Chapter 11 Evolution and Its Processes PDF

CHAPTER 11 Evolution and Its Processes FIGURE 11.1 The diversity of life on Earth is the result of evolution, a continuous process that is still occurring. (credit “wolf”: modification of work by Gary Kramer, USFWS; credit “coral”: modification of work by William Harrigan, NOAA; credit “river”: mod...

CHAPTER 11 Evolution and Its Processes FIGURE 11.1 The diversity of life on Earth is the result of evolution, a continuous process that is still occurring. (credit “wolf”: modification of work by Gary Kramer, USFWS; credit “coral”: modification of work by William Harrigan, NOAA; credit “river”: modification of work by Vojtěch Dostál; credit “protozoa”: modification of work by Sharon Franklin, Stephen Ausmus, USDA ARS; credit “fish” modification of work by Christian Mehlführer; credit “mushroom”, “bee”: modification of work by Cory Zanker; credit “tree”: modification of work by Joseph Kranak) CHAPTER OUTLINE 11.1 Discovering How Populations Change 11.2 Mechanisms of Evolution 11.3 Evidence of Evolution 11.4 Speciation 11.5 Common Misconceptions about Evolution INTRODUCTION All species of living organisms—from the bacteria on our skin, to the trees in our yards, to the birds outside—evolved at some point from a different species. Although it may seem that living things today stay much the same from generation to generation, that is not the case: evolution is ongoing. Evolution is the process through which the characteristics of species change and through which new species arise. 248 11 Evolution and Its Processes The theory of evolution is the unifying theory of biology, meaning it is the framework within which biologists ask questions about the living world. Its power is that it provides direction for predictions about living things that are borne out in experiment after experiment. The Ukrainian- born American geneticist Theodosius Dobzhansky famously wrote that “nothing makes sense in 1 biology except in the light of evolution.” He meant that the principle that all life has evolved and diversified from a common ancestor is the foundation from which we understand all other questions in biology. This chapter will explain some of the mechanisms for evolutionary change and the kinds of questions that biologists can and have answered using evolutionary theory. 11.1 Discovering How Populations Change LEARNING OBJECTIVES By the end of this section, you will be able to: Explain how Darwin’s theory of evolution differed from the current view at the time Describe how the present-day theory of evolution was developed Describe how population genetics is used to study the evolution of populations The theory of evolution by natural selection describes a mechanism for species change over time. That species change had been suggested and debated well before Darwin. The view that species were static and unchanging was grounded in the writings of Plato, yet there were also ancient Greeks that expressed evolutionary ideas. In the eighteenth century, ideas about the evolution of animals were reintroduced by the naturalist Georges-Louis Leclerc, Comte de Buffon and even by Charles Darwin’s grandfather, Erasmus Darwin. During this time, it was also accepted that there were extinct species. At the same time, James Hutton, the Scottish naturalist, proposed that geological change occurred gradually by the accumulation of small changes from processes (over long periods of time) just like those happening today. This contrasted with the predominant view that the geology of the planet was a consequence of catastrophic events occurring during a relatively brief past. Hutton’s view was later popularized by the geologist Charles Lyell in the nineteenth century. Lyell became a friend to Darwin and his ideas were very influential on Darwin’s thinking. Lyell argued that the greater age of Earth gave more time for gradual change in species, and the process provided an analogy for gradual change in species. In the early nineteenth century, Jean-Baptiste Lamarck published a book that detailed a mechanism for evolutionary change that is now referred to as inheritance of acquired characteristics. In Lamarck’s theory, modifications in an individual caused by its environment, or the use or disuse of a structure during its lifetime, could be inherited by its offspring and, thus, bring about change in a species. While this mechanism for evolutionary change as described by Lamarck was discredited, Lamarck’s ideas were an important influence on evolutionary thought. The inscription on the statue of Lamarck that stands at the gates of the Jardin des Plantes in Paris describes him as the “founder of the doctrine of evolution.” 