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CHAPTER 14 ORIGIN OF SPECIES
Figure 14.3B Habitat isolation: Courtship ritual in blue-footed boobies as a behavioral barrier between species.
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14.1 The origin of species is the source of biological diversity
Speciation is the emergence of new species Every time speciation occurs, the diversity of life increases The many millions of species on Earth have all arisen from an ancestral life form that lived around 3.6 billion years ago Copyright © 2009 Pearson Education, Inc.
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Figure 14.1 How speciation increases diversity.
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14.2 There are several ways to define a species
Taxonomy is the branch of biology that names and classifies species and groups them into broader categories Carolus Linnaeus developed the binomial system of naming organisms using physical characteristics to distinguish over 11,000 species Similarities between some species and variation within species can make defining species difficult Copyright © 2009 Pearson Education, Inc.
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14.2 There are several ways to define a species
The biological species concept defines a species as a population or group of populations whose members have the potential to interbreed in nature and produce fertile offspring Reproductive isolation prevents gene flow and maintains separate species Copyright © 2009 Pearson Education, Inc.
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14.2 There are several ways to define a species
The morphological species concept classifies organisms based on observable phenotypic traits It can be applied to asexual organisms, fossils, and in cases when we donít know about possible interbreeding There is some subjectivity in deciding which traits to use Copyright © 2009 Pearson Education, Inc.
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14.2 There are several ways to define a species
The ecological species concept defines a species by its ecological role or niche Consider the cichlids, which are similar in appearance but feed at different depths in the lake Copyright © 2009 Pearson Education, Inc.
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14.2 There are several ways to define a species
The phylogenetic species concept defines a species as a set of organisms representing a specific evolutionary lineage Morphological or DNA similarities or differences can be used to define a species Defining the amount of difference required to distinguish separate species is a problem Copyright © 2009 Pearson Education, Inc.
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14.3 Reproductive barriers keep species separate
Reproductive barriers serve to isolate a species gene pool and prevent interbreeding Reproductive barriers are categorized as prezygotic or postzygotic, depending on whether they function before or after zygotes form Copyright © 2009 Pearson Education, Inc.
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Table 14.3 Reproductive Barriers between Species.
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14.3 Reproductive barriers keep species separate
Prezygotic Barriers Prezygotic barriers prevent mating or fertilization between species In temporal isolation, two species breed at different times (seasons, times of day, years) In habitat isolation, two species live in the same general area but not in the same kind of place Copyright © 2009 Pearson Education, Inc.
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14.3 Reproductive barriers keep species separate
Prezygotic Barriers In behavioral isolation, there is little or no sexual attraction between species, due to specific behaviors In mechanical isolation, female and male sex organs are not compatible In gametic isolation, female and male gametes are not compatible Copyright © 2009 Pearson Education, Inc.
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14.3 Reproductive barriers keep species separate
Postzygotic Barriers Postzygotic barriers operate after hybrid zygotes are formed In reduced hybrid viability, most hybrid offspring do not survive In reduced hybrid fertility, hybrid offspring are vigorous but sterile In hybrid breakdown, the first-generation hybrids are viable and fertile, but the offspring of the hybrids are feeble or sterile The process of speciation depends on whether reproductive barriers prevent gene flow between populations Copyright © 2009 Pearson Education, Inc.
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Figure 14.3E Reduced hybrid fertility: A horse (above left) and a donkey (above right) may produce a hybrid, sterile offspring, a mule.
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14.4 In allopatric speciation, geographic isolation leads to speciation
In allopatric speciation, populations of the same species are geographically separated, separating their gene pools Changes in the allele frequencies of each population may be caused by natural selection, genetic drift, and mutation, unaffected by gene flow from other populations Copyright © 2009 Pearson Education, Inc.
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14.4 In allopatric speciation, geographic isolation leads to speciation
Gene flow between populations is initially prevented by a geographic barrier The Grand Canyon and Colorado River separate two species of antelope squirrels Copyright © 2009 Pearson Education, Inc.
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A. harrisi A. leucurus South North
Figure 14.4 Allopatric speciation of geographically isolated antelope squirrels. Ask students for other examples of suitable barriers, then make the point that the size and nature of the barrier vary from species to species. South North
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14.4 In allopatric speciation, geographic isolation leads to speciation
Likelihood of allopatric speciation increases when a population is small and isolated A small population may have a different gene pool due to the founder effect Genetic drift and natural selection may have a greater effect in a small population in a new habitat Copyright © 2009 Pearson Education, Inc.
