Macroevolution: Large Scale Changes Over Time

1. Chapter 25 The History of Life on Earth

2. Overview: Lost Worlds • Past organisms were very different from those now alive. • The fossil record shows macroevolutionarychanges over large time scales including • The emergence of terrestrial vertebrates • The origin of photosynthesis • Long-term impacts of mass extinctions.

3. MacroEvolution: Large Scale Changes Over Time

4. Concept 25.1: Conditions on early Earth made the origin of life possible • Chemical and physical processes on early Earth may have produced very simple cells through a sequence of stages: 1. Abiotic synthesis of small organic molecules. 2. Joining of these small molecules into macromolecules. 3. Packaging of molecules into “protobionts.” 4. Origin of self-replicating molecules.

5. Synthesis of Organic Compounds on Early Earth • Earth formed about 4.6 billion years ago, along with the rest of the solar system. • Earth’s early atmosphere likely contained water vapor and chemicals released by volcanic eruptions (nitrogen, nitrogen oxides, carbon dioxide, methane, ammonia, hydrogen, hydrogen sulfide). • A. I. Oparin and J. B. S. Haldane hypothesized that the early atmosphere was a reducing environment. • Stanley Miller and Harold Urey conducted lab experiments that showed that the abiotic synthesis of organic molecules in a reducing atmosphere is possible.

6. However, the evidence is not yet convincing that the early atmosphere was in fact reducing. • Instead of forming in the atmosphere, the first organic compounds may have been synthesized near submerged volcanoes and deep-sea vents. • Amino acids have also been found in meteorites.

7. Deep Sea Vents

8. Abiotic Synthesis of MacromoleculesMonomers --> Polymers • Small organic molecules polymerize when they are concentrated on hot sand, clay, or rock. • Replication and metabolism are key properties of life. • Protobiontsare aggregates of abiotically produced molecules surrounded by a membraneor membrane-like structure. • Protobionts exhibit simple reproduction and metabolism and maintain an internal chemical environment.

9. Experiments demonstrate that protobionts could have formed spontaneously from abiotically produced organic compounds. • For example, small membrane-bounded droplets called liposomes can form when lipids or other organic molecules are added to water.

10. Protobionts May Have Formed Spontaneously 20 µm Glucose-phosphate Glucose-phosphate Phosphatase Starch Amylase Phosphate Maltose (a) Simple reproduction by liposomes Maltose (b) Simple metabolism

11. Self-Replicating RNA and the Dawn of Natural Selection • The first genetic material was probably RNA, not DNA. • RNA molecules called ribozymeshave been found to catalyze many different reactions • For example, ribozymes can make complementary copies of short stretches of their own sequence or other short pieces of RNA.

12. Sedimentary Rocks and Fossils • Sedimentary strata reveal the relative ages of fossils. • The absolute ages of fossils can be determined by radiometric dating. • A “parent” isotope decays to a “daughter” isotope at a constant rate. • Each isotope has a known half-life, the time required for half the parent isotope to decay.

13. Sedimentary Rock Strata -- Fossils Rhomaleosaurus victor, a plesiosaur Present Dimetrodon 100 million years ago Casts of ammonites 175 200 270 300 Hallucigenia 4.5 cm 375 Coccosteus cuspidatus 400 1 cm Dickinsonia costata 500 525 2.5 cm 565 Stromatolites Tappania, a unicellular eukaryote 600 3,500 1,500 Fossilized stromatolite

14. Radiometric Dating Accumulating “daughter” isotope Fraction of parent isotope remaining 1/2 Remaining “parent” isotope 1/4 1/8 1/16 4 3 2 1 Time (half-lives)

15. Radiocarbon dating can be used to date fossils up to 75,000 years old. • For older fossils, some isotopes can be used to date sedimentary rock layers above and below the fossil. • The magnetism of rocks can provide dating information. • Reversals of the magnetic poles leave their record on rocks throughout the world.

