LECTURE PRESENTATIONS For CAMPBELL BIOLOGY NINTH EDITION Jane

LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 24 The Origin of Species Lectures by Erin Barley Kathleen Fitzpatrick © 2011 Pearson Education, Inc.

Overview: That “Mystery of Mysteries” • In the Galápagos Islands Darwin discovered plants and

Video: Galápagos Tortoise © 2011 Pearson Education, Inc.

Figure 24. 1

• 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 changes in allele frequency in a population over time • Macroevolution refers to broad patterns of evolutionary change above the species level © 2011 Pearson Education, Inc.

Animation: Macroevolution Right-click slide / select “Play” © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Figure 24. 2 (a) Similarity between different species (b) Diversity within a species

Figure 24. 2 a (a) Similarity between different species

Figure 24. 2 b (b) Diversity within a species

Figure 24. 2 c

Figure 24. 2 d

Figure 24. 2 e

Figure 24. 2 f

Figure 24. 2 g

Figure 24. 2 h

Figure 24. 2 i

Figure 24. 2 j

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 © 2011 Pearson Education, Inc.

Figure 24. 3_a Prezygotic barriers Habitat Isolation Temporal Isolation (a) Gametic Isolation Mechanical Isolation Behavioral Isolation Individuals of different species Postzygotic barriers MATING ATTEMPT (c) (d) (e) Reduced Hybrid Viability Reduced Hybrid Fertility Hybrid Breakdown VIABLE, FERTILE OFFSPRING FERTILIZATION (f) (g) (h) (i) (j) (b) (k) (l)

Figure 24. 3_b Prezygotic barriers Habitat Isolation Temporal Isolation Individuals of different species (a) MATING ATTEMPT (c) (d) (b) Gametic Isolation Mechanical Isolation Behavioral Isolation (e) (f) FERTILIZATION (g)

Figure 24. 3_c Postzygotic barriers Reduced Hybrid Viability Reduced Hybrid Fertility Hybrid Breakdown VIABLE, FERTILE OFFSPRING FERTILIZATION (h) (i) (j) (k) (l)

• 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 © 2011 Pearson Education, Inc.

Figure 24. 3 a (a)

Figure 24. 3 b (b)

• Habitat isolation: Two species encounter each other rarely, or not at all,

Figure 24. 3 c (c)

Figure 24. 3 d (d)

• Temporal isolation: Species that breed at different times of the day, different

Figure 24. 3 e (e)

• Behavioral isolation: Courtship rituals and other behaviors unique to a species are

Video: Albatross Courtship Ritual © 2011 Pearson Education, Inc.

Video: Giraffe Courtship Ritual © 2011 Pearson Education, Inc.

Video: Blue-footed Boobies Courtship Ritual © 2011 Pearson Education, Inc.

Figure 24. 3 f (f)

• Mechanical isolation: Morphological differences can prevent successful mating © 2011 Pearson Education,

Figure 24. 3 g (g)

• Gametic Isolation: Sperm of one species may not be able to fertilize

• Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile

Figure 24. 3 h (h)

• Reduced hybrid viability: Genes of the different parent species may interact and

Figure 24. 3 i (i)

Figure 24. 3 j (j)

Figure 24. 3 k (k)

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

Figure 24. 3 l (l)

• 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 © 2011 Pearson Education, Inc.

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” © 2011 Pearson Education, Inc.

Figure 24. 4 Grizzly bear (U. arctos) Polar bear (U. maritimus) Hybrid “grolar bear”

Figure 24. 4 a Grizzly bear (U. arctos)

Figure 24. 4 b Polar bear (U. maritimus)

Figure 24. 4 c Hybrid “grolar bear”

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 © 2011 Pearson Education, Inc.

• 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 © 2011 Pearson Education, Inc.

Concept 24. 2: Speciation can take place with or without geographic separation • Speciation

Figure 24. 5 (a) Allopatric speciation. A population forms a new species while geographically isolated from its parent population. (b) Sympatric speciation. A subset of a population forms a new species without geographic separation.

Allopatric (“Other Country”) Speciation • In allopatric speciation, gene flow is interrupted or reduced 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 © 2011 Pearson Education, Inc.

The Process of Allopatric Speciation • The definition of 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 © 2011 Pearson Education, Inc.

