Part III Evolution
Darwin and Evolution
Chapter
14
Outline
	14.1	Darwin’s Theory of Evolution
	•	Charles Darwin’s trip around the Southern Hemisphere aboard the HMS Beagle provided him with evidence that the Earth is very old and that evolution does occur.219
	•	Darwin said that today’s species are descended from common ancestors and that natural selection is a mechanism for adaptation to the environment.221
	•	Alfred Wallace came to the same conclusions as Darwin did regarding evolution.223
	14.2	Evidence for Evolution
	•	The fossil record, biogeography, comparative anatomy, and comparative biochemistry support the hypothesis of common descent.224–27
What do the many breeds of dogs, the rare buttercups of New Zealand, and a child’s resistant ear infection have in common? EVOLUTION! Without evolution—change in a line of descent over time—we wouldn’t see such a great variety of living things about us. But aside from its many benefits, evolution also sometimes causes problems for humans. 
Some bacteria have evolved to the point that they are resistant to the antibiotics once successfully used to cure the diseases they cause. For example, antibiotics originally cured bacterial ear infections within a few days. Unseen, however, were the one or two bacteria with just the right mutation to resist a particular drug. All the descendants of these bacteria were also resistant, causing the antibiotic to be useless as a cure for this type of ear infection. The antibiotic is considered the selective agent because it allowed the resistant bacteria to flourish while killing their relatives.
What was the selective agent for the many breeds of dogs available as pets today? Humans were, of course. Over the years, humans selected which dogs to mate to produce today’s varieties. This process is called artificial selection. 
In nature, the selective agent is the environment, and the process is called natural selection. Many microenvironments in New Zealand plus periodic isolation allowed 14 alpine species of buttercups to evolve at different times, but now they are found side by side throughout the same area. In fact, there are more species of buttercups on the two islands of New Zealand than on any of the continents. 
This chapter explains how different types of selection lead to evolution and begins by describing the early studies that contributed to our current knowledge of the evolutionary process, particularly the work of Charles Darwin.
14.1		Darwin’s Theory of Evolution
Charles Darwin, the father of evolution, was only 22 in 1831 when he accepted the position of naturalist aboard the HMS Beagle, a British naval ship about to sail around the world (Fig. 14.1). Darwin had a suitable background for his position. He was an ardent student of nature and had long been a collector of insects. His sensitive nature had prevented him from studying medicine, and he went to divinity school at Cambridge instead. Even so, he attended many lectures in both biology and geology, and he was also tutored in these subjects by a friend and teacher, the Reverend John Henslow. Darwin spent the summer of 1831 doing fieldwork with Adam Sedgwick, a geologist at Cambridge, before Henslow recommended him to the captain of the Beagle as a naturalist. The trip was to take five years, and the ship was to traverse the Southern Hemisphere, where we now know that life is most abundant and varied. Along the way, Darwin encountered forms of life very different from those in his native England.
Darwin’s major mission was to expand the navy’s knowledge of natural resources (e.g., water and food) in foreign lands. The captain also hoped Darwin would find evidence of the biblical account of creation as it was interpreted at that time. Instead, by the time the trip was over, Darwin had made observations that were inconsistent with current beliefs, and these observations led him to conclude that biological evolution occurs. 
Before Darwin
Prior to Darwin, lay people had an entirely different way of looking at the world (Table 14.1). They believed that the Earth was only a few thousand years old and that since the time of creation, species had remained exactly the same. Even so, studying the anatomy of organisms and then classifying them interested many investigators who wished to show that species were created to be suitable to their environment. Explorers and collectors traveled the world and brought back currently existing species to be classified and also fossils, the remains of once-living organisms often found in strata. Strata are layers of rock or sedimentary material (Fig. 14.2a). Figure 14.3 shows how strata come about in the sea. Even if they form in the sea, they can later be found on land. 
A noted zoologist at the time, Georges Cuvier, founded the science of paleontology, the study of fossils. Cuvier was faced with a problem. He believed in the fixity of species; yet, the Earth’s strata clearly showed a succession of different life-forms over time. To explain these observations, he hypothesized that a local catastrophe had caused a mass extinction whenever a new stratum of that region showed a new mix of fossils. After each catastrophe, the region was repopulated by species from surrounding areas, and this accounted for the appearance of new fossils in the new stratum. The result of all these catastrophes was change appearing over time. Some of Cuvier’s followers, who came to be called catastrophists, even suggested that worldwide catastrophes had occurred and that after each of these events, new sets of species had been created. 
In contrast to Cuvier, Jean-Baptiste de Lamarck, another biologist, hypothesized that evolution occurs, and he also said adaptation to the environment was the cause of diversity. Therefore, after studying the succession of life-forms in strata, Lamarck concluded that more complex organisms are descended from less complex organisms. To explain the process of adaptation to the environment, Lamarck supported the idea of inheritance of acquired characteristics, which proposes that use and disuse of a structure can bring about inherited change. One example Lamarck gave—and the one for which he is most famous—is that the long neck of a giraffe developed over time because animals stretched their necks to reach food high in trees and then passed on a long neck to their offspring (see Fig. 14.8). However, his hypothesis for the inheritance of acquired characteristics has never been substantiated by experimentation. For example, if acquired characteristics were inherited, people who become blind by accident would have blind children, and circumcised males would have boys that lack a foreskin from birth. Modern genetics explains why the idea of acquired characteristics cannot be substantiated. Phenotypic changes acquired during an organism’s lifetime do not result in genetic changes that can be passed to subsequent generations.
