Lesson 6.

Speciation

How do new species arise?

A species is often defined as a group of individuals that actually or potentially
interbreed in nature. In this sense, a species is the biggest gene pool possible under
natural conditions.

For example, these happy face spiders look different, but since they can interbreed,
they are considered the same species: Theridion grallator.

Happy face spider photos courtesy of Rosemary Gillespie, UC Berkeley.

Defining speciation
Speciation is a lineage-splitting event that produces two or more separate species.
Imagine that you are looking at a tip of the tree of life that constitutes a species of
fruit fly. Move down the phylogeny to where your fruit fly twig is connected to the
rest of the tree. That branching point, and every other branching point on the tree,
is a speciation event. At that point genetic changes resulted in two separate fruit fly
lineages, where previously there had just been one lineage. But why and how did it
happen?
 The branching points on this partial Drosophila phylogeny represent long past
speciation events. Here is one scenario that exemplifies how speciation can
happen:

 The scene: a population of wild fruit flies minding its own business on several
bunches of rotting bananas, cheerfully laying their eggs in the mushy fruit…

Disaster strikes: A hurricane washes the bananas and the immature fruit flies they
contain out to sea. The banana bunch eventually washes up on an island off the
coast of the mainland. The fruit flies mature and emerge from their slimy nursery
onto the lonely island. The two portions of the population, mainland and island,
are now too far apart for gene flow to unite them. At this point, speciation has not
occurred — any fruit flies that got back to the mainland could mate and produce
healthy offspring with the mainland flies.
The populations diverge: Ecological conditions are slightly different on the island,
and the island population evolves under different selective pressures and
experiences different random events than the mainland population does.
Morphology, food preferences, and courtship displays change over the course of
many generations of natural selection.

So we meet again: When another storm reintroduces the island flies to the

mainland, they will not readily mate with the mainland flies since they’ve evolved
different courtship behaviors. The few that do mate with the mainland flies,
produce inviable eggs because of other genetic differences between the two
populations. The lineage has split now that genes cannot flow between the
populations.
This is a simplified model of speciation by geographic isolation, but it gives an idea
of some of the processes that might be at work in speciation. In most real-life
cases, we can only put together part of the story from the available evidence.
However, the evidence that this sort of process does happen is strong.
New species arise through a process called speciation. In speciation, an
ancestral species splits into two or more descendant species that are
genetically different from one another and can no longer interbreed.

Darwin envisioned speciation as a branching event. In fact, he considered it

so important that he depicted it in the only illustration of his famous
book, On the Origin of Species, below left. A modern representation of
Darwin's idea is shown in the evolutionary tree of elephants and their
relatives, below right, which reconstructs speciation events during the
evolution of this group.
 Image credit: Formation of new species: Figure 3 by OpenStax College, Biology, 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 often divide the ways that speciation can occur into two broad
categories:

 Allopatric speciation—allo meaning other and patric meaning homeland—

involves geographic separation of populations from a parent species and
subsequent evolution.

 Sympatric speciation—sym meaning same and patric meaning homeland—

involves speciation occurring within a parent species remaining in one
location.

Let's take a closer look at these forms of speciation and how they work.

Allopatric speciation
In allopatric speciation, organisms of an ancestral species evolve into two or
more descendant species after a period of physical separation caused by a
geographic barrier, such as a mountain range, rockslide, or river.

Sometimes barriers, such as a lava flow, split populations by changing the

landscape. Other times, populations become separated after some members
cross a pre-existing barrier. For example, members of a mainland population
may become isolated on an island if they float over on a piece of debris.

Once the groups are reproductively isolated, they may undergo genetic

divergence. That is, they may gradually become more and more different in
their genetic makeup and heritable features over many generations. Genetic
divergence happens because of natural selection, which may favor different
traits in each environment, and other evolutionary forces like genetic drift.

