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Taxonomy Five Kingdoms R.H Whittaker 1969 Three Domains Evolutionary Trees Cladistics: Cladograms and molecular data What are the tools used by scientists to observe and understand evolutionary relationships? • • • • • • Artificial Selection 1. Artificial selection 2. Fossil record 3. Comparative anatomy 4. Comparative embryology 5. Comparative biochemistry 6. Biogeography 1 The fossil record: A gallery of fossils What about analogous structures? • Convergent evolution • Co-evolution Molecular Data and the Evolutionary Relationships of Vertebrates Homologous structures: anatomical signs of descent with modification Comparative Embryology: • “Ontogeny recapitulates phylogeny” - the appearance of ancestral structures in the embryos of modern descendants (Haeckel) Biogeography - Wallace’s line 2 Some major episodes in the history of life Evidence from biogeographical studies: Figure 25.8 Taxonomy starts with the latin binomial, and exhibits a hierarchical structure reflecting phylogenetic relationships among taxa or groups of organisms Systematics: connecting phylogeny and taxonomy Latin “binomial” or two-part name (genus and species) • Taxonomy = the science of classifying organisms • Phylogenetics = the study of phylogeny, or the history of evolutionary relationships among species Figure 25.9 The connection between classification and phylogeny: this phylogenetic tree organizes taxa into a hypothesized tree of evolutionary relatedness, with species at the tips of the branches. Perceived similarities can be misleading How are these phylogenetic trees constructed? Traditional Approach: Phenetics Classifications based on perceived overall similarity Jellyfish Starfish Human Which pair is most similar? Which pair is most closely related? 3 Perceived similarities can be misleading Jellyfish Starfish • Live in water • No backbone Human Perceived similarities can be misleading Jellyfish Starfish Human • Live on land • Have backbone Classification based on phylogeny • Phylogeny = patterns of ancestry • Cladistics introduced by Willi Hennig in 1950’s Classification based on phylogeny • Phylogeny = patterns of ancestry • Cladistics introduced by Willi Hennig in 1950’s • Goals of cladistics: – Base classifications on phylogeny – Recognize continuum of evolution – Make taxonomists “show their work”, make explicit hypotheses based on data they present Key principles of cladistics • Since organisms are related by descent, close relationship implies recent common ancestors. • Organisms differ genetically and pass on these differences (and resultant traits) to their ancestors. • Therefore, organisms which are closely related should share new features, acquired by evolution, that are not present in other groups. • These uniquely shared derived characters are the best guide to the evolutionary sequence. The more DERIVED characters two lineages share, the more closely related they are. 4 Derived characters are identified through comparison with an outgroup. • An outgroup is a taxon known (or assumed) to have diverged from a common ancestor less recently than any of the other taxa we wish to classify – Relies on other studies, theory, or intuition Adding an outgroup to our comparison Sponge Jellyfish Starfish Human Derived characters are identified through comparison with an outgroup. • An outgroup is a taxon known (or assumed) to have diverged from a common ancestor less recently than any of the other taxa we wish to classify • Then derived characters are those which are found in some of the taxa under consideration but not the outgroup. Derived characters must be absent from outgroup Sponges, jellyfish, and starfish all live in water and do not have backbones, therefore these are NOT derived characters. Derived characters found in only one taxon set it apart Sponge Jellyfish Starfish Human Backbones and living on land are unique to humans. 5 Derived characters found in only one taxon set it apart Sponge Jellyfish Starfish Human backbone Derived characters found in all taxa do not resolve relationships Sponge Jellyfish Starfish Derived characters found in all taxa do not resolve relationships Human Sponges have only one tissue layer. The other taxa all have multiple tissue layers. Shared, derived characters group related taxa together Sponges and jellyfish lack enclosed body cavities. (ancestral) Starfish and humans have enclosed body cavities. (derived) multiple tissue layers Shared, derived characters group related taxa together Sponges and jellyfish lack enclosed body cavities. (ancestral) Starfish and humans have enclosed body cavities. (derived) Shared derived characters are called SYNAPOMORPHIES. Shared, derived characters group related taxa together Sponge Jellyfish Starfish Human enclosed body cavities multiple tissue layers 6 We have just made a cladogram • Phylogenetic tree showing the order in which different lineages diverged from a common ancestor We have just made a cladogram • Phylogenetic tree showing the order in which different lineages diverged from a common ancestor • Each branch point is labeled with (a) diagnostic synapomorphy(ies) – This is what allows others to “check our work” We have just made a cladogram • Phylogenetic tree showing the order in which different lineages diverged from a common ancestor • Each branch point is labeled with (a) diagnostic synapomorphy(ies) • Each synapomorphy is found on one branch of the tree but not the other branch Synapomorphies mark each branch on a cladogram Sponge Jellyfish Starfish Human enclosed body cavities multiple tissue layers Figure 25.10 Note: Legitimate taxa, such as genera or families, are composed of monophyletic groups of species. These represent evolutionarily unified groups. What are shared, derived, homologous traits? •Monophyletic groups contain a common ancestor and all of its descendants. •Paraphyletic groups are missing one or more descendants, and polyphyletic groupings are missing a common ancestor. • Shared traits are similarities among organisms • Some similarities are homologous, i.e. shared because of inheritance from a common ancestor • Other similarities are analogous, i.e. due to convergent evolution, NOT from inheritance from a common ancestor •How do we construct hypothesized monophyletic groups? Using shared, derived, homologous traits. 7 Algae Fern Pine Cherry Algae Fern Pine Flowers Cherry Flowers Woody Stem Woody Stem Lives on land Lives on land Ancestral trait shared by all 4 taxa Photsoynthesizes Algae Fern Photsoynthesizes Pine Cherry Algae Fern Pine Cherry Loss of “broad leaves” “broad leaves” “broad leaves” Flowers Flowers Woody Stem Woody Stem “broad leaves” Lives on land salamander hawk wings Lives on land squirrel wings bat What it means if a taxa has an ancestral trait • For the character under question, the state of the character matches the character state in the outgroup • E.g., ferns display the ancestral trait for the character state: does it have woody stem • This does not mean that the fern is the ancestor to any of the other taxa 8