16. The Origin of New Groups of Organisms • Mammals belong to the group of animals called tetrapods. • The evolution of unique mammalian features through gradual modifications can be traced from ancestral synapsids through the present.

17. Evolution of Mammals Synapsid (300 mya) Temporal fenestra Key Articular Dentary Quadrate Squamosal Therapsid (280 mya) Reptiles (including dinosaurs and birds) Temporal fenestra EARLY TETRAPODS Dimetrodon Early cynodont (260 mya) Synapsids Very late cynodonts Temporal fenestra Earlier cynodonts Therapsids Later cynodont (220 mya) Mammals Very late cynodont (195 mya)

18. Concept 25.3: Key events in life’s history include the origins of single-celled and multicelled organisms and the colonization of land • The geologic recordis divided into the Archaean, the Proterozoic, and the Phanerozoic eons. • The Phanerozoic encompasses multicellular eukaryotic life. • The Phanerozoic is divided into three eras: the Paleozoic, Mesozoic, and Cenozoic. • Major boundaries between geological divisions correspond to extinction events in the fossil record.

19. Geologic Record

20. Geologic Time Table Ceno- zoic Meso- zoic Humans Paleozoic Colonization of land Animals Origin of solar system and Earth 4 1 Proterozoic Archaean Prokaryotes years ago Billions of 3 2 Multicellular eukaryotes Single-celled eukaryotes Atmospheric oxygen

21. The First Single-Celled Organisms = Prokaryotes • The oldest known fossils are stromatolites, rock-like structures composed of many layers of bacteria and sediment. • Present-day stromatolites are found in a few warm, shallow, salty bays. • If complex bacterial communities existed 3.5 billion years ago, it seems reasonable that life originated much earlier, perhaps 3.9 billion years ago, when Earth first cooled to a temperature at which liquid water could exist. • Prokaryotes were Earth’s sole inhabitants from 3.5 to about 2.1 billion years ago.

22. Photosynthesis and the Oxygen Revolution • Most atmospheric O2 is of biological origin, from the water-splitting step of photosynthesis. • When oxygenic photosynthesis first evolved, the free O2 it produced likely dissolved in the surrounding water. • When O2 reached a high enough concentration to react with dissolved iron, the iron precipitated as iron oxide, which accumulated as red layers of rock in banded iron formations.

23. Additional O2 dissolved in the seas and lakes until they were saturated with O2. • After this, the O2 finally began to “gas out” of water and enter the atmosphere. • About 2.7 billion years ago, oxygen began accumulating in the atmosphere and terrestrial rocks with oxidized iron formed. • Although oxygen accumulation was gradual between 2.7 and 2.2 billion years ago, it shot up to between 1% and 10% of current values shortly afterward.

24. About 2.7 billion years ago, O2 began accumulating in the atmosphere and rusting iron-rich terrestrial rocks.

25. This “oxygen revolution” had an enormous impact on life. • In certain chemical forms, oxygen attacks chemical bonds, inhibits enzymes, and damages cells. • The increase in atmospheric oxygen likely doomed many prokaryote groups. • Some species survived in habitats that remained anaerobic, where their descendents survive as obligate anaerobes. • Other species evolved mechanisms to use O2 in cellular respiration, which uses oxygen to help harvest the energy stored in organic molecules.

26. The First Eukaryotes • Eukaryotic cells differ in many respects from the smaller cells of bacteria and archaea. • Even the simplest single-celled eukaryote is far more complex in structure than any prokaryote, with a nuclear envelope, mitochondria, and endomembrane system.

27. The First Eukaryotes • How did the complex organization of the eukaryotic cell evolve from the simpler prokaryotic condition? • A process called endosymbiosis probably led to mitochondria and plastids (the general term for chloroplasts and related organelles). • The endosymbiont theory suggests that mitochondria and plastids were formerly small prokaryotes that began living within larger cells. • The prokaryotic ancestors of mitochondria and plastids probably gained entry to the host cell as undigested prey or internal parasites. • As they became increasingly interdependent, the host and endosymbionts became a single organism.

