Phylogeny and Systematics Chapter 25 Phylogeny and Systematics

History of Life (See Table 26.1) 4500 Eras Boundaries between units in the Geologic Time Scale are marked by dramatic biotic change A figure from Ch. 26… 4500 Origin of Earth

5 Kingdom classification system in use through the late 1900s

5 Kingdom classification system in use through the late 1900s gave way to Woese’s 3 Domains

5 Kingdom classification system in use through the late 1900s gave way to Woese’s 3 Domains and multiple Kingdoms

“Did King Philip Come Over For Gumbo Sunday?” Fig. 25.8 “Did King Philip Come Over For Gumbo Sunday?” Taxon (taxa) = the named taxonomic unit(s) at any level in this taxonomic hierarchy Each taxonomic level is more inclusive or comprehensive than the previous one (moving from species toward domains). Panthera = genus pardus = specific epithet that refers to one species in the genus Panthera

Linnaeus convinced us to use a hierarchical classification system Darwin provided us with the mechanism by which evolution results in descent with modification Taxonomy – naming & classifying organisms Systematics – naming & classifying organisms according to their evolutionary relationships These two contributions (from Linnaeus and Darwin) combine in the modern branches of biology known as taxonomy, systematics, phylogenetics, and systematic phylogenetics. Systematic Phylogenetics Phylogenetics – reconstructing the evolutionary relationships among organisms

Macroevolution & Phylogeny Phylogenetic tree – hypothesized genealogy traced back to the last common ancestor (i.e., the most recent) through hierarchical, dichotomous branching Note that this is a highly pruned tree! Cladistics – the principles that guide the production of phylogenetic trees, a.k.a., cladograms

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Node – branch point, speciation event Note that this is a highly pruned tree!

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! Lineage or clade – an entire branch

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! Lineage or clade – an entire branch

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! Lineage or clade – an entire branch

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! A clade is a monophyletic group, i.e., an ancestral species and all of its descendents

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! A clade is a monophyletic group, i.e., an ancestral species and all of its descendents

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! A clade is a monophyletic group, i.e., an ancestral species and all of its descendents

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! A paraphyletic group consists of an ancestor and some of its descendents

Macroevolution & Phylogeny Phylogenetic tree, phylogeny, or cladogram Note that this is a highly pruned tree! A polyphyletic group lacks the common ancestor of species in the group

Macroevolution & Phylogeny Note that this is a highly pruned tree! Fig. 25.9 Taxonomic groups often reflect true clades…

Macroevolution & Phylogeny However, tension sometimes exists between taxonomic tradition and cladistic hypotheses… Note that phylogenetic trees can be oriented any which way.

Macroevolution & Phylogeny E.g., If the Class Reptilia is to be monophyletic, birds must be included!

Macroevolution & Phylogeny How are phylogenetic trees constructed?

Macroevolution & Phylogeny The fossil record is especially valuable, and the only option for many extinct taxa

Macroevolution & Phylogeny However, we almost never have a continuous record from one species to the next

Macroevolution & Phylogeny Cladistic principles allow us to construct hypothesized phylogenetic trees

Cladistic Analysis Fossils provide morphological data for extinct species, whereas comparisons of multiple types of traits – including molecular – do so for extant species

Cladistic Analysis Similar characters (e.g., morphological, behavioral, molecular, etc. traits or features) suggest relatedness… Wasps [Hymenoptera]

Cladistic Analysis But, not all similarity derives from common ancestry! Mantisfly [Neuroptera] Convergent evolution can produce superficially similar traits that lack homology with one another

Cladistic Analysis Homologous characters share common ancestry Lack of similarity among taxa results from divergence

Cladistic Analysis Analogous characters do not share common ancestry Similarity among taxa results from convergence

Cladistic Analysis As a general rule, the more homologous characters shared by two species, the more closely they are related Sequences of DNA & RNA (nucleotides) and proteins (amino acids) are used as characters; as a general rule, the more recently two species shared a common ancestor, the more similar their sequences

Each nucleotide can be treated as a character Cladistic Analysis Each nucleotide can be treated as a character Character changes (mutations) from the ancestral to the derived state include: Substitutions …AGCTCTAGG… …AGCTATAGG… Insertions A = adenine; C = cytosine; G = guanine; T = thymine …AGCTCTAGG… Mutations …AGCTGATCTAGG… Deletions …AGCTCTAGG… …AGCTCTAGG…

All similar characters Cladistic Analysis Shared Primitive Characters (ancestral) Analogies All similar characters Homologies Shared Derived Characters(unique to a clade) This flowchart summarizes types of similarities among species… The sequence of branching in a cladogram then represents the sequence in which evolutionary novelties (shared derived characters) evolved

Ingroup = the group whose relationships we are trying to resolve Cladistic Analysis Ingroup vs. Outgroup Ingroup = the group whose relationships we are trying to resolve Fig. 25.11

Cladistic Analysis Ingroup vs. Outgroup Outgroup = a species (or group) known to have an older most recent common ancestor with the ingroup than the ingroup’s most recent common ancestor Fig. 25.11

Cladistic Analysis Ingroup vs. Outgroup An outgroup helps identify shared ancestral and shared derived characters (unique to a clade) Fig. 25.11

Parsimony & Occam’s Razor Cladistic Analysis Parsimony & Occam’s Razor The most parsimonious tree is the one that requires the fewest evolutionary events (appearance of shared derived characters) Fig. 25.11

Parsimony & Occam’s Razor Cladistic Analysis Parsimony & Occam’s Razor On this most parsimonious cladogram, each key character originated (evolved) once Fig. 25.11

Cladistic Analysis An example… See Fig. 25.15 for another example Outgroup Ingroup See Fig. 25.15 for another example

Cladistic Analysis An example… See Fig. 25.15 for another example Outgroup Ingroup Create potential topologies for the tree See Fig. 25.15 for another example

Cladistic Analysis An example… See Fig. 25.15 for another example Outgroup Ingroup Create potential topologies for the tree The first topology requires 7 changes, the second 9, and the third 8. Map the characters onto the trees Choose the most parsimonious tree See Fig. 25.15 for another example

Macroevolution & Phylogeny Sometimes the most parsimonious arrangement for one character is not the most accurate overall… Bird-mammal clade Lizard-bird clade Fig. 25.16

Macroevolution & Phylogeny Don’t confuse the age of a clade with the age of its component species African Rift Lake Cichlid Alligator Snapping Turtle ~ 200,000 yr ~ 20,000,000 yr Based on the cladogram, which species is likely to be older? Once a clade arises, speciation events can continue to occur within it.

Macroevolution & Phylogeny The overall trend is increasing diversity, with periodic episodes of extinction Fig. 26.8

Macroevolution & Phylogeny Cretaceous mass extinction Asteroid impacts may have caused mass extinction events Cretaceous mass extinction marks boundary between Cretaceous and Tertiary Periods. Permian mass extinction Extinction of >90% of species

Macroevolution & Phylogeny K-T impact event

Macroevolution & Phylogeny Continental drift is responsible for many cladogenic events & biogeograhic distribution patterns E.g., Proteaceae – a plant family that originated in Gondwana

Continental drift results from plate tectonics Fig. 26.19

Macroevolution & Phylogeny Some biogeographic similarities result from common ancestry (common descent) E.g., all bromeliads are found in the New World Tropics & Sub-Tropics

Macroevolution & Phylogeny Some biogeographic similarities result from convergent evolution (convergent adaptive modification) E.g., cactus of Americas compared to euphorbs of Africa Cactus Euphorb