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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 animals found nowhere else on Earth © 2011 Pearson Education, Inc. Figure 24.1 • ______________, the origin of new species, is at the focal point of evolutionary theory • Evolutionary theory must explain how _________ _______________and how populations evolve • _________________ consists of changes in allele frequency in a population over time • _________________refers to broad patterns of evolutionary change above the species level © 2011 Pearson Education, Inc. Concept 24.1: The biological species concept emphasizes reproductive isolation • ___________ is a Latin word meaning “kind” or “appearance” • Biologists compare ________________ ____________________________________ sequences when grouping organisms © 2011 Pearson Education, Inc. The Biological Species Concept • The ______________________states that a ________________ 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 • _________________ 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 Reproductive Isolation • ____________________ is the existence of biological factors (barriers) that impede two species from producing viable, fertile offspring • _____________ are the offspring of crosses between different species • Reproductive isolation can be classified by whether factors act __________ or __________ ___________________ © 2011 Pearson Education, Inc. 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) • _________________ block fertilization from occurring by: – ______________________________________ – ______________________________________ _______________ - ______________________________ if mating is successful © 2011 Pearson Education, Inc. • __________________: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers © 2011 Pearson Education, Inc. • ___________________: Species that breed at ________________________, different seasons, or different years cannot mix their gametes © 2011 Pearson Education, Inc. • ___________________: Courtship rituals and other behaviors unique to a species are effective barriers © 2011 Pearson Education, Inc. • _______________________: Morphological differences can prevent successful mating © 2011 Pearson Education, Inc. • ____________________: Sperm of one species may not be able to fertilize eggs of another species © 2011 Pearson Education, Inc. • ______________________prevent the hybrid zygote from developing into a viable, fertile adult: – Reduced hybrid ___________________ – Reduced hybrid ___________________ – ___________________________ © 2011 Pearson Education, Inc. • _____________________: Genes of the different parent species may interact and impair the hybrid’s development © 2011 Pearson Education, Inc. • _______________________: Even if hybrids are vigorous, they may be sterile © 2011 Pearson Education, Inc. • __________________: 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 ________________________ (including all prokaryotes) • The biological species concept emphasizes ___________________________ • However, gene flow can occur between distinct species – For example, grizzly bears and polar bears can mate to produce “______________” © 2011 Pearson Education, Inc. Figure 24.4 Grizzly bear (U. arctos) Polar bear (U. maritimus) Hybrid “grolar bear” Other Definitions of Species • Other species concepts emphasize the _________ within a species rather than the separateness of different species • The ______________________defines a species by structural features – It applies to sexual and asexual species but relies on ______________________________ © 2011 Pearson Education, Inc. • The __________________________views a species in terms of its ___________________ – It applies to sexual and asexual species and emphasizes the role of ____________________ • The _______________________defines a species as the ________________ group of individuals on a phylogenetic tree – It applies to sexual and asexual species, but it can be difficult to determine the ______________ required for separate species © 2011 Pearson Education, Inc. Concept 24.2: Speciation can take place with or without geographic separation • Speciation can occur in two ways: – ____________ speciation – ____________ speciation © 2011 Pearson Education, Inc. 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 ____________________, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations – For example, the flightless ____________ of the Galápagos likely originated from a flying species on the mainland © 2011 Pearson Education, Inc. The Process of Allopatric Speciation • The definition of _____________depends on the ability of a population to disperse – For example, a ________________ may create a barrier for small rodents, but not birds, coyotes, or pollen © 2011 Pearson Education, Inc. Figure 24.6 A. harrisii A. leucurus • Separate populations may evolve independently through _________________________________ • _______________________ may arise as a result of genetic divergence – For example, _________________ in the Bahamas comprise several isolated populations in different ponds © 2011 Pearson Education, Inc. Evidence of Allopatric Speciation • 15 pairs of sibling species of ___________ (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 • Regions with many geographic barriers typically have ___________than do regions with fewer barriers • Reproductive isolation between populations generally ____________________ as the distance between them increases – For example, reproductive isolation increases between dusky salamanders that live further apart © 2011 Pearson Education, Inc. • Barriers to reproduction are _____________; separation itself is not a biological barrier © 2011 Pearson Education, Inc. 