On the Origin of Species

1761: 2627:... I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life), together with subsequent modification and the multiplication of new forms. We can thus understand the high importance of barriers, whether of land or water, which separate our several zoological and botanical provinces. We can thus understand the localisation of sub-genera, genera, and families; and how it is that under different latitudes, for instance in South America, the inhabitants of the plains and mountains, of the forests, marshes, and deserts, are in so mysterious a manner linked together by affinity, and are likewise linked to the extinct beings which formerly inhabited the same continent ... On these same principles, we can understand, as I have endeavoured to show, why oceanic islands should have few inhabitants, but of these a great number should be endemic or peculiar; ... 2614:, starting with the observation that differences in flora and fauna from separate regions cannot be explained by environmental differences alone; South America, Africa, and Australia all have regions with similar climates at similar latitudes, but those regions have very different plants and animals. The species found in one area of a continent are more closely allied with species found in other regions of that same continent than to species found on other continents. Darwin noted that barriers to migration played an important role in the differences between the species of different regions. The coastal sea life of the Atlantic and Pacific sides of Central America had almost no species in common even though the 2907: 1795:. On 31 March Darwin wrote to Murray in confirmation, and listed headings of the 12 chapters in progress: he had drafted all except "XII. Recapitulation & Conclusion". Murray responded immediately with an agreement to publish the book on the same terms as he published Lyell, without even seeing the manuscript: he offered Darwin ⅔ of the profits. Darwin promptly accepted with pleasure, insisting that Murray would be free to withdraw the offer if, having read the chapter manuscripts, he felt the book would not sell well (eventually Murray paid £180 to Darwin for the first edition and by Darwin's death in 1882 the book was in its sixth edition, earning Darwin nearly £3000). 12589: 1638: 2693:

dependent on each other in so complex a manner, have all been produced by laws acting around us ... Thus, from the war of nature, from famine and death, the most exalted object which we are capable of conceiving, namely, the production of the higher animals, directly follows. There is grandeur in this view of life, with its several powers, having been originally breathed into a few forms or into one; and that, whilst this planet has gone cycling 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.

2598:'s findings that the earliest members of a class were a few simple and generalised species with characteristics intermediate between modern forms, and were followed by increasingly diverse and specialised forms, matching the branching of common descent from an ancestor. Patterns of extinction matched his theory, with related groups of species having a continued existence until extinction, then not reappearing. Recently extinct species were more similar to living species than those from earlier eras, and as he had seen in South America, and 3237: 3614: 1448:"warring of the species" in plants; he immediately envisioned "a force like a hundred thousand wedges" pushing well-adapted variations into "gaps in the economy of nature", so that the survivors would pass on their form and abilities, and unfavourable variations would be destroyed. By December 1838, he had noted a similarity between the act of breeders selecting traits and a Malthusian Nature selecting among variants thrown up by "chance" so that "every part of newly acquired structure is fully practical and perfected". 2343:), and Chapter V discusses what he called the effects of use and disuse; he wrote that he thought "there can be little doubt that use in our domestic animals strengthens and enlarges certain parts, and disuse diminishes them; and that such modifications are inherited", and that this also applied in nature. Darwin stated that some changes that were commonly attributed to use and disuse, such as the loss of functional wings in some island-dwelling insects, might be produced by natural selection. In later editions of 1485: 3098: 1041: 36: 194: 2484:

hybrids, varied greatly, especially among plants. Sometimes what were widely considered to be separate species produced fertile hybrid offspring freely, and in other cases what were considered to be mere varieties of the same species could only be crossed with difficulty. Darwin concluded: "Finally, then, the facts briefly given in this chapter do not seem to me opposed to, but even rather to support the view, that there is no fundamental distinction between species and varieties."