Darwin’s ideas were close to those of Lamarck. For example, Darwin said that living things share common characteristics because they have a common ancestry. One of the most unfortunate interpretations of this statement was that we are descended from apes. For Darwin, however, humans and apes share a common ancestry just as, say, you and your cousins can trace your ancestry back to the same grandparents. In contrast to Lamarck, Darwin’s observations led him to conclude that species are suited to the environment through no will of their own but by natural selection. He saw the process of natural selection as the means by which different species come about (see Fig. 14.8). 
Darwin’s Conclusions
Darwin’s conclusions that organisms are related through common descent and that adaptation to various environments results in diversity were based on several types of data, including his study of geology and fossils and also his study of biogeography. Biogeography is the distribution of life-forms on Earth.
Darwin’s Study of Geology and Fossils
Darwin took Charles Lyell’s Principles of Geology on the Beagle voyage. This book presented arguments to support a theory of geological change proposed by James Hutton. In contrast to the catastrophists, Hutton believed the Earth was subject to slow but continuous cycles of erosion and uplift. Weathering causes erosion; thereafter, dirt and rock debris are washed into the rivers and transported to oceans. When these loose sediments are deposited, strata results (see Fig. 14.3a). Then the strata, which often contain fossils, are uplifted from below sea level to form land. Hutton concluded that extreme geological changes can be accounted for by slow, natural processes, given enough time. Lyell went on to propose a theory of uniformitarianism, which stated that these slow changes occurred at a uniform rate. Hutton’s general ideas about slow and continual geological change are still accepted today, although modern geologists realize that rates of change have not always been uniform. Darwin was not a believer in the idea of uniform change, but he was convinced, as was Lyell, that the Earth’s massive geological changes are the result of slow processes, and that the Earth, therefore, must be very old.
On his trip, Darwin observed massive geological changes firsthand. When he explored what is now Argentina, he saw raised beaches for great distances along the coast. When he got to the Andes Mountains, he was impressed by their great height. In Chile, he found fossil marine shells inland, well above sea level, and witnessed the effects of an earthquake that caused the land to rise several feet. While Darwin was making geological observations, he also collected fossil specimens that differed somewhat from modern species (Fig. 14.4). Once Darwin accepted the supposition that the Earth must be very old, he began to think that there would have been enough time for descent with modification—that is, evolution, to occur. Living forms could be descended from extinct forms known only from the fossil record. It would seem that species were not fixed; instead, they change over time.
Darwin’s Study of Biogeography 
Darwin could not help but compare the animals of South America to those with which he was familiar in England. For example, instead of rabbits, he found the Patagonian hare in the grasslands of South America. The Patagonian hare has long legs and ears but the face of a guinea pig, a rodent native to South America (Fig. 14.5). Did the Patagonian hare resemble a rabbit because the two types of animals were adapted to the same type of environment—an outcome we call convergent evolution today? Both animals ate grass, hid in bushes, and moved rapidly using long hind legs. Did the Patagonian hare have the face of a guinea pig because of common descent with guinea pigs?
As he sailed southward along the eastern coast of the continent of South America, Darwin saw how similar species replaced each other. For example, the greater rhea (an ostrichlike bird) found in the north was replaced by the lesser rhea in the south. Therefore, Darwin reasoned that related species could be modified according to the environment. When he got to the Galápagos Islands, he found further evidence of this. The Galápagos Islands are a small group of volcanic islands off the western coast of South America. The few types of plants and animals found there were slightly different from species Darwin had observed on the mainland, and even more important, they also varied from island to island according to the particular environments.
FinchesAlthough some of the the finches on the Galápagos Islands seemed like mainland finches, others were quite different (Fig. 14.6). Today, there are ground-dwelling finches with different-sized beaks, depending on the size of the seeds they feed on, and a cactus-eating finch with a more pointed beak. The beak size of the tree-dwelling finches also varies, according to the size of their insect prey. The most unusual of the finches is a woodpecker-type finch. This bird has a sharp beak to chisel through tree bark but lacks the woodpecker’s long tongue that probes for insects. To make up for this, the bird carries a twig or cactus thorn in its beak and uses it to poke into crevices. Once an insect emerges, the finch drops this tool and seizes the insect with its beak.
Later, Darwin speculated as to whether all the different species of finches he had seen could have descended from a mainland finch. In other words, he wondered if a mainland finch was the common ancestor to all the types on the Galápagos Islands. Had speciation, formation of a new species, occurred because the islands allowed isolated populations of birds to evolve independently? Could the present-day species have resulted from accumulated changes occurring within each of these isolated populations?
TortoisesEach of the Galápagos Islands also seemed to have its own type of tortoise, and Darwin began to wonder if this could be correlated with a difference in vegetation among the islands. Long-necked tortoises seemed to inhabit only dry areas, where food was scarce, and most likely the longer neck was helpful in reaching cactuses. In moist regions with relatively abundant ground foliage, short-necked tortoises were found. Had an ancestral tortoise given rise to these different types, each adapted to a different environment? An adaptation is any characteristic that makes an organism more suited to its environment.
