The Lecture:

Before we can study speciation, we need to learn what a species is. Defining a species
turns out to be somewhat difficult. Currently, different biologists have developed a
variety of different definitions of species. At a conference I went to recently someone
had a list of over thirty definitions. We won't learn all of those! Instead, we will study
why it can be difficult to define species and then focus on two of the main current
definitions of species.

We care about what a species is for several reasons, both in terms of understanding
evolution and for more practical reasons. For the purpose of understanding evolution,
many biologists argue that a species is a fundamental unit of evolution -- that
evolution of characteristics occurs within a species so that a species can be thought of
as the unit that is evolving. Different species then evolve independently from one
another. The formation of new species is a major evolutionary event because it splits a
group that was evolving as a unit into more than one separately evolving units.
Practically, defining species can have legal implications with regard to the
Endangered Species Act. Whether or not we take action to conserve a particular group
may depend on whether or not it is considered to be a separate species from other,
similar groups. This can affect the future of these organisms and the economics of the
people who live in the area where these organisms occur.

Historically, before people started studying evolution, when species were thought to
be unchangeable, people defined species simply as all individuals of a certain type,
and would describe a type specimen of each species and categorize everything that
looked like that type as belonging to the same species and everything different as
belonging to other species. To some extent, this is still done -- biological museums
keep "type specimens" of species to help categorize them, but it is recognized that
there is a major problem with this kind of definition of species. We can't define
species based on one description of what they should look like because individuals
within species vary.

We have seen that there is variation within species; it is the basis for evolution within
species. To understand the problems we may have in defining species, we need to
consider some of the different ways in which species can vary. Some of these kinds of
variation are:

• Polymorphism refers to variation within a population where different

individuals in the population have distinctly different structures, colors,
biochemistry, etc. but clearly belong to the same population since they
reproduce with one another.
• Geographic variation refers to variation over geography -- that is, populations
in different areas look different from each other. There may not be any obvious
pattern to such variation but sometimes there is. One common pattern of
geographic variation is clinal variation. Clinal variation refers to a gradual
change in some feature across geography. For example, if you go from south to
north in the northern hemisphere you frequently find that populations of
mammals are smaller in the south and gradually as you go north you find that
within a species the individuals are larger and larger. Such a pattern of gradual
change is called acline so this form of variation is called clinal variation.
• Hybrid zones are geographic areas where two distinctly different forms of
organism contact each other and interbreed. Generally, outside this zone, the
two forms retain distinct differences from one another but within the zone
intermediate forms between the two occur as a result of interbreeding. For
example, a kind of woodpecker, flickers, show a hybrid zone in the United
States. Western forms are called "red-shafted flickers" because the undersides
of their wing and tail feathers (the "shafts") are red; eastern forms are called
"yellow-shafted flickers" because the undersides of their wing and tail feathers
are yellow. Flickers occur from the west coast to the east coast of the United
States; throughout the west they are red-shafted and throughout the east they
are yellow shafted. In the middle of the country there is an area just a few
hundred miles wide where the two forms both occur and where they reproduce
with each other. There, some flickers have red shafts, some yellow, and some
have some red and some yellow. There are also DNA markers that have been
identified such that all eastern forms are fixed for one DNA form and all
western forms are fixed for a different DNA form but in the hybrid zone both
forms occur.

Hybrid zones raise some interesting evolutionary questions. Since the two
forms can reproduce with each other, we would expect gene flow to occur from
one form to the other. If gene flow occurs, we would expect the traits from one
form to spread into the other form. We observe, however, that the forms on the
two sides of the hybrid zone retain fixed differences. How is this possible?
There are several hypotheses that could explain the maintenance of stable
hybrid zones -- that is, explain how it is possible for a hybrid zone to exist over
time without there being spread of the characteristics of the form on one side of
the zone into the form on the other side of the zone. Two of these hypotheses
are as follows:
 1. The environment may be such that hybrids (organisms produced by
reproduction of the two main forms) have high fitness in the area of the
hybrid zone but low fitness outside zone. If this is the case, then hybrids
that move out of the zone will have low fitness and selection against
them may counteract the effects of gene flow out of the hybrid zone.
2. The environment where the hybrid zone occurs may be very poor quality
habitat for either form or for hybrids. As a result, individuals may move
into the zone from either side, but will not reproduce well enough within
the zone to produce many individuals that move out of the zone. If this is
the case, there will be very little gene flow out of the zone and fixed
differences between the forms on either side of the zone can be
maintained.

