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.
