21: INTRODUCTION TO

EVOLUTION
 CHAPTER OVERVIEW

21: Introduction to Evolution

The theory of evolution is the unifying theory of biology, meaning it is the framework within which biologists ask questions about
the living world. Its power is that it provides direction for predictions about living things that are borne out in experiment after
experiment. The Ukrainian-born American geneticist Theodosius Dobzhansky famously wrote that “nothing makes sense in
biology except in the light of evolution” (Dobzhansky, 1964). He meant that the tenet that all life has evolved and diversified from
a common ancestor is the foundation from which we approach all questions in biology.
Evolution by natural selection describes a mechanism for how species change over time. That species change had been suggested
and debated well before Darwin began to explore this idea. The view that species were static and unchanging was grounded in the
writings of Plato, yet there were also ancient Greeks who expressed evolutionary ideas. In the eighteenth century, ideas about the
evolution of animals were reintroduced by the naturalist Georges-Louis Leclerc Comte de Buffon who observed that various
geographic regions have different plant and animal populations, even when the environments are similar. It was also accepted that
there were extinct species.
During this time, James Hutton, a Scottish naturalist, proposed that geological change occurred gradually by the accumulation of
small changes from processes operating like they are today over long periods of time. This contrasted with the predominant view
that the geology of the planet was a consequence of catastrophic events occurring during a relatively brief past. Hutton’s view was
popularized in the nineteenth century by the geologist Charles Lyell who became a friend to Darwin. Lyell’s ideas were influential
on Darwin’s thinking: Lyell’s notion of the greater age of Earth gave more time for gradual change in species, and the process of
change provided an analogy for gradual change in species. In the early nineteenth century, Jean-Baptiste Lamarck published a book
that detailed a mechanism for evolutionary change. This mechanism is now referred to as an inheritance of acquired
characteristics by which modifications in an individual are caused by its environment, or the use or disuse of a structure during its
lifetime, could be inherited by its offspring and thus bring about change in a species. While this mechanism for evolutionary
change was discredited, Lamarck’s ideas were an important influence on evolutionary thought.

Charles Darwin and Natural Selection

In the mid-nineteenth century, the actual mechanism for evolution was independently conceived of and described by two
naturalists: Charles Darwin and Alfred Russel Wallace. Importantly, each naturalist spent time exploring the natural world on
expeditions to the tropics. From 1831 to 1836, Darwin traveled around the world on H.M.S. Beagle, including stops in South
America, Australia, and the southern tip of Africa. Wallace traveled to Brazil to collect insects in the Amazon rainforest from 1848
to 1852 and to the Malay Archipelago from 1854 to 1862. Darwin’s journey, like Wallace’s later journeys to the Malay
Archipelago, included stops at several island chains, the last being the Galápagos Islands west of Ecuador. On these islands,
Darwin observed species of organisms on different islands that were clearly similar, yet had distinct differences. For example, the
ground finches inhabiting the Galápagos Islands comprised several species with a unique beak shape (Figure 21.1). The species on
the islands had a graded series of beak sizes and shapes with very small differences between the most similar. He observed that
these finches closely resembled another finch species on the mainland of South America. Darwin imagined that the island species
might be species modified from one of the original mainland species. Upon further study, he realized that the varied beaks of each
finch helped the birds acquire a specific type of food. For example, seed-eating finches had stronger, thicker beaks for breaking
seeds, and insect-eating finches had spear-like beaks for stabbing their prey.

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 Figure 21.1 : Darwin observed that beak shape varies among finch species. He postulated that the beak of an ancestral species had
adapted over time to equip the finches to acquire different food sources. (“Galápagos Island Finches” by OpenStax is licensed
under CC BY 4.0)
Wallace and Darwin both observed similar patterns in other organisms and they independently developed the same explanation for
how and why such changes could take place. Darwin called this mechanism natural selection. Natural selection, also known as
“survival of the fittest,” is the more prolific reproduction of individuals with favorable traits that survive environmental change
because of those traits; this leads to evolutionary change.
For example, a population of giant tortoises found in the Galapagos Archipelago was observed by Darwin to have longer necks
than those that lived on other islands with dry lowlands. These tortoises were “selected” because they could reach more leaves and
access more food than those with short necks. In times of drought when fewer leaves would be available, those that could reach
more leaves had a better chance to eat and survive than those that couldn’t reach the food source. Consequently, long-necked
tortoises would be more likely to be reproductively successful and pass the long-necked trait to their offspring. Over time, only
long-necked tortoises would be present in the population.
Natural selection, Darwin argued, was an inevitable outcome of three principles that operated in nature. First, most characteristics
of organisms are inherited, or passed from parent to offspring. Although no one, including Darwin and Wallace, knew how this
happened at the time, it was a common understanding. Second, more offspring are produced than are able to survive, so resources
for survival and reproduction are limited. The capacity for reproduction in all organisms outstrips the availability of resources to
support their numbers. Thus, there is competition for those resources in each generation. Both Darwin and Wallace’s understanding
of this principle came from reading an essay by the economist Thomas Malthus who discussed this principle in relation to human
populations. Third, offspring vary among each other in regard to their characteristics and those variations are inherited. Darwin and
Wallace reasoned that offspring with inherited characteristics that allow them to best compete for limited resources will survive and
have more offspring than those individuals with variations that are less able to compete. Because characteristics are inherited, these
traits will be better represented in the next generation. This will lead to change in populations over generations in a process that
Darwin called descent with modification. Ultimately, natural selection leads to greater adaptation of the population to its local
environment; it is the only mechanism known for adaptive evolution.
Papers by Darwin and Wallace presenting the idea of natural selection were read together in 1858 before the Linnean Society in
London. The following year Darwin’s book, On the Origin of Species, was published. His book outlined in considerable detail his
arguments for evolution by natural selection. It’s important to note that Darwin is a deeply problematic figure and that many of
his contributions to science are remembered because of his privilege and power. Darwin had many racist and offensive beliefs, and
the fact that he is often celebrated in biology classes is an example of systematic racism in the classroom. Scientists and educators
need to be doing a better job of recognizing prejudice in our materials and highlighting ways to promote racial equity.
Demonstrations of evolution by natural selection are time-consuming and difficult to obtain. One of the best examples has been
demonstrated in the very birds that helped to inspire Darwin’s theory: the Galápagos finches. Peter and Rosemary Grant and their
colleagues have studied Galápagos finch populations every year since 1976 and have provided important demonstrations of natural
selection. The Grants found changes from one generation to the next in the distribution of beak shapes with the medium ground
finch on the Galápagos island of Daphne Major. The birds have inherited variation in the bill shape with some birds having wide
deep bills and others having thinner bills. During a period in which rainfall was higher than normal because of an El Niño, the large
hard seeds that large-billed birds ate were reduced in number; however, there was an abundance of the small soft seeds which the
small-billed birds ate. Therefore, survival and reproduction were much better in the following years for the small-billed birds. In
the years following this El Niño, the Grants measured beak sizes in the population and found that the average bill size was smaller.
Since bill size is an inherited trait, parents with smaller bills had more offspring and the size of bills had evolved to be smaller. As
conditions improved in 1987 and larger seeds became more available, the trend toward smaller average bill size ceased.

