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Anu Singh-Cundy • Gary Shin Discover Biology SIXTH EDITION CHAPTER 16 The Evolutionary History of Life © 2015 W. W. Norton & Company, Inc. CHAPTER 16 The Evolutionary History of Life, Part 1 PUZZLING FOSSILS IN A FROZEN WASTELAND 16.1 Macroevolution: Large-Scale Body Changes 16.2 The Fossil Record: A Guide to the Past The fossil record is incomplete Fossils reveal that whales are closely related to a group of hoofed mammals 16.3 The History of Life on Earth The first single-celled organisms arose at least 3.5 billion years ago Multicellular life evolved about 650 million years ago Colonization of land followed the Cambrian explosion 16.4 The Effects of Plate Tectonics 16.5 Mass Extinctions: Worldwide Losses of Species CHAPTER 16 The Evolutionary History of Life, Part 2 16.6 Rapid Macroevolution through Differential Gene Expression 16.7 Phylogenetics: Reconstructing Evolutionary Relationships The Linnaean system of biological classification reflects evolutionary history BIOLOLGY MATTERS: IS MASS EXTINCTION UNDER WAY? APPLYING WHAT WE LEARNED: WHEN ANTARCTICA WAS GREEN Puzzling Fossils in a Frozen Wasteland • Paleontologists have uncovered signs indicating that life in Antarctica once included dinosaurs and other reptiles, mammals and flightless birds, ferns and enormous trees, amphibians, freshwater fishes, and aquatic beetles. The Earth Is Billions of Years Old • In the nineteenth century, Lord Kelvin estimated the age of Earth at no more than ~100 million years. • Darwin recognized that while natural selection is potent enough to generate adaptations in organisms, very large timescales would be necessary to explain the great diversity of body plans observed in nature. • Radioisotopes are unstable forms of elements that decay to more stable forms at a constant rate over time. • History of Earth as a 24-hour day: Dinosaurs evolved just before 11:00 PM, and humans, just a minute before midnight . The application of isotope studies to geology brought the current estimate of Earth’s age to approximately 4.6 billion years. The Climate and Geology of Our Planet Have Changed over Hundreds of Millions of Years, Causing Dramatic Changes in the Life-forms on Earth • In the time span since Earth was formed, known as geologic time, both the land and climate have gone through large changes over and over again. • Geological time provides sufficient time for the process of natural selection to create large-scale changes in organisms. Hot plumes of liquid rock from Earth’s interior can rise to the surface and push the continents away from each other, as at this rift in Iceland. • About 1.7 million species have been described, and millions more await discovery. • However, present-day species are thought to represent less than 1 percent of all the species that have ever lived because many life forms have become extinct. Macroevolution Describes Large-Scale Alterations in Form • Organisms differ radically in form, in embryological development, in body plans, in behavior, and in ecology. • Macroevolution refers to large-scale changes in organisms, generally occurring over millions of years. The body plan of whales has been radically altered by adaptations to aquatic life compared with the body plan of their terrestrial ungulate (hoofed) ancestors. The Fossil Record Documents the History of Life on Earth • • • Fossils are the preserved remains or impressions of individual organisms that lived in the past, and are often found in sedimentary rock. Fossils provide evidence that past organisms were unlike organisms alive today, that many forms have disappeared from Earth completely, and that life has evolved through time. Older fossils are found in deeper, older rock layers. The order in which organisms appear in the fossil record agrees with our understanding of evolution based on other evidence, providing strong support for evolution. A Fossil’s Age Can Be Estimated Using Radioisotopes • Carbon dating can reveal the age of relatively recent fossils. For a given amount of the radioisotope carbon-14 (14C), half of the total decays to the stable element carbon-12 every 5,730 years. By measuring the amount of 14C that remains in a fossil, scientists can estimate the age of the fossil. • Carbon-14 can be used to date only relatively recent fossils: too little 14C