BLG144: BIOLOGY II

Chapter 24: Speciation

Week # 3 – Part 1
Department of Chemistry & Biology
Tarushika Vasanthan, PhD
Faculty of Science
Department of Chemistry & Biology

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Introduction to Speciation

• If gene flow stops, allele frequencies in isolated

populations can diverge
• The populations begin to evolve independently
• Divergence may occur as a result of mutation,
natural selection, and genetic drift

• This genetic divergence may eventually result in

speciation

• Speciation is a splitting event that creates 2 or more

distinct species from an ancestral species
• Can occur gradually or abruptly
• When speciation is complete, a new branch has
been added to the tree of life

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How Are Species Defined and Identified?

• A species is defined as an evolutionarily independent population or group of populations

• Biologists commonly use four criteria for identifying species:

1. The biological species concept

2. The morphological species (morphospecies) concept
3. The ecological species concept
4. The phylogenetic species concept

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The Biological Species Concept

• According to the biological species concept, the main criterion for identifying species is
reproductive isolation:
• Results in lack of gene flow between populations
• Members of the populations do not interbreed, or fail to produce viable, fertile offspring
after mating

Western Meadowlark Eastern Meadowlark Pheidole barbata

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The Biological Species Concept

• Biologists categorize the mechanisms that stop gene flow between populations (i.e.,
reproductive isolation) as either
• Postzygotic isolation: The hybrid offspring do not survive or reproduce
• Hybrid viability
• Hybrid stability

• Prezygotic isolation: Individuals of different species are prevented from mating successfully
• Ecological
• Behavioral
• Temporal
• Mechanical
• Gametic

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Mechanisms of Reproductive Isolation

Prezygotic Isolation

Ecological Behavioral Temporal Mechanical Gametic

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The Biological Species Concept

• The biological species concept has disadvantages:

• Reproductive isolation cannot be evaluated in
• Fossils
• Species that reproduce asexually

• It cannot be applied to populations that do not overlap geographically

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The Morphospecies Concept

• According to the morphospecies concept, individual lineages differ in size, shape, or other
morphological feature
• Distinguishing features most likely arise if populations are independent and isolated from
gene flow

• It is widely applicable
• Useful when there is no data on the extent of gene flow
• Equally applicable to sexual, asexual, and fossil species

• Disadvantages:
• One polymorphic species may be classified as more than one species
• It cannot identify cryptic species that differ in non-morphological traits
• The features used to distinguish species under this concept are subjective

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The Phylogenetic Species Concept

• Identifies species based on evolutionary history

• Based on the rationale that all species are related by common ancestry

• On phylogenetic trees, a monophyletic group consists of an ancestral population plus all of its
descendants
• Also called a clade or lineage

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The Phylogenetic Species Concept

• Monophyletic groups are identified by

synapomorphies (“unique forms”):

• Homologous traits found in common

ancestor and descendants but
missing in more distant ancestor

• Example: Fur or milk-producing

glands in mammals

• Therefore, a species is defined as the

smallest monophyletic group on the tree
of life

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The Phylogenetic Species Concept

• The phylogenetic species concept has two distinct advantages:

1. It can be applied to any type of population (e.g., fossil, asexual, or sexual)

2. It is logical because different species have different synapomorphies due to lack of gene
flow and independent evolution

• The phylogenetic species concept also has disadvantages:

• Phylogenies are currently available for only a tiny (though growing) subset of populations
on the tree of life

• Critics point out that it would probably lead to recognition of many more species than
either of the other species concepts