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The book The Beak of the Finch: A Story of Evolution in Our Time by Jonathan Weiner is a wonderful exploration of Peter and
Rosemary Grant’s work. If you want to learn more, I highly recommend checking it out.

Processes and Patterns of Evolution

Natural selection can only take place if there is variation, or differences, among individuals in a population. Importantly, these
differences must have some genetic basis; otherwise, the selection will not lead to change in the next generation. This is critical
because variation among individuals can be caused by non-genetic reasons such as an individual being taller because of better
nutrition rather than different genes.
Genetic diversity in a population comes from two main mechanisms: mutation and sexual reproduction. Mutation, a change in
DNA, is the ultimate source of new alleles, or new genetic variation in any population. The genetic changes caused by mutation can
have one of three outcomes on the phenotype. A mutation affects the phenotype of the organism in a way that gives it reduced
fitness—lower likelihood of survival or fewer offspring. A mutation may produce a phenotype with a beneficial effect on fitness.
And, many mutations will also have no effect on the fitness of the phenotype; these are called neutral mutations. Mutations may
also have a whole range of effect sizes on the fitness of the organism that expresses them in their phenotype, from a small effect to
a great effect. Sexual reproduction also leads to genetic diversity: when two parents reproduce, unique combinations of alleles
assemble to produce the unique genotypes and thus phenotypes in each of the offspring.
A heritable trait that helps the survival and reproduction of an organism in its present environment is called an adaptation.
Scientists describe groups of organisms becoming adapted to their environment when a change in the range of genetic variation
occurs over time that increases or maintains the “fit” of the population to its environment. The webbed feet of platypuses are an
adaptation for swimming. The snow leopards’ thick fur is an adaptation for living in the cold. The cheetahs’ fast speed is an
adaptation for catching prey.
Whether or not a trait is favorable depends on the environmental conditions at the time. The same traits are not always selected
because environmental conditions can change. For example, consider a species of plant that grew in a moist climate and did not
need to conserve water. Large leaves were selected because they allowed the plant to obtain more energy from the sun. Large
leaves require more water to maintain than small leaves, and the moist environment provided favorable conditions to support large
leaves. After thousands of years, the climate changed, and the area no longer had excess water. The direction of natural selection
shifted so that plants with small leaves were selected because those populations were able to conserve water to survive the new
environmental conditions.
The evolution of species has resulted in enormous variation in form and function. Sometimes, evolution gives rise to groups of
organisms that become tremendously different from each other. When two species evolve in diverse directions from a common
point, it is called divergent evolution.
In other cases, similar phenotypes evolve independently in distantly related species. For example, flight has evolved in both bats
and insects, and they both have structures we refer to as wings, which are adaptations to flight. However, the wings of bats and
insects have evolved from very different original structures. This phenomenon is called convergent evolution, where similar traits
evolve independently in species that do not share a common ancestry. The two species came to the same function, flying, but did so
separately from each other.
These physical changes occur over enormous spans of time and help explain how evolution occurs. Natural selection acts on
individual organisms, which in turn can shape an entire species. Although natural selection may work in a single generation on an
individual, it can take thousands or even millions of years for the genotype of an entire species to evolve. It is over these large time
spans that life on earth has changed and continues to change.
21.1: Evidence of Evolution
21.2: Formation of New Species
21.3: Speciation

References
Dobzhansky, T. 1964. “Biology, Molecular and Organismic.” American Zoologist 4, no. 4: 449.
OpenStax, Biology. OpenStax CNX. June 23, 2020. [Link]
Evolution.

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Thumbnail: A silhouette of human evolution. (CC BY-SA 3.0; Tkgd2007 via Wikimedia Commons).

21: Introduction to Evolution is shared under a CC BY license and was authored, remixed, and/or curated by LibreTexts.

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21.1: Evidence of Evolution
The evidence for evolution is compelling and extensive. Looking at every level of organization in living systems, biologists see the
signature of past and present evolution. Darwin dedicated a large portion of his book, On the Origin of Species, to identifying
patterns in nature that were consistent with evolution, and since Darwin, our understanding has become clearer and broader.

