1. The Origin of Species chapter 24 Campbell and Reece

2. Speciation process by which one species splits into 2 or more species Speciation explains both the diversity of life and the unity of living things.

3. Speciation : forms bridge between: MICROEVOLUTION Evolutionary change below species level Example: ◦ change in allele frequencies in population over generations MACROEVOLUTION Evolutionary change above the species level Examples: ◦ origin of new group of organisms ◦ impact of mass extinctions

4. Biological Species Concept Species: a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring members of a species cannot produce viable, fertile offspring with other groups emphasizes the separateness of species due to reproductive barriers

5. What holds the gene pool of a species together? Gene Flow: transfer of alleles between populations of same species exchange of alleles tends to hold populations together genetically

6. Reproductive Isolation existence of biological barriers that keep members of 2 populations from interbreeding over long periods of time

7. Reproductive Isolation block gene flow between the species & limit formation of hybrids

8. Reproductive Isolation hybrids: offspring that result from the mating of individuals from 2 different species or from 2 true- breeding varieties of same species

9. Reproductive Isolation single barrier may not prevent all gene flow combination of several barriers can effectively isolate a species’ gene pool 2 categories: 1. prezygotic barriers 2. postzygotic barriers

10. Prezygotic Barriers block fertilization from happening by: 1. impeding members of different species from attempting to mate 2. preventing attempted mating from being completed successfully 3. hindering fertilization if mating was completed successfully

11. Postzygotic Barriers reproductive barrier that prevents hybrid zygotes produced by 2 different species from developing into viable, fertile adults 1. lethal developmental errors 2. infertility in viable offspring

12. Types of Prezygotic Reproductive Barriers 1. Habitat Isolation 2 species that occupy different habitats w/in same area may rarely interact example:

13. Types of Prezygotic Reproductive Barriers 2. Temporal Isolation: species breed during different times of day, different seasons, or different years eastern & western spotted skunks: 1 breeds in late summer other in late winter

14. Types of Prezygotic Reproductive Barriers 3. Behavioral Isolation: courtship rituals used to attract mates are effective barriers

15. Types of Prezygotic Reproductive Barriers 4. Mechanical Isolation: morphological differences prevent successful completion even if attempted

16. Types of Prezygotic Reproductive Barriers 5. Gametic Isolation: sperm of 1 species may not be able to fertilize egg of another: ◦ reproductive tract hostile to sperm ◦ sperm does not have enzymes to penetrate zona pellicida of another species

19. Types of Postzygotic Reproductive Barriers 1. Reduced Hybrid Viability: hybrids development or survival is impaired

20. Types of Postzygotic Reproductive Barriers 2. Reduced Hybrid Fertility: hybrids may develop and be healthy but they are not fertile

22. Types of Postzygotic Reproductive Barriers 3. Hybrid Breakdown: Some 1 st generation hybrids are fertile but those offspring are feeble or sterile

23. Species There is no single, universally applicable species concept that can define, explain, and identify all species. There are multiple ways to think about & define species.

24. Biological Species Limitations unable to use these characteristics on fossils of extinct species only applies to organisms that reproduce sexually only applies where there is no gene flow

25. Other Definitions of Species these dfns emphasize the unity w/in a species. 1. morphological species concept: characterizes a species by a structural feature applies to species that reproduce sexually or asexually how scientists distinguish most species disadvantage: subjective

26. Morphological Species Concept Problems: ◦ domestic dogs may look very different but are still same species ◦ mouse lemurs look very similar but there are 18 species of them ◦ grey mouse lemur lesser mouse lemur

27. Ecological Species Concept views species in terms of its niche ◦ the sum of how members of the species interact with the nonliving & living parts of their environment ◦ asexual or sexual species ◦ emphasizes role of disruptive NS as organisms adapt to different environmental conditions

28. Phylogenetic Species Concept defines species as smallest group of individuals that share a common ancestor, forming one branch on the “tree of life” determining degree of differences is difficult

29. Species There are >20 other ways to define species

30. Speciation can take place with or w/out geographic separation Speciation can occur in 2 main ways: 1. Allopatric Speciation 2. Sympatric Speciation

31. ALLOPATRIC SPECIATION “other country” gene flow is interrupted when population is divided into geographically isolated subpopulations

32. Allopatric Speciation Process once geographic separation has occurred, the separated gene pools will each have their own mutations NS & genetic drift may alter allele frequencies in different ways in each subpopulation group

34. Allopatric Speciation

35. Evidence of Allopatric Speciation are many studies & examples supporting this type speciation indirect support: regions that are isolated or highly subdivided have more species than regions w/out those features

