in six eukaryotic groups: animals, plants, fungi, and brown, red and green algae.
Most multicellular species remained small and simple until a great adaptive radiation event,
called the Cambrian explosion, beginning about 541 million years ago and lasting between
13 and 25 million years. This is a short time considering the drastic morphological and
physiological changes, and that most modern animal phyla first appeared during this time.
One of the most important evolutionary developments before the Cambrian explosion, that
enabled the rapid diversification, is the evolution of symmetrical body plans like radial (e.g.
sea stars), and the bilateral body plan (controlled by the Hox genes) that is organised into a

Page 46 of 50
left and right, front and back, top and bottom configuration. This development, together
with environmental changes like increased availability of oxygen and calcium, and the co-
evolution of species due to predator-prey interactions, may have all contributed to the
Cambrian explosion (Figure 60).

Figure 60 The Cambrian period saw a diversity of multicellular lifeforms, new adaptations, structures
and body plans, behaviours and interactions, and the origin of most of the currently existing animal
phyla. This period began with the Cambrian explosion, a period of major adaptive radiation.

The move to land

Most of the history of life was under water. After enough oxygen was available in the
atmosphere and the protective ozone layer had been formed, organisms still needed many
adaptations before they could live on land, including adaptations to breathe air, prevent
water loss from their bodies and their reproductive units (spores and eggs), and structural
support to counter the absence of support from water. The first land plants appeared at
about 500 million years ago, spreading inland from the water’s edge at about 450 million
years ago. Invertebrates followed, beginning about 416 million years ago, followed by
vertebrates at about 360 million years ago, likely starting with amphibians (Figure 61). The
intense natural selection of the new environment, and the many available ecological niches,
facilitated another adaptive radiation in animals, with new adaptations including muscles
and bones to withstand increased gravity effects, senses to better perceive sound and light
transmitted through air, and different ways to move and navigate on land.

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 Figure 61 An artist’s impression of a Devonian period ecosystem showing some of the first
vertebrates on the water’s edge. After the plants and invertebrates had adapted to life on land
and formed ecosystems, vertebrates followed to exploit the great variety of available niches and
abundant food. The first of these vertebrates were likely amphibian and used both land and
water.

Extinction
Extinction of species is a normal occurrence as species fail to adapt to environmental
changes; however, at least five times during the course of the history of life, environmental
changes were so dramatic that a large percentage of life on earth was wiped out in a very
short time (Figure 62). This is known as a mass extinction, and it can be caused by massive
meteorites, volcanic activity, tectonic plate movements and even large-scale wildfires,
which can cause global-scale climate change and changes in atmospheric and ocean
chemistry. The greatest known extinction occurred 252 million years ago at the end of the
Permian period, when the supercontinent Pangea was formed and massive volcanic
eruptions heated and polluted the atmosphere and acidified the ocean. 95% of marine
species and 70% of terrestrial vertebrates were wiped out.
A mass extinction opens many ecological niches, and when environmental conditions
improve, an adaptive radiation can take place to fill all these niches. For example, during the
time that a great diversity of dinosaurs dominated many niches on earth, mammals were
generally small and inconspicuous (Figure 63). The Cretaceous mass extinction wiped out
the dinosaurs and most of the other flying and swimming archosaurs – the crocodilians and
birds were the only archosaurs that survived. The mammals could then diversify to fill all the
niches that were left unexploited (Figure 64).

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Figure 62 Extinction is a normal constant occurrence; however, at least five times during the
course of the history of life, a mass extinction occurred when a large percentage of life on earth
was wiped out in a very short time.

Figure 63 Mammals existed during the time that a great diversity of dinosaurs dominated the
Earth, but they were mostly small insectivores and didn’t occupy many ecological niches.

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Figure 64 After the Cretaceous mass extinction which wiped out the dinosaurs and most other
archosaurs, the mammals underwent an adaptive radiation to fill all the niches that used to be
occupied by dinosaurs and other archosaurs.

~END~

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