Speciation and Macroevolution Chapter 20 Learning Objective 1 • What is the biological species concept? • List two potential problems with the concept Biological Species Concept • Species • • • • one or more populations members interbreed in nature produce fertile offspring do not interbreed with different species Sterile Hybrid Problems • Biological species concept applies only to sexually reproducing organisms • Individuals assigned to different species may occasionally successfully interbreed KEY CONCEPTS • According to the biological species concept, a species consists of individuals that can successfully interbreed with one another but not with individuals from other species Learning Objective 2 • What is the significance of reproductive isolating mechanisms? • Distinguish among different prezygotic and postzygotic barriers Reproductive Isolating Mechanisms • Restrict gene flow between species • Prezygotic barriers • • prevent fertilization from taking place Postzygotic barriers • prevent gene flow after fertilization has taken place (reproductive isolating mechanisms) Prezygotic Barriers (1) • Temporal isolation • • two species reproduce at different times of day, season, or year Habitat isolation • two closely related species live and breed in different habitats in same geographic area Temporal Isolation Fig. 20-2a, p. 430 Fig. 20-2b, p. 430 Leopard frog Mating activity Wood frog March 1 April 1 May 1 Fig. 20-2c, p. 430 Insert “Temporal isolation among cicadas” temporal_isolation.swf Prezygotic Barriers (2) • Behavioral isolation • • distinctive courtship behaviors prevent mating between species Mechanical isolation • incompatible structural differences in reproductive organs of similar species Behavioral Isolation Mechanical Isolation Prezygotic Barriers (3) • Gametic isolation • gametes from different species are incompatible because of molecular and chemical differences Postzygotic Barriers (1) • Hybrid inviability • • interspecific embryos die during development Hybrid sterility • prevents interspecific hybrids that survive to adulthood from reproducing successfully Hybrid Sterility Postzygotic Barriers (2) • Hybrid breakdown • prevents offspring of hybrids that survive to adulthood and successfully reproduce from reproducing beyond one or a few generations KEY CONCEPTS • The evolution of different species begins with reproductive isolation, in which two populations are no longer able to interbreed successfully Insert “Reproductive isolating mechanisms” isolating_mechanisms_m.swf Explore reproductive isolation by clicking on the figures in ThomsonNOW Learning Objective 3 • What is allopatric speciation? • Give an example Allopatric Speciation (1) • Evolution of a new species • • from ancestral population Population becomes geographically isolated from rest of species • subsequently diverges Allopatric Speciation (2) • More likely to occur if original isolated population is small • • makes genetic drift more significant Examples: • • • Death Valley pupfishes Kaibab squirrels Porto Santo rabbits Allopatric Speciation KEY CONCEPTS • In allopatric speciation, populations diverge into different species due to geographic isolation, or physical separation Insert “Allopatric speciation on an archipelago” archipelago.swf Explore allopatric speciation by clicking on the figures in ThomsonNOW. Learning Objective 4 • What is sympatric speciation? • Give plant and animal examples Sympatric Speciation • Does not require geographic isolation • More common in plants than animals Sympatric Speciation in Plants • Usually results from allopolyploidy • • polyploid individual (>2 sets of chromosomes) is hybrid derived from two species Examples: • • kew primroses hemp nettles Allopolyploidy in Plants Species A 2n=6 Species B 2n=4 P generation n=2 n=3 Gametes Hybrid AB F1 generation Fig. 20-9a, p. 435 No doubling of chromosome number Doubling of chromosome number 2 n = 10 Chromosomes either cannot pair or go through erratic meiosis Pairing now possible during meiosis n=5 No gametes or sterile gametes — no sexual reproduction possible Viable gametes — sexual reproduction possible (selffertilization) Fig. 20-9b, p. 435 Species A 2n=6 Species B 2n=4 P generation n=2 n=3 Gametes Hybrid AB F1 generation No doubling of chromosome number Doubling of chromosome number 2 n = 10 Chromosomes either Pairing now cannot pair or go through possible erratic meiosis n=5 during meiosis No gametes or sterile gametes — no sexual reproduction