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