Fossils
Fossils provide solid evidence that organisms from the past are not the same as those found today, and fossils show a progression
of evolution. Scientists determine the age of fossils and categorize them from all over the world to determine when the organisms
lived relative to each other. The resulting fossil record tells the story of the past and shows the evolution of form over millions of
years. For example, scientists have recovered highly detailed records showing the evolution of humans and horses (Figure 21.1.1).

Figure 21.1.1: In this (a) display, fossil hominids are arranged from oldest (bottom) to newest (top). As hominids evolved, the
shape of the skull changed. An artist’s rendition of (b) extinct species of the genus Equus reveals that these ancient species
resembled the modern horse (Equus ferus) but varied in size. (credit: “Fossil evidence” by OpenStax is licensed under CC BY 4.0)

Anatomy and Embryology

Another type of evidence for evolution is the presence of structures in organisms that share the same basic form. For example, the
bones in the appendages of a human, dog, bird, and whale all share the same overall construction (Figure 21.1.2) resulting from
their origin in the appendages of a common ancestor. Over time, evolution led to changes in the shapes and sizes of these bones in
different species, but they have maintained the same overall layout. Scientists call these synonymous parts homologous structures.

Figure 21.1.2: The similar construction of these appendages indicates that these organisms share a common ancestor. (Credit:
“homologous structures” by OpenStax is licensed under CC BY 4.0)
Some structures exist in organisms that have no apparent function at all and appear to be residual parts from a past common
ancestor. These unused structures without function are called vestigial structures. Other examples of vestigial structures are wings
on flightless birds, leaves on some cacti, and hind leg bones in whales.
Another evidence of evolution is the convergence of form in organisms that share similar environments. For example, species of
unrelated animals, such as the arctic fox and ptarmigan, living in the arctic region have been selected for seasonal white phenotypes
during winter to blend with the snow and ice (Figure 21.1.3). These similarities occur not because of common ancestry, but
because of similar selection pressures—the benefits of not being seen by predators.

21.1.1 [Link]
 Figure 21.1.3: The white winter coat of the (a) arctic fox and the (b) ptarmigan’s plumage are adaptations to their environments.
(Credit: “winter coats” by OpenStax is licensed under CC BY 4.0. Figure a: modification of work by Keith Morehouse)
Embryology, the study of the development of the anatomy of an organism to its adult form, also provides evidence of relatedness
between now widely divergent groups of organisms. Mutational tweaking in the embryo can have such magnified consequences in
the adult that embryo formation tends to be conserved. As a result, structures that are absent in some groups often appear in their
embryonic forms and disappear by the time the adult or juvenile form is reached. For example, all vertebrate embryos, including
humans, exhibit gill slits and tails at some point in their early development. These disappear in the adults of terrestrial groups but
are maintained in adult forms of aquatic groups such as fish and some amphibians. Great ape embryos, including humans, have a
tail structure during their development that is lost by the time of birth.

Biogeography
The geographic distribution of organisms (referred to as biogeography) on the planet follows patterns that are best explained by
evolution in conjunction with the movement of tectonic plates over geological time. Broad groups that evolved before the breakup
of the supercontinent Pangaea (about 200 million years ago) are distributed worldwide. Groups that evolved since the breakup
appear uniquely in regions of the planet, such as the unique flora and fauna of northern continents that formed from the
supercontinent Laurasia and of the southern continents that formed from the supercontinent Gondwana. The presence of members
of the plant family Proteaceae in Australia, southern Africa, and South America is best explained by their presence prior to the
southern supercontinent Gondwana breaking up.
The great diversification of marsupials in Australia and the absence of other mammals reflect Australia’s long isolation. Australia
has an abundance of endemic species—species found nowhere else—which is typical of islands whose isolation by expanses of
water prevents species from migrating. Over time, these species diverge evolutionarily into new species that look very different
from their ancestors that may exist on the mainland. The marsupials of Australia, the finches on the Galápagos, and many species
on the Hawaiian Islands are all unique to their one point of origin, yet they display distant relationships to ancestral species on
mainlands.

Molecular Biology
Like anatomical structures, the structures of the molecules of life reflect descent with modification. Evidence of a common ancestor
for all of life is reflected in the universality of DNA as the genetic material and in the near universality of the genetic code and the
machinery of DNA replication and expression. Fundamental divisions in life between the three domains are reflected in major
structural differences in otherwise conservative structures such as the components of ribosomes and the structures of membranes.
In general, the relatedness of groups of organisms is reflected in the similarity of their DNA sequences—exactly the pattern that
would be expected from descent and diversification from a common ancestor.
DNA sequences have also shed light on some of the mechanisms of evolution. For example, it is clear that the evolution of new
functions for proteins commonly occurs after gene duplication events that allow the free modification of one copy by mutation,
selection, or drift (changes in a population’s gene pool resulting from chance), while the second copy continues to produce a
functional protein.

Misconceptions of Evolution
Although the theory of evolution generated some controversy when it was first proposed, it was almost universally accepted by
biologists, particularly younger biologists, within 20 years after the publication of On the Origin of Species. Nevertheless, the
theory of evolution is a difficult concept, and misconceptions about how it works abound.

21.1.2 [Link]
Evolution Is Just a Theory
Critics of the theory of evolution dismiss its importance by purposefully confounding the everyday usage of the word “theory”
with the way scientists use the word. In science, a “theory” is understood to be a body of thoroughly tested and verified
explanations for a set of observations of the natural world. Scientists have a theory of the atom, a theory of gravity, and the theory
of relativity, each of which describes understood facts about the world. In the same way, the theory of evolution describes facts
about the living world. As such, a theory in science has survived significant efforts to discredit it by scientists. In contrast, a
“theory” in common vernacular is a word meaning a guess or suggested explanation; this meaning is more akin to the scientific
concept of “hypothesis.” When critics of evolution say evolution is “just a theory,” they are implying that there is little evidence
supporting it and that it is still in the process of being rigorously tested. This is a mischaracterization.

