Ecology and Evolution

Lecture 13
Macroevolution
 Macroevolution
◼ Macroevolution refers to large-scale
evolutionary changes that occur over
extended periods of time, leading to the
emergence of new species (speciation) or
extinction of species

◼ Unlike microevolution, which involves small,

incremental changes within a species or
population, macroevolution encompasses
significant transformations that can result in
the development of entirely new lineages.

◼ During the gradual separation of species,

two populations at various stages of
becoming a new species will exist

◼ Reproductive isolation is an important

component of the process of speciation.

◼ If individuals within a population mate with

one another but not with individuals of other
populations, they are an independent
evolutionary unit separate branches on the
tree of life.
 The Biological Species Concept
◼ The biological species concept is defined
as a population or group of populations
whose members have the potential to
breed with each other in nature to produce
viable, fertile offspring

◼ they cannot interbreed with other

populations (or if they do, the cannot
produce viable, fertile offspring with
members of other species).

◼ Two different species: look very similar, but

do not interbreed

◼ One species may look very diverse, but can

breed

◼ Species are based on the ability to

interbreed, not physical similarity.
 The Biological Species Concept Limitations

◼ The biological species concept

cannot be applied to

◼ Asexual organisms

◼ Fossils- one cannot test the

reproductive isolation of
morphologically similar fossils,
which are separated into
species based on morphology.

◼ Organisms about which little is

known regarding their
reproduction
 Preservation of the Species:
Reproductive Barriers
◼ The existence of biological factors that
impede members of two species from
producing viable, fertile hybrid

◼ Preserves genetic integrity (gene flow

prevented)

◼ Isolating mechanisms:
a. Prezygotic barriers
◼ Impede mating between species
or hinder the fertilization of ova if
members of different species
attempt to mate

b. Postzygotic barriers
◼ Often prevent the hybrid zygote
from developing into a viable,
fertile adult
 a) Prezygotic Barriers
◼ Prevent fertilization from taking
place (even when ranges
overlap)

◼ Interspecfic zygote never

produced

◼ Five types:
1. Habitat
2. Temporal
3. Behavioral
4. Mechanical
5. Gametic

◼ A single barrier may not block all

genetic exchange between
species, but a combination of
several barriers can effectively
isolate a species’ gene pool
 1) Habitat Isolation
◼ Habitat isolation. Two
organisms that use different
habitats (even in the same
geographic area) are unlikely
to encounter each other to
even attempt mating.

◼ Two species of garter snakes

in the genus Thamnophis
occur in the same areas.

◼ Because one lives mainly in

water and the other is
primarily terrestrial, they rarely
encounter each other.
 2) Temporal Isolation
◼ Temporal isolation. Two
species that breed during
different times of day, different
seasons, or different years
cannot mix gametes.

◼ The geographic ranges of the

western spotted skunk and the
eastern spotted skunk overlap.
However, they do not
interbreed because the former
mates in late summer and the
latter in late winter.
 3) Behavioral Isolation
◼ Behavioral isolation.
Many species use
elaborate courtship
behaviors unique to the
species to attract mates.

◼ In many species, elaborate

courtship displays identify
potential mates of the
correct species and
synchronize gonadal
maturation.
 4) Mechanical Isolation
◼ Closely related species may attempt to mate
but fail because they are anatomically
incompatible (size or incompatible genitalia
prevents mating)

◼ For example, mechanical barriers contribute

to the reproductive isolation of flowering
plants that are pollinated by insects or other
animals.

◼ The snail shells spiral in these two species

are in opposite directions, and therefore the
genital openings are not aligned.
 5) Gametic Isolation
◼ The gametes of two species do not form a zygote
because of incompatibilities preventing fertilization.

◼ In species with internal fertilization, the environment

of the female reproductive tract may not be
conducive to the survival of sperm from other
species.

◼ For species with external fertilization, gamete

recognition may rely on the presence of specific
molecules on the egg’s coat, which adhere only to
specific molecules on sperm cells of the same
species.

◼ A similar molecular recognition mechanism enables

a flower to discriminate between pollen of the same
species and pollen of a different species.

◼ Examples:
◼ sperm cannot penetrate the egg of the different
species, such as between sea urchins species, as
well as in many other animal species

◼ in plants, often pollen grains do not get the proper

signal to germinate when on a stigma of a different
species
b) Postzygotic barriers
◼ Reproductive isolating mechanisms that
prevent gene flow after fertilization has taken
place

1. Hybrid inviability
2. Hybrid sterility
3. Hybrid breakdown
 Postzygotic: Hybrid Inviability or
Reduced hybrid viability
◼ The most common type of
postzygotic barrier

◼ Most F1 do not complete

development and die
during the tadpole stage
and those that do are frail
and die early

◼ This is true for the

occasional hybrids
between frogs in the
genus Rana.
 Postzygotic: Hybrid Sterility
◼ Even if the hybrid offspring are
vigorous, the hybrids may be
infertile, and the hybrid cannot
backbreed with either parental
species.

◼ This infertility may be due to

problems in meiosis because
of differences in chromosome
number or structure.

◼ For example, while a mule, the

hybrid product of mating
between a horse and donkey,
is a robust organism, it cannot
mate (except very rarely) with
either horses or donkeys.
 Postzygotic: Hybrid Breakdown
◼ Hybrid breakdown is a type of reproductive failure
that appears after the F2 generation of crosses
between different species or subspecies.

◼ Some first-generation hybrids are fertile

◼ However, when they mate with another species

or with either parent species, offspring of the next
generation are feeble or sterile

◼ Example 1:
◼ 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 recessive alleles are small and
sterile.