Biologists generally agree that we do not want to categorize the different forms in a
polymorphic population as different species, because they reproduce with each other
and offspring of the same parents could have different forms. We generally do not
want to call different populations that show clinal variation different species since the
populations change gradually from area to area and individuals can reproduce with
individuals in slightly different neighboring populations, so there would be no clear
place to draw a line to distinguish species and overall the different populations can
clearly affect one another by reproducing with one another. But what about the
different forms on either side of a hybrid zone? On one hand, they retain distinct
differences from one another so it does not appear that they are affecting each other
evolutionarily; based on this argument, people would consider them as belonging to
different species. On the other hand, they reproduce with each other within the hybrid
zone, and their offspring are healthy and fertile, so it appears that they are very similar
and that they could affect each other, so based on this argument people would
consider them as belonging to the same species. Examples such as this show why it
can be difficult to categorize groups as species. We will discuss some of the main
concepts of what a species is and consider how they would categorize groups from
different examples as either the same or as different species.

Biologists differ on whether or not they view species as fundamental evolutionary

units. Some biologists argue that evolution occurs primarily within populations. There
is evidence in a number of species that different populations evolve fairly
independently from one another. If that is the case then it may not be important how
we define species; it is determining what groups make up the populations within
which evolution is occurring that is important. Other biologists point out that
populations do affect each other, genetically -- gene flow occurs -- and argue that we
want to define species as the groups that are evolving, the fundamental units of
evolution. These are the biologists who really care about species concepts. They do
not, however, all agree on which concept of species is the best. Two of the main
concepts of species are as follows:

1. The biological species concept (BSC): categorizes species as groups of

actually or potentially interbreeding organisms that are reproductively isolated
from other such groups. In other words, organisms are considered to belong to
the same species if they can potentially reproduce with each other and have
offspring that are healthy and fertile. Otherwise, they are considered to belong
to different species.
2. The phylogenetic species concept (PSC): categorizes a species as the smallest
group that can be diagnosed with a derived character state that clearly separates
it from other such groups. In other words, if you have a group of organisms that
all share a derived character state and that can not be subdivided into smaller
groups based on shared derived character states then you have a species.
Different species have different derived character states from each other.

Both these concepts of species try to identify the groups that will evolve as
independent evolutionary units. The biological species concept is based on the idea
that if gene flow can occur between groups (because they can reproduce and produce
healthy, fertile offspring) then these groups can affect each other genetically and
should be grouped into a single species. The phylogenetic species concept is based on
the idea that we can tell which groups are in fact affecting each other genetically by
their characteristics -- if they share a unique derived characteristic not found in other
groups, then they are not spreading this characteristic to other groups; if they do not
have the derived characteristics found in other groups, then the other groups are not
affecting them.

There are potential problems with each concept that are solved by the other concept.
These follow:

The biological species concept can not categorize asexual species (since they do not
reproduce with other individuals at all) or fossil species (since we can not tell what
fossils could have reproduced with what other fossils now that they're all dead.) The
phylogenetic species concept, since it is based on characteristics other than
reproduction, provides a basis for classifying such species.

The biological species concept lumps into a single species some groups that are very
ecologically and structurally different and that are apparently evolving separately. For
example, there is a group of plants on the Hawaiian islands called the silverswords:
some are vines, some are shrubs, some are small plants, some are trees. All forms can
apparently reproduce with all other forms, but the different forms are clearly distinct
from each other and evolving independently. They appear to be different species, but
the biological species concept would categorize all as belonging to the same species
because they can reproduce with each other. In contrast, the phylogenetic species
concept classifies them as different species because each group has its own unique
derived characteristics. Similarly, the biological species concept would categorize two
groups that are different from each other but form a hybrid zone, like the flickers
described above, as belonging to the same species despite their distinctive differences,
while the phylogenetic species concept would categorize them as different species
because of their distinctive differences. The phylogenetic concept may thus better
reflect, in these cases, groups that are evolving independently from each other.