remains to date fossils formed more than 70,000 years ago. • Other radioisotopes reveal the age of the more ancient fossils. Elements such as uranium-235, which has a half-life of 700 million years, can be used to date much older materials. • If a fossil does not contain any radioisotopes, methods like carbon or uranium dating enable us to determine an approximate date for the fossil by dating rocks found above and below the fossil. The Fossil Record Is Not Complete The fossil record contains many gaps because: • Most organisms decompose rapidly after death and do not form fossils. • Other have few hard parts and do not fossilize well for that reason. • Organisms may not live near areas where sediments typically form. • Those fossils that are formed can be destroyed by common geologic processes such as erosion and extreme heat or pressure. • Fossils are hard to find. It takes a unique set of circumstances to form and preserve fossils, and for fossils to be discovered. Fossils Reveal That Whales Are Closely Related to a Group of Hoofed Mammals • Whales have a relatively complete fossil record, which enables scientists to observe many transitional forms that document their evolutionary past. • Whales show the greatest DNA similarity to artiodactyls, which are even-toed, hoofed land mammals (such as hippos, camels, and giraffes). The History of Life on Earth: Single-Celled Life Appeared by 3.5 Billion Years Ago • The oldest known rocks on Earth (3.8 billion years old) contain carbon deposits that hint at life. • Cell-like structures have been found in stromatolites that formed 3.5 billion years ago. • DNA analysis also supports the idea that life had appeared on Earth by 3.5 billion years ago. Stromatolites are sediment mounds containing fossilized microorganisms. • The first life-forms were prokaryotes. • Eukaryotes are first seen in the fossil record at about 2.1 billion years ago. The Evolution of Oxygen-Producing Photosynthesis Was One of the Most Important Events in the History of Life on Earth • Earth’s atmosphere initially contained almost no oxygen. • Roughly 2.8 billion years ago, a group of bacteria evolved a type of photosynthesis that releases oxygen as a by-product. • As a result, oxygen began to accumulate in the atmosphere. • Eukaryotes, with their larger cells and therefore greater energy needs, evolved when oxygen reached at least 2–3 percent of present-day levels. • Multicellular life, with their even greater energy demands, evolved when oxygen levels reached presentday levels. Multicellular Life Evolved about 650 Million Years Ago • The first multicellular organisms—soft-bodied animals—evolved in the shallow seas that covered the Earth during the Precambrian period, about 650 mya. The Cambrian Period Witnessed Multiple Adaptive Radiations of Animal Life • • Starting in the early to middle Cambrian period, 530 mya, there was a dramatic increase in the diversity of animal life, known as the Cambrian explosion, which lasted only 5–10 million years. The presence of new predators is thought to have sped up the evolution of Cambrian herbivores. Colonization of Land Followed the Cambrian Explosion • • • • Green algae were the first organisms to colonize land, about 500 mya. Plants covered Earth by the end of the Devonian period, 360 mya. Fungi appeared on land at about the same time as plants; the two groups developed mutualisms. The first definite fossils of terrestrial animals are of spiders and millipedes and date from about 410 mya. Waterproofing, Efficient Transport Mechanisms, and Roots Helped Land Plants Acquire and Conserve Water Land plants evolved key innovations to deal with the challenges of terrestrial life: - Waterproof cuticle - Vascular systems for transport of water and nutrients - Structural support tissues (wood) -Leaves and roots of various kinds - Seeds - Tree growth forms - Specialized reproductive structures Amphibians, the First Vertebrates to Colonize Land, Are Thought to Have Descended from Lobe-Finned Fishes Reptiles Evolved from Amphibians and Were the First to Evolve the Amniotic Egg, a Major Event in the History of Life • Mammals, the dominant vertebrate group on land currently, evolved from reptiles roughly 220 mya. The Effects of Plate Tectonics • • • • Continents “float” on Earth’s mantle, a hot layer of semisolid rock. The slow movement of the