• But this may actually reflect the extent of life’s diversity

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Species Concept
Criterion for Identifying
Populations as Species
Biological Reproductive isolation between
populations (they don’t breed and
don’t produce viable, fertile
offspring)
Morphospecies Morphologically distinct
populations
Ecological Resources exploited; range of
environmental tolerances
Phylogenetic Smallest monophyletic group
on phylogenetic tree
Advantages
Reproductive isolation =
evolutionary independence
Widely applicable
Applicable to asexual
species
Widely applicable; based on
testable criteria
Disadvantages
Not applicable to asexual or
fossil species; difficult to
assess if populations do not
overlap geographically
Subjective (researchers often
disagree about how much or
what kinds of morphological
distinction indicate
speciation); misidentifies
polymorphic species; misses
cryptic species
Different species may have
similar ecological roles
Relatively few well-estimated
phylogenies are currently
available
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Species Definitions in Action: The Case of the Dusky Seaside Sparrow

• Subspecies are populations that

• Live in discrete geographic areas
• Have distinguishing features
• Are not distinct enough to be considered a separate species
• Can interbreed if geographical barriers are removed

• Traditionally named 6 subspecies of seaside sparrows

• Live in salt marshes along the Atlantic and Gulf coasts
• Populations are physically isolated from one another
• Differ by colouration and songs

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Species Definitions in Action: The Case of the Dusky Seaside Sparrow

• In 1980, only six individuals—all males—remained of the dusky seaside sparrow subspecies

• Biologists launched a conservation program under the Endangered Species Act

• The plan was to breed the males with females from the nearby subspecies
• The hybrid offspring would be used in a reintroduction program by preserving as much
genetic diversity as possible by reestablishing a population of dusky-like birds

• Phylogenetic analysis showed only two distinct monophyletic groups of seaside sparrows exist

• The dusky seaside sparrow was

• genetically indistinguishable from the other Atlantic Coast sparrows
• but different from sparrows from the Gulf Coast lineage
• by crossing females from the Gulf Coast lineage with dusky seaside males
• dusky seaside sparrow went extinct

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CHECK YOUR UNDERSTANDING

Two populations of birds with somewhat different coloration live on opposite sides of a
peninsula. The habitat between the populations is not suitable for these birds. When birds
from the two populations are brought together, they produce young whose appearance is
intermediate between the two parents. These offspring will breed with each other or with
birds from either parent population, and all offspring of these pairings appear intermediate
to various degrees. What keeps the two populations separate?

A) temporal reproductive isolation

B) lack of hybrid viability
C) gametic barrier
D) geographic reproductive isolation
E) behavioural reproductive isolation

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LEARNING OBJECTIVE

Explain the roles that gene flow, selection, genetic drift, and mutation play
in the process of speciation.

Define, compare, and contrast the four species concepts. Describe the
advantages and disadvantages of the four different sets of criteria used to
identify species.

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Isolation and Divergence in Allopatry

• Speciation begins when gene flow between populations is reduced or eliminated, causing
genetic isolation

• Genetic isolation happens when populations become geographically separated

• Populations that live in different areas are in allopatry

• Speciation that begins with geographic isolation is called allopatric speciation

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Isolation and Divergence in Allopatry

Geographic isolation occurs in one of two ways:

1. Dispersal
• A population moves to a new habitat, colonizes it, and founds a new population

2. Vicariance—the physical splitting of a habitat

• A physical barrier splits a population into subgroups that are physically isolated from
each other

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Isolation and Divergence in Allopatry

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Allopatric Speciation by Dispersal – Example

• Colonization events can lead to speciation

• Peter and Rosemary Grant compared parents and offspring from large ground finches that
remained on the island of Daphne Major (colonists) with those from the home island
(migrants)
• Colonists represented a new allopatric population, but could this have led to speciation?
• Average beak size in colonist population was much larger than those of migrant
population (Example of genetic drift)

• Natural selection favoured alleles associated with large beaks

• The Grants concluded that both mechanisms contributed to the change in beak size in the large
ground finches
• New population is not a separate species yet, because there is still some gene flow
• Over time, the populations could continue to diverge

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Allopatric Speciation by Vicariance – Example

