Charles Darwin (1809–1882) was born in Shrewsbury, in western England.

Enrolled at
Cambridge University. Soon after Darwin graduated, he joined HMS Beagle for a long voyage
around the world.
Darwin's most famous work, "On the Origin of Species," published in 1859, introduced the
theory of evolution by natural selection. Darwin spent most of his time on shore, observing
and collecting thousands of plants and animals while on the Beagle. He described features of
organisms that made them well suited to diverse environments, adaptations, inherited
characteristics of organisms that enhance their survival and reproduction in specific
environments.
His explanation of how adaptations arise centered on natural selection, a process in which
individuals that have certain inherited traits tend to survive and reproduce at higher rates
than other individuals because of those traits.
Scientist influenced Darwin
★ Georges Cuvier (Paleontologist): Cuvier was the first to demonstrate that the
different strata of rock in the Paris basin each had its own mammal fauna. He
speculated that each boundary between strata represented a sudden
catastrophic event, such as a flood, that had destroyed many of the species living
in that area in what is called catastrophism.
★ James Hutton proposed the theory of gradualism, that Earth’s geologic features
could be explained by gradual mechanisms, such as valleys being formed by
rivers.
★ Charles Lyell developed the theory of uniformitarianism, the processes by which
current geological features were created were slow, steady, and constant. The
forces molding the planet today have operated continuously throughout its
history.
★ Jean-Baptiste de Lamarck: believed that living things evolved in a continuously
upward direction, from dead matter, through simple to more complex forms,
toward human "perfection." Species didn't die out in extinctions, Lamarck
claimed. Instead, they changed into other species. According to Lamarck,
organisms altered their behavior in response to environmental change. Their
changed behavior, in turn, modified their organs, and their offspring inherited
those "improved" structures. According to Lamarck, organisms altered their
behavior in response to environmental change. Their changed behavior, in turn,
modified their organs, and their offspring inherited those "improved" structures.
Conversely, in Lamarck's view, a structure or organ would shrink or disappear if

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 used less or not at all. Driven by these heritable modifications, all organisms
would become adapted to their environments as those environments changed.
This was called the use and disuse principle. The second principle he suggested,
inheritance of acquired characteristics, stated that an organism could pass these
modifications to its offspring.
★ Alfred Russel Wallace: developed some of the most important ideas about natural
selection Wallace supplied Darwin with birds for his studies and decided to seek
Darwin’s help in publishing his own ideas on evolution. He sent Darwin his theory
in 1858, which, to Darwin’s shock, nearly replicated Darwin’s own.
★ Thomas Malthus: he believed that the human population was growing at a rate
that would be too large to be supported in the future. He thought this would lead
to a worldwide famine. He called this population problem a "struggle for
existence”.

Natural Selection
Darwin realized that explaining such adaptations was essential to understanding
evolution. His explanation of how adaptations arise centered on natural selection,
a process in which individuals that have certain inherited traits tend to survive and
reproduce at higher rates than other individuals because of those traits.
Darwin amassed evidence that three broad observations about nature
● the unity of life
● the diversity of life
● the match between organisms and their environments
resulted from descent with modification by natural selection.
In his book “ On the Origin of Species by Means of Natural Selection“ Darwin defined
evolution as descent with modification from a common ancestor, proposing that Earth’s
many species are descendants of ancestral species that were very different from those
alive today.
● All organisms are related through descent from a common ancestor that lived in
the remote past.
● Over evolutionary time, the descendents of that common ancestor have
accumulated diverse modifications, or adaptations, that allow them to survive and
reproduce in specific habitats.

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 ● Over long periods of time, descent
with modification has led to the
rich diversity of life we see today
● The history of life is like a tree,
with multiple branches from a
common trunk.
Closely related species, the twigs
on a common branch of the tree,
shared the same line of descent
until their recent divergence from
a common ancestor.
Artificial Selection
Darwin studied selective breeding of
domesticated plants and animals.
Humans have modified other species
over many generations by selecting and
breeding individuals that possess
desired traits, a process called artificial
selection. As a result of artificial
selection, crops, livestock animals, and
pets often bear little resemblance to
their wild ancestors. Darwin reasoned that if artificial selection can bring about dramatic
change in a relatively short period of time, then natural selection should be capable of
substantial modification of species over many hundreds of generations.
Darwin described four observations of nature, from which he drew two inferences.
○ Observation #1: Members of a population vary greatly in their traits.
○ Observation #2: Offspring inherit traits from their parents.
○ Observation #3: All species are capable of producing more offspring than the
environment can support.
○ Observation #4: Due to lack of food or other resources, many of these offspring do not
survive.
○ Inference #1: Individuals whose inherited traits give them a higher probability of
surviving and reproducing in a given environment than other individuals tend to leave
more offspring than other individuals.
○ Inference #2: This unequal ability of individuals to survive and reproduce will cause
favorable traits to accumulate over generations.

