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Charles Darwin: Evolutionary Theory, Past and Present The Evidence for Evolution by Natural Selection from C. Darwin, 1859, Origin of Species “There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved.” from C. Darwin, 1859, Origin of Species PREDICTIONS OF THE THEORY 1) Descent with modification, 2) From common ancestors, 3) Producing a branching tree of life, 4) Connected by intermediate species, 5) Characterized by extinction of less fit taxa, 6) And superior adaptation of those who survived. The variations acted on by selection are random, so there is no apparent direction or purpose evident in the history of life, but adaptation to environmental parameters, which governs natural selection, is anything but random. The biosphere is a marvel of myriad adaptations! THE EVIDENCE FOR EVOLUTION A. Fossil record evidence of speciation, adaptive radiation events, and extinctions. B. Observations of modern organisms - biogeography, homologous structures, developmental trends, molecular biology. C. Examples of ongoing evolution - Resistance to pesticides, the pathology of the AIDS virus, Galapagos Finches. EVIDENCE FROM THE FOSSIL RECORD Stratigraphy Stratigraphy Faunal succession P D P M C pC Extinction Stratigraphy Faunal succession P D P M C pC Extinction Paleogeography W. W. Norton. Modified from Wegener A, The Origin of the Continents and Oceans. Paleobiogeography (land dwellers) (seeds don’t spread far) Paleobiogeography ©Blackwell, 1990 (Sheldon, 1987) Microevolution Parallel trends in trilobite morphology Speciation in Brachiopods 1.7 measured by change in ornament through time 5.2 Definition of paleospecies? Average species duration? 1.1 2.0 Speciation Patterns Predicted B •Lack of intermediate forms •Arbitrary paleospecies definitions A Speciation Patterns Predicted Observed D B C •Species stability WHY? A What is the average species duration? 5 - 10 myr B A Normalizing Selection Habitat tracking BRANCHING PHYLOGENY EXHIBITS GRADUAL PHYLETIC EVOLUTION IN EOCENE MAMMAL LINEAGE Gingerich, P.D., 1974, Stratigraphic record of Early Eocene Hyopsodus and the geometry of mammalian phylogeny, Nature 248, 107 - 109. Lineage evolution Orthogenetic Trend? Or species selection? Macroevolution and the Evidence for Transitional Forms Macroevolutionary Events F A M T L,S C Mesozoic B C D K J AR T Paleozoic P P Birds (B) M Mammals (M) D Reptiles (T,L,S,C,D) S Amphibians (A) Fish (F) Macroevolution fish to amphibians Where are the missing links? Macroevolution - fish to amphibians Ichthyostega Eusthenopteron Fish Land Vertebrates Can we find a transitional form at the exact fish-to-amphibian transition? Tiktaalik Edward B. Daeschler, Neil H. Shubin and Farish A. Jenkins, Jr (6 April 2006). "A Devonian tetrapod-like fish and the evolution of the tetrapod body plan". Nature 440: 757–763. Shubin, Neil (2008). Your Inner Fish. Pantheon. ISBN 9780375424472. Theraspids Macroevolution Reptiles to mammals Mammal-like reptiles A M T L,S C B C D K J AR T P P M D S adaptation to terrestrial habitats Birds Mammals Reptiles Amphibians Macroevolution reptiles to birds A M C T L,S B C D K J AR T P P M D S Birds Mammals Reptiles Amphibians Macroevolution - reptiles to birds Adaptation to aerial habitats A M C T L,S B C D K J AR T P P M D S Birds Mammals Reptiles Amphibians Archaeopteryx lithographica Macroevolution - tetrapods to whales “Darwin seems to believe that a white bear …might be turned into a whale.” Evolution of marine mammals Re-adaptation to marine habitats Genetic evidence shows most closely related living land creature to Cetaceans is… Anthracotheriidae Adaptation to terrestrial habitats What were they waiting for? DINOSAURIA Radiation of all mammal clades The Cambrian Explosion