• When there is a physical barrier such as a mountain range

uplifting or a river splitting the geographic range of a
species, vicariance has occurred.
• Example: Researchers compared the DNA sequences of
trumpeters from different areas of the Amazon basin to the
geological events that occurred
• They found evidence of isolation of populations by
vicariance
• Initially, the formation of the Amazon split the
ancestral population
• The formation of the river systems then subdivided
the populations

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Isolation and Divergence in Sympatry

• Populations or species that live in the same geographic

area—close enough to interbreed—live in sympatry

• Researchers traditionally believed that speciation could not

occur among sympatric populations because:

• Gene flow would overwhelm any differences among

populations created by genetic drift and natural selection

• Recently, studies show that sympatric speciation can occur

• Speciation that occurs among populations within the
same geographical area

• Serves as a source of new branches on the tree of life

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Isolation and Divergence in Sympatry

• Sympatric speciation can be initiated by two types of events

1. External events
• Example: Disruptive selection based on different ecological niches or mate preferences
2. Internal events
• Example: Chromosomal mutations

• A niche is: the range of ecological resources that a species can use and the range of conditions it
can tolerate

• Even though sympatric populations are not geographically isolated,

• They may be reproductively isolated by adapting to different ecological niches via disruptive
selection

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Isolation and Divergence in Sympatry

• Apple maggot flies mate on apple fruits, and their

larvae use the apple for food source
• Hawthorn maggot flies feed & mate on hawthorn fruits
• Apple maggot flies originated from hawthorn flies after
apples introduced in North America

• Hybrid individuals do not orient to fruit scent as well as

their parents do, resulting in lower reproductive success
(since they do not find fruit and mates as well as do
parents)

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Sympatric Speciation by Polyploidization

• If populations become isolated, it is unlikely that mutation

alone could cause them to diverge appreciably since it occurs
at a low rate

• One type of mutation, polyploidy, is important in speciation

• Occurs when an error in meiosis or mitosis results in more than

two sets of chromosomes

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Sympatric Speciation by Polyploidization

Polyploids may be:

• Autopolyploid
• Individuals produced when a mutation results in a doubling of the chromosome number
• Chromosomes are all from the same species

• Allopolyploid
• Individuals are created when parents of different species mate and an error in mitosis
occurs, resulting in viable, nonsterile offspring
• Produces offspring with two different sets of chromosomes

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Autopolyploidy

• Less common than allopolyploidy

• Example: Researchers found a population of tetraploid (4n) maidenhair ferns within a normal
population
• These were offspring of a parent that produced diploid gametes and then self-fertilized

• Polyploid individuals like these are reproductively isolated from the original population

• Tetraploid individuals can successfully breed with other tetraploids but not with diploids,
because breeding between tetraploid and diploid individuals produces sterile offspring

• According to the biological species concept, speciation occurred in a single generation

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Allopolyploidy

• New tetraploid species may be created

when two diploid species hybridize

• Usually the offspring will be sterile

• If an error occurs during mitosis creating

diploid gametes in the adult offspring,
meiosis proceeds, and gametes fuse
during self-fertilization, a tetraploid
individual results

• Allopolyploid offspring has two copies

each of two sets of chromosomes

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Why is Speciation by Polyploidy so Common in Plants?

• Many diploid plant species have closely related polyploid species

• This supports the claim that speciation by polyploidy is important in plants

• Polyploids have higher levels of heterozygosity than do diploids

• Polyploids can tolerate higher levels of self-fertilization

• they are not as affected by inbreeding depression as are diploids

• Genes on duplicated chromosomes can diverge independently

• This increases genetic variation in the population

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What Happens When Isolated Populations Come into Contact?

