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How Biological Diversity Evolves MACROEVOLUTION Biology and Society: The Sixth Mass Extinction Over the past 600 million years the fossil record reveals five periods of extinction when 50–90% of living species suddenly died out. © 2010 Pearson Education, Inc. Figure 14.00 Our current rate of extinction, over the past 400 years, indicates that we may be living in, and contributing to, the sixth mass extinction period. Mass extinctions: Pave the way for the evolution of new and diverse forms, but Take millions of years for Earth to recover The most “famous” mass-extinction is the K-T Mass Extinction which included the extinction of the dinosaurs (except birds). This will be discussed in Chapter 17. MACROEVOLUTION AND THE DIVERSITY OF LIFE Macroevolution: Encompasses the major biological changes evident in the fossil record Includes the formation of new species THE ORIGIN OF SPECIES Species is a Latin word meaning: “Kind” or “Appearance.” What Is a Species? The biological species concept defines a species as “A group of populations whose members have the potential to interbreed and produce fertile offspring” It is also sometimes stated like this: “A group of populations whose members share a common gene pool and are potentially interbreeding” Speciation: Is the focal point of macroevolution May occur based on two contrasting patterns, non-branching and branching. In non-branching evolution: A population transforms but does not create a new species Actually, it may, but the hypothesis is usually untestable. In branching evolution, one or more new species branch from a parent species that may: Continue to exist in much the same form or Change considerably Similarity between different species Diversity within one species Figure 14.2 The biological species concept cannot be applied in all situations, including: Fossils Asexual organisms Reproductive Barriers/Isolating Mechanisms between Species What’s the logic of isolating mechanisms/reproductive barriers? ? Reproductive Barriers between Species Prezygotic barriers prevent mating or fertilization between species. Below are some examples you can see at MasteringBiology Video: Albatross Courtship Ritual Video: Blue-footed Boobies Courtship Ritual Video: Giraffe Courtship Ritual INDIVIDUALS OF DIFFERENT SPECIES Prezygotic Barriers Temporal isolation Habitat isolation Behavioral isolation MATING ATTEMPT Mechanical isolation Gametic isolation FERTILIZATION (ZYGOTE FORMS) Postzygotic Barriers Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown VIABLE, FERTILE OFFSPRING Figure 14.3 PREZYGOTIC BARRIERS Temporal Isolation Behavioral Isolation Mechanical Isolation Habitat Isolation Gametic Isolation Figure 14.4 Postzygotic barriers operate if: Interspecies mating occurs and Hybrid zygotes form INDIVIDUALS OF DIFFERENT SPECIES Prezygotic Barriers Temporal isolation Habitat isolation Behavioral isolation MATING ATTEMPT Mechanical isolation Gametic isolation FERTILIZATION (ZYGOTE FORMS) Postzygotic Barriers Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown VIABLE, FERTILE OFFSPRING Figure 14.3 Postzygotic barriers include: Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown POSTZYGOTIC BARRIERS Reduced Hybrid Viability Reduced Hybrid Fertility Horse Donkey Mule Figure 14.5 Hybrid Breakdown Mechanisms of Speciation A key event in the potential origin of a species occurs when a population is severed from other populations of the parent species. Species can form by: Allopatric speciation, due to geographic isolation Sympatric speciation, without geographic isolation Allopatric speciation Simpatric speciation Figure 14.6 Allopatric Speciation Geologic processes can: Fragment a population into two or more isolated populations Contribute to allopatric speciation Video: Lava Flow Video: Volcanic Eruption Video: Grand Canyon Ammospermophilus harrisii Ammospermophilus leucurus Figure 14.7 Speciation occurs only with the evolution of reproductive barriers between the isolated population and its parent population. Populations become allopatric Populations become sympatric Populations interbreed Gene pools merge: No speciation Geographic barrier Populations cannot interbreed Reproductive isolation: Speciation has occurred Time Figure 14.8 Sympatric Speciation Sympatric speciation occurs: While the new species and old species live in the same time and place If a genetic change produces a reproductive barrier between the new and old species Polyploids can: Originate from accidents during cell division Result from the hybridization of two parent species Many domesticated plants are the result of sympatric speciation, including: Oats Potatoes Bananas Peanuts Apples Coffee Wheat Domesticated Triticum monococcum (14 chromosomes) BB AA Wild Triticum (14 chromosomes) AB Sterile hybrid (14 chromosomes) T. turgidum Emmer wheat (28 chromosomes) AA BB DD Wild T. tauschii (14 chromosomes) ABD Sterile hybrid (21 chromosomes) AA BB DD T. aestivum Bread wheat (42 chromosomes) Figure 14.9-4 Figure 14.9a What Is the Tempo of Speciation? There are two contrasting models of the pace of evolution: The gradual model, in which