continents over time relative to one another is called plate tectonics. Plate tectonics can be caused by spreading of the sea floor or the collision and subsequent sinking of one plate into the mantle under the other plate. Patterns of plate tectonics have had dramatic effects on the history of life through its effects on climate and biogeographical isolation leading to speciation. Mass Extinctions: Worldwide Losses of Species • At several points in Earth’s history, the extinction rate has soared. • The fossil record shows that there have been five mass extinctions, periods of time during which great numbers of species went extinct throughout most of Earth. Each Mass Extinction Has Left a Permanent Mark on the History of Life Reconstruction of the head of a Mapusaurus. This dinosaur may have been the largest carnivore ever to walk the Earth. • The mass extinctions are thought to have been caused by such things as climate change, massive volcanic eruptions, and asteroid impacts. The Extinction of One or More Groups of Organisms Can Provide New Ecological and Evolutionary Opportunities for Other Groups of Organisms Allosaurus skeleton Effect of mass extinctions on the diversity of life: - First, entire groups of organisms perish, changing the history of life forever. - Second, mass extinctions enable surviving species to experience adaptive radiation, as the extinction of dominant groups of organisms provides new ecological and evolutionary opportunities for groups of organisms that previously were of relatively minor importance, dramatically altering the course of evolution. Rapid Macroevolution through Differential Gene Expression Altering how and when a homeotic gene is expressed in the developing embryo can radically change the body plan of an animal. • Dramatic macroevolutionary changes can occur over geological timescales because of its immense magnitude and also because geologic processes can alter the position of continents, cause shifts in atmospheric composition, and produce large-scale extinction events. • However, the relatively new field of evo-devo (evolutionary developmental biology) has demonstrated that changes in patterns of gene expression can also bring about dramatic changes in form. Macroevolution Does Not Necessarily Require Vast Timescales in Order to Produce Large Changes This legless lizard lacks legs because a homeotic gene for limb development is suppressed during its embryonic development. • A small alteration in the expression of a single homeotic gene can rapidly lead to a significant phenotypic change and introduce a completely novel vertebrate body plan. Differences in Patterns of Gene Expression Explain Many of the Differences between Chimps and Humans • Human and chimpanzee genomes are approximately 98 percent similar. • Humans have the same genes for hair growth that a chimpanzee has, but these genes are expressed only in certain areas of the body. • The same genes responsible for facial elongation during growth are found in both species, but they are expressed for a longer period of time in chimps than in humans, leading to their relatively longer faces. Atavistic Traits and Vestigial Traits Provide Evidence That Many Existing Body Plans Are the Product of Developmental Changes • An atavistic trait is one that represents a reversion to an ancestral state. • Being descended from organisms with tails, humans are sometimes born with tails because we possess the genes for tail development within our genomes. • Similarly, horses are occasionally born with three toes instead of one because ancestral horses had multiple toes. Similarities between Organisms Can Be Used to Infer Their Evolutionary Histories • Organizing and classifying the tremendous diversity of life on Earth is the focus of a discipline called taxonomy. • Scientists classify organisms into groups based on morphological or genetic similarities. • An evolutionary tree illustrates the evolutionary history of groups of organisms. • The goal of phylogenetics is to construct evolutionary trees that illustrate the patterns of species evolution. An Evolutionary Tree Maps the Relationship between Ancestral Groups and Their Descendants • Living things have diversified into many lines of descent, or lineages, that have evolved into many different types of organisms, some of them now extinct. • The node represents