SELECTION 

 Selection is the process by which those organisms which appear physically, physiologically
and behaviour better adapted to the environment survive and reproduce; those organisms not
so well adapted either fail to reproduce or die.
 The former organisms pass on their successful characteristics to the next generation, whereas
the latter do not.
 Selection depends on the existence of phenotypic variation within the population and is part
of the mechanism by which a species adapts to its environment.
 When a population increases in size, certain environmental factors become limiting such as
food availability in animals and light in the case of plants. This produces competition for
resources between members of population.
 Those organisms exhibiting characteristics which give them a competitive advantage will
obtain the resource, survive and reproduce.
 Organisms without these characteristics are at a disadvantage and may die before reproducing
 Both the environment and population size operate together to produce a selection pressure
which can vary in intensity.
 Therefore selection is the process determining which alleles are passed on to the next
generation by virtue of the relative advantage they show when expressed as phenotypes.
 Selection pressure can then be seen as a means of increasing or decreasing the spread of an
allele within the gene pool and these changes in allele frequency can lead to evolutionary
change.
 There are three types of selection process occurring in natural and artificial populations and
they are described as stabilising, directional and disruptive.
STABILISING SELECTION
 It occurs in populations and tends to eliminate extremes from the population.
 For example, there an optimum wing length for a hawk of a particular size with a certain
mode of life in a given environment.
 Stabilising selection will eliminate those hawks with wing spans larger or smaller than this
optimum length.
 Stabilising selection pressures do not promote evolutionary change but tend to maintain
phenotypic stability within the population from generation to generation. 

DIRECTIONAL SELECTION
 This form of selection operates in response to gradual changes in environmental conditions.
 It operates on the range of phenotypes existing within the population and exerts selection
pressure which moves the mean phenotype towards one phenotypic extreme.
  Once the mean phenotype coincides with the new optimum environmental conditions,
stabilising selection will take over.
 This kind of selection brings about evolutionary change by producing a selection pressure
which favours the increase in frequency of new alleles within the population.
 Directional selection forms the basis of artificial selection where the selective breeding of
phenotypes showing desirable traits increases the frequency of those phenotypes within the
population.

DISRUPTIVE SELECTION
 This is probably the rarest form of selection but can be very important in bringing about
evolutionary change.
 Fluctuating conditions within the environment, may favour the presence of more than one
phenotype within a population
 Selection pressures acting from within the population as a result of increased competition may
push the phenotypes away from the population mean towards the extremes of the population.
 This can split a population into  two subpopulations.
 If gene flow between the two subpopulations is prevented, each population may give rise to a
new species.
NATURAL SELECTION AS A MECHANISM OF EVOLUTION
 Evolution will forever be linked to the name of Charles Darwin (1809–1882)
 Charles Darwin was a British naturalist who proposed the theory of biological evolution by
natural selection.
 Darwin defined evolution as "descent with modification," the idea that species change over
time, give rise to new species, and share a common ancestor.
 The mechanism that Darwin proposed for evolution is natural selection.
 His theory of natural selection stated that: 
1. There is competition for resources between individuals of the same species that have the same
means to obtain those resources; animals compete for water, food, territories, nesting sites and
mates; plants compete for space, water, ions, light, carbon dioxide and, in some cases,
pollinators.
2. Competition leads to high death rates among young animals that starve, are eaten by predators
or die of disease; there is a similar high mortality among seedling plants that start growing in
unsuitable places, do not absorb enough water, ions, light or carbon dioxide, are eaten by
herbivores or are killed by disease 
3. In this struggle for existence the individuals best adapted to obtain resources survive to breed
and pass on their alleles. 
4. This differential survival means that populations consist of those individuals best adapted to
the conditions existing at any one time.
5. The best adapted animals are good at finding food, escaping from predators, resisting disease
and finding mates. 

