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Part of a series onCreationismHistory of creationism Neo-creationismTypes of creationismYoung Earth creationism Old Earth creationism Day-Age creationism Progressive creationism Gap creationism Intelligent designOther religious viewsHindu Islamic Jewish Deist PandeistCreation theologyCreation in Genesis Genesis as an allegory Framework interpretation Omphalos hypothesisCreation scienceBaraminology Flood geology Intelligent designControversyPolitics of creationism Public education History Teach the Controversy Associated articlesCreationism Portal v d e Part of the Biology series onEvolutionIntroductionMechanisms and processesAdaptation Genetic drift Gene flow Mutation Natural selection SpeciationResearch and historyEvidence Evolutionary history of life History Modern synthesis Social effect

Theory and fact Objections / ControversyEvolutionary biology fieldsCladistics Ecological genetics Evolutionary development Human evolution Molecular evolution Phylogenetics Population geneticsBiology Portal v d e The creationevolution controversy (also termed the creation vs. evolution debate or the origins debate) is a recurring theological and cultural-political dispute about the origins of the Earth, humanity, life, and the universe,[1] between the proponents of evolution, backed by scientific consensus, and those who espouse the validity and/or superiority of various literal interpretations of creation myth. The dispute particularly involves the field of evolutionary biology, but also the fields of geology, palaeontology, thermodynamics, nuclear physics and cosmology.[2] This debate is most prevalent in the United States but also exists in Europe and elsewhere.[3] It is often portrayed as part of the culture wars.[4] While the controversy has a long history,[5] today it is mainly over what constitutes good science,[6] with the politics of creationism primarily focusing on the teaching of creation and evolution in public education.[7] The debate also focuses on issues such as the definition of science (and of what constitutes scientific research and evidence), science education (and whether the teaching of the scientific consensus view should be 'balanced' by also teaching fringe theories), free speech, separation of Church and State, and theology (particularly how different Christian denominations interpret the Book of Genesis). Within the scientific community and academia the level of support for evolution is essentially universal,[8] while support for biblicallyliteral accounts or other creationist alternatives is very small among scientists, and virtually nonexistent among those in the relevant fields.[9] The debate is sometimes portrayed as being between science and religion. However, as the National Academy of Sciences states: Today, many religious denominations accept that biological evolution has produced the diversity of living things over billions of years of Earths history. Many have issued statements observing that evolution and the tenets of their faiths are compatible. Scientists and theologians have written eloquently about their awe and wonder at the history of the universe and of life on this planet, explaining that they see no conflict between their faith in God and the evidence for evolution. Religious denominations that do not accept the occurrence of evolution tend to be those that believe in strictly literal interpretations of religious texts.Science, Evolution, and Creationism, National Academy of Sciences[10] Contents * 1 History of the controversy o 1.1 Controversies in the age of Darwin o 1.2 Creationism * 1.2.1 The Butler Act and the Scopes monkey trial

* 1.2.2 Epperson v. Arkansas * 1.2.3 Daniel v. Waters o 1.3 Creation Science * 1.3.1 Court cases * 1.3.1.1 McLean v. Arkansas * 1.3.1.2 Edwards v. Aguillard o 1.4 Intelligent Design * 1.4.1 Controversy in recent times * 1.4.2 Kansas evolution hearings * 1.4.3 The Dover Trial * 2 Viewpoints o 2.1 Young Earth creationism o 2.2 Old Earth creationism o 2.3 Neo-Creationism o 2.4 Theistic evolution o 2.5 Naturalistic evolution * 3 Arguments relating to the definition and limits of science o 3.1 Definitions o 3.2 Limitations of the scientific endeavor o 3.3 Theory vs. fact o 3.4 Falsifiability o 3.5 Conflation of science and religion * 4 Disputes relating to science o 4.1 Biology * 4.1.1 Common descent * 4.1.1.1 Human evolution * 4.1.2 Macroevolution * 4.1.3 Transitional fossils o 4.2 Geology o 4.3 Other sciences * 4.3.1 Cosmology * 4.3.2 Nuclear physics o 4.4 Misrepresentations of science * 4.4.1 Quote mining * 5 Public policy issues o 5.1 Science education o 5.2 Freedom of speech * 6 Issues relating to religion o 6.1 Theological arguments o 6.2 Religion and historical scientists * 7 Forums for the controversy o 7.1 Debates o 7.2 Political lobbying o 7.3 In the media * 8 Outside the United States o 8.1 Europe o 8.2 Australia o 8.3 Islamic countries * 9 See also * 10 Footnotes * 11 References o 11.1 Published books and other resources * 12 External links o 12.1 Creationism as social policy o 12.2 Creationist beliefs * 12.2.1 Scientific rebuttals o 12.3 Evolution versus creationism debates History of the controversy

See also: History of evolutionary thought Controversies in the age of Darwin See also: Reaction to Darwin's theory

A satirical image of Charles Darwin as an ape from 1871 reflects part of the social controversy over whether humans and apes share a common lineage. The creation-evolution controversy originated in Europe and North America in the late eighteenth century when discoveries in geology led to various theories of an ancient earth, and fossils showing past extinctions prompted early ideas of evolution, notably Lamarckism. In England these ideas of continuing change were seen as a threat to the fixed social order, and were harshly repressed.[11] Conditions eased, and in 1844 the controversial Vestiges popularised transmutation of species. The scientific establishment dismissed it scornfully and the Church of England reacted with fury, but many Unitarians, Quakers and Baptists opposed to the privileges of the Established church favoured its ideas of God acting through laws. Publication of Charles Darwin's On the Origin of Species by Means of Natural Selection in 1859 brought scientific credibility to evolution, and made it more respectable.[12] There was intense interest in the religious implications of Darwin's book, but the Church of England's attention was largely diverted by theological controversy over higher criticism set out in Essays and Reviews by liberal Christian authors, some of whom expressed support for Darwin, as did many nonconformists. The Reverend Charles Kingsley openly supported the idea of God working through evolution. However, many Christians were opposed to the idea and even some of Darwin's close friends and supporters including Charles Lyell and Asa Gray could not accept some of his ideas.[13] Thomas Huxley, who strongly promoted Darwin's ideas while campaigning to end the dominance of science by the clergy, coined the term agnostic to describe his position that Gods existence is unknowable, and Darwin also took this position,[13] but evolution was also taken up by prominent atheists including Edward Aveling and Ludwig Bchner and criticised, in the words of one reviewer, as "tantamount to atheism."[14] By the end of the 19th century Roman Catholics guided by Pope Leo XIII accepted human evolution from animal ancestors while affirming that the human soul was directly created by God.[13] Creationists during this period were largely premillennialists, whose belief in Christ's return depended on a quasi-literal reading of the Bible.[15] However, they were not as concerned about geology, freely granting scientists any time they needed before the Garden of Eden to account for scientific observations, such as fossils and geological findings.[16] In the immediate post-Darwinian era, few scientists or clerics rejected the antiquity of the earth or the progressive nature of the fossil record.[17] Likewise, few attached geological significance to the Biblical flood, unlike subsequent creationists.[17] Evolutionary skeptics, creationist leaders and skeptical scientists were usually willing either to adopt a figurative reading of the first chapter of Genesis, or to allow that the six days of creation were not necessarily 24-hour days.[18] Creationism Main article: History of creationism See also: Creation and evolution in public education

In the United States of America Creationism was widely accepted and was considered a foundational truth, but there was no official resistance to evolution by mainline denominations.[13] Around the start of the 20th century some evangelical scholars had ideas accommodating evolution, such as B. B. Warfield who saw it as a natural law expressing Gods will. However, development of the eugenics movement led many Catholics to reject evolution.[13] In this enterprise they received little aid from conservative Christians in Britain and Europe. In Britain this has been attributed to their minority status leading to a more tolerant, less militant theological tradition. The main British Creationist movement in this period was the Evolution Protest Movement, formed in the 1930s.[19] The Butler Act and the Scopes monkey trial Main article: Scopes trial

Clarence Darrow and William Jennings Bryan chat in court during the Scopes trial. In the aftermath of World War I, the Fundamentalist-Modernist Controversy brought a surge of opposition to the idea of evolution, and following the campaigning of William Jennings Bryan several states introduced legislation prohibiting the teaching of evolution. By 1925, such legislation was being considered in 15 states, and passed in some states, such as Tennessee. The American Civil Liberties Union offered to defend anyone who wanted to bring a test case against one of these laws. John T. Scopes accepted, and he confessed to teaching his Tennessee class evolution in defiance of the Butler Act. The textbook in question was Hunter's Civic Biology (1914). The trial was widely publicized by H. L. Mencken among others, and is commonly referred to as the Scopes Monkey Trial. Scopes was convicted; however, the widespread publicity galvanized proponents of evolution. When the case was appealed to the Tennessee Supreme Court, the Court overturned the decision on a technicality (the judge had assessed the fine when the jury had been required to). Although it overturned the conviction, the Court decided that the law was not in violation of the First Amendment. The Court held, "We are not able to see how the prohibition of teaching the theory that man has descended from a lower order of animals gives preference to any religious establishment or mode of worship. So far as we know, there is no religious establishment or organized body that has in its creed or confession of faith any article denying or affirming such a theory." Scopes v. State 289 S.W. 363, 367 (Tenn. 1927). The interpretation of the Establishment clause up to that time was that Congress could not establish a particular religion as the State religion. Consequently, the Court held that the ban on the teaching of evolution did not violate the Establishment clause, because it did not establish one religion as the "State religion." As a result of the holding, the teaching of evolution remained illegal in Tennessee, and continued campaigning succeeded in removing evolution from school textbooks throughout the United States.[20] Epperson v. Arkansas Main article: Epperson v. Arkansas In 1968, the United States Supreme Court invalidated a forty year old Arkansas statute that prohibited the teaching of evolution in the public schools. A Little Rock high school biology teacher, Susan Epperson, filed suit charging the law violated the constitutional protection of free speech. The Little Rock Ministerial Association supported Epperson's challenge, declaring, "to use the Bible to support an irrational and an

archaic concept of static and undeveloping creation is not only to misunderstand the meaning of the Book of Genesis, but to do God and religion a disservice by making both enemies of scientific advancement and academic freedom."[21] The Court held that the United States Constitution prohibits a state from requiring, in the words of the majority opinion, "that teaching and learning must be tailored to the principles or prohibitions of any religious sect or dogma."[22] But the Supreme Court decision also suggested that creationism could be taught in addition to evolution.[23] Daniel v. Waters Main article: Daniel v. Waters Daniel v. Waters was a 1975 legal case in which the United States Court of Appeals for the Sixth Circuit struck down Tennessee's law regarding the teaching of "equal time" of evolution and creationism in public school science classes because it violated the Establishment clause of the US Constitution. Following this ruling, creationism was stripped of overt biblical references and renamed creation science, and several states passed legislative acts requiring that this be given equal time with teaching of evolution. Creation Science Main article: Creation Science As biologists grew more and more confident in evolution as the central defining principle of biology,[24] American membership in churches favoring increasingly literal interpretations of scripture rose, with the Southern Baptist Convention and Lutheran Church - Missouri Synod outpacing all other denominations.[25] With growth, these churches became better equipped to promulgate a creationist message, with their own colleges, schools, publishing houses, and broadcast media.[26] In 1961, the first major modern creationist book was published: Henry M. Morris and John C. Whitcomb Jr.'s The Genesis Flood. Morris and Whitcomb argued that creation was literally 6 days long, that humans lived concurrently with dinosaurs, and that God created each 'kind' of life individually.[27] On the strength of this, Morris became a popular speaker, spreading anti-evolutionary ideas at fundamentalist churches, colleges, and conferences.[28] Morris' Creation Science Research Center (CSRC) rushed publication of biology text books that promoted creationism, and also published other books such as Kelly Segrave's sensational Sons of God Return that dealt with UFOlogy, flood geology, and demonology against Morris' objections.[29] Ultimately, the CSRC broke up over a divide between sensationalism and a more intellectual approach, and Morris founded the Institute for Creation Research, which was promised to be controlled and operated by scientists.[30] During this time, Morris and others who supported flood geology adopted the scientific-sounding terms scientific creationism and creation science.[31] The flood geologists effectively co-opted "the generic creationist label for their hyperliteralist views".[32] Court cases McLean v. Arkansas Main article: McLean v. Arkansas In 1982 another case in Arkansas ruled that the Arkansas "Balanced Treatment for Creation-Science and Evolution-Science Act" was unconstitutional because it violated the establishment clause of the U.S. Constitution. Much of the transcript of the case was lost, including evidence from Francisco Ayala.

Edwards v. Aguillard Main article: Edwards v. Aguillard In the early 1980s, the Louisiana legislature passed a law titled the "Balanced Treatment for Creation-Science and Evolution-Science in Public School Instruction Act". The act did not require teaching either evolution or creationism as such, but did require that when evolutionary science was taught, so-called creation science had to be taught as well. Creationists had lobbied aggressively for the law, arguing that the act was about academic freedom for teachers, an argument adopted by the state in support of the act. Lower courts ruled that the State's actual purpose was to promote the religious doctrine of creation science, but the State appealed to the Supreme Court. The similar case in McLean v. Arkansas had also decided against creationism. Mclean v. Arkansas however was not appealed to the federal level, creationists instead thinking that they had better chances with Edwards v. Aguillard. In 1987 the Supreme Court of the United States ruled that the act was unconstitutional, because the law was specifically intended to advance a particular religion. At the same time, however, it held that "teaching a variety of scientific theories about the origins of humankind to school children might be validly done with the clear secular intent of enhancing the effectiveness of science instruction" leaving open the door for a handful of proponents of creation science to evolve their arguments into the iteration of creationism that came to be known as intelligent design.[33] Intelligent Design

The Discovery Institute's Center for the Renewal of Science and Culture used banners based on "The Creation of Adam" from the Sistine Chapel. Later it used a less religious image, then was renamed the Center for Science and Culture.[34] Main article: Intelligent design See also: Neo-creationism, Intelligent design movement, Teach the Controversy, and Critical Analysis of Evolution In response to Edwards v. Aguillard, the Neo-Creationist intelligent design movement was formed around the Discovery Institute's Center for Science and Culture. Its goal is to restate creationism in terms more likely to be well received by the public, policy makers, educators, and the scientific community, and makes the claim that "certain features of the universe and of living things are best explained by an intelligent cause, not an undirected process such as natural selection."[35] It has been viewed as a "scientific" approach to creationism by creationists, but is widely rejected as unscientific by the science community (see for example, list of scientific societies rejecting intelligent design). Controversy in recent times See also: Politics of creationism and Intelligent design in politics The controversy continues to this day, with the mainstream scientific consensus on the origins and evolution of life challenged by creationist organizations and religious groups who desire to uphold some form of creationism (usually young earth creationism, creation science, old earth creationism or intelligent design) as an alternative. Most of these groups are explicitly Christian, and more than one sees the debate as part of the Christian mandate to evangelize.[36] Some see science and religion as being diametrically opposed views which cannot be reconciled. More accommodating viewpoints, held by many mainstream churches and many scientists, consider science and religion to be separate categories of

thought, which ask fundamentally different questions about reality and posit different avenues for investigating it.[37] Public opinion in regards to the concepts of evolution, creationism, and intelligent design is fluctuating. More recently, the Intelligent Design movement has taken an antievolution position which avoids any direct appeal to religion. Scientists argue that Intelligent design does not represent any research program within the mainstream scientific community, and is essentially creationism.[38] Its leading proponent, the Discovery Institute, made widely publicised claims that it was a new science, though the only paper arguing for it published in a scientific journal was accepted in questionable circumstances and quickly disavowed in the Sternberg peer review controversy, with the Biological Society of Washington stating that it did not meet the journal's scientific standards, was a "significant departure" from the journal's normal subject area and was published at the former editor's sole discretion, "contrary to typical editorial practices".[39] President Bush commented endorsing the teaching of Intelligent design alongside evolution "I felt like both sides ought to be properly taught ... so people can understand what the debate is about."[40] Kansas evolution hearings Main article: Kansas evolution hearings In the push by intelligent design advocates to introduce intelligent design in public school science classrooms, the hub of the intelligent design movement, the Discovery Institute, arranged to conduct hearings to review the evidence for evolution in the light of its Critical Analysis of Evolution lesson plans. The Kansas Evolution Hearings were a series of hearings held in Topeka, Kansas 5 May to 12 May 2005. The Kansas State Board of Education eventually adopted the institute's Critical Analysis of Evolution lesson plans over objections of the State Board Science Hearing Committee, and electioneering on behalf of conservative Republican candidates for the Board.[41] On 1 August 2006, 4 of the 6 conservative Republicans who approved the Critical Analysis of Evolution classroom standards lost their seats in a primary election. The moderate Republican and Democrats gaining seats vowed to overturn the 2005 school science standards and adopt those recommended by a State Board Science Hearing Committee that were rejected by the previous board,[42] and on 13 February 2007, the Board voted 6 to 4 to reject the amended science standards enacted in 2005. The definition of science was once again limited to "the search for natural explanations for what is observed in the universe."[43] The Dover Trial Main article: Kitzmiller v. Dover Area School District Following the Edwards v. Aguillard decision by the Supreme Court of the United States, in which the Court held that a Louisiana law requiring that creation science be taught in public schools whenever evolution was taught was unconstitutional, because the law was specifically intended to advance a particular religion, creationists renewed their efforts to introduce creationism into public school science classes. This effort resulted in intelligent design, which sought to avoid legal prohibitions by leaving the source of creation an unnamed and undefined intelligent designer, as opposed to God.[44] This ultimately resulted in the "Dover Trial," Kitzmiller v. Dover Area School District, which went to trial on 26 September 2005 and was decided on 20 December 2005 in favor of the plaintiffs, who charged that a mandate that intelligent design be taught in public school science classrooms was an unconstitutional establishment

of religion. The 139 page opinion of Kitzmiller v. Dover was hailed as a landmark decision, firmly establishing that creationism and intelligent design were religious teachings and not areas of legitimate scientific research. Viewpoints Young Earth creationism Main article: Young Earth creationism See also: Creation science and Flood geology Young Earth creationism is the belief that the Earth was created by God within the last 10,000 years, literally as described in Genesis, within the approximate timeframe of biblical genealogies (detailed for example in the Ussher chronology). Young Earth creationists often believe that the Universe has a similar age as the Earth. Creationist cosmologies are attempts by some creationist thinkers to give the universe an age consistent with the Ussher chronology and other Young-Earth timeframes. This belief generally has a basis in a literal and inerrant interpretation of the Bible. Old Earth creationism Main article: Old Earth creationism See also: Gap creationism, Day-Age Creationism, and Progressive creationism Old Earth creationism holds that the physical universe was created by God, but that the creation event of Genesis is not to be taken strictly literally. This group generally believes that the age of the Universe and the age of the Earth are as described by astronomers and geologists, but that details of the evolutionary theory are questionable. Old Earth creationists interpret the creation accounts of Genesis in a number of ways, that each differ from the six, consecutive, 24-hour day creation of the literalist Young Earth Creationist view. Neo-Creationism Main article: Neo-Creationism See also: Intelligent design Neo-Creationists intentionally distance themselves from other forms of creationism, preferring to be known as wholly separate from creationism as a philosophy. Their goal is to restate creationism in terms more likely to be well received by the public, education policy makers and the scientific community. It aims to re-frame the debate over the origins of life in non-religious terms and without appeals to scripture, and to bring the debate before the public. Neo-creationists may be either Young Earth or Old Earth Creationists, and hold a range of underlying theological viewpoints (e.g. on the interpretation of the Bible). NeoCreationism currently exists in the form of the intelligent design movement, which has a 'big tent' strategy making it inclusive of many Young Earth Creationists (such as Paul Nelson and Percival Davis). Theistic evolution Main article: Theistic evolution See also: Naturalism (philosophy), Evolution and the Roman Catholic Church, and Clergy Letter Project Theistic evolution is the general view that, instead of faith being in opposition to biological evolution, some or all classical religious teachings about God and creation are compatible with some or all of modern scientific theory, including, specifically, evolution. It generally views evolution as a tool used by a creator god, who is both

the first cause and immanent sustainer/upholder of the universe; it is therefore well accepted by people of strong theistic (as opposed to deistic) convictions. Theistic evolution can synthesize with the day-age interpretation of the Genesis creation account; however most adherents consider that the first chapters of Genesis should not be interpreted as a "literal" description, but rather as a literary framework or allegory. This position does not generally exclude the viewpoint of methodological naturalism, a long standing convention of the scientific method in science. Theistic evolutionists have frequently been prominent in opposing creationism (including intelligent design). Notable examples have been biologist Kenneth R. Miller and theologian John Haught (both Catholics), who testified for the plaintiffs in Kitzmiller v. Dover Area School District. Another example is the Clergy Letter Project, an organization that has created and maintains a statement signed by American Christian clergy of different denominations rejecting creationism, with specific reference to points raised by intelligent design proponents. Theistic evolutionists have also been active in Citizens Alliances for Science that oppose the introduction of creationism into public school science classes (one example being evangelical Christian geologist Keith B. Miller, who is a prominent board member of Kansas Citizens for Science). Naturalistic evolution See also: Metaphysical naturalism Naturalistic evolution is the position of acceptance of biological evolution and of metaphysical naturalism (and thus rejection of theism and theistic evolution). This section requires expansion. Arguments relating to the definition and limits of science Critiques such as those based on the distinction between theory and fact are often leveled against unifying concepts within scientific disciplines. Principles such as uniformitarianism, Occam's Razor or parsimony, and the Copernican principle are claimed to be the result of a bias within science toward philosophical naturalism, which is equated by many creationists with atheism.[45] In countering this claim, philosophers of science use the term methodological naturalism to refer to the long standing convention in science of the scientific method. The methodological assumption is that observable events in nature are explained only by natural causes, without assuming the existence or nonexistence of the supernatural, and therefore supernatural explanations for such events are outside the realm of science.[46] Creationists claim that supernatural explanations should not be excluded and that scientific work is paradigmatically close-minded.[47] Because modern science tries to rely on the minimization of a priori assumptions, error, and subjectivity, as well as on avoidance of Baconian idols, it remains neutral on subjective subjects such as religion or morality.[48] Mainstream proponents accuse the creationists of conflating the two in a form of pseudoscience.[49] Definitions Fact: In science, an observation that has been repeatedly confirmed and for all practical purposes is accepted as "true." Truth in science, however, is never final, and what is accepted as a fact today may be modified or even discarded tomorrow. Hypothesis: A tentative statement about the natural world leading to deductions that can be tested. If the deductions are verified, it becomes more probable that the hypothesis is correct. If the deductions are incorrect, the original hypothesis can be abandoned or modified. Hypotheses can be used to build more complex