The biological species concept may group organisms that are not each others closest
relatives into a species. For example, in a group of birds called orioles, two forms, the
Baltimore oriole and the Bullock's oriole, can reproduce with each other and produce
healthy, fertile offspring. Phylogenetic analysis of these orioles, however, suggests
that Bullock's orioles are more closely related to other species of oriole, with which
they can not interbreed, than to Baltimore orioles. This kind of situation can arise
because the ability to interbreed is primitive within these species -- originally, they
belonged to the same species and reproduced with each other. More recently, some
species have derived traits such that they can't reproduce with other groups, but some
retain the primitive ability to reproduce with each other. Species categorized based on
the phylogenetic species concept should avoid this problem. They are categorized
based on derived characteristics and these, as you learned in the lectures on
systematics, reflect phylogenetic relatedness. So the phylogenetic concept should
group organisms into species in which all members are more closely related to each
other than to other species. The phylogenetic species concept thus reflects
evolutionary history better than does the biological species concept.

There are also problems with the phylogenetic species concept, though. The
phylogenetic species concept may divide species into groups based on characteristics
that do not have any clear biological relevance and appear to be very minor traits. The
phylogenetic concept may divide life up into an inconveniently large number of
species. Probably the most serious problem with the phylogenetic species concept is
that it may divide groups into species that are not going to remain separate species
because the groups can still reproduce with each other. Thus, the species defined by
the phylogenetic species concept may not be permanent -- they may merge together
over time. The biological species concept avoids these problems.

it is important to note that most of the time they agree on what they
define as species. Any time two groups are so different that they
 can not reproduce to produce healthy fertile offspring, both
concepts consider them to be separate species. The biological
concept considers them separate species because the ability to
reproduce is the criterion this concept uses to group organisms into
the same species. The phylogenetic concept considers them
separate species because if two groups can not reproduce with each
other then they must have some different derived traits that prevent
them from reproducing. So the two concepts only differ where there
are groups of organisms that can reproduce with each other and
produce healthy, fertile offspring but have distinctly different
derived traits. When this occurs, the biological concept lumps these
organisms into the same species, and the phylogenetic concept
splits them into separate species. This occurs among very closely
related groups that may just be in the process of speciating

Speciation, the evolution of two or more new species from one pre-existing species, will occur if
different groups within a species evolve to become different from each other -- so different that
they would be considered different species. How different that is depends on which species
concept you are using. Remember that the biological species concept says they have to be so
different that they cannot reproduce with each other to produce healthy, fertile offspring. The
phylogenetic species concept says they have to have evolved different derived character states
from each other.

Whether different groups within a species evolve differences depends on the main
forms of evolution we've already studied -- natural selection, genetic drift, and gene
flow. These affect speciation as follows.

Remember that gene flow tends to make populations similar to each other. Since
speciation requires that groups evolve differences from each other, gene flow tends to
prevent speciation from occurring.

Remember that genetic drift and natural selection tend to make different
populations different from each other. Since speciation requires that groups evolve
differences from each other, genetic drift and natural selection can result in the
evolution of differences among populations and therefore can cause speciation to
occur.

Cases Where the Biological Species Concept Applies

One advantage of the BSC is that it provides a reasonably unambiguous test that can
be applied to possible speciation events. Recall that under the BSC species are defined
as being reproductively isolated from other species. Demonstrating that a population
 is reproductively isolated (in a nontrivial way) from populations that it was formerly
able to interbreed with shows that speciation has occurred. In practice, it is also
necessary to show that at least one isolating mechanism with a hereditary basis is
present. After all, just because a pair of critters don't breed during an experiment
doesn't mean they can't breed or even that they won't breed. Debates about whether a
speciation event has occurred often turn on whether isolating mechanisms have been
produced.

4.1.1 Isolating Mechanisms

Mechanisms which produce reproductive isolation fall into two broad categories --
premating mechanisms and postmating mechanisms.

Premating isolating mechanisms operate to keep species separate before mating

occurs. Often they act to prevent mating altogether. Examples of premating
mechanisms include ecological, temporal, behavioral and mechanical mechanisms.

Ecological isolation occurs when species occupy or breed in different habitats. It is

important to be careful when claiming ecological isolation. For example, I have a
population of Dinobryon cylindricum (a colonial algal flagellate) growing in a culture
tube in an environmental chamber. It's been there for three years (which is a lot of
time in flagellate years! :-)). Even though there is no possibility that they will mate
with the D. cylindricum in Lake Michigan, it would be silly to assert that they
therefore constitute a separate species. Physical isolation alone does not constitute an
isolating mechanism with an hereditary basis.

