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Chapter 25 Phylogeny and Systematics
[object Object],[object Object],[object Object]
[object Object],[object Object],Figure 25.1
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[object Object],[object Object],Figure 25.2
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The Fossil Record ,[object Object],[object Object],[object Object],Figure 25.3 1   Rivers carry sediment to the  ocean. Sedimentary rock layers  containing fossils form on the  ocean floor. 2   Over time, new strata are  deposited, containing fossils  from each time period. 3   As sea levels change and the seafloor  is pushed upward, sedimentary rocks are  exposed. Erosion reveals strata and fossils. Younger stratum  with more recent  fossils Older stratum  with older fossils
[object Object],[object Object],[object Object],[object Object]
[object Object],[object Object],Figure 25.4a–g (a)  Dinosaur bones being excavated  from sandstone (g)  Tusks of a 23,000-year-old mammoth,  frozen whole in Siberian ice (e)  Boy standing in a 150-million-year-old  dinosaur track in Colorado (d)  Casts of ammonites,  about 375 million  years old (f)   Insects  preserved  whole in  amber (b)  Petrified tree in Arizona, about  190 million years old (c)  Leaf fossil, about 40 million years old
Morphological and Molecular Homologies ,[object Object],[object Object],[object Object],[object Object]
Sorting Homology from Analogy ,[object Object],[object Object]
[object Object],[object Object],Figure 25.5
[object Object],[object Object]
Evaluating Molecular Homologies ,[object Object],[object Object],Figure 25.6 C  C  A  T  C  A  G  A  G  T  C  C  C  C  A  T  C  A  G  A  G  T  C  C  C  C  A  T  C  A  G  A  G  T  C  C  C  C  A  T  C  A  G  A  G  T  C  C  G  T  A Deletion Insertion C  C  A  T  C  A  A  G  T  C  C  C  C  A  T  G  T  A  C  A  G  A  G  T  C  C  C  C  A  T  C  A  A  G  T  C  C  C  C  A  T  G  T  A  C  A  G  A  G  T  C  C  1 Ancestral homologous  DNA segments are  identical as species 1  and species 2 begin to  diverge from their  common ancestor. 2 Deletion and insertion  mutations shift what  had been matching  sequences in the two  species. 3 Homologous regions  (yellow) do not all align  because of these mutations. 4 Homologous regions  realign after a computer  program adds gaps in  sequence 1. 1 2 1 2 1 2 1 2 A  C  G  G  A  T  A  G  T  C  C  A  C  T  A  G  G  C  A  C  T  A T  C  A  C  C  G  A  C  A  G  G  T  C  T  T  T  G  A  C  T  A  G Figure 25.7
[object Object],[object Object],[object Object]
Binomial Nomenclature ,[object Object],[object Object],[object Object]
[object Object],[object Object],[object Object]
Hierarchical Classification ,[object Object],[object Object],Figure 25.8 Panthera pardus Panthera Felidae Carnivora Mammalia Chordata Animalia Eukarya Domain Kingdom Phylum Class Order Family Genus Species
Linking Classification and Phylogeny ,[object Object],[object Object],Figure 25.9 Panthera  pardus (leopard) Mephitis  mephitis  (striped skunk) Lutra lutra  (European  otter) Canis  familiaris  (domestic dog) Canis lupus  (wolf) Panthera Mephitis Lutra Canis Felidae Mustelidae Canidae Carnivora Order Family Genus Species
[object Object],[object Object],Leopard Domestic cat Common ancestor
[object Object],[object Object],Leopard Domestic cat Common ancestor Wolf
[object Object],[object Object],[object Object],[object Object],[object Object],[object Object],[object Object]
Cladistics ,[object Object],[object Object]
[object Object],[object Object],Figure 25.10a (a) Monophyletic.  In this tree, grouping 1,  consisting of the seven species B–H, is a  monophyletic group, or clade. A mono- phyletic group is made up of an  ancestral species (species B in this case)  and  all  of its descendant species. Only  monophyletic groups qualify as  legitimate taxa derived from cladistics. Grouping 1 D C E G F B A J I K H
[object Object],[object Object],Figure 25.10b (b) Paraphyletic.  Grouping 2 does not  meet the cladistic criterion: It is  paraphyletic, which means that it  consists of an ancestor (A in this case)  and  some , but not all, of that ancestor’s  descendants. (Grouping 2 includes the  descendants I, J, and K, but excludes  B–H, which also descended from A.) D C E B G H F J I K A Grouping 2
