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MACROEVOLUTION AND SPECIATION How did this flightless bird come to live on the isolated Galápagos Islands? Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Separation of the Species Macroevolution is best observed within the fossil record, Requires the origin of species, also called speciation. Speciation is the final result of changes in gene pool allelic and genotypic frequencies. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 2 Macroevolution: the origin of new taxonomic groups Speciation: the origin of new species 1- Anagenesis (phylogenetic evolution): accumulation of heritable changes 2- Cladogenesis (branching evolution): budding of new species from a parent species that continues to exist (basis of biological diversity) Species Definitions Evolutionary species concept distinguish species from one another based on: Morphological (structural) traits, Biological species concept relies primarily on reproductive isolation rather than trait differences to define a species. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 4 What is a species? Biological species concept (ErnstMayr): a population or group of populations whose members have the potential to interbreed and produce viable, fertile offspring (genetic exchange is possible and that is genetically isolated from other populations) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings How and why do new species originate? Species are created by a series of evolutionary processes populations become isolated geographically isolated reproductively isolated isolated populations evolve independently Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings PREZYGOTIC BARRIERS Reproductive Isolation (isolation of gene pools) Prezygotic barriers: impede mating between species or hinder the fertilization of the ova Habitat (snakes; water/terrestrial) Behavioral (fireflies; mate signaling) Temporal (salmon; seasonal mating) Mechanical (flowers; pollination anatomy) Gametic (frogs; egg coat receptors) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Habitat isolation: Species occur in same region, but occupy different habitats so rarely encounter each other 2 species of garter snake, Thamnophis, occur in same area, but one lives in water & other is terrestrial Leopon http://ahgertz.wordpress.com/2010/07/28/zedonk-isawesome-hybrid-animals/ lions & tigers could hybridize, but they live in different habitats: lions in grasslands tigers in rainforest Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes Eastern spotted skunk (R) & western spotted skunk (L) overlap in range but eastern mates in late winter & western mates in late summer Western spotted skunk (Spilogale gracilis) Eastern spotted skunk (Spilogale putorius) Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers Courtship ritual of bluefooted boobies Video: Albatross Courtship Ritual Video: Giraffe Courtship Ritual Video: Blue-footed Boobies Courtship Ritual Mechanical isolation: Morphological differences can prevent successful mating For many insects, male & female sex organs of closely related species do not fit together, preventing sperm transfer – lack of “fit” between sexual organs: hard to imagine for us… but a big issue for insects with different shaped genitals! Bradybaena with shells spiraling in opposite directions Gametic isolation: Sperm of one species may not be able to fertilize eggs of another species Sea urchins Sea urchins release sperm & eggs into surrounding waters where they fuse & form zygotes. Gametes of different species— red & purple —are unable to fuse. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings POSTZYGOTIC BARRIERS Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult: Zygote mortality (Reduced hybrid viability) Hybrid Sterility (Reduced hybrid fertility) F2 Fitness (Hybrid breakdown) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Zygote mortality (Reduced hybrid viability): Genes of the different parent species may interact and impair the hybrid development Ensatina hybrid Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Hybrid Sterility (Reduced hybrid fertility): Even if hybrids are vigorous, they may be sterile Donkey Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings F2 Fitness (Hybrid breakdown): Some firstgeneration hybrids are fertile, but when they mate with another species or with either parent species, offspring of the next generation are feeble or sterile Hybrid cultivated rice plants with stunted offspring (center) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings REPRODUCTIVE ISOLATION REVIEW Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Speciation can take place with or without a geographic barrier Geographic speciation can occur in two ways: Allopatric speciation Sympatric speciation (a) Allopatric speciation Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings (b) Sympatric speciation Allopatric (Other Country) Speciation In allopatric speciation, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations Evidence of Allopatric Speciation • Regions with many geographic barriers typically have more species than do regions with fewer barriers Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Allopatric Speciation Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Lake male River male Lake female River female a. Sockeye salmon at Pleasure Point Beach, Lake Washington b. Sockeye salmon in Cedar River .The river connects with Lake Washington. Salmon that matured at Pleasure Point Beach do not reproduce with those that matured in the Cedar River. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 23 Adaptive Radiation Adaptive Radiation When members of a species invade several new geographically separate environments The populations become adapted to the different environments Many new species evolve from the single ancestral species This is an example of allopatric speciation Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 24 Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takes place in geographically overlapping populations Caused by chromosomal abnormalities….abnormal number of chromosomes Nonrandom mating (lowers gene flow) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 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 • Many important crops (oats, cotton, potatoes, tobacco, and wheat) are polyploids Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Autopolyploidy Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Allopolyploidy Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Alloploidy Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. seeds diploid banana (2n) Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings no seeds polyploid banana 29 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 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 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 equilibrium 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 Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Fig. 24-17 (a) Punctuated pattern Time (b) Gradual pattern Speciation Rates The punctuated pattern in the fossil record and evidence from lab studies suggests that speciation can be rapid The interval between speciation events can range from 4,000 years (some cichlids) to 40,000,000 years (some beetles), with an average of 6,500,000 years Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Principles of Macroevolution Macroevolution Evolution at the species or higher level of classification Occurs gradually Evolutionists support a gradualistic model Speciation occurs after populations become isolated Each group continuing its own evolutionary pathway The gradualistic model suggests that it is difficult to indicate when speciation occurred Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 34 Principles of Macroevolution Macroevolution Some paleontologists believe that Species can appear quite suddenly Remain essentially unchanged phenotypically during a period of stasis (sameness) until they undergo extinction. Based on these findings, they developed a punctuated equilibrium model to explain the pace of evolution. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 35 Developmental Genes and Macroevolution Genes can bring about radical changes in body shapes and organs. Gene Expression Can Influence Development A change in gene expression could stop developmental process or continue it beyond its normal time. Using modern technology researchers discovered genes whose differential expression can bring about changes in body shapes and organs. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 36 Pax6 Gene and Eye Development (Left): © Carolina Biological Supply/Photo Researchers, Inc.; (Center): © Vol. OS02/PhotoDisc/Getty Images; (Right): © Aldo Brando/Peter Arnold, Inc. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings 37 Evolution is not goal-oriented An evolutionary trend does not mean that evolution is goal-oriented. Surviving species do not represent the peak of perfection. There is compromise & random chance involved as well Remember that for humans as well! Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings Evolution is not the survival of the fittest. Rather it is the survival of the just good enough. From Speciation to Macroevolution Macroevolution is the cumulative effect of many speciation and extinction events Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings