Systematics and the Phylogenetic Revolution Chapter 23

Introduction 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

Systematics Since fossil records are not complete, scientists rely on other types of evidence to establish the best hypothesis of evolutionary relationships Systematics: the study of evolutionary relationships Phylogeny: a hypothesis about patterns of relationship among species

Systematics 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

Systematics 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 Darwin called this process “descent with modification”

Phylogenies depict evolutionary relationships Systematics Phylogenies depict evolutionary relationships

Systematics Key to interpreting a phylogeny: look at how recently species share a common ancestor 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

Systematics Evolution can occur rapidly at one time and slowly at another (punctuated and gradual evolution)

Systematics Oscillating selection: Traits can evolve in one direction, then back the other way Evolution is not always divergent: convergent evolution Use similar habitats Similar environmental pressures Evolutionary reversal: process in which a species re-evolves the characteristics of an ancestral species

Cladistics Derived characteristic: similarity that is inherited from the most recent common ancestor of an entire group Ancestral: similarity that arose prior to the common ancestor of the group In 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

Cladistics Characters can be any aspect of the phenotype Morphology - Physiology Behavior - DNA Characters should exist in recognizable character states Example: Teeth in amniote vertebrates has two states, present in most mammals and reptiles and absence in birds and turtles

Cladistics Examples of ancestral versus 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

Cladistics Determination of ancestral versus derived First step in a manual cladistic analysis is to polarize the characters (are they ancestral or derived) Example: polarize “teeth” means to determine presence or absence in the most recent common ancestor

Cladistics Outgroup comparison is used to assign character polarity A species or group of species not a member of the group under study is designated as the outgroup Outgroup species do not always exhibit the ancestral condition

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

Cladistics Following the character state-outgroup method Presence of teeth in mammals and reptiles is ancestral Absence of teeth in birds and turtles is derived

Cladistics Construction of a cladogram Polarize characteristics Clade: 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: a derived character shared by clade members

Cladistics A simple cladogram is a nested set of clades Plesiomorphies: ancestral states Symplesiomorphies: shared ancestral states

Cladistics

Cladistics Homoplasy: a shared character state that has not been inherited from a common ancestor Results from convergent evolution Results from 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 Cladistics Parsimony and Homoplasy

Cladistics A Cladogram; DNA

Cladistics A Cladogram: DNA

Other Phylogenetic Methods Some characters evolve rapidly and principle of parsimony may be misleading Rate at which some parts of the DNA genome evolve Mutations in repetition sequences, not deleted by natural selection Statistical approaches Molecular clock: rate of evolution of a molecule is constant through time

Systematics and Classification Classification: how we place species and higher groups into the taxonomic hierarchy Genus, family, class.. Monophyletic group: includes the most recent common ancestor of the group and all of its descendants (clade) Paraphyletic group: includes the most recent common ancestor of the group, but not all its descendants

Systematics and Classification Polyphyletic group: does not include the most recent common ancestor of all members of the group Taxonomic hierarchies are based on shared traits, should reflect evolutionary relationships Why should you refer to birds as a type of dinosaur?

Systematics and Classification Monophyletic Group

Systematics and Classification Paraphyletic Group

Systematics and Classification Polyphyletic Group

Systematics and Classification Old plant classification system

Systematics and Classification New plant classification system

Systematics and Classification Phylogenetic species concept (PSC) Focuses on shared derived characters Biological species concept (BSC) Defines species as groups of interbreeding population that are reproductively isolated Phylogenetic species concept: species should be applied to groups of populations that have been evolving independently of other groups

Systematics and Classification BSC cannot be applied to allopatric populations PSC can be applied to allopatric populations PSC can be applied to both sexual and asexual species BSC can be applied only to sexual species PSC still controversial

Systematics and Classification Paraphyly and phylogenetic species concept

Comparative Biology Phylogenetics is the basis of all comparative biology Homologous structures are derived from the same ancestral source (e.g. dolphin flipper and horse leg) Homoplastic structures are not (e.g. wings of birds and dragonflies): -Parental care Dinosaurs, birds, crocodiles Homologous behavior

Parental care in dinosaurs and crocodiles Comparative Biology Parental care in dinosaurs and crocodiles

Comparative Biology Homoplastic convergence: saber teeth Occurred in different groups of extinct carnivores Similar body proportions (cat) Similar predatory lifestyle Most likely evolved independently at least 3 times

Distribution of saber-toothed mammals Comparative Biology Distribution of saber-toothed mammals

Comparative Biology 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 Comparative Biology Convergent evolution of conducting tubes

Comparative Biology Most complex characters evolve through a sequence of evolutionary changes Modern-day birds wings, feathers, light bones, breastbone Initial stages of a character evolved as an adaptation to some environmental selective pressure First featherlike structure evolved in theropod phylogeny Insulation or perhaps decoration

Comparative Biology

Comparative Biology Phylogenetic methods can be used to distinguish between competing hypotheses 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

Comparative Biology Increase through time in proportion of species whose larvae do not disperse

Comparative Biology Two processes could produce an increase in nondispersing larvae 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

Comparative Biology

Comparative Biology Analysis indicates: Evolutionary increase in nondispersing larvae through time may be a result of both a bias in the evolutionary direction and an increase in rate of diversification Lack of evolutionary reversal

Comparative Biology Loss of larval stage in marine invertebrates Nonreversible evolutionary change Marine limpets: show direct development has evolved many times 3 cases where evolution reversed and larval stage re-evolved

Comparative Biology

Comparative Biology Phylogenetics helps explain species diversification Use phylogenetic analysis to suggest and test hypotheses Species richness in beetles Coleoptera: 60% of all animals are insects and 80% of all insects are beetles Phytophaga: clade with most herbivorous beetles Family Nemonychidae: specialized on conifers since Jurassic

Evolutionary diversification of the Phytophaga Comparative Biology Evolutionary diversification of the Phytophaga

Comparative Biology Phylogenetic explanations for beetle diversification Not the evolution of herbivory Specialization on angiosperms a prerequisite for diversification Risen 5 times independently within herbivorous beetles Angiosperm specializing clade is more species-rich than the clade most closely related

Disease Evolution HIV evolved from a simian (monkey) viral counterpart SIV First recognized in 1980’s Current estimate: >39 million people infected; > 3 million die each year SIV found in 36 species of primates Does not usually cause illness in monkeys Around for more than a million years as SIV

Disease Evolution

Disease Evolution Phylogenetic analysis of HIV and SIV First: HIV descended from SIV All strains of HIV are nested within clades of SIV Second: a number of different strains of HIV exits Independent transfers from different primate species Each human strain is more closely related to a strain of SIV than to other HIV strains

Disease Evolution Third: humans have acquired HIV from different host species