CHAPTER 25 ORIGIN OF SPECIES

Prepared by

Brenda Leady, University of Toledo

1 Copyright (c) The McGraw-Hill Companies, Inc. Permission required for reprod

Species
A

group of organisms that maintains a distinctive set of attributes in nature

Macroevolution
 Evolutionary

changes that create new species and groups of species  Occurs by accumulation of microevolutionary changes (changes in a single gene)
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Known number of species about 1.4 million  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
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Amount of separation time for 2 populations
 Short

time – likely to be similar and considered the same species  Long time – more likely to show unequivocal differences

May find situations where some differences are apparent but difficult to decide if the 2 populations are truly different species
 Sometimes

use subspecies classification
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Species concepts
Phylogenetic  Biological  Evolutionary  Ecological

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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

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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

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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

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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
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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
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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
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Prezygotic barriers
 

Habitat isolation
 Geographic  Reproduce

barrier prevents contact

Temporal isolation
year at different times of the day or

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Behavioral isolation
 Behaviors

important in mate choice  Changes in song

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 Mechanical
Size

isolation

or incompatible genitalia prevents mating

 Gametic

isolation

Gametes

fail to unite successfully Important in species that release gametes into the water or air
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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

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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

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Patterns of speciation

Anagenesis
 Single

species transformed into a different species over the course of many generations of a species into 2 or more species

Cladogenesis
 Division

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

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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
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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 rate  Could have played a role in reproductive isolation

Hybridization
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

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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

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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
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Evo-Devo
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

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Developmental genes are key players in evolution
Changes in developmental genes affect traits that can be acted on by natural selection  Compare chicken and duck foot

 Due

to differences in expression of 2 cellsignaling proteins
BMP4 – causes cells to undergo apoptosis and die  Gremlin – inhibits the function of BMP4 and allows cell to survive
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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

feet

Speciation may have been promoted by geographical isolation of habitats

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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

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Three lines of evidence support the idea that Hox gene complexity has been instrumental in the evolution and speciation of animals with different body patterns
 Hox

genes are known to control body development  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
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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

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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 another  Paedomorphosis – retention of juvenile traits in an adult organism

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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

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