1. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings PowerPoint ® Lecture Presentations for Biology Eighth Edition Neil Campbell and Jane Reece Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp Chapter 24 The Origin of Species

2. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Overview: That “Mystery of Mysteries” In the Galápagos Islands Darwin discovered plants and animals found nowhere else on Earth Video: Galápagos Tortoise Video: Galápagos Tortoise

3. Fig. 24-1

4. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Speciation, the origin of new species, is at the focal point of evolutionary theory Evolutionary theory must explain how new species originate and how populations evolve Microevolution consists of adaptations that evolve within a population, confined to one gene pool Macroevolution refers to evolutionary change above the species level Animation: Macroevolution Animation: Macroevolution

5. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 24.1: The biological species concept emphasizes reproductive isolation Species is a Latin word meaning “kind” or “appearance” Biologists compare morphology, physiology, biochemistry, and DNA sequences when grouping organisms

6. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Biological Species Concept The biological species concept states that a species is a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring; they do not breed successfully with other populations Gene flow between populations holds the phenotype of a population together

7. Fig. 24-2 (a) Similarity between different species (b) Diversity within a species

8. Fig. 24-2a (a) Similarity between different species

9. Fig. 24-2b (b) Diversity within a species

10. Fig. 24-3 EXPERIMENT RESULTS Example of a gene tree for population pair A-B AllelePopulation Gene flow event 1 B B B B 5 6 7 2 3 4 A A A Allele 1 is more closely related to alleles 2, 3, and 4 than to alleles 5, 6, and 7. Inference: Gene flow occurred. Alleles 5, 6, and 7 are more closely related to one another than to alleles in population A. Inference: No gene flow occurred. Pair of populations with detected gene flow Estimated minimum number of gene flow events to account for genetic patterns Distance between populations (km) A-B K-L A-C B-C F-G G-I C-E 5 3 2–3 2 2 2 1–2 340 720 1,390 1,190 1,110 760 1,310

11. Fig. 24-3a EXPERIMENT Example of a gene tree for population pair A-B AllelePopulation Gene flow event 1B B B B 5 6 7 2 3 4 A A A Allele 1 is more closely related to alleles 2, 3, and 4 than to alleles 5, 6, and 7. Inference: Gene flow occurred. Alleles 5, 6, and 7 are more closely related to one another than to alleles in population A. Inference: No gene flow occurred.

12. Fig. 24-3b RESULTS Pair of populations with detected gene flow Estimated minimum number of gene flow events to account for genetic patterns Distance between populations (km) A-B K-L A-C B-C F-G G-I C-E 5 3 2–3 2 2 2 1–2 340 720 1,390 1,190 760 1,110 1,310

13. Fig. 24-3c Grey-crowned babblers

14. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Reproductive Isolation Reproductive isolation is the existence of biological factors (barriers) that impede two species from producing viable, fertile offspring Hybrids are the offspring of crosses between different species Reproductive isolation can be classified by whether factors act before or after fertilization

15. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Prezygotic barriers block fertilization from occurring by: – Impeding different species from attempting to mate – Preventing the successful completion of mating – Hindering fertilization if mating is successful

16. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Habitat isolation: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers

17. Fig. 24-4 Prezygotic barriers Habitat Isolation Individuals of different species Temporal Isolation Behavioral Isolation Mating attempt Mechanical Isolation Gametic Isolation Fertilization Reduced Hybrid ViabilityReduced Hybrid Fertility Postzygotic barriers Hybrid Breakdown Viable, fertile offspring (a) (b) (d) (c)(e) (f) (g)(h) (i) (j) (l) (k)

18. Fig. 24-4a Habitat IsolationTemporal Isolation Prezygotic barriers Behavioral Isolation Mating attempt Mechanical Isolation (f) (e) (c) (a) (b) (d) Individuals of different species

19. Fig. 24-4i Prezygotic barriers Gametic Isolation Fertilization Reduced Hybrid Viability Postzygotic barriers Reduced Hybrid FertilityHybrid Breakdown Viable, fertile offspring (g) (h) (i) (j) (l) (k)

20. Fig. 24-4b Prezygotic barriers Habitat Isolation Temporal Isolation Behavioral Isolation Mechanical Isolation Individuals of different species Mating attempt

21. Prezygotic barriers Fig. 24-4j Gametic Isolation Fertilization Reduced Hybrid Viability Reduced Hybrid Fertility Postzygotic barriers Hybrid Breakdown Viable, fertile offspring