Individuals Evolve
Evolution is the change in the genetic composition of a population over time, specifically over generations, resulting from
differential reproduction of individuals with certain alleles. Individuals do change over their lifetime, obviously, but this is called
development and involves changes programmed by the set of genes the individual acquired at birth in coordination with the
individual’s environment. When thinking about the evolution of a characteristic, it is probably best to think about the change of the
average value of the characteristic in the population over time. For example, when natural selection leads to a beak size change in
medium-ground finches in the Galápagos, this does not mean that individual beaks on living finches are changing. Instead, it means
that if one measures the average beak size among all individuals in the population at one time and then measures the average beak
size in the population several years later, the average value will be different as a result of evolution. Although some individuals
may survive from the first time to the second, they will still have the same beak size; however, there will be many new individuals
that contribute to the shift in average beak size.

Evolution Explains the Origin of Life

It is a common misunderstanding that evolution includes an explanation of life’s origins. The theory does not try to explain the
origin of life. The theory of evolution explains how populations change over time and how life diversifies. It does not shed light on
the beginnings of life including the origins of the first cells, which is how life is defined. The mechanisms of the origin of life on
Earth are a particularly difficult problem because it occurred a very long time ago, and presumably, it just occurred once.
However, once a mechanism of inheritance was in place in the form of a molecule like DNA either within a cell or pre-cell, these
entities would be subject to the principle of natural selection. More effective reproducers would increase in frequency at the
expense of inefficient reproducers. So while evolution does not explain the origin of life, it may have something to say about some
of the processes operating once pre-living entities acquired certain properties.

Organisms Evolve on Purpose

Statements such as “organisms evolve in response to a change in an environment” are quite common, but such statements can lead
to two types of misunderstandings. First, the statement must not be understood to mean that individual organisms evolve. The
statement is shorthand for “a population evolves in response to a changing environment.” However, a second misunderstanding
may arise by interpreting the statement to mean that evolution is somehow intentional. A changed environment results in some
individuals in the population, those with particular phenotypes, benefiting and therefore producing proportionately more offspring
than other phenotypes. This results in a change in the population if the characteristics are genetically determined.
It is also important to understand that the variation that natural selection works on is already in a population and does not arise in
response to an environmental change. For example, applying antibiotics to a population of bacteria will, over time, select a
population of bacteria that are resistant to antibiotics. The resistance, which is caused by a gene, did not arise by mutation because
of the application of the antibiotic. The gene for resistance was already present in the gene pool of the bacteria, likely at a low
frequency. The antibiotic, which kills the bacterial cells without the resistance gene, strongly selects individuals that are resistant,
since these would be the only ones that survived and divided. Experiments have demonstrated that mutations for antibiotic
resistance do not arise as a result of antibiotics.
In a larger sense, evolution is not goal-directed. Species do not become “better” over time; they simply track their changing
environment with adaptations that maximize their reproduction in a particular environment at a particular time. Evolution has no
goal of making faster, bigger, more complex, or even smarter species, despite the commonness of this kind of language in popular
discourse. What characteristics evolve in a species are a function of the variation present and the environment, both of which are

21.1.3 [Link]
constantly changing in a non-directional way. What trait is fit in one environment at one time may well be fatal at some point in the
future. This holds equally well for a species of insect as it does the human species.

Summary
Evolution is the process of adaptation through mutation which allows more desirable characteristics to be passed to the next
generation. Over time, organisms evolve more characteristics that are beneficial to their survival. For living organisms to adapt and
change to environmental pressures, genetic variation must be present. With genetic variation, individuals have differences in form
and function that allow some to survive certain conditions better than others. These organisms pass their favorable traits to their
offspring. Eventually, environments change, and what was once a desirable, advantageous trait may become an undesirable trait
and organisms may further evolve. Evolution may be convergent with similar traits evolving in multiple species or divergent with
diverse traits evolving in multiple species that came from a common ancestor. Evidence of evolution can be observed by means of
DNA code and the fossil record, and also by the existence of homologous and vestigial structures.

References
OpenStax, Biology. OpenStax CNX. June 23, 2020. [Link]
Evolution.

21.1: Evidence of Evolution is shared under a CC BY license and was authored, remixed, and/or curated by LibreTexts.

21.1.4 [Link]
21.2: Formation of New Species
Although all life on earth shares various genetic similarities, only certain organisms combine genetic information by sexual
reproduction and have offspring that can then successfully reproduce. Scientists call such organisms members of the same
biological species.

Species Concepts
It turns out that scientists don’t always agree on the definition of a species. The different ideas about what does and does not
constitute a species are referred to as species concepts. There are around twenty-six different species concepts, but four are the
most accepted. The biological species concept states that if two organisms are able to successfully breed and produce viable, fertile
offspring, then they are the same species. When populations of organisms cease to successfully breed (called reproductive
isolation) they are then considered separate species. The biological species concept works well for scientists studying living
creatures that have regular breeding patterns, such as insects or mammals. The morphological species concept states that if two
organisms are morphologically similar enough, then they are the same species. There is no guideline about how similar counts as
similar enough, so it is up to the researcher to make the judgment call. This concept works well for organisms that don’t breed
regularly (such as fungi) or that are no longer living. The genetic species concept states that if two organisms are genetically
similar enough, then they are the same species. Again, there is no rule about how genetically similar they need to be, so each
discipline determines its own limits. This concept works well for organisms that are very tiny, have unusual reproduction strategies,
and are not morphologically distinct (such as bacteria). The evolutionary species concept states that if two organisms’
evolutionary paths are similar enough, then they are the same species. This concept works well for long-dead organisms because
fossils cannot breed, often are lacking impressions of soft tissues, and usually don’t have enough DNA left to work with.