Natural Selection and Adaptation
Once Darwin decided that adaptations develop over time, he began to think about a mechanism by which adaptations might arise. Eventually, he proposed natural selection as the mechanism. Natural selection is a process in which steps 1–3 result in a population adapted to the environment (4):
	1.	The members of a population have heritable variations (Fig. 14.7).
	2.	The population produces more offspring than the resources of an environment can support.
	3.	The more adapted individuals survive and reproduce to a greater extent than those that lack the adaptations.
	4.	Across generations, a larger proportion of the population becomes adapted to the environment.
Notice that because natural selection utilizes only variations that happen to be provided by genetic changes, it lacks any directedness or anticipation of future needs. Natural selection is an ongoing process because the environment of living things is constantly changing. Extinction (loss of a species) can occur when previous adaptations are no longer suitable to a changed environment.
Organisms Have Variations
With reference to  step 1, Darwin emphasized that the members of a population vary in their functional, physical, and behavioral characteristics. Before Darwin, variations were considered imperfections that should be ignored since they were not important to the description of a species (see Table 14.1). Darwin, on the other hand, believed variations were essential to the natural selection process. Darwin suspected—but did not have the evidence we have today—that the occurrence of variations is completely random; they arise by accident and for no particular purpose. Also, new variations are more likely to be harmful than beneficial to an organism.
The variations that make adaptation to the environment possible are those that are passed on from generation to generation. The science of genetics was not yet well established, so Darwin was never able to determine the cause of variations or how they are passed on. Today, we realize that genes, together with the environment, determine the phenotypes of an organism. Mutations, along with chromosomal rearrangements and assortment of chromosomes during meiosis and fertilization, can cause new variations to arise.
Organisms Struggle to Exist
With reference to step 2, in Darwin’s time a socioeconomist named Thomas Malthus stressed the reproductive potential of human beings. He proposed that death and famine are inevitable because the human population tends to increase faster than the supply of food. Darwin applied this concept to all organisms and saw that the available resources were not sufficient to allow all members of a population to survive. He calculated the reproductive potential of elephants. Assuming a life span of about 100 years and a breeding span of 30 to 90 years, a single female probably bears no fewer than six young. If all these young were to survive and continue to reproduce at the same rate, after only 750 years, the descendants of a single pair of elephants would number about 19 million!
Each generation has the same reproductive potential as the previous generation. Therefore, there is a constant struggle for existence, and only certain members of a population survive and reproduce each generation.
Organisms Differ in Fitness
With reference to step 3, fitness is the reproductive success of an individual relative to other members of a population, in a particular environment. The most-fit individuals are the ones that capture a disproportionate amount of resources, and convert these resources into a larger number of viable offspring. Because organisms vary anatomically and physiologically, and because the challenges of local environments vary as well, what determines fitness varies for different populations. For example, among western diamondback rattlesnakes living on lava flows, the most fit are those that are black in color. But among those living on desert soil, the most fit are those with the typical light and dark-brown coloring. Background matching helps an animal both capture prey and avoid being captured; therefore, it is expected to lead to survival and increased reproduction.
Darwin noted that when humans help carry out artificial selection, breeders select animals with particular traits to reproduce. For example, prehistoric humans probably noted desirable variations among wolves and selected particular animals for breeding. Therefore, the desired traits increased in frequency in the next generation. This same process was -repeated many times over. The result today is the existence of many varieties of dogs, all descended from the wolf. In a similar way, several varieties of vegetables can be traced to a single ancestor. Chinese cabbage, brussels sprouts, and kohlrabi are all derived from a single species, Brassica oleracea. 
In nature, interactions with the environment determine which members of a population reproduce to a greater degree than other members. In contrast to artificial selection, the result of natural selection is not predesired. Natural selection occurs because certain members of a population happen to have a variation that allows them to survive and reproduce to a greater extent than other members. For example, any variation that increases the speed of a hoofed animal helps it escape predators and live longer; a variation that reduces water loss is beneficial to a desert plant; and one that increases the sense of smell helps a wild dog find its prey. Therefore, we expect the organisms with these characteristics to have increased fitness. Figure 14.8 contrasts Lamarck’s ideas with those of Darwin. 
Organisms Become Adapted
With reference to step 4 (see page 221), it takes time for adaptations to evolve. We rarely observe the process, but we can observe the end result. We can especially recognize an adaptation when unrelated organisms living in a particular environment display similar characteristics. For example, manatees, penguins, and sea turtles all have flippers, which help them move through the water—also an example of convergent evolution. Adaptations also account for why organisms are able to escape their predators (Fig. 14.9) or why they are suited to their way of life (Fig. 14.10). Natural selection causes adaptive traits to be increasingly represented in each succeeding generation. There are other processes of evolution aside from natural selection (see Chapter 15), but natural selection is the only process that results in adaptation to the environment.
Darwin and Wallace
After the HMS Beagle returned to England in 1836, Darwin waited more than 20 years to publish his book On the Origin of Species. During the intervening years, he used the scientific process to test his hypothesis that today’s diverse life-forms arose by descent from a common ancestor and that natural selection is a mechanism by which species can change and new species can arise. Darwin was prompted to publish his book after reading a similar hypothesis formulated by Alfred Russel Wallace. 
Wallace (1823–1913) was an English naturalist who, like Darwin, was a collector at home and abroad. He went on collecting trips, each of which lasted several years, to the Amazon and the Malay Archipelago. After studying the animals on every major island, he divided these islands into a western group, with animals like those of the Orient, and an eastern group, with animals like those of Australia. The dividing line between the islands is a narrow but deep strait now known as the Wallace Line. 
