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Chapter 24 The Origin of Species Species Hummingbirds of Costa Rica Species Biological Species Concept A population whose members can potentially interbreed in nature to produce viable, fertile offspring In practice, we recognize two populations as separate species if each has distinguishing characteristics relative to the other, and members of each population lack the potential to interbreed in nature with members of the other population Reproductive Barriers Isolate populations from one another A complete barrier would result in no gene flow An incomplete barrier could still reduce gene flow Reproductive Barriers Pre-zygotic vs. post-zygotic A complete barrier would result in no gene flow An incomplete barrier could still reduce gene flow Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Eurasian spoonbill Roseate spoonbill Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation North America Madagascar Central America Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Ecological isolation Rhagoletis pomonella Hawthorns or apples for oviposition Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Ecological isolation Behavioral (mate choice) isolation Galapagos Islands Red-footed booby Blue-footed boobies Satin Bowerbirds & their bowers Australia Tooth-billed Catbird’s leaf display Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Ecological isolation Behavioral (mate choice) isolation Temporal isolation Dog-day (17 year) Periodical (12 year) Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Ecological isolation Behavioral (mate choice) isolation Temporal isolation Mechanical isolation C. wapleri Thompson Creek, W. Feliciana Parish Reproductive Barriers Pre-zygotic Habitat isolation Geographic isolation Ecological isolation Behavioral (mate choice) isolation Temporal isolation Mechanical isolation Gametic isolation head acrosome tail midpiece ovum Buffon (1811) Reproductive Barriers Post-zygotic Reduced hybrid viability Reduced hybrid fertility Hybrid sterility (in the extreme) ♀ x = ♂ sterile Reproductive Barriers Post-zygotic Reduced hybrid viability Reduced hybrid fertility Hybrid sterility (in the extreme) Hybrid breakdown (reduced viability or fertility occurs in the offspring of hybrids) Reproductive Barriers Habitat isolation Temporal isolation Individuals of different species HABITAT ISOLATION Behavioral isolation Mechanical isolation Mating attempt TEMPORAL ISOLATION Figure 24.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings BEHAVIORAL ISOLATION MECHANICAL ISOLATION Reproductive Barriers Gametic isolation Reduce hybrid viability Fertilization GAMETIC ISOLATION REDUCED HYBRID VIABILITY Figure 24.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Reduce hybrid fertility Hybrid breakdown Viable fertile offspring REDUCED HYBRID FERTILITY HYBRID BREAKDOWN Speciation The process by which new species form Two requirements: Reproductive isolation of populations (gene flow sufficiently reduced) Genetic divergence (divergent evolution) Two main modes: Allopatric vs. sympatric Allopatric speciation Divergence occurs in geographic isolation Sympatric speciation Divergence occurs despite lack of geographic isolation Allopatric vs. sympatric speciation Figure 24.5 Sympatric speciation Rhagoletis pomonella Hawthorns or apples for oviposition Sympatric speciation Divergence occurs despite lack of geographic isolation Polyploids Organisms with extra sets of chromosomes; can result in sympatric speciation in one generation The origin of a polyploid individual is usually a mistake in meiosis or mitosis Autopolyploidy vs. allopolyploidy Sympatric speciation An autopolyploid Is an individual that has > two chromosome sets, all from a single species Failure of cell division in a cell of a growing diploid plant produces a tetraploid branch. Gametes produced by flowers on this branch will be diploid. If offspring are viable and fertile, it is a new biological species. 2n 2n = 6 4n = 12 Figure 24.8 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4n Sympatric speciation An allopolyploid Is an individual with > two chromosome sets, derived from different species Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Species A 2n = 4 Unreduced gamete with 7 chromosomes Viable fertile hybrid Meiotic error; chromosome number not reduced from 2n to n 2n = 10 Normal gamete n=3 Species B 2n = 6 Figure 24.9 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Normal gamete n=3 Polyploidy is especially common in plants From Micro- to MacroEvolution Microevolution – Population and species-level evolution, including speciation Macroevolution – Speciation and all broader-scale evolutionary patterns and processes Anagenesis Sufficient microevolutionary changes may occur that the earlier population is considered a separate species from the later population Figure 24.2 time Cladogenesis Branching evolution occurs as speciation events accumulate. The formation of each new branch is a macroevolutionary event. Anagenesis Sufficient microevolutionary changes may occur that the earlier population is considered a separate species from the later population Figure 24.2 time Cladogenesis Branching evolution occurs as speciation events accumulate The formation of each new branch is a macroevolutionary event Broad patterns across the resulting tree are also within the scope of macroevolution Adaptive Radiation One species gives rise to many others in a geologically short period of time E.g., Darwin’s finches Islands are good places to find adaptive radiations Adaptive Radiation E.g., Hawaiian silverswords Figure 24.12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Tempo of Evolution time Gradualism Figure 24.13 Punctuated Equilibrium Tempo of Evolution time Gradualism Figure 24.13 Punctuated Equilibrium Evolutionary Novelties Evolutionary novelties usually arise as modifications of existing traits Figure 24.14 Evolutionary Development (Evo-Devo) Slight alterations in development can have profound effects on adults Heterochrony = change in the rate or timing of development, relative to an ancestor E.g., paedomorphosis = heterochronic retention of juvenile structures in adulthood Evolutionary Development (Evo-Devo) Slight alterations in development can have profound effects on adults Heterotopy = change in the physical position of a trait during development, relative to an ancestor E.g., feathers from one part of a bird’s body may become expressed on another part Evolutionary Development (Evo-Devo) Allometric growth = relative growth rates of different body parts within a species during development Evolutionary Development (Evo-Devo) Allometric growth = relative growth rates of different body parts within a species during development E.g., allometric growth in horned scarab beetles; certain clusters of cells grow faster than others Evolutionary Development (Evo-Devo) Evolutionary Development (Evo-Devo) Phanaeus imperator Evolution is not “goal oriented” Evolution is a genetic response to the interaction between the individuals of a population and their current environment (which includes other individuals) Natural selection can only act on the phenotypic variation present, and selection can only cause evolutionary change if phenotypic variation results (at least in part) from genetic variation Evolution is not “goal oriented” Therefore, even if female tigers in a population would prefer blue males, blue tigers will never evolve if the population lacks variation that includes blue fur YES! But he exists only in my dreams. He’s not blue, but he’s better than nothing!