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Natural selection is a process in which individuals that have certain heritable traits
survive and reproduce at a higher rate than other individuals because of those
traits.
Over time, natural selection can increase the match between organisms and their
environment. If an environment changes, or if individuals move to a new environment,
natural selection may result in adaptation to these new conditions, sometimes giving
rise to new species.

Evidence of Evolution
1. The fossil record
The fossil record documents the pattern of evolution, showing that past
organisms differed from present-day organisms and that many species have
become extinct. Fossils also show the evolutionary changes that have occurred in
various groups of organisms. An example is the fossil record of cetaceans,
the mammalian order that includes whales, dolphins, and porpoises. Some of
these fossils provide support for a hypothesis that cetaceans are closely related to
even toed ungulates, a group that includes deer, pigs, camels, and cows.
A fossil that shows an intermediate state between an ancestral trait and that of its
later descendants is said to be a
transitional fossil.
For example, Pakicetus is a close relative
of ancient whales. Pakicetids lived on
land and had nostrils at the front of the
skull, as modern cows and sheep do.
The ancestors of whales probably
looked something like Pakicetus.
Scientists find evidence of this transition
in the fossils of Aetiocetus, which had
nostrils in the middle of its skull.
Fossil dating:
● Relative dating places fossils in a temporal sequence by noting their
positions in layers of rocks, known as strata. Fossils found in lower strata
were deposited earlier and thus must be older than fossils in higher strata.
● Numerical dating relies on radioactive elements, such as uranium,
potassium, rubidium and carbon. Radioactive elements decay, or convert to

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 a non-radioactive form, at rates that scientists have carefully observed.
Fossils less than about 50,000 years old can be dated directly using their
radioactive carbon content.
2. Homology
Evidence for evolution comes from similarities in the characteristics of different
organisms. Features inherited from common ancestors—even if their appearance
is quite different in close relatives— are known as homologies.
★ Anatomical Homology:
Homologous structures are similar physical features in organisms that
share a common ancestor, but the features serve completely different
functions.
Hummingbirds, whales, frogs, and rabbits all have differently shaped
forelimbs, reflecting their different lifestyles. But those different forelimbs
all share the same set of bones – starting from the shoulder, one bone (the
humerus), followed by two bones (the radius, and the ulna). These same
arm bones are seen in modern tetrapods (like us humans!) and in fossils all
the way back to the extinct fish Eusthenopteron. This sort of evidence helps
demonstrate the common ancestry of all tetrapods.

A vestigial structure is a feature that a species inherited from an ancestor but

that is now less elaborate and functional than in the ancestor. They are
features of an organism that are considered to have lost much or all of their
original function through evolution. For example, the skeletons of some snakes
and of fossil whales retain vestiges of the pelvis and leg bones of walking
ancestors.

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 ★ Molecular Homology
All organisms are made of cells. All species of life have the same basic genetic
machinery of RNA and DNA,and the genetic code is essentially universal. In
addition, the sequence of some proteins is similar across the tree of life.
★ Developmental Homology
Comparing early stages of
development in different
animal species reveals
additional anatomical
homologies not visible in
adult organisms. For
example, at some point in
their development, all
vertebrate embryos have a
tail located
posterior to (behind) the
anus, as well as structures
called pharyngeal (throat) arches. These homologous throat arches ultimately
develop into structures with very different functions, such as gills in fishes and
parts of the ears and throat in humans and other mammals.
3. Biogeography
Geographic distribution of species. The geographic distribution of organisms is
influenced by many factors, including continental drift. Two hundred fifty million
years ago, all of Earth’s landmasses joined to form a single large continent called
Pangaea. Pangaea broke apart 200 million years ago. Home → Lines of Evidence →
Distribution in time and space → Biogeography
For example, marsupial mammals (those with pouches such as the koala and
kangaroos) are found in the Americas as well as Australia and New Guinea.
Marsupials didn’t need a migration route from one part of the world to another;
they rode the continents to their present positions and continued to diversify and
move around on those separate continents.

