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 Learning Target: 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

3. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Speciation, the origin of new species, is at the focal point of evolutionary theory 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 Evolution theory

4. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 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 Species do not breed successfully with other populations Gene flow between populations holds the phenotype of a population together

5. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 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 can be classified by whether factors act before or after fertilization

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

7. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 1.Habitat isolation: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers 2.Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes 3.Mechanical isolation: Morphological differences can prevent successful mating Prezygotic barriers

8. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 4.Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers 5.Gametic isolation: Sperm of one species may not be able to fertilize eggs of another species 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 Prezygotic barriers continued

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

10. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 1.Reduced hybrid viability: Genes of the different parent species may interact and impair the hybrid’s development 2.Reduced hybrid fertility: Even if hybrids are vigorous, they may be sterile Postzygotic barriers Mule (sterile hybrid)

11. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 3.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 Postzygotic barriers continued

12. 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

13. 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 Other Definitions of Species

14. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Speciation can take place with or without geographic separation Speciation can occur in two ways 1. Allopatric speciation 2. Sympatric speciation

15. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 1. Allopatric (“Other Country”) Speciation In allopatric speciation, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations 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

16. A. harrisi A. leucurus Regions with many geographic barriers typically have more species than do regions with fewer barriers Evidence of Allopatric Speciation

17. 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

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

19. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 2. Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takes place in geographically overlapping populations

20. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Polyploidy 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 An allopolyploid is a species with multiple sets of chromosomes derived from different species Polyploidy is much more common in plants than in animals (oats, cotton, potatoes and wheat)

21. 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

22. 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

23. 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

24. 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

25. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hybrid zones is a region in which members of different species mate and produce hybrids 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

26. 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

27. 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

28. 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

29. Pundamilia nyerereiPundamilia pundamilia Pundamilia “turbid water,” hybrid offspring from a location with turbid water

30. 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

31. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 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 Broad patterns in speciation can be studied using the fossil record, morphological data, or molecular data

32. 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 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

33. Punctuated pattern Gradual pattern Time

34. 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

35. 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 Macroevolution is the cumulative effect of many speciation and extinction events