3464:. Historians write that most such political and economic commentators had only a superficial understanding of Darwin's scientific theory, and were as strongly influenced by other concepts about social progress and evolution, such as the Lamarckian ideas of Spencer and Haeckel, as they were by Darwin's work. Darwin objected to his ideas being used to justify military aggression and unethical business practices as he believed morality was part of fitness in humans, and he opposed 2619:

so commonly and notoriously the case." Darwin explained how a volcanic island formed a few hundred miles from a continent might be colonised by a few species from that continent. These species would become modified over time, but would still be related to species found on the continent, and Darwin observed that this was a common pattern. Darwin discussed ways that species could be dispersed across oceans to colonise islands, many of which he had investigated experimentally.

2347:, Darwin expanded the role attributed to the inheritance of acquired characteristics. Darwin also admitted ignorance of the source of inheritable variations, but speculated they might be produced by environmental factors. However, one thing was clear: whatever the exact nature and causes of new variations, Darwin knew from observation and experiment that breeders were able to select such variations and produce huge differences in many generations of selection. The 920: 12947: 11941: 749: 2053: 1969: 11951: 2268:" in "a large and metaphorical sense, including dependence of one being on another"; he gives examples ranging from plants struggling against drought to plants competing for birds to eat their fruit and disseminate their seeds. He describes the struggle resulting from population growth: "It is the doctrine of Malthus applied with manifold force to the whole animal and vegetable kingdoms." He discusses checks to such increase including complex 5187:, Quote: "There nonetheless are a few cases in which Darwin does discuss selection processes in which groups are the units, and these will be the focus of the present chapter. But even here it does not matter whether the groups are from different 'races' or from the same race. It is nests of honeybees that compete with each other, and human tribes that compete with other human tribes. For Darwin, the question of group selection had nothing 2544:, and fossil collections were very poor. Evolved local varieties which migrated into a wider area would seem to be the sudden appearance of a new species. Darwin did not expect to be able to reconstruct evolutionary history, but continuing discoveries gave him well-founded hope that new finds would occasionally reveal transitional forms. To show that there had been enough time for natural selection to work slowly, he cited the example of 2293: 2194:

reflect on the vast diversity of the plants and animals which have been cultivated, and which have varied during all ages under the most different climates and treatment, I think we are driven to conclude that this greater variability is simply due to our domestic productions having been raised under conditions of life not so uniform as, and somewhat different from, those to which the parent-species have been exposed under nature.

12975: 736: 10114: 2318:(1871). Natural selection was expected to work very slowly in forming new species, but given the effectiveness of artificial selection, he could "see no limit to the amount of change, to the beauty and infinite complexity of the coadaptations between all organic beings, one with another and with their physical conditions of life, which may be effected in the long course of time by nature's power of selection". Using a 2771:

Darwin making concessions and adding details to address his critics, and recommended the first edition. James T. Costa said that because the book was an abstract produced in haste in response to Wallace's essay, it was more approachable than the big book on natural selection Darwin had been working on, which would have been encumbered by scholarly footnotes and much more technical detail. He added that some parts of

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descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion, instead of the species being, as we see them, well defined?" Darwin attributed this to the competition between different forms, combined with the small number of individuals of intermediate forms, often leading to extinction of such forms.

2023:, as well as numerous explanatory notes giving her own answers to doubts that Darwin expressed. Darwin corresponded with Royer about a second edition published in 1866 and a third in 1870, but he had difficulty getting her to remove her notes and was troubled by these editions. He remained unsatisfied until a translation by Edmond Barbier was published in 1876. A Dutch translation by 3513:

which from then on read "life, with its several powers, having been originally breathed by the Creator into a few forms or into one". While some commentators have taken this as a concession to religion that Darwin later regretted, Darwin's view at the time was of God creating life through the laws of nature, and even in the first edition there are several references to "creation".