inferences and explanations. Law: A descriptive generalization about how some aspect of the natural world behaves under stated circumstances. Theory: In science, a well-substantiated explanation of some aspect of the natural world that can incorporate facts, laws, inferences, and tested hypotheses. National Academy of Sciences, Science and Creationism[50] Limitations of the scientific endeavor In science, explanations are limited to those based on observations and experiments that can be substantiated by other scientists. Explanations that cannot be based on empirical evidence are not a part of science. National Academy of Sciences, Science and Creationism[50] This section requires expansion. Theory vs. fact Main article: Evolution as theory and fact The argument that evolution is a theory, not a fact, has often been made against the exclusive teaching of evolution.[51] The argument is related to a common misconception about the technical meaning of "theory" that is used by scientists. In common usage, "theory" often refers to conjectures, hypotheses, and unproven assumptions. However, in science, "theory" usually means "a plausible or scientifically acceptable general principle or body of principles offered to explain phenomena."[52] Exploring this issue, paleontologist Stephen Jay Gould wrote:[53] Evolution is a theory. It is also a fact. And facts and theories are different things, not rungs in a hierarchy of increasing certainty. Facts are the world's data. Theories are structures of ideas that explain and interpret facts. Facts do not go away when scientists debate rival theories to explain them. Einstein's theory of gravitation replaced Newton's, but apples did not suspend themselves in mid-air, pending the outcome. And humans evolved from ape-like ancestors whether they did so by Darwin's proposed mechanism or by some other yet to be discovered. Falsifiability Philosopher of science Karl R. Popper set out the concept of falsifiability as a way to distinguish science and pseudoscience: Testable theories are scientific, but those that are untestable are not.[54] However, in Unended Quest, Popper declared "I have come to the conclusion that Darwinism is not a testable scientific theory but a metaphysical research programme, a possible framework for testable scientific theories," while pointing out it had "scientific character".[55] In what one sociologist derisively called "Popper-chopping,"[56] opponents of evolution seized upon Popper's definition to claim evolution was not a science, and claimed creationism was an equally valid metaphysical research program.[57] For example, Duane Gish, a leading Creationist proponent, wrote in a letter to Discover magazine (July 1981): "Stephen Jay Gould states that creationists claim creation is a scientific theory. While many Creationists claim creation is a scientific theory other Creationists have stated that neither creation nor evolution is a scientific theory (and each is equally religious)."[58][citation needed] Popper responded to news that his conclusions were being used by antievolutionary forces by affirming that evolutionary theories regarding the origins of life on earth were scientific because "their hypotheses can in many cases be tested."[59] However, creationists claimed that a key evolutionary concept, that all life on Earth is descended from a single common ancestor, was not mentioned as testable by Popper, and claimed it never would be.[60]

In fact, Popper wrote admiringly of the value of Darwin's theory.[61] Only a few years later, Popper changed his mind, and later wrote, "I still believe that natural selection works in this way as a research programme. Nevertheless, I have changed my mind about the testability and logical status of the theory of natural selection; and I am glad to have an opportunity to make a recantation".[62] Debate among some scientists and philosophers of science on the applicability of falsifiability in science continues.[63] However, simple falsifiability tests for common descent have been offered by some scientists: For instance, biologist and prominent critic of creationism Richard Dawkins and J.B.S. Haldane both pointed out that if fossil rabbits were found in the Precambrian era, a time before most similarly complex lifeforms had evolved, "that would completely blow evolution out of the water."[64][65] Falsifiability has also caused problems for creationists: In his 1982 decision McLean v. Arkansas Board of Education, Judge William R. Overton used falsifiability as one basis for his ruling against the teaching of creation science in the public schools, ultimately declaring it "simply not science."[66] Conflation of science and religion Many of the most vocal creationists blur the boundaries between criticisms of modern science, philosophy, and culture. They often conjoin their arguments focused on the science of evolution with doctrinal statements or evangelistic attempts. This can be a central focus of apologetics. For example, in explanation for his "struggle" against evolution, prominent creationist Ken Ham has declared "the Lord has not just called us to knock down evolution, but to help in restoring the foundation of the gospel in our society. We believe that if the churches took up the tool of Creation Evangelism in society, not only would we see a stemming of the tide of humanistic philosophy, but we would also see the seeds of revival sown in a culture which is becoming increasingly more pagan each day."[67] Ham teaches that a rejection of the biblical creation history undermines the relevancy of the Christian gospels and derivatively weakens the moral foundations of society. Disputes relating to science Many creationists vehemently oppose certain scientific theories in a number of ways, including opposition to specific applications of scientific processes, accusations of bias within the scientific community,[68] and claims that discussions within the scientific community reveal or imply a crisis. In response to perceived crises in modern science, creationists claim to have an alternative, typically based on faith, creation science, and/or intelligent design. The scientific community has responded by pointing out that their conversations are frequently misrepresented (e.g. by quote mining) in order to create the impression of a deeper controversy or crisis, and that the creationists' alternatives are generally pseudoscientific. Biology

A phylogenetic tree based on rRNA genes.

Disputes relating to evolutionary biology are central to the controversy between Creationists and the scientific community. The aspects of evolutionary biology disputed include common descent (and particularly

human evolution from common ancestors with other members of the Great Apes), macroevolution, and the existence of transitional fossils. Common descent Main article: Common descent See also: Evidence of common descent and Tree of life (science) [The] Discovery [Institute] presents common descent as controversial exclusively within the animal kingdom, as it focuses on embryology, anatomy, and the fossil record to raise questions about them. In the real world of science, common descent of animals is completely noncontroversial; any controversy resides in the microbial world. There, researchers argued over a variety of topics, starting with the very beginning, namely the relationship among the three main branches of life.John Timmer, Evolution: what's the real controversy?[69]A group of organisms is said to have common descent if they have a common ancestor. A theory of universal common descent based on evolutionary principles was proposed by Charles Darwin and is now generally accepted by biologists. The last universal common ancestor, that is, the most recent common ancestor of all currently living organisms, is believed to have appeared about 3.9 billion years ago. With a few exceptions (e.g. Michael Behe), the vast majority of Creationists reject this theory[citation needed]. Evidence of common descent includes evidence from fossil records, comparative anatomy, geographical distribution of species, comparative physiology and comparative biochemistry. Human evolution Wikibooks has a book on the topic of Introduction to PaleoanthropologyMain article: Human evolution See also: Paleoanthropology and Adam and Eve Human evolution is the study of the biological evolution of humans as a distinct species from its common ancestors with other animals. Analysis of fossil evidence and genetic distance are two of the means by which scientists understand this evolutionary history. Fossil evidence suggests that humans' earliest hominoid ancestors may have split from other primates as early as the late Oligocene, circa 2624 Ma, and that by the early Miocene, the adaptive radiation of many different hominoid forms was well underway.[70] Evidence from the molecular dating of genetic differences indicates that the gibbon lineage (family Hylobatidae) diverged between 18 and 12 Ma, and the orangutan lineage (subfamily Ponginae) diverged about 12 Ma. While there is no fossil evidence thus far clearly documenting the early ancestry of gibbons, fossil proto-orangutans may be represented by Sivapithecus from India and Griphopithecus from Turkey, dated to around 10 Ma. Molecular evidence further suggests that between 8 and 4 Ma, first the gorillas, and then the chimpanzee (genus Pan) split from the line leading to the humans.[71] We have no fossil record of this divergence, but distinctively hominid fossils have been found dating to 3.2 Ma (see Lucy) and possibly even earlier, at 6 or 7 Ma (see Touma).[72] Comparisons of chimpanzee and human DNA show the two are approximately 98.4 percent identical, and are taken as strong evidence of recent common ancestry.[73] Today, only one distinct human species survives, but many earlier species have been found in the fossil record, including Homo erectus, Homo habilis, and Homo neanderthalensis. Creationists dispute there is evidence of shared ancestry in the fossil evidence, and argue either that these are misassigned ape fossils (e.g. that Java man was a gibbon[74]) or too similar to modern humans to designate them as distinct or transitional forms.[75] However

Creationists frequently disagree where the dividing lines would be.[76] Creation myths (such as the Book of Genesis) frequently posit a first man (Adam, in the case of Genesis) as an alternative viewpoint to the scientific account. Creationists also dispute science's interpretation of genetic evidence in the study of human evolution. They argue that it is a "dubious assumption" that genetic similarities between various animals imply a common ancestral relationship, and that scientists are coming to this interpretation only because they have preconceived notions that such shared relationships exist. Creationists also argue that genetic mutations are strong evidence against evolutionary theory because the mutations required for major changes to occur would almost certainly be detrimental.[21] Macroevolution Main article: Macroevolution See also: Speciation Creationists have long argued against the possibility of Macroevolution. Macroevolution is defined by the scientific community to be evolution that occurs at or above the level of species. Under this definition, Macroevolution can be considered to be a fact, as evidenced by observed instances of speciation. Creationists however tend to apply a more restrictive, if vaguer, definition of Macroevolution, often relating to the emergence of new body forms or organs. The scientific community considers that there is strong evidence for even such more restrictive definitions, but the evidence for this is more complex. Recent arguments against (such restrictive definitions of) macroevolution include the Intelligent design arguments of Irreducible complexity and Specified complexity. However, neither argument has been accepted for publication in a peer-reviewed scientific journal, and both arguments have been rejected by the scientific community as pseudoscience. This section requires expansion. Transitional fossils Main article: Transitional fossil See also: List of transitional fossils, Bird evolution, and Evolution of the horse It is commonly stated by critics of evolution that there are no known transitional fossils.[77][78] This position is based on a misunderstanding of the nature of what represents a transitional feature. A common creationist argument is that no fossils are found with partially functional features. It is plausible, however, that a complex feature with one function can adapt a wholly different function through evolution. The precursor to, for example, a wing, might originally have only been meant for gliding, trapping flying prey, and/or mating display. Nowadays, wings can still have all of these functions, but they are also used in active flight.

Reconstruction of Ambulocetus natans As another example, Alan Haywood stated in Creation and Evolution that "Darwinists rarely mention the whale because it presents them with one of their most insoluble problems. They believe that somehow a whale must have evolved from an ordinary land-dwelling animal, which took to the sea and lost its legs ... A land mammal that was in the process of becoming a whale would fall between two stoolsit would not be fitted for life on land or at sea, and would have no hope for survival."[79] The evolution of whales has however been documented in considerable detail, with

Ambulocetus, described as looking like a three-metre long mammalian crocodile, as one of the transitional fossils. Although transitional fossils elucidate the evolutionary transition of one life-form to another, they only exemplify snapshots of this process. Due to the special circumstances required for preservation of living beings, only a very small percentage of all life-forms that ever have existed can be expected to be discovered. Thus, the transition itself can only be illustrated and corroborated by transitional fossils, but it will never be known in detail. However, progressing research and discovery managed to fill in several gaps and continues to do so. Critics of evolution often cite this argument as being a convenient way to explain off the lack of 'snapshot' fossils that show crucial steps between species. The theory of punctuated equilibrium developed by Stephen Jay Gould and Niles Eldredge is often mistakenly drawn into the discussion of transitional fossils. This theory, however, pertains only to welldocumented transitions within taxa or between closely related taxa over a geologically short period of time. These transitions, usually traceable in the same geological outcrop, often show small jumps in morphology between periods of morphological stability. To explain these jumps, Gould and Eldredge envisaged comparatively long periods of genetic stability separated by periods of rapid evolution. For example the change from a creature the size of a mouse, to one the size of an elephant, could be accomplished over 60,000 years, with a rate of change too small to be noticed over any human lifetime. 60,000 years is too small a gap to be identified or identifiable in the fossil record.[citation needed] Geology Main article: Flood Geology See also: Creation geophysics, Geochronology, and Age of the Earth Many believers in Young Earth Creationism a position held by the majority of proponents of Flood Geology accept biblical chronogenealogies (such as the Ussher chronology which in turn is based on the Masoretic version of the Genealogies of Genesis).[80][81] They believe that God created the universe approximately 6000 years ago, in the space of six days. Much of creation geology is devoted to debunking the dating methods used in anthropology, geology, and planetary science that give ages in conflict with the young Earth idea. In particular, creationists dispute the reliability of radiometric dating and isochron analysis, both of which are central to mainstream geological theories of the age of the Earth. They usually dispute these methods based on uncertainties concerning initial concentrations of individually considered species and the associated measurement uncertainties caused by diffusion of the parent and daughter isotopes. However, a full critique of the entire parameter-fitting analysis, which relies on dozens of radionuclei parent and daughter pairs, has not been done by creationists hoping to cast doubt on the technique. The consensus of professional scientific organisations worldwide is that no scientific evidence contradicts the age of approximately 4.5 billion years.[82] Young Earth creationists reject these ages on the grounds of what they regard as being tenuous and untestable assumptions in the methodology. Apparently inconsistent radiometric dates are often quoted to cast doubt on the utility and accuracy of the method. Mainstream proponents who get involved in this debate point out that dating methods only rely on the assumptions that the physical laws governing radioactive decay have not been violated since the sample was formed (harking back to Lyell's doctrine of uniformitarianism). They also point out that the "problems" that creationists publicly mentioned can be shown to either

not be problems at all, are issues with known contamination, or simply the result of incorrectly evaluating legitimate data. Other sciences Cosmology See also: Age of the universe Whilst Young Earth Creationists believe that the Universe was created approximately 6000 years ago, the current scientific consensus is that it is about 13.7 billion years old. The recent science of nucleocosmochronology is extending the approaches used for Carbon-14 dating to the dating of astronomical features. For example based upon this emerging science, the Galactic thin disk of the Milky Way galaxy is estimated to have been formed between 8.3 1.8 billion years ago.[83] Many other creationists, including Old Earth Creationists, do not necessarily dispute these figures. Nuclear physics See also: radiometric dating Creationists point to experiments they have performed, which they claim demonstrate that 1.5 billion years of nuclear decay took place over a short period of time, from which they infer that "billion-fold speed-ups of nuclear decay" have occurred, a massive violation of the principle that radioisotope decay rates are constant, a core principle underlying nuclear physics generally, and radiometric dating in particular.[84] The scientific community points to numerous flaws in these experiments, to the fact that their results have not been accepted for publication by any peer-reviewed scientific journal, and to the fact that the creationist scientists conducting them were untrained in experimental geochronology.[85][86] In refutation of young-Earth claims of inconstant decay rates affecting the reliability of radiometric dating, Roger C. Wiens, a physicist specialising in isotope dating states: There are only three quite technical instances where a half-life changes, and these do not affect the dating methods [under discussion][87]": 1. Only one technical exception occurs under terrestrial conditions, and this is not for an isotope used for dating. ... The artificially-produced isotope, beryllium-7 has been shown to change by up to 1.5%, depending on its chemical environment. ... [H]eavier atoms are even less subject to these minute changes, so the dates of rocks made by electron-capture decays would only be off by at most a few hundredths of a percent. 2. ... Another case is material inside of stars, which is in a plasma state where electrons are not bound to atoms. In the extremely hot stellar environment, a completely different kind of decay can occur. 'Bound-state beta decay' occurs when the nucleus emits an electron into a bound electronic state close to the nucleus. ... All normal matter, such as everything on Earth, the Moon, meteorites, etc. has electrons in normal positions, so these instances never apply to rocks, or anything colder than several hundred thousand degrees. ... 3. The last case also involves very fast-moving matter. It has been demonstrated by atomic clocks in very fast spacecraft. These atomic clocks slow down very slightly (only a second or so per year) as predicted by Einstein's theory of relativity. No rocks in our solar system are going fast enough to make a noticeable change in their dates. ... Roger C. Wiens, Radiometric Dating, A Christian Perspective[88] Misrepresentations of science

Quote mining Main article: Quote mining As a means to criticise mainstream science, creationists have been known to quote, at length, scientists who ostensibly support the mainstream theories, but appear to acknowledge criticisms similar to those of creationists.[89] However, almost universally these have been shown to be quote mines that do not accurately reflect the evidence for evolution or the mainstream scientific community's opinion of it, or highly out-ofdate.[90][91] Many of the same quotes used by creationists have appeared so frequently in Internet discussions due to the availability of cut and paste functions, that the TalkOrigins Archive has created "The Quote Mine Project" for quick reference to the original context of these quotations.[90] Public policy issues Science education Main article: Creation and evolution in public education See also: Teach the Controversy Creationists promote that evolution is a theory in crisis[92][93] with scientists criticizing evolution[94] and claim that fairness and equal time requires educating students about the alleged scientific controversy. Opponents, being the overwhelming majority of the scientific community and science education organizations,[95] reply that there is in fact no scientific controversy and that the controversy exists solely in terms of religion and politics.[92][96] The American Association for the Advancement of Science and other science and education professional organizations say that Teach the Controversy proponents seek to undermine the teaching of evolution[92][97] while promoting intelligent design,[98][99][100] and to advance an education policy for US public schools that introduces creationist explanations for the origin of life to public-school science curricula.[101][102] This viewpoint was supported by the December 2005 ruling in the Kitzmiller v. Dover Area School District trial.[102] George Mason University Biology Department introduced a course on the creation/evolution controversy, and apparently as students learn more about biology, they find objections to evolution less convincing, suggesting that teaching the controversy rightly as a separate elective course on philosophy or history of science, or "politics of science and religion," would undermine creationists criticisms, and that the scientific communitys resistance to this approach was bad public relations.[103] Freedom of speech Creationists have claimed that preventing them from teaching Creationism violates their right of Freedom of speech. However court cases (such as Webster v. New Lenox School District and Bishop v. Aronov) have upheld school districts' and universities' right to restrict teaching to a specified curriculum. Issues relating to religion See also: Relationship between religion and science and Evolution and the Roman Catholic Church Theological arguments

See also: Allegorical interpretations of Genesis and Evolutionary argument against naturalism This section requires expansion. Religion and historical scientists Creationists often argue that Christianity and literal belief in the Bible are either foundationally significant or directly responsible for scientific progress.[104] To that end, Institute for Creation Research founder Henry M. Morris has enumerated scientists such as astronomer and philosopher Galileo, mathematician and theoretical physicist James Clerk Maxwell, mathematician and philosopher Blaise Pascal, geneticist monk Gregor Mendel, and Isaac Newton as believers in a biblical creation narrative.[105] This argument usually involves scientists either who were no longer alive when evolution was proposed or whose field of study didn't include evolution. The argument is generally rejected as specious by those who oppose creationism.[106] Many of the scientists in question did some early work on the mechanisms of evolution, e.g., the Modern evolutionary synthesis combines Darwin's Evolution with Mendel's theories of inheritance and genetics. Though biological evolution of some sort had become the primary mode of discussing speciation within science by the late-19th century, it was not until the mid-20th century that evolutionary theories stabilized into the modern synthesis. Some of the historical scientists marshalled by creationists were dealing with quite different issues than any are engaged with today: Louis Pasteur, for example, opposed the theory of spontaneous generation with biogenesis, an advocacy some creationists describe as a critique on chemical evolution and abiogenesis. Pasteur accepted that some form of evolution had occurred and that the Earth was millions of years old.[107] The relationship between science and religion was not portrayed in antagonistic terms until the late-19th century, and even then there have been many examples of the two being reconcilable for evolutionary scientists.[108] Many historical scientists wrote books explaining how pursuit of science was seen by them as fulfillment of spiritual duty in line with their religious beliefs. Even so, such professions of faith were not insurance against dogmatic opposition by certain religious people. Some extensions to this creationist argument have included the incorrect suggestions that Einstein's deism was a tacit endorsement of creationism or that Charles Darwin converted on his deathbed and recanted evolutionary theory. Forums for the controversy Debates Many creationists and scientists engage in frequent public debates regarding the origin of human life, hosted by a variety of institutions. However, some scientists disagree with this tactic, arguing that by openly debating supporters of supernatural origin explanations (creationism and intelligent design), scientists are lending credibility and unwarranted publicity to creationists, which could foster an inaccurate public perception and obscure the factual merits of the debate.[109] For example, in May 2004 Dr. Michael Shermer debated creationist Kent Hovind in front of a predominately creationist audience. In Shermer's online reflection while he was explaining that he won the debate with intellectual and scientific evidence he felt it was "not an intellectual exercise," but rather it was "an emotional drama."[clarification needed][110] While receiving positive responses

from creationist observers, Shermer concluded "Unless there is a subject that is truly debatable (evolution v. creation is not), with a format that is fair, in a forum that is balanced, it only serves to belittle both the magisterium of science and the magisterium of religion."[110] (see: scientific method). Others, like evolutionary biologist Massimo Pigliucci, have debated Hovind, and have expressed surprise to hear Hovind try "to convince the audience that evolutionists believe humans came from rocks" and at Hovind's assertion that biologists believe humans "evolved from bananas."[111][clarification needed] Eugenie Scott of the National Center for Science Education, a non-profit organization dedicated to defending the teaching of evolution in the public schools, claimed debates are not the sort of arena to promote science to creationists.[110] Scott says that "Evolution is not on trial in the world of science," and "the topic of the discussion should not be the scientific legitimacy of evolution" but rather should be on the lack of evidence in creationism. Similarly, Stephen Jay Gould took a public stance against appearing to give legitimacy to creationism by debating its proponents. He noted during a Caltech lecture in 1985:[112] Debate is an art form. It is about the winning of arguments. It is not about the discovery of truth. There are certain rules and procedures to debate that really have nothing to do with establishing fact which creationists have mastered. Some of those rules are: never say anything positive about your own position because it can be attacked, but chip away at what appear to be the weaknesses in your opponent's position. They are good at that. I don't think I could beat the creationists at debate. I can tie them. But in courtrooms they are terrible, because in courtrooms you cannot give speeches. In a courtroom you have to answer direct questions about the positive status of your belief. We destroyed them in Arkansas. On the second day of the two-week trial we had our victory party! Political lobbying See also: Politics of creationism, Kansas evolution hearings, Santorum Amendment, and List of scientific societies rejecting intelligent design A wide range of organisations, on both sides of the controversy, are involved in lobbying in an attempt to influence political decisions relating to the teaching of evolution, at a number of levels. These include the Discovery Institute, the National Center for Science Education, the National Science Teachers Association, state Citizens Alliances for Science, and numerous national science associations and state Academies of Science.[113] This section requires expansion. In the media The controversy has been discussed in numerous newspaper articles, reports, op-eds and letters to the editor, as well as a number of radio and television programmes (including the PBS series, Evolution and Coral Ridge Ministries' Darwin's Deadly Legacy). This has led some commentators to express a concern at what they see as a highly inaccurate and biased understanding of evolution among the general public. Pulitzer Prizewinning journalist and writer Edward Humes states:[114] There are really two theories of evolution. There is the genuine scientific theory and there is the talk-radio pretend version, designed not to enlighten but to deceive and enrage. The talk-radio version had a packed town hall up in arms at the "Why Evolution Is Stupid" lecture. In this version of the theory, scientists supposedly believe that all life is accidental, a random crash of molecules that magically produced flowers, horses and humans -- a scenario as unlikely as a tornado in a junkyard assembling a 747. Humans come from monkeys in this theory, just popping into existence one day.