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14.5 In sympatric speciation, speciation takes place without geographic isolation
In sympatric speciation, new species may arise within the same geographic area as a parent species Gene flow between populations may be reduced by factors such as polyploidy, habitat differentiation, or sexual selection Copyright © 2009 Pearson Education, Inc.
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14.5 In sympatric speciation, speciation takes place without geographic isolation
Many plant species have evolved by polyploidy, the multiplication of the chromosome number due to errors in cell division A tetraploid (4n) plant can arise from a diploid parent Copyright © 2009 Pearson Education, Inc.
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Self- fertilization Parent species 2n = 6 Tetraploid cells 4n = 12
3 1 Self- fertilization Parent species 2n = 6 Tetraploid cells 4n = 12 Viable, fertile tetraploid species 4n = 12 Figure 14.5A Sympatric speciation by polyploidy within a single species. Consider customizing Figure 14.5A with the following labels: Label “diploid gametes”: A failure of cell division after chromosome duplication doubles a cell’s chromosomes. Label “viable, fertile tetraploid species”: Self-fertilization produces tetraploid zygotes that can self or mate with other 4n plants. Point out to your students that the result of polyploidy is instant speciation. The tetraploids cannot produce fertile offspring by mating with parent plants The result would be 3n offspring, which are sterile. Diploid gametes 2n = 6
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14.5 In sympatric speciation, speciation takes place without geographic isolation
Most polyploids arise from hybridization of two different species Haploid gametes from two different species combine to produce a sterile hybrid Why is the hybrid sterile? How can it reproduce? Chromosome duplications may produce a fertile polyploid species What is the chromosome number of the new species Copyright © 2009 Pearson Education, Inc.
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Chromosomes not homologous (cannot pair) Species A Gamete 2n = 4 n = 2
1 2 Species A 2n = 4 Gamete n = 2 3 Sterile hybrid n = 5 Viable, fertile hybrid species 2n = 10 Figure 14.5B Sympatric speciation producing a hybrid polyploid from two different species. The hybrid is sterile because its chromosomes cannot pair during meiosis. The hybrid may reproduce asexually. The polyploid’s chromosome number is equal to the sum of the diploid chromosome numbers of the parent species. Point out to your students that the new species is reproductively isolated from both parent species. Gamete n = 3 Species B 2n = 6
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14.5 In sympatric speciation, speciation takes place without geographic isolation
Sympatric speciation in animals more commonly occurs through habitat differentiation and sexual selection Remember the cichlids in Lake Victoria! Copyright © 2009 Pearson Education, Inc.
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14.6 EVOLUTION CONNECTION: Most plant species trace their origin to polyploid speciation
80% of all living plant species are the descendants of ancestors that formed by polyploid speciation Hybridization between two species accounts for most of these species What advantage might there be to hybridization? Polyploid food plants include oats, potatoes, bananas, peanuts, barley, plums, apples, sugarcane, coffee, and bread wheat Cotton is also polyploid Copyright © 2009 Pearson Education, Inc.
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14.6 EVOLUTION CONNECTION: Most plant species trace their origin to polyploid speciation
Wheat has been domesticated for 11,000 years It is the most widely cultivated plant in the world Bread wheat, Triticum aestivum, is a polyploid with 42 chromosomes Copyright © 2009 Pearson Education, Inc.
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Figure 14.6B The evolution of wheat.
AA BB Wild Triticum (14 chromo- somes) Triticum monococcum (14 chromosomes) 1 Hybridization AB Sterile hybrid (14 chromosomes) 2 Cell division error and self-fertilization AA BB DD T. turgidum Emmer wheat (28 chromosomes) T. tauschii (wild) (14 chromosomes) 3 Hybridization Figure 14.6B The evolution of wheat. Uppercase letters represent sets of chromosomes, not genes. Consider customizing Figure 14.6B with the following labels: Label first cross: A cultivated wheat (AA) hybridized with a weed (BB). Label first offspring: The AB hybrid was sterile. Label emmer wheat: An error in cell division and self-fertilization led to a new species (AABB). Label 2nd hybridization: The new species hybridized with a wild relative, producing a sterile hybrid (ABD). Label bread wheat: An error in cell division and self-fertilization led to bread wheat (AABBDD). ABD Sterile hybrid (21 chromosomes) 4 Cell division error and self-fertilization AA BB DD T. aestivum Bread wheat (42 chromosomes)
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14.8 Hybrid zones provide opportunities to study reproductive isolation
What happens when isolated populations renew contact? In hybrid zones, members of different species meet and mate to produce hybrid offspring Copyright © 2009 Pearson Education, Inc.
