Processes of Evolution Chapter 18 Part 2 18.6 Maintaining Variation Natural selection theory helps explain diverse aspects of nature, including differences between males and females, and the relationship between sickle-cell anemia and malaria Sexual Selection With sexual selection, some version of a trait gives an individual an advantage over others in attracting mates Distinct male and female phenotypes (sexual dimorphism) is one outcome of sexual selection Sexual Selection Balanced Polymorphism Balanced polymorphism • A state in which natural selection maintains two or more alleles at relatively high frequencies • Occurs when environmental conditions favor heterozygotes Example: Sickle cell anemia and malaria • HbA/HbS heterozygotes survive malaria more often than people who make only normal hemoglobin Sickle Cell Anemia and Malaria Fig. 18-13a, p. 287 Fig. 18-13b, p. 287 Fig. 18-13c, p. 287 18.7 Genetic Drift— The Chance Changes Genetic drift • A random change in allele frequencies over time • Can lead to a loss of genetic diversity, especially in small populations Fixation has occurred when all individuals in a population are homozygous for one allele Genetic Drift and Population Size Genetic Drift and Population Size Bottlenecks Bottleneck • A drastic reduction in population size brought about by severe pressure • After a bottleneck, genetic drift is pronounced when a few individuals rebuild a population • Example: Northern elephant seals The Founder Effect Founder effect • Genetic drift is pronounced when a few individuals start a new population Inbreeding • Breeding or mating between close relatives who share a large number of alleles • Example: Old Order Amish in Lancaster County, Pennsylvania (Ellis-van Creveld syndrome) 18.3-18.6 Key Concepts Patterns of Natural Selection Natural selection is a microevolutionary process Depending on the population and its environment, natural selection can shift or maintain the range of variation in heritable traits 18.8 Gene Flow Gene flow • Physical movement of alleles caused by individuals moving into and away from populations • Tends to counter the evolutionary effects of mutation, natural selection, and genetic drift on a population • Example: Movement of acorns by blue jays Gene Flow Between Oak Populations 18.7-18.8 Key Concepts Other Microevolutionary Processes With genetic drift, change can occur in a line of descent by chance alone Gene flow counters the evolutionary effects of mutation, natural selection, and genetic drift 18.9 Reproductive Isolation Speciation • Evolutionary process by which new species form • Reproductive isolating mechanisms are always part of the process Reproductive isolation • The end of gene exchange between populations • Beginning of speciation Four Butterflies, Two Species Reproductive Isolating Mechanisms Reproductive isolating mechanisms prevent interbreeding among species • Heritable aspects of body form, function, or behavior that arise as populations diverge • Prezygotic isolating mechanisms prevent pollination or mating • Postzygotic isolating mechanisms result in weak or infertile hybrids Prezygotic Isolating Mechanisms Temporal isolation Mechanical isolation Behavioral isolation Ecological isolation Gamete incompatibility Mechanical Isolation Behavioral Isolation Animation: Albatross courtship Postzygotic Isolation Mechanisms Reduced hybrid viability (ligers, tigons) • Extra or missing genes Reduced hybrid fertility (mules) • Robust but sterile offspring Hybrid breakdown • Lower fitness with successive generations Reproductive Isolating Mechanisms Different species! Prezygotic isolating mechanisms Temporal isolation: Individuals of different species reproduce at different times. Mechanical isolation: Individuals cannot mate or pollinate because of physical incompatibilities. Behavioral isolation: Individuals of different species ignore or do not get the required cues for sex. Ecological isolation: Individuals of different species live in different places They interbreed and never do meet up. anyway. Gamete incompatibility: Reproductive cells meet up, but no fertilization occurs. Zygotes form, but . . . Postzygotic isolating mechanisms Hybrid inviability: Hybrid embryos die early, or new individuals die before they can reproduce. Hybrid sterility: Hybrid individuals or their offspring do not make functional gametes. No offspring, sterile offspring, or weak offspring that die before reproducing Fig. 18-17, p. 290 Animation: Reproductive isolating mechanisms 18.10 Allopatric Speciation Allopatric speciation • A physical barrier arises and ends gene flow between populations • Genetic divergence results in speciation • Example: llamas, vicunas, and camels Allopatric Speciation The Inviting Archipelagos Winds or ocean currents carry a few individuals of mainland species to remote, isolated islands chains (archipelagos) such as Hawaii Habitats and selection pressures that differ within and between the islands foster divergences that result in allopatric speciation Allopatric Speciation on an Isolated Archipelago A A few individuals of a mainland species reach isolated island 1. In the new habitat, populations of their descendants diverge, and speciation occurs. B Later, a few individuals of a new species colonize nearby island 2. Speciation follows genetic divergence in the new habitat. C Genetically different descendants of the ancestral species may colonize islands 3 and 4 or even invade island 1. Genetic divergence and speciation may follow. Fig. 18-21a, p. 293 Akepa Akekee (L. Nihoa finch Palila(Loxioides Maui parrotbill (Loxops caeruleirostris) (Telespiza (Pseudonestor