Temporal isolation occurs when species breed at different times. This may be
different times of the year or different times of day.

Behavioral isolating mechanisms rely on organisms making a choice of whether to

mate and a choice of who to mate with. Differences in courtship behavior, for
instance, may be sufficient to prevent mating from occurring. A behavioral isolating
mechanism should result in some sort of positive assortative mating. Simply put,
positive assortative mating occurs when organisms that differ in some way tend to
mate with organism that are like themselves. For example, if blonds mate exclusively
with blonds, brunettes mate exclusively with brunettes, redheads mate exclusively
with redheads (and those of us without much hair don't get to mate :-() the human
population would exhibit a high degree of positive assortative mating. In most
examples in the literature when positive assortative mating is seen it is not this strong.
Positive assortative mating is especially important in discussions of sympatric
speciation.
Mechanical isolating mechanisms occur when morphological or physiological
differences prevent normal mating.

Postmating isolating mechanisms prevent hybrid offspring from developing or

breeding when mating does occur. There are also several examples of postmating
mechanisms.

Mechanical postmating isolating mechanisms occur in those cases where mating is

possible, but the gametes are unable to reach each other or to fuse. Mortality acts as an
isolating mechanism when the hybrid dies prior to maturity. Sterility of hybrids can
act as an isolating mechanism. Finally a reduction in the fitness of the hybrid
offspring can isolate two populations. This happens when the F1 hybrid is fertile but
the F2 hybrid has lower fitness than either of the parental species.

Models of Speciation:

The different models of speciation we will look at in this lecture are all based on
determining situations in which there will be little or no gene flow between groups so
that speciation can occur because of natural selection and genetic drift within groups,
or at situations in which the effects of natural selection may be strong enough to
counteract gene flow so that speciation occurs.

The main models of speciation are categorized based on geography -- that is, what are
the geographic locations, with respect to one another, of the groups that evolve to be
different species. There are three geographic categories of speciation. These are:

1. Allopatric speciation: speciation that occurs when the groups that evolve to be
separate species are in different geographic locations and are isolated
geographically from each other so that individuals cannot move between the
different locations.
2. Parapatric speciation: speciation that occurs when the groups that evolve to
be separate species are geographic neighbors; they are in different areas, but the
areas are next to each other and individuals can move between the areas.
3. Sympatric speciation: speciation that occurs when the groups that evolve to be
separate species occur together in the same geographic area.

Note that these definitions are based on geographic location -- they do not consider the process
through which speciation occurs. We will look now at the processes that can occur to cause each
of the geographic categories of speciation listed above.

During allopatric speciation there is a geographic barrier between areas that prevents
gene flow between areas from occurring. As a result, there is no gene flow to keep
populations similar to each other. Natural selection and genetic drift occurring in each
of the geographically isolated populations will cause those populations to evolve to be
different from each other -- eventually they will be so different that they evolve to be
different species.

The question we should ask about allopatric speciation is: how do different
populations of a species become geographically isolated from each other? There are
two main causes of geographic isolation:

1. Vicariance events are events that split the range of an existing species by
creating some kind of geographic barrier within the range of that species. These
are typically geological events. Examples include:
o the formation of glaciers during the ice ages, which divided the ranges of
species into smaller areas isolated by glaciers (huge mountains of ice.)
o continental drift -- all the continents on earth were once a single land
mass; as they separated from each other the species on them became
separated into different populations on the different continents.
o changes in water position and level -- rising water levels have flooded
low lying areas and turned high areas into islands isolated from each
other by water; rivers changing course can divide species that can't
disperse over water into isolated populations.
2. Dispersal events, also called founder events are events during which a small
number of individuals from the original geographic range of a species move to
a new area, previously unoccupied by that species, and start a new population
there. For example, a small number of individuals may find their way over a
mountain range, or be blown out to an island during a storm. After this
dispersal, there is no further contact between the original population and the
new population. The form of allopatric speciation that occurs after a dispersal
(founder) event is called founder effect speciation. It is not currently clear
how important founder effect speciation is as a form of speciation; some
evolutionary biologists have argued that it is an extremely common form of
speciation while others have argued that it is rare. We will consider the
theoretical arguments on both sides of this issue in the next lecture.

The basic process of allopatric speciation is similar for either cause of isolation of the
populations -- once the populations are isolated, gene flow is prevented and differences between
the isolated populations evolve through drift and selection.

