CHAPTER 24

THE ORIGIN OF SPECIES

Learning objectives

What Is a Species?
1. Define Ernst Mayrs biological species concept.
A species is a group of populations whose members have the potential to interbreed in
nature and produce viable, fertile offspring; they do not breed successfully with
members of other such groups.
o Ex. A business woman in NY is unlikely to meet a farmer in Cali but they could
have viable babies who develop into fertile adults. In contrast, humans and
chimpanzees remain distinct biological species, even where they live in the
same region because many factors keep them from interbreeding and
producing fertile offspring.
Emphasizes reproductive isolation, meaning species tend to only breed within the species in
order to produce viable fertile offspring.
2. Distinguish between prezygotic and postzygotic reproductive barriers.
Prezygotic reproductive barriers (before the zygote) block fertilization from
occurring: habitat, temporal, behavioral, mechanical, gametic isolation.
o Work in 1 of three ways:
1. Impeding different species from attempting to mate
2. Preventing the successful completion of mating
3. Hindering fertilization if mating is completed successfully
Postzygotic reproductive barriers (after the zygote) prevent the hybrid zygote from
developing into a viable, fertile adult.
o Work in three ways:
1. Reduce hybrid viability
2. Reduce hybrid fertility
3. Hybrid breakdown
Prezygotic barriers block fertilization from occurring in one of three ways all occurring before
fertilization. It can block fertilization by impeding the species from attempting to mate, by
preventing the successful completion of mating, and lastly by hindering fertilization if mating is
completed successfully.
Postzygotic barriers prevent the hybrid zygote from becoming a successful fertile adult by
reducing hybrid viability, meaning developmental errors that may reduce survival rate before
reproducing.
3. Describe five prezygotic reproductive barriers and give an example of each.
Habitat isolation occurs when two species rarely encounter one another, or at all, because they
occupy different habitats, even though not isolated by physical barriers.
o Ex. Africa has both lions and tigers that could interbreed, but lions live in grasslands
while tigers live in forests. The two would never encounter one another, therefore
never reproduce.
Temporal isolation occurs when species that could interbreed do not due to mating differences
in times of the day, seasons, or different years and cannot mix their gametes.
o Ex. Some crickets become sexually mature at different seasons, some in the springs
and other in autumn and therefore could not reproduce.
Behavioral isolation occurs when courtship rituals and other behaviors unique to a species are
effective reproductive barriers even between closely related species.
o Ex. Blue-footed boobies perform a courtship ritual unique to their species which
involved a high-step that draws the females attention to its bright blue feet.
Mechanical isolation occurs when morphological differences prevent successful mating
completion.
 o Ex. Plants who do not have the correct shape of flower to match up with a natural
pollinator will not receive a pollen transfer.
Gametic isolation occurs when sperms of one species may not be able to fertilize the eggs of
another species.
o For instance, sperm may not be able to survive in the reproductive tract of females of
the other species, or biochemical mechanisms may prevent the sperm from
penetrating the membrane surrounding the other species eggs.
Ex. sea urchins release their sperm and eggs into the water where they fuse
and form a zygote. It is difficult for gametes of other species such as the red
and purple urchins to fuse because proteins on the surface of the eggs and
sperm bind very poorly to each other.
1. Temporal isolation describes species who breed at different times of day, seasons, or years cannot mix
gametes with other species. Ex. Bullfrogs and red-legged frogs. 2. Behavior isolation describes the
barriers between species due to courtship rituals and other behaviors unique to that species. Ex. blue
footed-booby who dances for love will be ignored by a female bird who prefers to see salsa dancing. 3.
Mechanical isolation refers to reproductive organs not being compatible. Ex. A fly cannot mate with a
rodent. 4. Gametic isolation refers to sperm of one species not being compatible to with the egg of
another species failing to form a zygote. Ex. Sperm of a chimpanzee cannot penetrate the egg of a
human. 5. Habitat isolation refers to species not being able to mate living in different habitats or
preferred place of reproduction. Ex. certain bird species prefer to mate in trees, ground, etc preventing
this bird species to mate with other species who prefer a different location.
4. Explain how hybrid breakdown maintains separate species even if fertilization occurs.
Hybrid breakdown: some first-generation hybrids are viable (surviving/living successfully) and
fertile, but when they mate with one another or with either parent species, offspring of the
next generation are feeble or sterile.
Strains of cultivated rice have accumulated different mutant recessive alleles at two loci in the
course of their divergence from a common ancestor. Hybrids between them are vigorous and
fertile, but plants in the next generation that carry too many of these species recessive alleles
are small and sterile. Although these rice strains are not yet considered different species, they
have begun to be separated by postzygotic barriers.
Hybrids are usually less fit than their parent species, when fertilization is successful between hybrids it
allows for reinforcement of reproductive barriers. Reproductive barriers would then reduce the rise of
unfit hybrids and over time the overall rate of hybridization would decrease. In other cases fertile hybrids
conceive sterile or feeble offspring
5. Describe some limitations of the biological species concept.
The biological species concept emphasizes the absence of gene flow and cannot be applied to fossils
because sexual reproduction cannot be fossilized, or asexual organisms including all prokaryotes since
they reproduce by themselves.
However gene flow can occur between some distinct species
6. Define and distinguish among the following: ecological species concept, phylogenetic species
concept, and morphological species concept.
Ecological species concept defines a species in terms of its ecological niche, the sum of how
members of the species interact with the nonliving and living parts of their environment.
o Ex. two species of oak trees might differ in their size or in their ability to tolerate dry
conditions, yet still occasionally interbreed. Because they occupy different ecological
niches, these oaks would be considered separate species even though they are
connected by some gene flow.
o Can accommodate asexual and sexual species.
Phylogenetic species concept defines a species as the smallest group of individuals that share a
common ancestor, forming one branch on a phylogenetic tree.
o Applies to sexual and asexual species, but can be difficult to determine the degree of
difference required for separate species
Morphological species concept distinguishes a species by body shape and other structural
features.
 o Applies to asexual and sexual organisms and can also be used even without the
information on the extent of gene flow unlike ecological and phylogenetic species
concepts. Because Morphological species concept relies on subjective criteria, there
may be disagreements in which structural features distinguish a species.