Species and the Ability to Reproduce

A species is a group of individual organisms that interbreed and produce fertile, viable offspring. According to this definition, one
species is distinguished from another when, in nature, it is not possible for matings between individuals from each species to
produce fertile offspring.
Members of the same species share both external and internal characteristics, which develop from their DNA. The closer
relationship two organisms share, the more DNA they have in common, just like people and their families. People’s DNA is likely
to be more like their father or mother’s DNA than their cousin or grandparent’s DNA. Organisms of the same species have the
highest level of DNA alignment and therefore share characteristics and behaviors that lead to successful reproduction.
Species’ appearance can be misleading in suggesting an ability or inability to mate. For example, even though domestic dogs
(Canis lupus familiaris) display phenotypic differences, such as size, build, and coat, most dogs can interbreed and produce viable
puppies that can mature and sexually reproduce (Figure 21.2.1).

Figure 21.2.1: The (a) poodle and (b) cocker spaniel can reproduce to produce a breed known as (c) the cockapoo. (credit a:
modification of work by Sally Eller, Tom Reese; credit b: modification of work by Jeremy McWilliams; credit c: modification of
work by Kathleen Conklin. This image by OpenStax is licensed under CC BY 4.0)
In other cases, individuals may appear similar although they are not members of the same species. For example, even though bald
eagles (Haliaeetus leucocephalus) and African fish eagles (Haliaeetus vocifer) are both birds and eagles, each belongs to a separate
species group (Figure 21.2.2). If humans were to artificially intervene and fertilize the egg of a bald eagle with the sperm of an
African fish eagle and a chick did hatch, that offspring, called a hybrid (a cross between two species), would probably be infertile
—unable to successfully reproduce after it reached maturity. Different species may have different genes that are active in

21.2.1 [Link]
development; therefore, it may not be possible to develop a viable offspring with two different sets of directions. Thus, even though
hybridization may take place, the two species still remain separate.

Figure 21.2.2: The (a) African fish eagle is similar in appearance to the (b) bald eagle, but the two birds are members of different
species. (credit a: modification of work by Nigel Wedge; credit b: modification of work by U.S. Fish and Wildlife Service. This
image by OpenStax is licensed under CC BY 4.0 )
Populations of species share a gene pool: a collection of all the variants of genes in the species. Again, the basis for any changes in
a group or population of organisms must be genetic for this is the only way to share and pass on traits. When variations occur
within a species, they can only be passed to the next generation along two main pathways: asexual reproduction or sexual
reproduction. The change will be passed on asexually simply if the reproducing cell possesses the changed trait. For the changed
trait to be passed on by sexual reproduction, a gamete, such as a sperm or egg cell, must possess the changed trait. In other words,
sexually-reproducing organisms can experience several genetic changes in their body cells, but if these changes do not occur in a
sperm or egg cell, the changed trait will never reach the next generation. Only heritable traits can evolve. Therefore, reproduction
plays a paramount role for genetic change to take root in a population or species. In short, organisms must be able to reproduce
with each other to pass new traits to offspring.

Speciation
The biological definition of species, which works for sexually reproducing organisms, is a group of actually or potentially
interbreeding individuals. There are exceptions to this rule. Many species are similar enough that hybrid offspring are possible and
may often occur in nature, but for the majority of species, this rule generally holds. In fact, the presence in nature of hybrids
between similar species suggests that they may have descended from a single interbreeding species, and the speciation process may
not yet be completed.
Given the extraordinary diversity of life on the planet, there must be mechanisms for speciation: the formation of two species from
one original species. Darwin envisioned this process as a branching event and diagrammed the process in the only illustration found
in On the Origin of Species (Figure 21.2.3a). Compare this illustration to the diagram of elephant evolution (Figure 21.2.3b),
which shows that as one species changes over time, it branches to form more than one new species, repeatedly, as long as the
population survives or until the organism becomes extinct.

21.2.2 [Link]
 Figure 21.2.3: The only illustration in Darwin’s On the Origin of Species is (a) a diagram showing speciation events leading to
biological diversity. The diagram shows similarities to phylogenetic charts that are drawn today to illustrate the relationships of
species. (b) Modern elephants evolved from the Palaeomastodon, a species that lived in Egypt 35–50 million years ago. (credit: this
image by OpenStax is licensed under CC BY 4.0)
For speciation to occur, two new populations must be formed from one original population and they must evolve in such a way that
it becomes impossible for individuals from the two new populations to interbreed. Biologists have proposed mechanisms by which
this could occur that fall into two broad categories. Allopatric speciation (allo- = “other”; -patric = “homeland”) involves
geographic separation of populations from a parent species and subsequent evolution. Sympatric speciation (sym- = “same”; -
patric = “homeland”) involves speciation occurring within a parent species remaining in one location.
Biologists think of speciation events as the splitting of one ancestral species into two descendant species. There is no reason why
there might not be more than two species formed at one time except that it is less likely and multiple events can be conceptualized
as single splits occurring close in time.

