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Prepared by
Brenda Leady, University of Toledo

1 reprod
Copyright (c) The McGraw-Hill Companies, Inc. Permission required for
 Species
A group of organisms that maintains a
distinctive set of attributes in nature
 Macroevolution
 Evolutionarychanges that create new species
and groups of species
 Occurs by accumulation of microevolutionary
changes (changes in a single gene)

 Known number of species about 1.4
 Estimates of unidentified species range
from 2 to 100 million
 Difficulty in identifying a “species”
A single species may exist in 2 distinct
populations that are in the slow process of
evolving into 2 or more different species

 Amount of separation time for 2
 Short time – likely to be similar and
considered the same species
 Long time – more likely to show unequivocal
 May find situations where some
differences are apparent but difficult to
decide if the 2 populations are truly
different species
 Sometimes use subspecies classification

Species concepts
 Phylogenetic
 Biological
 Evolutionary
 Ecological

Phylogenetic species concept
 Species are identified by having a unique
combination of traits
 Historically used physical traits
 Now can use DNA sequences
 Advantage
 Can be applied to all organisms
 Drawbacks
 How many traits to consider, traits that vary in
a continuous way, choose degree of
dissimilarity to use, and members of the same
species can look very different while members
of a different species can look very similar 6
Biological species concept
 A species is a group of individuals whose
members have the potential to interbreed
with one another in nature to produce
viable, fertile offspring but cannot
successfully interbreed with members of
other species
 Reproductive isolation prevents breeding
with other species

 3 problems
 May be difficult to determine if 2 populations
reproductively isolated
 There are cases where 2 species can
interbreed but do not
 Cannot be applied to asexual species

Evolutionary species concept
 A species is derived from a single lineage
that is distinct from other lineages and has
its own evolutionary tendencies and
historical fate
 Lineage
 Genetic relationship between an individual or
group of individuals and its ancestors
 Drawback – no easy way to identify a
unique species because lineages difficult
to examine and quantitate
Ecological species concept
 Each species occupies an ecological niche
 Unique set of habitat resources that a species
requires, as well as its influence on the environment
and other species
 Within their own niche, members of a given
species compete with each other for survival
 If two organisms are very similar, their needs will
overlap, which results in competition
 Such competing individuals are likely to be of the
same species
 Useful for bacterial species
Reproductive isolating mechanisms

 Consequence of genetic changes as

species adapts to its environment
 Prezygotic barriers
 Prevent formation of zygote
 Postzygotic barriers
 Block development of viable, fertile individuals
 Interspecies hybrid- when 2 species do
produce offspring
Prezygotic barriers
 Habitat isolation
 Geographic barrier prevents contact
 Temporal isolation
 Reproduce at different times of the day or

 Behavioral
 Behaviors
important in mate
 Changes in song

 Mechanical isolation
Size or incompatible genitalia prevents
 Gametic isolation
Gametes fail to unite successfully
Important in species that release
gametes into the water or air

Postzygotic barriers
 Less common in nature because they are
more costly in terms of energy and
resources used
 Hybrid inviability – fertilized egg cannot
progress past an early embryo
 Hybrid sterility – interspecies hybrid viable
but sterile
 Mule example
 Hybrid breakdown – hybrids viable and
fertile but subsequent generations have
genetic abnormalities
Mechanisms in speciation
 Underlying cause of speciation is the
accumulation of genetic changes that
ultimately promote enough differences so
that we judge a population to constitute a
unique species

Patterns of speciation
 Anagenesis
 Single
species transformed into a different
species over the course of many generations
 Cladogenesis
 Division of a species into 2 or more species

Allopatric speciation
 Most prevalent method for cladogenesis
 Occurs when some members of a species
become geographically separated

 Can also occur when small population
moves to a new location that is
geographically separated founder effect
 Genetic drift and natural selection may quickly
lead to differences
 Adaptive radiation – single species evolves
into array of descendents that differ greatly in
habitat, form or behavior

Podos Found That an Adaptation to Feeding Also
May Have Promoted Reproductive Isolation
in Finches
 Darwin’s finches have different beak sizes
and shapes as adaptations to different
feeding strategies
 Podos analyzed songs to see if beak
morphology birds with larger beaks had
more narrow frequency range and/or trill
 Could have played a role in reproductive
 Prior to complete reproductive isolation,
the zones where two populations can
interbreed are known as hybrid zones
 As the two populations accumulate
different genetic changes, this may
decrease the ability of individuals from
different populations to mate with each
other in the hybrid zone
 Once gene flow through the hybrid zone is
greatly diminished, the two populations
are reproductively isolated 28
Sympatric speciation
 Occurs when members of a species that
initially occupy the same habitat within the
same range diverge into two or more
different species
 Tends to involve abrupt genetic changes
that quickly lead to the reproductive
isolation of a group of individuals
 Changes in chromosome number

Pace of speciation
 Gradualism
 each new species evolves continuously over long
spans of time
 large phenotypic differences that produce new
species are due to the accumulation of many small
genetic changes
 Punctuated equilibrium
 Tempo more sporadic
 Species in equilibrium for long periods and then short
rapid bursts of changes

 Evolutionary developmental biology
 Compares the development of different
organisms in an attempt to understand
ancestral relationships between organisms
and the developmental mechanisms that
bring about evolutionary change
 Involves the discovery of genes that
control development

Developmental genes are key
players in evolution
 Changes in developmental genes affect
traits that can be acted on by natural
 Compare chicken and duck foot
 Dueto differences in expression of 2 cell-
signaling proteins
 BMP4 – causes cells to undergo apoptosis and die
 Gremlin – inhibits the function of BMP4 and allows

cell to survive
 Mutations on the expression of BMP4 and
gremlin provided variation
 In terrestrial settings, nonwebbed feet are
an advantage
 Natural selection maintains nonwebbed feet
 In aquatic environments, webbed feet are
an advantage
 Natural selection would have favored webbed
 Speciation may have been promoted by
geographical isolation of habitats
Hox genes
 Found in all animals
 Genetic variation may have been critical
event in the formation of new body plans
 Number and arrangement of Hox genes
varies among different types of animals
 Increases in the number of Hox genes
may have led to greater complexity in
body structure
 Three lines of evidence support the idea
that Hox gene complexity has been
instrumental in the evolution and
speciation of animals with different body
 Hox genes are known to control body
 General trend for simpler animals to have
fewer Hox genes and Hox gene clusters
 Comparison of Hox gene evolution and
animal evolution bear striking parallels

Developmental genes that affect
growth rate
 Allometric growth – different parts of the
body grow at different rates with respect to
each other
 Compare growth of head between human
and chimpanzee

 Changes in growth rates can also affect
the developmental stage at which one
species reproduces compared to that of
another species
 Reproduction in the adult is observed at
an earlier stage in one species than
 Paedomorphosis – retention of juvenile
traits in an adult organism

The Study of the Pax6 Gene Indicates That Different
Types of Eyes Evolved from a Simpler Form

 Explaining how a complex organ comes into

existence is a major challenge
 Researchers have discovered many different
types of eyes
 Thought that eyes may have independently
arisen many different times during evolution
 Pax6 is a master control gene that controls
the expression of many other genes and
influences eye development
 Eyes of Drosophila and mammals are
evolutionarily derived from a modification of
an eye that arose once during evolution
 If Drosophila and mammalian eyes had
arisen independently, the Pax6 gene from
mice would not be expected to induce the
formation of eyes in Drosophila
 Hypothesized that the eyes from many
different species all evolved from a common
ancestral form consisting of, as proposed by
Darwin, one photoreceptor cell and one
pigment cell