As they diverge, the groups may evolve traits that act as prezygotic and/or
postzygotic barriers to reproduction. For instance, if one group evolves large
body size and the other evolves small body size, the organisms may not be
physically able to mate—a prezygotic barrier—if the populations are
reunited.

If the reproductive barriers that have arisen are strong—effectively

preventing gene flow—the groups will keep evolving along separate paths.
That is, they won't exchange genes with one another even if the geographical
barrier is removed. At this point, the groups can be considered separate
species.
Allopatric speciation

Allopatric speciation is just a fancy name for speciation by

geographic isolation, discussed earlier. In this mode of speciation, something
extrinsic to the organisms prevents two or more groups from mating with each
other regularly, eventually causing that lineage to speciate. Isolation might occur
because of great distance or a physical barrier, such as a desert or river, as shown
below.
Allopatric speciation can occur even if the barrier is a little “porous,” that is, even if
a few individuals can cross the barrier to mate with members of the other group. In
order for a speciation even to be considered “allopatric,” gene flow between the
soon-to-be species must be greatly reduced — but it doesn’t have to be reduced
completely to zero.

Case study: squirrels and the Grand Canyon

The Grand Canyon was gradually carved out by the Colorado River over
millions of years. Before it formed, only one species of squirrel inhabited the
area. As the canyon got deeper over time, it became increasingly difficult for
squirrels to travel between the north and south sides.

Image credit: Toroweap
sunrise by John Fowler, CC BY 2.0

Eventually, the canyon became too deep for the squirrels to cross and
a subgroup of squirrels became isolated on each side. Because the
squirrels on the north and south sides were reproductively isolated
from one another due to the deep canyon barrier, they eventually
diverged into different species.
 Image credit: left, image modified from Ammospermophilus harrisii by Ryan Johnston, CC BY 2.0; right,
image modified from Ammospermophilus leucurus by Jarek Tuszynski, CC BY-SA 3.0

Harris's antelope squirrel evolved on the south side of the Grand Canyon as a
result of allopatric speciation.

The white-tailed antelope squirrel evolved on the north side of the Grand
Canyon as a result of allopatric speciation.

Sympatric speciation
In sympatric speciation, organisms from the same ancestral species become
reproductively isolated and diverge without any physical separation.

At first, this idea may seem kind of weird, especially after thinking about
allopatric speciation. Why would groups of organisms in a population stop
interbreeding when they still live in the same place?

There are several ways that sympatric speciation can happen. However, one
mechanism that's quite common—in plants, that is!—involves chromosome
separation errors during cell division. 
Home → Modes of speciation → Sympatric speciation

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Sympatric speciation

Unlike the previous modes, sympatric speciation does not require

large-scale geographic distance to reduce gene flow between parts of a population. How could a
randomly mating population reduce gene flow and speciate? Merely exploiting a new niche may
automatically reduce gene flow with individuals exploiting the other niche. This may
occasionally happen when, for example, herbivorous insects try out a new host plant.

For example, 200 years ago, the ancestors of apple maggot flies laid their eggs only on
hawthorns — but today, these flies lay eggs on hawthorns (which are native to America) and
domestic apples (which were introduced to America by immigrants and bred). Females generally
choose to lay their eggs on the type of fruit they grew up in, and males tend to look for mates on
the type of fruit they grew up in. So hawthorn flies generally end up mating with other hawthorn
flies and apple flies generally end up mating with other apple flies. This means that gene flow
between parts of the population that mate on different types of fruit is reduced. This host shift
from hawthorns to apples may be the first step toward sympatric speciation — in fewer than 200
years, some genetic differences between these two groups of flies have evolved.

From left to right, apple maggot flies, apples, and hawthorns. Apple maggot flies photo used with
permission from University of California Statewide IPM Program, J. K. Clark, photographer; Apple photo
by Doug Wilson courtesy of USDA; Hawthorn photo © Erv Evans
Gene flow has been reduced between flies that feed on different food varieties, even though they both live
in the same geographic area.

