Systematics and phylogeny Organizing life • All organisms: – Are composed of one or more cells – Carry out metabolism – Transfer energy with ATP – Encode hereditary information in DNA • Tremendous diversity of life – Bacteria-----whales----sequoia trees • Biologists group organisms based on shared characteristics Taxonomy • Field of biology concerned with identifying and naming • Binomial system devised by Linnaeus • Classification is how species and higher groups are placed into the taxonomic hierarchy Systematics • Since fossil records are not complete, scientists rely on other types of evidence to establish the best hypothesis of evolutionary relationships • Systematics is the study of evolutionary relationships • Phylogeny is a hypothesis about the evolutionary relationships among groups Tree of life • Darwin envisioned that all species were descended from a single common ancestor • He depicted this history of life as a branching tree – Now called a cladogram Cladogram • Twigs of a tree represent existing species • Joining of twigs and branches reflects the pattern of common ancestry back in time to a single common ancestor Phylogenies depict evolutionary relationships Early trees • Similarity may not accurately predict evolutionary relationships – Early systematists relied on the expectation that the greater the time since two species diverged from a common ancestor, more different would be Evolution can happen quickly Evolution is not unidirectional Evolution is not always divergent Evolutionary reversal can occur Identifying inherited similarity • Derived characteristic is similarity that is inherited from the most recent common ancestor of an entire group • Ancestral characteristic is similarity that arose prior to the common ancestor of the group Modern systematics - cladistics • Only shared derived characters are considered informative about evolutionary relationships • To use the cladistic method, character variation must be identified as ancestral or derived Using characters • Characters can be any aspect of the phenotype – Morphology – Physiology – DNA – Behavior • Characters should exist in recognizable character states Ancestral vs. derived characters • Presence of hair is a shared derived feature of mammals • Presence of lungs in mammals is an ancestral feature; also present in amphibians and reptiles Determination of ancestral versus derived • First step in a manual cladistic analysis is to polarize the characters (are they ancestral or derived) – Outgroup comparison is used to assign character polarity Cladistics • When the group under study exhibits multiple character states, and one of those states is exhibited by the outgroup, then that state is ancestral and other states are derived • Most reliable if character state is exhibited by several different outgroups Teeth absence polarization • Presence of teeth in mammals and reptiles is ancestral (also found in outgroup of fish) • Absence of teeth in birds and turtles is derived Constructing a cladogram • Clade is a group of species that share a common ancestor as indicated by the possession of shared derived characters • Clades are evolutionary units and refer to a common ancestor and all descendants • Synapomorphy is a derived character shared by clade members Cladogram • A simple cladogram is a nested set of clades • Plesiomorphies are ancestral states • Symplesiomorphies are shared ancestral states Classifying vertebrates Complications in cladistics • Homoplasy is a shared character state that has not been inherited from a common ancestor – Results from convergent evolution and evolutionary reversal • If there are conflicts among characters, use the principle of parsimony which favors the hypothesis that requires the fewest assumptions Parsimony and homoplasy Building a cladogram with DNA Limitations of use of parsimony • Some characters evolve rapidly and principle of parsimony may be misleading • Rate of DNA evolution can be high – Mutations in non-functional sequences are not affected by natural selection, but are affected by genetic drift Statistical approaches • Neighbor-joining method • Maximum likelihood method • Bayesian method Molecular clock • Branches in a cladogram can be dated using the fact that the rate of a molecule is constant over time in a given group Systematics and classification • A monophyletic group includes the most recent common ancestor of the group and all of its descendants (clade) • A paraphyletic group includes the most recent common ancestor of the group, but not all its descendants • A polyphyletic group does not include the most recent common ancestor of all members of the group Mismatches • Taxonomic hierarchies are based on shared traits, should reflect evolutionary relationships • Difficulties changing current perception – i.e., dinosaurs as birds rather than reptiles Monophyletic group Paraphyletic group Polyphyletic group • Warm-blooded Old classification system New classification system Phylogenetic species concept (PSC) • Focuses on shared derived characters • Species should be applied to groups of populations that have been evolving independently of other groups Strengths of PSC • PSC can be applied to allopatric populations • PSC can be applied to both sexual and asexual species Weakness of PSC Comparative biology • Phylogenetics is the basis of all comparative biology • Homologous structures are derived from the same ancestral source • Homoplastic structures are not Homologous behavior Homoplastic convergence • Saber teeth – Occurred in different groups of extinct carnivores – Similar body proportions – Similar predatory lifestyle – Most likely evolved independently at least 3 times Distribution of saber-toothed mammals Homoplastic convergence • Plant conducting tubes – Sieve tubes facilitate long-distance transport of food that is essential for the survival of tall plants – Brown algae also have sieve elements – Closest ancestor a single-celled organism Convergent evolution of conducting tubes Evolution of complex characters occur in steps • Birds adapted for flight – wings, feathers, light bones, breastbone • Initial stages of a character may evolve as an adaptation to some environmental selective pressure different from current use – First feather-like structure evolved in theropod phylogeny for insulation or perhaps decoration Testing hypotheses with phylogenies • Larval dispersal in marine snails – Some snails produce microscopic larvae that drift in the ocean currents – Some species have larvae that settle to the ocean bottom and do not disperse – Fossils show increase in nondispersing snails Increase through time in proportion of species whose larvae do not disperse Competing hypotheses • Evolutionary change from dispersing to nondispersing occurs more often than change in the opposite direction • Species that are nondispersing speciate more frequently, or become extinct less frequently than dispersing species • The two processes would result in different phylogenetic patterns Hypothetical trees Which hypothesis is more likely? Loss of larval stage in marine invertebrates – non-reversible? Alternative hypothesis Species diversification • Use phylogenetic analysis to suggest and test hypotheses about why some groups have greater species richness than others Evolutionary diversification of the Phytophaga Evolution of disease • HIV evolved from a simian viral counterpart SIV – Current estimate: >39 million people infected; > 3 million die each year – SIV found in 36 species of primates Evolution of HIV • HIV descended from SIV • Independent transfers of different strains from different primate species Disease transmission • Evidence use to convict dentist for infecting patient