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The Evolution of Populations
The evolutionary impact of natural selection is apparent only in the changes in a
population of organisms over time → It is the population, not the individual, that evolves.
Microevolution is defined as a change in allele frequencies in a population over time.
Three mechanisms can cause allele frequencies to change:
● Natural selection
● Genetic drift (chance events that alter allele frequencies)
● Gene flow (the transfer of alleles between populations).
Natural selection is the only mechanism of adaptive evolution, improving the match
between organisms and their environment.
Individual variation occurs in all species, heritable variation makes evolution possible.
Genetic variation is produced by:
a. Mutation: is a change in the nucleotide sequence of an organism’s DNA. Only
mutations in cells that form gametes can be passed onto offspring. New genes
and new alleles originate only by mutation.
b. Sexual reproduction: produce the genetic variation that makes evolution
possible. Sexual reproduction produces unique combinations of alleles. Sexual
reproduction shuffles variant alleles and deals them at random to produce unique
individual genotypes (Three mechanisms contribute to the shuffling: crossing
over, independent assortment of chromosomes, and fertilization).
The Hardy-Weinberg equation
A population is a group of individuals that belong to
the same species, live in the same area, and interbreed
to produce fertile offspring.
A population’s gene pool is the combination of all the
genes (including alleles) present in a reproducing
population. Each allele has a frequency in the
population’s gene pool.
Allele Frequency: the number of times an allele occurs
in a gene pool, as a percentage of total occurrence of all
alleles for that gene in the gene pool.
The Hardy-Weinberg principle describes the gene pool
of a population that is not evolving. The Hardy-Weinberg principle states that the
frequencies of alleles and genotypes in a population’s gene pool will remain constant

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over generations unless acted upon by agents other than Mendelian segregation and
recombination of alleles.

Suppose that the individuals in a population not only donate gametes to the next
generation at random but also mate at random. In other words, all male-female
matings are equally likely. The allele frequencies in this population will not
change from one generation to the next. Its genotype frequencies, which can be
predicted from the allele frequencies, will also remain unchanged.

Five conditions must be met for a population to remain in Hardy-Weinberg equilibrium.

The Hardy-Weinberg principle describes a hypothetical population that is not evolving.
Real populations do evolve, however, and their allele and genotype frequencies do
change over time.
1. No mutations. Alteration of alleles or deletion or duplication of genes will modify
the gene pool.
2. Random mating. If individuals pick mates with certain genotypes, or if inbreeding
is common, the mixing of gametes will not be random and genotype frequencies
will change.
3. No natural selection. Differential survival or reproductive success among
genotypes will alter allele frequencies.
4. Extremely large population. In small populations, chance fluctuations in the gene
pool will cause allele frequencies to change over time. These random changes are
called genetic drift.
5. No gene flow. Gene flow, the transfer of alleles due to the migration of individuals
or gametes between populations, will change the frequencies of alleles.
Deviation from any of these conditions results in evolutionary change, which is common
in natural populations.

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The rate of evolutionary change in many populations is so slow that they appear to be
close to equilibrium.
The three mechanisms that directly alter allele frequencies to bring about evolutionary
change are
● Natural selection Individuals with variations better suited to the environment
tend to produce more offspring (reproductive success) than those with variations
that are less well suited. Alleles are passed onto the next generation in
frequencies different from their relative frequencies in the present population.
● Genetic drift occurs when changes in gene frequencies from one generation to
another occur because of chance events (sampling errors) that occur in small
populations. In a large population, allele frequencies do not change from
generation to generation by chance alone.
The founder effect occurs when a new population is started by a small number of
individuals who do not represent the gene pool of the larger source population.

The bottleneck effect occurs when the numbers of individuals in a large

population are drastically reduced by a disaster.

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 ● gene flow: is the transfer of alleles among populations due to the migration of
fertile individuals or gametes. Gene flow, like mutation, can introduce new alleles
into a population. Because gene flow can occur at a higher rate than mutation, it
can have a greater effect on allele frequencies
Once gene flow or mutation introduces a new allele to a population, natural
selection may cause the new allele to increase (as in the insecticide-resistance
alleles in mosquitoes) or decrease (as in the copper-tolerance alleles in bent
grass).

There are three modes of selection: directional, disruptive, and stabilizing.

