Lecture 5

Origin of species
• What is a species?
• Reproductive barriers
• Allopatric & Sympatric speciation
• Hybrid Zones

Readings: Campbell Biology, 3rd Canadian

edition
– Chapter 24: p. 530-544, 547 (2nd Can ed.; 530-
 The Origin
of Species

Darwin, 1859
 What is a species?
• Difficult to make universal
definition

• e.g. Morphological species concept:

– Based on morphological similarity
– Historically, most commonly used concept
– How much difference is enough?

• Molecular sequence similarity now

also used (e.g. ‘DNA-barcoding’)
 Biological species concept
Inter-fertility: Populations that
interbreed to produce fertile
offspring

Reproductive Isolation: Do not

normally successfully interbreed in
nature with other species
= No/few ‘Hybrids’

Not always applicable, but usually

 Speciation

• ‘Reproductive
barriers’ inhibit gene
flow between
populations,
allowing

Time
evolutionary
divergence

‘Speciation event’
 Reproductive barriers

Fertilization
Potential
Parent 1 Mating Birth, Healthy,
Development,
etc Fertile
Potential Adult
Parent 2 Offspring

Prezygotic Barriers - Postzygotic Barriers

Act before - Act after
fertilization fertilization
 Prezygotic barriers
Examples:
1) Habitat Isolation
2) Temporal Isolation

3) Behavioural Isolation

4) Mechanical Isolation

5) Gametic isolation (gamete incompatibility)

See textbook pages 538-539 for definitions and examples

 Postzygotic barriers

Examples:
1) Hybrid inviability
2) Hybrid infertility (sterility)

3) [Hybrid breakdown]

See textbook page 539 for definitions and examples

Hybrid infertility (sterility): e.g. Mules

Horse Donkey

Incompatible
chromosome
Mule: organisations
Vigorous,
Textbook page 539:
but Infertile
learn as example!
Evolution of reproductive
barriers

May arise ‘accidentally’ as result of

evolution in isolation

May evolve through natural selection to

reduce inter-species mating that lowers
reproductive success.
(e.g. ‘Reinforcement’: more later)
 Speciation
• Allopatric speciation - Geographic
barrier blocks gene flow between
populations

• Sympatric speciation - New

species arise within range of
parent population
Allo: ‘other’
Sym- (Syn-) : ‘together’
 Allopatric Speciation
1) Geographic barrier separates populations
(A barrier forms,
or migrants cross existing barrier, founding
new population)

2) Evolutionary change
on one or both sides

Time
 Geographic
Barrier (e.g.)

Isthmus of Panama
Formed 10-3 million
years ago, isolating
populations of many
marine species

Images: Anker et al. (2007) Zootaxa 1577: 41–60

 Evolution after
separation
• Natural selection to different
environments (i.e. adaptive
evolution)

• Genetic drift
(especially in small isolated
populations?)
e.g.

Large
Ancestral Migrants found new small
Population population in peripheral
habitat (e.g. island)
 If contact re-established
between evolved allopatric
populations..
1) Complete reproductive barriers evolved:
Populations now classic biological species

Time
..or.. 2) Partial reproductive barriers
evolved: Formation of hybrids where
species contact
Fusion:
Hybrids form readily; have high
fitness: the incipient species
merge into one again.

Reinforcement:
Hybrids have low fitness: Natural
selection strengthens barriers -
Hybridisation gradually ends 
Two good Biological Species

Long-lasting hybrid zone:

(e.g. hybrids have variable fitness;
or are uncommon)
 Sympatric speciation
Requires a barrier to gene flow
within a geographic region.

Some examples:
• Host switching by specialist herbivores or parasites
[See textbook’s apple maggot fly example if interested - p.545-546]

• Disruptive selection (lecture 4)

favouring evolution of reproductive barriers
between individuals with different phenotypes?
• Polyploid speciation, especially in plants
 Polyploid speciation
Allopolyploid example (one mechanism)

Species A
Gamete
Viable but infertile Viable fertile
(sterile) hybrid hybrid

2n = 6

Species B Gamete Mitotic

(or meiotic)
error doubles 2n = 10
chromosome
number

2n = 4
New fertile hybrid, reproductively isolated
from both Species A and Species B
Allopolyploid speciation is an
example of ‘hybrid speciation’

Species A

New Species

Species B
Time

* Common in plants
* Exception to a strict “tree-of-life”
 Speciation and
tree-of-life

Each branching on the

tree-of-life began with
a speciation event

Time
 Summary
• The biological species concept is based on inter-fertility
and reproductive isolation
• Reproductive barriers between species are prezygotic (e.g.
behavioural isolation) or postzygotic (e.g. hybrid infertility).
• Allopatric speciation is the divergence of geographically
separated populations to (begin to) form distinct species.
• Incipient species that re-contact often form hybrids:
• Hybrids can allow gene pools to fuse again; or
reinforcement can gradually suppress hybrid formation.
• Sympatric speciation (i.e. without geographic separation) is
also significant
• In Allopolyploid speciation there are two sympatric
parental species, and a third, new, hybrid species arises. It
is common in some plant groups.