The evidence against Darwin is overwhelming, the purveyors of talk-radio evolution rail, yet scientists embrace his ideas because they want to promote atheism. Outside the United States

Views on human evolution in other countries While the controversy has been prominent in the United States, it has flared up in other countries as well.[115][116][117] Europe Europeans have often regarded the creation-evolution controversy as an American matter.[116] However, in recent years the conflict has become an issue in a variety of countries including Germany, The United Kingdom, Italy, the Netherlands, Poland and Serbia.[116][117][118][119] On 17 September 2007 the Committee on Culture, Science and Education of the Parliamentary Assembly of the Council of Europe issued a report on the attempt by American inspired creationists to promote creationism in European schools. It concludes "If we are not careful, creationism could become a threat to human rights which are a key concern of the Council of Europe.... The war on the theory of evolution and on its proponents most often originates in forms of religious extremism which are closely allied to extreme right-wing political movements... some advocates of creationism are out to replace democracy by theocracy."[120] Australia With declining church attendance, there has been some growth in fundamentalist and pentecostal Christian denominations.[121] Under the former Queensland state government of Joh Bjelke-Petersen, in 1980 lobbying was so successful that Queensland allowed the teaching of creationism as science to school children. Public lectures have been given in rented rooms at Universities, by visiting American speakers, and speakers with doctorates purchased by mail from Florida sites.[122] One of the most acrimonious aspects of the Australian debate was featured on the science television program Quantum, about a long-running and ultimately unsuccessful court case by Ian Plimer, Professor of Geology at Melbourne University, against an ordained minister, Dr. Allen Roberts, who had claimed that there were remnants of Noah's Ark in eastern Turkey. Although the court found that Dr Roberts had made false and misleading claims, they were not made in the course of trade or commerce, so the case failed.[123] Islamic countries See also: Islamic creationism In recent times, the controversy has become more prominent in Islamic countries.[124] Currently, in Egypt evolution is taught in schools but Saudi Arabia and Sudan have both banned the teaching of evolution in schools.[115] Creation science has also been heavily promoted in Turkey and in immigrant communities in Western Europe, primarily by Harun Yahya.[117] See also * Articles related to the creation-evolution controversy * Allegorical interpretations of Genesis * Anti-intellectualism * Clergy Letter Project * Creation science * Creationism

* * * * * * * * * * * * * * * * * *

Evidence of common descent Evolution and the Roman Catholic Church Evolution Sunday Evolutionary origin of religions Hindu views on evolution History of the creation-evolution controversy Intelligent design Jainism and non-creationism Jewish views on evolution Level of support for evolution List of participants in the creation-evolution controversy Mormonism and evolution Natural theology Objections to evolution Politics of creationism Project Steve Relationship between religion and science Teach the Controversy

Footnotes 1. ^ See Hovind 2006, for example. 2. ^ An Index to Creationist Claims , Mark Isaak, Talkorigins Archive,Copyright 2006. 3. ^ Curry, Andrew (27 February 2009). "Creationist Beliefs Persist in Europe". Science 323 (5918): 1159. doi:10.1126/science.323.5918.1159. PMID 19251601. "News coverage of the creationism-versus-evolution debate tends to focus on the United States ... But in the past 5 years, political clashes over the issue have also occurred in countries all across Europe. ... "This isn't just an American problem," says Dittmar Graf of the Technical University of Dortmund, who organized the meeting". 4. ^ Larson 2004, p. 247-263 Chapter titled Modern Culture Wars. See also Ruse 1999, p. 26, who writes "One thing that historians delighted in showing is that, contrary to the usually held tale of science and religion being always opposed...religion and theologically inclined philosophy have frequently been very significant factors in the forward movement of science." 5. ^ Numbers 1992, p. 3-240 6. ^ See: o Peters & Hewlett 2005, p. 1; o Kitzmiller v. Dover Area School District, page 20 7. ^ See: o Battle on Teaching Evolution Sharpens, Peter Slevin, Washington Post, Monday, 14 March 2005, Page A01; o The Political Design of Intelligent Design, Russell D. Renka, 16 November 2005; o Politicized Scholars Put Evolution on the Defensive, Jody Wilgoren, The New York Times, 21 April 2005 o The Newest Evolution of Creationism, Barbara Forrest, Natural History, April, 2002, page 80; o Kitzmiller vs. Dover Area School District, pages 7-9, also pages 64-90 8. ^ Myers 2006; NSTA 2007; IAP 2006; AAAS 2006; and Pinholster 2006; Ruling, Kitzmiller v. Dover page 83 9. ^ Larson 2004, p. 258 "Virtually no secular scientists accepted the doctrines of creation science; but that did not deter creation scientists from advancing scientific arguments for their position." See also Martz & McDaniel 1987, p. 23, a Newsweek article which states "By one count there are some 700 scientists (out of a total of 480,000 U.S. earth and life

scientists) who give credence to creation-science, the general theory that complex life forms did not evolve but appeared 'abruptly'." 10. ^ Committee on Revising Science and Creationism, A View from the National Academy of Sciences, National Academy of Sciences and Institute of Medicine of the National Academies (2008). Science, Evolution, and Creationism. National Academy of Sciences. p. 12. ISBN 0-309-10586-2. http://www.nap.edu/catalog.php?record_id=11876. 11. ^ Desmond & Moore 1991, p. 34-35 12. ^ See": o van Wyhe 2006; o Desmond & Moore 1991, p. 321-323, 503-505. 13. ^ a b c d e AAAS Evolution Dialogues: Science, Ethics and Religion study guide (pdf) 14. ^ See: o Hodge 1874, p. 177; o Numbers 1992, p. 14; o Burns, Ralph, Lerner, & Standish 1982, p. 965; o Huxley 1902 15. ^ Numbers 1992, p. 14 16. ^ Numbers 1992, p. 14-15 17. ^ a b Numbers 1992, p. 17 18. ^ Numbers 1992, p. 18, noting that this applies to published or public skeptics. Many or most Christians may have held on to a literal six days of creation, but these views were rarely expressed in books and journals. Exceptions are also noted, such as literal interpretations published by Eleazar Lord (1788-1871) and David Nevins Lord (1792-1880). However, the observation that evolutionary critics had a relaxed interpretation of Genesis is supported by specifically enumerating: Louis Agassiz (1807-1873); Arnold Henry Guyot (1807-1884); John William Dawson (1820-1899); Enoch Fitch Burr (1818-1907); George D. Armstrong (18131899); Charles Hodge, theologian (1797-1878); James Dwight Dana (18131895); Edward Hitchcock, clergyman and respected Amherst College geologist, (1793-1864); Reverend Herbert W. Morris (1818-1897); H. L. Hastings (1833?-1899); Luther T. Townsend (1838-1922; Alexander Patterson, Presbyterian evangelist who published The Other Side of Evolution Its Effects and Fallacy 19. ^ Numbers(2006) p161 20. ^ See: o s:Kitzmiller v. Dover Area School District/2:Context#Page 19 of 139; o Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. (pdf) A Position Paper from the Center for Inquiry, Office of Public Policy Barbara Forrest. May, 2007; o TalkOrigins Archive: Post of the Month: March 2006, The History of Creationism by Lenny Flank. 21. ^ a b Nelkin, Dorothy (2000), The Creation Controversy: Science or Scripture in Schools, New York: iUniverse, pp. 242, ISBN 0-595-00194-7 22. ^ Epperson et al. v. Arkansas, 393 U.S. 97 (U.S. Supreme Court 196811-12). 23. ^ Larson, Edward J. (2003), Trial and Error: The American Controversy Over Creation and Evolution, Oxford University Press, pp. 276, ISBN 0195154703 24. ^ Larson 2004, p. 248,250, see also Dobzhansky 1973 25. ^ Larson 2004, p. 251 26. ^ Larson 2004, p. 252 27. ^ Larson 2004, p. 255,Numbers 1992, p. xi,200-208 28. ^ Larson 2004, p. 255 29. ^ Numbers 1992, p. 284-285 30. ^ Numbers 1992, p. 284-6

31. ^ Quoting Larson 2004, p. 255-256: "Fundamentalists no longer merely denounced Darwinism as false; they offered a scientific-sounding alternative of their own, which they called either 'scientific creationism (as distinct from religious creationism) or 'creation science' (as opposed to evolution science." 32. ^ Larson 2004, p. 254-255, Numbers 1998, p. 5-6 33. ^ Ruling, Kitzmiller v. Dover Area School District pp 7-9. 34. ^ "NCSE Resource". Evolving Banners at the Discovery Institute. 200208-29. http://ncseweb.org/creationism/general/evolving-banners-atdiscovery-institute. Retrieved on 2009-04-07. 35. ^ "Top Questions-1.What is the theory of intelligent design?". Discovery Institute. http://www.discovery.org/csc/topQuestions.php#questionsAboutIntelligentDe sign. Retrieved on 2007-05-13. . 36. ^ Verderame 2007,Simon 2006 37. ^ Dewey 1994, p. 31, and Wiker 2003, summarizing Gould. 38. ^ Larson 2004, p. 258 "Virtually no secular scientists accepted the doctrines of creation science; but that did not deter creation scientists from advancing scientific arguments for their position." See also Martz & McDaniel 1987, p. 23, a Newsweek article which states "By one count there are some 700 scientists (out of a total of 480,000 U.S. earth and life scientists) who give credence to creation-science, the general theory that complex life forms did not evolve but appeared 'abruptly'." 39. ^ Statement from the Council of the Biological Society of Washington 40. ^ Bumiller 2005, Peters & Hewlett 2005, p. 3 41. ^ Some question group's move with elections nearing 6News Lawrence, Lawrence Journal-World. 7 July 2006. 42. ^ Evolutions foes lose ground in Kansas MSNBC, 2 August 2006. 43. ^ Evolution of Kansas science standards continues as Darwin's theories regain prominence The Associated Press, via the International Herald Tribune, 13 February 2007. 44. ^ The "Evolution" of Creationism Timeline: how creationism has "evolved". People for the American Way. 45. ^ Johnson 1998, Hodge 1874, p. 177, Wiker 2003, Peters & Hewlett 2005, p. 5--Peters and Hewlett argue that the atheism of many evolutionary supporters must be removed from the debate 46. ^ Lenski 2000, p. Conclusions 47. ^ Johnson 1998 48. ^ Einstein 1930, p. 1-4 49. ^ Dawkins 1997 50. ^ a b Free Executive Summary, Science and Creationism: A View from the National Academy of Sciences, Second Edition, Steering Committee on Science and Creationism, National Academy of Sciences, 1999, ISBN 978-0309-06406-4. 51. ^ Johnson 1993, p. 63, Tolson 2005, Moran 1993 ; Selman v. Cobb County School District. US District Court for the Northern District of Georgia (2005); Talk. Origins; Bill Moyers et al, 2004. "Now with Bill Moyers." PBS. Accessed 2006-01-29. Interview with Richard Dawkins 52. ^ Merriam-Webster online dictionary. www.m-w.com 53. ^ Gould 1981 54. ^ See: o Number 1992, p. 247; o Wilkins, John S, Evolution and Philosophy: Is Evolution Science, and What Does 'Science' Mean?, TalkOrigins Archive 55. ^ Popper 1976, p. 168 and 172 quoted in Kofahl 1981 56. ^ Unknown sociologist quoted in Numbers 1992, p. 247 57. ^ Kofahl 1989 as quoted by Numbers 1992, p. 247 58. ^ Lewin 1982

59. ^ Numbers(2006) p274 60. ^ Kofahl 1981, p. 873 61. ^ Talkorigins summary of Karl Popper attitudes towards evolution 62. ^ See: o Natural selection and the emergence of mind, Karl Popper, Dialectica 32(3/4): 339355, 1978 o Did Popper refute evolution?, Massimo Pigliucci, Skeptical Inquirer, Sept-Oct 2004 63. ^ Ruse 1999, p. 13-37, which discusses conflicting ideas about science among Karl Popper, Thomas Samuel Kuhn, and their disciples. 64. ^ As quoted by Wallis 2005, p. 32. Also see Dawkins 1986 and Dawkins 1995 65. ^ Wallis 2005, p. 6 Dawkins quoting Haldane 66. ^ Dorman 1996 67. ^ Ham, Ken. Creation Evangelism (Part II of Relevance of Creation). Creation Magazine '6'(2):17, November 1983. 68. ^ Johnson 1993, p. 69 where Johnson cites three pages spent in Issac Asimov's New Guide to Science that take creationists to task, while only spending one half page on evidence of evolution. 69. ^ Evolution: what's the real controversy?, John Timmer, Nobel Intent, 7 May 2008 70. ^ Stringer, Chris; Andrews, Peter (2005), The complete world of human evolution, London: Thames & Hudson, pp. 240, ISBN 0-500-05132-1 71. ^ Relethford, John (2003), Reflections of our past: how human history is revealed in our genes, Boulder, Colo: Westview Press, pp. 257, ISBN 08133-3958-8 72. ^ "Touma the Human Ancestor: Skull of Oldest Known Hominid Unearthed in Chad". NPR: All Things Considered. http://www.npr.org/programs/atc/features/2002/july/toumai/index.html. Retrieved on 2009-02-21. 73. ^ Chimps are human, gene study implies New Scientist, website, 19 May 2003 74. ^ Was Java Man a gibbon?, Jim Foley, TalkOrigins website, 30 April 2003. 75. ^ See disputes over the classification of Neanderthals in The Counter-Creationism Handbook, Mark Isaak, University of California Press (2007), ISBN 978-0520249264 76. ^ Comparison of all skulls, Jim Foley, TalkOrigins website, 8 August 2005. 77. ^ Scientific Creationism, Henry M. Morris, 1985, pp. 78-90 78. ^ Life--How Did It Get Here?, Watchtower Bible and Tract Society, 1985, pp. 57-59 79. ^ Haywood, Alan (1985) Creation and Evolution.Triangle Books, London. Quoted in Hooking Leviathan by Its Past, Stephen Jay Gould 80. ^ Biblical chronogenealogies 81. ^ The Meaning of the Chronogenealogies of Genesis 5 and 11 82. ^ IAP Statement on the Teaching of Evolution, Interacademy Panel on Global Issues, 21 June 2006. 83. ^ Del Peloso, E.F.; Da Silva, L.; De Mello, G.F.P.; Arany-prado, L.I. (2005). "The age of the Galactic thin disk from Th/Eu nucleocosmochronology". A&A 434: 301308. doi:10.1051/00046361:20047060. 84. ^ Nuclear Decay: Evidence For A Young World, D. Russell Humphreys, Impact, Number 352, October 2002. 85. ^ Young-Earth Creationist Helium Diffusion "Dates" Fallacies Based on Bad Assumptions and Questionable Data, Kevin R. Henke, TalkOrigins website, Original version: 17 March 2005, Revision: 24 November 2005.

86. ^ R.A.T.E: More Faulty Creation Science from The Institute for Creation Research, J. G. Meert, Gondwana Research, The Official Journal of the International Association for Gondwana, 13 November 2000 (updated 6 February 2003). 87. ^ Dating methods discussed were potassium-argon dating, argon-argon dating, rubidium-strontium dating, samarium-neodymium dating, lutetiumhafnium, rhenium-osmium dating, and uranium-lead dating. 88. ^ Radiometric Dating, A Christian Perspective, Roger C. Wiens, American Scientific Affiliation, p20-21 89. ^ Dobzhansky 1973 90. ^ a b Pieret 2006 91. ^ o Isaak, Mark (2004). "Index to Creationist Claims: Claim CA113". Talk.origins. http://www.toarchive.org/indexcc/CA/CA113.html. Retrieved on 2007-12-27. o Dunford, Mike (2007-07-02). "A new (mis)take on an old paper (and other posts)". [[The Panda's Thumb (blog)|]]. http://www.pandasthumb.org/archives/quote_mines/. Retrieved on 2007-1226. o Myers, PZ (2004-09-11). "I'm shocked, shocked to find that quote mining is going on in there!". Pharyngula (blog). http://pharyngula.org/index/weblog/comments/im_shocked_shocked_to_find_th at_quote_mining_is_going_on_in_there. Retrieved on 2007-12-27. 92. ^ a b c "Some bills seek to discredit evolution by emphasizing socalled "flaws" in the theory of evolution or "disagreements" within the scientific community. Others insist that teachers have absolute freedom within their classrooms and cannot be disciplined for teaching nonscientific "alternatives" to evolution. A number of bills require that students be taught to "critically analyze" evolution or to understand "the controversy." But there is no significant controversy within the scientific community about the validity of the theory of evolution. The current controversy surrounding the teaching of evolution is not a scientific one." AAAS Statement on the Teaching of Evolution American Association for the Advancement of Science. 16 February 2006 (PDF file) 93. ^ Ruling, Kitzmiller v. Dover Area School District, page 89 94. ^ "That this controversy is one largely manufactured by the proponents of creationism and intelligent design may not matter, and as long as the controversy is taught in classes on current affairs, politics, or religion, and not in science classes, neither scientists nor citizens should be concerned." Intelligent Judging Evolution in the Classroom and the Courtroom George J. Annas, New England Journal of Medicine, Volume 354:2277-2281 25 May 2006 95. ^ See: 1) List of scientific societies rejecting intelligent design 2) Kitzmiller v. Dover page 83. The Discovery Institute's Dissent From Darwin Petition has been signed by about 500 scientists. The AAAS, the largest association of scientists in the U.S., has 120,000 members, and firmly rejects intelligent design and denies that there is a legitimate scientific controversy. More than 70,000 Australian scientists and educators condemn teaching of intelligent design in school science classes. Statements from Scientific Organizations on the status intelligent design and other forms of creationism. 96. ^ "That this controversy is one largely manufactured by the proponents of creationism and intelligent design may not matter, and as long as the controversy is taught in classes on current affairs, politics, or religion, and not in science classes, neither scientists nor citizens should be concerned." Intelligent Judging Evolution in the Classroom and the Courtroom George J. Annas, New England Journal of Medicine, Volume 354:2277-2281 25 May 2006

97. ^ "In summary, the disclaimer singles out the theory of evolution for special treatment, misrepresents its status in the scientific community, causes students to doubt its validity without scientific justification, presents students with a religious alternative masquerading as a scientific theory, directs them to consult a creationist text as though it were a science resource, and instructs students to forgo scientific inquiry in the public school classroom and instead to seek out religious instruction elsewhere." Ruling - disclaimer, pg. 49 Kitzmiller v. Dover Area School District. 98. ^ "ID's home base is the Center for Science and Culture at Seattle's conservative Discovery Institute. Meyer directs the center; former Reagan adviser Bruce Chapman heads the larger institute, with input from the Christian supply-sider and former American Spectator owner George Gilder (also a Discovery senior fellow). From this perch, the ID crowd has pushed a "teach the controversy" approach to evolution that closely influenced the Ohio State Board of Education's recently proposed science standards, which would require students to learn how scientists "continue to investigate and critically analyze" aspects of Darwin's theory." Chris Mooney. The American Prospect. 2 December 2002 Survival of the Slickest: How anti-evolutionists are mutating their message 99. ^ Teaching Intelligent Design: What Happened When? by William A. Dembski"The clarion call of the intelligent design movement is to "teach the controversy." There is a very real controversy centering on how properly to account for biological complexity (cf. the ongoing events in Kansas), and it is a scientific controversy." 100. ^ Nick Matzke's analysis shows how teaching the controversy using the Critical Analysis of Evolution model lesson plan is a means of teaching all the intelligent design arguments without using the intelligent design label.No one here but us Critical Analysis-ists... Nick Matzke. The Panda's Thumb, 11 July 2006 101. ^ "has the effect of implicitly bolstering alternative religious theories of origin by suggesting that evolution is a problematic theory even in the field of science." . . . The effect of Defendants actions in adopting the curriculum change was to impose a religious view of biological origins into the biology course, in violation of the Establishment Clause. Kitzmiller v. Dover Area School District, Conclusion, page 134 102. ^ a b "ID's backers have sought to avoid the scientific scrutiny which we have now determined that it cannot withstand by advocating that the controversy, but not ID itself, should be taught in science class. This tactic is at best disingenuous, and at worst a canard. The goal of the IDM is not to encourage critical thought, but to foment a revolution which would supplant evolutionary theory with ID."Kitzmiller v. Dover Area School District, whether ID is science, page 89 103. ^ AAAS Dialogue on Science, Ethics, and Religion, 20 April 2006, Emmett Holman, Associate Professor of Philosophy from George Mason University, retrieved 2007-04-29 104. ^ Woods 2005, p. 67-114, Chapter Five: The Church and Science 105. ^ Morris 1982 106. ^ Index to Creationist Claims - Claim CA114 edited by Mark Isaak. 2005 107. ^ Index to Creationist Claims - Claim CA114.22 edited by Mark Isaak. 2005 108. ^ [1] Science and religion: Conflicts & occasional agreements 109. ^ Why I Won't Debate Creationists, Richard Dawkins, Reason : In the News, richarddawkins.net, the official Richard Dawkins website, Monday, 15 May 2006.