Modes of Speciation
7. Distinguish between allopatric and sympatric speciation.
In allopatric speciation gene flow is interrupted when a population is divided into
geographically isolated subpopulations.
Ex. water level in a lake may subside resulting in two or more smaller lakes that
are now home to separate populations.
o Allopatric speciation can also occur without geographic remodeling.
Ex. such as when individuals colonize a remote area and their descendants
become geographically isolated from the parent population.
o Importance of allopatric speciation is also suggested by the fact that regions that are
isolated or highly subdivided by barriers typically have more species than do otherwise
similar regions that lack such features.
o Reproductive isolation between two populations generally increases as the geographic
distance between them increases, a finding consistent with allopatric speciation.
o While geographic isolation prevents interbreeding between members of allopatric
speciation, physical separation is NOT a biological barrier to reproduction.
o Biological barriers prevent interbreeding when members of different populations come
into contact with one another.
Sympatric speciation occurs in populations that live in the same geographic area (no physical
barriers).
o Although members remain in contact with each other (ongoing gene flow) making
sympatric speciation less common than allopatric speciation, sympatric speciation can
occur if gene flow is reduced by such factors as polyploidy, sexual selection, and
habitat differentiation (same factors can also promote allopatric speciation).
o Can also be driven by sexual selection.
Cichlids experiment.
o Can also occur when a subpopulation exploits a habitat or resource not used by the
parent population (a switch to a habitat or food source not used by the parent
population).
Ex. apple maggot fly and pear fly.
8. Define allopatric speciation. Describe the mechanisms that may lead to genetic divergence of
isolated gene pools.
In allopatric speciation gene flow is interrupted when a population is divided into
geographically isolated subpopulations.
Ex. water level in a lake may subside resulting in two or more smaller lakes that
are now home to separate populations.
o Allopatric speciation can also occur without geographic remodeling.
Ex. such as when individuals colonize a remote area and their descendants
become geographically isolated from the parent population.
o Importance of allopatric speciation is also suggested by the fact that regions that are
isolated or highly subdivided by barriers typically have more species than do otherwise
similar regions that lack such features.
o Reproductive isolation between two populations generally increases as the geographic
distance between them increases, a finding consistent with allopatric speciation.
o While geographic isolation prevents interbreeding between members of allopatric
speciation, physical separation is NOT a biological barrier to reproduction.
 o Biological barriers prevent interbreeding when members of different populations come
into contact with one another.
The mechanisms that may lead to genetic divergence of isolated gene pools occurs once
geographic separation has occurred and the separated gene pools may diverge. Different
mutations arise, natural ways selection, and genetic drift may alter allele frequencies in
different ways in the separated populations. Reproductive isolation then evolves as a by-
product of the genetic divergence that results from selection or drift.
9. Explain how reproductive barriers evolve.
Reproductive barriers evolve in many ways however, they are initiated with a form of species
separation followed by genetic drift/ sexual selection/ or other forms of gene flow block to
eventual divergence and isolation.
10. Define sympatric speciation and explain how polyploidy can cause reproductive isolation.
Sympatric speciation occurs in populations that live in the same geographic area (no physical
barriers).
o Although members remain in contact with each other (ongoing gene flow) making
sympatric speciation less common than allopatric speciation, sympatric speciation can
occur if gene flow is reduced by such factors as polyploidy, sexual selection, and
habitat differentiation (same factors can also promote allopatric speciation).
o Can also be driven by sexual selection.
Cichlids experiment.
o Can also occur when a subpopulation exploits a habitat or resource not used by the
parent population (a switch to a habitat or food source not used by the parent
population).
Ex. apple maggot fly and pear fly.
Polyploidy: the presence of extra sets of chromosomes due to accidents during cell division.
o Polyploidy can cause reproductive isolation by creating a new species that is no longer
able to reproduce with other species but other accidental polyploidy species.
Ex. Failure of cell division in a plant can double a cell's chromosome number
from 2n to 4n. This new plant can produce fertile offspring by self-pollinating
or by mating with plants like it. This leads to these new 4n plants to be
reproductively isolated from 2n plants of the original population.
11. Distinguish between autopolyploid and an allopolyploid speciation and describe examples of each.
Autopolyploid is an organism that has more than two chromosome sets that are all derived
from a single species.
o Ex. In plants a tetraploid plant (4n) can be produced from a 2n plant who did not cell
divide properly. The new tetraploid species can now only self-pollinate or mate with
other tretraploids.
Allopolyploid speciation occurs when two different species interbreed and produce a hybrid
offspring who then doubles its chromosomes and makes a new species that is sterile.
o Ex. Oats, cotton, potatoes, and wheat. These plants are fertile when mating with each
other, but cannot interbreed with either parent species representing a new biological
species.
12. Explain how habitat differentiation has led to sympatric speciation in North American maggot flies.
The fly's original habitat was the native hawthorn tree, when new apple trees were introduced
some populations colonized on those rather than the hawthorn trees. The Apple maggot flies
normally mate on or near their host plant resulting in a prezygotic barrier, habitat isolation,
between populations that fed on apples and populations that fed on hawthorn trees. Because
applies mature more quickly than hawthorn fruit, favored apple-feeding flies with rapid
development creating a temporal isolation between the two subspecies providing a prezygotic
restriction to gene flow between the two subpopulations.
13. Explain how sexual selection has led to sympatric adaptive radiation in the cichlids of Lake Victoria.
Explain how the process of speciation may be reversing, due to pollution in this lake.
Genetic evidence shows that the many species of cichlids originated from a small number of
colonizing species 100,000 years ago all from different rivers feeding into the lake. Sexual
selection for mates of different colors has likely contributed to speciation in cichlids fish in Lake
Victoria. Pollution in this lake maybe causing female fish to mistakenly mate with different
colored cichlids because they are unable to distinguish from their own species in polluted
water causing speciation to reverse.

