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How did we get from Linnaean Taxonomy… 2 3 5 6 8 Bacteria Archaea Archezoans Euglenoids Chrysophytes Green Algae Brown Algae Red algae Slime Molds True Fungi Bryophytes Tracheophytes Protozoans Myxozoans Multicellular Animals

To a tree-like concept of organisms? Extant Multicellular Animals Myxozoans Protozoans Tracheophytes Bryophytes True Fungi Slime Molds Red algae Brown Algae Green Algae Chrysophytes Euglenoids Archezoans Archaea Bacteria 0.5 Land! Animals 1 Multicellular 2 First Eukaryotes Extinct 3 Cyanobacterial Oxygen Long Time with Prokaryotes only 4 BYBP Origin of Life Original Cell

Emil Hans (Willi) Hennig German entomologist Hennig developed a mechanism (cladistics) to find the pathways of evolution among related organisms. It is based not only on what one sees, but on many kinds of evidence, including molecular sequences. The pathways are determined by virtue of shared derived characteristics (synapomorphies). Rather than putting organisms into Linnean taxonomic “boxes,” the cladistics process shows the pathway of evolution. Emil Hans (Willi) Hennig German entomologist 1913-1976 http://scienceblogs.com/evolvingthoughts/upload/2007/04/Willi_Hennig2.jpg

Evidence Categories History - clearer recently, more obscure anciently Fossils - stratigraphic depth, isotope decay, etc. Chemical - metabolic products such as O2, Ss Molecular - DNA sequence alterations, etc. Developmental sequences - onto- phylo- geny Biogeography - Pangea, Gondwana & Laurasia

How do we find the Evolution Pathway? Phylogenetic Systematics Inferences from comparison of extant organisms Characters-Attributes of the organism Anatomy Morphology Development Physiology Macromolecule Sequences Polarizing Character States Plesiomorphies-Ancient, shared by descendants Apomorphies-More-recent derivatives Synapomorphy-Shared among related organisms Autapomorphy-Found only in one organism Use of outgroup to compare to ingroup

Typical Cladogram Present Extant A Extant B A’ A” A’” This branching of evolution is called cladogenesis (aka: macroevolution). Extinct Transitional Forms A’ is the common ancestor of extant A and extant B Time This straight line of evolution is called anagenesis (aka: microevolution). Common Ancestor Ancient

5 species of frogs found locally What could have made them speciate? http://zoology.unh.edu/faculty/litvaitis/Research/woodfrog.jpg http://www.dnr.state.wi.us/org/caer/ce/eek/critter/amphibian/images/northernLeopardFrog.jpg https://www.denix.osd.mil/denix/Public/Library/NCR/PhotoGallery/NR-Bull-frog.jpg Rana sylvatica Wood Frog Rana sphenocephala Leopard Frog http://www.turtletrack.org/Issues03/Co05172003/Art/Green_Frog.jpg Rana catesbeiana Bull Frog http://www.wingwatchers.com/images2/pickfrog1.jpg What could have made them speciate? Rana palustris Pickerel Frog Rana clamitans Green Frog

Mating Activity Leopard frog Pickerel frog Wood frog Green frog Could the size of the pond, availability of tadpole food, etc. be pivotal? Which frog was mating in mid-March? Which frog was mating in early May? When were three species of frog mating? Why do you think Green and Bull frogs mated so much later? Leopard frog Pickerel frog Wood frog Green frog Bull frog Mating Activity March April May June July Figure 1. Hypothetical phenology of frog mating behavior in a pond in the north temperate zone.