110. ^ a b c Shermer, Michael (2004-05-10). "Then a Miracle Occurs: An Obstreperous Evening with the Insouciant Kent Hovind, Young Earth Creationist and Defender of the Faith". eSkeptic Online. http://www.skeptic.com/eskeptic/04-05-10.html#miracle. Retrieved on 200702-11. 111. ^ Massimo Pigliucci. Denying Evolution: Creationism, Scientism, and the Nature of Science. (Sinauer, 2002): ISBN 0878936599 page 102. 112. ^ Shermer, Michael. 'Why People Believe Weird Things', Owl Books, 2002. Paperback ed, p. 153. 113. ^ Statements from Scientific Organizations, NCSE 114. ^ Unintelligent designs on Darwin, Edward Humes, Pittsburgh TribuneReview 115. ^ a b Pitock, Todd (06 2007). "Science and Islam". Discover: 3645. 116. ^ a b c Gregory Katz (2008-02-16). "Clash Over Creationism Is Evolving In Europe's Schools". Associated Press. http://www2.tbo.com/content/2008/feb/16/na-clash-over-creationism-isevolving-in-europes-s. Retrieved on 2008-02-17. / 117. ^ a b c Taner Edis. "Cloning Creationism in Turkey". RNCSE 19 (6): 30-35. National Center for Science Education. http://ncseweb.org/rncse/19/6/cloning-creationism-turkey. Retrieved on 2008-02-17. 118. ^ "Serbia reverses Darwin suspension". BBC. 2004-09-09. http://news.bbc.co.uk/1/hi/world/europe/3642460.stm. Retrieved on 200802-17. 119. ^ Roger Highfield (2007-02-10). "Creationists rewrite natural history". The Telegraph. http://www.telegraph.co.uk/earth/main.jhtml?view=DETAILS&grid=&xml=/earth /2007/10/02/scihist102.xml. Retrieved on 2008-02-17. 120. ^ New Scientist 10 November 2007, p. 72 121. ^ Christianity Pentecostalism Australian Broadcasting Corporation 122. ^ Plimer, Ian "Telling lies for God- Reason versus Creationism", (Random House) 123. ^ "Telling Lies for God"? - One Man's Crusade, accessed 2008-02-05, Quantum. See transcript link for detail. 124. ^ "Evolution and religion: In the beginning". The Economist. 200704-19. http://www.economist.com/world/displaystory.cfm?story_id=9036706. Retrieved on 2007-04-25. This article gives a worldwide overview of recent developments on the subject of the controversy. References * AAAS, American Association for the Advancement of Science (2006-02-16), Statement on the Teaching of Evolution, aaas.org, <http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf>. Retrieved on 2007-01-14 * Bumiller, Elisabeth (2005), "Bush Remarks Roil Debate on Teaching of Evolution", The New York Times (no. 2005-08-03), <http://www.nytimes.com/2005/08/03/politics/03bush.html?ex=1280721600&en= 8bbf73d2f5204260&ei=5088&partner=r>. Retrieved on 2007-02-03 * Burns, Edward M.; Philip Lee Ralph & Robert E. Lerner et al. (1982), World Civilizations Their History and Their Culture (Sixth ed.), W.W. Norton & Company, ISBN 0-393-95077-8 * Dawkins, Richard (1986), The Blind Watchmaker, W. W. Norton & Company, Inc., ISBN 0-393-31570-3 * Dawkins, Richard (1995), River Out of Eden, Basic Books, ISBN 0-46506990-8 * Dawkins, Richard (January 1997), "Is Science a Religion?", Humanist, <http://www.thehumanist.org/humanist/articles/dawkins.html>. Retrieved on 2007-01-30

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* Moran, Laurence (1993), Evolution is a Fact and a Theory, TalkOrigins Archive Foundation, <http://www.toarchive.org/faqs/cre-error.html>. Retrieved on 2007-01-23 * Morris, Henry M. (1982-01-01), "Bible-Believing Scientists of the Past", Impact (Institute for Creation Research) 103, <http://www.icr.org/index.php?module=articles&action=view&ID=185>. Retrieved on 2007-01-20 * Myers, P.Z. (2006-06-18), "Ann Coulter: No Evidence for Evolution?", Pharyngula (ScienceBlogs), <http://scienceblogs.com/pharyngula/2006/06/ann_coulter_no_evidence_for_e v.php>. Retrieved on 2007-09-12 * NSTA, National Science Teachers Association (2007), An NSTA Evolution Q&A, http://www.nsta.org/publications/evolution.aspx, retrieved on 200802-01 Archive link * Numbers, Ronald L. (1992), Creationists: The Evolution of Scientific Creationism, Alfred A. Knopf, Inc., pp. 224, ISBN 0-679-40104-0 * Numbers, Ronald (2006-11-30). The Creationists: From Scientific Creationism to Intelligent Design, Expanded Edition. Harvard University Press. p. 624 pages. ISBN 0674023390. * Numbers, Ronald L. (1998-11-15), Darwinism Comes to America, Harvard University Press, 224, ISBN 0674193121 * Peters, Ted & Martinez Hewlett (2005-12-22), The Evolution Controversy: Who's Fighting with Whom about What?, Pacific Luthern Theological Seminary, <http://www.plts.edu/docs/EvolutionBrief2.pdf>. Retrieved on 2007-01-28 * Pieret, John (2006), The Quote Mine Project Or, Lies, Damned Lies and Quote Mines, TalkOrigins Archive Foundation, <http://www.toarchive.org/faqs/quotes/mine/project.html>. Retrieved on 2007-01-23 * Pinholster, Ginger (2006-02-19), AAAS Denounces Anti-Evolution Laws as Hundreds of K-12 Teachers Convene for 'Front Line' Event, aaas.org, <http://www.aaas.org/news/releases/2006/0219boardstatement.shtml>. Retrieved on 2007-01-14 * Popper, Karl (1976), Unended Quest: An Intellecutal Autobiography, Open Court Publishing Co., ISBN 0-87548-366-6 * Popper, Karl (1980), "Letter to the Editor", New Scientist 87 * Ruse, Michael (1999-04-30), Mystery of Mysteries: Is Evolution a Social Construction, Harvard University Press, 320, ISBN 0-674-46706-X * Simon, Stephanie (2006), "Their Own Version of a Big Bang: Those who believe in creationism -- children and adults -- are being taught to challenge evolution's tenets in an in-your-face way.", Los Angeles Times (no. 2006-02-11), <http://philosophy.tamucc.edu/article.pl?sid=06/02/12/1727208&mode=thread >. Retrieved on 2007-02-03 * Tolson, Jay (2005-09-05), Religion in America, "Religion in America: Intelligent Design on Trial", U.S. News & World Report, <http://www.usnews.com/usnews/culture/articles/050928/28religion.htm>. Retrieved on 2007-01-6 * van Wyhe, John (2006), Charles Darwin: gentleman naturalist: A biographical sketch, <http://darwin-online.org.uk/darwin.html>. Retrieved on 2007-01-24 * Verderame, John (2007), Creation evangelism: cutting through the excess, answersingenesis.org, <http://www.answersingenesis.org/docs2001/0510news.asp>. Retrieved on 2007-02-07 * Wallis, Claudia (2005-08-07), "The Evolution Wars", Time Magazine, <http://www.time.com/time/magazine/article/0,9171,1090909-1,00.html> Retrieved on 2007-01-31

* Wiker, Benjamin D. (July/August 2003), "Does Science Point to God? Part II: The Christian Critics", Crisis Politics, Culture, and the Church (CRISIS Magazine), <http://www.crisismagazine.com/julaug2003/feature1.htm>. Retrieved on 2007-01-21 * Woods, Thomas E., Jr. (2005), How the Catholic Church Built Western Civilization, Regnery Publishing, Inc., ISBN 0-89526-038-7 Published books and other resources * Burian, RM: 1994. Dobzhansky on Evolutionary Dynamics: Some Questions about His Russian Background; in Adams, Mark A. (1994). The Evolution of Theodosius Dobzhansky: essays on his life and thought in Russia and America. Princeton, N.J: Princeton University Press. ISBN 0-691-03479-6. * Butler, Samuel (2007). Evolution, Old And New: Or The Theories Of Buffon, Dr. Erasmus Darwin And Lamarck, As Compared With That Of Charles Darwin (1911). Kessinger Publishing, LLC. ISBN 0548641323. * Beer, Gillian; Darwin, Charles (1996). The origin of species. Oxford [Oxfordshire]: Oxford University Press. ISBN 019283438X. * Gould, Stephen Jay; Dobzhansky, Theodosius Grigorievich (1982). Genetics and the origin of species. New York: Columbia University Press. ISBN 0231054750. * Henig, Robin Marantz (2000). The monk in the garden: the lost and found genius of Gregor Mendel, the father of genetics. Boston: Houghton Mifflin. ISBN 0395977657. * Kutschera U, Niklas KJ (2004). "The modern theory of biological evolution: an expanded synthesis". Naturwissenschaften 91 (6): 25576. doi:10.1007/s00114-004-0515-y. PMID 15241603. * Mayr, Ernst (1985). The Growth of Biological Thought: Diversity, Evolution, and Inheritance. Cambridge, Mass: Belknap Press. ISBN 0674364465. * Miller, James H. (2001). An evolving dialogue: theological and scientific perspectives on evolution. Valley Forge, Pa: Trinity Press International. ISBN 1-56338-349-7. * Morris, HR (1963). The Twilight of Evolution. Grand Rapids, MI: Baker Pub Group. ISBN 0801058627. * Numbers, Ronald L. (1993). The creationists. Berkeley: University of California Press. ISBN 0520083938. * Pennock RT (2003). "Creationism and intelligent design". Annu Rev Genomics Hum Genet 4: 14363. doi:10.1146/annurev.genom.4.070802.110400. PMID 14527300. * Sagan, Carl (1997). The demon-haunted world: science as a candle in the dark. [New York: Ballantine Books. ISBN 0345409469. * Scott, E.C. (1997). "Antievolution And Creationism In The United States". Annual Review of Anthropology 26 (1): 263289. doi:10.1146/annurev.anthro.26.1.263. * Maynard Smith, "The status of neo-darwinism," in "Towards a Theoretical Biology" (C.H. Waddington, ed., University Press, Edinburgh, 1969. * Hull, D.L. (1999). "The Use and Abuse of Sir Karl Popper". Biology and Philosophy 14 (4): 481504. doi:10.1023/A:1006554919188. http://www.springerlink.com/index/T4220X33T0284444.pdf. Retrieved on 2008-04-11. * Strobel, Lee (2004). The case for a Creator: a journalist investigates scientific evidence that points toward God. Grand Rapids, Mich: Zondervan/Willow Creek Resources. ISBN 0310240506. External links * Gallup public opinion poll in regards to the concepts of Evolution, Creationism, and Intelligent Design as of May 2007

* Data by country regarding the percentage of the population that believes in evolution Creationism as social policy * Isaac Asimov, The "Threat" of Creationism. Creationist beliefs * Answers in Genesis, Young Earth Creationism * Reasons to Believe, Old Earth Creationism Scientific rebuttals * Talk.origins Index to Creationist Claims Evolution versus creationism debates Location and linkYearPro-evolutionPro-creationismStanford University1994Will ProvinePhillip E. JohnsonNova on-line1996Kenneth R. MillerPhillip E. JohnsonFiring Line1997Kenneth R. Miller, Michael Ruse, Eugenie Scott & Barry LynnPhillip E. Johnson, Michael Behe, David Berlinski & William F. BuckleyUniversity of London (transcript)2007Lewis WolpertSteve FullerRetrieved from "http://en.wikipedia.org/wiki/Creation%E2%80%93evolution_controversy" Categories: Evolution | Creationism | Religion and science | Intelligent design controversies Hidden categories: Articles to be expanded since June 2008 | All articles to be expanded | All articles with unsourced statements | Articles with unsourced statements since September 2008 | Articles with unsourced statements since April 2008 | All pages needing cleanup | Wikipedia articles needing clarification from March 2008 Views * Article * Discussion * Edit this page * History Personal tools * Log in / create account Navigation * Main page * Contents * Featured content * Current events * Random article Search Principio del formulario Final del formulario Interaction * About Wikipedia * Community portal * Recent changes * Contact Wikipedia * Donate to Wikipedia * Help Toolbox * What links here * Related changes * Upload file * Special pages * Printable version

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Natural selection From Wikipedia, the free encyclopedia Jump to: navigation, search For other uses, see Natural selection (disambiguation). Natural selection is the process by which favorable heritable traits become more common in successive generations of a population of reproducing organisms, and unfavorable heritable traits become less common, due to differential reproduction of genotypes. Natural selection acts on the phenotype, or the observable characteristics of an organism, such that individuals with favorable phenotypes are more likely to survive and reproduce than those with less favorable phenotypes. The phenotype's genetic basis, the genotype associated with the favorable phenotype, will increase in frequency over the following generations. Over time, this process may result in adaptations that specialize organisms for particular ecological niches and may eventually result in the emergence of new species. In other words, natural selection is a mechanism by which evolution may take place within a population of organisms. Natural selection is one of the cornerstones of modern biology. The term was introduced by Charles Darwin in his groundbreaking 1859 book On the Origin of Species[1] in which natural selection was described by analogy to artificial selection, a process by which animals with traits considered desirable by human breeders are systematically favored for reproduction. The concept of natural selection was originally developed in the absence of a valid theory of inheritance; at the time of Darwin's writing, nothing was known of modern genetics. Although Gregor Mendel, the father of modern genetics, was a contemporary of Darwin's, his work would lie in obscurity until the early 20th century. The union of traditional Darwinian evolution with subsequent discoveries in classical and molecular genetics is termed the modern evolutionary synthesis. Although other mechanisms of molecular evolution, such as the neutral theory advanced by Motoo Kimura, have been identified as important causes of genetic diversity, natural selection remains the single primary explanation for adaptive evolution. Contents * 1 General principles o 1.1 Nomenclature and usage o 1.2 Fitness o 1.3 Types of selection o 1.4 Sexual selection * 2 Antibiotic resistance o 2.1 Directionality of selection o 2.2 Selection and genetic variation * 2.2.1 Mutation selection balance * 2.2.2 Genetic linkage * 3 Evolution by means of natural selection o 3.1 Speciation * 4 Historical development o 4.1 Pre-Darwinian theories o 4.2 Darwin's theory o 4.3 Modern evolutionary synthesis * 5 Impact of the idea o 5.1 Social and psychological theory o 5.2 Information and systems theory * 6 See also * 7 References * 8 Further reading

* 9 External links General principles

Darwin's illustrations of beak variation in the finches of the Galpagos Islands, which hold 13 closely related species that differ most markedly in the shape of their beaks. The beak of each species is suited to its preferred food, suggesting that beak shapes evolved by natural selection. See also character displacement, adaptive radiation, divergent evolution. See also: Genotype-phenotype distinction. Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is determined by an organism's genetic make-up (genotype) and the environment in which the organism lives. Often, natural selection acts on specific traits of an individual, and the terms phenotype and genotype are used narrowly to indicate these specific traits. When different organisms in a population possess different versions of a gene for a certain trait, each of these versions is known as an allele. It is this genetic variation that underlies phenotypic traits. A typical example is that certain combinations of genes for eye color in humans which, for instance, give rise to the phenotype of blue eyes. (On the other hand, when all the organisms in a population share the same allele for a particular trait, and this state is stable over time, the allele is said to be fixed in that population.) Some traits are governed by only a single gene, but most traits are influenced by the interactions of many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on the phenotype; together, these genes can produce a continuum of possible phenotypic values.[2] Nomenclature and usage The term "natural selection" has different definitions in different contexts. In simple terms, "natural selection" is most often defined to operate on heritable traits, but can sometimes refer to the differential reproductive success of phenotypes regardless of whether those phenotypes are heritable. Natural selection is "blind" in the sense that individuals' level of reproductive success is a function of the phenotype and not of whether or to what extent that phenotype is heritable. Following Darwin's primary usage[1] the term is often used to refer to both the consequence of blind selection and to its mechanisms.[3][4] It is sometimes helpful to explicitly distinguish between selection's mechanisms and its effects; when this distinction is important, scientists define "natural selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce," without regard to whether the basis of the selection is heritable. This is sometimes referred to as 'phenotypic natural selection.'[5] Traits that cause greater reproductive success of an organism are said to be selected for whereas those that reduce success are selected against. Selection for a trait may also result in the selection of other correlated traits that do not themselves directly influence fitness. This may occur as a result of pleiotropy or gene linkage.[6] Fitness Main article: Fitness (biology) The concept of fitness is central to natural selection. However, as with Natural selection above, there is serious divergence of opinion over the precise meaning of the term, and Richard Dawkins manages in his later books to avoid it entirely. (He devotes a chapter of his The Extended

Phenotype to discussing the various senses in which the term is used.) Although fitness is sometimes colloquially understood as a quality that promotes survival of a particular individual - as illustrated in the well-known phrase survival of the fittest - modern evolutionary theory defines fitness in terms of individual reproduction. The basis of this approach is: if an organism lives half as long as others of its species, but has twice as many offspring surviving to productive adulthood, its genes will become more common in the adult population of the next generation. This is known as differential reproduction. Though natural selection acts on individuals, its average effect on all individuals with a particular genotype corresponds to the fitness of that genotype. Very low-fitness genotypes cause their bearers to have few or no offspring on average; examples include many human genetic disorders like cystic fibrosis. Conditions like sickle-cell anemia may have low fitness in the general human population, but because it confers immunity from malaria, it has high fitness value in populations which have high malaria infection rates. Broadly speaking, an organism's fitness is a function of its alleles' fitnesses. Since fitness is an averaged quantity, however, it is possible a favorable mutation may arise in an individual that does not survive to adulthood for unrelated reasons. Types of selection Natural selection can act on any phenotypic trait, and selective pressure can be produced by any aspect of the environment, including mates and conspecifics, or members of the same species. However, this does not imply that natural selection is always directional and results in adaptive evolution; natural selection often results in the maintenance of the status quo by eliminating less fit variants. The unit of selection can be the individual or it can be another level within the hierarchy of biological organisation, such as genes, cells, and kin groups. There is still debate about whether natural selection acts at the level of groups or species to produce adaptations that benefit a larger, non-kin group. Selection at a different level such as the gene can result in an increase in fitness for that gene, while at the same time reducing the fitness of the individuals carrying that gene, in a process called intragenomic conflict. Overall, the combined effect of all selection pressures at various levels determines the overall fitness of an individual, and hence the outcome of natural selection.