the most recent common ancestor of the two lineages in question—that is, the most immediate ancestor that both lineages share. • A given ancestor and all its descendants make up a clade, or branch, on an evolutionary tree. An evolutionary tree clusters the groups that are most closely related on neighboring branches. Shared Derived Traits Are Evolutionary Novelties That Reveal Relatedness among Species • The most useful characteristics for discerning evolutionary relationships are unique features that are found in a group’s most recent common ancestor. • Shared derived traits are evolutionary novelties shared by an ancestor and its descendants but not seen in groups that are not direct descendants of that ancestor. The Linnaean System of Biological Classification • The Linnaean hierarchy is a system of biological classification devised by a Swedish naturalist named Carolus Linnaeus. • The species is the smallest unit (lowest level) of classification in the Linnaean hierarchy. • Closely related species are grouped together to form a genus (plural, genera). • Using these two categories in the hierarchy, each type of organism is given a unique, two-word Latin name, called its scientific name. In the Linnaean Hierarchy, Each Species Is Placed in Successively Larger and More Inclusive Categories Beyond the Genus • Closely related genera are grouped together into a family. • Closely related families are grouped together into an order. • Closely related orders are grouped together into a class. • Closely related classes are grouped together into a phylum. • Closely related phylum are grouped together into a kingdom. BIOLOGY MATTERS: IS MASS EXTINCTION UNDER WAY? • The Red List produced by the International Union for Conservation of Nature (IUCN) identifies the world’s threatened species. To be defined as such, a species must face a high to extremely high risk of extinction in the wild. • About 30 percent of the species evaluated by the IUCN are threatened. • The IUCN evaluates only about 4 percent of the world’s 1.7 million described species. • Some experts believe that current species loss is approaching rates seen in some of the previous mass extinctions. The Iberian lynx (Lynx pardinus) is the most critically endangered feline in the world. About 100 individuals survive in small populations in Spain. Number of Species at Risk of Extinction in Some Major Groups of Organisms NOTE: Based on the IUCN Red List, 2013. MAMMALS BIRDS REPTILES AMPHIBIANS FISHES MOLLUSKS OTHER PLANTS TOTAL Canada 11 16 5 1 36 5 10 2 86 United States 36 78 37 56 236 301 265 270 1,279 APPLYING WHAT WE LEARNED: WHEN ANTARCTICA WAS GREEN • Antarctic fossils include Cambrian marine organisms, land plants, birds, and mammals, revealing that great changes have occurred over vast stretches of time. • About 40 million years ago, Antarctica broke away from the other continental masses and started drifting toward the South Pole. • • A circumpolar cold current contributed to the formation of the Antarctic ice cap, and the last of the alpine flowers died out, along with nearly all other life-forms, about 14 million years ago. The same continental movements that brought destruction to terrestrial life in Antarctica sowed the seeds of evolutionary diversity elsewhere. List of Key Terms: Chapter 16 atavistic trait (p. 370) Cambrian explosion (p. 364) class (p. 372) Cretaceous extinction (p. 368) domain (p. 372) evo-devo (p. 369) family (p. 372) genus (p. 372) geologic time (p. 359) homeotic gene (p. 369) kingdom (p. 372) lineage (p. 371) Linnaean hierarchy (p. 372) macroevolution (p. 359) mantle (p. 365) mass extinction (p. 366) most recent common ancestor (p. 371) node (p. 371) order (p. 372) Permian extinction (p. 366) phylogenetics (p. 371) phylum (p. 372) plate tectonics (p. 365) radioisotope (p. 359) scientific name (p. 372) shared derived trait (p. 372) species (p. 372) supercontinent (p. 365) taxon (p. 373) taxonomy (p. 371) The coelocanth, a living fossil Class Quiz, Part 1 What was the first significant source of the oxygen that began to accumulate in Earth’s atmosphere? A. Anaerobic bacteria B. Photosynthesizing prokaryotes C. Minerals in Earth’s mantle D. Land plants Class Quiz, Part 2 Which of the following were present on Earth 3.0 billion years ago? A. B. C. D. Prokaryotes Eukaryotes Multicellular organisms Mammals