big changes (speciations) occur by the steady accumulation of many small changes The punctuated equilibria model, in which there are Long periods of little change, equilibrium, punctuated by Abrupt episodes of speciation Punctuated model Time Graduated model Figure 14.10 THE EVOLUTION OF BIOLOGICAL NOVELTY What accounts for the evolution of biological novelty? Adaptation of Old Structures for New Functions Birds: Are derived from a lineage of earthbound reptiles Evolved flight from flightless ancestors Wing claw (like reptile) Teeth (like reptile) Feathers Long tail with many vertebrae (like reptile) Fossil Artist’s reconstruction Figure 14.11 An exaptation: Is a structure that evolves in one context, but becomes adapted for another function Is a type of evolutionary remodeling Exaptations can account for the gradual evolution of novel structures. Bird wings are modified forelimbs that were previously adapted for non-flight functions, such as: Thermal regulation Courtship displays Camouflage The first flights may have been only glides or extended hops as the animal pursued prey or fled from a predator. Evo-Devo: Development and Evolutionary Novelty A subtle change in a species’ developmental program can have profound effects, changing the: Rate Timing Spatial pattern of development © 2010 Pearson Education, Inc. Evo-devo, evolutionary developmental biology, is the study of the evolution of developmental processes in multicellular organisms. Paedomorphosis: Is the retention into adulthood of features that were solely juvenile in ancestral species Has occurred in the evolution of Axolotl salamanders Humans Animation: Allometric Growth Gills Figure 14.12 Chimpanzee fetus Chimpanzee adult Human fetus Human adult (paedomorphic features) Figure 14.13 Homeotic genes are master control genes that regulate: When structures develop How structures develop Where structures develop Mutations in homeotic genes can profoundly affect body form. EARTH HISTORY AND MACROEVOLUTION Macroevolution is closely tied to the history of the Earth. Geologic Time and the Fossil Record The fossil record is: The sequence in which fossils appear in rock strata An archive of macroevolution Figure 14.14 Figure 14.14a Figure 14.14b Figure 14.14c Figure 14.14d Figure 14.14e Geologists have established a geologic time scale reflecting a consistent sequence of geologic periods. Animation: The Geologic Record Animation: Macroevolution Table 14.1 Table 14.1a Table 14.1b Table 14.1c Table 14.1d Fossils are reliable chronological records only if we can determine their ages, using: The relative age of fossils, revealing the sequence in which groups of species evolved, or The absolute age of fossils, requiring other methods such as radiometric dating Radiometric dating: Is the most common method for dating fossils Is based on the decay of radioactive isotopes Helped establish the geologic time scale Carbon-14 radioactivity (as % of living organism’s C-14 to C-12 ratio) Radioactive decay of carbon-14 100 75 50 25 0 0 5.6 11.2 16.8 22.4 28.0 33.6 39.2 44.8 50.4 Time (thousands of years) How carbon-14 dating is used to determine the vintage of a fossilized clam shell Carbon-14 in shell Figure 14.15 Plate Tectonics and Macroevolution The continents are not locked in place. Continents drift about the Earth’s surface on plates of crust floating on a flexible layer called the mantle. The San Andreas fault is: In California At a border where two plates slide past each other Figure 14.16 About 250 million years ago: Plate movements formed the supercontinent Pangaea The total amount of shoreline was reduced Sea levels dropped The dry continental interior increased in size Many extinctions occurred Cenozoic Present 65 Eurasia Africa India South America Madagascar Mesozoic Laurasia Paleozoic 251 million years ago 135 Antarctica Figure 14.17 About 180 million years ago: Pangaea began to break up Large continents drifted increasingly apart Climates changed The organisms of the different biogeographic realms diverged Plate tectonics explains: Why Mesozoic reptiles in Ghana (West Africa) and Brazil look so similar How marsupials were free to evolve in isolation in Australia Mass Extinctions and Explosive Diversifications of Life The fossil record reveals that five mass extinctions have occurred over the last 600 million years. The Permian mass extinction: Occurred at about the time the merging continents formed Pangaea (250 million years ago) Claimed about 96% of marine species © 2010 Pearson Education, Inc. The Cretaceous extinction: Occurred at the end of the Cretaceous period, about 65 million years ago Included the extinction of all the dinosaurs except birds Permitted the rise of mammals The Process of Science: Did a Meteor Kill the Dinosaurs? Observation: About 65 million years ago, the fossil record shows that: The climate cooled Seas were receding Many plant species died out Dinosaurs (except birds) became extinct A thin layer of clay rich in iridium was deposited © 2010 Pearson Education, Inc. Question: Is the iridium layer the result of fallout from a huge cloud of dust that billowed into the atmosphere when a large meteor or asteroid hit