• If divergence has occurred and prezygotic isolation exists, then mating between the populations
is rare, gene flow is minimal, and the populations continue to diverge

• When prezygotic isolation does not exist, populations may successfully interbreed

• Gene flow then occurs and may erase distinctions between the two populations

• Other possible outcomes are reinforcement, development of hybrid zones, and speciation by
hybridization

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Reinforcement

• If two populations have diverged extensively and are genetically distinct, the fitness of hybrid
offspring will be lower than the parents’ fitness

• If populations are well adapted to different habitats, then hybrid offspring will not be well
adapted to either habitat

• When postzygotic isolation occurs, there is strong natural selection against interbreeding

• Natural selection for traits that prevent interbreeding among populations is called
reinforcement

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Reinforcement

• Some of the best data comes from laboratory studies of Drosophila

• Closely related sympatric species will seldom mate with one another in the lab

• Allopatric species are often willing to mate with one another in the lab

• This is the pattern expected by reinforcement

• Natural selection can act to reduce mating between species only when their ranges overlap

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Hybrid Zones

• Sometimes the hybrid offspring of diverged populations can mate and produce viable fertile
offspring

• A hybrid zone is a geographic area where interbreeding between two populations occurs and
hybrid offspring are common

• Depending on the hybrid offspring fitness and extent of breeding between parental species,
hybrid zones can be
• Narrow or wide
• Long or short-lived
• Stable in one place or move over time

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Hybrid Zones

• Example: Townsend’s warblers & hermit warblers hybridize extensively where their ranges
overlap; Hybrid offspring have intermediate characteristics relative to the two parental
species

• Data from mitochondrial (mt) DNA show

• Most hybrids form when Townsend’s warbler males mate with hermit warbler females
• Townsend’s males invade hermit territories, drive off hermit males & mate with hermit
females
• In some regions, many Townsend’s warblers have hermit mtDNA

• This result suggests that hermit warblers were once as far north as Alaska
• Townsend warblers have taken over their range, causing the hybrid zone to move south
• The hybrid zone may continue moving south, eventually leading to the extinction of the
hermit warbler

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Hybrid Zones

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New Species Through Hybridization

• Two species may interbreed and produce hybrid offspring that survive, reproduce, and possess
a unique combination of traits adaptive to their particular environment
• A new species might be the result

• Example: Hybridization between two species of sunflowers, which had diverged previously,
produced a new species
• This new species had unique combinations of alleles from each parental species and
therefore different characteristics

• The new species was able to grow in drier habitats

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New Species Through Hybridization

Experimental Evidence That New Species Can Originate in Hybridization Events

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New Species Through Hybridization

• Secondary contact of two populations can produce a dynamic range of possible outcomes
• Fusion of the populations
• Reinforcement of divergence
• Founding of hybrid zones
• Extinction of one population
• Formation of new species

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New Species Through Hybridization
Possible Outcomes of Secondary Contact Between Populations
Process
Fusion of the populations The two populations freely interbreed.
Reinforcement of If hybrid offspring have low fitness, natural selection
divergence favours the evolution of traits that prevent
interbreeding between the populations.
Hybrid zone formation There is a well-defined geographic area where
hybridization occurs. This area may move over time
or be stable.
Extinction of one If one population or species is a better competitor
population for shared resources, then the poorer competitor
may be driven to extinction.
Creation of new species If the combination of genes in hybrid offspring
allows them to occupy distinct habitats or use novel
resources, they may form a new species.
Example
Occurs whenever populations of the same species
come into contact.
Appears to be common in fruit fly species that
occupy the same geographic areas.
Many stable hybrid zones have been described;
the hybrid zone between hermit and Townsend’s
warblers appears to have moved over time.
Townsend’s warblers may be driving hermit
warblers to extinction.
Hybridization between sunflowers gave rise to a
new species with unique characteristics.
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CHECK YOUR UNDERSTANDING

If a plant species with 2n = 14 forms an allopolyploid with a plant species with 2n = 18,
what would be the likely diploid (2n) number of the allopolyploid?

A) 16
B) 28
C) 32
D) 36

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LEARNING OBJECTIVE

Describe the processes of allopatric speciation and sympatric speciation

and give an example of each process.

Explain how polyploidy can lead to speciation.

Predict what may happen when two divergent populations come into
contact again under various circumstances.

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