Natural selection and

speciation

Chapter 9 of text
 By the end of this lecture you should be able to:
• explain how environmental factors act as forces of natural selection
with reference to antibiotics, Biston betularia, Trinidadian guppies
and the Dominican anole;
• explain how natural selection may be an agent of constancy or an
agent of change with reference to directional, disruptive and
stabilising selection;
• discuss natural selection as a mechanism of evolution, discussing
Darwin’s theory, its observations and conclusions;
• discuss the biological species concept and its limitations;
• explain the process of speciation, including the role of isolating
mechanisms (reproductive, geographic, behavioural and temporal)
and with reference to named examples of allopatric and sympatric
speciation.
 Natural selection
• Natural selection: is the process by which organisms that are better
adapted to their environment survive and breed while those less well
adapted fail to do so. Those that are adapted and so survive to
reproductive age will be the ones that pass on their favourable alleles to
the next generation.
• This causes evolution to occur.
• Evolution: change in the characteristics of organisms over time.
• Charles Darwin and Alfred Wallace in 1856 independently developed the
theory of evolution by natural selection.
• This theory grew out of four observations and three logical deductions
from these observations.
 Natural selection
Table of Darwin’s observations and deductions
Observations Deductions
• All organisms over-reproduce (i.e. • There is competition for survival –
more offspring are produced than are the ‘struggle for existence’
required to keep the population at a • Individuals with characteristics
steady state) that best adapt them for their
• Population numbers tend to remain environment are most likely to
fairly constant over long periods of survive and reproduce.
time. • If these characteristics can be
• Organisms within a species vary. inherited, then the organisms will
• Some of these variations are pass the characteristics on to their
inherited. offspring.
 Natural selection in action

• Antibiotic resistance in bacteria

• Pesticide resistance in insects
• Industrial melanism
• Trinidadian guppies
• Dominican anole
Antibiotic resistance in bacteria
• Staphylococcus aureus
• Methicillin-resistant Staphylococcus aureus (MRSA)
Pesticide resistance in insects
 Industrial melanism

Pre 1800s
Light form common
throughout Britain
Dark arose by mutation-
very rare

Pale form most abundant in non-

industrial areas.
Dark forms most abundant in
areas suffering from heavy air-
pollution.
 Experiments on peppered moths
• Bernard Kettlewell investigated the effect of predation by birds as the selective agent in
maintaining populations of peppered moths in the 1950s and 1970s.
• He trapped and reared melanic (dark) and non-melanic (light) forms, marked them and then
released them in a wood in an industrial area of Birmingham and a wood in a rural area where
the trees were covered in lichen.
• After a few days he used a moth trap to recapture the moths. He obtained the following data:

Describe and explain the

data shown in the table.
 Guppies in Trinidad
• Guppies, Poecilia reticulata, are small fish that live throughout the
Caribbean.
• In Trinidad the effects of predation on populations of these fish in the
Aripo river system have been studied by David Reznick.
• The guppies living in rivers with predators grow faster and mature at
smaller sizes, they also reproduce at a younger age than those in
rivers without predators. WHY?

• Reason: Guppies in rivers with predators breed as soon as possible

because they are at risk of being eaten.
 Experiments on guppies in Trinidad
• Investigations of the guppies involved moving individuals from each
population and putting them into rivers which did not have guppies.
• Some of these new rivers had predators in them, others did not.
• So populations were moved from rivers with predators to rivers without
predators and from rivers without predators to those with predators.
• As a control, populations were moved from rivers with predators to
different rivers with predators and the same was done for populations
from rivers lacking predators.
• The fish were monitored for eleven years.
• Those in rivers without predators the body size increased and sexual
maturation occurred later.
• In rivers with predators the fish were smaller and matured earlier.
• The guppy populations changed to maximise their chances of reproduction
by adopting different strategies depending on the presence or absence of
predators.
 Dominican anole
• The Dominican anole, Anolis oculatus, is endemic to Dominica. These tree-dwelling lizards live
in a variety of habitats from coastal woodlands to rainforests.

Montane Ecotype Anole is found in high

North Ecotype Anole is found in the most arid
part of Dominica in low, scrub-like woodland on
the northwest coast.
• It can have the most complex markings
• Its ground colour is predominantly gray or
pale or yellow brown
elevation rain forests located in central
Dominica
• It has a deep green ground colour, which
matches the moss-covered tree trunks
• It has the largest size and small, bluish
white spots
 Dominican anole
• Common garden rearing experiments on the Dominican anole suggest that the characters
showing geographic variation have genetic variance.

