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Speciation 
Chapter 13
Evolution 
• Microevolution—Relatively short term 
changes in ALLELE FREQUENCIES within a 
population or species 
• Macroevolution—Large scale evolutionary 
changes 
– Over long periods 
– Small scale microevolution leads to 
macroevolution
Biological Species Concept 
• Species—Group of populations whose 
members have the potential to interbreed in 
nature and produce viable, fertile offspring, 
but do not produce viable offspring with 
members of other groups. 
• What maintains species? 
– Gene flow 
– Reproductive isolation (biological)
Other Species Definitions 
• Morphological Species Concept 
– Characterizes by body shape and other structures 
• Ecological Species Concept 
– Characterizes in terms of ecological niche 
• Phylogenetic Species Concept 
– Smallest group of organisms that share a common 
ancestor
Reproductive Isolation 
• Prezygotic Barriers—block fertilization from 
happening 
– Prevent different species from attempting to mate 
– Prevent an attempted mating from being successful 
– Hindering fertilization 
• Postzygotic Barriers—occur after zygote is 
formed 
– Developmental errors 
– Birth defects cause death 
– Infertile offspring
Prezygotic Reproductive Isolating 
Barriers 
• Behavioral isolation—Absence of cross-attraction between individuals of 
separate species inhibiting any courtship initiation or individual behavior 
during copulation does not allow normal fertilization to occur. 
• Ecological isolation—Variations in the ecology of species give rise to 
barriers: 
– Habitat isolation—Even when living in the same common locality, species 
occupy diverse habitats due to different biological or genetic tendencies 
thereby limiting gene flow during breeding seasons. 
– Temporal isolation—Species living in the same area have different breeding 
seasons preventing gene flow. 
– Pollinator isolation—Flowering plants have varying interactions with 
pollinators thereby reducing gene flow. 
• Mechanical isolation—Reproductive structures are incompatible between 
species inhibiting copulation or pollination. 
• Gametic isolation–Gametes that are transferred during copulation or 
pollination are ineffectual for fertilization due to problems with storage or 
transfer of gametes or because of conspecific pollen or sperm partiality.
Prezygotic Barriers—Habitat Isolation
Prezygotic Barriers—Temporal 
Isolation
Prezygotic Barriers—Behavioral 
Isolation 
• http://www.fiddlercrab.info/video/u_heteropleura02.html 
• http://www.fiddlercrab.info/video/u_ornata01.html
Prezygotic Barriers—Mechanical 
Isolation
Prezygotic Barriers—Gametic Isolation
Postzygotic Reproductive Isolating 
Barriers 
• Ecological unviability—Although normal development 
occurs, hybrids cannot find a suitable ecological niche 
thereby lowering viability. 
• Hybrid unviability—Hybrid species have developmental 
issues causing complete or incomplete lethality. 
• Behavioral sterility—Although normal gametogenesis 
occurs, hybrids are less fertile and typically exhibit 
phenotypes or courtship behaviors that make them less 
desirable mates. 
• Hybrid sterility—Hybrids can have developmental problems 
of the reproductive organs or gametes, or can suffer from 
physiological or neurological issues that prevent effective 
courtship.
Postzygotic Barriers—Reduced Fertility
Speciation 
• Development of a new species through a 
variety of factors 
• Rate of speciation depends on generation 
time, environmental conditions, etc. 
• Can be caused by a change in just 1 gene or a 
set of genes causing some sort of isolation
Types of Speciation 
• Allopatric Speciation—gene flow is interrupted when a 
population is divided into geographically isolated 
subpopulations 
• Parapatric Speciation—occurs when part of a population 
enters a new habitat bordering the range of the parent 
species 
– Some gene flow may occur between populations in border zone 
• Sympatric Speciation—occurs in populations that live in 
the same geographic area 
– Less common than allopatric speciation 
– Happens when gene flow is diminished by: 
• Polyploidy 
• Habitat differentiation 
• Sexual selection
Types of Speciation
Allopatric 
Speciation
Parapatric 
Speciation 
Ephedra californica 
Ephedra trifurca
Sympatric Speciation 
Apple maggot flies (Rhagoletis pomonella) on domestic apple (left) and on much smaller 
native hawthorn fruits (right). Flies that emerge from a given host generally return to 
mate and lay eggs on the same type of fruit.
How Does Speciation Occur? 