Now let's consider parapatric speciation. Since individuals can move between the
populations in this case, gene flow occurs. We have seen that gene flow tends to make speciation
unlikely. For speciation to occur in such a situation, there must be strong selection to counteract
the effect of gene flow. It is argued that parapatric speciation may occur where there is an abrupt
change in the environment over a geographic border. The result is that forms that have high
fitness in one area have low fitness in the neighboring area, and hybrids between these forms do
not have high fitness in either area. For example, there are areas where there is a sudden change
in soil type from one place to the next. Plants that have high fitness in one soil type have low
fitness in the other. This results in strong natural selection against the forms from other areas and
this may counteract the effects of potential gene flow enough so that the forms in the different
areas evolve, through natural selection, into different species.

Note that the form of selection occurring in this case is disruptive selection in that
one extreme form has high fitness in one environment, the opposite extreme has high fitness in
the other environment, and intermediates do not do well anywhere so have the lowest fitness.

Note that parapatric speciation differs from allopatric in the process that occurs -- for parapatric
speciation to occur there must be strong disruptive selection since that is what counteracts the
effects of gene flow. For allopatric speciation, gene flow is not possible because of geographic
barriers, and genetic drift or weaker forms of natural selection can lead, over time, to speciation
-- strong disruptive selection is not required. Because of the requirement for an environment that
will result in strong disruptive selection across a geographic boundary, parapatric speciation is
generally thought to be less common than allopatric speciation.

Finally, let us consider sympatric speciation. Sympatric speciation was initially

considered to be much less common than allopatric speciation because if groups are
occurring within the same area it seems likely that gene flow will occur between
them, and this will tend to prevent speciation. It has been recognized, however, that
there are situations in which sympatric speciation can occur. Two main ways
sympatric speciation can occur are:

1. Ecological isolation in which different groups occur in the same geographic area but
do not contact each other because of some ecological difference that prevents gene flow
between the groups. For example, parasitic forms of a species that occur, and reproduce,
in or on different host species will not have gene flow between them. Flower forms that
attract different pollinators and are only pollinated by certain specialist pollinators will
not have gene flow between them. Species with small body size may specialize on
different parts of the habitat -- for example, some on small plants, some on treetops -- and
as a result not contact each other. In any situation like this, it is possible for there to be
different forms within the same geographic area that do not have gene flow between
them. In the absence of gene flow, natural selection and genetic drift will tend to make
the groups different over time, and speciation can occur. Note that the process of
speciation in this case is similar to what occurs in allopatric speciation; the difference is
that for allopatric speciation the barriers to gene flow are geographic, while in this case
there are ecological barriers to gene flow within the same geographic range
2. strong disruptive selection within an area could potentially result in sympatric
speciation if selection for the extreme forms and against intermediates is sufficiently
strong so that crosses between the extreme forms result in few surviving offspring and
therefore little gene flow between the two extremes. Note that this process of speciation
 is similar to what occurs during parapatric speciation; the difference is that for parapatric
speciation the disruptive selection occurs across a geographic boundary, while in this
case it is within one area.

With regard to all of these speciation models, an unanswered question in evolutionary biology is
how much is speciation driven by strong selection, such as active selection for different forms of
traits in different groups, versus how much occurs gradually through drift and gradual
accumulation of minor differences between groups as a result of weak selection. One approach
that is being taken to study such questions is based on analyses of genes that affect quantitative
traits that differ between closely related species. The approach involves mapping the location of
quantitative trait loci (genes responsible for quantitative traits) on chromosomes; this is referred
to as QTL analysis. QTL analyses are based on making hybrids between two species, and then
finding genetic markers that are statistically associated with particular parental phenotypes.
Your textbook discusses an example of this in two species of monkeyflower, one which is pink
and pollinated by bumblebees, and the other of which is red and pollinated by hummingbirds
(this study inspired the red monkeyflower example in the first computer assignment.) Bradshaw,
Schemske, and others have created hybrids between these species by cross-pollinating them by
hand and have found genetic markers associated with traits important in pollination, such as
color of the flowers, shape of the flowers, and nectar production. Their results suggest that a
relatively small number of genetic loci that are associated with these important adaptive traits
and that differ between the species. They suggest that speciation may occur as a result of
selection for these important adaptive traits that result in reproductive isolation (because they
attract different pollinators.) We do not know at this point how general this result is:

Founder Effect And Chromosomal Speciation:

In the last lecture, you learned that one of the ways that allopatric speciation can occur is through
a founder or dispersal event -- that is, an event where a small number of individuals disperse to
a new location, previously unoccupied by that species. Some evolutionary biologists have argued
that such dispersal events are likely to result in rapid evolution in the newly founded population
and are, as a result, particularly likely to result in speciation as the new population evolves to
become a different species from the original population. Others, however, have argued that most
speciation results from vicariance events and that dispersal events are not likely to result in the
kind of evolution that will lead to new species evolving. We will consider one argument that
suggests that founder events are likely to cause speciation and one argument that suggests that
founder events are unlikely to cause speciation. Further tests of these hypotheses are necessary
before we can evaluate which is correct.
First, we will consider an argument that founder events are NOT likely to cause
speciation. According to this hypothesis:

• Newly founded populations, resulting from a dispersal event, are small, so there
will be high levels of genetic drift in newly founded populations.
• The main result of genetic drift is loss of genetic variation, so newly founded
populations will quickly lose genetic variation.
• Genetic variation is required for evolutionary change to occur, so newly
founded populations, once they have lost genetic variation, will not evolve
much. As a result, they are unlikely to evolve major differences from the
original populations and are not very likely evolve to be different species from
the original populations.

Now let us consider the argument that founder events ARE likely to result in speciation.
According to this hypothesis:

• Newly founded populations, resulting from a dispersal event, are small, so there
will be high levels of genetic drift in newly founded populations.
• Newly founded populations will grow, so that genetic drift will only be a major
evolutionary process at first. As a result, there will not be loss of too much
genetic variation.
• While the population is small, and genetic drift is the major form of evolution
occurring, traits will evolve at random.
• As a result of random evolutionary change in traits, trait combinations may
evolve away from the adaptive peaks where they originally occurred. This
assumes that there is an adaptive landscape with several adaptive peaks (note: if
you have forgotten about adaptive landscapes, you should review them at this
point.Click here to review that lecture.) Remember that populations will not
evolve from one adaptive peak to another through natural selection, because of
the low fitness adaptive valleys in between the adaptive peaks. Since genetic
drift is random, it CAN result in evolution away from the closest adaptive peak.
• Once the population grows large enough so that genetic drift is no longer the
major process of evolution and natural selection also occurs, the population is
likely to be near different adaptive peaks from the original population (because
it went through the period of random evolution through genetic drift.) Natural
selection will then result in evolution of the population to whatever the closest
adaptive peak now is.
• Since the population is likely to be at a different adaptive peak from the
original population, it has gone through significant evolutionary change. It is
thus likely to have evolved to be a different species from the original
population.
Note that both arguments are based on an initial phase of genetic drift that is predicted because
populations are likely to be started (founded) by only a small number of individuals so genetic
drift should be the main form of evolution occurring initially in these populations. They differ in
the predicted effect of genetic drift. If genetic drift occurs for a long enough time so that genetic
variation is lost, then these new populations are not likely to evolve to be different species. If,
however, genetic drift only occurs for a short period of time, and the population grows large
enough so that genetic drift becomes less important relatively quickly, then genetic drift could
cause some initial random genetic changes in the population that will subsequently make it more
likely to evolve to be very different from the original population so that the two populations will
be different species. Both arguments are plausible; both have been modeled theoretically. We
need more data on what really happens in populations and how speciation events have most
likely occurred to evaluate which is more likely to be generally true.

For the rest of the lecture we will consider a different topic of relevance to speciation.
We have considered speciation so far as occurring as a result of relatively small scale
genetic changes that build up, through natural selection and genetic drift, so that
populations evolve to become different species. There are also, however, larger scale
mutations that may cause much more rapid speciation. We will consider these now.

We will define chromosomal mutations as large-scale changes in chromosomal

structure or number. These can occur in several ways; we will consider three main
kinds of chromosomal mutation:

1. Fusions occur when two small chromosomes combine to form one larger
chromosome. A common way in which this occurs is for two chromosomes
with centromeres very near the end of the chromosome to fuse into one large
chromosome with the centromere in the middle, as shown in the following
picture. Note that when chromosomes become fused like this, the total number
of chromosomes is decreased.
 2. Inversions occur when a large segment of a chromosome is spliced out,
reversed in order, and then spliced back into a chromosome. This is illustrated
in the following picture, where the letters represent genes on the chromosome.
An inversion generally involves several genes and changes the order of genes
on the chromosome.