◼ Example 2:
◼ The offspring of hybrid copepods have less
potential for survival or reproduction

◼ Example 3
◼ crosses between sunflower species result in
80% hybrid breakdown in the F 2 generation
 How Species Arise
i. Modes of Speciation:
Possible evolutionary
mechanisms that explain
how millions of species
are on earth today

ii. Rates of Speciation

iii. Macroevolutionary
changes
 i) Mode of Speciation
◼ Speciation can occur in two
ways

1. Allopatric speciation
◼ geographic separation of
populations restricts gene
flow.

2. Sympatric speciation
◼ speciation occurs in
geographically overlapping
populations when
biological factors, such as
◼ chromosomal changes
◼ nonrandom mating
◼ reduce gene flow.
 Allopatric (“Other Country”)
Speciation
◼ Speciation by geographic
isolation

◼ Gene flow is interrupted or

reduced when a population is
divided into two or more
geographically isolated
subpopulations

◼ Mountain ranges, glaciers, land

bridges, or splintering of lakes
may divide one population into
isolated groups.

◼ Once geographic separation has

occurred
◼ One or both populations may
undergo evolutionary change
during the period of separation
 Sympatric (“Same Country”) Speciation

◼ In sympatric speciation
◼ a new species can
originate in the
geographic midst of
the parent species.

◼ Sympatric speciation
does not require large-
scale geographic
distance to reduce
gene flow between
parts of a population.
 Sympatric Speciation in Animals
◼ Reproductive isolation can result
when genetic factors cause
individuals to exploit resources not
used by the parent.

◼ Example is the North American

maggot fly, Rhagoletis pomonella.

◼ Before the arrival of apples from

Europe, it was found mainly in
hawthorns trees

◼ Because apples mature more quickly

than hawthorn fruit, the apple-feeding
flies have been selected for more
rapid development and now show
temporal isolation from the
hawthorn-feeding maggot flies.

◼ Speciation is underway.
 Rates of Speciation
◼ In the fossil record, two different
models have been developed to
explain rates of species evolution
as observed in the fossil record
models:

1. Gradualism: changes in
species is slow and gradual,
occurring in small periodic
changes in the gene pool

2. Punctuated equilibrium: many

species appear as new forms
rather suddenly (in geologic
terms)
• Short time meaning thousands
of years!
 Gradual Refinement of Existing
Structures
◼ The fossil record indicates that light
bones predated flight.

◼ Therefore, they must have had

some function on the ground,
perhaps as a light frame for agile,
bipedal dinosaurs.

◼ Once flight became an advantage,

natural selection would have
remodeled the skeleton to better fit
their additional function.

◼ The wing-like forelimbs and

feathers that increased the surface
area of these forelimbs were co-
opted for flight after functioning in
some other capacity, such as
courtship, thermoregulation, or
camouflage.
 Genes that control development play a
major role in evolution
◼ Heterochrony, an evolutionary change in
the rate or timing of developmental events,
has led to many striking evolutionary
transformations.

◼ Change relative rates of growth even slightly,

and you can change the adult form
substantially.
◼ i.e. fetal skulls human & chimpanzee

◼ Allometric growth: varied rates of growth

for different parts of the body

◼ Ex: the heads of newborn babies are big

compared to their little bodies. As the child
grows, the arms & legs grow faster than the
head.

◼ If this pattern changes, can have big change

in morphology
 Heterochrony
◼ Heterochrony has also played a
part in the evolution of salamander
feet
◼ responsible for differences in
the feet of tree-dwelling versus
ground-dwelling salamanders.

◼ The feet of the tree-dwellers are

adapted for climbing vertically, with
shorter digits and more webbing.

◼ This modification may have evolved

due to mutations in the alleles
that control the timing of foot
development.

◼ Stunted feet may have resulted if

regulatory genes switched off foot
growth early.
 A Phylogenetic Tree
◼ A phylogenetic tree is a
diagram that represents
the evolutionary
relationships among
various species or
groups based on their:
◼ Morphological
◼ Genetic
◼ Behavioral
characteristics
 How To Read a Phylogenetic
Tree?
◼ The root represent the theoretical
common ancestor that gave rise to all
organism on the tree

◼ The tree’s branching structure

represents evolutionary divergences,
with each node indicating a common
ancestor from which descendant
species evolved.

◼ The length of the branches can

sometimes indicate the amount of
evolutionary change or time.

◼ The tips of the branches correspond to

current or extinct species.

◼ A group of taxa that includes a

common ancestor and all of its
descendants is called a monophyletic
group. (aka clade)

◼ A group that excludes one or

more descendants is a paraphyletic
group
 Phylogenetic Tree
Based on Morphological Characters
◼ Phylogenetic trees based on
morphological characters are
constructed by analyzing the
physical traits of different
organisms to determine their
evolutionary relationships.

◼ Scientists compare various

characteristics, such as bone
structures, leaf shapes, or body
patterns, across species to
identify similarities and
differences.

◼ These traits are then used to

infer how species are related,
with more similar traits
suggesting closer evolutionary
relationships.
 Phylogenetic Tree
Based on Molecular Characters
◼ Phylogenetic trees based on
molecular characters are
constructed by analyzing the
DNA, RNA, or protein
sequences of different
organisms.

◼ Scientists compare these

molecular sequences to
identify similarities and
differences, which reflect
evolutionary relationships.

◼ By aligning and comparing

genetic material, researchers
can determine how closely
related species are, based on
the degree of sequence
similarity or divergence.
Are Mammals More Closely
Related to Amphibians or to Birds?