2659:. He says, "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions?" This made no sense under doctrines of independent creation of species, as even 7177:, p. 60, "In this book, he was completely silent on the subject of human origins, although he did refer in several places to mankind as an example of biological details. The only words he allowed himself—and these out of a sense of duty that he must somewhere refer to human beings–were gnomic in their brevity. 'Light will be thrown on the origin of man and his history'." 1704:

a vengeance, ... forestalled" and he would "of course, at once write and offer to send 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 publication putting together Wallace's pages with extracts from Darwin's 1844 Essay and his 1857 letter to Gray should be presented at the

2721:, and his introduction ridicules that work as failing to provide a viable mechanism. Therefore, the first four chapters lay out his case that selection in nature, caused by the struggle for existence, is analogous to the selection of variations under domestication, and that the accumulation of adaptive variations provides a scientifically testable mechanism for evolutionary 1738:, but was concerned about "how it can be made scientific for a Journal, without giving facts, which would be impossible." He asked Hooker how many pages would be available, but "If the Referees were to reject it as not strictly scientific I would, perhaps publish it as pamphlet." He began his "abstract of Species book" on 20 July 1858, while on holiday at 2107:, as naturalist, I was much struck with certain facts in the distribution of the inhabitants of South America, and in the geological relations of the present to the past inhabitants of that continent. These facts seemed to me to throw some light on the origin of species—that mystery of mysteries, as it has been called by one of our greatest philosophers. 7193:, Quote: "There is no exception to the rule that every organic being naturally increases at so high a rate, that if not destroyed, the earth would soon be covered by the progeny of a single pair. Even slow-breeding man has doubled in twenty-five years, and at this rate, in a few thousand years, there would literally not be standing room for his progeny." 3559:, and it was believed that animals did not have spiritual qualities. This conflict could be reconciled by supposing there was some supernatural intervention on the path leading to humans, or viewing evolution as a purposeful and progressive ascent to mankind's position at the head of nature. While many conservative theologians accepted evolution, 2001:, but the German translation published in 1860 imposed Bronn's own ideas, adding controversial themes that Darwin had deliberately omitted. Bronn translated "favoured races" as "perfected races", and added essays on issues including the origin of life, as well as a final chapter on religious implications partly inspired by Bronn's adherence to 3345:, but Darwin's ideas had little impact in France, where any scientists supporting evolutionary ideas opted for a form of Lamarckism. The intelligentsia in Russia had accepted the general phenomenon of evolution for several years before Darwin had published his theory, and scientists were quick to take it into account, although the 2276:

immigration of others and, where suitable variations occurred, descendants of some species became adapted to new conditions. He remarks that the artificial selection practised by animal breeders frequently produced sharp divergence in character between breeds, and suggests that natural selection might do the same, saying:

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resembled Darwin's: he endorsed natural selection, rejected the inheritance of acquired characters, remained silent on the origin of variation, and identified "the altruism of bees, the regeneration of tissue, the effects of exercise, and the inheritance of disadvantageous traits" as puzzles presented by the theory.

1877:, and the phrase "by the Creator" added to the closing sentence. 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, with a number of sentences rewritten or added and an introductory appendix, 1873:, ensuring that the book promptly reached a large number of subscribers to the library. The second edition of 3,000 copies was quickly brought out on 7 January 1860, and incorporated numerous corrections as well as a response to religious objections by the addition of a new epigraph on page ii, a quotation from 2766:, it avoided the narrative style of the historical novel and cosmological speculation, though the closing sentence clearly hinted at cosmic progression. Darwin had long been immersed in the literary forms and practices of specialist science, and made effective use of his skills in structuring arguments. 7213:, Quote: "What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions?" 3252:

in the spring of 1860. Du Bois-Reymond was a committed supporter, securing Darwin an honorary degree from the University of Breslau, teaching his theory to students at the University of Berlin, and defending his name to paying audiences across Germany and The Netherlands. Du Bois-Reymond's exposition

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It seemed to me sufficient to indicate, in the first edition of my 'Origin of Species,' that by this work 'light would be thrown on the origin of man and his history;' and this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on

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Some other statements in the book are quietly effective at pointing out the implication that humans are simply another species, evolving through the same processes and principles affecting other organisms. For example, in Chapter III: "Struggle for Existence" Darwin includes "slow-breeding man" among

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was to include a "note on Man", but when Wallace enquired in December 1857, Darwin replied; "You ask whether I shall discuss 'man';—I think I shall avoid whole subject, as so surrounded with prejudices, though I fully admit that it is the highest & most interesting problem for the naturalist." On

