1. CAMPBELL BIOLOGY IN FOCUS © 2014 Pearson Education, Inc. Urry Cain Wasserman Minorsky Jackson Reece Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge 19 Descent with Modification

2. © 2014 Pearson Education, Inc. Overview: Endless Forms Most Beautiful  Lepidopteran insects (moths and butterflies) have many features in common including a juvenile feeding stage called a caterpillar  Lepidopteran species also have many features that are distinct from each other in both the caterpillar and adult forms

3. © 2014 Pearson Education, Inc.  Lepidopterans illustrate three key observations about life  The fit between organisms and their environment  The shared characteristics (unity) of life  The diversity of life

4. © 2014 Pearson Education, Inc. Figure 19.1

5. © 2014 Pearson Education, Inc.  A new era of biology began in 1859 when Charles Darwin published The Origin of Species  The Origin of Species focused biologists’ attention on the great diversity of organisms

6. © 2014 Pearson Education, Inc.  Darwin noted that current species are descendants of ancestral species  Evolution can be defined by Darwin’s phrase descent with modification  Evolution can be viewed as both a pattern and a process

7. © 2014 Pearson Education, Inc.  Darwin’s revolutionary ideas had deep historical roots Concept 19.1: The Darwinian revolution challenged traditional views of a young Earth inhabited by unchanging species

8. © 2014 Pearson Education, Inc. Figure 19.2

9. © 2014 Pearson Education, Inc. Scala Naturae and Classification of Species  The Greek philosopher Aristotle viewed species as fixed and arranged them on a scala naturae  The Old Testament holds that species were individually designed by God and therefore perfect

10. © 2014 Pearson Education, Inc.  Carolus Linnaeus interpreted organismal adaptations as evidence that the Creator had designed each species for a particular purpose  Linnaeus was the founder of taxonomy, the branch of biology concerned with classifying organisms  He developed the binomial format for naming species (for example, Homo sapiens)

11. © 2014 Pearson Education, Inc. Ideas About Change over Time  The study of fossils helped to lay the groundwork for Darwin’s ideas  Fossils are remains or traces of organisms from the past, usually found in sedimentary rock, which appears in layers or strata

12. Video: Grand Canyon

13. © 2014 Pearson Education, Inc. Figure 19.3 Younger stratum with more recent fossils Older stratum with older fossils

14. © 2014 Pearson Education, Inc.  Paleontology, the study of fossils, was largely developed by French scientist Georges Cuvier  Cuvier speculated that each boundary between strata represents a catastrophe that destroyed many species

15. © 2014 Pearson Education, Inc.  Geologists James Hutton and Charles Lyell perceived that changes in Earth’s surface can result from slow, continuous actions still operating today  Lyell further proposed that the mechanisms of change are constant over time  This view strongly influenced Darwin’s thinking

16. © 2014 Pearson Education, Inc. Lamarck’s Hypothesis of Evolution  Lamarck hypothesized that species evolve through use and disuse of body parts and the inheritance of acquired characteristics  The mechanisms he proposed are unsupported by evidence

17. © 2014 Pearson Education, Inc. Figure 19.4

18. © 2014 Pearson Education, Inc.  Some doubt about the permanence of species preceded Darwin’s ideas Concept 19.2: Descent with modification by natural selection explains the adaptations of organisms and the unity and diversity of life

19. © 2014 Pearson Education, Inc. Darwin’s Research  As a boy and into adulthood, Charles Darwin had a consuming interest in nature  Darwin first studied medicine (unsuccessfully) and then theology at Cambridge University  After graduating, he took an unpaid position as naturalist and companion to Captain Robert FitzRoy for a five-year around-the-world voyage on the Beagle

20. © 2014 Pearson Education, Inc. The Voyage of the Beagle  During his travels on the Beagle, Darwin collected specimens of South American plants and animals  He observed that fossils resembled living species from the same region, and living species resembled other species from nearby regions  He experienced an earthquake in Chile and observed the uplift of rocks

21. © 2014 Pearson Education, Inc.  Darwin was influenced by Lyell’s Principles of Geology and thought that Earth was more than 6,000 years old  His interest in geographic distribution of species was kindled by a stop at the Galápagos Islands west of South America  He hypothesized that species from South America had colonized the Galápagos and speciated on the islands