22. Fig. 24-4c (a) Water-dwelling Thamnophis

23. Fig. 24-4d (b) Terrestrial Thamnophis

24. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes

25. Fig. 24-4e (c) Eastern spotted skunk (Spilogale putorius)

26. Fig. 24-4f (d) Western spotted skunk (Spilogale gracilis)

27. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers Video: Blue-footed Boobies Courtship Ritual Video: Blue-footed Boobies Courtship Ritual Video: Giraffe Courtship Ritual Video: Giraffe Courtship Ritual Video: Albatross Courtship Ritual Video: Albatross Courtship Ritual

28. Fig. 24-4g (e) Courtship ritual of blue- footed boobies

29. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Mechanical isolation: Morphological differences can prevent successful mating

30. Fig. 24-4h (f) Bradybaena with shells spiraling in opposite directions

31. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Gametic isolation: Sperm of one species may not be able to fertilize eggs of another species

32. Fig. 24-4k (g) Sea urchins

33. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult: – Reduced hybrid viability – Reduced hybrid fertility – Hybrid breakdown

34. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Reduced hybrid viability: Genes of the different parent species may interact and impair the hybrid’s development

35. Fig. 24-4l (h) Ensatina hybrid

36. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Reduced hybrid fertility: Even if hybrids are vigorous, they may be sterile

37. Fig. 24-4m (i) Donkey

38. Fig. 24-4n ( j) Horse

39. Fig. 24-4o (k) Mule (sterile hybrid)

40. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hybrid breakdown: Some first-generation hybrids are fertile, but when they mate with another species or with either parent species, offspring of the next generation are feeble or sterile

41. Fig. 24-4p (l) Hybrid cultivated rice plants with stunted offspring (center)

42. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Limitations of the Biological Species Concept The biological species concept cannot be applied to fossils or asexual organisms (including all prokaryotes)

43. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Other Definitions of Species Other species concepts emphasize the unity within a species rather than the separateness of different species The morphological species concept defines a species by structural features – It applies to sexual and asexual species but relies on subjective criteria

44. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The ecological species concept views a species in terms of its ecological niche – It applies to sexual and asexual species and emphasizes the role of disruptive selection The phylogenetic species concept: defines a species as the smallest group of individuals on a phylogenetic tree – It applies to sexual and asexual species, but it can be difficult to determine the degree of difference required for separate species

45. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 24.2: Speciation can take place with or without geographic separation Speciation can occur in two ways: – Allopatric speciation – Sympatric speciation

46. Fig. 24-5 (a) Allopatric speciation (b) Sympatric speciation

47. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Allopatric (“Other Country”) Speciation In allopatric speciation, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations

48. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Process of Allopatric Speciation The definition of barrier depends on the ability of a population to disperse Separate populations may evolve independently through mutation, natural selection, and genetic drift

49. Fig. 24-6 A. harrisi A. leucurus

50. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Evidence of Allopatric Speciation Regions with many geographic barriers typically have more species than do regions with fewer barriers

51. Fig. 24-7 Mantellinae (Madagascar only): 100 species Rhacophorinae (India/Southeast Asia): 310 species Other Indian/ Southeast Asian frogs Millions of years ago (mya) 1 23 1 2 3 100 80 60 40 20 0 88 mya65 mya 56 mya India Madagascar

52. Fig. 24-7a Mantellinae (Madagascar only): 100 species Rhacophorinae (India/Southeast Asia): 310 species Other Indian/ Southeast Asian frogs Millions of years ago (mya) 100 80 6040 200 1 23

53. Fig. 24-7b India 88 mya65 mya 56 mya Madagascar 12 3

54. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Reproductive isolation between populations generally increases as the distance between them increases

55. Fig. 24-8 Geographic distance (km) Degree of reproductive isolation 0 0 50100150 250200 300 0.5 1.0 1.5 2.0

56. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Barriers to reproduction are intrinsic; separation itself is not a biological barrier

57. Fig. 24-9 EXPERIMENT RESULTS Initial population Some flies raised on starch medium Mating experiments after 40 generations Some flies raised on maltose medium Female Starch Maltose population 1 population 2 Male Starch Maltose Male Starch population 1 population 2 22 8 20 918 12 15 Mating frequencies in experimental group Mating frequencies in control group

58. Fig. 24-9a EXPERIMENT Initial population Some flies raised on starch medium Mating experiments after 40 generations Some flies raised on maltose medium

59. Fig. 24-9b RESULTS Female Starch Maltose population 1 population 2 Male Starch Maltose Male Starch population 1 population 2 22 820 918 12 15 Mating frequencies in experimental group Mating frequencies in control group

60. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takes place in geographically overlapping populations

61. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Polyploidy Polyploidy is the presence of extra sets of chromosomes due to accidents during cell division An autopolyploid is an individual with more than two chromosome sets, derived from one species

62. Fig. 24-10-1 2n = 64n = 12 Failure of cell division after chromosome duplication gives rise to tetraploid tissue.

63. Fig. 24-10-2 2n = 64n = 12 Failure of cell division after chromosome duplication gives rise to tetraploid tissue. 2n2n Gametes produced are diploid..

