22 The Origin of Species

Aim: How can we describe the phenomenon of speciation Aim: How can we describe the phenomenon of speciation? Do Now: How can two organisms that were originally the same species, become two different species? Homework: Read chapter 22 and complete concept questions.

Speciation is the process by which one species splits into two or more species Speciation explains the features shared between organisms due to inheritance from their recent common ancestor 3

Speciation forms a conceptual bridge between microevolution and macroevolution Microevolution consists of changes in allele frequency in a population over time Macroevolution refers to broad patterns of evolutionary change above the species level 4

Concept 22.1: 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 5

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 populations together genetically 6

(a) Similarity between different species Figure 22.2 Figure 22.2 The biological species concept is based on the potential to interbreed rather than on physical similarity (a) Similarity between different species (b) Diversity within a species 7

Reproductive Isolation Reproductive isolation is the existence of biological 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 barriers act before or after fertilization 8

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 9

Habitat isolation Temporal isolation Behavioral isolation Figure 22.3a Prezygotic barriers Habitat isolation Temporal isolation Behavioral isolation MATING ATTEMPT (a) (c) (e) Figure 22.3a Exploring reproductive barriers (part 1: prezygotic barriers) (d) (b) 10

Habitat isolation: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers 11

Figure 22.3aa (a) Figure 22.3aa Exploring reproductive barriers (part 1a: habitat isolation, water) 12

Figure 22.3ab (b) Figure 22.3ab Exploring reproductive barriers (part 1b: habitat isolation, terrestrial) 13

Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes 14

Figure 22.3ac (c) Figure 22.2ac Exploring reproductive barriers (part 1c: temporal isolation, winter) 15

Figure 22.3ad (d) Figure 22.2ad Exploring reproductive barriers (part 1d: temporal isolation, summer) 16

Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers 17

Figure 22.3ae (e) Figure 22.ae Exploring reproductive barriers (part 1e: behavioral isolation) 18

Mechanical isolation Gametic isolation Figure 22.3b Prezygotic barriers Mechanical isolation Gametic isolation MATING ATTEMPT FERTILIZATION (f) (g) Figure 22.3b Exploring reproductive barriers (part 2: prezygotic barriers) 19

Mechanical isolation: Morphological differences prevent successful mating 20

Figure 22.3bf (f) Figure 22.3bf Exploring reproductive barriers (part 2f: mechanical isolation) 21

Gametic isolation: Sperm of one species may not be able to fertilize eggs of another species 22

Figure 22.3bg (g) Figure 22.3bg Exploring reproductive barriers (part 2g: gametic isolation) 23

Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult by: Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown 24

Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown Figure 22.3c Postzygotic barriers Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown VIABLE, FERTILE OFFSPRING FERTILIZATION (h) (i) (l) (j) Figure 22.3c Exploring reproductive barriers (part 3: postzygotic barriers) (k) 25

Reduced hybrid viability: Genes of the different parent species may interact and impair the hybrid’s development or survival 26

Figure 22.3ch (h) Figure 22.3ch Exploring reproductive barriers (part 3h: reduced hybrid viability) 27

Reduced hybrid fertility: Even if hybrids are vigorous, they may be sterile 28

Figure 22.3ci (i) Figure 22.3ci Exploring reproductive barriers (part 3i: reduced hybrid fertility) 29

Figure 22.3cj (j) Figure 22.3cj Exploring reproductive barriers (part 3j: reduced hybrid fertility) 30

Figure 22.3ck (k) Figure 22.3ck Exploring reproductive barriers (part 3k: reduced hybrid fertility) 31

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 32

Figure 22.3cl (l) Figure 22.3cl Exploring reproductive barriers (part 3l: hybrid breakdown) 33

Aim: How can we continue to describe the phenomenon of speciation Aim: How can we continue to describe the phenomenon of speciation? Do Now: Other that molecular similarities, what other information can be used to define a species?

Prezygotic barriers Habitat isolation Temporal isolation Behavioral Figure 22.3d Prezygotic barriers Habitat isolation Temporal isolation Behavioral isolation MATING ATTEMPT Mechanical isolation Gametic isolation MATING ATTEMPT FERTILIZATION Figure 22.3d Exploring reproductive barriers (part 4: art summary) Postzygotic barriers Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown VIABLE, FERTILE OFFSPRING FERTILIZATION 35

Limitations of the Biological Species Concept The biological species concept cannot be applied to fossils or asexual organisms (including all prokaryotes) The biological species concept emphasizes absence of gene flow However, gene flow can occur between distinct species For example, grizzly bears and polar bears can mate to produce “grolar bears” 36

Grizzly bear (U. arctos) Polar bear (U. maritimus) Figure 22.4 Grizzly bear (U. arctos) Figure 22.4 Hybridization between two species of bears in the genus Ursus Hybrid “grolar bear” Polar bear (U. maritimus) 37

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 38

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 39

Concept 22.2: Speciation can take place with or without geographic separation Speciation can occur in two ways: Allopatric speciation Sympatric speciation 40