28. The prokaryotic ancestors of mitochondria and plastids probably gained entry to the host cell as undigested prey or internal parasites. • In the process of becoming more interdependent, the host and endosymbionts would have become a single organism. • Serial endosymbiosissupposes that mitochondria evolved before plastids through a sequence of endosymbiotic events.

29. Key evidence supporting an endosymbiotic origin of mitochondria and plastids: • Similarities in inner membrane structures and functions. • These organelles transcribe and translate their own DNA. • Their ribosomes are more similar to prokaryotic than eukaryotic ribosomes.

30. The Origin of Multicellularity • The evolution of eukaryotic cells allowed for a greater range of unicellular forms. • A second wave of diversification occurred when multicellularity evolved and gave rise to algae, plants, fungi, and animals. • Comparisons of DNA sequences date the common ancestor of multicellular eukaryotes to 1.5 billion years ago. • The oldest known fossils of multicellular eukaryotes are of small algae that lived about 1.2 billion years ago.

31. The Origin of Multicellularity • Why were multicellular eukaryotes so limited in size, diversity, and distribution until the late Proterozoic? • Geologic evidence suggests that a series of severe ice ages gripped Earth from 750 to 580 million years ago. • At times during this period, glaciers covered all of Earth’s landmasses, and the seas were largely iced over. • According to the “snowball Earth” hypothesis, life would have been confined to deep-sea vents and hot springs or to equatorial regions of the ocean that lacked ice cover.

32. The Cambrian Explosion • The first major diversification of multicellular eukaryotic organisms corresponds to the time of the thawing of snowball Earth, about 575 million years ago. • Many phyla of animals appear suddenly in the fossil record in a phenomenon known as the Cambrian explosion. • Sponges, cnidarians (the phylum that includes sea anemones), and molluscs appear in older rocks from the late Proterozoic. • Prior to the Cambrian explosion, animals were small, soft-bodied, or both, and there was little evidence of predation. • Within 10 million years, predators longer than 1 meter emerged, with claws and other predatory structures. Prey showed defensive adaptations, such as sharp spines and heavy body armor.

33. Proterozoic Fossils that may be animal embryos (SEM) (a) Two-cell stage (b) Later stage 200 µm 150 µm

34. The Colonization of Land • Fungi, plants, and animals began to colonize land about 500 million years ago. • Plants and fungi likely colonized land together by 420 million years ago. • Arthropods and tetrapods are the most widespread and diverse land animals. • Tetrapods evolved from lobe-finned fishes around 365 million years ago.

35. Concept 25.4: The rise and fall of dominant groups reflect continental drift, mass extinctions, and adaptive radiations • At three points in time, the land masses of Earth have formed a supercontinent: 1.1 billion, 600 million, and 250 million years ago. • Earth’s continents move slowly over the underlying hot mantle through the process of continental drift. • Oceanic and continental plates can collide, separate, or slide past each other. • Interactions between plates cause the formation of mountains and islands, and earthquakes.

36. Earth - Plate Tectonics: Continental Drift North American Plate Eurasian Plate Crust Caribbean Plate Philippine Plate Juan de Fuca Plate Arabian Plate Indian Plate Cocos Plate Mantle South American Plate Pacific Plate Nazca Plate African Plate Outer core Australian Plate Inner core Antarctic Plate Scotia Plate (b) Major continental plates (a) Cutaway view of Earth

37. Consequences of Continental Drift • Formation of the supercontinentPangaeaabout 250 million years ago had many effects: • A reduction in shallow water habitat • A colder and drier climate inland • Changes in climate as continents moved toward and away from the poles • Changes in ocean circulation patterns leading to global cooling.

