The Origin of Species (full title: On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life) by British naturalist Charles Darwin, first published on 24 November 1859, is one of the pivotal works in scientific history and arguably the pre-eminent work in biology.
In it, Darwin makes "one long argument", with copious empirical examples as support, for his theory that organisms gradually evolve not individually but in "groups" (now called populations) through the process of natural selection, a mechanism the book effectively introduced to the public. The work presents detailed scientific evidence that Darwin had accumulated on the Voyage of the Beagle in the 1830s and since his return, painstakingly laying out his theory and refuting the doctrine of "Created kinds", which underlay the then widely accepted theories of Creation biology.
The book is quite readable even for the non-specialist and attracted widespread interest on publication. Although its ideas are supported by an overwhelming body of scientific evidence and are widely accepted by scientists today, they are still highly controversial in some parts of the world, particularly among some non-scientists who perceive them to contradict various religious texts (the "creation-evolution controversy").
Theory in a nutshell
- Species have great fertility. They make more offspring than can grow to adulthood.
- Populations remain roughly the same size, with modest fluctuations.
- Food resources are limited, but are relatively stable over time.
- An implicit struggle for survival ensues.
- In sexually reproducing species, generally no two individuals are identical.
- Some of these variations directly impact the ability of an individual to survive in a given environment.
- Much of this variation is inheritable.
- Individuals less suited to the environment are less likely to survive and less likely to reproduce, while individuals more suited to the environment are more likely to survive and more likely to reproduce.
- The individuals that survive are most likely to leave their inheritable traits to future generations.
- This slowly effected process results in populations that adapt to the envirionment over time, and ultimately, after interminable generations, the creations of new varieties, and ultimately, new species.
The idea of biological evolution was around long before Darwin published The Origin, and was set out in Classical times by the Greek and Roman atomists, notably Lucretius. However, Christian thought in Medieval Europe involved complete faith in the ancient Biblical teachings of creation according to Genesis. Its concepts including "Created kinds" were interpreted by the priesthood as theology, then the Protestant Reformation widened access to the Bible and brought more literal interpretations. Natural history exploring the wonders of God's works made many discoveries and naturalists such as Carolus Linnaeus categorised an enormous number of species. A new belief developed that the original pair of every species had been brought into existence by God not so long ago. By the time of Darwin's birth in 1809, it was widely believed in England that both the natural world and the hierarchical social order were held stable, fixed by God's will, with nothing happening purely naturally and spontaneously.
The idea that fossils were the remains of extinct species, first put forward by Robert Hooke in the mid seventeenth century, gradually gained acceptance and several competing theories of geology were put forward, notably James Hutton's uniformitarian theory of 1785 which envisioned gradual change over aeons of time. Some individuals put forward evolutionary concepts. By 1796 Charles Darwin's grandfather Erasmus Darwin had proposed ideas of common descent with organisms "acquiring new parts" in response to stimuli then passing these changes to their offspring. In 1809 Jean-Baptiste Lamarck developed a similar theory, with "needed" traits being acquired through use then passed on. At this time the word evolution (from the Latin word "evolutio", meaning "unroll like a scroll") was used to refer to an orderly sequence of events, particularly one in which the outcome was somehow contained within it from the start, so Lamarck avoided using this word for his concept in which traits were acquired during an organism's life, and the term transmutation came into use.
Such ideas were seen in Britain as attacking the social order, already threatened by the aftermath of the American and French Revolutions. In England, natural history was dominated by the universities which trained clergy for the Church of England in William Paley's natural theology which sought evidence of beneficial "design" by a Creator. British naturalists adopted Georges Cuvier's explanation of the fossil record by catastrophism, the concept that animals and plants were periodically annihilated as a result of natural catastrophes and that their places were taken by new species created ex nihilo (out of nothing), modifying it to support the biblical account of Noah's flood. However Lamarck's ideas were taken up by Radicals who wanted to overturn the establishment and extend the vote to the lower classes.
Inception of Darwin's theory
Charles Darwin's education at the University of Edinburgh gave him direct involvement in Robert Edmund Grant's evolutionist developments of the ideas of Erasmus Darwin and Jean-Baptiste Lamarck. Then at Cambridge University his theology studies convinced him of William Paley's argument of "design" by a Creator while his interest in natural history was increased by the botanist John Stevens Henslow and the geologist Adam Sedgwick, both of whom believed strongly in divine creation. During the Voyage of the Beagle Charles Darwin became convinced by Charles Lyell's uniformitarianism, and puzzled over discrepancies between Lyell's uniformitarian idea that each species had its "centre of creation" and the evidence he saw. On his return Richard Owen showed that fossils Darwin had found were of extinct species related to current species in the same locality, and John Gould startlingly revealed that completely different birds from the Galápagos Islands were species of finches distinct to each island.
By early 1837 Darwin was speculating on transmutation in a series of secret notebooks. He investigated the breeding of domestic animals, consulting William Yarrell and reading a pamphlet by Yarrell's friend Sir John Sebright which commented that "A severe winter, or a scarcity of food, by destroying the weak and the unhealthy, has all the good effects of the most skilful selection." At the zoo in 1838 he had his first sight of an ape, and the orang-utan's antics impressed him as being "just like a naughty child" which from his experience of the natives of Tierra del Fuego made him think that there was little gulf between man and animals despite theological doctrines that only mankind possessed a soul.
In late September 1838 he began reading the 6th edition of Malthus's Essay on the Principle of Population which reminded him of Malthus's statistical proof that human populations breed beyond their means and compete to survive, at a time when he was primed to apply these ideas to animal species. Darwin applied to his search for the Creator's laws the Whig social thinking of struggle for survival with no hand-outs. By December 1838 he was seeing a similarity between breeders selecting traits and a Malthusian Nature selecting from variants thrown up by chance so that "every part of newly acquired structure is fully practised and perfected", thinking this "the most beautiful part of my theory".
First writings on the theory
Darwin was well aware of the implication the theory had for the origin of humanity and the real danger to his career and reputation as an eminent geologist of being convicted of blasphemy. He worked in secret to consider all objections and prepare overwhelming evidence supporting his theory. He increasingly wanted to discuss his ideas with his colleagues, and in January 1842 sent a tentative description of his ideas in a letter to Lyell, who was then touring America. Lyell, dismayed that his erstwhile ally had become a Transmutationist, noted that Darwin "denies seeing a beginning to each crop of species".
Despite problems with illness, Darwin formulated a 35 page "Pencil Sketch" of his theory in June 1842 then worked it up into a larger "essay". The botanist Joseph Dalton Hooker became Darwin's mainstay, and late in 1845 Darwin offered his "rough Sketch" for comments without immediate success, but in January 1847 when Darwin was particularly ill Hooker took away a copy of the "Sketch". After some delays he sent a page of notes, giving Darwin the calm critical feedback that he needed. Darwin made a huge study of barnacles which established his credentials as a biologist and provided more evidence supporting his theory.
In the spring of 1856 Lyell drew Darwin's attention to a paper on the "introduction" of species written by Alfred Russel Wallace, a naturalist working in Borneo, and urged Darwin to publish to establish priority. Darwin was now torn between the desire to set out a full and convincing account and the pressure to quickly produce a short paper. He ruled out exposing himself to an editor or counsel which would have been required to publish in an academic journal. On 14 May 1856 he began a "sketch" account and, by July, had decided to produce a full technical treatise on species.
Darwin pressed on, overworking, and was throwing himself into his work with his book on Natural Selection well under way, when on 18 June 1858 he received a parcel from Wallace enclosing about twenty pages describing an evolutionary mechanism, an unexpected response to Darwin's recent encouragement, with a request to send it on to Lyell. Darwin wrote to Lyell that "your words have come true with a vengeance,... forestalled" and he would, "of course, at once write and offer to send [it] to any journal" that Wallace chose, adding that "all my originality, whatever it may amount to, will be smashed". Lyell and Hooker agreed that a joint paper should be presented at the Linnean Society, and on 1 July 1858 the Wallace and Darwin papers entitled respectively On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection were read out, to surprisingly little reaction.
Darwin now worked hard on an "abstract" trimmed from his Natural Selection, writing much of it from memory. Lyell made arrangements with the publisher John Murray, who agreed to publish the manuscript sight unseen, and to pay Darwin two-thirds of the net proceeds. Darwin had decided to call his book An Abstract of an Essay on the Origin of Species and Varieties through Natural Selection, but with Murray's persuasion it was eventually reduced to the snappier On the Origin of Species through Natural Selection.
Publication of The Origin
The Origin was first published on 24 November 1859, price fifteen shillings, and was oversubscribed, so that all 1250 copies were claimed by booksellers that day. The second edition came out on 7 January 1860, and added "by the Creator" into the closing sentence, so that from then on it read "There is grandeur in this view of life, with its several powers, having been originally breathed by the Creator into a few forms or into one; and that, whilst this planet has gone circling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being evolved."
During Darwin's lifetime the book went through six editions, with cumulative changes and revisions to deal with counter-arguments raised. The third edition came out in 1861, and the fourth in 1866, each with an increasing number of sentences rewritten or added. The fifth edition published on 10 February 1869 incorporated more changes again, and for the first time included Herbert Spencer's phrase "survival of the fittest".
In January 1871 Mivart published On the Genesis of Species, the cleverest and most devastating critique of natural selection in Darwin's lifetime. Darwin took it personally and from April to the end of the year made extensive revisions to the Origin, using the word "evolution" for the first time and adding a new chapter to refute Mivart. He told Murray of working men in Lancashire clubbing together to buy the 5th edition at fifteen shillings, and he wanted a new cheap edition to make it more widely available.
Darwin's theory, as presented
Darwin opened his argument by pointing to the results of domestication, mostly through artificial selection (though environmental changes, such as more food and protection from predators, were also factors). Comparing domesticated and wild varieties, Darwin showed that the nineteenth-century definition of species was chiefly a matter of opinion, since the discovery of new linking forms often degraded species to varieties.
In his introduction, Darwin sets out the essence of his theory:
As many more individuals of each species are born than can possibly survive; and as, consequently, there is a frequently recurring struggle for existence, it follows that any being, if it vary however slightly in any manner profitable to itself, under the complex and sometimes varying conditions of life, will have a better chance of surviving, and thus be naturally selected. From the strong principle of inheritance, any selected variety will tend to propagate its new and modified form.
Darwin did not suggest that every variation and every character must have a selection value. However, he pointed out that, because of our ignorance of animal physiology and its relationship with the environment, it was extremely rash to set down any characters as valueless to their owners. It is even more important to notice that he did not suggest that every individual with a favorable variation must be selected, or that the selected or favored animals were better or higher, but merely that they were more adapted to their surroundings.
Darwin further infers that natural selection, if carried far enough, makes changes in a population, eventually leading to new species. He puts forward myriad observations as demonstrations of this, and also claims that the fossil record can be interpreted as supporting these observations. Darwin imagined it might be possible that all life is descended from an original species from ancient times. Modern DNA evidence is consistent with this idea.
In later editions of his book, starting in the fifth edition, in some instances where Darwin used the wording, Natural Selection, this was modified to "Natural Selection, or the Survival of the Fittest" in varying ways, borrowing the term survival of the fittest from Herbert Spencer.
Variation and heredity
One of the chief difficulties for Darwin and other naturalists in his time was that there was no agreed-upon model of heredity — in fact, the idea of heredity had not been completely separated conceptually from the idea of the development of the organism. Darwin himself saw variation and heredity as two essentially antagonistic forces, with most heredity working to preserve the fixity of a type rather than acting as the agent of species variability. Darwin's own model of heredity worked out in later works, which he dubbed "Pangenesis", was a mixture of a number of different ideas about heredity at the time. It contained what are now considered to be essentially Lamarckian aspects, whereby the effects of use and dis-use of different parts of the body in the parent could be transmitted to the child. Beyond this, it was essentially a model of "blended" heredity, by which the contributions of two parents (in the form of particles he called "gemmules") were roughly equal. Darwin was confident that even in this model, over long periods of time species would still be able to evolve.
It was not until the early 20th century that a model of heredity would become completely integrated with a model of variation, with the advent of the modern evolutionary synthesis known as neo-Darwinism. It is a common trope in the history of evolution and genetics written by scientists, rather than historians, to claim that Darwin's lack of an adequate model of heredity was the source of suspicion about his theory, but later historians of science have adequately documented the fact that this was not the source of most objections to Darwin, and that later scientists, such as Karl Pearson and the biometric school, could develop compelling models of evolution by natural selection with even a relatively simple "blending" model of heredity such as that used by Darwin.
Compatibility with Lamarckian inheritance
Contrarily to a common opinion, Darwin did not rule out at first the possibility of inherited acquired traits, and even mentions it explicitly in chapter 7:
"When the first tendency was once displayed, methodical selection and the inherited effects of compulsory training in each successive generation would soon complete the work; and unconscious selection is still at work, as each man tries to procure, without intending to improve the breed, dogs which will stand and hunt best. On the other hand, habit alone in some cases has sufficed; no animal is more difficult to tame than the young of the wild rabbit; scarcely any animal is tamer than the young of the tame rabbit; but I do not suppose that domestic rabbits have ever been selected for tameness; and I presume that we must attribute the whole of the inherited change from extreme wildness to extreme tameness, simply to habit and long-continued close confinement".
Nevertheless, there is a distinction between Lamarck's theory and Darwin's one: while Darwin's theory stays valid whether acquired traits are transmitted or not, Lamarck's theory becomes inoperative if acquired traits cannot be transmitted.
At the time of publication of Darwin's book, the educated public generally held the belief that science was a "friend of humanity" and that the natural world was orderly. This belief was based in part on advances made by Frenchman Louis Pasteur, who, in 1859, finally laid to rest the theory of spontaneous generation, Cite error: Invalid
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After publication, evolution was discussed and debated extensively. As well as attracting attention from naturalists and learned religious people, Huxley's "working-men's lectures" proved very popular and the 6th edition was halved in price, increasing sales and more widely disseminating the theory. The book was highly controversial, as it contradicted the then-prevailing theories of establishment scientists and theologians, as well as popularly held religious beliefs. The origin of species conflicted with a literal interpretation of the biblical creation accounts in the Book of Genesis. The religious controversy was fueled in part by one of Darwin's most vigorous defenders, Thomas Henry Huxley, who opined that Christianity is "a compound of some of the best and some of the worst elements of Paganism and Judaism, moulded in practice by the innate character of certain people of the Western World. Cite error: Invalid
<ref> tag. Tag has more than one name associated with reference. Perhaps the most uncompromising of the evolutionist philosophers was the German Ernst Heinrick Haeckel, a professor of biology, who dogmatically affirmed that nothing spiritual exists, and instead asserted that all life descended from protoplasm that spontaneously combined from essential protoplasmic elements in antiquity. The antireligious implications of Darwin's theory and their dizzying effect on the middle classes of Europe and United States was softened by the so-called Social Darwinists, such as Herbert Spencer, who promoted the virtues of social competition in fields outside biology.
Not all of the debates were based on religion, however. Many of the debates did not center around Darwin's specifically proposed mechanism for evolution — natural selection — but rather on the concept of evolution in general. Though Darwin himself was too sickly to defend his work in public, four of his close scientific friends took up the cause of promoting Darwin's work and defending it against critics. Chief among these were Huxley, who argued for the evidence of evolution in anatomical morphology, and Joseph Dalton Hooker, the Royal botanist at Kew Gardens. In the United States, Asa Gray worked in close correspondence with Darwin to assure the theory's spread despite the opposition of one of the most prominent scientists in the country at the time, Louis Agassiz, and helped to facilitate American publication of the book.
Scientific reaction to Darwin's theory was mixed. Many well-respected members of the scientific community, such as the aforementioned Agassiz and the anatomist Richard Owen, came out strongly against Darwin's work. On the whole, though, Darwin was successful in convincing many scientists, especially of the younger generations, that evolution had happened in one form or another. Over the course of the next two decades, most scientists and educated lay-people would come to believe that evolution had occurred. Natural selection, though, did not find wide support, and was actively attacked and relatively unpopular until its revival during the creation of the modern evolutionary synthesis in the 1920 and 1930s. Similarly, Darwin's notion that evolution occurred gradually was also often attacked, and many of the evolutionary theories which flourished during what Peter J. Bowler has called the "eclipse of Darwinism" were forms of "saltationism", in which new species arose through "jumps" rather than gradual adaptation.
With the 1874 publication of, "What is Darwinism?", the theologian Charles Hodge argued that Darwin's theories were tantamount to atheism.Cite error: Invalid
<ref> tag. Tag has more than one name associated with reference. This is an argument that had been made by many almost immediately after Darwin's first publication. Evolutionary origin of species contradicts the literal readings of many religious accounts; therefore, those who accepted the theory grew more skeptical of the Bible or other religious sources. As Hodge pointed out, evolution does not seem to originate from a divine source, and some viewed God as a less powerful force in the universe.
Darwin's theory changed the way humans saw themselves and their world. If one accepted that humans were descended from animals, it became clear that humans also are animals. The natural world took on a darker tinge in the minds of many, as animals in the wild are understood to be in a constant state of deadly competition with one another. The world was also seen in a less permanent fashion; since the world was apparently much different millions of years ago, it dawned on many that the impact of human beings would lessen and perhaps disappear altogether over time.
From the 1860s up until the 1930s, Darwinian "selectionist" evolution was not universally accepted by scientists, while evolution of some form generally was (a variety of evolutionary theories competed for scientific approval, including neo-Darwinism, neo-Lamarckism, orthogenesis, and mutation theory). In the 1930s, the work of a number of biologists and statisticians (especially R. A. Fisher) created the modern synthesis of evolution, which merged Darwinian selection theory with sophisticated statistical understandings of Mendelian genetics.
Today, the overwhelming majority of scientists in the fields of earth and life sciences (over 99.8%) consider Darwin's theory correct. "Often, claims that scientists reject evolution or support creationism are exaggerated or fraudulent. Many scientists doubt some aspects of evolution, especially recent hypotheses about it. All good scientists are skeptical about evolution (and everything else) and open to the possibility, however remote, that serious challenges to it may appear. Creationists frequently seize such expressions of healthy skepticism to imply that evolution is highly questionable. They fail to understand that the fact that evolution has withstood many years of such questioning really means it is about as certain as facts can get." A significant proportion of non-scientists in the United States and a few other countries disagree, often on religious grounds.
Misconceptions, and comparison to Wallace's theory
Darwin was not the first to form evolutionary theories. The idea that species might evolve, i.e., that current species have arisen from previous ones, was already under discussion at the time, and several mechanisms had been proposed dating back to Anaximander's theory of aquatic decent in the sixth century B.C., as well as Jean-Baptiste Lamarck's 1809 hypothesis on the inheritance of acquired characteristics, which was well known at the time of publication.
Darwin's achievements were fourfold: Firstly, to propose a credible mechanism (natural selection); secondly, to provide a great deal of new evidence for evolution; thirdly, to present his ideas in a compelling book; and fourthly, to ally with other highly motivated and influential biologists and philosophers in a concerted effort to publicize and advocate his ideas. On every point, Darwin was successful.
Like many great scientists, Darwin did not invent his theory from the ground up. He seized upon earlier research to create a comprehensive and defendable theory. As he subsequently acknowledged, others before him published brief statements outlining the principle of natural selection, but he was not aware of these little known statements until after publication of the Origin. Instead, he and Wallace put forth the first convincing and coherent mechanism of evolution: natural selection. Darwin's work, through its long list of facts and its support by prominent naturalists, established for most that evolution of some form did occur—that there was no fixity of species—even if there was considerable disagreement on the mechanism. Also contrary to a common understanding, Darwin did not invent the phrase "survival of the fittest", but added this in the 5th edition of The Origin of Species, giving due credit to the philosopher Herbert Spencer (who had introduced the phrase in his Principles of Biology of 1864) and usually using the phrase "Natural Selection, or the Survival of the Fittest". Other aspects of Darwin's overall theory which themselves evolved over time were: common descent, sexual selection, gradualism, and pangenesis.
Darwin's explanation of natural selection was slightly different from that given by Wallace. Darwin used comparison to selective breeding and artificial selection as a means for understanding natural selection. No such connection between selective breeding and natural selection was made by Wallace; he expressed it simply as a basic process of nature and did not think the phenomena were in any way related. On Wallace's own first edition of The Origin of Species, he crossed out every instance of the phrase "natural selection" and replaced with it Spencer's "survival of the fittest." He also ruled out much of the ideas of Lamarckian inheritance present in Darwin's work, calling it "quite unnecessary." Darwin and Wallace would disagree on many substantive issues later in their lives especially, most bitterly on the question of whether human consciousness had itself evolved (to Darwin's horror, Wallace eventually turned against this and towards spiritualism).
According to Ernst Mayr, Darwin's evolutionary thinking rests on a rejection of essentialism, which assumes the existence of some perfect, essential form for any particular class of existent, and treats differences between individuals as imperfections or deviations away from the perfect essential form. Darwin embraced instead what Mayr calls "population thinking", which denies the existence of any essential form and holds that a class is the conceptualization of the numerous unique individuals. Individuals, in short, are real in an objective sense, while the class is an abstraction, an artifact of epistemology. This emphasis on the importance of individual differences is necessary if one believes that the mechanism of evolution, natural selection, operates on individual differences.
Mayr claims essentialism had dominated Western thinking for two thousand years, and that Darwin's theories thus represent an important and radical break from traditional Western philosophy. Ripples of Darwin's thought can now be seen in fields such as economics and complexity theory, suggesting that Darwin's influence extends well beyond the field of biology.
The historian of science Peter J. Bowler has suggested that many of the "implications" attributed to Darwinism had little to do with Darwin's theories themselves. Many of the so-called "Darwinists" of the late-nineteenth century, such as Herbert Spencer and Ernst Haeckel, were actually very non-Darwinian in many aspects of their thought and theory, and even the biggest supporters of Darwin, such as Thomas Henry Huxley, were suspicious as to whether natural selection was really what caused evolution. Nevertheless, Darwin became quickly identified with evolution in general and hailed as the figurehead of many conceptual changes in both science and society, whether or not all of these ideas were stated explicitly or at all in Darwin's work itself.
- William Benjamin Carpenter Darwin on the Origin of Species. National Review 10: December 1859 188-214
- Thomas Henry Huxley Time and life: Mr Darwin's "Origin of species." Macmillan's Magazine 1: 1859 142-148.
- Richard Owen Review of Darwin's Origin of Species, Edinburgh Review, 3, April 1860, pp. 487-532.]
- Thomas Henry Huxley Review of The Origin of Species, Westminster Review 17 (n.s.) April 1860, pp. 541-70.
- Samuel Wilberforce (Review of) ‘On the origin of species’, Quarterly Review, June 1860, pp. 225-264.
- Andrew Murray On Mr Darwin's theory of the origin of species. Proceedings of the Royal Society of Edinburgh 4: 1860 274-291.
- Fleeming Jenkin 'Review of Darwin's The origin of species' The North British Review, June 1867, 46, pp. 277-318.
- ↑ Freeman 1977
- ↑ Mayr 1982
- ↑ Darwin 1859, Chapter XIV. Recapitulation and Conclusion.
- ↑ Moore 2006
- ↑ Desmond & Moore 1991, p. 34-35.
- ↑ Browne 1995, p. 129, 139-140.
- ↑ Desmond & Moore 1991, p. 295.
- ↑ Darwin 1859, p. 5.
- ↑ Bowler 1989
- ↑ 10.0 10.1 10.2 10.3 10.4 Burns, Edward M.; Philip Lee Ralph, Robert E. Lerner, Standish Meacham (1982). World Civilizations Their History and Their Culture. W.W. Norton & Company, 964-965. ISBN 0-393-95077-8.
- ↑ Huxley, Thomas H. (1902). Collected Essays, V. Kessinger Publishing. ISBN 978-1417973729.
- ↑ Hodge, Charles (1874). What is Darwinism?. Scribner, Armstrong, and Company. ASIN B0006AEEMO.
- ↑ http://www.talkorigins.org/indexcc/CA/CA111.html
- ↑ Kenneth, Chang. "Few Biologists but Many Evangelicals Sign Anti-Evolution Petition", New York Times, 2006-02-21. Retrieved on 2006-12-21. “But random interviews with 20 people who signed the [Discovery Institute's] petition and a review of the public statements of more than a dozen others suggest that many are evangelical Christians, whose doubts about evolution grew out of their religious beliefs.”
- ↑ http://www.religioustolerance.org/ev_publi.htm
- ↑ Bowler 1989
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Basic topics in evolutionary biology
|Evidence of evolution|
|Processes of evolution: adaptation - macroevolution - microevolution - speciation|
|Population genetic mechanisms: selection - genetic drift - gene flow - mutation|
|Evo-devo concepts: phenotypic plasticity - canalisation - modularity|
|Modes of evolution: anagenesis - catagenesis - cladogenesis|
|History: History of evolutionary thought - Charles Darwin - The Origin of Species - modern evolutionary synthesis|
|Other subfields: ecological genetics - human evolution - molecular evolution - phylogenetics - systematics|
|List of evolutionary biology topics | Timeline of evolution|