1 Theodosius Dobzhansky. “Biology, Molecular and Organismic.” American Zoologist 4, no. 4 (1964): 449. Access for free at openstax.org  11.1 Discovering How Populations Change 249 Charles Darwin and Natural Selection The actual mechanism for evolution was independently conceived of and described by two naturalists, Charles Darwin and Alfred Russell Wallace, in the mid-nineteenth century. Importantly, each spent time exploring the natural world on expeditions to the tropics. From 1831 to 1836, Darwin traveled around the world on H.M.S. Beagle, visiting South America, Australia, and the southern tip of Africa. Wallace traveled to Brazil to collect insects in the Amazon rainforest from 1848 to 1852 and to the Malay Archipelago from 1854 to 1862. Darwin’s journey, like Wallace’s later journeys in the Malay Archipelago, included stops at several island chains, the last being the Galápagos Islands (west of Ecuador). On these islands, Darwin observed species of organisms on different islands that were clearly similar, yet had distinct differences. For example, the ground finches inhabiting the Galápagos Islands comprised several species that each had a unique beak shape (Figure 11.2). He observed both that these finches closely resembled another finch species on the mainland of South America and that the group of species in the Galápagos formed a graded series of beak sizes and shapes, with very small differences between the most similar. Darwin imagined that the island species might be all species modified from one original mainland species. In 1860, he wrote, “Seeing this gradation and diversity of structure in one small, intimately related group of birds, one might really fancy that from an original paucity of birds in this archipelago, one species had been taken and modified 2 for different ends.” FIGURE 11.2 Darwin observed that beak shape varies among finch species. He postulated that the beak of an ancestral species had adapted over time to equip the finches to acquire different food sources. This illustration shows the beak shapes for four species of ground finch: 1. Geospiza magnirostris (the large ground finch), 2. G. fortis (the medium ground finch), 3. G. parvula (the small tree finch), and 4. Certhidea olivacea (the green-warbler finch). Wallace and Darwin both observed similar patterns in other organisms and independently conceived a mechanism to explain how and why such changes could take place. Darwin called this mechanism natural selection. Natural selection, Darwin argued, was an inevitable outcome of three principles that operated in nature. First, the characteristics of organisms are inherited, or passed from parent to offspring. Second, more offspring are produced than are able to survive; in other words, resources for survival and reproduction are limited. The capacity for reproduction in all organisms outstrips the availability of resources to support their numbers. Thus, there is a competition for those resources in each generation. Both Darwin and Wallace’s understanding of this principle came from reading an essay by the economist Thomas Malthus, who discussed this principle in relation to human populations. Third, offspring vary among each other in regard to their characteristics and those variations are inherited. Out of these three principles, Darwin and Wallace reasoned that offspring with inherited characteristics that allow them to best compete for limited resources will survive and have more offspring than those individuals with variations that are less able to compete. Because characteristics are inherited, these traits will be better represented in the next generation. This will lead to change in populations over generations in a process that Darwin 2 Charles Darwin, Journal of Researches into the Natural History and Geology of the Countries Visited during the Voyage of H.M.S. Beagle Round the World, under the Command of Capt. Fitz Roy, R.N, 2nd. ed. (London: John Murray, 1860), http://www.archive.org/details/ journalofresea00darw. 250 11 Evolution and Its Processes called “descent with modification.” Papers by Darwin and Wallace (Figure 11.3) presenting the idea of natural selection were read together in 1858 before the Linnaean Society in London. The following year Darwin’s book, On the Origin of Species, was published, which outlined in considerable detail his arguments for evolution by natural selection. FIGURE 11.3 (a) Charles Darwin and (b) Alfred Wallace wrote scientific papers on natural selection that were presented together before the Linnean Society in 1858. Demonstrations of evolution by natural selection can be time consuming. One of the best demonstrations has been in the very birds that helped to inspire the theory, the Galápagos finches. Peter and Rosemary Grant and their colleagues have studied Galápagos finch populations every year since 1976 and have provided important demonstrations of the operation of natural selection. The Grants found changes from one generation to the next in the beak shapes of the medium ground finches on the Galápagos island of Daphne Major. The medium ground finch feeds on seeds. The birds have inherited variation in the bill shape with some individuals having wide, deep bills and others having thinner bills. Large-billed birds feed more efficiently on large, hard seeds, whereas smaller billed birds feed more efficiently on small, soft seeds. During 1977, a drought period altered vegetation on the island. After this period, the number of seeds declined dramatically: the decline in small, soft seeds was greater than the decline in large, hard seeds. The large-billed birds were able to survive better than the small-billed birds the following year. The year following the drought when the Grants measured beak sizes in the much-reduced population, they found that the average bill size was larger (Figure 11.4). This was clear evidence for natural selection (differences in survival) of bill size caused by the availability of seeds. The Grants had studied the inheritance of bill sizes and knew that the