CAMBRIAN EXPLOSION Sudden appearance of all extant body plans, as well some completely novel plans never seen again. Total diversity far exceeding that of modern taxa. Ediacarans Innovation Intense selection Innovation Burgess Shale Fauna Intense selection Large scale trends in the history of life Phanerozoic Evolutionary History Ordovician radiation Cambrian radiation Mesozoic radiation Sepkoski’s Evolutionary Faunas A. Marine Invertebrates B. Vascular Plants C. Tetrapods No new body plans Angiosperms Gymnosperms Turtles-mammals Diapsids-Dinosaurs Pteridophytes Labyrinthodonts Anapsids Synapsids Low diversity scavengers Trilobites, Inarticulate Brachiopods Generalized deposit feeding Sessile benthos Brachiopods, Bryozoa, Molluscs Rise of vertebrates (Fish) Dominant epifaunal suspension feeders Mobile benthos Molluscs, Echinoderms Marine reptiles Marine mammals Complex food webs, many more guilds, durophagous predators and deep tiering Evolution of Durophagous Predators Shell-crushing fish Marine reptiles Evidence for Natural Selection THE FOSSIL RECORD shows… Proof of extinction (ultimate selection) Evidence of adaptive radiation (lineages) Transitional or intermediate forms Large scale trends in the history of life No directionality, only adaptation and contingency B D Triassic Permian Equus Consider the significance of the statement “no reversals have been found.” Descent with modification PREDICTS that there should be no reversals in the history of life - in other words, the occurrence of a mammal in the Precambrian would invalidate evolutionary theory as we know it. The History of Life is Contingent! But how good is the fossil record? •Overall disadvantages Fragmentary and full of gaps Subject to spatial/temporal biases Environments - terrestrial vs. marine Taxa - invertebrate vs. vertebrate Historical - recent vs. deep past How biased? Diversity “The Pull of the Recent” amount of exposed rock of a given time period But this doesn’t explain Sepkoski’s evolutionary faunas… Exposure Equil model ©Blackwell, 1990 # of paleontologists working within a time period Sampling ©Blackwell, 1990 Exceptional Preservation: Fossil Lagerstätten ©Natl. Geographic But the more common situation… Is the fossil record a representative sample? “…light will be thrown on the origin of man.” ©Natl. Geographic African Rift Valley EVIDENCE FROM MODERN ORGANISMS Island Biogeography The role of geographic isolation Galapagos Islands Adaptation & Speciation STRUCTURAL ADAPTATIONS RELATED TO LIFE HABIT AND RESOURCE EXPLOITATION ©Benjamin Cummings Publ. 1979 Galapagos Finches ‘Island’ Biogeography & the Founder Effect ©Sci. American, 1978 “topographic islands” & isolation since the Last Glacial Maximum (20 Ka). Biogeography Ostrich Rhea Emu (land dwellers) Kiwi Penguin (seeds don’t spread far) Convergent evolution Analogous Structures Anatomical structures that perform the same function in different biological species and evolved from different structures in unrelated lineages Analogous structures reflect convergent evolution …reflecting the constraints of adaptation to a particular set of environmental conditions. ©Sci. American, 1978 ANALOGOUS STRUCTURES Homologous Structures Anatomical structures that perform the same function in different biological species and evolved from the same structure in some ancestor species HOMOLOGOUS STRUCTURES REPTILES BIRDS Bone Differences reflect ADAPTATION Sternum Skin ©Sci. American, 1978 Forelimb Homology in mammal forearms …adaptation to different life habits Diversification of CLAMS from common ancestor (HAM) de ep er ©A. Seilacher, 1987 thicker A great example of niche partitioning VESTIGIAL ORGANS Human hair follicle VESTIGIAL ORGANS Hind limbs Neoteny in primates Barnacle life histories Homeotic genes ©Sci. American. 