Allopatric Speciation
A geographically continuous population has a gene pool that is relatively homogeneous. Gene flow, the movement of alleles across
the range of the species, is relatively free because individuals can move and then mate with individuals in their new location. Thus,
the frequency of an allele at one end of a distribution will be similar to the frequency of the allele at the other end. When
populations become geographically discontinuous, that free-flow of alleles is prevented. When that separation lasts for a period of
time, the two populations are able to evolve along different trajectories. Thus, their allele frequencies at numerous genetic loci
gradually become more and more different as new alleles independently arise by mutation in each population. Typically,
environmental conditions, such as climate, resources, predators, and competitors for the two populations will differ causing natural
selection to favor divergent adaptations in each group.
Isolation of populations leading to allopatric speciation can occur in a variety of ways: a river forming a new branch, erosion
forming a new valley, a group of organisms traveling to a new location without the ability to return, or seeds floating over the ocean
to an island. The nature of the geographic separation necessary to isolate populations depends entirely on the biology of the
organism and its potential for dispersal. If two flying insect populations took up residence in separate nearby valleys, chances are,
individuals from each population would fly back and forth continuing gene flow. However, if two rodent populations became
divided by the formation of a new lake, continued gene flow would be unlikely; therefore, speciation would be more likely.
Biologists group allopatric processes into two categories: dispersal and vicariance. Dispersal is when a few members of a species
move to a new geographical area, and vicariance is when a natural situation arises to physically divide organisms.
Scientists have documented numerous cases of allopatric speciation taking place. For example, along the west coast of the United
States, two separate sub-species of spotted owls exist. The northern spotted owl has genetic and phenotypic differences from its
close relative: the Mexican spotted owl, which lives in the south (Figure 21.2.4).

21.2.3 [Link]
 Figure 21.2.4: The northern spotted owl and the Mexican spotted owl inhabit geographically separate locations with different
climates and ecosystems. The owl is an example of allopatric speciation. (credit “northern spotted owl”: modification of work by
John and Karen Hollingsworth; credit “Mexican spotted owl”: modification of work by Bill Radke. “spotted owls” by OpenStax is
licensed under CC BY 4.0)
Additionally, scientists have found that the further the distance between two groups that once were the same species, the more
likely it is that speciation will occur. This seems logical because as the distance increases, the various environmental factors would
likely have less in common than locations in close proximity. Consider the two owls: in the north, the climate is cooler than in the
south; the types of organisms in each ecosystem differ, as do their behaviors and habits; also, the hunting habits and prey choices of
the southern owls vary from the northern owls. These variances can lead to evolved differences in the owls, and speciation likely
will occur.

Adaptive Radiation
In some cases, a population of one species disperses throughout an area, and each finds a distinct niche or isolated habitat. Over
time, the varied demands of their new lifestyles lead to multiple speciation events originating from a single species. This is called
adaptive radiation because many adaptations evolve from a single point of origin; thus, causing the species to radiate into several
new ones. Island archipelagos like the Hawaiian Islands provide an ideal context for adaptive radiation events because water
surrounds each island which leads to geographical isolation for many organisms. The Hawaiian honeycreeper illustrates one
example of adaptive radiation. From a single species, called the founder species, numerous species have evolved, including the six
shown in Figure 21.2.5.

21.2.4 [Link]
 Figure 21.2.5: The honeycreeper birds illustrate adaptive radiation. From one original species of bird, multiple others evolved,
each with its own distinctive characteristics. (credit: “honeycreepers” by OpenStax is licensed under CC BY 4.0)
Notice the differences in the species’ beaks. Evolution in response to natural selection based on specific food sources in each new
habitat led to the evolution of a different beak suited to the specific food source. The seed-eating bird has a thicker, stronger beak
which is suited to break hard nuts. The nectar-eating birds have long beaks to dip into flowers to reach the nectar. The insect-eating
birds have beaks like swords, appropriate for stabbing and impaling insects. Darwin’s finches are another example of adaptive
radiation in an archipelago.

Sympatric Speciation
Can divergence occur if no physical barriers are in place to separate individuals who continue to live and reproduce in the same
habitat? The answer is yes. The process of speciation within the same space is called sympatric speciation; the prefix “sym” means
same, so “sympatric” means “same homeland” in contrast to “allopatric” meaning “other homeland.” A number of mechanisms for
sympatric speciation have been proposed and studied.
One form of sympatric speciation can begin with a serious chromosomal error during cell division. In a normal cell division event
chromosomes replicate, pair up, and then separate so that each new cell has the same number of chromosomes. However,
sometimes the pairs separate and the end cell product has too many or too few individual chromosomes in a condition called
aneuploidy (Figure 21.2.6).

Figure 21.2.6: Aneuploidy results when the gametes have too many or too few chromosomes due to nondisjunction during
meiosis. In the example shown here, the resulting offspring will have 2n+1 or 2n-1 chromosomes. (credit: “Aneuploidy” by
OpenStax is licensed under CC BY 4.0)

21.2.5 [Link]
Polyploidy is a condition in which a cell or organism has an extra set, or sets, of chromosomes. Scientists have identified two main
types of polyploidy that can lead to reproductive isolation of an individual in the polyploidy state. Reproductive isolation is the
inability to interbreed. In some cases, a polyploid individual will have two or more complete sets of chromosomes from its own
species in a condition called autopolyploidy (Figure 21.2.7). The prefix “auto-” means “self,” so the term means multiple
chromosomes from one’s own species. Polyploidy results from an error in meiosis in which all of the chromosomes move into one
cell instead of separating.

Figure 21.2.7: Autopolyploidy results when mitosis is not followed by cytokinesis. (credit: “Autopolyploidy” by OpenStax is
licensed under CC BY 4.0)
For example, if a plant species with 2n = 6 produces autopolyploid gametes that are also diploid (2n = 6, when they should be n =
3), the gametes now have twice as many chromosomes as they should have. These new gametes will be incompatible with the
normal gametes produced by this plant species. However, they could either self-pollinate or reproduce with other autopolyploid
plants with gametes having the same diploid number. In this way, sympatric speciation can occur quickly by forming offspring with
4n called a tetraploid. These individuals would immediately be able to reproduce only with those of this new kind and not those of
the ancestral species.
The other form of polyploidy occurs when individuals of two different species reproduce to form a viable offspring called an
allopolyploid. The prefix “allo-” means “other” (recall from allopatric): therefore, an allopolyploid occurs when gametes from two
different species combine. Figure 21.2.8 illustrates one possible way an allopolyploid can form. Notice how it takes two
generations, or two reproductive acts, before the viable fertile hybrid results.