surviving large-billed birds would tend to produce offspring with larger bills, so the selection would lead to evolution of bill size. Subsequent studies by the Grants have demonstrated selection on and evolution of bill size in this species in response to changing conditions on the island. The evolution has occurred both to larger bills, as in this case, and to smaller bills when large seeds became rare. FIGURE 11.4 A drought on the Galápagos island of Daphne Major in 1977 reduced the number of small seeds available to finches, causing many of the small-beaked finches to die. This caused an increase in the finches’ average beak size between 1976 and 1978. Variation and Adaptation Natural selection can only take place if there is variation, or differences, among individuals in a population. Access for free at openstax.org  11.1 Discovering How Populations Change 251 Importantly, these differences must have some genetic basis; otherwise, selection will not lead to change in the next generation. This is critical because variation among individuals can be caused by non-genetic reasons, such as an individual being taller because of better nutrition rather than different genes. Genetic diversity in a population comes from two main sources: mutation and sexual reproduction. Mutation, a change in DNA, is the ultimate source of new alleles or new genetic variation in any population. An individual that has a mutated gene might have a different trait than other individuals in the population. However, this is not always the case. A mutation can have one of three outcomes on the organisms’ appearance (or phenotype): A mutation may affect the phenotype of the organism in a way that gives it reduced fitness—lower likelihood of survival, resulting in fewer offspring. A mutation may produce a phenotype with a beneficial effect on fitness. Many mutations, called neutral mutations, will have no effect on fitness. Mutations may also have a whole range of effect sizes on the fitness of the organism that expresses them in their phenotype, from a small effect to a great effect. Sexual reproduction and crossing over in meiosis also lead to genetic diversity: when two parents reproduce, unique combinations of alleles assemble to produce unique genotypes and, thus, phenotypes in each of the offspring. A heritable trait that aids the survival and reproduction of an organism in its present environment is called an adaptation. An adaptation is a “match” of the organism to the environment. Adaptation to an environment comes about when a change in the range of genetic variation occurs over time that increases or maintains the match of the population with its environment. The variations in finch beaks shifted from generation to generation providing adaptation to food availability. Whether or not a trait is favorable depends on the environment at the time. The same traits do not always have the same relative benefit or disadvantage because environmental conditions can change. For example, finches with large bills were benefited in one climate, while small bills were a disadvantage; in a different climate, the relationship reversed. Patterns of Evolution The evolution of species has resulted in enormous variation in form and function. When two species evolve in different directions from a common point, it is called divergent evolution. Such divergent evolution can be seen in the forms of the reproductive organs of flowering plants, which share the same basic anatomies; however, they can look very different as a result of selection in different physical environments, and adaptation to different kinds of pollinators (Figure 11.5). FIGURE 11.5 Flowering plants evolved from a common ancestor. Notice that the (a) dense blazing star and (b) purple coneflower vary in appearance, yet both share a similar basic morphology. (credit a, b: modification of work by Cory Zanker) In other cases, similar phenotypes evolve independently in distantly related species. For example, flight has evolved in both bats and insects, and they both have structures we refer to as wings, which are adaptations to flight. The wings of bats and insects, however, evolved from very different original structures. When similar structures arise through evolution independently in different species it is called convergent evolution. The wings of bats and insects are called analogous structures; they are similar in function and appearance, but do not share an origin in a common ancestor. Instead they evolved independently in the two lineages. The wings of a hummingbird and an 252 11 Evolution and Its Processes ostrich are homologous structures, meaning they share similarities (despite their differences resulting from evolutionary divergence). The wings of hummingbirds and ostriches did not evolve independently in the hummingbird lineage and the ostrich lineage—they descended from a common ancestor with wings. The Modern Synthesis The mechanisms of inheritance, genetics, were not understood at the time Darwin and Wallace were developing their idea of natural selection. This lack of understanding was a stumbling block to comprehending many aspects of evolution. In fact, blending inheritance was the predominant (and incorrect) genetic theory of the time, which made it difficult to understand how natural selection might operate. Darwin and Wallace were unaware of the