Natural selection can alter the frequency distribution of heritable traits in three ways,
depending on which phenotypes in a population are favored. The three modes of
selection are called directional, disruptive, and stabilizing selection.
Directional selection is most common during periods of environmental change or when
members of a population migrate to a new habitat with different environmental
conditions.
Directional selection shifts the frequency curve for a phenotypic character in one
direction by favoring individuals who deviate from the average.
○ For example, fossil evidence indicates that the average size of black bears in
Europe increased during each glacial period, only to decrease again during
the warmer interglacial periods.
o Large bears have a smaller surface-to-volume ratio and are better
at conserving body heat during periods of extreme cold.
Disruptive selection occurs when environmental conditions favor individuals at both
extremes of the phenotypic range over those with intermediate phenotypes.
o For example, two distinct bill types are present in Cameroon’s
black-bellied seed crackers.
o Larger-billed birds are more efficient in feeding on hard seeds, and
smaller-billed birds are more efficient in feeding on soft seeds.
o Birds with intermediate bills are relatively inefficient at cracking
both types of seeds and thus have lower relative fitness.
Stabilizing selection favors intermediate variants and acts against extreme
phenotypes. Stabilizing selection reduces variation and maintains the status quo
for a trait.
o Human birth weight is subject to stabilizing selection.

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 o Babies much larger or smaller than 3–4 kg have higher infant
mortality than average sized babies.

Natural selection is the only evolutionary mechanism that consistently leads to

adaptive evolution.
Sexual selection results in sexual dimorphism, is the systematic difference in form
between individuals of different sex in the same species. For example, in some
species, including many mammals, the male is larger than the female. In others,
such as some spiders, the female is larger than the male.Males and females may
differ in size, coloration, and ornamentation.
● Intrasexual selection is direct competition among individuals of one sex for
mates of the opposite sex.
○ Competition may take the form of direct physical battles between
individuals, usually males. For example, a single male may patrol a
group of females and prevent other males from mating with them.

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 ● Intersexual selection, or mate choice, occurs when members of one sex
(usually females) are choosy in selecting their mates from individuals of the
other sex.

Speciation
Speciation, the process by which one species splits into two or more species. When one
species splits into two, the species that result share many characteristics because they
are descended from this common ancestor. It is the evolution of a new species from the
pre-existing one by gradual modification.

Macroevolution, the broad pattern of evolution above the species level. An example of
macroevolutionary change is the origin of new groups of organisms, such as mammals
or flowering plants, through a series of speciation events
A species is a group of populations whose members have the potential to interbreed in
nature and produce viable, fertile offspring. The formation of a new species depends
on reproductive isolation—the existence of biological factors (barriers) that obstruct
members of two species from interbreeding and producing viable, fertile offspring. Such
barriers block gene flow between the species and limit the formation of hybrids,
offspring that result from an interspecific mating.
Mechanisms of Speciation
★ Prezygotic barriers (“before the zygote”) block fertilization from occurring
○ Habitat Isolation: Two species that occupy different habitats within the
same area may encounter each other rarely, if at all.
○ Temporal Isolation: Species that breed during different times of the day,
different seasons, or different years cannot mix their gametes.
○ Behavioral Isolation: Courtship rituals that attract mates and other
behaviors unique to a species.
○ Mechanical Isolation: morphological differences prevent successful
fertilization
○ Gametic Isolation: gametes (egg and sperm) come into contact, but no
fertilization takes place.
★ Postzygotic barriers: Postzygotic barriers prevent a hybrid zygote from
developing into a viable, fertile adult. The mule is a typical example. Differences in
chromosome number or arrangement of genes on chromosomes usually result in

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 postzygotic isolation because chromosomes may not pair normally during mitosis
or meiosis.

○ Reduced Hybrid Viability: offspring that have a lower rate of survival than
the parent species. This reduces the likelihood of the hybrid offspring
surviving to sexual maturity.
○ Reduced Hybrid Fertility: Hybridization normally produces sterile offspring.
○ Hybrid Breakdown: Some first-generation hybrids are viable and fertile, but
when they mate with one another or with either parent species, offspring of
the next generation are feeble or sterile.
Causes of Speciation
Allopatric speciation
In allopatric speciation, gene flow is interrupted when a
population is divided into geographically isolated
subpopulations.
Sympatric speciation
Caused by a reproductive barrier that isolates a subset
of a population from the remainder of the population in
the same area, gene flow to and from the isolated
subpopulation is blocked. This can occur as a result of
● Polyploidy, a condition in which an organism has
extra sets of chromosomes
● Sexual selection
● natural selection that results from
● A switch to a habitat or food source not used by the parent population.
Adaptive Radiation
Adaptive radiation is a type of speciation that occurs when a single ancestral species
rapidly gives rise to a variety of new species as each adapts to a specific environment.
Many instances of adaptive radiation involve sympatric speciation following the removal
of a competitor, a predator, or a change in the environment. When competition is
reduced, it results in ecological release. This is an opportunity for a species to expand its
use of resources within habitats that now have less competition. Allopatric speciation
can also cause a population to undergo adaptive radiation.
Rate of Evolution and Speciation