The life cycle of a sexually reproducing organism. Various components of natural selection are indicated for each life stage.[7] Natural selection occurs at every life stage of an individual. An individual organism must survive until adulthood before it can reproduce, and selection of those that reach this stage is called viability selection. In many species, adults must compete with each other for mates via sexual selection, and success in this competition determines who will parent the next generation. When individuals can reproduce more than once, a longer survival in the reproductive phase increases the number of offspring, called survival selection. The fecundity of both females and males (for example, giant sperm in certain species of Drosophila[8]) can be limited via fecundity selection. The viability of produced gametes can differ, while intragenomic conflicts such as meiotic drive between the haploid gametes can result in gametic or genic selection. Finally, the union of some combinations of eggs and sperm might be more compatible than others; this is termed compatibility selection. Sexual selection

Main article: Sexual selection It is also useful to make a mechanistic distinction between ecological selection and the narrower term sexual selection. Ecological selection covers any mechanism of selection as a result of the environment (including relatives, e.g. kin selection, and conspecifics, e.g. competition, infanticide), while sexual selection refers specifically to competition between conspecifics for mates.[9] Sexual selection can be intrasexual, as in cases of competition among individuals of the same sex in a population, or intersexual, as in cases where one sex controls reproductive access by choosing among a population of available mates. Most commonly, intrasexual selection involves male-male competition and intersexual selection involves female choice of suitable males, due to the generally greater investment of resources for a female than a male in a single offspring organism. However, some species exhibit sex-role reversed behavior in which it is males that are most selective in mate choice; the best-known examples of this pattern occur in some fishes of the family Syngnathidae, though likely examples have also been found in amphibian and bird species.[10] Some features that are confined to one sex only of a particular species can be explained by selection exercised by the other sex in the choice of a mate, for example, the extravagant plumage of some male birds. Similarly, aggression between members of the same sex is sometimes associated with very distinctive features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual selection is often associated with sexual dimorphism, including differences in body size between males and females of a species.[11] Antibiotic resistance

Schematic representation of how antibiotic resistance is enhanced by natural selection. The top section represents a population of bacteria before exposure to an antibiotic. The middle section shows the population directly after exposure, the phase in which selection took place. The last section shows the distribution of resistance in a new generation of bacteria. The legend indicates the resistance levels of individuals. A well-known example of natural selection in action is the development of antibiotic resistance in microorganisms. Since the discovery of penicillin in 1928 by Alexander Fleming, antibiotics have been used to fight bacterial diseases. Natural populations of bacteria contain, among their vast numbers of individual members, considerable variation in their genetic material, primarily as the result of mutations. When exposed to antibiotics, most bacteria die quickly, but some may have mutations that make them slightly less susceptible. If the exposure to antibiotics is short, these individuals will survive the treatment. This selective elimination of maladapted individuals from a population is natural selection. These surviving bacteria will then reproduce again, producing the next generation. Due to the elimination of the maladapted individuals in the past generation, this population contains more bacteria that have some resistance against the antibiotic. At the same time, new mutations occur, contributing new genetic variation to the existing genetic variation. Spontaneous mutations are very rare, and advantageous mutations are even rarer. However, populations of bacteria are large enough that a few individuals will have beneficial mutations. If a new mutation reduces their susceptibility to an antibiotic, these individuals are more likely to survive when next confronted with that antibiotic. Given enough time,

and repeated exposure to the antibiotic, a population of antibioticresistant bacteria will emerge. The widespread use and misuse of antibiotics has resulted in increased microbial resistance to antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been described as a 'superbug' because of the threat it poses to health and its relative invulnerability to existing drugs.[12] Response strategies typically include the use of different, stronger antibiotics; however, new strains of MRSA have recently emerged that are resistant even to these drugs.[13] This is an example of what is known as an evolutionary arms race, in which bacteria continue to develop strains that are less susceptible to antibiotics, while medical researchers continue to develop new antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms races are not necessarily induced by man; a well-documented example involves the elaboration of the RNA interference pathway in plants as means of innate immunity against viruses.[14] Directionality of selection When some component of a trait is heritable, selection will alter the frequencies of the different alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three classes, on the basis of its effect on allele frequencies.[15] Directional selection occurs when a certain allele has a greater fitness than others, resulting in an increase in frequency of that allele. This process can continue until the allele is fixed and the entire population shares the fitter phenotype. It is directional selection that is illustrated in the antibiotic resistance example above. Far more common is stabilizing selection (also known as purifying selection), which lowers the frequency of alleles that have a deleterious effect on the phenotype - that is, produce organisms of lower fitness. This process can continue until the allele is eliminated from the population. Purifying selection results in functional genetic features, such as protein-coding genes or regulatory sequences, being conserved over time due to selective pressure against deleterious variants. Finally, a number of forms of balancing selection exist, which do not result in fixation, but maintain an allele at intermediate frequencies in a population. This can occur in diploid species (that is, those that have two pairs of chromosomes) when heterozygote individuals, who have different alleles on each chromosome at a single genetic locus, have a higher fitness than homozygote individuals that have two of the same alleles. This is called heterozygote advantage or overdominance, of which the best-known example is the malarial resistance observed in heterozygous humans who carry only one copy of the gene for sickle cell anemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which favors genotypes that depart from the average in either direction (that is, the opposite of overdominance), and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of other phenotypes in the population. The principles of game theory have been applied to understand the fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal altruism.[16][17] Selection and genetic variation A portion of all genetic variation is functionally neutral in that it produces no phenotypic effect or significant difference in fitness; the

hypothesis that this variation accounts for a large fraction of observed genetic diversity is known as the neutral theory of molecular evolution and was originated by Motoo Kimura. Neutral variation was once thought to encompass most of the genetic variation in non-coding DNA, which was hypothesized to be composed of "junk DNA". However, more recently, the functional roles of non-coding DNA, such as the regulatory and developmental functions of RNA gene products, has been studied in depth;[18] large parts of non-protein-coding DNA sequences are highly conserved under strong purifying selection and thus do not vary much from individual to individual, indicating that mutations in these regions have deleterious consequences.[19][20] When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency of such variation. As a result, the genetic variation at those sites will be higher than at sites where variation does influence fitness.[15] Mutation selection balance Natural selection results in the reduction of genetic variation through the elimination of maladapted individuals and consequently of the mutations that caused the maladaptation. At the same time, new mutations occur, resulting in a mutation-selection balance. The exact outcome of the two processes depends both on the rate at which new mutations occur and on the strength of the natural selection, which is a function of how unfavorable the mutation proves to be. Consequently, changes in the mutation rate or the selection pressure will result in a different mutation-selection balance. Genetic linkage Genetic linkage occurs when the loci of two alleles are linked, or in close proximity to each other on the chromosome. During the formation of gametes, recombination of the genetic material results in reshuffling of the alleles. However, the chance that such a reshuffle occurs between two alleles depends on the distance between those alleles; the closer the alleles are to each other, the less likely it is that such a reshuffle will occur. Consequently, when selection targets one allele, this automatically results in selection of the other allele as well; through this mechanism, selection can have a strong influence on patterns of variation in the genome. Selective sweeps occur when an Allele becomes more common in a population as a result of positive selection. As the prevalence of one allele increases, linked Alleles can also become more common, whether they are neutral or even slightly deleterious. This is called genetic hitchhiking. A strong selective sweep results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are essentially the only ones that exist in the population. Whether a selective sweep has occurred or not can be investigated by measuring linkage disequilibrium, or whether a given haplotype is overrepresented in the population. Normally, genetic recombination results in a reshuffling of the different alleles within a haplotype, and none of the haplotypes will dominate the population. However, during a selective sweep, selection for a specific allele will also result in selection of neighbouring alleles. Therefore, the presence of strong linkage disequilibrium might indicate that there has been a 'recent' selective sweep, and this can be used to identify sites recently under selection. Background selection is the opposite of a selective sweep. If a specific site experiences strong and persistent purifying selection, linked variation will tend to be weeded out along with it, producing a region in the genome of low overall variability. Because background selection is a

result of deleterious new mutations, which can occur randomly in any haplotype, it produces no linkage disequilibrium. Evolution by means of natural selection Main articles: Evolution and Darwinism A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation, is the presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of mutations, recombinations and alterations in the karyotype (the number, shape, size and internal arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or highly disadvantageous, but large effects are very rare. In the past, most changes in the genetic material were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a synonymous substitution. However, recent research suggests that many mutations in non-coding DNA do have slight deleterious effects.[19][20] Although both mutation rates and average fitness effects of mutations are dependent on the organism, estimates from data in humans have found that a majority of mutations are slightly deleterious.[21]

The exuberant tail of the peacock is thought to be the result of sexual selection by females. This peacock is an albino it carries a mutation that makes it unable to produce melanin. Selection against albinos in nature is intense because they are easily spotted by predators or are unsuccessful in competition for mates, and so these mutations are usually rapidly eliminated by natural selection. By the definition of fitness, individuals with greater fitness are more likely to contribute offspring to the next generation, while individuals with lesser fitness are more likely to die early or fail to reproduce. As a result, alleles which on average result in greater fitness become more abundant in the next generation, while alleles which generally reduce fitness become rarer. If the selection forces remain the same for many generations, beneficial alleles become more and more abundant, until they dominate the population, while alleles with a lesser fitness disappear. In every generation, new mutations and recombinations arise spontaneously, producing a new spectrum of phenotypes. Therefore, each new generation will be enriched by the increasing abundance of alleles that contribute to those traits that were favored by selection, enhancing these traits over successive generations.

X-ray of the left hand of a ten year old boy with polydactyly. Some mutations occur in so-called regulatory genes. Changes in these can have large effects on the phenotype of the individual because they regulate the function of many other genes. Most, but not all, mutations in regulatory genes result in non-viable zygotes. Examples of nonlethal regulatory mutations occur in HOX genes in humans, which can result in a cervical rib[22] or polydactyly, an increase in the number of fingers or toes.[23] When such mutations result in a higher fitness, natural selection will favor these phenotypes and the novel trait will spread in the population. Established traits are not immutable; traits that have high fitness in one environmental context may be much less fit if environmental conditions change. In the absence of natural selection to preserve such a trait, it will become more variable and deteriorate over time, possibly resulting in a vestigial manifestation of the trait. In many circumstances, the apparently vestigial structure may retain a limited

functionality, or may be co-opted for other advantageous traits in a phenomenon known as preadaptation. A famous example of a vestigial structure, the eye of the blind mole rat, is believed to retain function in photoperiod perception.[24] Speciation Speciation requires selective mating, which result in a reduced gene flow. Selective mating can be the result of, for example, a change in the physical environment (physical isolation by an extrinsic barrier), or by sexual selection resulting in assortative mating. Over time, these subgroups might diverge radically to become different species, either because of differences in selection pressures on the different subgroups, or because different mutations arise spontaneously in the different populations, or because of founder effects - some potentially beneficial alleles may, by chance, be present in only one or other of two subgroups when they first become separated. A lesser-known mechanism of speciation occurs via hybridization, well-documented in plants and occasionally observed in species-rich groups of animals such as cichlid fishes.[25] Such mechanisms of rapid speciation can reflect a mechanism of evolutionary change known as punctuated equilibrium, which suggests that evolutionary change and particularly speciation typically happens quickly after interrupting long periods of stasis. Genetic changes within groups result in increasing incompatibility between the genomes of the two subgroups, thus reducing gene flow between the groups. Gene flow will effectively cease when the distinctive mutations characterizing each subgroup become fixed. As few as two mutations can result in speciation: if each mutation has a neutral or positive effect on fitness when they occur separately, but a negative effect when they occur together, then fixation of these genes in the respective subgroups will lead to two reproductively isolated populations. According to the biological species concept, these will be two different species. Historical development Main articles: History of evolutionary thought, Inception of Darwin's theory, and Development of Darwin's theory

The modern theory of natural selection derives from the work of Charles Darwin in the nineteenth century. Pre-Darwinian theories Several ancient philosophers expressed the idea that nature produces a huge variety of creatures, apparently randomly, and that only those creatures survive that manage to provide for themselves and reproduce successfully; well-known examples include Empedocles[26] and his intellectual successor, Lucretius,[27] while related ideas were later refined by Aristotle.[28] The struggle for existence was later described by al-Jahiz in the 9th century.[29][30] Such classical arguments were reintroduced in the 18th century by Pierre Louis Maupertuis[31] and others, including Charles Darwin's grandfather Erasmus Darwin. While these forerunners had an influence on Darwinism, they later had little influence on the trajectory of evolutionary thought after Charles Darwin. Until the early 19th century, the prevailing view in Western societies was that differences between individuals of a species were uninteresting departures from their Platonic ideal (or typus) of created kinds. However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act continuously over long periods of time

to produce radical changes in the Earth's landscape. The success of this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny, virtually imperceptible changes in successive generations could produce consequences on the scale of differences between species. Early 19th century evolutionists such as Jean Baptiste Lamarck suggested the inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by an organism during its lifetime could be inherited by that organism's progeny, eventually causing transmutation of species.[32] This theory has come to be known as Lamarckism and was an influence on the anti-genetic ideas of the Stalinist Soviet biologist Trofim Lysenko.[33] Darwin's theory In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a trait], if useful, is preserved".[34] The concept was simple but powerful: individuals best adapted to their environments are more likely to survive and reproduce. As long as there is some variation between them, there will be an inevitable selection of individuals with the most advantageous variations. If the variations are inherited, then differential reproductive success will lead to a progressive evolution of particular populations of a species, and populations that evolve to be sufficiently different eventually become different species.[35] Darwin's ideas were inspired by the observations that he had made on the Beagle voyage, and by the work of two political economists. The first was the Reverend Thomas Malthus, who in An Essay on the Principle of Population, noted that population (if unchecked) increases exponentially whereas the food supply grows only arithmetically; thus inevitable limitations of resources would have demographic implications, leading to a "struggle for existence" as a divinely ordained law "in order to rouse man into action, and form his mind to reason" for the greater good despite the "partial evil" limiting population.[36] The second was Adam Smith who, in The Wealth of Nations, identified a regulating mechanism in free markets, which he referred to as the "invisible hand", which suggests that prices self-adjust according to supplies and demand.[37] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the "struggle for existence" in nature and it struck him that as population outgrew resources, "favourable variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be the formation of new species."[38] Once he had his theory "by which to work", Darwin was meticulous about gathering and refining evidence as his "prime hobby" before making his idea public. He was in the process of writing his "big book" to present his researches when the naturalist Alfred Russel Wallace independently conceived of the principle and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker decided (without Wallace's knowledge) to present his essay together with unpublished writings which Darwin had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-discovery of the principle in July 1858.[39] Darwin published a detailed account of his evidence and conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others notably William Charles Wells in 1813, and Patrick Matthew in 1831 had proposed similar ideas, but had neither developed them nor presented them in notable scientific publications.[40]

Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding, which he called artificial selection; in his early manuscripts he referred to a 'Nature' which would do the selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has been the main, but not exclusive means of modification."[41] In a letter to Charles Lyell in September 1860, Darwin regretted the use of the term 'Natural Selection', preferring the term 'Natural Preservation'.[42] For Darwin and his contemporaries, natural selection was essentially synonymous with evolution by natural selection. After the publication of The Origin of Species, educated people generally accepted that evolution had occurred in some form. However, natural selection remained controversial as a mechanism, partly because it was perceived to be too weak to explain the range of observed characteristics of living organisms, and partly because even supporters of evolution balked at its 'unguided' and non-progressive nature,[43] a response that has been characterized as the single most significant impediment to the idea's acceptance.[44] However, some thinkers enthusiastically embraced natural selection; after reading Darwin, Herbert Spencer introduced the term survival of the fittest, which became a popular summary of the theory.[45] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer, of the Survival of the Fittest, is more accurate, and is sometimes equally convenient."[46] Although the phrase is still often used by nonbiologists, modern biologists avoid it because it is tautological if fittest is read to mean functionally superior and is applied to individuals rather than considered as an averaged quantity over populations.[47] Modern evolutionary synthesis Main article: Modern evolutionary synthesis Only after the integration of a theory of evolution with a complex statistical appreciation of Austrian monk Gregor Mendel's 're-discovered' laws of inheritance did natural selection become generally accepted by scientists. The work of Ronald Fisher (who developed the language of mathematics and natural selection in terms of the underlying genetic processes),[3] J.B.S. Haldane (who introduced the concept of the 'cost' of natural selection),[48] Sewall Wright (who elucidated the nature of selection and adaptation),[49] Theodosius Dobzhansky (who established the idea that mutation, by creating genetic diversity, supplied the raw material for natural selection),[50] William Hamilton (who conceived of kin selection), Ernst Mayr (who recognised the key importance of reproductive isolation for speciation)[51] and many others formed the modern evolutionary synthesis. This synthesis cemented natural selection as the foundation of evolutionary theory, where it remains today. Impact of the idea Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th century thought. Perhaps the most radical claim of the theory of evolution through natural selection is that "elaborately constructed forms, so different from each other, and dependent on each other in so complex a manner" evolved from the simplest forms of life by a few simple principles. This claim inspired some of Darwin's most ardent supportersand provoked the most profound opposition. The radicalism of natural selection, according to Stephen Jay Gould,[52] lay in its power to "dethrone some of the deepest and most

traditional comforts of Western thought". In particular, it challenged long-standing beliefs in such concepts as a special and exalted place for humans in the natural world and a benevolent creator whose intentions were reflected in nature's order and design. Social and psychological theory The social implications of the theory of evolution by natural selection also became the source of continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he showed that free competition, the struggle for existence, which the economists celebrate as the highest historical achievement, is the normal state of the animal kingdom".[53] Interpretation of natural selection as necessarily 'progressive', leading to increasing 'advances' in intelligence and civilisation, was used as a justification for colonialism and policies of eugenics, as well as broader sociopolitical positions now described as Social Darwinism. Konrad Lorenz won the Nobel Prize in Physiology or Medicine in 1973 for his analysis of animal behavior in terms of the role of natural selection (particularly group selection). However, in Germany in 1940, in writings that he subsequently disowned, he used the theory as a justification for policies of the Nazi state. He wrote "... selection for toughness, heroism, and social utility...must be accomplished by some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-induced degeneracy. The racial idea as the basis of our state has already accomplished much in this respect."[54] Others have developed ideas that human societies and culture evolve by mechanisms that are analogous to those that apply to evolution of species.[55] More recently, work among anthropologists and psychologists has led to the development of sociobiology and later evolutionary psychology, a field that attempts to explain features of human psychology in terms of adaptation to the ancestral environment. The most prominent such example, notably advanced in the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain is adapted to acquire the grammatical rules of natural language.[56] Other aspects of human behavior and social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender roles, have been hypothesized to have similar origins as adaptations to the early environment in which modern humans evolved. By analogy to the action of natural selection on genes, the concept of memes - "units of cultural transmission", or culture's equivalents of genes undergoing selection and recombination - has arisen, first described in this form by Richard Dawkins[57] and subsequently expanded upon by philosophers such as Daniel Dennett as explanations for complex cultural activities, including human consciousness.[58] Extensions of the theory of natural selection to such a wide range of cultural phenomena have been distinctly controversial and are not widely accepted.[59] Information and systems theory In 1922, Alfred Lotka proposed that natural selection might be understood as a physical principle which could be energetically quantified,[60] a concept that was later developed by Howard Odum as the maximum power principle whereby evolutionary systems with selective advantage maximise the rate of useful energy transformation. Such concepts are sometimes relevant in the study of applied thermodynamics. The principles of natural selection have inspired a variety of computational techniques, such as "soft" artificial life, that simulate

selective processes and can be highly efficient in 'adapting' entities to an environment defined by a specified fitness function.[61] For example, a class of heuristic optimization algorithms known as genetic algorithms, pioneered by John Holland in the 1970s and expanded upon by David E. Goldberg,[62] identify optimal solutions by simulated reproduction and mutation of a population of solutions defined by an initial probability distribution.[63] Such algorithms are particularly useful when applied to problems whose solution landscape is very rough or has many local minima. See also Co-evolution Gene-centered view of evolution Negative selection Unit of selection Artificial selection

* * * * *

References 1. ^ a b Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life John Murray, London; modern reprint Charles Darwin, Julian Huxley (2003). The Origin of Species. Signet Classics. ISBN 0-451-52906-5. Published online at The complete work of Charles Darwin online: On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. 2. ^ Falconer DS & Mackay TFC (1996) Introduction to Quantitative Genetics Addison Wesley Longman, Harlow, Essex, UK ISBN 0-582-24302-5 3. ^ a b Fisher RA (1930) The Genetical Theory of Natural Selection Clarendon Press, Oxford 4. ^ Works employing or describing this usage: Endler JA (1986). Natural Selection in the Wild. Princeton, New Jersey: Princeton University Press. ISBN 0-691-00057-3. Williams GC (1966). Adaptation and Natural Selection. Oxford University Press. 5. ^ Works employing or describing this usage: Lande R & Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210-26 Futuyma DJ (2005) Evolution. Sinauer Associates, Inc., Sunderland, Massachusetts. ISBN 0-87893-187-2 Haldane, J.B.S. 1953. The measurement of natural selection. Proceedings of the 9th International Congress of Genetics. 1: 480-487 6. ^ Sober E (1984; 1993) The Nature of Selection: Evolutionary Theory in Philosophical Focus University of Chicago Press ISBN 0-226-76748-5 7. ^ Modified from Christiansen FB (1984) The definition and measurement of fitness. In: Evolutionary ecology (ed. Shorrocks B) pp65-79. Blackwell Scientific, Oxford by adding survival selection in the reproductive phase 8. ^ Pitnick S & Markow TA (1994) Large-male advantage associated with the costs of sperm production in Drosophila hydei, a species with giant sperm. Proc Natl Acad Sci USA 91:9277-81; Pitnick S (1996) Investment in testes and the cost of making long sperm in Drosophila. Am Nat 148:57-80 9. ^ Andersson, M (1995). Sexual Selection. Princeton, New Jersey: Princeton University Press. ISBN 0-691-00057-3. 10. ^ Eens M, Pinxten R. (2000). Sex-role reversal in vertebrates: behavioural and endocrinological accounts. Behav Processes 51(1-3):135147. PMID 11074317 11. ^ Barlow GW. (2005). How Do We Decide that a Species is Sex-Role Reversed? The Quarterly Review of Biology 80(1):2835. PMID 15884733 12. ^ "MRSA Superbug News". http://www.inboxrobot.com/news/MRSASuperbug. Retrieved on 2006-05-06.