Class Quiz, Part 3 We can infer from this evolutionary tree that A. black bears do not belong on the clade that includes the walrus, the elephant seal, and the giant panda. B. walruses are the most advanced among these mammals. C. the most recent common ancestor of the gray wolf and domestic dogs evolved more recently than the most recent common ancestor of the coyote and the gray wolf. Class Quiz, Part 4 The figure shows the Linnaean hierarchy for the Chinese rose, Rosa chinensis. Which of the statements that follow is true? A. The genus that this rose belongs to is “chinensis.” B. All roses belong to the genus Rosaceae. C. The order in which roses are placed is “Rosa.” D. The order in which roses are placed is “Rosales.” The Linnaean Hierarchy for Rosa Chinensis Relevant Art from Other Chapters All art files from the book are available in JPEG and PPT formats online Milestones in the History of Life on Earth, Part 1 Milestones in the History of Life on Earth, Part 2 Milestones in the History of Life on Earth, Part 3 16.1 Concept Check, Part 1 1. What is macroevolution? ANSWER: Macroevolution is large-scale evolution in which the effect of natural selection is seen as radical changes in body plan, not simply a change in allele frequencies. 16.1 Concept Check, Part 2 2. Why is the concept of geologic time so important to our understanding of macroevolution? ANSWER: A short timescale would make it difficult to argue convincingly that natural selection could shape the entire biodiversity seen on Earth. 16.2 Concept Check, Part 1 1. What is a fossil? ANSWER: A fossil is the preserved traces of an organism. Fossils may include petrified bone, impressions cast in stone, or any other remnant of a once-living organism. 16.2 Concept Check, Part 2 2. Why is the fossil record so incomplete? ANSWER: The conditions for fossil preservation are relatively rare. Some organisms are more likely to be fossilized than others (for example, bony versus softbodied animals). 16.3 Concept Check, Part 1 1. Why was it significant that early bacteria evolved the ability to carry out photosynthesis? ANSWER: Oxygen is a by-product of photosynthesis, and increasing oxygen in the atmosphere set the stage for the evolution of eukaryotes, a group that includes all multicellular life. 16.3 Concept Check, Part 2 2. How did the Cambrian explosion change life on Earth? ANSWER: The Cambrian explosion marks the appearance of most of the major living animal phyla, many of which were the world’s first predators. 16.4 Concept Check, Part 1 1. Why is the age of Earth a key factor in discussions of plate tectonics? ANSWER: Because the continents move so slowly, the effects of plate tectonics are apparent only over long timescales 16.4 Concept Check, Part 2 2. What are two ways that plate tectonics can affect biodiversity? ANSWER: The movement of continents can geographically isolate populations, leading to allopatric speciation (see Section 15.2). It can also alter climate by changing ocean currents. 16.5 Concept Check, Part 1 1. Given the long history of Earth, why have so few mass extinctions been recorded? ANSWER: The causes of mass extinctions are global in scale and catastrophic in magnitude. Such events are inherently rare. 16.5 Concept Check, Part 2 2. What is the impact of mass extinction events on biodiversity? ANSWER: Many species are lost in mass extinction events, but their loss opens up ecological niches permitting the adaptive radiation of successor species. 16.6 Concept Check, Part 1 1. What is a homeotic gene? ANSWER: A homeotic gene controls the embryological development of a body structure. 16.6 Concept Check, Part 2 2. Explain why the discovery of homeotic genes had a significant impact on evolutionary thought. ANSWER: Prior to the discovery of homeotic genes, macroevolutionary change was thought to occur solely over long periods of time. Changes in homeotic gene expression, however, have shown that significant alterations in body plans can occur in a single generation. 16.7 Concept Check, Part 1 1. In evolutionary terms, what happens at a node? ANSWER: A node represents a speciation event in which one lineage splits into two. 16.7 Concept Check, Part 2 2. List the eight taxonomic tiers of Linnaean classification, beginning with the most inclusive. ANSWER: Domain, kingdom, phylum, class, order, family, genus, species