22. Video: Albatross Courtship

23. Video: Boobies Courtship

24. Video: Galápagos Islands

25. Video: Marine Iguana

26. Video: Sea Lion

27. Video: Soaring Hawk

28. Video: Tortoise

29. © 2014 Pearson Education, Inc. Figure 19.5 HMS Beagle at sea Darwin in 1840, after his return from the voyage NORTH AMERICA Malay Archipelago SOUTH AMERICA Great Britain ATLANTIC OCEAN PACIFIC OCEAN EUROPE AFRICA AUSTRALIA PACIFIC OCEAN Cape of Good Hope Equator Tasmania New Zealand Brazil Argentina Cape Horn Chile Andes Mtns. Genovesa PACIFIC OCEAN Equator Pinta Marchena Fernandina Florenza Pinzón Santiago Daphne Islands Santa Cruz Santa Fe Isabela Española San Cristobal Kilometers 040 20 The Galápagos Islands

30. © 2014 Pearson Education, Inc. Figure 19.5a NORTH AMERICA Malay Archipelago SOUTH AMERICA Great Britain ATLANTIC OCEAN PACIFIC OCEAN EUROPE AFRICA AUSTRALIA PACIFIC OCEAN Cape of Good Hope Equator Tasmania New Zealand Brazil Argentina Cape Horn Chile Andes Mtns.

31. © 2014 Pearson Education, Inc. Figure 19.5b Darwin in 1840, after his return from the voyage

32. © 2014 Pearson Education, Inc. Figure 19.5c Genovesa PACIFIC OCEAN Equator Pinta Marchena Fernandina Florenza Pinzón Santiago Daphne Islands Santa Cruz Santa Fe Isabela Española San Cristobal Kilometers 0 40 20 The Galápagos Islands

33. © 2014 Pearson Education, Inc. Figure 19.5d HMS Beagle at sea

34. © 2014 Pearson Education, Inc. Darwin’s Focus on Adaptation  In reassessing his observations, Darwin perceived adaptation to the environment and the origin of new species as closely related processes  From studies made years after Darwin’s voyage, biologists have concluded that this is what happened to the Galápagos finches

35. © 2014 Pearson Education, Inc. Figure 19.6 (b) Seed-eater (a) Cactus-eater(c) Insect-eater

36. © 2014 Pearson Education, Inc. Figure 19.6a (a) Cactus-eater

37. © 2014 Pearson Education, Inc. Figure 19.6b (b) Seed-eater

38. © 2014 Pearson Education, Inc. Figure 19.6c (c) Insect-eater

39. © 2014 Pearson Education, Inc.  In 1844, Darwin wrote an essay on natural selection as the mechanism of descent with modification but did not introduce his theory publicly  Natural selection is a process in which individuals with favorable inherited traits are more likely to survive and reproduce  In June 1858, Darwin received a manuscript from Alfred Russell Wallace, who had developed a theory of natural selection similar to Darwin’s  Darwin quickly finished The Origin of Species and published it the next year

40. © 2014 Pearson Education, Inc. Figure 19.7

41. © 2014 Pearson Education, Inc. Figure 19.7a

42. © 2014 Pearson Education, Inc. Figure 19.7b

43. © 2014 Pearson Education, Inc. Ideas from The Origin of Species  Darwin explained three broad observations about life  The unity of life  The diversity of life  The match between organisms and their environment

44. © 2014 Pearson Education, Inc. Descent with Modification  Darwin never used the word evolution in the first edition of The Origin of Species  The phrase descent with modification summarized Darwin’s perception of the unity of life  The phrase refers to the view that all organisms are related through descent from an ancestor that lived in the remote past

45. © 2014 Pearson Education, Inc.  In the Darwinian view, the history of life is like a tree with branches representing life’s diversity  Fossils of extinct species help to “fill in” the morphological gaps between present-day groups

46. © 2014 Pearson Education, Inc. Figure 19.8

47. © 2014 Pearson Education, Inc. Figure 19.9 Moeritherium † † Barytherium Hyracoidea (Hyraxes) Sirenia (Manatees and relatives) † Deinotherium † Mammut † Platybelodon † Stegodon Elephas maximus (Asia) Mammuthus † Loxodonta cyclotis (Africa) Loxodonta africana (Africa) Millions of years ago Years ago 6034 245.52 10 4 0