genetics work by Austrian monk Gregor Mendel, which was published in 1866, not long after publication of On the Origin of Species. Mendel’s work was rediscovered in the early twentieth century at which time geneticists were rapidly coming to an understanding of the basics of inheritance. Initially, the newly discovered particulate nature of genes made it difficult for biologists to understand how gradual evolution could occur. But over the next few decades genetics and evolution were integrated in what became known as the modern synthesis—the coherent understanding of the relationship between natural selection and genetics that took shape by the 1940s and is generally accepted today. In sum, the modern synthesis describes how evolutionary pressures, such as natural selection, can affect a population’s genetic makeup, and, in turn, how this can result in the gradual evolution of populations and species. The theory also connects this gradual change of a population over time, called microevolution, with the processes that gave rise to new species and higher taxonomic groups with widely divergent characters, called macroevolution. Population Genetics Recall that a gene for a particular character may have several variants, or alleles, that code for different traits associated with that character. For example, in the ABO blood type system in humans, three alleles determine the particular blood-type carbohydrate on the surface of red blood cells. Each individual in a population of diploid organisms can only carry two alleles for a particular gene, but more than two may be present in the individuals that make up the population. Mendel followed alleles as they were inherited from parent to offspring. In the early twentieth century, biologists began to study what happens to all the alleles in a population in a field of study known as population genetics. Until now, we have defined evolution as a change in the characteristics of a population of organisms, but behind that phenotypic change is genetic change. In population genetic terms, evolution is defined as a change in the frequency of an allele in a population. Using the ABO system as an example, the frequency of one of the alleles, IA, is the number of copies of that allele divided by all the copies of the ABO gene in the population. For example, a study in 3 Jordan found a frequency of IA to be 26.1 percent. The IB, I0 alleles made up 13.4 percent and 60.5 percent of the alleles respectively, and all of the frequencies add up to 100 percent. A change in this frequency over time would constitute evolution in the population. There are several ways the allele frequencies of a population can change. One of those ways is natural selection. If a given allele confers a phenotype that allows an individual to have more offspring that survive and reproduce, that allele, by virtue of being inherited by those offspring, will be in greater frequency in the next generation. Since allele frequencies always add up to 100 percent, an increase in the frequency of one allele always means a corresponding decrease in one or more of the other alleles. Highly beneficial alleles may, over a very few generations, become “fixed” in this way, meaning that every individual of the population will carry the allele. Similarly, detrimental alleles may be swiftly eliminated from the gene pool, the sum of all the alleles in a population. Part of the study of population genetics is tracking how selective forces change the allele frequencies in a population over time, which can give scientists clues regarding the selective forces that may be operating on a given population. The studies of changes in wing coloration in the peppered moth from mottled white to dark in response to soot-covered tree trunks and then back to mottled white when factories stopped producing so much soot is a classic example of studying evolution in natural populations (Figure 11.6). 3 Sahar S. Hanania, Dhia S. Hassawi, and Nidal M. Irshaid, “Allele Frequency and Molecular Genotypes of ABO Blood Group System in a Jordanian Population,” Journal of Medical Sciences 7 (2007): 51-58, doi:10.3923/jms.2007.51.58 Access for free at openstax.org  11.1 Discovering How Populations Change 253 FIGURE 11.6 As the Industrial Revolution caused trees to darken from soot, darker colored peppered moths were better camouflaged than the lighter colored ones, which caused there to be more of the darker colored moths in the population. In the early twentieth century, English mathematician Godfrey Hardy and German physician Wilhelm Weinberg independently provided an explanation for a somewhat counterintuitive concept. Hardy’s original explanation was in response to a misunderstanding as to why a “dominant” allele, one that masks a recessive allele, should not increase in frequency in a population until it eliminated all the other alleles. The question resulted from a common confusion about what “dominant” means, but it forced Hardy, who was not even a biologist, to point out that if there are no factors that affect an allele frequency those frequencies will remain constant from one generation to the next. This principle is now known as the Hardy-Weinberg equilibrium. The theory states that a population’s allele and genotype frequencies are inherently stable—unless some kind of evolutionary force is acting on the population, the population