Darwin’s finches
 Darwin collected specimens of finches, from the islands he visited. 
 They are now known as Darwin’s finches; he collected around 14 species from the Galápagos
Islands. 
 At first Darwin did not appreciate the significance of the Finches, he was content to simply
collect samples and continue on his voyage. It was not until his return to England that it
dawned on him that he had taken home various different, but related, species of Finch. This
led him to the realization that:
 “Seeing this gradation and diversity of structure in one small, intimately related group of
birds, one might really fancy that from an original paucity of birds in this archipelago, one
species had been taken and modified for different ends.”
 From these observations Darwin’s famous Theory of Evolution was born. Human knowledge
was changed forever, and the Galapagos Finch was renamed the Darwin Finch in his honor.
 Scientists believe that Darwin’s Finches are descended from the Grassquit.
 Over 2 million years ago the small Grassquit made an epic journey of more than 1000Km
across the seas. Perhaps through flight, or more likely a combination of flight and rest on
foliage drifting on the sea currents. Miraculously, the Grassquit somehow found it’s way to
the small volcanic islands of the Galapagos Archipelago, far into the Pacific Ocean.
  Each island has distinct food sources and habitat. On one island the birds can feed easily on
seeds from abundant plants, while on the other island cactuses are the common food source.
 It is here that the story starts to get more interesting. Natural Selection comes into play. On
the first island, a short, stout beak was beneficial for eating seeds from the ground. So birds
with this beak type were more likely to survive, mate, and pass the short beak trait onto their
chicks. On the second island, a longer pointed beak was more useful to pierce cactus fruit.
Likewise this pointy beak trait meant a higher chance of survival, and was passing genetically
onto their chicks.
  Now let’s fast forward over hundreds of thousands of breeding generations. The advantageous
beak traits on each island have been passed down to young birds many thousands of times. So
over a long period of time, the size and shape of bird beaks between the two islands has
become significantly different. The birds on each island are now sufficiently different in
appearance and behavior that they can be recognised as seperate species. Not only that, they
are also now both different from the original pioneering Grasquit. So the same bird species
that arrived to Galapagos has now evolved into two different, unique species.
 As such, Darwin Finches are an exceptional example of the process of evolution of species.
Back in the day, this was a very controversial discovery, that went against the widely
accepted belief of god as the one creator.
 Definitions:
1. Evolution – in terms of living organisms, it may be defined as the development of
differentiated organisms from pre-existing, less differentiated organisms over the
course of time (The basic idea of biological evolution is that species of organisms
change over time.)
2. Natural Selection – Selection is the process by which those organisms which appear
physically, physiologically and behaviourly better adapted to the environment survive
and reproduce; those organisms not so well adapted either fail to reproduce or die.

HOW ENVIRONMENTAL FACTORS ACT AS FORCES OF NATURAL SELECTION

 Individuals may produce more offspring than their environment can support, and some die
because of factors such as predation, food shortage or disease. 
 These factors are known as environmental selection pressures and they determine which
individuals will do best at surviving and reproducing.
 In short, selection pressures therefore affect an organism’s ability to survive in a given
environment
 Types of selection pressures include:
1. Resource availability – Presence of sufficient food, habitat (shelter / territory) and mates
2. Environmental conditions – Temperature, weather conditions or geographical access
3. Biological factors – Predators and pathogens (diseases)

EXAMPLES OF SELECTION PRESSURE IN ACTION

1. Antibiotic resistance in bacteria 
 Antibiotic resistance evolves naturally via natural selection through random
mutation
 The antibiotic action is an environmental pressure; those bacteria which have a
mutation allowing them to survive will live on to reproduce.
 They will then pass this trait to their offspring, which will be a fully resistant
generation.

2. Industrial melanism in the peppered moth, Biston betularia 

 The peppered moth, Biston betularia, is found throughout the British Isles. It flies
at night and during the day settles on trees where it is camouflaged against
lichens that grow on bark in unpolluted parts of the country. 
 There are two very distinct forms within this species. One has fairly light
coloured wings speckled with black that makes it look as if its white wings have
been dusted with black pepper. This form is well camouflaged against the lichens
that grow on trees. 
 The other form is black and is known as the melanic form. As soon as any
melanic moths appeared they were likely to be eaten by predatory birds since
they were so obvious against the speckled background on trees. They were eaten
before they had the chance to reproduce. 
 In the middle of the 18th century, the environment changed significantly. With
the industrial revolution came severe air pollution from the burning of coal.
Smoke contains sulphur dioxide, which killed most of the lichen species, and
soot, which was deposited on the trees. 
 In woodlands around Manchester people noticed more and more of the melanic
peppered moths which were now camouflaged against the trees. 
 The speckled moths were easily spotted by birds and eaten while the melanics
survived and reproduced. The melanics left more offspring than the speckled
variety so that by the beginning of the 20th century the melanic form made up
over 90% of the population in woodlands around industrial cities. 
 The composition of populations in rural areas with no pollution has not changed,
although in eastern England where there is little pollution the melanic variety
increased as pollution was carried by the prevailing wind. 
 Now, with the decline of heavy industry in the UK and the introduction of Clean
Air Acts there is much less pollution and the population has changed with the
speckled form increasing in numbers and the numbers of melanics decreasing. In
this example, bird predators are acting as the agent of natural selection although
their effect was due to pollution in the environment as a consequence of human
activity.
 Note - In this example, bird predators are acting as the agent of natural selection
although their effect was due to pollution in the environment as a consequence of
human activity.

