IN THE BEGINNING - Welby Thomas Cox, Jr.

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has traits that give it an advantage in a particular environment, that organism will be more likely to survive and pass on those traits. As differences accumulate over generations, populations of organisms diverge from their ancestors.

      

CHARLES DARWIN

      Charles Darwin arrived at many of his insights into evolution by studying the variations among species on the Galápagos Islands off the coast of Ecuador.

      Darwin's original hypothesis has undergone extensive modification and expansion, but the central concepts stand firm. Studies in genetics and molecular biology—fields unknown in Darwin's time—have explained the occurrence of the hereditary variations that are essential to natural selection. Genetic variations result from changes, or mutations, in the nucleotide sequence of DNA, the molecule that genes are made from. Such changes in DNA now can be detected and described with great precision.

      Genetic mutations arise by chance. They may or may not equip the organism with better means for surviving in its environment. But if a gene variant improves adaptation to the environment (for example, by allowing an organism to make better use of an available nutrient, or to escape predators more effectively—such as through stronger legs or disguising coloration), the organisms carrying that gene are more likely to survive and reproduce than those without it. Over time, their descendants will tend to increase, changing the average characteristics of the population. Although the genetic variation on which natural selection works is based on random or chance elements, natural selection itself produces "adaptive" change—the very opposite of chance.

      Scientists also have gained an understanding of the processes by which new species originate. A new species is one in which the individuals cannot mate and produce viable descendants with individuals of a preexisting species. The split of one species into two often starts because a group of individuals becomes geographically separated from the rest. This is particularly apparent in distant remote islands, such as the Galápagos and the Hawaiian archipelago, whose great distance from the Americas and Asia means that arriving colonizers will have little or no opportunity to mate with individuals remaining on those continents. Mountains, rivers, lakes, and other natural barriers also account for geographic separation between populations that once belonged to the same species.

      Once isolated, geographically separated groups of individuals become genetically differentiated as a consequence of mutation and other processes, including natural selection. The origin of a species is often a gradual process, so that at first the reproductive isolation between separated groups of organisms is only partial, but it eventually becomes complete. Scientists pay special attention to these intermediate situations because they help to reconstruct the details of the process and to identify particular genes or sets of genes that account for the reproductive isolation between species.

      A particularly compelling example of speciation involves the 13 species of finches studied by Darwin on the Galápagos Islands, now known as Darwin's finches. The ancestors of these finches appear to have immigrated from the South American mainland to the Galápagos. Today the different species of finches on the island have distinct habitats, diets, and behaviors, but the mechanisms involved in speciation continue to operate. A research group led by Peter and Rosemary Grant of Princeton University has shown that a single year of drought on the islands can drive evolutionary changes in the finches. Drought diminishes supplies of easily cracked nuts but permits the survival of plants that produce larger, tougher nuts. Droughts thus favor birds with strong, wide beaks that can break these tougher seeds, producing populations of birds with these traits. The Grants have estimated that if droughts occur about once every 10 years on the islands, a new species of finch might arise in only about 200 years.

      The different species of finches on the Galápagos Islands, now known as Darwin's finches, have different-sized beaks that have evolved to take advantage of distinct food sources.

      The following sections consider several aspects of biological evolution in greater detail, looking at paleontology, comparative anatomy, biogeography, embryology, and molecular biology for further evidence supporting evolution.

       The Fossil Record

      Although it was Darwin, above all others, who first marshaled convincing evidence for biological evolution, earlier scholars had recognized that organisms on Earth had changed systematically over long periods of time. For example, in 1799 an engineer named William Smith reported that, in undisrupted layers of rock, fossils occurred in a definite sequential order, with more modern-appearing ones closer to the top. Because bottom layers of rock logically were laid down earlier and thus are older than top layers, the sequence of fossils also could be given a chronology from oldest to youngest. His findings were confirmed and extended in the 1830s by the paleontologist William Lonsdale, who recognized that fossil remains of organisms from lower strata were more primitive than the ones above. Today, many thousands of ancient rock deposits have been identified that show corresponding successions of fossil organisms.

      Thus, the general sequence of fossils had already been recognized before Darwin conceived of descent with modification. But the paleontologists and geologists before Darwin used the sequence of fossils in rocks not as proof of biological evolution, but as a basis for working out the original sequence of rock strata that had been structurally disturbed by earthquakes and other forces.

      In Darwin's time, paleontology was still a rudimentary science. Large parts of the geological succession of stratified rocks were unknown or inadequately studied.

      Weathering has exposed layers of sedimentary rock near the Praia River in Utah

      Darwin, therefore, worried about the rarity of intermediate forms between some major groups of organisms.

      Today, many of the gaps in the paleontological record have been filled by the research of paleontologists. Hundreds of thousands of fossil organisms, found in well-dated rock sequences, represent successions of forms through time and manifest many evolutionary transitions. As mentioned earlier, microbial life of the simplest type was already in existence 3.5 billion years ago. The oldest evidence of more complex organisms (that is, eucaryotic cells, which are more complex than bacteria) has been discovered in fossils sealed in rocks approximately 2 billion years old. Multicellular organisms, which are the familiar fungi, plants, and animals, have been found only in younger geological strata. The following list presents the order in which increasingly complex forms of life appeared:

       Life Form

       Millions of Years Since First Known Appearance (Approximate)

      Microbial (procaryotic cells): 3,500

      Complex (eucaryotic cells): 2,000

      First multicellular animals: 670

      Shell-bearing animals: 540

      Vertebrates (simple fishes): 490

      Amphibians: 350

      Reptiles: 310

      Mammals: 200

      Nonhuman primates: 60

      Earliest apes: 25

      Australopithecine ancestors of humans: 5

      Modern humans: 0.15 (150,000 years)

      So many intermediate forms have been discovered between fish and amphibians, between amphibians and reptiles, between reptiles and mammals, and along the primate lines of descent that it often is difficult to identify categorically when the transition occurs from one to another particular species. Actually, nearly all fossils can be regarded as intermediates in some sense; they are life forms that come between the forms that preceded them and those that followed.

      The fossil record thus provides consistent evidence of systematic change through time—of descent with modification. From this huge body of evidence,

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