would carry the same alleles in the same proportions generation after generation. Individuals would, as a whole, look essentially the same and this would be unrelated to whether the alleles were dominant or recessive. The four most important evolutionary forces, which will disrupt the equilibrium, are natural selection, mutation, genetic drift, and migration into or out of a population. A fifth factor, nonrandom mating, will also disrupt the Hardy-Weinberg equilibrium but only by shifting genotype frequencies, not allele frequencies (unless the allele contributes toward increased or decreased reproductive potential). In nonrandom mating, individuals are more likely to mate with individuals with specific phenotypes rather than at random. Since nonrandom mating does not change allele frequencies, it does not cause evolution directly. Natural selection has been described. Mutation creates one allele out of another one and changes an allele’s frequency by a small, but continuous amount each generation. Each allele is generated by a low, constant mutation rate that will slowly increase the allele’s frequency in a population if no other forces act on the allele. If natural selection acts against the allele, it will be removed from the population at a low rate leading to a frequency that results from a balance between selection and mutation. This is one reason that genetic diseases remain in the human population at very low frequencies. If the allele is favored by selection, it will increase in frequency. Genetic drift causes random changes in allele frequencies when populations are small. Genetic drift can often be important in evolution, as discussed in the next section. Finally, if two populations of a species have different allele frequencies, migration of individuals between them will cause frequency changes in both populations. As it happens, there is no population in which one or more of these processes are not operating, so populations are always evolving, and the Hardy-Weinberg equilibrium will never be exactly observed. However, the Hardy-Weinberg principle gives scientists a baseline expectation for allele frequencies in a non-evolving population to which they can compare evolving populations and thereby infer what evolutionary forces might be at play. The population is evolving if the frequencies of alleles or genotypes deviate from the value expected from the Hardy-Weinberg principle. Darwin identified a special case of natural selection that he called sexual selection. Sexual selection affects an individual’s ability to mate and thus produce offspring, and it leads to the evolution of dramatic traits that often appear maladaptive in terms of survival but persist because they give their owners greater reproductive success. Sexual selection occurs in two ways: through intrasexual selection, as male–male or female–female competition for mates, and through intersexual selection, as female or male selection of mates. Male–male competition takes the form of conflicts between males, which are often ritualized, but may also pose significant threats to a male’s survival. Sometimes the competition is for territory, with females more likely to mate with males with higher quality territories. Female choice occurs when females choose a male based on a particular trait, such as feather colors, the performance of a mating dance, or the building of an elaborate structure. In some cases male–male competition and female choice combine in the mating process. In each of these cases, the traits selected for, such as fighting ability or feather color and length, become enhanced in the males. In general, it is thought that sexual selection can 254 11 Evolution and Its Processes proceed to a point at which natural selection against a character’s further enhancement prevents its further evolution because it negatively impacts the male’s ability to survive. For example, colorful feathers or an elaborate display make the male more obvious to predators. 11.2 Mechanisms of Evolution LEARNING OBJECTIVES By the end of this section, you will be able to: Describe the four basic causes of evolution: natural selection, mutation, genetic drift, and gene flow Explain how each evolutionary force can influence the allele frequencies of a population The Hardy-Weinberg equilibrium principle says that allele frequencies in a population will remain constant in the absence of the four factors that could change them. Those factors are natural selection, mutation, genetic drift, and migration (gene flow). In fact, we know they are probably always affecting populations. Natural Selection Natural selection has already been discussed. Alleles are expressed in a phenotype. Depending on the environmental conditions, the phenotype confers an advantage or disadvantage to the individual with the phenotype relative to the other phenotypes in the population. If it is an advantage, then that individual will likely have more offspring than individuals with the other phenotypes, and this will mean that the allele behind the phenotype will have greater representation in the next generation. If conditions remain the same, those offspring, which are carrying the same allele, will also benefit. Over time, the allele will increase in frequency in the population. Mutation Mutation is a source of new alleles in a population. Mutation is a change in the