The Dominican anole

 The Dominican anole, Anolis oculatus, is endemic to Dominica. 
 It is a habitat generalist, utilizing a wide variety of habitats from dry seasonal
woodland and scrub to rainforest, and within each habitat it occupies a wide range
of structural niches, frequently being found on the ground as well as more than 5
metres above it on tree trunks.

 If we zoom into the various environments, we’ll discover something interesting.

Each habitat contains a population of the same anole species, but with distinct
characteristics in color, size, and scalation. 
 Much of the observed variation appears to be associated with habitat differences.
thus suggesting selection by geographically varying environmental conditions
as one possible cause.
  The long legs of the trunk-ground anoles enable them to move faster on the ground
than the short-legged twig anoles. 
 The long-legged adaptation helps the trunk-ground anoles not only catch prey on
the ground but also avoid predators that live in their habitats. 
 However, when placed in the habitat of the twig anoles, where twig anoles can
move easily with their short legs, the trunk-ground anoles are clumsy. 
 Also the canopy anoles, with their large toe pads, are adapted to living on leaf
surfaces in the canopy.
 Physical differences among anole species, for example in tails, limb proportions,
and toe-pad size, evolved as a result of natural selection in different environments
 There is therefore a strong correlation between selection and which habitat the
anoles occupied.
 This is an interesting observation, showing natural selection in real time. 

TRINIDADIAN GUPPIES
The Trinidadian guppy (Poecilia reticulata) is an excellent species for studying
the relationship between adaptive evolution across different environmental
circumstances because:
1. It matures rapidly (one generation = 3-4 months)
2. It inhabits different ecological environments that can be easily manipulated
 On Trinidad, guppies live in streams, or portions of streams, that can differ in
the species of predators that the guppies have to contend with. Some streams are
high-predation environments, others  low-predation.
 Guppies from high-predation environments experience much higher mortality
rates than do guppies in low-predation environments. High mortality is
associated with the following characteristics, all of which have a genetic basis:
1. Earlier maturity
2. Greater investment of resources in reproduction
3. More and smaller offspring.
 We have found that mortality rates can be manipulated by:
1. Transplanting guppies from high-predation localities into sites from which they
and their predators had previously been excluded by natural waterfalls, thus
lowering mortality rates. 
2. Introducing predators into low-predation sites, thus increasing mortality rates. 
Such experiments have shown that species evolve as predicted by theory. We
have also found that evolution by natural selection can be remarkably fast.

THE BIOLOGICAL SPECIES CONCEPT AND ITS LIMITATIONS

 The biological species concept defines a species as members of populations that actually or
potentially interbreed in nature, not according to similarity of appearance. 
 The biological species concept has been prevalent in the evolutionary literature for the last several
decades and is emphasized in many biology courses. 
 It is probably the species concept most familiar to biologists 
 Species defined by the biological species concept have also been championed as units of
conservation
 Despite the long historical acceptance of the biological species concept, it has become
controversial because a growing number of evolutionary biologists have found the biological
species concept unworkable in a wide variety of situations. for example:
1. A fundamental drawback to this concept is that it is exclusively defined in terms of sexual
reproduction. Asexual taxa are obviously excluded from this concept, but it is also true
that many species capable of sexual reproduction cannot be easily accommodated within
the framework of the biological species concept.
2. What do you do with organisms that occasionally form hybrids with one another. Around
10% of birds and butterfly species produce hybrids in the wild, although each species
usually does so very rarely (maybe 1/1000 or less). Plants are especially well known for
their tendency to hybridize (probably well over 20%).
3. Specimens collected may be dead and some are known from single specimens
4. Fossil species cannot breed.
5. It is very difficult to observe mating behaviour in the wild, let alone check the offspring
to see if they are fertile

ISOLATING MECHANISMS 
An isolating mechanism is a means of producing and maintaining reproductive isolation
within a population.  