1978 DEVELOPMENTAL EVIDENCE ©Benjamin Cummings Publ. 1979 ©S.J. Gould, 1989, Norton Publ. CLASSIFICATION CLADISTICS Derived Primitive Systematics based on grouping of shared primitive vs. shared derived characters to establish ancestordescendent relationships. Cladistic Phylogeny Derived Primitive Molecular Phylogeny based on DNA sequencing of Galapagos Finches Common ancestor Molecular Phylogeny DNA Protein Galapagos islands Finch BMP4 African lakes Cichlid National Geographic, Feb. 2009 Molecular Phylogeny ©National Acad. Sci., 1998 Divergence the in gene for Cytochrome C Molecular Phylogeny 16S Ribosomal RNA sequence ©Natl. Acad. Sci., 1998 Homologies of molecular biology? A. The genetic code is the same in all known organisms. B. The “Central Dogma” of cellular metabolism is the same in all known organisms. C. The fundamental biochemical agents that mediate metabolism (hemoglobin, cytochrome c, etc.) are found in a very wide range of known organisms. EXAMPLES OF ONGOING EVOLUTION ©Natl. Geographic Artificial selection Microevolution Peppered Moths Variations in dominant coloration develop at the population level as environmental conditions (tree color) change. The selecting mechanism is predation by birds. ©Sci. American, 1978 Due to environmental changes in Britian since the Industrial Revolution… The humble Soapberry bug Galapagos Finches DROUGHT DROUGHT DROUGHT DROUGHT Resistance to pesticides Mosquitos in India AIDS The Global AIDS Epidemic Selected areas shown w/total # infected 15-45 yr age group 2006: >38 million infected 2050: estimated >90 million 2.9 million died dead HIV mechanism 1.Virion stage. 2. HIV gp120 prot. binds to host CD4 & coreceptor. 3. HIV RNA, RT, I, & P enter host. RT- Reverse Transcriptase I - Integrase P - Protease 4. RT synthesizes HIV DNA from HIV RNA template. 5. I splices HIV DNA into host genome, it gets transcribed to mRNA by host. 6.HIV mRNA is translated to HIV proteins by host ribosomes; new generation of virions assembles, & buds from host. Progression of HIV infection in typical untreated patient Immune system collapses Evolution of the HIV population w/in a single patient SHAMELESS PROMOTION ~ 95 Ma Modified from Sewall et al., 2007 Cretaceous Ocean Anoxic Events Tibet Carbon isotope excursion at the Cenomanian - Turonian Boundary δ13C ‰ PDB Li, et al., 2006 secondary mass extinction Carbonate TOM Hasegawa et al., 2002 SUN H2O + CO2 <=> CH2O + O2 Photosynthesis Respiration CO2 PLANKTON H2O ventilation O2 CO2 ventilation H2O + CO2 <=> CH2O + O2 CO2 -O2 -O2 reduced ventilation reduced ventilation BLACK SHALE M.m. S.g. N.j. Rock Canyon Anticline, near Pueblo, CO δ13C Eicher & Diner, 1991 Hartland SH Bridge Creek LS Cenomanian Turonian W.d P.f. V.b. Cenomanian-Turonian stratotype Detailed correlation of marine record to coeval terrestrial strata. Bald Knoll Mine, UT Development of improved time scale for marine C-T Roberts & Kirschbaum, 1995 Marine-terrestrial linkage of OAE 2 Cuticle morphotyping Lauraceae ( Stomatal Index = SD SDSD + ED x 100 + ED ) X 100 pCO2 = 1 Stomatal Index SD = # stomata per mm2 C. c. M. mosbyense S. gracile Cenomanian N. juddii W. dev. Turonian SD vs SI – CO2 concentrations Modern Eocene RCO2 = 4x? RCO2=1x RCO2 = 3x - 2x Cretaceous Modern and Eocene Data from McElwain, 1988 CONCLUSIONS •The evidence for descent with modification, common ancestors, speciation to produce a branching tree of life, intermediate species, extinction, and superior adaptation is abundant and uncontestable. •Evolution by means of natural selection is the central unifying principle of the biosphere… “nothing in biology makes sense except in the light of evolution.” (T. Dobzhansky) •The biosphere IS a marvel of myriad adaptations! •And there is indeed, grandeur in this view of life.