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In terms of how quickly speciation occurs, two patterns are currently observed: the
gradual speciation model and the punctuated equilibrium model.
In the gradual speciation model, species diverge gradually over time in small steps.
In the punctuated equilibrium model, a new species changes quickly from the parent
species and then remains largely unchanged for long periods of time afterward. This
early change model is called punctuated equilibrium, because it begins with a
punctuated or periodic change and then remains in balance afterward.

Patterns of Evolution

1. Convergent evolution is said to occur when a biological trait evolves in two

unrelated species as a result of exposure to similar environments.
Dolphins and tuna do not share a recent common ancestor, yet they both have a
dorsal fin for swimming. The ability to swim has evolved in dolphins and tuna
independently and, so, has resulted in two different, although similar-looking,
solutions to the requirements of moving through water with the greatest amount
of efficiency.
2. Divergent evolution refers to the process by which interbreeding species
diverged into two or more evolutionary groups. It means that these groups of
species used to be similar and related. However, they became more and more
dissimilar through time.
3. Coevolution: the process of reciprocal evolutionary change that occurs between
pairs of species or among groups of species as they interact with one another. The
activity of each species that participates in the interaction applies selection
pressure on the others. Coevolution is frequently seen in pairs of species that
interact closely:
a. Herbivore and plants
b. Pollinator and flower
c. Predator and prey
d. Parasite and host

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Conditions on Early Earth
Direct evidence of life on early Earth comes from fossils of microorganisms that lived 3.5
billion years ago. Observations and experiments in chemistry, geology, and physics have led
scientists to hypothesize that chemical and physical processes could have produced simple
cells through a sequence of four main stages:
1. Stage 1-Organic Monomers: The abiotic
(nonliving) synthesis of small organic molecules,
such as amino acids and nitrogenous bases
2. Stage 2-Organic Polymers: The joining of these
small molecules into macromolecules, such as
proteins and nucleic acids
3.Stage 3-Protocells: organic polymers become
enclosed in a membrane to form precursor of
the first cell
4. Stage 4-Living Cells: The origin of self-
replicating molecules that eventually made
inheritance possible
Our planet formed 4.6 billion years ago,
condensing from a vast cloud of dust and rocks
that surrounded the young sun.
The first atmosphere had little oxygen and was
probably thick with water vapor, along with
various compounds released by volcanic
eruptions, including nitrogen and its oxides,
carbon dioxide, methane, ammonia, and
hydrogen.
As Earth cooled, the water vapor condensed into
oceans, and much of the hydrogen escaped into
space
Stage 1-Organic Monomers
Abiotic Synthesis: is the process of chemical evolution forming organic molecules from
inorganic materials.
★ Oparin-Haldane hypothesis (primordial soup hypothesis): Russian chemist A. I. Oparin
and British scientist J. B. S. Haldane independently hypothesized that Earth’s early

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 atmosphere was a reducing (electronadding) environment, in which organic
compounds could have formed from simpler molecules.
★ Miller-Urey Experiment: Stanley Miller, working with Harold Urey, tested the Oparin-
Haldane hypothesis by creating laboratory conditions comparable to those that
scientists at the time thought existed on early Earth. His apparatus yielded a variety of
amino acids found in organisms today, along with other organic compounds.
★ Another hypothesis is that organic compounds were first produced in deep-sea
hydrothermal vents, areas on the seafloor where heated water and minerals gush
from Earth’s interior into the ocean. Some of these vents release water so hot (300–
400°C) that organic compounds formed there may have been unstable.
★ Another source of organic molecules may have been meteorites.
Stage 2-Organic Polymers: Abiotic Synthesis of Macromolecules
The abiotic synthesis of RNA monomers can occur
spontaneously from simple precursor molecules.
Amino acids polymerize when exposed to dry heat,
heat from the sun could polymerize small
polypeptides called proteinoids which have catalytic
activity.
Stage 3- Protocells
Life is defined by two properties: accurate
replication and metabolism. Neither property can
exist without the other. Self-replicating molecules
and a metabolism-like source of the building blocks
must have appeared together.
The necessary conditions may have been provided
by protobionts.