Isolating mechanisms 
The following below are examples of reproductive isolation mechanisms
1. Mechanical isolation – occurs in animals where differences in genetalia prevent
successful copulation and in plants where related species of flowers are pollinated
by  different organisms.
Example: The Bucket Orchid and the Orchid Bee
Male Orchid Bees obtain a wax from the orchid that they use to make a substance to
attract female bees.
The anatomy of the Bucket Orchid allows pollination only by this species of bee.
This partnership is so precise that if either species became scarce or extinct, the
other would follow

2. Ecological isolation – Two closely related species may occupy different habitats within
similar regions. They are ecologically isolated if their habitat preferences lower their
probability of mating.
Example: The Red-legged Frog (Rana draytonii) tends to breed  in large ponds.
The Yellow-legged Frog (Rana boylii) breeds almost exclusively in fast-moving
streams.
Their habitat preferences contribute to their reproductive isolation.

3. Temporal isolation – Two related species occupying the same geographical range
may have different periods of sexual activity or breeding seasons.
Example : Closely related Fruit Flies in Hawaii
Drosophila persimilis breeds in early morning.
Closely related Drosophila pseudoobscura breeds in the afternoon

4. Behavioural isolation – occurs where animals exhibit courtship patterns, mating

only results if the courtship display by one sex results in acceptance by the other sex.
Example: The blue-footed booby (Sula nebouxii), for instance, shares its habitat
with several other species of the genus Sula, but despite all their similarities, it never
 mates with them. In this case, behavioral isolation is facilitated by the elaborate
courtship ritual that is unique to this species. The female blue-footed booby selects a
mating partner only after watching the entire courtship ritual

Geographical isolation – Geographical isolation refers to the physical barriers that

exist that keep two species from mating
Example: For example, the Kaibab squirrel is a subspecies of the Abert’s squirrel
that formed when a small population became isolated on the north rim of the Grand
Canyon. Separate gene pools formed, and genetic changes in one group were not
passed on to the other.

SPECIATION 
 This is the process by which one or more species arise from previously existing species.
 Types of speciation:
1. ALLOPATRIC SPECIATION 
2. SYMPATRIC SPECIATION
 Allopatric speciation occurs when populations of a species are physically separated and are
in two different geographical areas. Sympatric speciation occurs when the two or more
populations are in the same geographical area.
1. Allopatric speciation is characterized by the occurrence, at some stage, of spatial separation.
Geographical barriers such as mountain ranges, seas and rivers or habitat preferences, may
produce a barrier to gene flow because of spatial separation.  In their separate niches, the two
groups go their own evolutionary ways, accumulating different gene mutations, being
subjected to different selective pressures, experiencing different historical events, finally
becoming incapable of interbreeding should they ever come together again. 

Examples: 
I. Grand Canyon Squirrels -When Arizona's Grand Canyon formed, squirrels and other small
mammals that had once been part of a single population could no longer contact and
reproduce with each other across this new geographic barrier. They could no longer
interbreed. The squirrel population underwent allopatric speciation. Today, two separate
squirrel species inhabit the north and south rims of the canyon. On the other hand, birds and
other species that could easily cross this barrier continued to interbreed and were not divided
into separate populations.
II. Galapagos Finches - A major example of allopatric speciation occurred in the Galapagos
finches that Charles Darwin studied. There are about 15 different species of finches on the
Galapagos islands, and they each look different and have specialized beaks for eating
different types of foods, such as insects, seeds, and flowers. All of these finches came from a
common ancestor species that must have emigrated to the different islands. Once populations
were established on the islands, they became isolated from each other, eventually becoming
different species.

2. Sympatric Speciation – does not involve geographical separation of the population at the
time n which genetic isolation occurs. It requires the development of some form of
reproductive isolating mechanism which has arisen by selection within a geographically
confined area. 

Examples
I. Africa's Lake Victoria is home to one of evolution's greatest experiments. In its waters, what
began as a single lineage belonging to the cichlid family of fishes has since given rise to a
 dazzling array of forms. But the Lake Victoria cichlids far surpass Darwin's finches in the
astonishing speed with which they diversified: the more than 500 species that live there and
only there today all evolved within the past 15,000 to 10,000 years—an eyeblink in geologic
terms—compared with the 14 finch species that evolved over several million years.

II. Apple maggot flies - 200 years ago, the ancestors of apple maggot flies laid their eggs only
on hawthorns — but today, these flies lay eggs on hawthorns (which are native to America)
and domestic apples (which were introduced to America by immigrants and bred). Females
generally choose to lay their eggs on the type of fruit they grew up in, and males tend to look
for mates on the type of fruit they grew up in. So hawthorn flies generally end up mating with
other hawthorn flies and apple flies generally end up mating with other apple flies. This
means that gene flow between parts of the population that mate on different types of fruit is
reduced. This host shift from hawthorns to apples may be the first step toward sympatric
speciation — in fewer than 200 years, some genetic differences between these two groups of
flies have evolved.