While traveling, Wallace also wrote an essay called “On the Law Which Has Regulated the Introduction of New Species.” In this essay he said that “every species has come into existence coincident both in time and space with a preexisting closely allied species.” A year later, after reading Malthus’s treatise on human population increase, Wallace conceived the idea of “survival of the fittest.” He quickly completed an essay proposing natural selection as an agent for evolutionary change and sent it to Darwin for comment. Darwin was stunned. Here was the hypothesis he had formulated, but had never dared to publish. He told his friend and colleague, Charles Lyell, that Wallace’s ideas were so similar to his own that even Wallace’s “terms now stand as heads of my chapters.”
Darwin suggested that Wallace’s paper be published immediately, even though Darwin himself as yet had nothing in print. Lyell and others who knew of Darwin’s detailed work substantiating the process of natural selection suggested that a joint paper be read to the Linnean Society, a renowned scientific society. The title of Wallace’s section was “On the Tendency of Varieties to Depart Indefinitely from the Original Type.” Darwin presented an abstract of a paper he had written in 1844 and an abstract of On the Origin of Species, which was published in 1859. 
14.2		Evidence for Evolution
The Theory of Evolution states that all living things have a common ancestor, but each is adapted to a particular way of life. Many lines of evidence consistently support the hypothesis that organisms are related through common descent. A hypothesis becomes a scientific theory only when a variety of evidence made by independent investigators supports the hypothesis. The theory of evolution is a unifying principle in biology because it can explain so many different observations in various fields of biology. The theory of evolution has the same status in biology that the germ theory of disease has in medicine.
Darwin cited much of the evidence for evolution we will discuss, except that he had no knowledge of the biochemical data that became available after his time.
Fossil Evidence
The fossil record is the history of life recorded by remains from the past. Fossils are at least 10,000 years old and include such items as pieces of bone, impressions of plants pressed into shale, and even insects trapped in tree resin (which we know as amber). For the last three centuries, paleontologists have studied fossils in the Earth’s strata all over the world and have pieced together the story of past life.
The fossil record is rich in information. One of its most striking patterns is a succession of life-forms from the simple to the more complex. Occasionally this pattern is reversed, showing that evolution is not unidirectional. Catas-trophists offered an explanation for the extinction and subsequent replacement of one group of organisms by another group, but they never could explain successive changes that link groups of organisms historically. Particularly interesting are the fossils that serve as transitional links between groups. Even in Darwin’s day, scientists knew of the Archaeopteryx fossils, which are intermediate between reptiles and birds (Fig. 14.11). The dinosaur-like skeleton of these fossils has reptilian features, including jaws with teeth, and a long, jointed tail, but Archaeopteryx also had feathers and wings.
Other transitional links between fossil vertebrates have more recently been found, including the amphibious fish Eustheopteron, the reptile-like amphibian Seymouria, and the mammal-like reptiles, or therapsids. These fossils allow us to deduce that fishes evolved before amphibians, which evolved before reptiles, which evolved before both birds and mammals, in the history of life. Recently, four-legged aquatic mammals, the mesonychids, were discovered that provide important insights concerning the evolution of whales from land-living, hoofed ancestors (Fig. 14.12). The fossilized whale Ambulocetus may have been amphibious, walking on land and still swimming in the sea. Rodhocetus swam with an up-and-down tail motion like modern whales do; its reduced hind limbs could not have helped in swimming.  
Biogeographical Evidence
Biogeography is the study of the distribution of plants and animals in different places throughout the world. Such distributions are consistent with the hypothesis that, when forms are related, they evolved in one locale and then spread to accessible regions. Therefore, you would expect a different mix of plants and animals whenever geography separates continents, islands, or seas. As previously mentioned, Darwin noted that South America lacked rabbits, even though the environment was quite suitable for them. He concluded that no rabbits lived in South America because rabbits evolved somewhere else and had no means of reaching South America.
To take another example, both cactuses and euphorbia are plants adapted to a hot, dry environment—both are succulent, spiny, flowering plants. Why do cactuses grow in North American deserts and euphorbia grow in African deserts, when each would do well on the other continent? It seems obvious that they just happened to evolve on their respective continents.
The islands of the world are home to many unique species of animals and plants that are found no place else, even when the soil and climate are the same. Why do so many species of finches live on the Galápagos Islands when these same species are not on the mainland? The reasonable explanation is that an ancestral finch migrated to all the different islands. Then geographic isolation allowed the ancestral finch to evolve into a different species on each island.
Also, long ago South America, Antarctica, and Australia were connected (see Fig. 16.13a). Marsupials (pouched mammals) and placental mammals arose at this time, but today marsupials are plentiful only in Australia and placental mammals are plentiful in South America. Why are marsupials only plentiful in Australia (Fig. 14.13)? After marsupials arose, Australia separated and drifted away, and marsupials were free to evolve into many different forms because they had no competition from placental mammals. In the Americas, the placentals were able to successfully compete against the marsupials, and the opossum is the only marsupial in the Americas. In some cases, placental and marsupial mammals physically resemble each other—for example, the pouched marsupial mouse and the harvest mouse, the marsupial mole and the common mole, the marsupial wombat and the marmot, and the tasmanian wolf and the wolf. This supports the hypothesis that evolution is influenced by the environment and by the mix of plants and animals in a particular continent—that is, by biogeography, not by design.