bailleui) Mamane coccineus) Insects, ultima) xanthophrys) Insects, spiders spiders, some Insects, buds, seeds ripped Rips dry branches from buds nectar; high seeds, flowers, from pods; for insect larvae, twisted apart buds, mountain rain seabird eggs; pupae, caterpillars; by bill, some flowers, forest rocky or mountain forest nectar; high some berries, shrubby slopes with open canopy, mountain rain insects; high dense underbrush forest mountain dry forests Apapane (Himatione sanguinea) Nectar, especially of ohialehua flowers; caterpillars and other insects; spiders; high mountain forests Fig. 18-21b, p. 293 Poouli Maui Alauahio Kauai Amakihi (Melamprosops (Paroreomyza (Hemignathus phaeosoma) montana) kauaiensis) Bark-picker; Tree snails, Bark or leaf insects in insects, some insects, spiders, nectar; high understory; nectar; high last known mountain rain mountain rain forest male died in forest 2004 Akiapolaau (Hemignathus munroi) Probes, digs insects from big trees; high mountain rain forest Akohekohe (Palmeria dolei) Mostly nectar from flowering trees, some insects, pollen; high mountain rain forest Iiwi (Vestiaria coccinea) Mostly nectar (ohia flowers, lobelias, mints), some insects; high mountain rain forest Fig. 18-21c, p. 293 Animation: Allopatric speciation on an archipelago 18.11 Other Speciation Models Populations sometimes speciate even without a physical barrier that blocks gene flow • Sympatric speciation • Parapatric speciation Sympatric Speciation In sympatric speciation, new species form within a home range of an existing species, in the absence of a physical barrier A change in chromosome number (polyploidy) can cause instant speciation On Lord Howe Island, species of palms are reproductively isolated Sympatric Speciation in Wheat T. turgidum (wild emmer) Triticum monococcum (einkorn) 14AA Unknown spontaneous species of chromosome Triticum doubling X 14BB A By 11,000 years ago, humans were cultivating wild wheats. Einkorn has a diploid chromosome number of 14 (two sets of 7). It probably hybridized with another wild wheat species having the same number of chromosomes. 14AB T. tauschii (a wild relative) 28AABB X B About 8,000 years ago, the allopolyploid wild emmer originated from an AB hybrid wheat plant in which the chromosome number doubled. Wild emmer is tetraploid, or AABB; it has two sets of 14 chromosomes. There is recently renewed culinary interest in emmer, also called farro. 14DD T. aestivum (one of the common bread wheats) 42AABBDD C AABB emmer probably hybridized with T. tauschii, a wild relative of wheat. Its diploid chromosome number s 14 (two sets of 7 DD). Common bread wheats have a chromosome number of 42 (six sets of 7 AABBDD). Fig. 18-22, p. 294 Animation: Sympatric speciation in wheat Sympatric Speciation in Palms Parapatric Speciation In parapatric speciation, populations in contact along a common border evolve into distinct species Hybrids in the contact zone are less fit than individuals on either side Parapatric Speciation T. barretti hybrid zone T. anophthalmus Fig. 18-24c, p. 295 Different Speciation Models 18.9-18.11 Key Concepts How Species Arise Speciation varies in its details, but it typically starts after gene flow ends Microevolutionary events that occur independently lead to genetic divergences, which are reinforced as reproductive isolation mechanisms evolve 18.12 Macroevolution Macroevolution • Large-scale patterns of evolutioary change • Includes patterns of change such as one species giving rise to multiple species, the origin of major groups, and major extinction events Coevolution Two species in close ecological contact act as agents of selection on each other (coevolution) • Predator and prey • Host and parasite • Pollinator and flower Over time, the two species may come to depend on each other Coevolution proboscis nectar tube 10 cm Fig. 18-25, p. 296 Stasis and Exaptation Stasis • A lineage exists for millions of years with little or no change (e.g. coelacanth) Exaptation (preadaptation) • Some complex traits in modern species held different adaptive value in ancestral lineages (e.g. feathers in birds and dinosaurs) Adaptive Radiation Adaptive radiation • A burst of speciation that occurs when a lineage encounters a new set of niches Key innovation • A structural or functional adaptation that allows individuals to exploit their habitat in a new way Extinction Extinction • The irrevocable loss of a species from Earth Mass extinctions • Extinctions of many lineages, followed by adaptive radiations • Five catastrophic events in which the majority of species on Earth disappeared Adaptive Radiation of Mammals Following the K-T Extinction Eomaia scansoria Cenozoic Mesozoic Fig. 18-26a, p. 297 rabbits rodents primates armadillos shrews, other insectivores; bats carnivores whales, dolphins deer, other artiodactyls horses, other perissodactyls anteaters platypus, other monotremes kangaroos, other marsupials elephants, other proboscideans manatees Evolutionary Theory Evolutionary biologists try to explain how all species are related by descent from common ancestors Genetic change is the basis of evolution, but many biologists disagree about how it occurs 18.12 Key Concepts Macroevolutionary Patterns Patterns of genetic change that involve more than one species are called macroevolution Recurring patterns of macroevolution include exaptation, adaptive radiation, and extinction Animation: Models of speciation Animation: Morphological differences within a species Animation: Simulation of genetic drift Animation: Temporal isolation among cicadas Video: Rise of the super rats Video: Humpback whales Video: Salamander gills