[object Object],[object Object],Figure 25.10c Grouping 3 (c) Polyphyletic.  Grouping 3 also fails the  cladistic test. It is polyphyletic, which  means that it lacks the common ancestor  of (A) the species in the group. Further- more, a valid taxon that includes the  extant species G, H, J, and K would  necessarily also contain D and E, which  are also descended from A. D C B E G F H A J I K
Shared Primitive and Shared Derived Characteristics ,[object Object],[object Object]
[object Object],[object Object],[object Object]
[object Object],[object Object]
Outgroups ,[object Object],[object Object]
[object Object],[object Object],[object Object],[object Object]
[object Object],[object Object],Figure 25.11a, b Salamander TAXA Turtle Leopard Tuna Lamprey Lancelet (outgroup) 0 0 0 0 0 1 0 0 0 0 1 1 0 0 0 1 1 1 0 0 1 1 1 1 0 1 1 1 1 1 Hair Amniotic (shelled) egg Four walking legs Hinged jaws Vertebral column (backbone) Leopard Hair Amniotic egg Four walking legs Hinged jaws Vertebral column Turtle Salamander Tuna Lamprey Lancelet (outgroup) (a) Character table.  A 0 indicates that a character is absent; a 1  indicates that a character is present. (b) Cladogram.  Analyzing the distribution of these  derived characters can provide insight into vertebrate  phylogeny. CHARACTERS
Phylogenetic Trees and Timing ,[object Object],[object Object]
Phylograms ,[object Object],[object Object],Figure 25.12 Drosophila Lancelet Amphibian Fish Bird Human Rat Mouse
Ultrametric Trees ,[object Object],[object Object],Figure 25.13 Drosophila Lancelet Amphibian Fish Bird Human Rat Mouse Cenozoic Mesozoic Paleozoic Proterozoic 542 251 65.5 Millions of years ago
Maximum Parsimony and Maximum Likelihood ,[object Object],[object Object],[object Object]
[object Object],[object Object]
[object Object],Figure 25.14 Human Mushroom Tulip 40% 40% 0 30% 0 Human Mushroom Tulip (a) Percentage differences between sequences 0
[object Object],Figure 25.14 Tree 1: More likely (b) Comparison of possible trees Tree 2: Less likely 15% 5% 15% 20% 5% 10% 15% 25%
[object Object],[object Object],Figure 25.15a APPLICATION   In considering possible phylogenies for a group of species, systematists compare molecular data for the species. The most efficient way to study the various phylogenetic hypotheses is to begin by first considering the most parsimonious—that is, which hypothesis requires the fewest total evolutionary events (molecular changes) to have occurred. TECHNIQUE   Follow the numbered steps as we apply the principle of parsimony to a hypothetical phylogenetic problem involving four closely related bird species. Species I Species II Species III Species IV I II III IV I III II IV I IV II III Sites in DNA sequence Three possible phylogenetic hypothese 1 2 3 4 5 6 7 A G G G G G T G G G A G G G G A G G A A T G G A G A A G I II III IV I II III IV A G G G G G G Bases at  site 1 for  each species Base-change event 1   First, draw the possible phylogenies for the species  (only 3 of the 15 possible trees relating these four  species are shown here). 2   Tabulate the molecular data for the species (in this simplified  example, the data represent a DNA sequence consisting of  just seven nucleotide bases). 3   Now focus on site 1 in the DNA sequence. A single base- change event, marked by the crossbar in the branch leading  to species I, is sufficient to account for the site 1 data. Species