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Later chapters provide evidence that evolution has occurred, supporting the idea of branching, adaptive evolution without directly proving that selection is the mechanism. Darwin presents supporting facts drawn from many disciplines, showing that his theory could explain a myriad of observations from

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It is interesting to contemplate an entangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and

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Chapter XII continues the discussion of biogeography. After a brief discussion of freshwater species, it returns to oceanic islands and their peculiarities; for example on some islands roles played by mammals on continents were played by other animals such as flightless birds or reptiles. The summary

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Chapter VIII addresses the idea that species had special characteristics that prevented hybrids from being fertile in order to preserve separately created species. Darwin said that, far from being constant, the difficulty in producing hybrids of related species, and the viability and fertility of the

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In a section on "organs of little apparent importance", Darwin discusses the difficulty of explaining various seemingly trivial traits with no evident adaptive function, and outlines some possibilities such as correlation with useful features. He accepts that we "are profoundly ignorant of the causes

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But how, it may be asked, can any analogous principle apply in nature? I believe it can and does apply most efficiently, from the simple circumstance that the more diversified the descendants from any one species become in structure, constitution, and habits, by so much will they be better enabled to

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It can be seen here that Darwin attributes the greater variation amongst individuals of domestic varieties compared to their progenitor populations in nature as being due to their "conditions of life" (environment) being "not so uniform as", and "somewhat different from" those of the parent species".

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recognised the implications of Wallace's paper and its possible connection to Darwin's work, although Darwin did not, and in a letter written on 1–2 May 1856 Lyell urged Darwin to publish his theory to establish priority. Darwin was torn between the desire to set out a full and convincing account and

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were possible. His strategy established that evolution through natural laws was worthy of scientific study, and by 1875, most scientists accepted that evolution occurred but few thought natural selection was significant. Darwin's scientific method was also disputed, with his proponents favouring the

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I might have adduced for this same purpose the differences between the races of man, which are so strongly marked; I may add that some little light can apparently be thrown on the origin of these differences, chiefly through sexual selection of a particular kind, but without here entering on copious

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has described the book as written in everyday language for a wide audience, but noted that Darwin's literary style was uneven: in some places he used convoluted sentences that are difficult to read, while in other places his writing was beautiful. Quammen advised that later editions were weakened by

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While the book was readable enough to sell, its dryness ensured that it was seen as aimed at specialist scientists and could not be dismissed as mere journalism or imaginative fiction. Though Richard Owen did complain in the Quarterly Review that the style was too easy for a serious work of science.

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The final chapter, "Recapitulation and Conclusion", reviews points from earlier chapters, and Darwin concludes by hoping that his theory might produce revolutionary changes in many fields of natural history. He suggests that psychology will be put on a new foundation and implies the relevance of his

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All the foregoing rules and aids and difficulties in classification are explained, if I do not greatly deceive myself, on the view that the natural system is founded on descent with modification; that the characters which naturalists consider as showing true affinity between any two or more species,

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In the sixth edition Darwin inserted a new chapter VII (renumbering the subsequent chapters) to respond to criticisms of earlier editions, including the objection that many features of organisms were not adaptive and could not have been produced by natural selection. He said some such features could

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and the construction of hexagonal cells by honey bees. Darwin noted that some species of slave-making ants were more dependent on slaves than others, and he observed that many ant species will collect and store the pupae of other species as food. He thought it reasonable that species with an extreme

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Murray's response was favourable, and a very pleased Darwin told Lyell on 30 March that he would "send shortly a large bundle of M.S. but unfortunately I cannot for a week, as the three first chapters are in three copyists' hands". He bowed to Murray's objection to "abstract" in the title, though he

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and Gilolo). It enclosed twenty pages describing an evolutionary mechanism, a response to Darwin's recent encouragement, with a request to send it on to Lyell if Darwin thought it worthwhile. The mechanism was similar to Darwin's own theory. Darwin wrote to Lyell that "your words have come true with

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The book was written for non-specialist readers and attracted widespread interest upon its publication. Darwin was already highly regarded as a scientist, so his findings were taken seriously and the evidence he presented generated scientific, philosophical, and religious discussion. The debate over