• Gradualism—one species gradually 
transforming into another through a series of 
intermediate forms 
– Evolutions occurs in small, incremental changes 
over MANY generations 
– Should be able to see in fossil record but we do 
not – intermediate forms NOT present in fossils 
• Fossil record incomplete or; 
• Missing links too rare in fossils or; 
• See next slide 
Gradualism
How Does Speciation Occur? 
• Punctuated Equilibrium—relatively brief 
bursts of rapid evolution interrupting long 
periods of little change 
– Fits with allopatric speciation 
– Can occur during adaptive radiation—population 
inhabiting a patchy environment gives rise to 
multiple specialized forms in short time period 
• Common in island groups
Punctuated Equilibrium
GULF 
OF 
MEXICO 
Florida 
ATLANTIC OCEAN 
CUBA 
PUERTO 
RICO 
JAMAICAHISPANIOLA 
Adaptive 
Radiation A. loysiana 
Upper trunk/canopy 
A. evermanni 
Midtrunk 
Lower trunk/ground 
A. cristatellus 
Grass/bush 
A. pulchellus
Adaptive 
Radiation
Extinction 
• A species goes extinct when ALL of its 
members die. 
• Many factors can cause—climate, 
reproductive barriers, disease. 
• No matter what it is, it is a FAILURE of that 
species to adapt to the new conditions 
• More prevalent in small, less genetically 
diverse populations
Types of Extinction 
• Background Extinction—gradual loss of species 
over time. 
– Loss of habitat, small climate change, etc. 
• Mass Extinction—many number of species 
disappeared over a relatively short period of 
time. 
– Open up new habitats for adaptive radiation 
– Two theories: 
• Impact Theory—meteor, comet crashes into earth making it 
inhabitable for the life there 
• Movement of Earth’s Crust—causes dramatic environmental 
changes
Taxonomy 
• How we name and classify organisms 
• Binomial Nomenclature 
– First name—Genus 
– Second name—Species 
– Homo sapiens 
• Organisms are classified based on a hierarchy 
of more specific categories
Phylogeny 
• Phylogeny—Depiction of species relationships 
based on descent from shared ancestors 
• Phylogenetic Trees—pictures of these 
relationships 
– Anatomical features of fossils and living creatures 
– Behaviors 
– Physiological adaptations 
– Molecular sequences
Cladistics 
• Phylogenetic system based on ancestral and 
derived characteristics 
– Ancestral Characteristics—inherited attributes and 
RESEMBLE those of ancestor 
– Derived Characteristics—features that are different 
from ancestors group 
– Monophyletic—Group of organisms with 1 common 
ancestor and ALL the descendants 
– Paraphyletic—Group of organisms with 1 common 
ancestor and some, but not all descendants 
– Polyphyletic—Group of organisms that EXCLUDES the 
most recent common ancestor
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 
Clades 
Taxa at branch 
tips 
Birds 
Nonavian 
dinosaurs 
Crocodiles 
Lizards 
and snakes 
Turtles 
Mammals 
Amphibians 
Time 
Node 
(common 
ancestor) 
Last 
common 
ancestorRoot 
of tree 
Cladogram
Cladogram
Cladogram
Constructing Cladograms 
• Light bones 
• 3-toed foot 
• Wishbone 
• Breastbone 
• Loss of 4th 
and 5th digits 
• Down like 
feathers 
• Longer arms 
and hands 
• Complex 
feathers 
• Arms as long or 
longer than legs 
• Feathers 
support flight 
1 
2 
3 
4 
5 
Coelophysis 
SinosauropteryxArchaeopteryx 
Last 
common 
ancestor of 
theropods 
Allosaurus Protarchaeopteryx Modern birds 
Coelophysis 
Allosaurus 
Sinosauropteryx 
Protarchaeopteryx 
Archaeopteryx 
Modern birds 
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 
1 
2 
3 
4 
5
Constructing Cladograms
Cladistics
Phylogenetics and Cladistics Cheat 
Sheet 
Term Definition 
Ancestral characters a trait inherited from the ancestor of a clade 
Clade a group consisting of an ancestor and all its descendants—a single "branch" on the "tree of life” 
Cladistics the science that tries to reconstruct phylogenetic trees and thus discover clades 
Cladogram tree-shaped diagrams; the result of cladistic analyses 
Derived characters a trait that has evolved—not from the common ancestor of the clade 
Monophyletic group a taxon (group of organisms) which forms a clade 
Outgroup a monophyletic group of organisms that serve as a reference group when determining the evolutionary 
relationship among three or more monophyletic groups of organisms 
Paraphyletic group a group that consists of all the descendants of the last common ancestor of the group's members minus a 
small number of monophyletic groups of descendants 
Phylogenetic tree a branching diagram or "tree" showing the inferred evolutionary relationships among various biological 
species or other entities—their phylogeny—based upon similarities and differences in their physical or genetic 