Hybrid Zones
14. Define the term hybrid zone. Describe the three outcomes for hybrid zones over time.
Hybrid zone refers to the region where members of different species meet and mate,
producing an offspring of mixed ancestry.
o Three possible outcomes for the hybrid zone over time include:
1. Reinforcement of reproductive barriers, hybrids are less fit than the parent
species and the rate of hybridization decreases
2. Fusion of species where species come together and breed so frequently,
eventually fuse into one species due to their increasingly alike gene pool.
3. Stability can occur when hybridization continually occurs in many cases
because those offspring can survive or reproduce better than the members of
the parent species, in some cases continual hybridization occurs even though
they are strongly selected against.
15. Describe examples that illustrate rapid and gradual speciation events.
Rapid speciation appears to have reproduced the wild sunflower Helianthus anomalus.
Evidence suggests this species originated by the hybridization of two other sunflower species.
This sunflower changed most as it branched from its parent species and only changed a little
more for the rest of its life showing a more gradual speciation.
Apple maggot fly and Lake Victoria cichlid suggest new species can arise rapidly once
divergence begins. Gradual speciation occurred in the shrimp.
16. Explain how a small number of genetic changes may lead to speciation in plants and animals.
A small number of genetic changes may lead to speciation in plants and animals quickly if those
genes happen to create a mechanical reproductive barrier preventing them from reproducing
or reproducing viable offspring.
In other organisms the speciation process is influenced by larger numbers of genes and gene
interactions.
o Ex. genetic interactions among a certain number of loci can contribute to hybrid
sterility between fruit fly species.