36. Drosophila Experiment

37. Sympatric Speciation “same country” occurs in populations in same geographic area less common than allopatric occurs if gene flow is reduced by factors like: ◦ polyploidy ◦ habitat differentiation ◦ sexual selection

38. POLYPLOIDY means extra sets of chromosomes can occur in animals gray tree frog ( Hyla versicolor) ◦ around Great Lakes

39. Polyploidy much more common in plants estimate: 80% of today’s plants species have ancestors that formed by polyploid speciation 2 forms

40. 1. Autopolyploid individual has >2 chromosome sets all derived from a single species plant polyploidy

41. Plant Polyploidy tetraploid plant can produce fertile tetraploid offspring by self- pollinating or mating with other tetraploids

42. 2. Allopolyploid 2 different species interbreed making a hybrid hybrid reproduces asexually over generations sterile hybrid  fertile polyploid (called an allopolyploid)

43. Allopolyploids can breed with each other but not with either of their parents so are a new species rare: 5 new plant species since 1850 documented Mimulus peregrinus

44. Allopolyploids include many agricultural crops Triticum aestivum (bread wheat) has 6 sets chromosomes (2 pair from each of 3 parents), an allohexaploid 1 st polyploidy event probably occurred ~8,000 yrs ago as spontaneous hybrid

45. Top 2 parents bottom: Triticum aestivum

46. Allopolyploids plant geneticists “create” new polyploids making hybrids with desired characteristics use chemicals that induce meiotic & mitotic errors

47. Habitat Differentiation Sympatric speciation can occur when genetic factors enable a subpopulation to exploit a habitat or resource used by the parent population

48. Rhagoletis pomonella North American apple maggot fly original habitat was the native hawthorn tree

49. Habitat Differentiation apples mature faster than hawthorn fruit, NS has favored flies with rapid development have an allele that benefits flies that feed off only 1 or the other not both (post-zygotic barrier to reproduction) flies feeding on apple trees now show temporal isolation from flies still eating hawthorn fruit (prezygotic restriction to gene flow)

50. Sexual Selection can also drive sympatric speciation: cichlid fish Pundamilia pundamilia

51. Cichlid Fish >600 species found in Lake Victoria originated in past 100 000 yrs hypothesis : subgroups of original population adapted to different food sources  genetic divergence female preference for mates may also be a factor: 1 species breeding males have blue back another species has orange back

52. Breeding Cichlids Colors

53. Sexual Selection Study placed the 2 subspecies in same tank used monochromatic orange light so both appeared very similar females bred with either conclusion: mate choice by females is based on male coloration so it’s the main reproductive barrier (prezygotic behavior)

54. Allopatric & Sympatric Speciation are the 2 main modes of speciation Allopatric geographic isolation NS genetic drift sexual selection Sympatric requires emergence of a reproductive barrier that isolates a subgroup less common polyploidy sexual selection

55. Hybrid Zones A region in which members of different species meet & mate, producing at least some offspring of mixed ancestry. Hybrids are more likely to die as embryos & show a variety of morphological abnormalities

56. Hybrids of yellow-bellied & fire-bellied toads Bombino bambino Bombino variegata Hybrids are more likely to die as embryos & show a variety of morphological abnormalities

58. Hybrid Zones

59. Hybrid Zones over Time barriers between parent species may increase (strengthening of reproductive barriers) eventually hybrids no longer formed

60. Hybrid Zones over Time Or barrier weakens 2 species fuse

61. Hybrid Zone over Time 3 rd possibility: status quo is stabilized and hybrids continue to be produced

62. Speciation Time Course Patterns in the Fossil Record ◦ Punctuated Equilibrium: geologic periods of apparent stasis then sudden dramatic changes demonstrated by some species ◦ other species only show gradual changes coined Gradualism

63. Punctuated Equilibrium

64. Punctuated Equilibrium & Gradualism

65. Speciation Rates

66. Hybrid sunflower that diverged reproductively from parents to form new species over short time period F1 generation 5% fertility F5 generation >90% fertile

67. Genetics of Speciation fundamental question for biologists: How many genes must change to form a new species?

68. Single Gene Examples found in a few cases: Euhadra ◦ snail : single gene resulted in mechanical barrier to reproduction

69. Small # of Genes 2 monkey flower species have different pollinators = strong barrier to cross-pollination Mimulus lewisii Mimulus cardinalis

70. Large #s of Genes Hybrid of 2 species of fruit fly: Drosophila pseudoobscura results from gene interactions among at least 4 loci

71. Large #s of Genes Western sunflower: hybrid species thrives in dry sand but 2 parents live in nearby moister environments