Figure 24. 6 A. harrisii A. leucurus

Figure 24. 6 a A. harrisii

Figure 24. 6 b A. leucurus

Figure 24. 6 c

• Separate populations may evolve independently through mutation, natural selection, and genetic drift • Reproductive isolation may arise as a result of genetic divergence – For example, mosquitofish in the Bahamas comprise several isolated populations in different ponds © 2011 Pearson Education, Inc.

Figure 24. 7 (a) Under high predation (b) Under low predation

Figure 24. 7 a

Figure 24. 7 b

Evidence of Allopatric Speciation • 15 pairs of sibling species of snapping shrimp (Alpheus) are separated by the Isthmus of Panama • These species originated 9 to 13 million years ago, when the Isthmus of Panama formed and separated the Atlantic and Pacific waters © 2011 Pearson Education, Inc.

Figure 24. 8 A. formosus A. nuttingi Atlantic Ocean Isthmus of Panama Pacific Ocean A. panamensis A. millsae

Figure 24. 8 a

Figure 24. 8 b Atlantic Ocean Isthmus of Panama Pacific Ocean

Figure 24. 8 c A. formosus

Figure 24. 8 d A. panamensis

Figure 24. 8 e A. nuttingi

Figure 24. 8 f A. millsae

• Regions with many geographic barriers typically have more species than do regions with fewer barriers • Reproductive isolation between populations generally increases as the distance between them increases – For example, reproductive isolation increases between dusky salamanders that live further apart © 2011 Pearson Education, Inc.

Degree of reproductive isolation Figure 24. 9 2. 0 1. 5 1. 0 0. 5 0 0 50 100 150 200 250 Geographic distance (km) 300

• Barriers to reproduction are intrinsic; separation itself is not a biological barrier

EXPERIMENT Initial population of fruit flies (Drosophila pseudoobscura) Some flies raised on maltose medium Some flies raised on starch medium Mating experiments after 40 generations RESULTS Female 22 9 8 20 Male Maltose Starch population 1 population 2 Number of matings in experimental group Starch population 2 population 1 Starch Female Maltose Figure 24. 10 18 15 12 15 Number of matings in control group

Figure 24. 10 a EXPERIMENT Initial population of fruit flies (Drosophila pseudoobscura) Some flies raised on maltose medium Some flies raised on starch medium Mating experiments after 40 generations

Figure 24. 10 b RESULTS Female Maltose 22 9 8 20 Number of matings in experimental group Starch population 2 population 1 Starch population 1 population 2 Male Maltose Starch Female 18 15 12 15 Number of matings in control group

Sympatric (“Same Country”) Speciation • In sympatric speciation, speciation takes place in geographically overlapping

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 © 2011 Pearson Education, Inc.

• An allopolyploid is a species with multiple sets of chromosomes derived from

Figure 24. 11 -1 Species A 2 n = 6 Normal gamete n=3 Species B 2 n = 4 Meiotic error; chromosome number not reduced from 2 n to n Unreduced gamete with 4 chromosomes

Figure 24. 11 -2 Species A 2 n = 6 Normal gamete n=3 Species B 2 n = 4 Meiotic error; chromosome number not reduced from 2 n to n Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes

Figure 24. 11 -3 Species A 2 n = 6 Normal gamete n=3 Species B 2 n = 4 Meiotic error; chromosome number not reduced from 2 n to n Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Normal gamete n=3 Unreduced gamete with 7 chromosomes

Figure 24. 11 -4 Species A 2 n = 6 Normal gamete n=3 Species B 2 n = 4 Meiotic error; chromosome number not reduced from 2 n to n Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Normal gamete n=3 Unreduced gamete with 7 chromosomes New species: viable fertile hybrid (allopolyploid) 2 n = 10

• Many important crops (oats, cotton, potatoes, tobacco, and wheat) are polyploids ©

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 © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Figure 24. 12 EXPERIMENT Normal light P. pundamilia P. nyererei Monochromatic orange light

Figure 24. 12 a Normal light P. pundamilia

Figure 24. 12 b Normal light P. nyererei

Figure 24. 12 c Monochromatic orange light P. pundamilia

Figure 24. 12 d Monochromatic orange light P. nyererei

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 © 2011 Pearson Education, Inc.

• 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 © 2011 Pearson Education, Inc.