64. Fig. 24-10-3 2n = 64n = 12 Failure of cell division after chromosome duplication gives rise to tetraploid tissue. 2n2n Gametes produced are diploid.. 4n4n Offspring with tetraploid karyotypes may be viable and fertile.

65. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings An allopolyploid is a species with multiple sets of chromosomes derived from different species

66. Fig. 24-11-1 Species A 2n = 6 Normal gamete n = 3 Meiotic error Species B 2n = 4 Unreduced gamete with 4 chromosomes

67. Fig. 24-11-2 Species A 2n = 6 Normal gamete n = 3 Meiotic error Species B 2n = 4 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes

68. Fig. 24-11-3 Species A 2n = 6 Normal gamete n = 3 Meiotic error Species B 2n = 4 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Unreduced gamete with 7 chromosomes Normal gamete n = 3

69. Fig. 24-11-4 Species A 2n = 6 Normal gamete n = 3 Meiotic error Species B 2n = 4 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Unreduced gamete with 7 chromosomes Normal gamete n = 3 Viable fertile hybrid (allopolyploid) 2n = 10

70. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Polyploidy is much more common in plants than in animals Many important crops (oats, cotton, potatoes, tobacco, and wheat) are polyploids

71. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Habitat Differentiation Sympatric speciation can also result from the appearance of new ecological niches For example, the North American maggot fly can live on native hawthorn trees as well as more recently introduced apple trees

72. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Sexual Selection Sexual selection can drive sympatric speciation Sexual selection for mates of different colors has likely contributed to the speciation in cichlid fish in Lake Victoria

73. Fig. 24-12 EXPERIMENT Normal light Monochromatic orange light P. pundamilia P. nyererei

74. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Allopatric and Sympatric Speciation: A Review In allopatric speciation, geographic isolation restricts gene flow between populations Reproductive isolation may then arise by natural selection, genetic drift, or sexual selection in the isolated populations Even if contact is restored between populations, interbreeding is prevented

75. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings In sympatric speciation, a reproductive barrier isolates a subset of a population without geographic separation from the parent species Sympatric speciation can result from polyploidy, natural selection, or sexual selection

76. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 24.3: Hybrid zones provide opportunities to study factors that cause reproductive isolation A hybrid zone is a region in which members of different species mate and produce hybrids

77. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Patterns Within Hybrid Zones A hybrid zone can occur in a single band where adjacent species meet Hybrids often have reduced fitness compared with parent species The distribution of hybrid zones can be more complex if parent species are found in multiple habitats within the same region

78. Fig. 24-13 EUROPE Fire-bellied toad range Hybrid zone Yellow-bellied toad range Yellow-bellied toad, Bombina variegata Fire-bellied toad, Bombina bombina Allele frequency (log scale) Distance from hybrid zone center (km) 4030 20 10 0 0.01 0.1 0.5 0.9 0.99

79. Fig. 24-13a Yellow-bellied toad, Bombina variegata

80. Fig. 24-13b Fire-bellied toad, Bombina bombina

81. Fig. 24-13c Fire-bellied toad range Yellow-bellied toad range Hybrid zone Allele frequency (log scale) Distance from hybrid zone center (km) 403020 10 0 0.01 0.1 0.5 0.9 0.99

82. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hybrid Zones over Time When closely related species meet in a hybrid zone, there are three possible outcomes: – Strengthening of reproductive barriers – Weakening of reproductive barriers – Continued formation of hybrid individuals

83. Fig. 24-14-1 Gene flow Population (five individuals are shown) Barrier to gene flow

84. Fig. 24-14-2 Gene flow Population (five individuals are shown) Barrier to gene flow Isolated population diverges

85. Fig. 24-14-3 Gene flow Population (five individuals are shown) Barrier to gene flow Isolated population diverges Hybrid zone Hybrid

86. Fig. 24-14-4 Gene flow Population (five individuals are shown) Barrier to gene flow Isolated population diverges Hybrid zone Hybrid Possible outcomes: Reinforcement OR Fusion Stability

87. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Reinforcement: Strengthening Reproductive Barriers The reinforcement of barriers occurs when hybrids are less fit than the parent species Over time, the rate of hybridization decreases Where reinforcement occurs, reproductive barriers should be stronger for sympatric than allopatric species

88. Fig. 24-15 Sympatric male pied flycatcher Allopatric male pied flycatcher Pied flycatchers Collared flycatchers Number of females (none) Females mating with males from: Own species Other species Sympatric males Own species Other species Allopatric males 0 4 8 12 16 20 24 28

89. Fig. 24-15a Sympatric male pied flycatcher Allopatric male pied flycatcher

90. Fig. 24-15b Pied flycatchers Collared flycatchers 28 24 20 16 12 8 4 0 (none) Number of females Females mating with males from: Own species Other species Sympatric males Own species Other species Allopatric males

91. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fusion: Weakening Reproductive Barriers If hybrids are as fit as parents, there can be substantial gene flow between species If gene flow is great enough, the parent species can fuse into a single species

92. Fig. 24-16 Pundamilia nyerereiPundamilia pundamilia Pundamilia “turbid water,” hybrid offspring from a location with turbid water

93. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Stability: Continued Formation of Hybrid Individuals Extensive gene flow from outside the hybrid zone can overwhelm selection for increased reproductive isolation inside the hybrid zone In cases where hybrids have increased fitness, local extinctions of parent species within the hybrid zone can prevent the breakdown of reproductive barriers

94. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Concept 24.4: Speciation can occur rapidly or slowly and can result from changes in few or many genes Many questions remain concerning how long it takes for new species to form, or how many genes need to differ between species

95. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings The Time Course of Speciation Broad patterns in speciation can be studied using the fossil record, morphological data, or molecular data

96. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Patterns in the Fossil Record The fossil record includes examples of species that appear suddenly, persist essentially unchanged for some time, and then apparently disappear Niles Eldredge and Stephen Jay Gould coined the term punctuated equilibrium to describe periods of apparent stasis punctuated by sudden change The punctuated equilibrium model contrasts with a model of gradual change in a species’ existence

97. Fig. 24-17 (a) Punctuated pattern (b) Gradual pattern Time

98. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Speciation Rates The punctuated pattern in the fossil record and evidence from lab studies suggests that speciation can be rapid The interval between speciation events can range from 4,000 years (some cichlids) to 40,000,000 years (some beetles), with an average of 6,500,000 years

99. Fig. 24-18 (a) The wild sunflower Helianthus anomalus H. anomalus (b) The genetic composition of three chromosomes in H. anomalus and in experimental hybrids Chromosome 1 Chromosome 2 Chromosome 3 Experimental hybrid Key Region diagnostic for parent species H. petiolaris Region diagnostic for parent species H. annuus Region lacking information on parental origin

100. Fig. 24-18a (a) The wild sunflower Helianthus anomalus

101. Fig. 24-18b (b) The genetic composition of three chromosomes in H. anomalus and in experimental hybrids Region lacking information on parental origin Region diagnostic for parent species H. petiolaris Region diagnostic for parent species H. annuus Key Experimental hybrid Chromosome 3 Chromosome 2 Chromosome 1 H. anomalus

102. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Studying the Genetics of Speciation The explosion of genomics is enabling researchers to identify specific genes involved in some cases of speciation Depending on the species in question, speciation might require the change of only a single allele or many alleles

103. Fig. 24-19

104. Fig. 24-20 (a) Typical Mimulus lewisii(b) M. lewisii with an M. cardinalis flower-color allele (c) Typical Mimulus cardinalis(d) M. cardinalis with an M. lewisii flower-color allele

105. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings From Speciation to Macroevolution Macroevolution is the cumulative effect of many speciation and extinction events

106. Fig. 24-UN1 Original population Allopatric speciation Sympatric speciation

107. Fig. 24-UN2 Ancestral species: Triticum monococcum (2n = 14) AABB Wild Triticum (2n = 14) Product: AA BB DD T. aestivum (bread wheat) (2n = 42) Wild T. tauschii (2n = 14) DD

108. Fig. 24-UN3

109. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings You should now be able to: 1.Define and discuss the limitations of the four species concepts 2.Describe and provide examples of prezygotic and postzygotic reproductive barriers 3.Distinguish between and provide examples of allopatric and sympatric speciation 4.Explain how polyploidy can cause reproductive isolation 5.Define the term hybrid zone and describe three outcomes for hybrid zones over time