Figure 21.2.8: Alloploidy results when two species mate to produce viable offspring. In the example shown, a normal gamete
from one species fuses with a polyploid gamete from another. Two matings are necessary to produce viable offspring. (credit:
“Alloploidy” by OpenStax is licensed under CC BY 4.0)
The cultivated forms of wheat, cotton, and tobacco plants are all allopolyploids. Although polyploidy occurs occasionally in
animals, it takes place most commonly in plants. (Animals with any of the types of chromosomal aberrations described here are
unlikely to survive and produce normal offspring.) Scientists have discovered more than half of all plant species studied relate back
to a species that evolved through polyploidy. With such a high rate of polyploidy in plants, some scientists hypothesize that this
mechanism takes place more as an adaptation than as an error.

21.2.6 [Link]
Reproductive Isolation
Given enough time, the genetic and phenotypic divergence between populations will affect characters that influence reproduction:
if individuals of the two populations were to be brought together, mating would be less likely, but if mating occurred, offspring
would be non-viable or infertile. Many types of diverging characters may affect the reproductive isolation, the ability to interbreed,
of the two populations.
Reproductive isolation can take place in a variety of ways. Scientists organize them into two groups: prezygotic barriers and
postzygotic barriers. Recall that a zygote is a fertilized egg: the first cell of the development of an organism that reproduces
sexually. Therefore, a prezygotic barrier is a mechanism that blocks reproduction from taking place; this includes barriers that
prevent fertilization when organisms attempt reproduction. A postzygotic barrier occurs after zygote formation; this includes
organisms that don’t survive the embryonic stage and those that are born sterile.
Some types of prezygotic barriers prevent reproduction entirely. Many organisms only reproduce at certain times of the year, often
just annually. Differences in breeding schedules, called temporal isolation, can act as a form of reproductive isolation. For
example, two species of frogs inhabit the same area, but one reproduces from January to March, whereas the other reproduces from
March to May (Figure 21.2.9).

Figure 21.2.9: These two related frog species exhibit temporal reproductive isolation. (a) Rana aurora breeds earlier in the year
than (b) Rana boylii. (credit a: modification of work by Mark R. Jennings, USFWS; credit b: modification of work by Alessandro
Catenazzi. “frog reproductive isolation” by OpenStax is licensed under CC BY 4.0)
In some cases, populations of a species move or are moved to a new habitat and take up residence in a place that no longer overlaps
with the other populations of the same species. This situation is called habitat isolation. Reproduction with the parent species
ceases, and a new group exists that is now reproductively and genetically independent. For example, a cricket population that was
divided after a flood could no longer interact with each other. Over time, the forces of natural selection, mutation, and genetic drift
will likely result in the divergence of the two groups.
Behavioral isolation occurs when the presence or absence of a specific behavior prevents reproduction from taking place. For
example, male fireflies use specific light patterns to attract females. Various species of fireflies display their lights differently. If a
male of one species tried to attract the female of another, she would not recognize the light pattern and would not mate with the
male.
Other prezygotic barriers work when differences in their gamete cells (eggs and sperm) prevent fertilization from taking place; this
is called a gametic barrier. Similarly, in some cases closely related organisms try to mate, but their reproductive structures simply
do not fit together. For example, damselfly males of different species have differently shaped reproductive organs. If one species
tries to mate with the female of another, their body parts simply do not fit together. (Figure 21.2.10).

Figure 21.2.10 : The shape of the male reproductive organ varies among male damselfly species, and is only compatible with the
female of that species. Reproductive organ incompatibility keeps the species reproductively isolated. (credit: “damselfly
reproductive organ” by OpenStax is licensed under CC BY 4.0)
In plants, certain structures aimed to attract one type of pollinator simultaneously prevent a different pollinator from accessing the
pollen. The tunnel through which an animal must access nectar can vary widely in length and diameter, which prevents the plant
from being cross-pollinated with a different species (Figure 21.2.11).

21.2.7 [Link]
 Figure 21.2.11 : Some flowers have evolved to attract certain pollinators. The (a) wide foxglove flower is adapted for pollination
by bees, while the (b) long, tube-shaped trumpet creeper flower is adapted for pollination by hummingbirds. (credit: “pollination”
by OpenStax is licensed under CC BY 4.0)
When fertilization takes place and a zygote forms, postzygotic barriers can prevent reproduction. Hybrid individuals in many cases
cannot form normally in the womb and simply do not survive past the embryonic stages. This is called hybrid inviability because
the hybrid organisms simply are not viable. In another postzygotic situation, reproduction leads to the birth and growth of a hybrid
that is sterile and unable to reproduce offspring of their own; this is called hybrid sterility.

Habitat Influence on Speciation

Sympatric speciation may also take place in ways other than polyploidy. For example, consider a species of fish that lives in a lake.
As the population grows, competition for food also grows. Under pressure to find food, suppose that a group of these fish had the
genetic flexibility to discover and feed off another resource that was unused by the other fish. What if this new food source was
found at a different depth of the lake? Over time, those feeding on the second food source would interact more with each other than
the other fish; therefore, they would breed together as well. The offspring of these fish would likely behave as their parents: feeding
and living in the same area and keeping them separate from the original population. If this group of fish continued to remain
separate from the first population, eventually sympatric speciation might occur as more genetic differences accumulated between
them.
This scenario does play out in nature, as do others that lead to reproductive isolation. One such place is Lake Victoria in Africa,
famous for its sympatric speciation of cichlid fish. Researchers have found hundreds of sympatric speciation events in these fish,
which have not only happened in great number, but also over a short period of time. Figure 21.2.12 shows this type of speciation
among a cichlid fish population in Nicaragua. In this locale, two types of cichlids live in the same geographic location but have
come to have different morphologies that allow them to eat various food sources.