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 Starch population 1 population 2 Number of matings in experimental group Starch Starch population 2 population 1 Starch Starch Male Female Maltose Figure 24.10 18 15 12 15 Number of matings in control group Sympatric (“Same Country”) Speciation • In ___________________, speciation takes place in geographically overlapping populations © 2011 Pearson Education, Inc. Polyploidy • __________________ is the presence of extra sets of chromosomes due to accidents during cell division • Polyploidy is much more common in _______ than in animals • An __________________ is an individual with more than two chromosome sets, derived from one species © 2011 Pearson Education, Inc. • An __________________ is a species with multiple sets of chromosomes derived from different species © 2011 Pearson Education, Inc. Figure 24.11-4 Species A 2n = 6 Normal gamete n=3 Species B 2n = 4 Meiotic error; chromosome number not reduced from 2n 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) 2n = 10 • Many important crops (oats, cotton, potatoes, tobacco, and wheat) are ___________________ © 2011 Pearson Education, Inc. Habitat Differentiation • _____________________ 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 • _________________ 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 Allopatric and Sympatric Speciation: A Review • In _________________ speciation, geographic isolation restricts gene flow between populations • Reproductive isolation may then arise by _______ ___________________in the isolated populations • Even if contact is restored between populations, interbreeding is ___________________ © 2011 Pearson Education, Inc. • In _____________ speciation, a reproductive barrier isolates a subset of a population without geographic separation from the parent species • Sympatric speciation can result from ________ ________________________________________ © 2011 Pearson Education, Inc. Concept 24.3: Hybrid zones reveal factors that cause reproductive isolation • A _______________ is a region in which members of different species mate and produce hybrids • Hybrids are the result of mating between species with ____________________________________ © 2011 Pearson Education, Inc. Patterns Within Hybrid Zones • A hybrid zone can occur in a single band where ______________________ – 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 Fire-bellied toad, Bombina bombina Yellow-bellied toad, Bombina variegata Frequency of B. variegata-specific allele Yellow-bellied toad range 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) • Hybrids often have __________________ compared with parent species • The distribution of hybrid zones can be more complex if parent species are found in _________ within the same region © 2011 Pearson Education, Inc. Hybrid Zones over Time • When closely related species meet in a hybrid zone, there are three possible outcomes: – __________________ – __________________ – __________________ © 2011 Pearson Education, Inc. Figure 24.14-4 Possible outcomes: Isolated population diverges Hybrid zone Reinforcement OR Fusion OR Gene flow Population Barrier to gene flow Hybrid individual Stability Reinforcement: Strengthening Reproductive Barriers • The ________________ of barriers occurs when _________________ than the parent species • Over time, the rate of hybridization ___________ • Where reinforcement occurs, reproductive barriers should be ________________ 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 (none) 0 Own species Other species Female mate choice Own species Other species Female mate choice Fusion: Weakening Reproductive Barriers • If hybrids are as fit as parents, there can be substantial _______________between species • If gene flow is great enough, the parent species can _________________________ • 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 _______________ _________________________ © 2011 Pearson Education, Inc. Figure 24.16 Pundamilia nyererei Pundamilia pundamilia 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 _________________ 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 __________ it takes for new species to form, or _____________ _______________ need to differ between species © 2011 Pearson Education, Inc. The Time Course of Speciation • Broad patterns in speciation can be studied using the fossil record, morphological data, or molecular data © 2011 Pearson Education, Inc. Patterns in the Fossil Record • The ______________ 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 ______________to describe periods of apparent stasis punctuated by sudden change • The punctuated equilibrium model ____________ 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 _______________ – For example, the sunflower Helianthus anomalus originated from the hybridization of two other sunflower species © 2011 Pearson Education, Inc. Figure 24.19 EXPERIMENT H. annuus gamete H. petiolarus gamete F1 experimental hybrid (4 of the 2n = 34 chromosomes are shown) RESULTS H. anomalus Chromosome 1 Experimental hybrid H. anomalus Chromosome 2 Experimental hybrid • The interval between speciation events can range from __________ years (some cichlids) to ______ _____________ years (some beetles), with an average of ______ million years © 2011 Pearson Education, Inc. Studying the Genetics of Speciation • A fundamental question of evolutionary biology persists: ___________________ when a new species forms? • Depending on the species in question, speciation might require the change of only a _________ ________________________________ • 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________________ , which influences pollinator preference • Pollination that is dominated by either hummingbirds or bees can lead to ___________ _____________________of the flowers • In other species, speciation can be influenced by __________________ and _________________ © 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 From Speciation to Macroevolution • _________________ is the cumulative effect of many speciation and extinction events © 2011 Pearson Education, Inc.