Allopatric (“Other Country”) Speciation In allopatric speciation, gene flow is interrupted when a population is divided into geographically isolated subpopulations. For example, the flightless cormorant of the Galápagos likely originated from a flying species on the mainland. 41

The Process of Allopatric Speciation The definition of a geographic barrier depends on the ability of a population to disperse. For example, a canyon may create a barrier for small rodents, but not birds, coyotes, or pollen. 42

Reproductive isolation may arise as a result of genetic divergence. Separate populations may evolve independently through mutation, natural selection, and genetic drift. Reproductive isolation may arise as a result of genetic divergence. For example, mosquito fish in the Bahamas comprise several isolated populations in different ponds. 43

body shape that enables rapid bursts of speed (a) Under low predation: Figure 22.6 Under high predation: body shape that enables rapid bursts of speed (a) Under low predation: body shape that favors long, steady swimming (b) Figure 22.6 Reproductive isolation as a by-product of selection 44

Evidence of Allopatric Speciation Fifteen pairs of sister species of snapping shrimp (Alpheus) are separated by the Isthmus of Panama These species originated from 9 million to 3 million years ago, when the Isthmus of Panama formed and separated the Atlantic and Pacific waters 45

A. formosus A. nuttingi ATLANTIC OCEAN Isthmus of Panama PACIFIC OCEAN Figure 22.7 A. formosus A. nuttingi ATLANTIC OCEAN Isthmus of Panama PACIFIC OCEAN Figure 22.7 Allopatric speciation in snapping shrimp (Alpheus) A. panamensis A. millsae 46

Regions with many geographic barriers typically have more species than do regions with fewer barriers Reproductive isolation between populations generally increases as the geographic distance between them increases 47

Barriers to reproduction are intrinsic; separation itself is not a biological barrier Intrinsic reproductive barriers can develop in experimentally isolated populations 48

Initial population of fruit flies (Drosophila pseudoobscura) Figure 22.8 Experiment Initial population of fruit flies (Drosophila pseudoobscura) Some flies raised on starch medium Some flies raised on maltose medium Mating experiments after 40 generations Results Female Female Starch population 1 Starch population 2 Starch Maltose Figure 22.8 Inquiry: Can divergence of allopatric populations lead to reproductive isolation? Starch population 1 Starch 22 9 18 15 Male Male population 2 Starch Maltose 8 20 12 15 Number of matings in experimental group Number of matings in control group 49

Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takes place in populations that live in the same geographic area Sympatric speciation occurs when gene flow is reduced between groups that remain in contact through factors including Polyploidy Habitat differentiation Sexual selection 50

Polyploidy Polyploidy is the presence of extra sets of chromosomes due to accidents during cell division Polyploidy is much more common in plants than in animals An autopolyploid is an individual with more than two chromosome sets, derived from one species The offspring of matings between autopolyploids and diploids have reduced fertility 51

Tetraploid cell 4n  12 New species (4n) Figure 22.UN01 Cell division error 2n  6 Tetraploid cell 4n  12 Figure 22.UN01 In-text figure, autopolyploid, p. 425 2n New species (4n) Gametes produced by tetraploids 52

Allopolyploids cannot interbreed with either parent species An allopolyploid is a species with multiple sets of chromosomes derived from different species Allopolyploids cannot interbreed with either parent species 53

Meiotic error; chromosome number not reduced from 2n to n Figure 22.9-1 Species A 2n  6 Species B 2n  4 Meiotic error; chromosome number not reduced from 2n to n Normal gamete n  3 Unreduced gamete with 4 chromosomes Figure 22.9-1 One mechanism for allopolyploid speciation in plants (step 1) 54

Meiotic error; chromosome number not reduced from 2n to n Figure 22.9-2 Species A 2n  6 Species B 2n  4 Meiotic error; chromosome number not reduced from 2n to n Normal gamete n  3 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Figure 22.9-2 One mechanism for allopolyploid speciation in plants (step 2) 55

Meiotic error; chromosome number not reduced from 2n to n Figure 22.9-3 Species A 2n  6 Species B 2n  4 Meiotic error; chromosome number not reduced from 2n to n Normal gamete n  3 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Normal gamete n  3 Figure 22.9-3 One mechanism for allopolyploid speciation in plants (step 3) Unreduced gamete with 7 chromosomes 56

Meiotic error; chromosome number not reduced from 2n to n Figure 22.9-4 Species A 2n  6 Species B 2n  4 Meiotic error; chromosome number not reduced from 2n to n Normal gamete n  3 Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Normal gamete n  3 Figure 22.9-4 One mechanism for allopolyploid speciation in plants (step 4) Unreduced gamete with 7 chromosomes New species: viable fertile hybrid (allopolyploid) 2n  10 57

Allopatric speciation: forms a new species while Figure 22.5 Figure 22.5 Two main modes of speciation Allopatric speciation: forms a new species while geographically isolated. (a) (b) Sympatric speciation: a subset forms a new species without geographic separation. 58

Aim: How can we continue to describe the phenomenon of speciation Aim: How can we continue to describe the phenomenon of speciation? Do Now: Describe the difference between allopatric and sympatric speciation.