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interpreted natural selection as an instrument of God's design, with the cleric Charles Kingsley seeing it as "just as noble a conception of Deity". In the second edition of January 1860, Darwin quoted Kingsley as "a celebrated cleric", and added the phrase "by the Creator" to the closing sentence,

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The book aroused international interest and a widespread debate, with no sharp line between scientific issues and ideological, social and religious implications. Much of the initial reaction was hostile, in a large part because very few reviewers actually understood his theory, but Darwin had to be

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review of 3 December 1859 commented, "Much of Mr. Darwin's volume is what ordinary readers would call 'tough reading;' that is, writing which to comprehend requires concentrated attention and some preparation for the task. All, however, is by no means of this description, and many parts of the book

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was only a few miles wide. His explanation was a combination of migration and descent with modification. He went on to say: "On this principle of inheritance with modification, we can understand how it is that sections of genera, whole genera, and even families are confined to the same areas, as is

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Owing to this struggle for life, any variation, however slight and from whatever cause proceeding, if it be in any degree profitable to an individual of any species, in its infinitely complex relations to other organic beings and to external nature, will tend to the preservation of that individual,

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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

1810:, briefly stating how these illustrated Darwin's general principles and preparing the way for the larger work expected shortly: "Every body is interested in pigeons." Darwin responded that this was impractical: he had only the last chapter still to write. In September the main title still included " 10402: 7365:

the derivation of any particular species is never discussed, yet I thought it best, in order that no honourable man should accuse me of concealing my views, to add that by the work in question 'light would be thrown on the origin of man and his history.' It would have been useless and injurious to

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was published in Feb. 1871. As soon as I had become, in the year 1837 or 1838, convinced that species were mutable productions, I could not avoid the belief that man must come under the same law. Accordingly I collected notes on the subject for my own satisfaction, and not for a long time with any

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Another difficulty, related to the first one, is the absence or rarity of transitional varieties in time. Darwin commented that by the theory of natural selection "innumerable transitional forms must have existed," and wondered "why do we not find them embedded in countless numbers in the crust of

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is arbitrary, with experts disagreeing and changing their decisions when new forms were found. He concludes that "a well-marked variety may be justly called an incipient species" and that "species are only strongly marked and permanent varieties". He argues for the ubiquity of variation in nature.

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with short legs), and (2) ubiquitous small differences (example: slightly shorter or longer bill of pigeons). Both types of hereditary changes can be used by breeders. However, for Darwin the small changes were most important in evolution. In this chapter Darwin expresses his erroneous belief that

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has determined that the idea that Darwin delayed publication only dates back to the 1940s, and Darwin's contemporaries thought the time he took was reasonable. Darwin always finished one book before starting another. While he was researching, he told many people about his interest in transmutation

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Various biographers have proposed that Darwin avoided or delayed making his ideas public for personal reasons. Reasons suggested have included fear of religious persecution or social disgrace if his views were revealed, and concern about upsetting his clergymen naturalist friends or his pious wife

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Secondly, is it possible that an animal having, for instance, the structure and habits of a bat, could have been formed by the modification of some animal with wholly different habits? Can we believe that natural selection could produce, on the one hand, organs of trifling importance, such as the

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In his autobiography, Darwin said he had "gained much by my delay in publishing from about 1839, when the theory was clearly conceived, to 1859; and I lost nothing by it". On the first page of his 1859 book he noted that, having begun work on the topic in 1837, he had drawn up "some short notes"

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Chapter VI begins by saying the next three chapters will address possible objections to the theory, the first being that often no intermediate forms between closely related species are found, though the theory implies such forms must have existed. As Darwin noted, "Firstly, why, if species have

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The book was widely translated in Darwin's lifetime, but problems arose with translating concepts and metaphors, and some translations were biased by the translator's own agenda. Darwin distributed presentation copies in France and Germany, hoping that suitable applicants would come forward, as

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Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers, —ants making slaves, —the larvæ of ichneumonidæ feeding within the live bodies of caterpillars, —not as specially endowed or