South Ecotype Anole is

found in the south and
southwest coast
• It is the smallest in size
and the palest in
colour and markings
• Colour: Light tan to
Atlantic Ecotype Anole is found along yellow ground colour
most of the Atlantic (East) coast of because of the Sulphur
Dominica which is wetter than the west in the soil
• Intermediate size
• Orange to chocolate brown ground
colour
 What are the factors that keep the population size in check?
• These factors are divided into
biotic or abiotic factors.
• Biotic factors: caused by other
living organisms such as predation,
competition for food or infection
by pathogens.
• Abiotic factors: caused by non-
living components of
environment-such as water supply
or nutrient levels.
 Selection pressures
• The environmental factors that limit the population of a species are
called selection pressures.
• Selection pressures include:
• Competition for food
• Competition for a space in which to live, breed and rear young
• Need for light, water, oxygen, etc
• Climate changes, e.g. temperature, rainfall, wind/water currents
• Predation
• Disease

NB: The selection pressures determine the characteristics that would be

best suited and most likely to survive in a population.
 Main types of selection
• Directional – favours individuals
that vary in one direction from
the average of the population;
e.g. antibiotic resistance in
bacteria, peppered moth
• Disruptive – favours individuals
at the extremes rather than
those around the average of the
population; e.g. beak size
• Stabilising – favours the average
individuals; e.g. body mass of
human children at birth or sickle
cell anaemia
 Sickle cell anaemia and protection against malaria
• The parasite, Plasmodium, that causes malaria
spends part of its life cycle in the red blood cells.
• Red blood cells that show sickling die prematurely.
Therefore, if they are infected with the malarial
parasite, the parasite will also die without
reproducing.
• In persons who are heterozygous for sickle cell, the
presence of the malarial parasite in the red blood
cells will also cause the red blood cells to rupture
prematurely, so the parasite cannot reproduce.
• So in countries where malaria is an important
infectious disease, the majority of the population will
be heterozygous for sickle cell anaemia.
 Speciation
• Speciation is the evolution of new species from existing
ones.
• A species is a group of individuals with similar features which
are capable of breeding with one another to produce fertile
offspring.
• Individuals which do not interbreed to produce fertile
offspring under normal circumstances are regarded as
reproductively isolated and they belong to separate species.
 Limitations of the biological species concept
• Can only be used for organisms that reproduce sexually hence cannot
be used to determine if asexually reproducing organisms belong to
the same or different species.
• In nature the coyote (Canis latrans) and the wolf (Canis lupus) do not
normally interbreed and are therefore classed as different species
even though they are capable of interbreeding and producing fertile
offspring in captivity.
• Similarly tigers and lions are reproductively isolated and belong to
different species: male tiger + female lion → tiglons or tigons (fertile)
while male lions + female tiger → ligers (infertile)
• Donkeys and horses can interbreed but produce a mule (infertile) so
they belong to different species
 How new species are formed
• New species can arise in two different ways:

• Reproductive isolation followed by genetic change due to

natural selection (occur during allopatric and sympatric
speciation).

• Cross fertilisation between individuals of two different

species (occur during sympatric speciation only).
 Two main forms of speciation

• Allopatric – production of a
new species as a result of
geographical isolation

• Sympatric – production of
a new species due to
reproductive isolation
without any geographical
separation
 Allopatric speciation
• If individuals of a species migrate to occupy a new area, they are
exposed to different selection pressures compared with the area that
they have left.

• Over time this may lead to changes in the isolated population to the
extent that they cannot interbreed with the original population.

• E.g. a snapping shrimp, Alpheus, were separated when the isthmus of

Panama was formed 3 million years ago.

• Populations on either side of the isthmus are very similar, but when
males and females from these two populations are put together they
‘snap’ at one another and do not mate.
 Sympatric speciation

• Speciation may occur within a population.

• Usually this is an abrupt change in a species, so that individuals

are not able to interbreed.

• All of the reproductive isolating mechanisms apply except

geographical.
 Forms of reproductive isolating mechanisms
• Geographical – populations are isolated by physical barriers
such as oceans, mountain ranges, rivers, etc.
• Ecological – populations inhabit different habitats within the
same area and so individuals rarely meet.
• Temporal – the breeding seasons of each population do not
coincide and so they do not interbreed.
• Reproductive – the breeding seasons coincide but
anatomical differences may prevent interbreeding
• Behavioural – the breeding seasons overlap but differences
in courtship rituals do not stimulate mating and prevents
interbreeding
Summary of the isolating
mechanisms that lead to
Allopatric and Sympatric
speciation
North Caribbean
ecotype
Summary

Montane ecotype
Atlantic ecotype
South Caribbean
ecotype

• Organisms with certain inherited characteristics are more likely to survive and
reproduce than organisms which do not have these characteristics.
• There are three types of natural selection; stabilising, directional and disruptive.
• Speciation is a process by which one species evolve from existing species.
• Forms of speciation include allopatric and sympatric.