Figure 21.2.12 : Cichlid fish from Lake Apoyeque, Nicaragua, show evidence of sympatric speciation. Lake Apoyeque, a crater
lake, is 1800 years old, but genetic evidence indicates that the lake was populated only 100 years ago by a single population of
cichlid fish. Nevertheless, two populations with distinct morphologies and diets now exist in the lake, and scientists believe these
populations may be in an early stage of speciation. (credit: “Cichlid fish” by OpenStax is licensed under CC BY 4.0)

Summary
Speciation occurs along two main pathways: geographic separation (allopatric speciation) and through mechanisms that occur
within a shared habitat (sympatric speciation). Both pathways isolate a population reproductively in some form. Mechanisms of
reproductive isolation act as barriers between closely related species, enabling them to diverge and exist as genetically independent
species. Prezygotic barriers block reproduction prior to the formation of a zygote, whereas postzygotic barriers block reproduction
after fertilization occurs. For a new species to develop, something must cause a breach in the reproductive barriers. Sympatric
speciation can occur through errors in meiosis that form gametes with extra chromosomes (polyploidy). Autopolyploidy occurs
within a single species, whereas allopolyploidy occurs between closely related species.

21.2.8 [Link]
References
OpenStax, Biology. OpenStax CNX. June 23, 2020. [Link]
Evolution.

21.2: Formation of New Species is shared under a CC BY license and was authored, remixed, and/or curated by LibreTexts.

21.2.9 [Link]
21.3: Speciation
Speciation occurs over a span of evolutionary time, so when a new species arises, there is a transition period during which the
closely related species continue to interact.

Reconnection
After speciation, two species may recombine or even continue interacting indefinitely. Individual organisms will mate with any
nearby individual with whom they are capable of breeding. An area where two closely related species continue to interact and
reproduce, forming hybrids, is called a hybrid zone. Over time, the hybrid zone may change depending on the fitness of the
hybrids and the reproductive barriers (Figure 21.3.1). If the hybrids are less fit than the parents, reinforcement of speciation occurs,
and the species continue to diverge until they can no longer mate and produce viable offspring. If reproductive barriers weaken,
fusion occurs and the two species become one. Barriers remain the same if hybrids are fit and reproductive: stability may occur and
hybridization continues.

Figure 21.3.1: After speciation has occurred, the two separate but closely related species may continue to produce offspring in an
area called the hybrid zone. Reinforcement, fusion, or stability may result, depending on reproductive barriers and the relative
fitness of the hybrids. (credit: “speciation diagram” by OpenStax is licensed under CC BY 4.0)

Exercise 21.3.1

If two species eat a different diet but one of the food sources is eliminated and both species are forced to eat the same foods,
what change in the hybrid zone is most likely to occur?

Answer
Fusion is most likely to occur because the two species will interact more and similar traits in food acquisition will be
selected.

Hybrids can be either less fit than the parents, more fit, or about the same. Usually, hybrids tend to be less fit; therefore, such
reproduction diminishes over time, nudging the two species to diverge further in a process called reinforcement. This term is used
because the low success of the hybrids reinforces the original speciation. If the hybrids are as fit or more fit than the parents, the
two species may fuse back into one species (Figure 21.3.1). Scientists have also observed that sometimes two species will remain
separate but also continue to interact to produce some hybrid individuals; this is classified as stability because no real net change is
taking place.

Varying Rates of Speciation

Scientists around the world study speciation, documenting observations both of living organisms and those found in the fossil
record. As their ideas take shape and as research reveals new details about how life evolves, they develop models to help explain
rates of speciation. In terms of how quickly speciation occurs, two patterns are currently observed: gradual speciation model and
punctuated equilibrium model.

21.3.1 [Link]
In the gradual speciation model, species diverge gradually over time in small steps. In the punctuated equilibrium model, a new
species undergoes changes quickly from the parent species and then remains largely unchanged for long periods of time afterward
(Figure 21.3.2). This early change model is called punctuated equilibrium because it begins with a punctuated or periodic change
and then remains in balance afterward. While punctuated equilibrium suggests a faster tempo, it does not necessarily exclude
gradualism.

Figure 21.3.2: In (a) gradual speciation, species diverge at a slow, steady pace as traits change incrementally. In (b) punctuated
equilibrium, species diverge quickly and then remain unchanged for long periods of time. (credit: “gradual speciation and
punctuated equilibrium” by OpenStax is licensed under CC BY 4.0)
The primary influencing factor on changes in speciation rate is environmental conditions. Under some conditions, selection occurs
quickly or radically. Consider a species of snails that had been living with the same basic form for many thousands of years. Layers
of their fossils would appear similar for a long time. When a change in the environment takes place—such as a drop in the water
level—a small number of organisms are separated from the rest in a brief period of time, essentially forming one large and one tiny
population. The tiny population faces new environmental conditions. Because its gene pool quickly became so small, any variation
that surfaces and that aids in surviving the new conditions becomes the predominant form.

Summary
Speciation is not a precise division: overlap between closely related species can occur in areas called hybrid zones. Organisms
reproduce with other similar organisms. The fitness of these hybrid offspring can affect the evolutionary path of the two species.
Scientists propose two models for the rate of speciation: one model illustrates how a species can change slowly over time; the other
model demonstrates how change can occur quickly from a parent generation to a new species. Both models continue to follow the
patterns of natural selection.

References
OpenStax, Biology. OpenStax CNX. June 25, 2020. [Link]
Evolution.

21.3: Speciation is shared under a CC BY license and was authored, remixed, and/or curated by LibreTexts.
18.3: Reconnection and Rates of Speciation by OpenStax is licensed CC BY 4.0.

21.3.2 [Link]