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When we look to the individuals of the same variety or sub-variety of our older cultivated plants and animals, one of the first points which strikes us, is, that they generally differ much more from each other, than do the individuals of any one species or variety in a state of nature. When we

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While Darwin had been somewhat coy about human origins, not identifying any explicit conclusion on the matter in his book, he had dropped enough hints about human's animal ancestry for the inference to be made, and the first review claimed it made a creed of the "men from monkeys" idea from

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Chapter IV details natural selection under the "infinitely complex and close-fitting ... mutual relations of all organic beings to each other and to their physical conditions of life". Darwin takes as an example a country where a change in conditions led to extinction of some species,

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were controversial as they conflicted with the beliefs that species were unchanging parts of a designed hierarchy and that humans were unique, unrelated to other animals. The political and theological implications were intensely debated, but transmutation was not accepted by the scientific

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as a book of wide general interest became associated with ideas of social reform. Its proponents made full use of a surge in the publication of review journals, and it was given more popular attention than almost any other scientific work, though it failed to match the continuing sales of

2388:, he struggled with the theoretical difficulty that novel individual variations would tend to blend into a population. However, inherited variation could be seen, and Darwin's concept of selection working on a population with a range of small variations was workable. It was not until the 1514:

used evidence from the fossil record and embryology to support the claim that living things had progressed from the simple to the more complex over time. But it proposed a linear progression rather than the branching common descent theory behind Darwin's work in progress, and it ignored

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Individuals less suited to the environment are less likely to survive and less likely to reproduce; individuals more suited to the environment are more likely to survive and more likely to reproduce and leave their heritable traits to future generations, which produces the process of

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with a first printing of 1250 copies. The book had been offered to booksellers at Murray's autumn sale on Tuesday 22 November, and all available copies had been taken up immediately. In total, 1,250 copies were printed but after deducting presentation and review copies, and five for

2564:. Combining this with an estimate of recent rates of sedimentation and erosion, Darwin calculated that erosion of The Weald had taken around 300 million years. The initial appearance of entire groups of well-developed organisms in the oldest fossil-bearing layers, now known as the 5168:

The full title employs the term 'race' only in the broad biological use of the word, which refers to varieties throughout organic life; however, speculation about the implications of his views specifically for the question of the human races began almost as soon as the book was

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published a more developed theory in 1809. Both envisaged that spontaneous generation produced simple forms of life that progressively developed greater complexity, adapting to the environment by inheriting changes in adults caused by use or disuse. This process was later called

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allegorically rather than as a literal historical account; organisms were described by their mythological and heraldic significance as well as by their physical form. Nature was widely believed to be unstable and capricious, with monstrous births from union between species, and

10117: 1845:. He used the term broadly, and as well as discussions of "the several races, for instance, of the cabbage" and "the hereditary varieties or races of our domestic animals and plants", there are three instances in the book where the phrase "races of man" is used, referring to 5191:

to do with 'race.' Still, writing in the heyday of empire, Darwin saw European nations outcompeting the nations, kingdoms, and tribes that occupy the rest of the globe. In this one very salient example, Darwin did see races struggling with each other. In any event, the word

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affirmed that he did not want to expose his ideas to review by an editor as would have been required to publish in an academic journal. He began a "sketch" account on 14 May 1856, and by July had decided to produce a full technical treatise on species as his "big book" on

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By early October, he began to "expect my abstract will run into a small volume, which will have to be published separately." Over the same period, he continued to collect information and write large fully detailed sections of the manuscript for his "big book" on Species,

1714:, by Wallace and Darwin respectively, were read out but drew little reaction. While Darwin considered Wallace's idea to be identical to his concept of natural selection, historians have pointed out differences. Darwin described natural selection as being analogous to the 5196:

in Darwin's subtitle needs to be understood very broadly; it encompasses competition among individuals, competition among groups in the same 'race,' and competition from groups from different 'races.' This is a much broader meaning than the word 'race' tends to have

2813:"In the distant future I see open fields for far more important researches. Psychology will be based on a new foundation, that of the necessary acquirement of each mental power and capacity by gradation. Light will be thrown on the origin of man and his history." 7382:, Quote: "During many years it has seemed to me highly probable that sexual selection has played an important part in differentiating the races of man; but in my 'Origin of Species' (first edition, p. 199) I contented myself by merely alluding to this belief." 2697:

Darwin added the phrase "by the Creator" from the 1860 second edition onwards, so that the ultimate sentence begins "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".