3. Polyploidy occurs when the total number of chromosome sets increases in

number -- typically what occurs is that all chromosomes are duplicated. A
species that was originally diploid (two copies of every chromosomes; 2N)
would, as a result, become tetraploid, (4N) with four copies of every
chromosome.

These chromosomal mutations can result in speciation, as follows. Suppose an individual is

produced with one of these chromosomal mutations. It will have all of the genes needed for
normal development. In the case of polyploidy, there will be more copies of all of these genes,
but they will occur in the same proportions as they did in the original individuals so there will be
no imbalance in the doses of the proteins coded by these genes. As a result, all of these
chromosomal mutations are likely to produce healthy individuals -- they will not be harmful to
the health of the individuals.

When an individual with a chromosomal mutation reproduces with an individual

without the chromosomal mutation, however, the offspring, while healthy, may be
sterile. Consider how this works with polyploidy. Suppose the polyploid individual is
4N. It reproduces with an original 2N individual. When the 4N individual produces
gametes, through meiosis, the gametes have half as many chromosomes as the
individual so they are 2N. The gametes of the diploid individual, produced through
meiosis, are haploid (1N.) As a result, fusion of the 2N gamete and the 1N gamete
results in an offspring that is 3N.
The 3N individual, the hybrid between the 2N and 4N forms, should be healthy
because (as noted above) there is no imbalance in the dosages of proteins coded by its
genes (it has all the protein products of the genes in the right proportions -- none is out
of proportion.) However, when its cells undergo meiosis to produce gametes, there are
likely to be major problems. The key event in meiosis that allows production of
gametes with half as many chromosomes as the original cells is the pairing of
homologous chromosomes. This 3N individual has 3 homologous chromosomes for
each type of chromosome. When they try to pair, three things can not form pairs, so
different parts of the chromosomes pair with parts of the other chromosomes, and
when they separate they are likely to break. The result is that the daughter cells get
broken chromosomes and may too many copies of some genes and no copies of other
genes. They will not be viable gametes. So these 3N individuals (and, in general, any
individual with an odd number of chromosomes) will be sterile because it cannot
produce viable gametes because chromosomes can not pair correctly during meiosis.

So this means when an individual that is polyploid reproduces with an individual with
the original number of chromosomes, the offspring of this cross are healthy but sterile.
By the biological species concept, the polyploid is therefore a different species from
the original -- the two can reproduce but their offspring are sterile. By the
phylogenetic species concept, being a polyploid means having a trait (in this case
polyploidy) that is derived and different from the original form, so it would be a
different species from the original by the phylogenetic species concept, too.

There is one problem we have to consider, though. What happens when a polyploid
individual is produced in a normal, diploid population? If it reproduces with the
diploids, the offspring will be sterile. The first polyploid produced through a mutation
would have only diploids to reproduce with. You would think, as a result, that this
form would quickly die out. It would, if the only way it can reproduce is sexually. If,
however, this species can reproduce asexually, then a polyploid can reproduce more
polyploids, like itself, asexually. These polyploids can then reproduce sexually with
one another, and their offspring will have the same, even number of chromosomes
that they do so they will be fertile as well as healthy. In this case, a new sexually
reproducing species can be produced.

We would generally expect, then, that polyploidy could result in new sexually
reproducing species primarily in species that can undergo both sexual and asexual
reproduction. In fact, we observe that in the flowering plants, a group in which most
species can undergo both sexual and asexual reproduction, many closely related
species differ in the number of chromosome sets, suggesting that they have been
produced as a result of polyploidy. Evidence of polyploidy is much rarer in animals,
which often can not reproduce asexually. If polyploidy occurred in a species that can
reproduce only sexually, it would quickly die out -- all the polyploid's offspring, with
the diploid members of the population, would be sterile.

The other forms of chromosomal mutation (inversion, fusion) can also result in
situations in which offspring of the original form and the mutated form are sterile.
There are some other complicating factors in these situations, though (not all
inversions or fusions have this effect, and they may be involved in speciation for other
reasons.) We will not consider these complications in this course. For this course,
focus on polyploidy as the main example of how chromosomal mutations can result in
speciation.