DNA sequence of the gene. A mutation can change one allele into another, but the net effect is a change in frequency. The change in frequency resulting from mutation is small, so its effect on evolution is small unless it interacts with one of the other factors, such as selection. A mutation may produce an allele that is selected against, selected for, or selectively neutral. Harmful mutations are removed from the population by selection and will generally only be found in very low frequencies equal to the mutation rate. Beneficial mutations will spread through the population through selection, although that initial spread is slow. Whether or not a mutation is beneficial or harmful is determined by whether it helps an organism survive to sexual maturity and reproduce. It should be noted that mutation is the ultimate source of genetic variation in all populations—new alleles, and, therefore, new genetic variations arise through mutation. Genetic Drift Another way a population’s allele frequencies can change is genetic drift (Figure 11.7), which is simply the effect of chance. Genetic drift is most important in small populations. Drift would be completely absent in a population with infinite individuals, but, of course, no population is this large. Genetic drift occurs because the alleles in an offspring generation are a random sample of the alleles in the parent generation. Alleles may or may not make it into the next generation due to chance events including mortality of an individual, events affecting finding a mate, and even the events affecting which gametes end up in fertilizations. If one individual in a population of ten individuals happens to die before it leaves any offspring to the next generation, all of its genes—a tenth of the population’s gene pool—will be suddenly lost. In a population of 100, that 1 individual represents only 1 percent of the overall gene pool; therefore, it has much less impact on the population’s genetic structure and is unlikely to remove all copies of even a relatively rare allele. Imagine a population of ten individuals, half with allele A and half with allele a (the individuals are haploid). In a stable population, the next generation will also have ten individuals. Choose that generation randomly by flipping a coin ten times and let heads be A and tails be a. It is unlikely that the next generation will have exactly half of each allele. There might be six of one and four of the other, or some different set of frequencies. Thus, the allele frequencies have changed and evolution has occurred. A coin will no longer work to choose the next generation (because the odds are no longer one half for each allele). The frequency in each generation will drift up and down on what is known as a random walk until at one point either all A or all a are chosen and that allele is fixed from that point on. This could take a very long time for a large population. This simplification is not very biological, but it can be shown that real populations behave this way. The effect of drift on frequencies is greater the smaller a population Access for free at openstax.org  11.2 Mechanisms of Evolution 255 is. Its effect is also greater on an allele with a frequency far from one half. Drift will influence every allele, even those that are being naturally selected. VISUAL CONNECTION FIGURE 11.7 Genetic drift in a population can lead to the elimination of an allele from a population by chance. In each generation, a random set of individuals reproduces to produce the next generation. The frequency of alleles in the next generation is equal to the frequency of alleles among the individuals reproducing. Do you think genetic drift would happen more quickly on an island or on the mainland? Genetic drift can also be magnified by natural or human-caused events, such as a disaster that randomly kills a large portion of the population, which is known as the bottleneck effect that results in a large portion of the gene pool suddenly being wiped out (Figure 11.8). In one fell swoop, the genetic structure of the survivors becomes the genetic structure of the entire population, which may be very different from the pre-disaster population. The disaster must be one that kills for reasons unrelated to the organism’s traits, such as a hurricane or lava flow. A mass killing caused by unusually cold temperatures at night, is likely to affect individuals differently depending on the alleles they possess that confer cold hardiness. FIGURE 11.8 A chance event or catastrophe can reduce the genetic variability within a population. 