48. © 2014 Pearson Education, Inc. Figure 19.9a Moeritherium † † Barytherium Hyracoidea (Hyraxes) Sirenia (Manatees and relatives) † Deinotherium † Mammut

49. © 2014 Pearson Education, Inc. Figure 19.9b † Platybelodon † Stegodon Elephas maximus (Asia) Mammuthus † Loxodonta cyclotis (Africa) Loxodonta africana (Africa) Millions of years agoYears ago 6034 245.52 10 4 0

50. © 2014 Pearson Education, Inc. Artificial Selection, Natural Selection, and Adaptation  Darwin noted that humans have modified other species by selecting and breeding individuals with desired traits, a process called artificial selection  Darwin argued that a similar process occurs in nature

51. © 2014 Pearson Education, Inc. Figure 19.10 Wild mustard Kohlrabi Cabbage Kale Broccoli Selection for leaves Selection for stems Selection for flowers and stems Selection for axillary (side) buds Selection for apical (tip) bud Brussels sprouts

52. © 2014 Pearson Education, Inc. Figure 19.10a Wild mustard

53. © 2014 Pearson Education, Inc. Figure 19.10b Kale

54. © 2014 Pearson Education, Inc. Figure 19.10c Brussels sprouts

55. © 2014 Pearson Education, Inc. Figure 19.10d Cabbage

56. © 2014 Pearson Education, Inc. Figure 19.10e Broccoli

57. © 2014 Pearson Education, Inc. Figure 19.10f Kohlrabi

58. © 2014 Pearson Education, Inc.  Darwin drew two inferences from two observations  Observation #1: Members of a population often vary in their inherited traits

59. © 2014 Pearson Education, Inc. Figure 19.11

60. © 2014 Pearson Education, Inc.  Observation #2: All species can produce more offspring than the environment can support, and many of these offspring fail to survive and reproduce

61. Video: Sea Horses

62. © 2014 Pearson Education, Inc. Figure 19.12 Spore cloud

63. © 2014 Pearson Education, Inc.  Inference #1: Individuals whose inherited traits give them a higher probability of surviving and reproducing in a given environment tend to leave more offspring than other individuals

64. © 2014 Pearson Education, Inc.  Inference #2: This unequal ability of individuals to survive and reproduce will lead to the accumulation of favorable traits in the population over generations

65. © 2014 Pearson Education, Inc.  Darwin was influenced by Thomas Malthus, who noted the potential for human population to increase faster than food supplies and other resources  If some heritable traits are advantageous, these will accumulate in a population over time, and this will increase the frequency of individuals with these traits  This process explains the match between organisms and their environment

66. © 2014 Pearson Education, Inc. Natural Selection: A Summary  Individuals with certain heritable traits survive and reproduce at a higher rate than other individuals  Over time, natural selection increases the match between organisms and their environment  If an environment changes over time, natural selection may result in adaptation to these new conditions and may give rise to new species

67. © 2014 Pearson Education, Inc. Figure 19.13 (a) A flower mantid in Malaysia (b) A leaf mantid in Borneo

68. © 2014 Pearson Education, Inc. Figure 19.13a (a) A flower mantid in Malaysia

69. © 2014 Pearson Education, Inc. Figure 19.13b (b) A leaf mantid in Borneo

70. © 2014 Pearson Education, Inc.  Note that individuals do not evolve; populations evolve over time  Natural selection can only increase or decrease heritable traits that vary in a population  Adaptations vary with different environments

71. © 2014 Pearson Education, Inc. Concept 19.3: Evolution is supported by an overwhelming amount of scientific evidence  New discoveries continue to fill the gaps identified by Darwin in The Origin of Species  There are four types of data that document the pattern of evolution  Direct observations  Homology  The fossil record  Biogeography

72. © 2014 Pearson Education, Inc. Direct Observations of Evolutionary Change  Two examples provide evidence for natural selection: natural selection in response to introduced plant species and the evolution of drug-resistant bacteria

73. © 2014 Pearson Education, Inc. Natural Selection in Response to Introduced Plant Species  Soapberry bugs use their “beak” to feed on seeds within fruits  In southern Florida soapberry bugs feed on balloon vine with larger fruit; they have longer beaks  In central Florida they feed on goldenrain tree with smaller fruit; they have shorter beaks  Correlation between fruit size and beak size has also been observed in Louisiana, Oklahoma, and Australia