38. History of Continental Drift Present Cenozoic Eurasia North America Africa 65.5 India South America Madagascar Australia Antarctica Laurasia 135 Mesozoic Gondwana Millions of years ago Pangaea 251 Paleozoic

39. The break-up of Pangaea lead to allopatric speciation. • The current distribution of fossils reflects the movement of continental drift. Similarity of fossils in parts of South America and Africa supports the idea that these continents were formerly attached. • The fossil record shows that most species that have ever lived are now extinct. • At times, the rate of extinction has increased dramatically and caused a mass extinction. • In each of the five mass extinction events, more than 50% of Earth’s species became extinct.

40. Five Big Mass Extinctions 800 20 700 600 15 500 Number of families: 400 Total extinction rate (families per million years): 10 300 200 5 100 0 0 Mesozoic Paleozoic Cenozoic Era Period E C Tr C O S D P J P N 200 145 65.5 0 542 488 444 416 359 299 251 Time (millions of years ago)

41. The Permian extinction defines the boundary between the Paleozoic and Mesozoic eras. • This mass extinction caused the extinction of about 96% of marine animal species and might have been caused by volcanism, which lead to global warming, and a decrease in oceanic oxygen. • The Cretaceous mass extinction 65.5 million years ago separates the Mesozoic from the Cenozoic. • Organisms that went extinct include about half of all marine species and many terrestrial plants and animals, including most dinosaurs.

42. Massive Meterorite Impact Evidence • The presence of iridium in sedimentary rocks suggests a meteorite impact about 65 million years ago. • The Chicxulub crater off the coast of Mexico is evidence of a meteorite that dates to the same time.

43. Evidence of Meteroite Impact NORTH AMERICA Chicxulub crater Yucatán Peninsula

44. Is a Sixth Mass Extinction Under Way? Consequences … • Scientists estimate that the current rate of extinction is 100 to 1,000 times the typical background rate. • Data suggest that a sixth human-caused mass extinction is likely to occur unless dramatic action is taken. • Mass extinction can alter ecological communities and the niches available to organisms. • It can take from 5 to 100 million years for diversity to recover following a mass extinction. • Mass extinction can pave the way for adaptive radiations.

45. Adaptive Radiations - New Environmental Opportunities … • Adaptive radiationis the evolution of diversely adapted species from a common ancestor upon introduction to new environmental opportunities. • Mammals underwent an adaptive radiation after the extinction of terrestrial dinosaurs. • The disappearance of dinosaurs (except birds) allowed for the expansion of mammals in diversity and size. • Other notable radiations include photosynthetic prokaryotes, large predators in the Cambrian, land plants, insects, and tetrapods.

46. World-Wide Adaptive Radiations Ancestral mammal Monotremes (5 species) ANCESTRAL CYNODONT Marsupials (324 species) Eutherians (placental mammals; 5,010 species) 50 200 250 100 150 0 Millions of years ago

47. Regional Adaptive Radiations • Adaptive radiations can occur when organisms colonize new environments with little competition. • The Hawaiian Islands are one of the world’s great showcases of adaptive radiation.

48. Hawaiian Islands -- Regional Adaptive Radiations Close North American relative, the tarweed Carlquistia muirii 1.3 million years MOLOKAI KAUAI 5.1 million years Dubautia laxa MAUI OAHU 3.7 million years Argyroxiphiumsandwicense LANAI HAWAII 0.4 million years Dubautia waialealae Dubautia scabra Dubautia linearis

49. Major changes in body form result from changes in the sequences and regulation of developmental genes • Studying genetic mechanisms of change can provide insight into large-scale evolutionary change. • Genes that program development control the rate, timing, and spatial pattern of changes in an organism’s form as it develops into an adult. • Heterochronyis an evolutionary change in the rate or timing of developmental events. • It can have a significant impact on body shape. • The contrasting shapes of human and chimpanzee skulls are the result of small changes in relative growth rates.

50. Allometric Growth 15 Newborn 5 Adult 2 Age (years) (a) Differential growth rates in a human Chimpanzee adult Chimpanzee fetus Human adult Human fetus (b) Comparison of chimpanzee and human skull growth