Species

and Speciation
BIOLOGICAL SPECIES
CONCEPT
What is Species?
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.

Happy Face Spider /Theridion grallator

…in nature, there are lots
of places where it is
difficult to apply this
definition.
For example, many bacteria
reproduce mainly
asexually. The bacterium
shown at right is
reproducing asexually, by
binary fission.
….cannot be easily applied
to organisms that
reproduce only or mainly
asexually.
Many plants, and some animals, form hybrids
in nature.
Hooded crows and carrion crows look
different, and largely mate within their own
groups—but in some areas, they hybridize.
Should they be considered the same
species or separate species?
Many modern biologists, particularly zoologists,
define species according to the "biological
species concept," a definition that has been
forcibly advocated by Ernst Mayr.

According to this definition, a species is a group

of organisms reproductively isolated from other
organisms.

In other words, a species is a group of organisms

that interbreed and produce fertile offspring only
with one another.
 Biological Species Concept
The biological species concept defines a
species as members of populations that
actually or potentially interbreed in nature,
not according to similarity of appearance.
Although appearance is helpful in identifying
species, it does not define species.
…..appearance isn’t everything

Organisms may appear to be alike and be different species.

For example, Western meadowlarks (Sturnella neglecta) and
Eastern meadowlarks (Sturnella magna) look almost
identical to one another, yet do not interbreed with each
other—thus, they are separate species according to this
definition.

The Western meadowlark (left) and the Eastern meadowlark

(right) appear to be identical, and their ranges overlap, but
their distinct songs prevent interbreeding.
Organisms may look
different and yet be
the same species.
They are sisters—two
ants of the species
Pheidole barbata,
fulfilling different
roles in the same
colony.
 Adding to the problem

We already pointed out two of the difficulties

with the biological species concept:
what do you do with asexual organisms,
and
what do you do with organisms that
occasionally form hybrids with one
another?
Other difficulties include:
What is meant by “potentially interbreeding?”

If a population of frogs were divided by a

freeway, as shown below, that prevented the
two groups of frogs from interbreeding, should
we designate them as separate species?

Probably not—but how distantly separated do

they have to be before we draw the line?
Ring species are species with
a geographic distribution that
forms a ring and overlaps at
the ends.
The many subspecies of
Ensatina salamanders in
California exhibit subtle
morphological and genetic
differences all along their
range.
They all interbreed with their
immediate neighbors with
one exception: where the
extreme ends of the range
overlap in Southern
California, E. klauberi and E.
eschscholtzii do not
interbreed.
So where do we mark the point
of speciation?
Other Species Concepts

The biological species concept has its

limitations (although it works well for many
organisms and has been very influential in
the growth of evolutionary theory).

In order to address some of these

limitations, many other “species concepts”
have been proposed, such as:
Recognition species concept: a
species is a set of organisms that can
recognize each other as potential
mates.

Note: Even though these two frogs have been

prevented from mating, the fact that they
recognize each other as potential mates makes
them the same species under the recognition
species concept.
Phenetic species concept: a species is
a set of organisms that are
phenotypically similar and that look
different from other sets of organisms.

According to this concept, phenotypic similarity is all that

matters in recognizing separate species. Since the frogs
depicted here look the same—even though they are
prevented from mating with each other—they would be
considered the same species according to the phenetic
species concept.
Phylogenetic species concept: a species is a
“tip” on a phylogeny, that is, the smallest set of
organisms that share an ancestor and can be
distinguished from other such sets.
Under this definition, a ring species is a single
species that encompasses a lot of phenotypic
variation.
 What is 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:
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.
Causes of Speciation
Geographic Isolation
In the fruit fly example, some fruit fly larvae
were washed up on an island, and
speciation started because populations
were prevented from interbreeding by
geographic isolation.
Geographic isolation is a common way
for the process of speciation to begin:
rivers change course, mountains rise,
continents drift, organisms migrate,
and what was once a continuous
population is divided into two or more
smaller populations
Reduction of Gene Flow
Speciation might also happen in a population with no
specific extrinsic barrier to gene flow. In a situation in
which a population extends over a broad geographic
range, and mating throughout the population is not
random.
Individuals in the far west would have zero chance of
mating with individuals in the far eastern end of the
range.
So, gene flow was reduced, but not total
isolation. This may or may not be sufficient to
cause speciation.
Speciation would require different selective
pressures at opposite ends of the range,
which would alter gene frequencies in groups
at different ends of the range so much that
they would not be able to mate if they were
reunited.
Most biologists basically accept the
biological species concept, but argue
that it is really an idealized concept,
since only rarely is an actual test for
reproductive compatibility performed.

Thus, in practice, nearly all biologists

think of species as morphologically
distinct groups of organisms.