74. © 2014 Pearson Education, Inc.  In all cases, beak size has evolved in populations that feed on introduced plants with fruits that are smaller or larger than the native fruits  These cases are examples of evolution by natural selection  In Florida this evolution in beak size occurred in less than 35 years

75. © 2014 Pearson Education, Inc. Figure 19.14 Soapberry bug with beak inserted in balloon vine fruit Beak length (mm) On native species, balloon vine (southern Florida) On introduced species, goldenrain tree (central Florida) Museum-specimen average Field Study Results Beak Number of individuals 6 78 9 1110 0 2 4 6 8 0 2 4 6 8

76. © 2014 Pearson Education, Inc. Figure 19.14a Soapberry bug with beak inserted in balloon vine fruit Field Study

77. © 2014 Pearson Education, Inc. Figure 19.14b Beak length (mm) On native species, balloon vine (southern Florida) On introduced species, goldenrain tree (central Florida) Museum-specimen average Results Beak Number of individuals 6 78 9 1110 0 2 4 6 8 0 2 4 6 8

78. © 2014 Pearson Education, Inc. Figure 19.14c Beak length (mm) On native species, balloon vine (southern Florida) On introduced species, goldenrain tree (central Florida) Museum-specimen average Results Number of individuals 6 78 9 1110 0 2 4 6 8 0 2 4 6 8

79. © 2014 Pearson Education, Inc. The Evolution of Drug-Resistant Bacteria  The bacterium Staphylococcus aureus is commonly found on people’s skin or in their nasal passages  Methicillin-resistant S. aureus (MRSA) strains are dangerous pathogens  S. aureus became resistant to penicillin in 1945, two years after it was first widely used  S. aureus became resistant to methicillin in 1961, two years after it was first widely used

80. © 2014 Pearson Education, Inc.  Methicillin works by inhibiting a protein used by bacteria in their cell walls  MRSA bacteria use a different protein in their cell walls  When exposed to methicillin, MRSA strains are more likely to survive and reproduce than nonresistant S. aureus strains  MRSA strains are now resistant to many antibiotics

81. © 2014 Pearson Education, Inc. Figure 19.15 Year Chromosome map of S. aureus clone USA300 Annual hospital admissions with MRSA (thousands) ’05 400 ’93 ’94’95 ’96 ’97’98’99 ’00 ’01 ’02 ’03’04 350 300 200 250 100 150 50 0 Increased gene exchange (within species) and toxin production Increased disease severity Ability to colonize hosts Methicillin resistance Key to adaptations 250,000 base pairs 500,000 750,000 2,000,000 1,750,000 1,500,000 1,250,000 1,000,000 2,250,000 2,500,000 2,750,000 1

82. © 2014 Pearson Education, Inc. Figure 19.15a Chromosome map of S. aureus clone USA300 Increased gene exchange (within species) and toxin production Increased disease severity Ability to colonize hosts Methicillin resistance Key to adaptations 250,000 base pairs 500,000 2,000,000 1,750,000 1,500,000 1,250,000 1,000,000 2,250,000 2,500,000 2,750,000 1 750,000

83. © 2014 Pearson Education, Inc. Figure 19.15b Year Annual hospital admissions with MRSA (thousands) ’05 400 ’93 ’94’95 ’96 ’97’98’99 ’00 ’01 ’02 ’03’04 350 300 200 250 100 150 50 0

84. © 2014 Pearson Education, Inc.  Natural selection does not create new traits, but edits or selects for traits already present in the population  The local environment determines which traits will be selected for or selected against in any specific population

85. © 2014 Pearson Education, Inc. Homology  Evolution is a process of descent with modification  Related species can have characteristics with underlying similarity that function differently  Homology is similarity resulting from common ancestry

86. © 2014 Pearson Education, Inc. Anatomical and Molecular Homologies  Homologous structures are anatomical resemblances that represent variations on a structural theme present in a common ancestor

87. © 2014 Pearson Education, Inc. Figure 19.16 Humerus Carpals Radius Human Ulna Metacarpals Phalanges WhaleCatBat

88. © 2014 Pearson Education, Inc.  Comparative embryology reveals anatomical homologies not visible in adult organisms  Vestigial structures are remnants of features that served important functions in the organism’s ancestors