Concept 24. 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 © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Figure 24. 13 EUROPE Fire-bellied toad range Hybrid zone Yellow-bellied toad, Bombina variegata Frequency of B. variegata-specific allele Yellow-bellied toad range Fire-bellied toad, Bombina bombina 0. 99 Hybrid zone 0. 9 Yellow-bellied toad range 0. 5 Fire-bellied toad range 0. 1 0. 01 40 10 0 20 10 20 30 Distance from hybrid zone center (km)

Figure 24. 13 a EUROPE Fire-bellied toad range Hybrid zone Yellow-bellied toad range

Frequency of B. variegata-specific allele Figure 24. 13 b 0. 99 Hybrid zone 0. 9 Yellow-bellied toad range 0. 5 Fire-bellied toad range 0. 1 0. 01 40 30 10 0 20 10 20 Distance from hybrid zone center (km)

Figure 24. 13 c Fire-bellied toad, Bombina bombina

Figure 24. 13 d Yellow-bellied toad, Bombina variegata

• 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 © 2011 Pearson Education, Inc.

Hybrid Zones over Time • When closely related species meet in a hybrid zone,

Figure 24. 14 -1 Gene flow Population Barrier to gene flow

Figure 24. 14 -2 Isolated population diverges Gene flow Population Barrier to gene flow

Figure 24. 14 -3 Isolated population diverges Gene flow Population Barrier to gene flow Hybrid zone Hybrid individual

Figure 24. 14 -4 Possible outcomes: Isolated population diverges Hybrid zone Reinforcement OR Fusion Gene flow Population OR Barrier to gene flow Hybrid individual Stability

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 – For example, in populations of flycatchers, males are more similar in allopatric populations than sympatric populations © 2011 Pearson Education, Inc.

Figure 24. 15 Females choosing between these males: 28 Number of females 24 Females choosing between these males: Sympatric pied male Allopatric pied male Sympatric collared male Allopatric collared male 20 16 12 8 4 0 (none) Own species Other species Female mate choice

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 • 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 © 2011 Pearson Education, Inc.

Figure 24. 16 Pundamilia nyererei Pundamilia pundamilia Pundamilia “turbid water, ” hybrid offspring from a location with turbid water

Figure 24. 16 a Pundamilia nyererei

Figure 24. 16 b Pundamilia pundamilia

Figure 24. 16 c Pundamilia “turbid water, ” hybrid offspring from a location with turbid water

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 © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

The Time Course of Speciation • Broad patterns in speciation can be studied using

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 equilibria 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 © 2011 Pearson Education, Inc.

Figure 24. 17 (a) Punctuated pattern Time (b) Gradual pattern

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 © 2011 Pearson Education, Inc.

Figure 24. 18

Figure 24. 19 EXPERIMENT H. annuus gamete H. petiolarus gamete F 1 experimental hybrid (4 of the 2 n = 34 chromosomes are shown) RESULTS H. anomalus Chromosome 1 Experimental hybrid H. anomalus Chromosome 2 Experimental hybrid

Figure 24. 19 a EXPERIMENT H. annuus gamete H. petiolarus gamete F 1 experimental hybrid (4 of the 2 n = 34 chromosomes are shown)

Figure 24. 19 b RESULTS H. anomalus Chromosome 1 Experimental hybrid H. anomalus Chromosome 2 Experimental hybrid

• The interval between speciation events can range from 4, 000 years (some

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 © 2011 Pearson Education, Inc.

• 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 © 2011 Pearson Education, Inc.

Figure 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

Figure 24. 20 a (a) Typical Mimulus lewisii

Figure 24. 20 b (b) M. lewisii with an M. cardinalis flower-color allele

Figure 24. 20 c (c) Typical Mimulus cardinalis

Figure 24. 20 d (d) M. cardinalis with an M. lewisii flower-color allele

From Speciation to Macroevolution • Macroevolution is the cumulative effect of many speciation and

Figure 24. UN 01 Cell division error 2 n = 6 Tetraploid cell 4 n = 12 2 n 2 n Gametes produced by tetraploids New species (4 n)

Figure 24. UN 02 Original population Allopatric speciation Sympatric speciation

Figure 24. UN 03 Ancestral species: Triticum monococcum (2 n = 14) Wild Triticum (2 n = 14) Product: T. aestivum (bread wheat) (2 n = 42) Wild T. tauschii (2 n = 14)

Figure 24. UN 04