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New species Ancestral species Gene flow
3 Hybrid zone 1 2 4 Gene flow Gene flow Figure 14.8A Formation of a hybrid zone. Four populations are connected by gene flow. A barrier to gene flow separates one population, which has diverged from the other three. Gene flow is reestablished in the hybrid zone. Hybrid Population (five individuals are shown) Barrier to gene flow
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Reinforcement Fusion Stability
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allopatric population Pied flycatcher from sympatric population
populations Sympatric populations Male collared flycatcher Male pied flycatcher Figure 14.8B Reinforcement of reproductive barriers. Pied flycatcher from allopatric population Pied flycatcher from sympatric population
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14.8 Hybrid zones provide opportunities to study reproductive isolation
What may happen in a hybrid zone? Reinforcement: If hybrids are less fit than parent species, natural selection strengthens reproductive barriers Fusion: Weak reproductive barriers between the two species, with considerable gene flow, reverses speciation and two species become one again Copyright © 2009 Pearson Education, Inc.
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14.8 Hybrid zones provide opportunities to study reproductive isolation
Stability: Many hybrid zones are stable, continuing to produce hybrids; this allows some gene flow between populations, but each species maintains its own integrity Which of these three outcomes—reinforcement, fusion, or stability—is happening to the Pundamilia species of cichlids in Lake Victoria? Copyright © 2009 Pearson Education, Inc.
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14.9 TALKING ABOUT SCIENCE: Peter and Rosemary Grant study the evolution of Darwin’s finches
Peter and Rosemary Grant have worked on medium ground finches on tiny, isolated, uninhabited Daphne Major in the Galapágos Islands for 35 years Medium ground finches and cactus finches occasionally interbreed Hybrid offspring have intermediate bill sizes and survive well during wet years, when there are plenty of soft, small seeds around During dry years, hybrids are outcompeted by both parental types Copyright © 2009 Pearson Education, Inc.
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14.9 TALKING ABOUT SCIENCE: Peter and Rosemary Grant study the evolution of Darwin’s finches
The occasional hybridization between finch species introduces new genes into both populations During drought years, hybrids die out This keeps medium ground finches and cactus finches on separate evolutionary paths Copyright © 2009 Pearson Education, Inc.
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Tool-using insect-eater (woodpecker finch)
Cactus-seed-eater (cactus finch) Figure 14.10A Examples of differences in beak shape and size in Galápagos finches, each adapted for a specific diet. Tool-using insect-eater (woodpecker finch) Seed-eater (medium ground finch)
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14.10 Adaptive radiation may occur when new opportunities arise
In adaptive radiation, many diverse species evolve from a common ancestor Adaptive radiations occur When a few organisms colonize new unexploited areas After a mass extinction Adaptive radiations are linked to new opportunities: lack of competitors, varying habitats and food sources, evolution of new structures Copyright © 2009 Pearson Education, Inc.
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14.11 Speciation may occur rapidly or slowly
What is the total length of time between speciation events (between formation of a species and subsequent divergence of that species)? In a survey of 84 groups of plants and animals, the time ranged from 4,000 to 40 million years Overall, the time between speciation events averaged 6.5 million years and rarely took less than 50,000 years Copyright © 2009 Pearson Education, Inc.
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Time Figure 14.11A Punctuated equilibrium model.
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Figure 14.11B Gradualism model.
Time
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You should now be able to
Explain how the diverse assemblage of cichlid species evolved in Lake Victoria; explain why many of these species no longer exist Compare the definitions, advantages, and disadvantages of the different species concepts Describe five types of prezygotic barriers and three types of postzygotic barriers that prevent populations belonging to closely related species from interbreeding Copyright © 2009 Pearson Education, Inc.
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You should now be able to
Explain how geographical processes can fragment populations and lead to speciation Explain how sympatric speciation can occur, noting examples in plants and animals Explain why polyploidy is important to modern agriculture; explain how modern wheat evolved Explain how reproductive barriers might evolve in isolated populations of organisms Copyright © 2009 Pearson Education, Inc.
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You should now be able to
Explain how hybrid zones are useful in the study of reproductive isolation Explain the conditions that can lead to adaptive radiation Copyright © 2009 Pearson Education, Inc.
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