89. © 2014 Pearson Education, Inc. Figure 19.17 Human embryo Chick embryo (LM) Pharyngeal arches Post-anal tail

90. © 2014 Pearson Education, Inc. Figure 19.17a Chick embryo (LM) Pharyngeal arches Post-anal tail

91. © 2014 Pearson Education, Inc. Figure 19.17b Human embryo Pharyngeal arches Post-anal tail

92. © 2014 Pearson Education, Inc.  Examples of homologies at the molecular level are genes shared among organisms inherited from a common ancestor  Homologous genes can be found in organisms as dissimilar as humans and bacteria

93. © 2014 Pearson Education, Inc. A Different Cause of Resemblance: Convergent Evolution  Convergent evolution is the evolution of similar, or analogous, features in distantly related groups  Analogous traits arise when groups independently adapt to similar environments in similar ways  Convergent evolution does not provide information about ancestry

94. © 2014 Pearson Education, Inc. Figure 19.18 AUSTRALIA NORTH AMERICA Sugar glider Flying squirrel

95. © 2014 Pearson Education, Inc. The Fossil Record  The fossil record provides evidence of  The extinction of species  The origin of new groups  Changes within groups over time

96. © 2014 Pearson Education, Inc. Figure 19.19 Cetaceans and even-toes ungulates Most mammals (d) Odocoileus (deer) (c) Sus (pig)(a) Canis (dog) (b) Pakicetus

97. © 2014 Pearson Education, Inc. Figure 19.19a (a) Canis (dog)

98. © 2014 Pearson Education, Inc. Figure 19.19b (b) Pakicetus

99. © 2014 Pearson Education, Inc. Figure 19.19c (c) Sus (pig)

100. © 2014 Pearson Education, Inc. Figure 19.19d (d) Odocoileus (deer)

101. © 2014 Pearson Education, Inc.  Fossils can document important transitions  For example, the transition from land to sea in the ancestors of cetaceans

102. © 2014 Pearson Education, Inc. Figure 19.20 Hippopotamuses Common ancestor of cetaceans Other even-toed ungulates Living cetaceans Millions of years ago Tibia Foot 70 Key Femur Pelvis 60 50 40 30 2010 0 Pakicetus † Rodhocetus † Dorudon †

103. © 2014 Pearson Education, Inc. Figure 19.20a Common ancestor of cetaceans Living cetaceans Millions of years ago Tibia Foot Key Femur Pelvis 50 40 30 2010 0 Pakicetus † Rodhocetus † Dorudon †

104. © 2014 Pearson Education, Inc. Figure 19.UN01 Diacodexis, an early even-toed ungulate 20 cm

105. © 2014 Pearson Education, Inc. Biogeography  Biogeography, the geographic distribution of species, provides evidence of evolution  Earth’s continents were formerly united in a single large continent called Pangaea but have since separated by continental drift  An understanding of continent movement and modern distribution of species allows us to predict when and where different groups evolved

106. © 2014 Pearson Education, Inc.  Endemic species are species that are not found anywhere else in the world  Islands have many endemic species that are often closely related to species on the nearest mainland or island  Darwin explained that species on islands gave rise to new species as they adapted to new environments

107. © 2014 Pearson Education, Inc. What Is Theoretical About Darwin’s View of Life?  In science, a theory accounts for many observations and explains and integrates a great variety of phenomena  The predictions of a scientific theory must stand up to continual testing by experimentation and observation  Darwin’s theory of evolution by natural selection integrates diverse areas of biological study and stimulates many new research questions  Ongoing research adds to our understanding of evolution

108. © 2014 Pearson Education, Inc. Figure 19.UN02a Guppies transplanted Pools with pike-cichlids and guppies Pools with killifish, but no guppies prior to transplant

109. © 2014 Pearson Education, Inc. Figure 19.UN02b Source population Number of colored spots Transplanted population Source population Transplanted population Area of colored spots (mm 2 ) 12 10 8 6 4 2 0 12 10 8 6 4 2 0

110. © 2014 Pearson Education, Inc. Figure 19.UN03 Observations Inferences and Over time, favorable traits accumulate in the population. Organisms produce more offspring than the environment can support. Individuals in a population vary in their heritable characteristics. Individuals that are well suited to their environment tend to leave more offspring than other individuals.

111. © 2014 Pearson Education, Inc. Figure 19.UN04