13. ^ Schito GC (2006). "The importance of the development of antibiotic resistance in Staphylococcus aureus". Clin Microbiol Infect 12 Suppl 1: 38. doi:10.1111/j.1469-0691.2006.01343.x. PMID 16445718. [1] 14. ^ Lucy A, Guo H, Li W, Ding S (2000). "Suppression of posttranscriptional gene silencing by a plant viral protein localized in the nucleus". EMBO J 19 (7): 167280. PMID 10747034. 15. ^ a b Rice SH. (2004). Evolutionary Theory: Mathematical and Conceptual Foundations. Sinauer Associates: Sunderland, Massachusetts, USA. ISBN 0-87893-702-1 See esp. ch. 5 and 6 for a quantitative treatment. 16. ^ Hamilton WD. (1964). The genetical evolution of social behaviour I and II. Journal of Theoretical Biology 7: 1-16 and 17-52. PMID 5875341 PMID 5875340 17. ^ Trivers RL. (1971). The evolution of reciprocal altruism. Q Rev Biol 46: 35-57. 18. ^ He L, Hannon GJ. (2004). MicroRNAs: small RNAs with a big role in gene regulation. Nat Rev Genet 5(7):522-31. PMID 15211354 19. ^ a b Kryukov GV, Schmidt S & Sunyaev S (2005) Small fitness effect of mutations in highly conserved non-coding regions. Human Molecular Genetics 14:2221-9 20. ^ a b Bejerano G, Pheasant M, Makunin I, Stephen S, Kent WJ, Mattick JS & Haussler D (2004) Ultraconserved elements in the human genome. Science 304:1321-5 21. ^ Eyre-Walker A, Woolfit M, Phelps T. (2006). The distribution of fitness effects of new deleterious amino acid mutations in humans. Genetics 173(2):891-900. PMID 16547091 22. ^ Galis F (1999) Why do almost all mammals have seven cervical vertebrae? developmental constraints, Hox genes, and cancer. J Exp Zool 285:19-26 23. ^ Zakany J, FromentalRamain C, Warot X & Duboule D (1997) Regulation of number and size of digits by posterior Hox genes: a dose-dependent mechanism with potential evolutionary implications. Proc Natl Acad Sci USA 94:13695-700 24. ^ Sanyal S, Jansen HG, de Grip WJ, Nevo E, de Jong WW. (1990). The eye of the blind mole rat, Spalax ehrenbergi. Rudiment with hidden function? Invest Ophthalmol Vis Sci. 1990 31(7):1398-404. PMID 2142147 25. ^ Salzburger W, Baric S, Sturmbauer C. (2002). Speciation via introgressive hybridization in East African cichlids? Mol Ecol 11(3): 619625. PMID 11918795 26. ^ Empedocles, On Nature, Book II 27. ^ Lucretius, De rerum natura, Book V 28. ^ Aristotle, Physics, Book II, Chapters 4 and 8 29. ^ Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of the American Philosophical Society 84 (1), p. 71-123. 30. ^ Mehmet Bayrakdar (Third Quarter, 1983). "Al-Jahiz And the Rise of Biological Evolutionism", The Islamic Quarterly. London. 31. ^ Maupertuis, Pierre Louis (1748). "Derivation of the laws of motion and equilibrium from a metaphysical principle (Original French text)". Histoire de l'academie des sciences et belle lettres de Berlin 1746: 267 294. 32. ^ Chevalier de Lamarck J-B, de Monet PA (1809) Philosophie Zoologique 33. ^ Joravsky D. (1959). Soviet Marxism and Biology before Lysenko. Journal of the History of Ideas 20(1):85-104. 34. ^ Darwin 1859, p. 61 35. ^ Darwin 1859, p. 5 36. ^ T. Robert Malthus (1798). "An Essay on the Principle of Population". June 29, 1999.

http://www.faculty.rsu.edu/~felwell/Theorists/Malthus/Essay.htm#112. Retrieved on 2008-11-03. 37. ^ Orrell, David (2007) Apollo's Arrow Toronto: HarperCollins Publishers Ltd. [2] 38. ^ Charles Darwin; ed. Nora Barlow (1958). "The autobiography of Charles Darwin 1809-1882". London: Collins. 120. http://darwinonline.org.uk/content/frameset?viewtype=text&itemID=F1497&pageseq=124. Retrieved on 2008-11-03. 39. ^ Wallace, Alfred Russel (1870) Contributions to the Theory of Natural Selection New York: Macmillan & Co. [3] 40. ^ Darwin 1861, p. xiii 41. ^ Darwin 1859, p. 6 42. ^ "Darwin Correspondence Online Database: Darwin, C. R. to Lyell, Charles, 28 September 1860". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2931.html. Retrieved on 2006-05-10. 43. ^ Eisley L. (1958). Darwin's Century: Evolution and the Men Who Discovered It. Doubleday & Co: New York, USA. 44. ^ Kuhn TS. [1962] (1996). The Structure of Scientific Revolution 3rd ed. University of Chicago Press: Chicago, Illinois, USA. ISBN 0-22645808-3 45. ^ "Pioneers of Psychology [2001 Tour - School of Education & Psychology"]. http://educ.southern.edu/tour/who/pioneers/spencer.html. Retrieved on 2007-08-29. Maurice E. Stucke. "Better Competition Advocacy". http://works.bepress.com/cgi/viewcontent.cgi?article=1000&context=maurice _stucke. Retrieved on 2007-08-29. "Herbert Spencer in his Principles of Biology of 1864, vol. 1, p. 444, wrote This survival of the fittest, which I have here sought to express in mechanical terms, is that which Mr. Darwin has called natural selection, or the preservation of favoured races in the struggle for life." 46. ^ Darwin 1872, p. 49. 47. ^ Mills SK, Beatty JH. [1979] (1994). The Propensity Interpretation of Fitness. Originally in Philosophy of Science (1979) 46: 263-286; republished in Conceptual Issues in Evolutionary Biology 2nd ed. Elliott Sober, ed. MIT Press: Cambridge, Massachusetts, USA. pp3-23. ISBN 0-26269162-0. 48. ^ Haldane JBS (1932) The Causes of Evolution; Haldane JBS (1957) The cost of natural selection. J Genet 55:511-24([4]. 49. ^ Wright S (1932) The roles of mutation, inbreeding, crossbreeding and selection in evolution Proc 6th Int Cong Genet 1:35666 50. ^ Dobzhansky Th (1937) Genetics and the Origin of Species Columbia University Press, New York. (2nd ed., 1941; 3rd edn., 1951) 51. ^ Mayr E (1942) Systematics and the Origin of Species Columbia University Press, New York. ISBN 0-674-86250-3 52. ^ The New York Review of Books: Darwinian Fundamentalism (accessed May 6, 2006) 53. ^ Engels F (1873-86) Dialectics of Nature 3d ed. Moscow: Progress, 1964 [5] 54. ^ Quoted in translation in Eisenberg L (2005) Which image for Lorenz? Am J Psychiatry 162:1760 [6] 55. ^ e.g. Wilson, DS (2002) Darwin's Cathedral: Evolution, Religion, and the Nature of Society. University of Chicago Press, ISBN 0-226-90134-3 56. ^ Pinker S. [1994] (1995). The Language Instinct: How the Mind Creates Language. HarperCollins: New York, NY, USA. ISBN 0-06-097651-9 57. ^ Dawkins R. [1976] (1989). The Selfish Gene. Oxford University Press: New York, NY, USA, p.192. ISBN 0-19-286092-5

58. ^ Dennett DC. (1991). Consciousness Explained. Little, Brown, and Co: New York, NY, USA. ISBN 0-316-18066-1 59. ^ For example, see Rose H, Rose SPR, Jencks C. (2000). Alas, Poor Darwin: Arguments Against Evolutionary Psychology. Harmony Books. ISBN 0609605135 60. ^ Lotka AJ (1922a) Contribution to the energetics of evolution [PDF] Proc Natl Acad Sci USA 8:14751 Lotka AJ (1922b) Natural selection as a physical principle [PDF] Proc Natl Acad Sci USA 8:1514 61. ^ Kauffman SA (1993) The Origin of order. Self-organization and selection in evolution. New York: Oxford University Press ISBN 0-19507951-5 62. ^ Goldberg DE. (1989). Genetic Algorithms in Search, Optimization and Machine Learning. Addison-Wesley: Boston, MA, USA 63. ^ Mitchell, Melanie, (1996), An Introduction to Genetic Algorithms, MIT Press, Cambridge, MA. Further reading * For technical audiences o Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University Press. ISBN 0-674-00613-5. o Maynard Smith, John (1993). The Theory of Evolution: Canto Edition. Cambridge University Press. ISBN 0-521-45128-0. o Popper, Karl (1978) Natural selection and the emergence of mind. Dialectica 32:339-55. See [7] o Sober, Elliott (1984) The Nature of Selection: Evolutionary Theory in Philosophical Focus. University of Chicago Press. o Williams, George C. (1966) Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thought. Oxford University Press. o Williams George C. (1992) Natural Selection: Domains, Levels and Challenges. Oxford University Press. * For general audiences o Dawkins, Richard (1996) Climbing Mount Improbable. Penguin Books, ISBN 0-670-85018-7. o Dennett, Daniel (1995) Darwin's Dangerous Idea: Evolution and the Meanings of Life. Simon & Schuster ISBN 0-684-82471-X. o Gould, Stephen Jay (1997) Ever Since Darwin: Reflections in Natural History. Norton, ISBN 0-393-06425-5. o Jones, Steve (2001) Darwin's Ghost: The Origin of Species Updated. Ballantine Books ISBN 0-345-42277-5. Also published in Britain under the title Almost like a whale: the origin of species updated. Doubleday. ISBN 1-86230-025-9. o Lewontin, Richard (1978) Adaptation. Scientific American 239:212-30 o Weiner, Jonathan (1994) The Beak of the Finch: A Story of Evolution in Our Time. Vintage Books, ISBN 0-679-73337-X. * Historical o Zirkle C (1941). Natural Selection before the "Origin of Species", Proceedings of the American Philosophical Society 84 (1), p. 71-123. o Kohm M (2004) A Reason for Everything: Natural Selection and the English Imagination. London: Faber and Faber. ISBN 0-571-22392-3. For review, see [8] van Wyhe J (2005) Human Nature Review 5:1-4 External links * The Origin of Species by Charles Darwin - Chapter 4,Natural Selection * Natural Selection- Modeling for Understanding in Science Education, University of Wisconsin * Natural Selection from University of Berkeley education website v d e

Basic topics in evolutionary biologyEvidence of common descentProcesses of evolutionAdaptation Macroevolution Microevolution SpeciationPopulation genetic mechanismsNatural selection Genetic drift Gene flow MutationEvolutionary developmental biology (Evo-devo) conceptsPhenotypic plasticity Canalisation ModularityThe evolution ofDNA Sex Aging Intelligence The Ear The Eye Flight Plants Fungi Life Humans Dolphins and whales Birds Evolution of the horse Spider evolution Evolution of sirenians Insect evolution Evolution of Mollusca Evolution of dinosaursModes of speciationAnagenesis Catagenesis CladogenesisHistoryHistory of evolutionary thought Charles Darwin On the Origin of Species Modern evolutionary synthesis Gene-centered view of evolution Life (classification trees)Other subfieldsEcological genetics Molecular evolution Phylogenetics SystematicsList of evolutionary biology topics Timeline of evolution v d e Topics in population geneticsKey conceptsHardy-Weinberg law Genetic linkage Linkage disequilibrium Fisher's fundamental theorem Neutral theory Price equationSelectionNatural Sexual Artificial EcologicalEffects of selection on genomic variationGenetic hitchhiking Background selectionGenetic driftSmall population size Population bottleneck Founder effect Coalescence Balding-Nichols ModelFoundersR.A. Fisher J. B. S. Haldane Sewall WrightRelated topicsEvolution Microevolution Evolutionary game theory Fitness landscape Genetic genealogyList of evolutionary biology topicsRetrieved from "http://en.wikipedia.org/wiki/Natural_selection" Categories: Selection | Evolution | Evolutionary biology | Ecological processes

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Adaptation From Wikipedia, the free encyclopedia Population geneticsBiology Portal v d e Adaptation is one of the basic phenomena of biology.[1] It is the process, which takes place under natural selection, whereby an organism becomes better suited to its habitat.[2] Also, the term may refer to some characteristic which stands out as being especially significant in the organism's survival. For example, the adaptation of horses' teeth to the grinding of grass, or their capacity to run fast to escape predators. Contents * 1 General principles * 2 Brief history * 3 Types of adaptation o 3.1 Changes in habitat * 3.1.1 Habitat tracking * 3.1.2 Genetic change o 3.2 Intimate relationships: co-adaptations * 3.2.1 Mimicry o 3.3 The basic machinery: internal adaptations o 3.4 Compromise and conflict between adaptations * 4 Shifts in function o 4.1 Pre-adaptations o 4.2 Co-option of existing traits: exaptation * 5 Related issues o 5.1 Non-adaptive traits o 5.2 Fitness landscapes; drift o 5.3 Extinction * 5.3.1 Co-extinction o 5.4 Flexibility, acclimatization, learning * 5.4.1 Flexibility * 5.4.2 Acclimatization * 5.4.3 Learning o 5.5 Function and teleonomy * 5.5.1 Function * 5.5.2 Teleonomy * 6 References * 7 See also General principles The significance of an adaptation can only be understood in relation to the total biology of the species. Julian Huxley [3] Adaptation is, first of all, a process, rather than a physical part of a body. The distinction may be seen in an internal parasite (such as a fluke), where the bodily structure is greatly simplified, but the organism is highly adapted to its environment. In such cases critical adaptations take place in the life-cycle, which is often quite complex.[4] However, as a practical term, adaptation is often used for the product: those features of a species which result from the process. Here one does find structures which were formerly adaptive, but are so no longer. Vestigial organs, such as the human caecum (appendix), are examples. Adaptation may be seen as one aspect of a two-stage process. First, there is speciation (species-splitting or cladogenesis), caused by geographical isolation or some other mechanism.[5][6] Second, there follows adaptation, driven by natural selection. Something like this must have happened with Darwin's finches, and there are many other examples. The present favourite is the evolution of cichlid fish in African lakes, where the question of reproductive isolation is much more complex.[7][8]

Another great principle is that an organism must be viable at all stages of its development and at all stages of its evolution. This is obviously true, and it follows that there are constraints on the evolution of development, behaviour and structure of organisms. The main constraint, over which there has been much debate, is the requirement that changes in the system during evolution should be relatively small changes, because the body systems are so complex and interlinked. This is a sound principle, though there may be rare exceptions: polyploidy in plants is common[9], and the symbiosis of micro-organisms that formed the eukaryota is a more exotic example.[10] All adaptations help organisms survive in their ecological niches.[11] These adaptations may be structural, behavioral or physiological. Structural adaptations are physical features of an organism (shape, body covering, defensive or offensive armament); and also the internal organization). Behavioural adaptations are inherited behaviour chains and the ability to learn: they may be inherited in detail (instincts), or a tendency for learning may be inherited (see neuropsychology). Examples: searching for food, mating, vocalizations. Physiological adaptations permit the organism to perform special functions (for instance, making venom, secreting slime, phototropism); but also more general functions such as growth and development, temperature regulation, ionic balance and other aspects of homeostasis. Adaptation, then, affects all aspects of the life of an organism. Brief history Main article: History of evolutionary thought Adaptation as a fact of life has been accepted by all the great thinkers who have tackled the world of living organisms. It is their explanations of how adaptation arises that separates these thinkers. A few of the most significant ideas:[12] * Empedocles did not believe that adaptation required a final cause (~ purpose), but "came about naturally, since such things survived". Aristotle, however, did believe in final causes. * In natural theology, adaptation was interpreted as the work of a deity, even as evidence for the existence of God. William Paley believed that organisms were perfectly adapted to the lives they lead, an argument that shadowed Leibnitz, who had argued that God had brought about the best of all possible worlds. Voltaire's Dr Pangloss[13] is a parody of this optimistic idea, and Hume also argued against design.[14] The Bridgewater Treatises are a product of natural theology, though some of the authors managed to present their work in a fairly neutral manner. The series was lampooned by Robert Knox, who held quasi-evolutionary views, as the Bilgewater Treatises. Darwin broke with the tradition by emphasising the flaws and limitations which occurred in the animal and plant worlds.[15]

Lamarck * Lamark. His is a proto-evolutionary theory of the inheritance of acquired traits, whose main purpose is to explain adaptations. He proposed a tendency for organisms to become more complex, moving up a ladder of progress, plus "the influence of circumstances", usually expressed as use and disuse. His ideas and those of Geoffroy, fail because they cannot be reconciled with heredity. This was known even before Mendel by medical men interested in human races (Wells, Lawrence), and especially by Weismann. Many other students of natural history, such as Buffon, accepted adaptation, and some also accepted evolution, without voicing their opinions as to the mechanism. This illustrates the real merit of Darwin and Wallace, and secondary figures such as Bates, for pushing forward a

point of view whose merit had only been glimpsed previously. A century later, experimental field studies and breeding experiments by such as Ford and Dobzhansky produced evidence that natural selection was not only the 'engine' behind adaptation, but was a much stronger force than had previously been thought.[16][17] Types of adaptation Adaptation is the heart and soul of evolution. Niles Eldredge [18] Changes in habitat Before Darwin, adaptation was seen as a fixed relationship between an organism and its habitat. It was not appreciated that as the climate changed, so did the habitat; and as the habitat changed, so did the biota. Also, habitats are subject to changes in their biota: for example, invasions of species from other areas. The relative numbers of species in a given habitat are always changing. Change is the rule, though much depends on the speed and degree of the change. When the habitat changes, three main things may happen to a resident population: habitat tracking, genetic change or extinction. In fact, all three things may occur in sequence. Of the three effects, only genetic change brings about adaptation. It is also possible that some limited change may take place without immediate genetic changes. Populations differ in their phenotypic plasticity, which is the ability of an organism with a given genotype to change its phenotype in response to changes in its habitat, or to its move to a different habitat.[19][20] Habitat tracking When a habitat changes, the most common thing to happen is that the resident population moves to another locale which suits it; this is the typical response of flying insects or oceanic organisms, who have wide (though not unlimited) opportunity for movement.[21] This common response is called habitat tracking. It is one explanation put forward for the periods of apparent stasis in the fossil record (the punctuated equilibrium thesis).[22] Genetic change Genetic change is what occurs in a population when natural selection acts on the genetic variability of the population. By this means, the population adapts genetically to its circumstances, irrespective of whether it does so by means of visible structures or cryptic physiological activity. Survival is the criterion. It is now clear that habitats and biota do frequently change. Therefore, it follows that the process of adaptation is never finally complete.[23] Over time, it may happen that the environment changes little, and the species comes to fit its surroundings better and better. On the other hand, it may happen that changes in the environment occur relatively rapidly, and then the species becomes less and less well adapted. Seen like this, adaptation is a genetic tracking process, which goes on all the time to some extent, but especially when the population cannot or does not move to another, less hostile area. Also, to a greater or lesser extent, the process affects every species in a particular ecosystem. Fitness (an organism's capacity to propagate its genes) is not a static concept.[24][25] Van Valen thought that even in a stable environment, competing species had to constantly adapt to maintain their relative standing. This became known as the Red Queen hypothesis. Intimate relationships: co-adaptations Main article: Co-adaptation In co-evolution, where the existence of one species is tightly bound up with the life of another species, new or 'improved' adaptations which occur in one species are often followed by the appearance and spread of

corresponding features in the other species. There are many examples of this; the idea emphasises that the life and death of living things is intimately connected, not just with the physical environment, but with the life of other species. These relationships are intrinsically dynamic, and may continue on a trajectory for millions of years, as has the relationship between flowering plants and insects (pollination).

Pollinator constancy: these two honeybees, active at the same time and place, selectively visit flowers from only one species, as can be seen by the colour of the pollen in their baskets * Predator-prey * Parasite-host * Infection-resistance * Symbiosis * Mutualism * Mimicry * Pollination syndrome * Co-extinction The gut contents, wing structures, and mouthpart morphologies of fossilized beetles and flies suggest that they acted as early pollinators. The association between beetles and angiosperms during the early Cretaceous period led to parallel radiations of angiosperms and insects into the late Cretaceous. The evolution of nectaries in late Cretaceous flowers signals the beginning of the mutualism between hymenopterans and angiosperms.[26] Mimicry Main article: Mimicry

A & B are real wasps; rest are mimics: 3 hoverflies, one beetle. Henry Walter Bates' work on Amazonian butterflies led him to develop the first scientific account of mimicry, especially the kind of mimicry which bears his name: Batesian mimicry.[27] This is the mimicry by a palatable species of an unpalatable or noxious species. A common example seen in temperate gardens is the hover-fly, many of which though bearing no sting mimic the warning colouration of hymenoptera (wasps and bees). Such mimicry does not need to be perfect to improve the survival of the palatable species.[28] Bates, Wallace and Mller believed that Batesian and Mllerian mimicry provided evidence for the action of natural selection, a view which is now standard amongst biologists.[29] All aspects of this situation can be, and have been, the subject of research.[30] Field and experimental work on these ideas continues to this day; the topic connects strongly to speciation, genetics and development.[31] * More on mimicry: Warning Colour and Mimicry Lecture outline from University College London The basic machinery: internal adaptations There are some types of adaptation which are of a general nature, to do with the overall co-ordination of the systems in the body. Such adaptations may have significant consequences. Examples, in vertebrates, would be temperature regulation, or improvements in brain function, or an effective immune system. An example in plants would be the development of the reproductive system in flowering plants.[32] The acquisition of such major adaptations has often served as the spark for adaptive radiation, and huge success for long periods of time for a whole group of animals or plants. Compromise and conflict between adaptations

It is a profound truth that Nature does not know best; that genetical evolution... is a story of waste, makeshift, compromise and blunder. Peter Medawar [33] All adaptations have a downside: horse legs are great for running on grass, but they can't scratch their backs; mammals' hair helps temperature, but offers a niche for ectoparasites; the only flying penguins do is under water. Adaptations serving different functions may be mutually destructive. Compromise and make-shift occur widely, not perfection. Selection pressures pull in different directions, and the adaptation that results is some kind of compromise.[34] Since the phenotype as a whole is the target of selection, it is impossible to improve simultaneously all aspects of the phenotype to the same degree. Ernst Mayr [35] Consider the antlers of the Irish elk, (often supposed to be far too large; in deer antler size has an allometric relationship to body size). Obviously antlers serve positively for defence against predators, and to score victories in the annual rut. But they are costly in terms of resource. Their size during the last glacial period presumably depended on the relative gain and loss of reproductive capacity in the population of elks during that time.[36] Another example: camouflage to avoid detection is destroyed when vivid colors are displayed at mating time. Here the risk to life is counterbalanced by the necessity for reproduction. The peacock's ornamental train (grown anew in time for each mating season) must reduce his manoeuverability and flight, and is hugely conspicuous; also, its growth costs food resources. Darwin's explanation was in terms of sexual selection: "it depends on the advantage which certain individuals have over other individuals of the same sex and species, in exclusive relation to reproduction." [37] The kind of sexual selection represented by the peacock is called 'mate choice', with an implication that the process selects the more fit over the less fit, and so has survival value.[38] The existence of sexual selection was for a long time in abeyance, but has been rehabilitated.[39] The conflict between the size of the human foetal brain at birth, (which cannot be larger than about 400ccs, else it will not get through the mother's pelvis) and the size needed for an adult brain (about 1400ccs), means the brain of a newborn child is quite immature. The most vital things in human life (locomotion, speech) just have to wait while the brain grows and matures. That is the result of the birth compromise. Much of the problem comes from our upright bipedal stance, without which our pelvis could be shaped more suitably for birth. Neanderthals had a similar problem.[40][41][42] Shifts in function Adaptation and function are two aspects of one problem. Julian Huxley [43] Pre-adaptations This occurs when a species or population has characteristics that are ideally suited for conditions which have not yet arisen. For example, the polyploid rice-grass Spartina townsendii is better adapted than either of its parent species to their own habitat of saline marsh and mudflats.[44] White Leghorn fowl are markedly more resistant to vitamin B deficiency than other breeds.[45] On a plentiful diet there is no difference, but on a restricted diet this preadaptation could be decisive. Pre-adaptation may occur because a natural population carries a huge quantity of genetic variability.[46] In diploid eukaryotes, this is a consequence of the system of sexual reproduction, where mutant alleles get partially shielded, for example, by the selective advantage of

heterozygotes. Micro-organisms, with their huge populations, also carry a great deal of genetic variability. The first experimental evidence of the pre-adaptive nature of genetic variants in micro-organisms was provided by Salvador Luria and Max Delbrck who developed fluctuation analysis, a method to show the random fluctuation of pre-existing genetic changes that conferred resistance to phage in the bacterium Escherichia coli. Co-option of existing traits: exaptation Main article: Exaptation The classic example is the ear ossicles of mammals, which we know from palaeontological and embrological studies originated in the upper and lower jaws and the hyoid of their Synapsid ancestors, and further back still were part of the gill arches of early fish.[47][48] We owe this esoteric knowledge to the comparative anatomists, who, a century ago, were at the cutting edge of evolutionary studies.[49] The word exaptation was coined to cover these shifts in function, which are surprisingly common in evolutionary history.[50] The origin of wings from feathers that were originally used for temperature regulation is a more recent discovery (see feathered dinosaurs). Related issues Non-adaptive traits Some traits appear to be not adaptive, that is, selectively neutral. There may be various causes: the utility of a trait is lost and does not now appear adaptive; the utility of a trait is unknown; the trait is a consequence of another trait that is adaptive (the Spandrel idea). Of course, a trait may have been adaptive at some point in an organism's evolutionary history, but habitats change, leading to adaptations becoming redundant or even a hindrance (maladaptations). Such adaptations are termed vestigial. The utility of adaptations will ebb and flow. Fitness landscapes; drift Main article: Fitness landscape Main article: Genetic drift Sewall Wright's explanation for evolutionary stasis was that organisms come to occupy adaptive peaks. In order to evolve to another, higher peak, the species would first have to pass through a valley of maladaptive intermediate stages. This could happen by genetic drift if the population were small enough. This was Wright's shifting balance theory of evolution.[51] There has been much skepticism among evolutionary biologists as to whether these rather delicate conditions hold often in natural populations.[52] Ronald Fisher felt that most populations in nature were too large for these effects of genetic drift to be important.[53] Extinction Main article: Extinction If a population cannot move or change sufficiently to preserve its longterm viability, then obviously, it will become extinct, at least in that locale. The species may or may not survive in other locales. Species extinction occurs when the death rate over the entire species (population, gene pool ...) exceeds the birth rate for a long enough period for the species to disappear. It was an observation of Van Valen that groups of species tend to have a characteristic and fairly regular rate of extinction.[54] Co-extinction Main article: Co-extinction Just as we have co-adaptation, there is also co-extinction. Co-extinction refers to the loss of a species due to the extinction of another; for example, the extinction of parasitic insects following the loss of their hosts. Co-extinction can also occur when a flowering plant loses its

pollinator, or through the disruption of a food chain.[55] "Species coextinction is a manifestation of the interconnectedness of organisms in complex ecosystems ... While co-extinction may not be the most important cause of species extinctions, it is certainly an insidious one".[56] Flexibility, acclimatization, learning Flexibility deals with the relative capacity of an organism to maintain themselves in different habitats: their degree of specialization. Acclimatization is a term used for automatic physiological adjustments during life; learning is the term used for improvement in mental performance during life. In biology these terms are preferred, not adaptation, for changes during life which improve the performance of individuals. These adjustments are not inherited genetically by the next generation. Their time-span is a single lifetime. Adaptation, on the other hand, occurs over many generations; it is a gradual process caused by natural selection which changes the genetic make-up of a population so the collective performs better in its niche. Flexibility To a greater or lesser extent, all living things can adjust to circumstances. Their degree of flexibility is inherited, and it varies (to some extent) between individuals. A highly specialized anmimal or plant lives only in a well-defined habitat, eats a specific type of food and cannot survive if its needs are not met. Many herbivores are like this; extreme examples are Koalas (eucalyptus); pandas (bamboo). A generalist, on the other hand, eats a range of food, and can survive in many different conditions. Examples are: Man, rats; crabs; many carnivores. The tendency to behave in a specialized or exploratory manner is inherited It is an adaptation. Rather different is developmental flexibility. "An animal or plant is developmentally flexible if when it is raised or transferred to new conditions it develops so that it is better fitted to survive in the new circumstances." [57] Once again, there are huge differences between species, and these capacities to be flexible are inherited. Acclimatization If humans move to a higher altitude, respiration and physical exertion become a problem, but after spending time in high altitude conditions they acclimatize to the pressure by increasing production of red blood corpuscles. The ability to acclimatize is an adaptation, but not the acclimatization itself. Fecundity goes down, but deaths from some tropical diseases also goes down. Over a longer period of time, some people will reproduce better at these high altitudes than others. They will contribute more heavily to later generations. Gradually the whole population becomes adapted to the new conditions. This we know takes place, because the performance of long-term communities at higher altitude is significantly better than the performance of new arrivals, even when the new arrivals have had time to make physiological adjustments.[58] Some kinds of acclimatization happen so rapidly that they are better called reflexes. The rapid colour changes in some flatfish, cephalopods, chameleons are examples.[59] Learning Social learning is supreme for humans, and is possible for quite a few mammals and birds: of course, that does not involve genetic transmission except to the extent that the capacities are inherited. Similarly, the capacity to learn is an inherited adaptation, but not what is learnt; the capacity for human speech is inherited, but not the details of language. Function and teleonomy Adaptation raises some issues concerning how biologists use key terms such as function.

Function To say something has a function is to say something about what it does for the organism, obviously. It also says something about its history: how it has come about. A heart pumps blood: that is its function. It also emits sound, which is just an ancillary side-effect. That is not its function. The heart has a history (which may be well or poorly understood), and that history is about how natural selection formed and maintained the heart as a pump. Every aspect of an organism that has a function has a history. Now, an adaptation must have a functional history: therefore we expect it must have undergone selection caused by relative survival in its habitat. It would be quite wrong to use the word adaptation about a trait which arose as a by-product.[60][61] It is widely regarded as unprofessional for a biologist to say something like "A wing is for flying", although that is their normal function. A biologist would be conscious that sometime in the remote past feathers on a small reptile had the function of retaining heat, and later many wings were not used for flying (e.g. penguins). So, the biologist would rather say that the wings on a bird or an insect usually had the function of aiding flight. That would carry the connotation of being an adaptation with a history of evolution by natural selection. Teleonomy Teleonomy is a term invented to describe the study of goal-directed functions which are not guided by the conscious forethought of man or any supernatural entity. It is contrasted with Aristotle's teleology, which has connotations of intention, purpose and foresight. Evolution is teleonomic; adaptation hoards hindsight rather than foresight. The following is a definition for its use in biology: Teleonomy: The hypothesis that adaptations arise without the existence of a prior purpose, but by the action of natural selection on genetic variability.[62] The term may have been suggested by Colin Pittendrigh in 1958;[63] it grew out of cybernetics and self-organising systems. Ernst Mayr, George C. Williams and Jacques Monod picked up the term and used it in evolutionary biology.[64][65][66][67] Philosophers of science have also commented on the term. Ernest Nagel analysed the concept of goal-directedness in biology;[68] and David Hull commented on the use of teleology and teleonomy by biologists: Haldane can be found remarking, "Teleology is like a mistress to a biologist: he cannot live without her but hes unwilling to be seen with her in public." Today the mistress has become a lawfully wedded wife. Biologists no longer feel obligated to apologize for their use of teleological language; they flaunt it. The only concession which they make to its disreputable past is to rename it teleonomy.[69] References 1. ^ Williams, George C. 1966. Adaptation and natural selection: a critique of some current evolutionary thought. Princeton. "Evolutionary adaptation is a phenomenon of pervasive importance in biology." p5 2. ^ The Oxford Dictionary of Science defines it as "Any change in the structure or functioning of an organism that makes it better suited to its environment". 3. ^ Huxley, Julian 1942. Evolution the modern synthesis. Allen & Unwin, London. p449 4. ^ Price P.W. 1980. The evolutionary biology of parasites. Princeton. 5. ^ Mayr E. 1963. Animal species and evolution. Harvard. 6. ^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance. Harvard. p562566 7. ^ Salzburger W., Mack T., Verheyen E., Meyer A. (2005). "Out of Tanganyika: Genesis, explosive speciation, key-innovations and

phylogeography of the haplochromine cichlid fishes" (PDF). BMC Evolutionary Biology 5 (17): 17. doi:10.1186/1471-2148-5-17. http://www.biomedcentral.com/content/pdf/1471-2148-5-17.pdf. 8. ^ Kornfield, Irv; Smith, Peter (November 2000). "African Cichlid Fishes: Model Systems for Evolutionary Biology". Annual Review of Ecology and Systematics 31: 163. doi:10.1146/annurev.ecolsys.31.1.163. http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.ecolsys.31.1. 163. 9. ^ Stebbins, G. Ledyard, Jr. 1950. Variation and evolution in plants. Columbia. Polyploidy, chapters 8 and 9. 10. ^ Margulis, Lynn (ed) 1991. Symbiosis as a source of evolutionary innovation: speciation and morphogenesis MIT. ISBN 0-262-13269-9 11. ^ Hutchinson G. Evelyn 1965. The ecological theatre and the evolutionary play. Yale. The niche is the central concept in evolutionary ecology; see especially part II The niche: an abstractly inhabited hypervolume. p2678 12. ^ references and details in their articles 13. ^ in Candide, ou l'optimisme 14. ^ Sober, Elliott 1993. Philosophy of biology. Oxford. Chapter 2 15. ^ Darwin, Charles. 1872. The origin of species. 6th edition, p397: Rudimentary, atrophied and aborted organs. 16. ^ Provine, William 1986. Sewall Wright and evolutionary biology. University of Chicago Press. 17. ^ Ford E.B. 1975. Ecological genetics, 4th ed. Chapman and Hall, London. 18. ^ Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate. Wiley N.Y. p33 19. ^ Price TD, Qvarnstrm A & Irwin DE 2003. The role of phenotypic plasticity in driving genetic evolution. Proc. Biol. Sci. 270 p14331440. 20. ^ Price T.D. 2006. Phenotypic plasticity, sexual selection and the evolution of colour patterns. J Exp Biol. 209 p23682376 21. ^ Eldredge, Niles 1986. Time frames: the rethinking of Darwinian evolution and the theory of punctuated equilibria. p136, Of glaciers and beetles. 22. ^ Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate. Wiley, N.Y. p64 23. ^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance. Harvard. Harvard. p481 and sequence tells how Darwin's ideas on adaptation developed as he came to appreciate it as "a continuing dynamic process" (bottom p483). 24. ^ Sterelny K. & Griffiths P.E. 1999. Sex and death: an introduction to philosophy of biology. University of Chicago Press. p217 ISBN O-22677304-3 25. ^ Freeman S. & Herron J.C. 2007. Evolutionary analysis. Pearson Education. p364 ISBN 0-13-227584-8 26. ^ Stebbins, G. Ledyard, Jr. 1974. Flowering plants: evolution above the species level. Harvard. 27. ^ Carpenter GDH and Ford EB 1933. Mimicry. Methuen, London. 28. ^ Wickler W. 1968. Mimicry in plants and animals. World University Library, London. 29. ^ Moon H.P. 1976. Henry Walter Bates FRS 1825-1892: explorer, scientist and darwinian. Leicestershire Museums, Leicester. 30. ^ Ruxton GD, Sherratt TN and Speed MP 2004. Avoiding attack: the evolutionary ecology of crypsis, warning signals and mimicry. Oxford. 31. ^ Mallet, James 2001. The speciation revolution. J Evolutionary Biology 14, 887-8.

32. ^ Stebbins, G. Ledyard, Jr. 1974. Flowering plants: evolution above the species level. Harvard. Contains an extensive analysis of the evolution of adaptations in the radiation of Angiosperms. 33. ^ Medawar, Peter 1960. The future of Man. Methuen, London. 34. ^ Jacob, Francois 1977. Evolution and tinkering. Science 196 1161 1166. 35. ^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance. Harvard. p589 36. ^ It is, of course, not possible to test selective pressures on extinct populations in any direct way. GOULD, STEPHEN J. (1974): Origin and function of 'bizarre' structures - antler size and skull size in 'Irish Elk', Megaloceros giganteus. Evolution 28(2): 191-220. doi:10.2307/2407322 (First page text) 37. ^ Darwin, Charles 1871. The Descent of Man and selection in relation to sex. Murray, London. 38. ^ The case was treated by Fisher R.A. 1930. Genetical theory of natural selection. Oxford. p134139. 39. ^ Cronin, Helen 1991. The ant and the peacock: altruism and sexual selection from Darwin to the present day. Cambridge. 40. ^ Rosenberg K.R. 2005. The evolution of modern human childbirth. Am J. Physical Anthropology 35, p89124. 41. ^ Friedlander, Nancy & Jordan, David K. 1995. Obstetric implications of Neanderthal robusticity and bone density. Human Evolution (Florence) 9: 331-342. 42. ^ Miller, Geoffrey 2007. Brain evolution. In Gangestad S.W. and Simpson J.A. (eds) The evolution of mind: fundamental questions and controversies. Guildford. 43. ^ Huxley, Julian 1942. Evolution the modern synthesis. Allen & Unwin, London. p417 44. ^ Huskins C.L. 1931. The origin of Spartina townsendii. Genetica 12, 531. 45. ^ Lamoreux W.F and Hutt F.B. 1939. Breed differences in resistance to a deficiency in vitamin B1 in the fowl. J. Agric. Res. Washington 58, 307315. 46. ^ [Dobzhansky T.] 1981. Dobzhansky's genetics of natural populations. eds Lewontin RC, Moore JA, Provine WB and Wallace B. Columbia University Press N.Y. 47. ^ Egdar F. Allin and James A. Hopson 1992. Evolution of the auditory system in Synapsida ("Mammal-like reptiles" and primitive mammals) as seen in the fossil record. Section IV (Mammals), Chapter 28, pages 587614 in The evolutionary biology of hearing edited by Douglas B. Webster, Richard R. Fay, and Arthur N. Popper. Springer-Verlag. ISBN 0-387-975888. 48. ^ Neil Shubin 2008. Your Inner Fish: a journey into the 3.5-billionyear history of the human body Pantheon Books 2008. ISBN 978-0-375-424472. Chapter 10, "Ears" 49. ^ Panchen, Alec. 1992. Classification, evolution and the nature of biology. Cambridge. Chapter 4 Homology and the evidence for evolution. 50. ^ Gould, Stephen Jay and Elizabeth S. Vrba 1982. Exaptation a missing term in the science of form. Paleobiology 8, 1, 415. 51. ^ Wright, Sewall 1932. The roles of mutation, inbreeding, crossbreeding, and selection in evolution. In Proceedings of the Sixth International Congress on Genetics, p355366. 52. ^ [Dobzhansky T.] 1981. Dobzhansky's genetics of natural populations. eds Lewontin RC, Moore JA, Provine WB and Wallace B. Columbia University Press N.Y. 53. ^ Provine, William 1986. Sewall Wright and evolutionary biology. University of Chicago Press.

54. ^ Van Valen L. 1973. A new evolutionary law. Evolutionary Theory 1, 130. 55. ^ Darwin in the Origin of Species tells the story of "a web of complex relations" involving heartsease (Viola tricolor), red clover (Trifolium pratense, humble-bees (bumblebees), mice and cats. Origin, 6th edition, p57. 56. ^ Koh, Lian Pih. 2004. Science, 305, 5690, 1632-1634, 10 September 2004. 57. ^ Maynard Smith J. 1993. The theory of evolution. Cambridge. 3rd ed, p33. 58. ^ Moore Lorna G. and Regensteiner Judith G. 1983. Adaptation to high altitude. Ann. Rev. Anthropology 12, p285304. 59. ^ Maynard Smith uses the term physiologically versatile for such animals. Maynard Smith J. 1993. The theory of evolution. Cambridge. 3rd ed, p32. 60. ^ Sober, Elliott 1993. Philosophy of biology. Oxford. p8586 61. ^ Williams, George C. 1966. Adaptation and natural selection: a critique of some current evolutionary thought. Princeton. p810 62. ^ "The hypothesis that adaptations arise without the existence of a prior purpose, but by chance may change the fitness of an organism." Oxford Dictionary of Zoology. But one might question the word chance, since natural selection, by its operation in particular habitats, is not a random process (it may be a stochastic or probabilistic process, however). 63. ^ Pittendrigh C.S. 1958. Adaptation, natural selection and behavior. In A. Roe and George Gaylord Simpson (eds) Behavior and evolution. Yale. 64. ^ Mayr, Ernst 1965. Cause and effect in biology. In D. Lerner (ed) Cause and effect. Free Press, New York. p3350. 65. ^ Mayr, Ernst 1988. Toward a new philosophy of biology. Chapter 3 "The multiple meanings of teleological". 66. ^ Williams, George C. 1966. Adaptation and natural selection; a critique of some current evolutionary thought. Chapter 9. Princeton. 67. ^ Monod, Jacques 1971. Chance and necessity: an essay on the natural philosophy of modern biology. Knopf, New York. ISBN 0-394-46615-2 68. ^ Nagel, E. 1977. Teleology revisited: goal-directed processes in biology. Journal of Philosophy 74: 261301. 69. ^ Hull D.L. 1982. Philosophy and biology. In G. Flistad (ed) Philosophy of Science Nijhoff. See also Evolutionary biology portal A planthopper mimics a leaf * Adaptive radiation * Co-adaptation * Co-evolution * Ecological trap * Exaptation * Intragenomic conflict * Maladaptation * Mimicry * Polymorphism (biology) v d e Basic topics in evolutionary biologyEvidence of common descentProcesses of evolutionAdaptation Macroevolution Microevolution SpeciationPopulation genetic mechanismsNatural selection Genetic drift Gene flow MutationEvolutionary developmental biology (Evo-devo) conceptsPhenotypic plasticity Canalisation ModularityThe evolution ofDNA Sex Aging Intelligence The Ear The Eye Flight Plants Fungi Life Humans Dolphins and whales

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Selection From Wikipedia, the free encyclopedia Jump to: navigation, search This article does not cite any references or sources. Please help improve this article by adding citations to reliable sources (ideally, using inline citations). Unsourced material may be challenged and removed. (November 2006) For other uses, see Selection (disambiguation). Phylogenetics Population genetics Biology Portal v d e In the context of evolution, certain traits or alleles of a species may be subject to selection. Under selection, individuals with advantageous or "adaptive" traits tend to be more successful than their peers reproductively--meaning they contribute more offspring to the succeeding generation than others do. When these traits have a genetic basis, selection can increase the prevalence of those traits, because offspring will inherit those traits from their parents. When selection is intense and persistent, adaptive traits become universal to the population or species, which may then be said to have evolved. Contents * 1 Overview * 2 Types and subtypes o 2.1 Patterns of selection o 2.2 Mechanisms of selection * 3 Further reading Overview Whether or not selection takes place depends on the conditions in which the individuals of a species find themselves. Adults, juveniles, embryos, and even eggs and sperm may undergo selection. Factors fostering selection include limits on resources (nourishment, habitat space, mates) and the existence of threats (predators, disease, adverse weather). Biologists often refer to such factors as selective pressures. Natural selection is the most familiar type of selection by name. The breeding of dogs, cows and horses, however, represents "artificial selection." Subcategories of natural selection are also sometimes distinguished. These include sexual selection, ecological selection, stabilizing selection, disruptive selection and directional selection (more on these below). Selection occurs only when the individuals of a population are diverse in their characteristics--or more specifically when the traits of individuals differ with respect to how well they equip them to survive or exploit a particular pressure. In the absence of individual variation, or when variations are selectively neutral, selection does not occur. Meanwhile, selection does not guarantee that advantageous traits or alleles will become prevalent within a population. Through genetic drift, such traits may become less common or disappear. In the face of selection even a so-called deleterious allele may become universal to the members of a species. This is a risk primarily in the case of "weak" selection (e.g. an infectious disease with only a low mortality rate) or small populations. Though deleterious alleles may sometimes become established, selection may act "negatively" as well as "positively." Negative selection decreases the prevalence of traits that diminish individuals' capacity to

succeed reproductively (i.e. their fitness), while positive selection increases the prevalence of adaptive traits. In biological discussions, traits subject to negative selection are sometimes said to be "selected against," while those under positive selection are said to be "selected for," as in the sentence Desert conditions select for drought tolerance in plants and select against shallow root architectures. Types and subtypes Selection is hierachically classified into natural and artificial selection. Natural selection is further subclassified into ecological and sexual selection Patterns of selection Aspects of selection may be divided into effects on a phenotype and their causes. The effects are called patterns of selection, and do not necessarily result from particular causes (mechanisms); in fact each pattern can arise from a number of different mechanisms. Stabilizing selection favors individuals with intermediate characteristics while its opposite, disruptive selection, favors those with extreme characteristics; directional selection occurs when characteristics lie along a phenotypic spectrum and the individuals at one end are more successful; and balancing selection is a pattern in which multiple characteristics may be favored. Mechanisms of selection Distinct from patterns of selection are mechanisms of selection; for example, disruptive selection often is the result of disassortative sexual selection, and balancing selection may result from frequencydependent selection and overdominance. Further reading * Bell, Graham (1997). Selection: The Mechanism of Evolution. New York: Chapman & Hall. pp. 699 p. ISBN 041205521X. (2nd edition published in 2008 by Oxford University Press, 553 p., ISBN 0198569726) v d e Topics in population genetics

Key concepts Hardy-Weinberg law Genetic linkage Linkage disequilibrium Fisher's fundamental theorem Neutral theory Price equation

Selection Natural Sexual Artificial Ecological

Effects of selection on genomic variation Genetic hitchhiking Background selection

Genetic drift Small population size Population bottleneck Founder effect Coalescence Balding-Nichols Model

Founders R.A. Fisher J. B. S. Haldane Sewall Wright

Related topics Evolution Microevolution Evolutionary game theory Fitness landscape Genetic genealogy

List of evolutionary biology topics Retrieved from "http://en.wikipedia.org/wiki/Selection" Categories: Evolutionary biology | Population genetics | Selection Hidden categories: Articles lacking sources from November 2006 | All articles lacking sources * This page was last modified on 23 March 2009, at 01:18 (UTC). * All text is available under the terms of the GNU Free Documentation License. (See Copyrights for details.) Wikipedia is a registered trademark of the Wikimedia Foundation, Inc., a U.S. registered 501(c)(3) tax-deductible nonprofit charity. * Privacy policy * About Wikipedia * Disclaimers

Speciation From Wikipedia, the free encyclopedia Jump to: navigation, search Phylogenetics Population geneticsBiology Portal v d e Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages.[1][2] Whether speciation is achieved normally via genetic drift or natural selection is the subject of much ongoing discussion. There are four geographic modes of speciation in nature, based on the extent to which speciating populations are geographically isolated from one another: allopatric, peripatric, parapatric, and sympatric. Speciation may also be induced artificially, through animal husbandry or laboratory experiments. Observed examples of each kind of speciation are provided throughout.[3] Contents * 1 Natural speciation o 1.1 Allopatric o 1.2 Peripatric o 1.3 Parapatric o 1.4 Sympatric * 1.4.1 Speciation via polyploidization * 1.4.2 Speciation via hybrid formation o 1.5 Reinforcement (Wallace effect) * 2 Artificial speciation * 3 Genetics o 3.1 Hybrid speciation o 3.2 Gene transposition as a cause o 3.3 Interspersed repeats o 3.4 Human speciation * 4 See also * 5 References * 6 Further reading * 7 External links Natural speciation All forms of natural speciation have taken place over the course of evolution, though it still remains a subject of debate as to the relative importance of each mechanism in driving biodiversity.[4]

The three-spined stickleback (Gasterosteus aculeatus) A good example of natural speciation is observed repeatedly in the threespined stickleback, a marine fish which frequently undergoes parapatric speciation into new freshwater colonies at the mouths of rivers. Over typically 10,000 generations, the stickleback may lose their pelvic hind fins, gain tougher bony armor in their scales, and lighten in color as they adapt to life in fresh water.[5] There is debate as to the rate at which speciation events occur over geologic time. While some evolutionary biologists claim that speciation events have remained relatively constant over time, some palaeontologists such as Niles Eldredge and Stephen Jay Gould have argued that species usually remain unchanged over long stretches of time, and that speciation occurs only over relatively brief intervals, a view known as punctuated equilibrium. Allopatric

Comparison of allopatric, peripatric, parapatric and sympatric speciation. Main article: allopatric speciation During allopatric speciation, a population splits into two geographically isolated allopatric populations (for example, by habitat fragmentation due to geographical change such as mountain building or social change such as emigration). The isolated populations then undergo genotypic and/or phenotypic divergence as they (a) become subjected to dissimilar selective pressures or (b) they independently undergo genetic drift. When the populations come back into contact, they have evolved such that they are reproductively isolated and are no longer capable of exchanging genes. Observed instances Island genetics, the tendency of small, isolated genetic pools to produce unusual traits, has been observed in many circumstances, including insular dwarfism and the radical changes among certain famous island chains, like Komodo and Galpagos, the latter having given rise to the modern expression of evolutionary theory, after being observed by Charles Darwin. Perhaps the most famous example of allopatric speciation is Darwin's Galpagos Finches. Peripatric Main article: peripatric speciation In peripatric speciation, new species are formed in isolated, small peripheral populations which are prevented from exchanging genes with the main population. It is related to the concept of a founder effect, since small populations often undergo bottlenecks. Genetic drift is often proposed to play a significant role in peripatric speciation. Observed instances * Mayr bird fauna * The Australian bird Petroica multicolor * Reproductive isolation occurs in populations of Drosophila subject to population bottlenecking The London Underground mosquito is a variant of the mosquito Culex pipiens which entered in the London Underground in the nineteenth century. Evidence for its speciation include genetic divergence, behavioral differences, and difficulty in mating.[6] Parapatric Main article: parapatric speciation In parapatric speciation, the zones of two diverging populations are separate but do overlap. There is only partial separation afforded by geography, so individuals of each species may come in contact or cross the barrier from time to time, but reduced fitness of the heterozygote leads to selection for behaviours or mechanisms which prevent breeding between the two species. Ecologists refer to parapatric and peripatric speciation in terms of ecological niches. A niche must be available in order for a new species to be successful. Observed instances * The three-spined stickleback[5] * Ring species o The Larus gulls form a ring species around the North Pole. o The Ensatina salamanders, which form a ring round the Central Valley in California. o The Greenish Warbler (Phylloscopus trochiloides), around the Himalayas.

* the grass Anthoxanthum has been known to undergo parapatric speciation in such cases as mine contamination of an area. Sympatric This date:March, 2008 needs additional citations for verification. Please help improve this article by adding reliable references (ideally, using inline citations). Unsourced material may be challenged and removed. (March 2008)Main article: sympatric speciation In sympatric speciation, species diverge while inhabiting the same place. Often cited examples of sympatric speciation are found in insects which become dependent on different host plants in the same area. However, the existence of sympatric speciation as a mechanism of speciation is still hotly contested. People have argued that the evidences of sympatric speciation are in fact examples of micro-allopatric, or heteropatric speciation. The most widely accepted example of sympatric speciation is that of the cichlids of Lake Nabugabo in East Africa, which is thought to be due to sexual selection. Sympatric speciation refers to the formation of two or more descendant species from a single ancestral species all occupying the same geographic location. Until recently, there has a been a dearth of hard evidence that supports this form of speciation, with a general feeling that interbreeding would soon eliminate any genetic differences that might appear. But there has been at least one recent study that suggests that sympatric speciation has occurred in Tennessee cave salamanders.[7] The three-spined sticklebacks, freshwater fishes, that have been studied by Dolph Schluter (who received his Ph.D. for his work on Darwin's finches with Peter Grant) and his current colleagues in British Columbia, provide an intriguing example that is best explained by sympatric speciation. They have found: * Two different species of three-spined sticklebacks in each of five different lakes. o a large benthic species with a large mouth that feeds on large prey in the littoral zone o a smaller limnetic species with a smaller mouth that feeds on the small plankton in open water. * DNA analysis indicates that each lake was colonized independently, presumably by a marine ancestor, after the last ice age. * DNA analysis also shows that the two species in each lake are more closely related to each other than they are to any of the species in the other lakes. * Nevertheless, the two species in each lake are reproductively isolated; neither mates with the other. * However, aquarium tests showed that o the benthic species from one lake will spawn with the benthic species from the other lakes and o likewise the limnetic species from the different lakes will spawn with each other. o These benthic and limnetic species even display their mating preferences when presented with sticklebacks from Japanese lakes; that is, a Canadian benthic prefers a Japanese benthic over its close limnetic cousin from its own lake. * Their conclusion: in each lake, what began as a single population faced such competition for limited resources that o disruptive selection competition favoring fishes at either extreme of body size and mouth size over those nearer the mean coupled with o assortative mating each size preferred mates like it - favored a divergence into two subpopulations exploiting different food in different parts of the lake.

o The fact that this pattern of speciation occurred the same way on three separate occasions suggests strongly that ecological factors in a sympatric population can cause speciation. Sympatric speciation driven by ecological factors may also account for the extraordinary diversity of crustaceans living in the depths of Siberia's Lake Baikal. Speciation via polyploidization Speciation via polyploidy: A diploid cell undergoes failed meiosis, producing diploid gametes, which self-fertilize to produce a tetraploid zygote. Polyploidy is a mechanism often attributed to causing some speciation events in sympatry. Not all polyploids are reproductively isolated from their parental plants, so an increase in chromosome number may not result in the complete cessation of gene flow between the incipient polyploids and their parental diploids (see also hybrid speciation). Polyploidy is observed in many species of both plants and animals. In fact, it has been proposed that all of the existing plants and most of the animals are polyploids or have undergone an event of polyploidization in their evolutionary history. However, reproduction is often by parthenogenesis since polyploid animals are often sterile. Rare instances of polyploid mammals are known, but most often result in prenatal death. Speciation via hybrid formation See Hybrid speciation section under the Genetics heading below. Reinforcement (Wallace effect) Reinforcement is the process by which natural selection increases reproductive isolation.[8] It may occur after two populations of the same species are separated and then come back into contact. If their reproductive isolation was complete, then they will have already developed into two separate incompatible species. If their reproductive isolation is incomplete, then further mating between the populations will produce hybrids, which may or may not be fertile. If the hybrids are infertile, or fertile but less fit than their ancestors, then there will be no further reproductive isolation and speciation has essentially occurred (e.g., as in horses and donkeys.) The reasoning behind this is that if the parents of the hybrid offspring each have naturally selected traits for their own certain environments, the hybrid offspring will bear traits from both, therefore would not fit either ecological niche as well as the parents did. The low fitness of the hybrids would cause selection to favor assortative mating, which would control hybridization. This is sometimes called the Wallace effect after the evolutionary biologist Alfred Russel Wallace who suggested in the late 19th century that it might be an important factor in speciation.[9] If the hybrid offspring are more fit than their ancestors, then the populations will merge back into the same species within the area they are in contact. Reinforcement is required for both parapatric and sympatric speciation. Without reinforcement, the geographic area of contact between different forms of the same species, called their "hybrid zone," will not develop into a boundary between the different species. Hybrid zones are regions where diverged populations meet and interbreed. Hybrid offspring are very common in these regions, which are usually created by diverged species coming into secondary contact. Without reinforcement the two species would have uncontrollable inbreeding. Reinforcement may be induced in artificial selection experiments as described below.

Artificial speciation New species have been created by domesticated animal husbandry, but the initial dates and methods of the initiation of such species are not clear. For example, domestic sheep were created by hybridisation, and no longer produce viable offspring with Ovis orientalis, one species from which they are descended.[10] Domestic cattle, on the other hand, can be considered the same species as several varieties of wild ox, gaur, yak, etc., as they readily produce fertile offspring with them.[11] The best-documented creations of new species in the laboratory were performed in the late 1980s. William Rice and G.W. Salt bred fruit flies, Drosophila melanogaster, using a maze with three different choices of habitat such as light/dark and wet/dry. Each generation was placed into the maze, and the groups of flies which came out of two of the eight exits were set apart to breed with each other in their respective groups. After thirty-five generations, the two groups and their offspring were isolated reproductively because of their strong habitat preferences: they mated only within the areas they preferred, and so did not mate with flies that preferred the other areas.[12] The history of such attempts is described in Rice and Hostert (1993).[13] Diane Dodd was also able to show how mating preferences can develop from reproductive isolation in Drosophila pseudoobscura fruit flies after only eight generations using different food types, starch and maltose.[14] Dodd's experiment has been easy for many others to replicate, including with other kinds of fruit flies and foods.[15] Genetics Few speciation genes have been found. They usually involve the reinforcement process of late stages of speciation. In 2008 a speciation gene causing reproductive isolation was reported.[16] It causes hybrid sterility between related subspecies. Hybrid speciation Main article: Hybrid speciation Hybridization between two different species sometimes leads to a distinct phenotype. This phenotype can also be fitter than the parental lineage and as such natural selection may then favor these individuals. Eventually, if reproductive isolation is achieved, it may lead to a separate species. However, reproductive isolation between hybrids and their parents is particularly difficult to achieve and thus hybrid speciation is considered an extremely rare event. The Mariana Mallard arose from hybrid speciation. Hybridization without change in chromosome number is called homoploid hybrid speciation. It is considered very rare but has been shown in Heliconius butterflies [17] and sunflowers. Polyploid speciation, which involves changes in chromosome number, is a more common phenomenon, especially in plant species. Gene transposition as a cause Theodosius Dobzhansky, who studied fruit flies in the early days of genetic research in 1930s, speculated that parts of chromosomes that switch from one location to another might cause a species to split into two different species. He mapped out how it might be possible for sections of chromosomes to relocate themselves in a genome. Those mobile sections can cause sterility in inter-species hybrids, which can act as a speciation pressure. In theory, his idea was sound, but scientists long debated whether it actually happened in nature. Eventually a competing

theory involving the gradual accumulation of mutations was shown to occur in nature so often that geneticists largely dismissed the moving gene hypothesis.[18] However, 2006 research shows that jumping of a gene from one chromosome to another can contribute to the birth of new species.[19] This validates the reproductive isolation mechanism, a key component of speciation.[20] Interspersed repeats Main article: Interspersed repeat Interspersed repetitive DNA sequences function as isolating mechanisms. These repeats protect newly evolving gene sequences from being overwritten by gene conversion, due to the creation of non-homologies between otherwise homologous DNA sequences. The non-homologies create barriers to gene conversion. This barrier allows nascent novel genes to evolve without being overwritten by the progenitors of these genes. This uncoupling allows the evolution of new genes, both within gene families and also allelic forms of a gene. The importance is that this allows the splitting of a gene pool without requiring physical isolation of the organisms harboring those gene sequences. Human speciation Humans have genetic similarities with chimpanzees and gorillas, suggesting common ancestors. Analysis of genetic drift and recombination using a Markov model suggests humans and chimpanzees speciated apart 4.1 million years ago.[21] See also * Species problem * Heteropatry * Chronospecies * Frozen plasticity * Koinophilia References 1. ^ Cook, O. F. 1906. Factors of species-formation. Science 23:506-507. 2. ^ Cook, O. F. 1908. Evolution without isolation. American Naturalist 42:727-731. 3. ^ Observed Instances of Speciation by Joseph Boxhorn. Retrieved 28 October 2006. 4. ^ J.M. Baker (2005). "Adaptive speciation: The role of natural selection in mechanisms of geographic and non-geographic speciation". Studies in History and Philosophy of Biological and Biomedical Sciences 36: 303326. doi:10.1016/j.shpsc.2005.03.005. available online 5. ^ a b Kingsley, D.M. (January 2009) "From Atoms to Traits," Scientific American, p. 57 6. ^ Katharine Byrne and Richard A Nichols (1999) "Culex pipiens in London Underground tunnels: differentiation between surface and subterranean populations" 7. ^ MATTHEW L. NIEMILLER, BENJAMIN M. FITZPATRICK, BRIAN T. MILLER (2008). "Recent divergence with gene flow in Tennessee cave salamanders (Plethodontidae: Gyrinophilus) inferred from gene genealogies". Molecular Ecology 17 (9): 22582275. available online 8. ^ Ridley, M. (2003) "Speciation - What is the role of reinforcement in speciation?" adapted from Evolution 3rd edition (Boston: Blackwell Science) tutorial online 9. ^ Ollerton, J. "Flowering time and the Wallace Effect" (PDF). Heredity, August 2005.

http://oldweb.northampton.ac.uk/aps/env/lbrg/journals/papers/OllertonHere dityCommentary2005.pdf. Retrieved on 2007-05-22. 10. ^ Hiendleder S., et al. (2002) "Molecular analysis of wild and domestic sheep questions current nomenclature and provides evidence for domestication from two different subspecies" Proceedings of the Royal Society B: Biological Sciences 269:893-904 11. ^ Nowak, R. (1999) Walker's Mammals of the World 6th ed. (Baltimore: Johns Hopkins University Press) 12. ^ Rice, W.R. and G.W. Salt (1988). "Speciation via disruptive selection on habitat preference: experimental evidence". The American Naturalist 131: 911917. doi:10.1086/284831. 13. ^ W.R. Rice and E.E. Hostert (1993). "Laboratory experiments on speciation: What have we learned in forty years?". Evolution 47: 1637 1653. doi:10.2307/2410209. 14. ^ Dodd, D.M.B. (1989) "Reproductive isolation as a consequence of adaptive divergence in Drosophila pseudoobscura." Evolution 43:13081311. 15. ^ Kirkpatrick, M. and V. Ravign (2002) "Speciation by Natural and Sexual Selection: Models and Experiments" The American Naturalist 159:S22S35 DOI 16. ^ http://www.sciencemag.org/cgi/content/short/323/5912/376 17. ^ Mavarez, J.; Salazar, C.A., Bermingham, E., Salcedo, C., Jiggins, C.D. , Linares, M. (2006). "Speciation by hybridization in Heliconius butterflies". Nature 441: 868. doi:10.1038/nature04738. 18. ^ University of Rochester Press Releases 19. ^ Masly, John P., Corbin D. Jones, Mohamed A. F. Noor, John Locke, and H. Allen Orr (September 2006). "Gene Transposition as a Cause of Hybrid Sterility in Drosophila". Science 313 (5792): 14481450. doi:10.1126/science.1128721. PMID 16960009. http://www.sciencemag.org/cgi/content/short/313/5792/1448. Retrieved on 2007-03-18. 20. ^ Minkel, J.R. (September 8, 2006) "Wandering Fly Gene Supports New Model of Speciation" Science News 21. ^ Hobolth A, Christensen OF, Mailund T, Schierup MH (2007) "Genomic Relationships and Speciation Times of Human, Chimpanzee, and Gorilla Inferred from a Coalescent Hidden Markov Model." PLoS Genet 3(2): e7 (doi:10.1371/journal.pgen.0030007) Further reading * Coyne, J. A. & Orr, H. A. (2004). Speciation. Sunderlands, Massachusetts: Sinauer Associates, Inc. ISBN 0-87893-089-2. * Grant, V. (1981). Plant Speciation (2nd Edit. ed.). New York: Columbia University Press. ISBN 0-231-05113-1. * Mayr, E. (1963). Animal Species and Evolution. Harvard University Press. ISBN 0-674-03750-2 * White, M. J. D. (1978). Modes of Speciation. San Francisco, California: W. H. Freeman and Company. ISBN 0-716-70284-3. * Dedicated issue of Philosophical Transactions B on Speciation in microorganisms is freely available. External links * Observed Instances of Speciation from the Talk.Origins Frequently Asked Questions * Speciation, and * Evidence for Speciation from Understanding Evolution by the University of California Museum of Paleontology * Speciation from John Hawks' Anthropology Weblog - paleoanthropology, genetics, and evolution * Speciation in the context of evolution

v d e SpeciationBasic conceptsSpecies Cline Chronospecies SpeciationModes of speciationAllopatric Heteropatric Peripatric Parapatric Sympatric Polyploidy PaleopolyploidyAuxiliary mechanismsSexual selection Assortative mating Punctuated equilibriumIntermediate stagesHybrid Ring species Haldane's rule v d e Basic topics in evolutionary biologyEvidence of common descentProcesses of evolutionAdaptation Macroevolution Microevolution SpeciationPopulation genetic mechanismsNatural selection Genetic drift Gene flow MutationEvolutionary developmental biology (Evo-devo) conceptsPhenotypic plasticity Canalisation ModularityThe evolution ofDNA Sex Aging Intelligence The Ear The Eye Flight Plants Fungi Life Humans Dolphins and whales Birds Evolution of the horse Spider evolution Evolution of sirenians Insect evolution Evolution of Mollusca Evolution of dinosaursModes of speciationAnagenesis Catagenesis CladogenesisHistoryHistory of evolutionary thought Charles Darwin On the Origin of Species Modern evolutionary synthesis Gene-centered view of evolution Life (classification trees)Other subfieldsEcological genetics Molecular evolution Phylogenetics SystematicsList of evolutionary biology topics Timeline of evolutionRetrieved from "http://en.wikipedia.org/wiki/Speciation" Categories: Ecology | Evolution | Evolutionary biology | Speciation Hidden categories: Articles needing additional references from March 2008 * This page was last modified on 1 April 2009, at 23:53 (UTC). * All text is available under the terms of the GNU Free Documentation License. (See Copyrights for details.) Wikipedia is a registered trademark of the Wikimedia Foundation, Inc., a U.S. registered 501(c)(3) tax-deductible nonprofit charity. * Privacy policy * About Wikipedia * Disclaimers