characteristics 
Polyphyletic group a group characterized by one or more homoplasies: character states which have converged or reverted so as 
to appear to be the same but which have not been inherited from common ancestors 
Systemactics the study of the diversification of living forms, both past and present, and the relationships among living 
things through time
The World Around Us

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Speciation

  • 2. Evolution • Microevolution—Relatively short term changes in ALLELE FREQUENCIES within a population or species • Macroevolution—Large scale evolutionary changes – Over long periods – Small scale microevolution leads to macroevolution
  • 3. Biological Species Concept • Species—Group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring, but do not produce viable offspring with members of other groups. • What maintains species? – Gene flow – Reproductive isolation (biological)
  • 4. Other Species Definitions • Morphological Species Concept – Characterizes by body shape and other structures • Ecological Species Concept – Characterizes in terms of ecological niche • Phylogenetic Species Concept – Smallest group of organisms that share a common ancestor
  • 5. Reproductive Isolation • Prezygotic Barriers—block fertilization from happening – Prevent different species from attempting to mate – Prevent an attempted mating from being successful – Hindering fertilization • Postzygotic Barriers—occur after zygote is formed – Developmental errors – Birth defects cause death – Infertile offspring
  • 6. Prezygotic Reproductive Isolating Barriers • Behavioral isolation—Absence of cross-attraction between individuals of separate species inhibiting any courtship initiation or individual behavior during copulation does not allow normal fertilization to occur. • Ecological isolation—Variations in the ecology of species give rise to barriers: – Habitat isolation—Even when living in the same common locality, species occupy diverse habitats due to different biological or genetic tendencies thereby limiting gene flow during breeding seasons. – Temporal isolation—Species living in the same area have different breeding seasons preventing gene flow. – Pollinator isolation—Flowering plants have varying interactions with pollinators thereby reducing gene flow. • Mechanical isolation—Reproductive structures are incompatible between species inhibiting copulation or pollination. • Gametic isolation–Gametes that are transferred during copulation or pollination are ineffectual for fertilization due to problems with storage or transfer of gametes or because of conspecific pollen or sperm partiality.
  • 9. Prezygotic Barriers—Behavioral Isolation • http://www.fiddlercrab.info/video/u_heteropleura02.html • http://www.fiddlercrab.info/video/u_ornata01.html
  • 12. Postzygotic Reproductive Isolating Barriers • Ecological unviability—Although normal development occurs, hybrids cannot find a suitable ecological niche thereby lowering viability. • Hybrid unviability—Hybrid species have developmental issues causing complete or incomplete lethality. • Behavioral sterility—Although normal gametogenesis occurs, hybrids are less fertile and typically exhibit phenotypes or courtship behaviors that make them less desirable mates. • Hybrid sterility—Hybrids can have developmental problems of the reproductive organs or gametes, or can suffer from physiological or neurological issues that prevent effective courtship.
  • 14. Speciation • Development of a new species through a variety of factors • Rate of speciation depends on generation time, environmental conditions, etc. • Can be caused by a change in just 1 gene or a set of genes causing some sort of isolation
  • 15. Types of Speciation • Allopatric Speciation—gene flow is interrupted when a population is divided into geographically isolated subpopulations • Parapatric Speciation—occurs when part of a population enters a new habitat bordering the range of the parent species – Some gene flow may occur between populations in border zone • Sympatric Speciation—occurs in populations that live in the same geographic area – Less common than allopatric speciation – Happens when gene flow is diminished by: • Polyploidy • Habitat differentiation • Sexual selection
  • 18. Parapatric Speciation Ephedra californica Ephedra trifurca
  • 19. Sympatric Speciation Apple maggot flies (Rhagoletis pomonella) on domestic apple (left) and on much smaller native hawthorn fruits (right). Flies that emerge from a given host generally return to mate and lay eggs on the same type of fruit.