Before the first true cell arose, a protocell (protobiont), would have emerged—a structure that
is characterized by having an outer membrane. After all, life requires chemical reactions to
take place within a boundary, protecting them from disruption of conditions.
The first membranes were likely made of fatty acids.
The first membrane hypothesis: lipids tend to organize themselves into doubled layered
bubbles when placed in water in the size of cells called liposomes. Perhaps liposomes with a
phospholipid membrane engulfed early molecules that had enzymatic, even replicative,

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abilities. The liposomes would have protected the molecules from their surroundings and
concentrated them, so that they could react (and evolve) quickly and efficiently.
Stage 4: Evolution of self replicating system
The first genetic material was most likely RNA, not DNA. Ribozymes are RNA molecules which
are responsible for synthesizing proteins. Natural selection on the molecular level has
produced ribozymes capable of self-replication.
Occasional copying errors create mutations; ; selection screens these mutations for the most
stable or the best at self-replication. Its descendents will be a family of closely related RNA
sequences, differing due to copying errors. Some copying errors will result in molecules that
are more stable or more capable of self-replication.
A protobiont with self-replicating, catalytic RNA could grow, split, and pass on its RNA
molecules to its daughters.
Double-stranded DNA is a more stable molecule, and it can be replicated more accurately.
Once DNA appeared, RNA molecules began to take on their modern roles as intermediates in
the translation of genetic programs.

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18
The Fossil Record
Sedimentary rocks are the richest source of fossils

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fossils have accumulated in sedimentary rock layers, called strata. While the order of fossils in
rock strata tells us the sequence in which the fossils were laid down—their relative ages—it
does not tell us their actual (absolute) ages.
Index fossil Index fossils are the fossils that are commonly found as well as widely distributed
and are the fossils limited in the period. These are helpful in the determination of the age of
organic rocks as well as other fossil assemblages.
Radiometric dating, which is based on the decay of radioactive isotopes. In this process, a
radioactive “parent” isotope decays to a “daughter” isotope at a characteristic rate.
The rate of decay is expressed by the half-life, the time required for 50% of the parent isotope
to decay. Each type of radioactive isotope has a characteristic half-life, which is not affected
by temperature, pressure, or other environmental variables. For example, carbon-14 decays
relatively quickly; its half-life is 5,730 years. Uranium-238 decays slowly; its half-life is 4.5
billion years.

When the organism dies, it stops accumulating carbon, and the amount of carbon-12 in its
tissues does not change over time. However, the carbon-14 that it contains at the time of
death slowly decays into another element, nitrogen-14. Thus, by measuring the ratio of
carbon-14 to carbon-12 in a fossil, we can determine the fossil’s age. This method works
for fossils up to about 75,000 years old; fossils older than that contain too little carbon-14 to
be detected with current techniques. Radioactive isotopes with longer half-lives are used to
date older fossils.
The sedimentary rocks in which fossils are found tend to be composed of sediments of
differing ages. Paleontologists can determine the absolute age of fossils sandwiched between
layers of volcanic rock.
● Example: paleontologists might measure the amount of potassium-40 in those
layers. Potassium-40 has a half-life of 1.3 billion years old. If the two surrounding
rock layers were determined to be 525 and 535 million years old, the fossils likely
represent organisms that lived about 530 million years ago.
Fossils provide evidence of the origin of new groups of organisms. Along with amphibians
and reptiles, mammals are considered tetrapods, with four limbs. Mammals with unique
anatomical features that fossilize readily have a good fossil record.
● The lower jaw of most tetrapods is composed of several bones. In mammals, each
side is composed of a single dentary bone.
● The hinge between the upper and lower jaws is composed of a different set of
bones in mammals and other tetrapods.