4 Continuing the comparison of bases at  sites 2, 3, and 4 reveals that each of  these possible trees requires a total of four base-change events (marked again by crossbars). Thus, the first four sites in this DNA sequence do not help us identify the most parsimonious tree. 5   After analyzing sites 5 and 6, we find that the first tree requires fewer evolutionary events than the other two trees (two base changes versus four). Note that in these diagrams, we assume that the common ancestor had GG at sites 5 and 6. But even if we started with an AA ancestor, the first tree still would require only two changes, while four changes would be required to make the other hypotheses work. Keep in mind that parsimony only considers the total number of events, not the particular nature of the events (how likely the particular base changes are to occur). 6   At site 7, the three trees also differ in the number of evolutionary events required to explain the DNA data. RESULTS   To identify the most parsimonious tree, we total all the base-change events noted in steps 3–6 (don’t forget to include the changes for site 1, on the facing page). We conclude that the first tree is the most parsimonious of these three possible phylogenies. (But now we must complete our search by investigating the 12 other possible trees.) Two base changes Figure 25.15b I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III GG GG AA AA GG AA GG GG AA GG AA GG GG GG GG AA GG AA GG GG GG T G T G T T T T T G G T G T T G G T T T T 10 events 9 events 8 events
Phylogenetic Trees as Hypotheses ,[object Object],[object Object]
[object Object],[object Object],Figure 25.16a, b Lizard Four-chambered heart Bird Mammal Lizard Four-chambered heart Bird Mammal Four-chambered heart (a) Mammal-bird clade (b) Lizard-bird clade
[object Object],[object Object],[object Object]
Gene Duplications and Gene Families ,[object Object],[object Object]
[object Object],[object Object],[object Object],Figure 25.17a Ancestral gene Speciation Orthologous genes (a)
[object Object],[object Object],[object Object],Figure 25.17b (b) Ancestral gene Gene duplication Paralogous genes
Genome Evolution ,[object Object],[object Object],[object Object],[object Object]
[object Object]
Molecular Clocks ,[object Object],[object Object]
Neutral Theory ,[object Object],[object Object],[object Object]
Difficulties with Molecular Clocks ,[object Object],[object Object]
Applying a Molecular Clock: The Origin of HIV ,[object Object],[object Object],[object Object],[object Object]
The Universal Tree of Life  ,[object Object],[object Object],[object Object],Figure 25.18 Bacteria Eukarya Archaea 4 Symbiosis of chloroplast ancestor with ancestor of green plants 3 Symbiosis of mitochondrial ancestor with ancestor of eukaryotes 2 Possible fusion of bacterium and archaean, yielding ancestor of eukaryotic cells 1 Last common ancestor of all living things 4 3 2 1 1 2 3 4 0 Billion years ago Origin of life

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Chapter25

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  • 41. 4 Continuing the comparison of bases at sites 2, 3, and 4 reveals that each of these possible trees requires a total of four base-change events (marked again by crossbars). Thus, the first four sites in this DNA sequence do not help us identify the most parsimonious tree. 5 After analyzing sites 5 and 6, we find that the first tree requires fewer evolutionary events than the other two trees (two base changes versus four). Note that in these diagrams, we assume that the common ancestor had GG at sites 5 and 6. But even if we started with an AA ancestor, the first tree still would require only two changes, while four changes would be required to make the other hypotheses work. Keep in mind that parsimony only considers the total number of events, not the particular nature of the events (how likely the particular base changes are to occur). 6 At site 7, the three trees also differ in the number of evolutionary events required to explain the DNA data. RESULTS To identify the most parsimonious tree, we total all the base-change events noted in steps 3–6 (don’t forget to include the changes for site 1, on the facing page). We conclude that the first tree is the most parsimonious of these three possible phylogenies. (But now we must complete our search by investigating the 12 other possible trees.) Two base changes Figure 25.15b I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III I II III IV I III II IV I IV II III GG GG AA AA GG AA GG GG AA GG AA GG GG GG GG AA GG AA GG GG GG T G T G T T T T T G G T G T T G G T T T T 10 events 9 events 8 events
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