Anatomical Evidence
Darwin was able to show that a common descent hypothesis offers a plausible explanation for vestigial structures and anatomical similarities among organisms. 
Vestigial structures are anatomical features that are fully developed in one group of organisms but reduced and nonfunctional in other similar groups. Most birds, for example, have well-developed wings used for flight. However, some bird species (e.g., ostrich) have greatly reduced wings and do not fly. -Similarly, whales (see Fig. 14.12) and snakes have no use for hind limbs, and yet extinct whales and even some present-day snakes have remnants of a pelvic girdle and legs. Humans have a -tailbone but no tail. The presence of vestigial structures can be explained by the common descent hypothesis. Vestigial structures occur because organisms inherit their anatomy from their ancestors; they are traces of an organism’s evolutionary history.
Vertebrate forelimbs are used for flight (birds and bats), orientation during swimming (whales and seals), running (horses), climbing (arboreal lizards), or swinging from tree branches (monkeys). Yet all vertebrate forelimbs contain the same sets of bones organized in similar ways, despite their dissimilar functions (Fig. 14.14). The most plausible explanation for this unity is that the basic forelimb plan belonged to a common ancestor, and then the plan became modified in the succeeding groups as each continued along its own evolutionary pathway. Anatomically similar structures explainable by inheritance from a common ancestor are called homologous structures. In contrast, analogous structures serve the same function, but are not constructed similarly, and therefore could not have a common ancestry. The wings of birds and insects are analogous structures. 
The homology shared by vertebrates extends to their embryological development (Fig. 14.15). At some time during development, all vertebrates have a postanal tail and exhibit paired pharyngeal pouches. In fishes and amphibian larvae, these pouches develop into functioning gills. In humans, the first pair of pouches becomes the cavity of the middle ear and the auditory tube. The second pair becomes the tonsils, while the third and fourth pairs become the thymus and parathyroid glands. Why should pharyngeal pouches, which have lost their original function, develop and then become modified in terrestrial vertebrates? The most likely explanation is that fishes are ancestral to other vertebrate groups. Anatomical and developmental homologies are independent evidences of a shared common ancestor and an evolutionary relationship between groups of organisms. 
Biochemical Evidence
Almost all living organisms use the same basic biochemical molecules, including DNA (deoxyribonucleic acid), ATP (adenosine triphosphate), and many identical or nearly identical enzymes. Further, organisms utilize the same DNA triplet code and the same 20 amino acids in their proteins. Now that we know the sequence of DNA bases in the genomes of many organisms, it has become clear that humans share a large number of genes with much simpler organisms. 
Also of interest, evo-devo researchers (evolutionary developmental biologists) have found that many developmental genes are shared in animals ranging from worms to humans. It appears that life’s vast diversity has come about by only slight differences in the same genes. The result has been widely divergent body plans. For example, a similar gene in arthropods and vertebrates determines the back to front axis. But, although the base sequences are similar, the genes have opposite effects. In arthropods, such as fruit flies and crayfish, the nerve cord is toward the front, whereas in vertebrates, such as chicks and humans, the nerve cord is toward the back.
When the degree of similarity in DNA base sequences or the degree of similarity in amino acid sequences of proteins is examined, the data are as expected, assuming common descent. Cytochrome c is a molecule used in the electron transport chain of all the organisms shown in Figure 14.16. Data regarding differences in the amino acid sequence of cytochrome c show that the sequence in a human differs from that in a monkey by only one amino acid, from that in a duck by 11 amino acids, and from that in Candida, a yeast, by 51 amino acids. These data are consistent with other data regarding the anatomical similarities of these organisms, and therefore their relatedness.
The Chapter in Review
Summary
14.1		Darwin’s Theory of Evolution
Charles Darwin took a position as a naturalist aboard the HMS Beagle and made a trip around the world, largely in the Southern Hemisphere.
Before Darwin
Before Darwin, people believed that the Earth was young, that species didn’t change, that any variations were imperfections, and that observations could substantiate this view.
	•	Cuvier was an early paleontologist who believed that species do not change. He observed species come and go in the fossil record, and he said that these changes were due to catastrophies. 
	•	Lamarck was a zoologist who hypothesized that evolution and adaptation to the environment do occur. He suggested that acquired characteristics could be inherited. For example, he said that giraffes stretched their necks to reach food in trees, and then this change was inherited by the next generation.
Darwin’s Conclusions
Darwin’s conclusions based on geology and fossils are:
	•	The Earth is very old, giving time for evolution to occur.
	•	Living things are descended from extinct forms known only from the fossil record.
Darwin’s conclusions based on biogeography are:
	•	Living things evolve where they are. This explains, for example, why South America has the Patagonian hare, while England has the rabbit. 
	•	Living things are adapted to local environments. This explains why there are many types of finches and tortoises in the Galápagos Islands.
Natural Selection and Adaptation 
According to Darwin, the result of natural selection is a population adapted to its local environment:
Darwin and Wallace
Alfred Russel Wallace was a naturalist who, like Darwin, traveled to other continents in the Southern Hemisphere. He also collected evidence of common descent, and his reading of Malthus caused him to develop the same mechanism for adaptation (natural selection) as Darwin. Darwin’s work was more thorough, as evidenced by his book On the Origin of Species.