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 ● Mammals also have a unique set of three bones to transmit sound in the middle
ear,compared with one such bone in other tetrapods.
● Finally, mammalian teeth are differentiated into incisors, canines, and cusped
molars.
Key events in life’s history

● The oldest known fossils are 3.5-billion-year-old stromatolites, rocklike structures

composed of layers of cyanobacteria and sediment.
● Prokaryotes dominated evolutionary history from 3.5 to 2.0 billion years ago.
● Most atmospheric O2 is of biological origin, from the water-splitting step of
photosynthesis.
● When O2 reached a high enough concentration it dissolved in the seas and lakes
until they were saturated with O2.
● After this, the O2 finally began to “gas out” of water and enter the atmosphere
The increase in atmospheric oxygen likely doomed many prokaryote groups. Other species
evolved mechanisms to use O2 in cellular respiration, which uses oxygen to help harvest the
energy stored in organic molecules.
A process called endosymbiosis probably led to mitochondria and plastids (the general term
for chloroplasts and related organelles). The endosymbiotic hypothesis suggests that
mitochondria and plastids were formerly small prokaryotes that began living within larger
cells. As they became increasingly interdependent, the host and endosymbionts became a
single organism. The hypothesis of serial endosymbiosis supposes that mitochondria evolved
before plastids.

A great deal of evidence supports an endosymbiotic origin of plastids and mitochondria.

○ The inner membranes of both organelles have enzymes and transport systems that are homologous
to those in the plasma membranes of modern prokaryotes.
○ Both mitochondria and plastids replicate by a splitting process similar to prokaryotic binary fission.
○ Like prokaryotes, each organelle has a single, circular DNA molecule that is not associated with
histone.
○ These organelles contain tRNAs, ribosomes, and other molecules needed to transcribe and translate
their DNA into protein.
○ Ribosomes of mitochondria and plastids are similar to prokaryotic ribosomes in size, nucleotide
sequence, and sensitivity to antibiotics

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A great range of eukaryotic unicellular forms evolved to create the diversity of present-day
unicellular forms, The oldest known fossils of multicellular eukaryotes are relatively small
algae that lived 1.2 billion years ago.
Geologic evidence suggests that a series of severe ice ages gripped Earth from 750 to 580
million years ago.
○ At times during this period, glaciers covered all of Earth’s landmasses, and the seas were largely iced
over.
○ According to the “snowball Earth” hypothesis, life would have been confined to deep sea vents and hot
springs or to equatorial regions of the ocean that lacked ice cover. The first major
diversification of multicellular eukaryotic organisms corresponds to the time of the thawing of
snowball Earth, about 575 million years ago

Phylogeny is the evolutionary history of a species or group of species.

Organisms share genes, metabolic pathways, and structural pathways with their close
relatives (common descent).
Taxonomy is the science of naming, describing and classifying organisms.
The Linnaean system, first formally proposed by Carolus Linnaeus in the 18th century, has
two main characteristics.
1. Each species has a two-part name.
2. Species are organized hierarchically into broader and broader groups of organisms.
Under the binomial system, each species is assigned a two-part Latinized name, a binomial.
● The first part of the name, the genus, is the closest group to which a species
belongs.
● The second part, the specific epithet, refers to one species within each genus.
● The first letter of the genus name is capitalized, and both the genus and species
names are italicized and Latinized.
A hierarchical classification groups species into increasingly inclusive taxonomic categories.
Species that appear to be closely related are grouped into the same genus.
Genera are grouped into progressively broader categories: family, order, class, phylum,
kingdom, and domain.
Each taxonomic level is more comprehensive than the previous one. The named taxonomic
unit at any level is called a taxon.

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The evolutionary history of a group of organisms can be represented in a diagram called a
phylogenetic tree.
The difficulty of aligning Linnaean classification with phylogeny has led systematists to
propose that classification be based entirely on evolutionary relationships.
● Evolutionary relationships are often represented as a series of dichotomies
showing the divergence of two evolutionary lineages from a common ancestor.
● Sister taxa are groups of organisms that share an immediate common ancestor.
● A rooted tree includes the last common ancestor to all taxa in the tree.
● The sequence of branching in a tree reflects patterns of descent and does not
indicate the absolute ages of particular species. 2. A taxon in a phylogenetic tree
did not evolve from an adjacent taxon. Rather, both taxa evolved from a common
ancestor.
Biologists place species into groups called clades, each of which includes an ancestral species
and all of its descendents. A valid clade is monophyletic, consisting of an ancestral species
and all its descendents.
● A character is any feature that a particular taxon possesses.
● A shared derived character is unique to a particular clade.
● A shared ancestral character is found not only in the clade being analyzed but also
in older clades
● The species making up the ingroup display a mixture of shared ancestral and
shared derived characters.

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