Polyploidy
Polyploidy is the condition of having more than two full sets of
chromosomes. Unlike humans and other animals, plants are often tolerant of
changes in their number of chromosome sets, and an increase in chromosome
sets, a.k.a. ploidy, can be an instant recipe for plant sympatric speciation.

How could polyploidy lead to speciation? As an example, let’s consider the

case where a tetraploid plant—4n, having four chromosome sets—suddenly
pops up in a diploid population—2n, having two chromosome sets.

Such a tetraploid plant might arise if chromosome separation errors

in meiosis produced a diploid egg and a diploid sperm that then met up to
make a tetraploid zygote. This process is shown in a general schematic
below.
Image credit: modified from Polyploidization by Ilmari Karonen, public domain

When the tetraploid plant matures, it will make diploid, 2n, eggs and sperm.
These eggs and sperm can readily combine with other diploid eggs and sperm
via self-fertilization, which is common in plants, to make more tetraploids.

On the other hand, the diploid eggs and sperm may or may not combine
effectively with the haploid, 1n, eggs and sperm from the parental species.
Even if the diploid and haploid gametes do get together to produce a triploid
plant with three chromosome sets, this plant would likely be sterile because
its three chromosome sets could not pair up properly during meiosis.
Because the tetraploid plants and the diploid species from which they came
cannot produce fertile offspring together, we consider them two separate
species. This means that speciation occurred after just a single generation!

Speciation by polyploidy is common in plants but rare in animals. In general,

animal species are much less likely to tolerate changes in ploidy. For
instance, human embryos that are triploid or tetraploid are non-viable—they
cannot survive.

Sympatric speciation without polyploidy

Can sympatric speciation, speciation without geographical separation, occur
by mechanisms other than polyploidy? There’s some debate about how
important or common a mechanism it is, but the answer appears to be yes, at
least in some cases. For instance, sympatric speciation may take place when
subgroups in a population use different habitats or resources, even though
those habitats or resources are in the same geographical area.

One classic example is the North American apple maggot fly. As the name
suggests, North American apple maggot flies, like the one pictured below,
can feed and mate on apple trees. The original host plant of these flies,
however, was the hawthorn tree. It was only when European settlers
introduced apple trees about 200 years ago that some flies in the population
started to exploit apples as a food source instead.

Image credit: Rhagoletis pomonella.jpg by

Joseph Berger, CC BY 3.0

The flies that were born in apples tended to feed on apples and mate with
other flies on apples, while the flies born on hawthorns tended to similarly
stick with hawthorns. In this way, the population was effectively divided into
two groups with limited gene flow between them, even though there was no
reason an apple fly couldn't go over to a hawthorne tree, or vice versa.

Over time, the population diverged into two genetically distinct groups with
adaptations, features arising by natural selection, that were specific for apple
and hawthorne fruits. For instance, the apple and hawthorne flies emerge at
different times of year, and this genetically specified difference synchronizes
them with the emergence date of the fruit on which they live.
 Some interbreeding still occurs between the apple-specialized flies and the
hawthorne-specialized flies, so they are not yet separate species. However,
many scientists think this is a case of sympatric speciation in progress.

Quiz:
Q1: 

Which scientist spent many years gathering evidence from natural populations to

support the theory of speciation?

 A Wallace
 B Mendel
 C Franklin
 D Watson
 E Darwin

Q2: 

If a population cannot adapt to a new environment, it becomes extinct. Which of the

following correctly defines extinction?

 A Most individuals of a species are dead.

 B The genetic variation of a population is drastically reduced.
 C All individuals of a species are permanently lost.
 D The death rate of a population is higher than the birth rate.
 E A species is at risk of not having any surviving members.

Q3: 

In speciation, what conditions change and provide the selective pressure?

 A Communicative
 B Temporary
 C Environmental
 D Phenotypic
 E Genetic

Q4: 

Fill in the blanks: In isolated populations, different______  mean different ______ are

advantageous and are therefore selected for.