  • 20. How Does Speciation Occur? • Gradualism—one species gradually transforming into another through a series of intermediate forms – Evolutions occurs in small, incremental changes over MANY generations – Should be able to see in fossil record but we do not – intermediate forms NOT present in fossils • Fossil record incomplete or; • Missing links too rare in fossils or; • See next slide 
  • 22. How Does Speciation Occur? • Punctuated Equilibrium—relatively brief bursts of rapid evolution interrupting long periods of little change – Fits with allopatric speciation – Can occur during adaptive radiation—population inhabiting a patchy environment gives rise to multiple specialized forms in short time period • Common in island groups
  • 24. GULF OF MEXICO Florida ATLANTIC OCEAN CUBA PUERTO RICO JAMAICAHISPANIOLA Adaptive Radiation A. loysiana Upper trunk/canopy A. evermanni Midtrunk Lower trunk/ground A. cristatellus Grass/bush A. pulchellus
  • 26. Extinction • A species goes extinct when ALL of its members die. • Many factors can cause—climate, reproductive barriers, disease. • No matter what it is, it is a FAILURE of that species to adapt to the new conditions • More prevalent in small, less genetically diverse populations
  • 27. Types of Extinction • Background Extinction—gradual loss of species over time. – Loss of habitat, small climate change, etc. • Mass Extinction—many number of species disappeared over a relatively short period of time. – Open up new habitats for adaptive radiation – Two theories: • Impact Theory—meteor, comet crashes into earth making it inhabitable for the life there • Movement of Earth’s Crust—causes dramatic environmental changes
  • 28. Taxonomy • How we name and classify organisms • Binomial Nomenclature – First name—Genus – Second name—Species – Homo sapiens • Organisms are classified based on a hierarchy of more specific categories
  • 29.
  • 30. Phylogeny • Phylogeny—Depiction of species relationships based on descent from shared ancestors • Phylogenetic Trees—pictures of these relationships – Anatomical features of fossils and living creatures – Behaviors – Physiological adaptations – Molecular sequences
  • 31. Cladistics • Phylogenetic system based on ancestral and derived characteristics – Ancestral Characteristics—inherited attributes and RESEMBLE those of ancestor – Derived Characteristics—features that are different from ancestors group – Monophyletic—Group of organisms with 1 common ancestor and ALL the descendants – Paraphyletic—Group of organisms with 1 common ancestor and some, but not all descendants – Polyphyletic—Group of organisms that EXCLUDES the most recent common ancestor
  • 32. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Clades Taxa at branch tips Birds Nonavian dinosaurs Crocodiles Lizards and snakes Turtles Mammals Amphibians Time Node (common ancestor) Last common ancestorRoot of tree Cladogram
  • 35. Constructing Cladograms • Light bones • 3-toed foot • Wishbone • Breastbone • Loss of 4th and 5th digits • Down like feathers • Longer arms and hands • Complex feathers • Arms as long or longer than legs • Feathers support flight 1 2 3 4 5 Coelophysis SinosauropteryxArchaeopteryx Last common ancestor of theropods Allosaurus Protarchaeopteryx Modern birds Coelophysis Allosaurus Sinosauropteryx Protarchaeopteryx Archaeopteryx Modern birds Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 1 2 3 4 5
  • 38. Phylogenetics and Cladistics Cheat Sheet Term Definition Ancestral characters a trait inherited from the ancestor of a clade Clade a group consisting of an ancestor and all its descendants—a single "branch" on the "tree of life” Cladistics the science that tries to reconstruct phylogenetic trees and thus discover clades Cladogram tree-shaped diagrams; the result of cladistic analyses Derived characters a trait that has evolved—not from the common ancestor of the clade Monophyletic group a taxon (group of organisms) which forms a clade Outgroup a monophyletic group of organisms that serve as a reference group when determining the evolutionary relationship among three or more monophyletic groups of organisms Paraphyletic group a group that consists of all the descendants of the last common ancestor of the group's members