Speciation – Chapter 18

Introduction
• Evolution - the process by which the characteristics of species change
and by which new species come into being.
• Theory of evolution - unifying theory of biology.
‒ Framework within which biologists ask questions about the living world.

Speciation
• For sexually reproducing organisms, a species is defined as a group
of actually or potentially interbreeding organisms that can produce viable,
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fertile offspring.
• Speciation - the formation of two species from one original species.
• For speciation to occur:
‒ Two new populations must form from a single original population.

‒ The new populations must evolve in such a way that it becomes impossible

for members of the two populations to interbreed.

• Two categories of speciation:
‒ Allopatric speciation - speciation due to a geographic
separation of a population from its parent population.
‒ Sympatric speciation - speciation occurring within a parent
species while remaining in one location.
Speciation Through Geographic Separation
• Geographically continuous population:
‒ Relatively homogenous gene pool.
‒ Relatively free gene flow due to the ability of individuals to
move and mate with individuals in a new location.
‒ Allele frequencies at one end of the distribution will be similar to
allele frequencies at the other end.
• Geographically discontinuous population:
 ‒ Free flow of alleles is prevented due to separation.
‒ Two populations will evolve along different trajectories when the separation

lasts for a period of time.

‒ Allele frequencies at many genetic loci will become increasingly

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different as new alleles independently arise by mutation in
each population.
‒ Differing environmental conditions between the two groups will cause

natural selection to favor divergent adaptations in each group.

• Reproductive Isolation - the inability of members of two populations to interbreed.
‒ Genetic and phenotypic divergence will, over time, likely affect

characteristics that influence reproduction.

 If the two populations were brought back together, mating would be
less likely or if mating occurred, the offspring would be non-viable or
infertile.
‒ Mechanisms of Reproductive Isolation

 Prezygotic barriers - operate before fertilization/blocks

reproduction from taking place.
 Postzygotic barriers - operate after fertilization/occurs
after zygote formation.
‒ Prezygotic Barriers

 Temporal isolation - differences in breeding schedules.

• Example: two frog species live in the same geographic area,
but one species reproduces January to March, the other from
March to May.
 Habitat isolation - one population moves to a new
habitat that no longer overlaps with the parent population.
• Example: Two cricket species occupy the same geographic
region, but one species prefers sandy soils and the other
prefers loamy soils.
  Behavioral isolation - the presence or absence of a
specific behavior prevents reproduction.
• Example: Male fireflies use light patterns to attract females, and
different species have unique, specific patterns. Females would
not recognize the light pattern of a male of a different species.
 Gametic isolation - fertilization does occur because
differences in egg and sperm from different species cannot merge.
 Mechanical isolation
• Two forms:
o Mating organs incompatible.
 Example: different species of damselflies have
anatomical incompatibilities that prevent mating
between species.
o Pollinators incompatible.
 Different plant species have structures to attract
particular pollinators or to prevent incompatible
pollinators from accessing nectar. This prevents
cross pollination across species.
‒ Postzygotic Barriers

 Hybrid infertility - Hybrid offspring are unable to

reproduce due to sterility.
• Meiosis does not occur in the germ cells due to parents
contributing different numbers of chromosomes.
 Hybrid inviability - Hybrid offspring do not survive.
• Offspring either do not develop past the embryonic stages or
die before reaching reproductive maturity due to incompatibility
of the parental genes.
  Hybrid breakdown - First generation hybrid offspring are
viable and fertile, but the second generation hybrid offspring have
reduced fitness.
• Two categories of allopatric processes:
‒ Dispersal - if a few members of a species move to a new
geographical area.
‒ Vicariance - if a natural situation arises to physically divide
organisms.
• Numerous examples of allopatric speciation have been documented.
‒ Two subspecies of spotted owls, one in the northwestern US and the other

in the southwestern US and regions of Mexico.

 Differences in climate between the two ecosystems.
 Different sources of prey.
 Differences in behavior and hunting habits.
• Adaptive radiation - situation in which a population of one
species disperses throughout an area and each member finds a distinct
niche/habitat; over time, multiple speciation events can occur.
‒ From one point of origin, the founder species radiates into several new

species.
 Island chains such as the Hawaiian Islands are ideal for adaptive
radiation due to the geographic isolation of each island being
surrounded by water.

Speciation Without Geographic Separation

• Mechanisms for sympatric speciation

‒ Polyploidy - a condition in which a cell or organism has an

extra set or sets of chromosomes.
‒ Two types:

 Autopolyploidy - individual will have two or more complete

sets of chromosomes from its own species.
 • Results from an error in meiosis in which all of the
chromosomes move into a single cell instead of separating.
 Allopolyploidy - occurs when gametes from two different
species combine to form an offspring with two or more complete sets
of chromosomes.
• Can form when a normal gamete from one species fuses with
polyploid gamete from a different species.
o It takes two generations before a viable fertile hybrid is
produced.
‒ Sympatric speciation can also occur in habitats with distinct ecological

zones that cause isolation as some members of the same population

exploit different parts of the habitat.
 Cichlids from Lake Victoria and Nicaragua.
‒ Other examples include host specificity of parasitic species.

Reconnection
• After speciation, two species may recombine or even continue
________________
interacting indefinitely.
• Hybrid zones: area where two closely related species interact and
interbreed.
• After speciation has occurred, the two separate but closely related species may
continue to produce offspring in an area called the hybrid zone.
• Reinforcement, fusion, or stability may result, depending on reproductive barriers
and the relative fitness of the hybrids.
• Reinforcement : process where hybrids are less fit, and thus
there is a nudge towards further divergence between the two species.
• Fusion : Reproductive barriers weaken until the two
species recombine to again become a single species.
• Stability : Fit hybrids continue to be produced.
Rates of Speciation
• How quickly does speciation occur?
• Two current models:
‒ Gradual speciation (aka phyletic gradualism): species diverge
gradually through time with small, incremental changes in traits.
‒ Punctuated equilibrium : species exhibit a large change in a
relatively short period of time followed by long periods of stasis.