Grand Canyon squirrels: Sciurus aberti kaibabensis Sciurus aberti aberti ©1996 Norton Presentation Maker, W. W. Norton & Company North Rim South Rim

Model of geographic speciation: interbreeding between populations decreases Kaibab Squirrel North Rim one species with unrestricted interbreeding erosion begins Grand Canyon prevents interbreeding Abert Squirrel South Rim Time (10,000 years)

Typical Cladogram Present Extant A Extant C Extant B A’ A’ A B constitute a clade A” C are a grade (is paraphyletic) A” Time A” C A’ A B are a clade (is monophyletic) A’” A’ A B is the sister group of C Common Ancestor Ancient

Typical Cladogram Present Extant A Extant C Extant B Extant D Extant E D A””’ E are a ? clade A”” Extinct! Common ancestor + A””’ D E are a ? grade A” Time A’” The ABC clade may be, say, a genus. The DE clade may be another genus… in the same family Common Ancestor Ancient The ABCDE clade would be the family

Typical Cladogram Present Extant A Extant B Extant C Extant D Extant E On the other hand… A””’ Extinct! AB are a genus C is a monotypic genus DE are a genus A” Time A”” A’” ABC might be one family DE are in another family Common Ancestor Ancient ABCDE might constitute an order

Protists are polyphyletic (unnatural taxon) Extant Multicellular Animals Myxozoans Protozoans Tracheophytes Bryophytes True Fungi Slime Molds Red algae Brown Algae Green Algae Chrysophytes Euglenoids Archezoans Archaea Bacteria Animals and Fungi are a clade! Plants are a clade (monophyletic) Extinct Eukaryotic organisms are a clade Prokaryotic organisms are a grade (paraphyletic) Living organisms are part of one clade (monophyletic) Original Cell

This is the tree-of-life cladogram used by your textbook Fig. 29.8 Pg 601 It varies in some minor ways, mostly additions. It shows “uncertainty” as a multifurcation.

Bacterial Phyla Extant Extinct Gram negative Gram positive Rhodopseudomonads Purple nonsulfur Extant Mycoplasmas Actinomycetes Prochlorophytes Clostrids Spirochetes Green Sulfur Cyanobacteria Purple Sulfur Desulfovibrios Myxobacteria Rikettsias +Para- site +Chl B -Cell Wall +Parasite H2S +µtubule H2O H2S +Chl A +gliding motility H2O +NAD +bacteriochlorophyll Gram negative +NADP Extinct Each simple line on our tree of life is, in fact, branches with twigs! Each blue hatch mark shows an evolutionary step…mostly biochem! Gram positive Original Cell

This is the textbook’s version of the Prokaryotic Cladogram Fig. 28.12 Pg 577 It differs mostly by addition of the Archaea.

A dendrogram is not a cladogram http://genomics.senescence.info/evolution/tree.gif

Is this a cladogram? http://www.nature.com/nature/journal/v428/n6982/images/428475a-f2.2.jpg

Patterns of evolution Time Divergent Parallel Convergent http://www.mobot.org/MOBOT/Madagasc/phytogeographic/gondwana.jpg Patterns of evolution Divergent Parallel Convergent Time

A cladogram of marsupials http://blogs.discovermagazine.com/gnxp/files/2010/08/0.jpeg

Notice how the marsupials of Australia are a clade (monophyletic). This is an area cladogram in which the cladogram is stretched over geographic domains. Notice how the marsupials of Australia are a clade (monophyletic). It is thought that this represents a single colonization followed by adaptive radiation (rapid speciation). http://blogs.discovermagazine.com/gnxp/files/2010/08/journal.pbio.1000436.g002.png

Marsupials are convergent with placentals of other continents: Marsupial mouse: Sminthopsis dolichura ©1996 Norton Presentation Maker, W. W. Norton & Company

Marsupials are convergent with placentals of other continents: Marsupial glider: Petauroides volans ©1996 Norton Presentation Maker, W. W. Norton & Company

Marsupials are convergent with placentals of other continents: Marsupial tiger cat: Dasyurus maculatus ©1996 Norton Presentation Maker, W. W. Norton & Company

Marsupials are convergent with placentals of other continents: Cuscus: Phalanger maculatus ©1996 Norton Presentation Maker, W. W. Norton & Company