14.2		Evidence for Evolution
A theory in science is a concept supported by much evidence, and the Theory of Evolution is supported by several types of evidence:
	•	Fossil record indicates the history of life in general and allows us to trace the descent of a particular group. 
	•	Biogeography shows that the distribution of organisms on Earth is explainable by assuming organisms evolved in one locale. 
	•	Anatomy and development of organisms reveals homologies explainable by common ancestry. 
	•	Biochemical molecules of the same type occur in all organisms. Differences indicate the degree of relatedness. 
Thinking Scientifically
	1.	Recent research indicates that genes involved in male reproduction (sperm production and morphology) evolve at a faster rate than other genes in primates. Explain why this makes sense, based on natural selection.
	2.	Geneticists often compare DNA sequences of genes among organisms to determine their rate of evolution, called the molecular clock. The molecular clock has been found to run at different rates for different genes. That is, some genes evolve faster than others. Explain why the DNA sequence in one gene might change faster over time than that in another gene.
Testing Yourself
Choose the best answer for each question.
	1.	_________ developed the idea that acquired characteristics can be inherited.
	a. Darwin	d. Sedgwick
	b. Lamarck	e. Cuvier
	c. Hutton
	2.	The major feature Darwin noticed that differed among species of Galápagos finches was
	a. coloration.	c. leg length.
	b. beak size.	d. length of tail feathers.
	3.	Which of the following is not an example of natural selection?
	a.	
Insect populations exposed to pesticides become resistant to the chemicals.
	b.	
Plant species that produce fragrances to attract pollinators produce more offspring.
	c.	
Rabbits that sprint quickly are more likely to escape predation.
	d.	
On a tree, leaves that grow in the shade are larger than those that grow in the sun.
	4.	The variations necessary for natural selection
	a. occur randomly.
	b. are influenced by environment.
	c. can be caused by mutation.
	d. can be caused by recombination during meiosis.
	e. All of these are correct.
	5.	Which is most likely to be favored during natural selection, but not artificial selection?
	a. fast seed germination rate
	b. short generation time
	c. efficient seed dispersal
	d. lean pork meat production
	6.	Natural selection is the only process that results in
	a. genetic variation.
	b. adaptation to the environment.
	c. phenotypic change.
	d. competition among individuals in a population.
	7.	Why was it helpful to Darwin to learn that Lyell concluded the Earth was very old?
	a.		An old Earth has more fossils than a new Earth.
	b.	
It meant there was enough time for evolution to have occurred slowly.
	c.	
There was enough time for the same species to spread out into all continents.
	d.	Darwin said artificial selection occurs slowly.
	e.	All of these are correct. 
	8.	In the following diagram, contrast Lamarck’s proposal with Darwin’s proposal by matching the phrases in the key to the letters in the diagram.
Key:
		Originally, giraffe neck length varied.
	Giraffes stretched their necks in order to reach food.
	Struggle to exist causes long-necked giraffes to have the most 
	offspring.
	Originally, giraffes had short necks.
	Today most giraffes have long necks. (used twice)
	9.	All the finches on the Galápagos Islands 
	a.	are unrelated but descended from a common ancestor.
	b.	
are descended from a common ancestor, and therefore related.
	c.	rarely compete for the same food source.
	d.	Both a and c are correct. 
	e.	Both b and c are correct.
	10.	Evolution is considered a
	a.	
hypothesis because it is supported by data from the fossil record.
	b. hypothesis because it is supported by multiple types of data.
	c. theory because it is supported by data from the fossil record.
	d. theory because it is supported by multiple types of data.
	11.	Catastrophists were not able to explain
	a. multiple extinctions.
	b. the replacement of one group of organisms by another.
	c.	
successive changes that link groups of organisms in the fossil record.
	d. More than one of these are correct.
For questions 12–15, match the description with the type of evidence for evolution it supports, as listed in the key. Answers can be used more than once.
Key:
	a. biogeographical
	b. anatomical
	c. biochemical
	12.	The genetic code is the same for all organisms. 
	13.	The human kneebone and spine were derived from ancestral structures that supported four-legged animals. 
	14.	The South American continent lacks rabbits even though the environment is quite suitable.
	15.	The amino acid sequence of human hemoglobin is more similar to that of rhesus monkeys than to that of mice. 
	16.	Fossils that serve as transitional links allow scientists to 
	a. determine how prehistoric animals interacted with each other.
	b. deduce the order in which various groups of animals arose.
	c. relate climate change to evolutionary trends.
	d. determine why evolutionary changes occur.
	17.	Among vertebrates, the flipper of a dolphin and the fin of a tuna are
	a. homologous structures.
	b. homogeneous structures.
	c. analogous structures.
	d. reciprocal structures.
	18.	Which of these pairs is mismatched?
	a. Charles Darwin—natural selection
	b. 	Cuvier—series of catastrophes explains the fossil record
	c. Lamarck—uniformitarianism
	d. All of these are correct.
	19.	According to the inheritance of acquired characteristics hypothesis,
	a.	
if a man loses his hand, then his children will also be missing a hand.
	b.	
changes in phenotype are passed on by way of the genotype to the next generation.
	c.	
organisms are able to bring about a change in their phenotype.
	d.	evolution is striving toward particular traits.
	e.	All of these are correct. 