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supported directed evolution, and compiled scientific and religious objections to natural selection. In response, Darwin made considerable changes to the sixth edition. The problems of the age of the Earth and heredity were only resolved in the 20th century.

7339:, Quote: "During many years I collected notes on the origin or descent of man, without any intention of publishing on the subject, but rather with the determination not to publish, as I thought that I should thus only add to the prejudices against my views." 1760: 1227:

clergymen, and science became a search for these laws. Geologists adapted catastrophism to show repeated worldwide annihilation and creation of new fixed species adapted to a changed environment, initially identifying the most recent catastrophe as the

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who explained beauty in plumage by design. Since 1858, Huxley had emphasised anatomical similarities between apes and humans, contesting Owen's view that humans were a separate sub-class. Their disagreement over human origins came to the fore at the

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Chapter X examines whether patterns in the fossil record are better explained by common descent and branching evolution through natural selection, than by the individual creation of fixed species. Darwin expected species to change slowly, but not at

895:. Within two decades, there was widespread scientific agreement that evolution, with a branching pattern of common descent, had occurred, but scientists were slow to give natural selection the significance that Darwin thought appropriate. During " 1992:

of New York, who got their edition out in mid-January 1860, and the other two withdrew. In a May letter, Darwin mentioned a print run of 2,500 copies, but it is not clear if this referred to the first printing only, as there were four that year.

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He later erroneously elaborates that changed "conditions of life" act on the reproductive organs to generate greater variability in the progeny. Even after 1860 when Darwin read the correct reason for greater variation in domestic varieties in

2444:. Darwin concludes: "If it could be demonstrated that any complex organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my theory would absolutely break down. But I can find out no such case." 7282:: 'Fuegians & Brazil, climate & habits of life so different good instance of how fixed races are, in face of very different external conditions. The slowness of any changes explained by constitutions selection & sexual selection'. 6500:, Quote: "We are profoundly ignorant of the causes producing slight and unimportant variations; and we are immediately made conscious of this by reflecting on the differences in the breeds of our domesticated animals in different countries" 2817:

Discussing this in January 1860, Darwin assured Lyell that "by the sentence I show that I believe man is in same predicament with other animals. Many modern writers have seen this sentence as Darwin's only reference to humans in the book;

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had accused Newton of introducing "occult qualities & miracles". The fourth edition in 1866 had further revisions. The fifth edition, published on 10 February 1869, incorporated more changes and for the first time included the phrase

3567:", defined narrowly as including rejection of design, was atheism though he accepted that Asa Gray did not reject design. Asa Gray responded that this charge misrepresented Darwin's text. By the early 20th century, four noted authors of 3507:

Natural theology was not a unified doctrine, and while some such as Louis Agassiz were strongly opposed to the ideas in the book, others sought a reconciliation in which evolution was seen as purposeful. In the Church of England, some

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dependency on slave workers had evolved in incremental steps. He suggested that bees that make hexagonal cells evolved in steps from bees that made round cells, under pressure from natural selection to economise wax. Darwin concluded:

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and will generally be inherited by its offspring ... I have called this principle, by which each slight variation, if useful, is preserved, by the term of Natural Selection, in order to mark its relation to man's power of selection.

1980:, an American colleague of Darwin, negotiated with a Boston publisher for publication of an authorised American version, but learnt that two New York publishing firms were already planning to exploit the absence of international 1519:. Darwin read it soon after publication, and scorned its amateurish geology and zoology, but he carefully reviewed his own arguments after leading scientists, including Adam Sedgwick, attacked its morality and scientific errors. 1457:

was completed. As he recalled in his autobiography, he had "at last got a theory by which to work", but it was only in June 1842 that he allowed himself "the satisfaction of writing a very brief abstract of my theory in pencil".