minus a small number of monophyletic groups of descendants Phylogenetic tree a branching diagram or "tree" showing the inferred evolutionary relationships among various biological species or other entities—their phylogeny—based upon similarities and differences in their physical or genetic characteristics Polyphyletic group a group characterized by one or more homoplasies: character states which have converged or reverted so as to appear to be the same but which have not been inherited from common ancestors Systemactics the study of the diversification of living forms, both past and present, and the relationships among living things through time

Editor's Notes

  1. “Prezygotic Reproductive Isolating Barriers” https://en.wikipedia.org/wiki/Isolation_by_distance#Prezygotic_Reproductive_Isolating_Barriers ++
  2. Left: “Western Terrestrial Garter Snake Juvie” by James Bettaso. (Public Domain). https://commons.wikimedia.org/wiki/File:Western_terrestrial_garter_snake_juvie.jpg Right: “Aquatic Garter Snake” by SarahStierch. Licensed under a CC-BY-SA 3.0 Unported license. https://commons.wikimedia.org/wiki/File:Aquatic_Garter_Snake_-_Stierch_B.jpg
  3. Left: “Spilogale Putorius” by the National Park Service. (Public Domain). http://commons.wikimedia.org/wiki/File:Spilogale_putorius_(2).jpg Right: “Spilogale Gracilis” by Brian Kentosh. Released into the public domain by the copyright holder. http://commons.wikimedia.org/wiki/File:Spilogale_gracilis_amphiala.jpg
  4. “Fiddler Crab” by NOAA. (Public Domain). http://en.wikipedia.org/wiki/File:Fiddler_crab.jpg
  5. Image from Boundless
  6. “Mushroom coral” by Brocken Inaglory. Licensed under a CC-BY-SA 3.0 Unported license. https://en.wikipedia.org/wiki/File:Muchroom_coral.JPG
  7. “Postzygotic Reproductive Isolating Barriers” https://en.wikipedia.org/wiki/Isolation_by_distance#Postzygotic_Reproductive_Isolating_Barriers
  8. Top left: “AAV Stute” by Seviwurst. Licensed under a CC-BY-SA 2.0 Germany license. http://commons.wikimedia.org/wiki/File:AAV-Stute-2.jpg Top right: “Donkey 1 ARP” by Adrian Pingstone. Released into public domain by the copyright holder. http://commons.wikimedia.org/wiki/File:Donkey_1_arp_750px.jpg Bottom: “Maultier Grau” by Summi. Licensed under a CC-BY-SA 3.0 Unported license. http://commons.wikimedia.org/wiki/File:Maultier_grau.jpg
  9. “Speciation modes” by Ilmari Karonen. Licensed under a CC-BY-SA 3.0 Unported license. https://en.wikipedia.org/wiki/File:Speciation_modes.svg
  10. Image from Boundless
  11. Top: “Ephedra californica” by Stan Shebs. Licensed under a CC-BY-SA 3.0 Unported license. http://en.wikipedia.org/wiki/File:Ephedra_californica_1.jpg Map: “US Locator Blank” by Papayoung. Licensed under a CC-BY-SA 3.0 Unported license. https://commons.wikimedia.org/wiki/File:US_Locator_Blank.svg Bottom: “Ephedra trifurca” by Dcotetzee. Licensed under a CC-0 1.0 license. http://commons.wikimedia.org/wiki/File:Ephedra_trifurca.jpg
  12. Image and caption from Boundless
  13. Image from Boundless
  14. Image from Boundless
  15. REPLACE IMAGE
  16. Image from Boundless
  17. Images from Boundless
  18. REPLACE IMAGE
  19. “Morphology Molecular Genetics” by Wang Y, Zhao J, Yang J, Zhou Z, Chen G, Liu Y. Licensed under a CC-BY 4.0 International license. https://commons.wikimedia.org/wiki/File:Morphology-Molecular-Genetics-and-Bioacoustics-Support-Two-New-Sympatric-Xenophrys-Toads-(Amphibia-pone.0093075.g002.jpg
  20. “Artiodactylamorpha” by C. Buell and L. Betti-Nash. Licensed under a CC-BY 2.5 Generic license. https://commons.wikimedia.org/wiki/File:Artiodactylamorpha.png
  21. REPLACE IMAGE
  22. Image from Boundless
  23. “Monophyly, Paraphyly, Poplyphyly” by Petter Bockman. Licensed under a CC-BY-SA 3.0 Unported license.
  24. “Clade” http://en.wikipedia.org/wiki/Clade “Monophyly” http://en.wikipedia.org/wiki/Monophyly “Outgroup” http://en.wikipedia.org/wiki/Outgroup_(cladistics) “Paraphyly” http://en.wikipedia.org/wiki/Paraphyly “Phylogenetic tree” http://en.wikipedia.org/wiki/Phylogenetic_tree “Polyphyly” http://en.wikipedia.org/wiki/Polyphyly “Systematics” http://en.wikipedia.org/wiki/Systematics
  25. “Phylogenentic Tree of Life” by NASA. (Public Domain). https://commons.wikimedia.org/wiki/File:Phylogenetic_tree.svg