	20.	Organisms
	a. compete with other members of their species.
	b. differ in fitness.
	c. are adapted to their environment.
	d. are related by descent from common ancestors.
	e. All of these are correct. 
	21.	DNA nucleotide differences between organisms
	a. indicate how closely related organisms are.
	b. indicate that evolution occurs.
	c. explain why there are phenotypic differences.
	d. are to be expected.
	e. All of these are correct.
	22.	The fossil record offers direct evidence for common descent because you can
	a. see that the types of fossils change over time.
	b. sometimes find common ancestors.
	c. trace the ancestry of a particular group.
	d. trace the biological history of living things.
	e. All of these are correct. 
Go to www.mhhe.com/maderessentials for more quiz questions.
Bioethical Issue
The term “theory” in science is reserved for those ideas that scientists have found to be all-encompassing because they are based on data collected in a number of different fields. Evolution is a scientific theory. So is the cell theory, which says that all organisms are composed of cells, and so is the atomic theory, which says that all matter is composed of atoms. No one argues that schools should teach alternatives to the cell theory or the atomic theory. Yet, confusion reigns over the use of the expression “the theory of evolution.”
No wonder most scientists in our country are dismayed when state legislatures or school boards rule that teachers must put forward a variety of “theories” on the origin of life, including one that runs contrary to the mass of data that supports the theory of evolution. An organization in California called the Institute for Creation Research advocates that students be taught an “intelligent-design theory,” which says that DNA could never have arisen without the involvement of an “intelligent agent,” and that gaps in the fossil record mean that species arose fully developed with no antecedents.
Since no purely religious ideas can be taught in the schools, the advocates for an intelligent-design theory are careful not to mention the Bible or any strictly religious ideas. Still, teachers who have a solid scientific background do not feel comfortable teaching an intelligent-design theory because it does not meet the test of a scientific theory. Science is based on hypotheses that have been tested by observation and/or experimentation. A scientific theory has stood the test of time—that is, no hypotheses have been supported by observation and/or experimentation that run contrary to the theory. On the contrary, the Theory of Evolution is supported by data collected in such --wide-ranging fields as development, anatomy, geology, and biochemistry.
The polls consistently show that nearly half of all Americans prefer to believe the Old Testament account of creation. That, of course, is their right. But should schools be required to teach an intelligent-design theory that traces its roots back to the Old Testament and is not supported by observation and experimentation?
Understanding the Terms
adaptation221
analogous structure226
artificial selection222
biogeography219
evolution215
fitness222
fossil record224
fossils217
homologous structure226
natural selection221
speciation220
strata217
uniformitarianism219
vestigial structure226
Match the terms to these definitions:
a.	_______________	Structure that is similar in different types of organisms explainable by a common ancestry. 
b.	_______________	Lyell’s theory that geological changes occur slowly and at a uniform rate.
c.	_______________	Darwin’s proposed mechanism for evolutionary change.
d.	_______________	In an organism, remains of a nonfunctional structure that was once functional in an ancestor of the organism.
e.	_______________	Trait that helps an organism be better adapted to its environment.

Figure 14.2Fossils and strata.
a. When highways cut through mountains, it is often possible to see a number of strata, layers of rock or sedimentary material that contain fossils (b). The oldest fossils are in the lowest stratum.
Figure 14.1Voyage of the HMS Beagle.
The map shows the journey of the HMS Beagle around the world. As Darwin traveled along the east coast of South America, he noted that a bird called a rhea looked like the African ostrich. On the Galápagos Islands, marine iguanas, found no other place on Earth, use their large claws to cling to rocks and their blunt snouts for eating seaweed. 
Check Your Progress
	1.	Describe Lamarck’s idea of inheritance of acquired characteristics.
	2.	Why did Lyell think the Earth was very old?
Answers:1. Use and disuse of structures can produce heritable changes.
2. Massive geological changes have occurred but at a slow rate.
Figure 14.3Formation of strata.
a. This diagram shows how water brings sediments into the sea; the sediments then become compacted to form a stratum (sing.). Fossils are often trapped in strata, and as a result of a later geological upheaval, the strata may be located on land. b. Fossil remains of freshwater snails, Turritella, in a stratum.
Figure 14.4A glyptodont compared to an armadillo.
a. The giant armadillo-like glyptodont is known only by the study of its fossil remains. Darwin found such fossils and came to the conclusion that this extinct animal must be related to living armadillos. The glyptodont weighed 2,000 kilograms. b. A modern armadillo weighs about 4.5 kilograms.
Check Your Progress
	1.	
Explain why Darwin’s observations about Galápagos finches were significant to him.
	2.	Outline Darwin’s theory of natural selection.
Answers:1. The finches were all descended from a mainland finch, and yet they were adapted differently.2. Members of a population have heritable variations. More offspring are produced than resources can support. The most adapted individuals survive and reproduce. Across generations, a larger proportion of a species becomes more adapted to the environment.
Figure 14.7Variations in shells of a marine snail, Liguus fascitus.
For Darwin, variations such as these in a species of snails were highly significant and were required in order for natural selection to result in adaptation to the environment.
Figure 14.6Galápagos finches.
Each of the present-day 13 species of finches has a beak adapted to a particular way of life. a. The heavy beak of the large ground-dwelling finch is suited to a diet of seeds. b. The beak of the warbler finch is suited to feeding on insects found among ground vegetation or caught in the air. 
c. The long, somewhat decurved beak and split tongue of the cactus finch are suited to probing cactus flowers for nectar.