possible Viable gametes — sexual reproduction possible (selffertilization) Stepped Art Fig. 20-9b, p. 435 Sympatric Speciation Sympatric Speciation in Animals • Fruit maggot flies Sympatric Speciation in Animals • Cichlids Fig. 20-12a, p. 437 Fig. 20-12b, p. 437 Fig. 20-12c, p. 437 KEY CONCEPTS • In sympatric speciation, populations become reproductively isolated from one another despite living in the same geographic area Insert “Sympatric speciation in wheat” wheat_speciation.swf Explore sympatric speciation by clicking on the figure in ThomsonNOW. Learning Objective 5 • Debate the pace of evolution by representing the views of either punctuated equilibrium or gradualism Evolution • Punctuated equilibrium model • • • evolution proceeds in spurts short periods of active speciation interspersed with long periods of stasis Gradualism model • populations slowly diverge from one another by accumulation of adaptive characteristics Punctuated Equilibrium and Gradualism Stasis Slow, gradual changes Extinction of original species Stasis Stasis Time Time Divergence is sudden, with rapid changes Stasis (little change) Structural changes Divergence is gradual Structural changes Fig. 20-13, p. 438 KEY CONCEPTS • Speciation may require millions of years but sometimes occurs much more quickly Learning Objective 6 • What is macroevolution? Macroevolution • Large-scale phenotypic changes in populations • • in taxonomic groups species level and higher new species, genera, families, orders, classes, phyla, kingdoms, or domains KEY CONCEPTS • The evolution of species and higher taxa is known as macroevolution Learning Objective 7 • Discuss novel features of macroevolution, including preadaptations, allometric growth, and paedomorphosis Macroevolution • Includes evolutionary novelties • • due to changes during development Slight changes in regulatory genes • cause major structural changes in organism Preadaptations • Structures originally fulfilled one role • • changed and adapted for different role Example: feathers Allometric Growth • Varied rates of growth for different parts of body • • causes overall changes in shape of organism Examples: • • ocean sunfish male fiddler crab Allometric Growth • Ocean sunfish Tail approx. 1 mm Newly hatched ocean sunfish Adult ocean sunfish Fig. 20-14a, p. 440 Fig. 20-14b, p. 440 Fig. 20-14b, p. 440 Paedomorphosis • Juvenile characteristics retained in adult • • due to changes in timing of development Example: adult axolotl salamanders • with external gills and tail fins Paedomorphosis • Salamander Learning Objective 8 • What is the macroevolutionary significance of adaptive radiation and extinction? Adaptive Radiation (1) • Diversification of ancestral species into many new species • Adaptive zones • new ecological opportunities not exploited by ancestral organism Adaptive Radiation (2) • When many adaptive zones are empty • • colonizing species diversify and exploit them Example: Hawaiian honeycreepers and silverswords • after ancestors colonized Hawaiian Islands Adaptive Radiation • Hawaiian honeycreepers Rips away bark to find insects Maui parrot bill Kauai Sips flower nectar ‘I‘iwi Oahu Forages among leaves and branches Maui creeper Maui Chisels holes in bark to get insects Akiapolaau Extinct Sipped flower nectar Black mamo Extinct Habits unknown Ula-ai-hawane Hawaii Picks food from cracks in the bark Akialoa Sips flower nectar Apapane Feeds on snails and Sips flower nectar invertebrates Poo-uli Crested honeycreeper Fig. 20-16, p. 441 Adaptive Radiation • Hawaiian silverswords Extinction (1) • Death of a species • When species become extinct • • adaptive zones they occupied become vacant allows other species to evolve and fill zones Extinction (2) • Background extinction • • continuous, low-level extinction of species Mass extinction • • extinction of numerous species, higher taxonomic groups in both terrestrial and marine environments Mass Extinction Common ancestor of birds and saurischians Archosaur common ancestor Fig. 20-18a, p. 443 Birds Saurischians (dinosaurs, extinct) Ornithischians (dinosaurs, extinct) Pterosaurs (flying reptiles, extinct) Crocodilians (alligators, crocodiles) Fig. 20-18b, p. 443 Paleozoic Permian Triassic Mesozoic Jurassic Theropods (carnivorous saurischians) Archosaurs Cenozoic Cretaceous Tertiary / Quaternary Birds Sauropods (herbivorous saurischians) Common ancestor Stegosaurs and other ornithischians Crocodilians Pterosaurs Crocodiles Fig. 20-18b, p. 443