 A genomes, genes
 B alleles, environments
 C conditions, environments
 D environments, alleles
 E alleles, genes

Q5: 

What is the process by which two species evolve from a single-origin species due to
isolation?

 A Specialization
 B Differentiation
 C Classification
 D Speciation
 E Determination

Q6: 

In speciation, which of the following options best describes when the separated
population and the original population are considered to be two distinct species?

 A When individuals from each group specialize to eat a certain food source
 B When individuals from each group look different enough
 C When there has been one set of offspring from every individual in the new group
 D When individuals from each group can no longer reproduce to create fertile
offspring
 E When individuals from each group can no longer mate
Q7: 

Which scientific term is given to the following definition: A group of similar

organisms that reproduce to give fertile offspring?

 A Kingdom
 B Species
 C Genus
 D Class
 E Family
Which of the following is a correct definition of speciation?
An evolutionary process that leads to the formation of a new species. The process by which
a species goes extinct, allowing a new species a chance to live in a new habitat.

When a species has a genetic defect, making it a brand new species. When one species
has a genetic mutation, allowing it to breed with another species.

 Which of the following is an example of speciation?

a. A population of a frog species becomes geographically isolated from its original
population. Over time, new traits are favored best suited to their new environment.
Over many generations, this population of frog is no longer be able to breed with the
original population of frog.
b. One individual frog living in a pond is moved to a new area. This frog finds a closely
related frog to mate with. They reproduce and their offspring is a new species.
c. A flood separates a population of a frog species. They have a new environment to live
in. They survive and reproduce in a new area. Their offspring are a new species.
d. A group of frogs migrate to a new area to seek food. They reproduce and populate the
new area. Over many generations they keep reproducing.

11. Speciation events are triggered by interruptions in gene flow. Distinguish between allopatric
speciation and sympatric speciation.

12. What type of speciation is caused by a formidable barrier such as the Grand Canyon?

13. How is the example of the snapping shrimp (Alpheus) evidence of allopatric speciation?
 14. Sympatric speciation occurs in populations that live in the same geographic area. How is this
possible?

15. Your response to question 14 should have included polyploidy, habitat differentiation, and
sexual selection. Choose either polyploidy or habitat differentiation to explain sympatric speciation.

16. Define sexual selection and then explain how sexual selection can drive sympatric speciation.

2. Which of the following is NOT true in the formation of a new species?

a. A mutation causes a population to breed with a different species.
b. A population needs to become reproductively isolated.
c. Reproductive isolation can occur by the formation of a mountain range.
d. If an isolated population has new environmental conditions new traits can be favored
eventually leading to the inability to reproduce with the original population.

es cookies to 

Key points
 The differences in characteristics between individuals of the same species is called variation.

 Some variation is passed on from parents to offspring, via genes during reproduction. This

is inherited variation.

 Some variation is the result of differences in the surroundings, or what an individual does. This is
called environmental variation.

 Genetic and environmental variation

 Characteristics in an individual organism are caused by both genetic

and environmental variation.

 For example, the weight of a dog is caused partly by its genes -

inherited - and partly by what it eats - environmental. The height of a
sunflower is caused partly by its genes and partly by how much light
and water it gets.
Quiz - Genetic or environmental? Or Both?
1.

2.

3.
4. Milk production?
5. Color of flowerhorn
6. Position of flowers (axial or terminal)
7. Being a good basketball player
8. Shape of a flower
9. Cactus sharp spines
 10.

Select a trait to view the relevant environmental factors. Choose one to learn more.

Analyzing and Interpreting Data; Argumentation

Students analyze data from actual studies and generate a claim based on those data.

References:

https://www.khanacademy.org/science/ap-biology/natural-selection/
speciation/a/species-speciation

https://www.bbc.co.uk/bitesize/topics/zpffr82/articles/z6s26yc#:~:text=The
%20differences%20in%20characteristics%20between,or%20what%20an
%20individual%20does.

https://evolution.berkeley.edu/modes-of-speciation/sympatric-speciation/