Figure 14.5The Patagonian hare.
This animal has the face of a guinea pig and is native to South America, which has no native rabbits. The Patagonian hare has long legs and other adaptations similar to those of rabbits.
Check Your Progress
Explain the occurrence of variations among members of a population.
Answer:Genetic variation arises by mutation and by assortment of chromosomes and recombination of alleles during sexual reproduction. 
Check Your Progress
What is the main difference between artificial selection and natural selection? 
Answer:Artificial selection is carried out by humans, and natural selection is carried out by environmental forces, other than humans.
Figure 14.10Adaptations of the vampire bat.
Vampire bats of the rain forests of Central and South America have the adaptations of a nocturnal, winged predator. The bat uses its enlarged ears and echolocation to locate prey in the dark. It bites its prey with two pointed incisors. Saliva containing an anticoagulant called draculin runs into the bite; the bat then licks the flowing blood. The vampire bat’s forelimbs are modified to form wings, and it roosts by using its grasping hind limbs.   
Figure 14.8Mechanism of evolution.
This diagram contrasts Jean-Baptiste Lamarck’s proposal of acquired characteristics with Charles Darwin’s proposal of natural selection. Only natural selection is supported by data.
Figure 14.13Marsupials of Australia.
Each type of marsupial in Australia is adapted to a different way of life. All of them presumably evolved from a common ancestor that entered Australia some 60 million years ago.
Figure 14.12Evolution of whales.
The discoveries of Ambulocetus and Rodhocetus have filled in the gaps in the evolution of whales from extinct hoofed mammals that lived on land to the ocean-dwelling mammal we know today.
Figure 14.16Significance of biochemical differences.
The branch points in this diagram tell the number of amino acids that differ between human cytochrome c and the organisms depicted. These biochemical data are consistent with those provided by a study of the fossil record and comparative anatomy.
Figure 14.14Significance of structural similarities.
Although the specific design details of vertebrate forelimbs are different, the same bones are present (color-coded here). This unity of plan is evidence of a common ancestor.
Figure 14.15Significance of developmental similarities.
At this comparable developmental stage, a chick embryo and a pig embryo have many features in common, which suggests they evolved from a common ancestor.
Check Your Progress
	1.	Explain how vestigial structures support the theory of evolution.
	2.	Contrast homologous structures with analogous structures.
	3.	With reference to Figure 14.16, why would it make sense that the most distantly related species have greater structural differences and greater DNA differences?
Answers:1. These structures probably arose from an ancestor that needed them, but they are no longer useful.2. Homologous structures share an anatomical similarity that reflects a common ancestry. Analogous structures have a common function but do not serve as evidence of a common ancestry.3. DNA determines the structure and function of organisms.
Due to artificial selection, you can choose from more than 150 breeds of dogs. 
Only in Hawaii did honey-creepers evolve into over 16 species. 

Table 14.1	Contrast of Worldviews	

Pre-Darwinian View	Post-Darwinian View	
1.	The Earth is relatively young—age is measured in thousands of years. 	1.	The Earth is relatively old—age is measured in billions of years.	
2.	Each species is specially created; species don’t change, and the number of species remains the same. 	2.	Species are related by descent—it is possible to piece together a history of life on Earth, showing lines of descent.	
3.	Adaptation to the environment is the work of a creator who 
decided the structure and function of each type of organism. 
Any variations are imperfections.	3.	Adaptation to the environment is the interplay of random variations and environmental conditions.	
4.	Observations are supposed to substantiate the prevailing 
worldview.	4.	Observation and experimentation are used to test hypotheses, including hypotheses about evolution.	

Coarse-haired wombat, nocturnal and living in burrows

Like dogs and honey-creepers, bacteria evolve. A growing problem in hospitals is the number of bacteria that have become resistant to antibiotics.
Kangaroo, a herbivore of plains and forests

marine iguanas
Charles Darwin
Check Your Progress
	1.	Describe the pre-Darwinian view of the world.
	2.	
Describe Cuvier’s contribution to evolutionary theory.
Answers:1. The Earth is only thousands of years old. Species do not change over time. A creator is responsible for adaptation to the environment. Observations should support the current view of the world.2. Cuvier founded the science of paleontology. Fossils are found in strata, and a change in species can be observed over time.
Figure 14.9Adaptations of the alligator bug.
The alligator bug of the Brazilian rain forest has antipredator adaptations. a. The insect blends into its background, but if discovered, the false head, which resembles a miniature alligator, may frighten a predator. b. If the predator is not frightened, the insect suddenly reveals huge false eyespots on its hindwings.
Check Your Progress
	1.	Explain why evolution is no longer considered a hypothesis.
	2.	
Explain how biogeographical information about Galápagos finches supports the theory of evolution.
Answers:1. Evolution is supported by many diverse and independent lines of evidence.2. The finches show that a common ancestor can give rise to other species through adaptation to unique environments—in this case, the environment on different islands.
Figure 14.11Re-creation Archaeopteryx.
The fossil record suggests that Archaeopteryx had a feather-covered, reptilian-type tail, which shows up well in this artist’s representation. (Red labels  reptilian characteristics; 
green labels  bird characteristics.)