Earth? Hypothesis: The mass extinction 65 million years ago was caused by the impact of an extraterrestrial object. Prediction: A huge impact crater of the right age should be found somewhere on Earth’s surface. Results: Near the Yucatán Peninsula, a huge impact crater was found that: Dated from the predicted time Was about the right size Was capable of creating a cloud that could have blocked enough sunlight to change the Earth’s climate for months Figure 14.18-1 Figure 14.18-2 Chicxulub crater Figure 14.18-3 CLASSIFYING THE DIVERSITY OF LIFE Systematics focuses on: Classifying organisms Determining their evolutionary relationships Taxonomy is the: Identification of species Naming of species Classification of species Some Basics of Taxonomy Scientific names ease communication by: Unambiguously identifying organisms Making it easier to recognize the discovery of a new species Carolus Linnaeus (1707–1778) proposed the current taxonomic system based upon: A two-part name for each species (binomial nomenclature) A hierarchical classification of species into broader groups of organisms Naming Species Each species is assigned a two-part name or binomial, consisting of: The genus A name unique for each species The scientific name for humans is Homo sapiens, a two part name, italicized and latinized, and with the first letter of the genus capitalized. Hierarchical Classification Species that are closely related are placed into the same genus. Leopard (Panthera pardus) Tiger (Panthera tigris) Lion (Panthera leo) Jaguar (Panthera onca) Figure 14.19 The taxonomic hierarchy extends to progressively broader categories of classification, from genus to: Family Order Class Phylum Kingdom Domain Species Panthera pardus Genus Panthera Leopard (Panthera pardus) Family Felidae Order Carnivora Class Mammalia Phylum Chordata Kingdom Animalia Domain Eukarya Figure 14.20 Classification and Phylogeny The goal of systematics is to reflect evolutionary relationships. Biologists use phylogenetic trees to: Depict hypotheses about the evolutionary history of species Reflect the hierarchical classification of groups nested within more inclusive groups Order Family Felidae Genus Species Panthera Panthera pardus (leopard) Mephitis Mephitis mephitis (striped skunk) Carnivora Mustelidae Lutra Lutra lutra (European otter) Canis latrans (coyote) Canidae Canis Canis lupus (wolf) Figure 14.21 Sorting Homology from Analogy Homologous structures: Reflect variations of a common ancestral plan Are the best sources of information used to Develop phylogenetic trees Classify organisms according to their evolutionary history Convergent evolution: Involves superficially similar structures in unrelated organisms Is based on natural selection Similarity due to convergence: Is called analogy, not homology Can obscure homologies Systematics Molecular systematics: Compares DNA and amino acid sequences between organisms Can reveal evolutionary relationships Some fossils are preserved in such a way that DNA fragments can be extracted for comparison with living organisms. Figure 14.22 The Cladistic Revolution Cladistics is the scientific search for clades. A clade: Consists of an ancestral species and all its descendants Forms a distinct branch in the tree of life Iguana Outgroup (reptile) Duck-billed platypus Hair, mammary glands Gestation Kangaroo Ingroup (mammals) Beaver Long gestation Figure 14.23 Cladistics has changed the traditional classification of some organisms, including the relationships between: Dinosaurs Birds Crocodiles Lizards Snakes Lizards and snakes Crocodilians Pterosaurs Common ancestor of crocodilians, dinosaurs, and birds Ornithischian dinosaurs Saurischian dinosaurs Birds Figure 14.24 Classification: A Work in Progress Linnaeus: Divided all known forms of life between the plant and animal kingdoms Prevailed with his two-kingdom system for over 200 years In the mid-1900s, the two-kingdom system was replaced by a five-kingdom system that: Placed all prokaryotes in one kingdom Divided the eukaryotes among four other kingdoms In the late 20th century, molecular studies and cladistics led to the development of a three-domain system, recognizing: Two domains of prokaryotes (Bacteria and Archaea) One domain of eukaryotes (Eukarya) Animation: Classification Schemes Domain Bacteria Earliest organisms Domain Archaea The protists (multiple kingdoms) Kingdom Plantae Domain Eukarya Kingdom Fungi Kingdom Animalia Figure 14.25 Evolution Connection: Rise of the Mammals Mass extinctions: Have repeatedly occurred throughout Earth’s history Were followed by a period of great evolutionary change © 2010 Pearson Education, Inc. Fossil evidence indicates that: Mammals first appeared about 180 million years ago The number of mammalian species Remained steady and low in number until about 65 million years ago and then Greatly increased after most of the dinosaurs became extinct Ancestral mammal Extinction of dinosaurs Reptilian ancestor Monotremes (5 species) Marsupials (324 species) Eutherians (5,010 species) 250 65 50 Millions of years ago 200 150 100 0 American black bear Figure 14.26 Bacteria Earliest organisms Archaea Eukarya Figure 14.UN3