476 Chapter 18 | Evolution and the Origin of Species

second misunderstanding may arise by interpreting the statement to mean that the 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 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 antibiotic.
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 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.

18.2 | Formation of New Species

By the end of this section, you will be able to:
• Define species and describe how species are identified as different
• Describe genetic variables that lead to speciation
• Identify prezygotic and postzygotic reproductive barriers
• Explain allopatric and sympatric speciation
• Describe adaptive radiation

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 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 18.9).

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

Chapter 18 | Evolution and the Origin of Species 477

Figure 18.9 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)

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 18.10). 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 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 18.10 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)

Populations of species share a gene pool: a collection of all the variants of genes in the species. Again, the basis to
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 18.11a). Compare this illustration to the diagram of elephant
478 Chapter 18 | Evolution and the Origin of Species

evolution (Figure 18.11b), 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.

Figure 18.11 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.

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 18.12).

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

Chapter 18 | Evolution and the Origin of Species 479

Figure 18.12 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)

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 18.13.
480 Chapter 18 | Evolution and the Origin of Species

Figure 18.13 The honeycreeper birds illustrate adaptive radiation. From one original species of bird, multiple others
evolved, each with its own distinctive characteristics.

Notice the differences in the species’ beaks in Figure 18.13. Evolution in response to natural selection based on specific
food sources in each new habitat led to 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.

Click through this interactive site (http://openstaxcollege.org/l/bird_evolution) to see how island birds evolved in
evolutionary increments from 5 million years ago to today.

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 18.14).

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

Chapter 18 | Evolution and the Origin of Species 481

Figure 18.14 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

Which is most likely to survive, offspring with 2n+1 chromosomes or offspring with 2n-1 chromosomes?

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 18.15). 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 18.15 Autopolyploidy results when mitosis is not followed by cytokinesis.

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 18.16 illustrates one possible way an allopolyploid can form. Notice how it
takes two generations, or two reproductive acts, before the viable fertile hybrid results.
482 Chapter 18 | Evolution and the Origin of Species

Figure 18.16 Alloploidy results when two species mate to produce viable offspring. In the example shown, a normal
gamete from one species fuses with a polyploidy gamete from another. Two matings are necessary to produce viable
offspring.

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 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.

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 18.17).

Figure 18.17 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)

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

Chapter 18 | Evolution and the Origin of Species 483

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 (Figure 18.18).

Figure 18.18 Speciation can occur when two populations occupy different habitats. The habitats need not be far apart.
The cricket (a) Gryllus pennsylvanicus prefers sandy soil, and the cricket (b) Gryllus firmus prefers loamy soil. The two
species can live in close proximity, but because of their different soil preferences, they became genetically isolated.

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 18.19).

Figure 18.19 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.

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 18.20).

Figure 18.20 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 humming birds.

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
484 Chapter 18 | Evolution and the Origin of Species

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. Offspring of these fish would likely
behave as their parents: feeding and living in the same area and keeping 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 18.21 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 18.21 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.

18.3 | Reconnection and Rates of Speciation

By the end of this section, you will be able to:
• Describe pathways of species evolution in hybrid zones
• Explain the two major theories on rates of 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 who they are capable of breeding with. 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 18.22). 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.

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10

Chapter 18 | Evolution and the Origin of Species 485

Figure 18.22 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.

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?

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 18.23). 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.
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 18.23). 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.
486 Chapter 18 | Evolution and the Origin of Species

Figure 18.23 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.

Which of the following statements is false?

a. Punctuated equilibrium is most likely to occur in a small population that experiences a rapid change in
its environment.
b. Punctuated equilibrium is most likely to occur in a large population that lives in a stable climate.
c. Gradual speciation is most likely to occur in species that live in a stable climate.
d. Gradual speciation and punctuated equilibrium both result in the divergence of species.

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.

Visit this website (http://openstaxcollege.org/l/snails) to continue the speciation story of the snails.

This OpenStax book is available for free at http://cnx.org/content/col11448/1.10