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 60

Sexual Selection Sexual selection can drive sympatric speciation Sexual selection for mates of different colors has likely contributed to speciation in cichlid fish in Lake Victoria 61

Monochromatic orange light Normal light P. pundamilia Figure 22.10 Experiment Monochromatic orange light Normal light P. pundamilia Figure 22.10 Inquiry: Does sexual selection in cichlids result in reproductive isolation? P. nyererei 62

Concept 22.3: Hybrid zones reveal factors that cause reproductive isolation A hybrid zone is a region in which members of different species mate and produce hybrids Hybrids are the result of mating between species with incomplete reproductive barriers 63

Patterns Within Hybrid Zones A hybrid zone can occur in a single band where adjacent species meet For example, two species of toad in the genus Bombina interbreed in a long and narrow hybrid zone 64

B. variegata-specific allele Distance from hybrid zone center (km) Figure 22.11 Fire-bellied toad range Fire-bellied toad, Bombina bombina Hybrid zone Yellow-bellied toad range 0.99 Hybrid zone 0.9 Figure 22.11 A narrow hybrid zone for Bombina toads in Europe B. variegata-specific allele Frequency of 0.5 Yellow-bellied toad range Fire-bellied toad range Yellow-bellied toad, Bombina variegata 0.1 0.01 40 30 20 10 10 20 Distance from hybrid zone center (km) 65

Hybrids often have reduced fitness compared with parent species The distribution of hybrid zones can be more complex if parent species are found in patches within the same region 66

Hybrid Zones over Time When closely related species meet in a hybrid zone, there are three possible outcomes: Reinforcement Fusion Stability 67

Gene flow Population Barrier to gene flow Figure 22.12-1 Figure 22.12-1 Formation of a hybrid zone and possible outcomes for hybrids over time (step 1) Population Barrier to gene flow 68

Isolated population diverges. Gene flow Population Barrier to Figure 22.12-2 Isolated population diverges. Gene flow Figure 22.12-2 Formation of a hybrid zone and possible outcomes for hybrids over time (step 2) Population Barrier to gene flow 69

Isolated population diverges. Hybrid zone Gene flow Population Figure 22.12-3 Isolated population diverges. Hybrid zone Gene flow Figure 22.12-3 Formation of a hybrid zone and possible outcomes for hybrids over time (step 3) Population Barrier to gene flow Hybrid individual 70

Isolated population diverges. Possible outcomes: Hybrid zone Figure 22.12-4 Isolated population diverges. Possible outcomes: Hybrid zone Reinforcement Fusion Gene flow Figure 22.12-4 Formation of a hybrid zone and possible outcomes for hybrids over time (step 4) Population Barrier to gene flow Hybrid individual Stability 71

Reinforcement occurs when hybrids are less fit than the parent species Natural selection strengthens (reinforces) reproductive barriers, and, over time, the rate of hybridization decreases Where reinforcement occurs, reproductive barriers should be stronger for sympatric than for allopatric species 72

This might be causing the fusion of many species Fusion of the parent species into a single species may occur if hybrids are as fit as parents, allowing substantial gene flow between species For example, researchers think that pollution in Lake Victoria has reduced the ability of female cichlids to distinguish males of different species This might be causing the fusion of many species 73

Pundamilia pundamilia Figure 22.13 Pundamilia nyererei Pundamilia pundamilia Figure 22.13 Fusion: the breakdown of reproductive barriers Pundamilia “turbid water,” hybrid offspring from a location with turbid water 74

In a stable hybrid zone, hybrids continue to be produced over time Stability of the hybrid zone may be achieved if extensive gene flow from outside the hybrid zone can overwhelm selection for increased reproductive isolation inside the hybrid zone In a stable hybrid zone, hybrids continue to be produced over time 75

Concept 22.4: 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 76

The Time Course of Speciation Broad patterns in speciation can be studied using the fossil record, morphological data, or molecular data 77

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 These periods of apparent stasis punctuated by sudden change are called punctuated equilibria The punctuated equilibrium model contrasts with a model of gradual change in a species’ existence 78

(a) Punctuated model Time (b) Gradual model Figure 22.14 Figure 22.14 Two models for the tempo of speciation, based on patterns observed in the fossil record (b) Gradual model 79

Speciation Rates The punctuated pattern in the fossil record and evidence from lab studies suggest that speciation can be rapid For example, the sunflower Helianthus anomalus originated from the hybridization of two other sunflower species and quickly diverged into a new species 80

The interval between speciation events can range from 4,000 years (some cichlids) to 40 million years (some beetles), with an average of 6.5 million years 81

Studying the Genetics of Speciation A fundamental question of evolutionary biology persists: How many genes change when a new species forms? Depending on the species in question, speciation might require the change of only a single allele or many alleles For example, in Japanese Euhadra snails, the direction of shell spiral affects mating and is controlled by a single gene 82

In monkey flowers (Mimulus), two loci affect flower color, which influences pollinator preference Pollination that is dominated by either hummingbirds or bees can lead to reproductive isolation of the flowers In other species, speciation can be influenced by larger numbers of genes and gene interactions 83

From Speciation to Macroevolution Macroevolution is the cumulative effect of many speciation and extinction events 84