256 11 Evolution and Its Processes Another scenario in which populations might experience a strong influence of genetic drift is if some portion of the population leaves to start a new population in a new location, or if a population gets divided by a physical barrier of some kind. In this situation, those individuals are unlikely to be representative of the entire population which results in the founder effect. The founder effect occurs when the genetic structure matches that of the new population’s founding fathers and mothers. The founder effect is believed to have been a key factor in the genetic history of the Afrikaner population of Dutch settlers in South Africa, as evidenced by mutations that are common in Afrikaners but rare in most other populations. This is likely due to a higher-than-normal proportion of the founding colonists, which were a small sample of the original population, carried these mutations. As a result, the population expresses unusually high incidences of Huntington’s disease (HD) and Fanconi anemia (FA), a genetic disorder known to cause 4 bone marrow and congenital abnormalities, and even cancer. LINK TO LEARNING Visit this site (http://openstax.org/l/genetic_drift2) to learn more about genetic drift and to run simulations of allele changes caused by drift. Gene Flow Another important evolutionary force is gene flow, or the flow of alleles in and out of a population resulting from the migration of individuals or gametes (Figure 11.9). While some populations are fairly stable, others experience more flux. Many plants, for example, send their seeds far and wide, by wind or in the guts of animals; these seeds may introduce alleles common in the source population to a new population in which they are rare. FIGURE 11.9 Gene flow can occur when an individual travels from one geographic location to another and joins a different population of the species. In the example shown here, the brown allele is introduced into the green population. 11.3 Evidence of Evolution LEARNING OBJECTIVES By the end of this section, you will be able to: Explain sources of evidence for evolution Define homologous and vestigial structures The evidence for evolution is compelling and extensive. Looking at every level of organization in living systems, biologists see the signature of past and present evolution. Darwin dedicated a large portion of his book, On the Origin of Species, identifying patterns in nature that were consistent with evolution and since Darwin our understanding has become clearer and broader. Fossils Fossils provide solid evidence that organisms from the past are not the same as those found today; fossils show the gradual evolutionary changes over time. Scientists determine the age of fossils and categorize them all over the world to determine when the organisms lived relative to each other. The resulting fossil record tells the story of the past, and shows the evolution of form over millions of years (Figure 11.10). For example, highly detailed fossil 4 A. J. Tipping et al., “Molecular and Genealogical Evidence for a Founder Effect in Fanconi Anemia Families of the Afrikaner Population of South Africa,” PNAS 98, no. 10 (2001): 5734-5739, doi: 10.1073/pnas.091402398. Access for free at openstax.org  11.3 Evidence of Evolution 257 records have been recovered for sequences of species in the evolution of whales and modern horses. The fossil record of horses in North America is especially rich and many contain transition fossils: those showing intermediate anatomy between earlier and later forms. The fossil record extends back to a dog-like ancestor some 55 million years ago that gave rise to the first horse-like species 55 to 42 million years ago in the genus Eohippus. The series of fossils tracks the change in anatomy resulting from a gradual drying trend that changed the landscape from a forested one to a prairie. Successive fossils show the evolution of teeth shapes and foot and leg anatomy to a grazing habit, with adaptations for escaping predators, for example in species of Mesohippus found from 40 to 30 million years ago. Later species showed gains in size, such as those of Hipparion, which existed from about 23 to 2 million years ago. The fossil record shows several adaptive radiations in the horse lineage, which is now much reduced to only one genus, Equus, with several species. FIGURE 11.10 This illustration shows an artist’s renderings of these species derived from fossils of the evolutionary history of the horse and its ancestors. The species depicted are only four from a very diverse lineage that contains many branches, dead ends, and adaptive radiations. One of the trends, depicted here is the evolutionary tracking of a drying climate and increase in prairie versus forest habitat reflected in forms that are more adapted to grazing and predator escape through running. Przewalski's horse is one of a few living species of horse. Anatomy and Embryology Another type of evidence for evolution is the presence of structures in organisms that share the same basic form. For example, the bones in the appendages of a human, dog, bird, and whale all share the same overall construction (Figure 11.11). That similarity results from their origin in the appendages of a common ancestor. Over time, evolution led to changes in the shapes and sizes of these bones in different species, but they have maintained the same overall layout, evidence of descent from a common ancestor. Scientists call these synonymous parts homologous structures. Some structures exist in organisms that have no apparent function at all, and appear to be residual parts from a past ancestor. For example, some snakes have pelvic bones despite having no legs because they descended from reptiles that did have legs. These unused structures without function are called vestigial structures. Other examples of vestigial structures are wings on flightless birds (which may have other functions), leaves on some cacti, traces of pelvic bones in whales, and the sightless eyes of cave animals. FIGURE 11.11 The similar construction of these appendages indicates that these organisms share a common ancestor.