7222:, Quote: "The framework of bones being the same in the hand of a man, wing of a bat, fin of the porpoise, and leg of the horse … at once explain themselves on the theory of descent with slow and slight successive modifications." 2421:

The chapter then deals with whether natural selection could produce complex specialised structures, and the behaviours to use them, when it would be difficult to imagine how intermediate forms could be functional. Darwin said:

2123:. He mentions his years of work on his theory, and the arrival of Wallace at the same conclusion, which led him to "publish this Abstract" of his incomplete work. He outlines his ideas, and sets out the essence of his theory: 9752: 5713: 7231:

Darwin, C. R. Notebook C, CUL-DAR122.- Transcribed by Kees Rookmaaker. (Darwin Online), notes from de Beer, Gavin ed. 1960. Darwin's notebooks on transmutation of species. Part II. Second notebook (February to July 1838).

4866:-orthodox, than the subject makes inevitable. That I do not discuss origin of man.— That I do not bring in any discussions about Genesis &c, & only give facts, & such conclusions from them, as seem to me fair. 2647:

are those which have been inherited from a common parent, and, in so far, all true classification is genealogical; that community of descent is the hidden bond which naturalists have been unconsciously seeking, ...

2358:

Breeding of animals and plants showed related varieties varying in similar ways, or tending to revert to an ancestral form, and similar patterns of variation in distinct species were explained by Darwin as demonstrating

7453:"[review of] On the origin of species, by means of natural selection; or the preservation of favoured races in the struggle for life. By Charles Darwin, M. A., F.R.S. London, 1860. Quarterly Review 108: 225–264" 2439:

as examples of how bats might have evolved from non-flying ancestors. He discussed various simple eyes found in invertebrates, starting with nothing more than an optic nerve coated with pigment, as examples of how the

6827:, Quote: "When the views entertained in this volume on the origin of species, or when analogous views are generally admitted, we can dimly foresee that there will be a considerable revolution in natural history. ..." 3539:

was among those accepting evolution but attacking Darwin's naturalistic mechanism. Eventually it was realised that supernatural intervention could not be a scientific explanation, and naturalistic mechanisms such as

2223:

Historians have noted that naturalists had long been aware that the individuals of a species differed from one another, but had generally considered such variations to be limited and unimportant deviations from the

1629:

without causing outrage. He firmly intended to publish, but it was not until September 1854 that he could work on it full-time. His 1846 estimate that writing his "big book" would take five years proved optimistic.

2737:(true cause) if three things could be demonstrated: its existence in nature, its ability to produce the effects of interest, and its ability to explain a wide range of observations. This reflected the influence of 10334: 1577:. He conducted empirical research focusing on difficulties with his theory. He studied the developmental and anatomical differences between different breeds of many domestic animals, became actively involved in 9692:

An Essay on the Principle of Population: A View of its Past and Present Effects on Human Happiness; with an Inquiry into Our Prospects Respecting the Future Removal or Mitigation of the Evils which It Occasions

5406:"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." 2322:

and calculations, he indicates the "divergence of character" from original species into new species and genera. He describes branches falling off as extinction occurred, while new branches formed in "the great

1657:, claimed that patterns in the geographical distribution of living and fossil species could be explained if every new species always came into existence near an already existing, closely related species. 1476:

voyage. He tentatively wrote of his ideas to Lyell in January 1842; then in June he roughed out a 35-page "Pencil Sketch" of his theory. Darwin began correspondence about his theorising with the botanist

3407:

By the mid-1870s, most scientists accepted evolution, but relegated natural selection to a minor role as they believed evolution was purposeful and progressive. The range of evolutionary theories during

3452:(1872), attempted to extend the idea of natural selection to competition between nations and between human races. Such ideas were incorporated into what was already an ongoing effort by some working in 3379:'s insistence that a trivial characteristic with no adaptive advantage could not be developed by selection. Darwin conceded that these could be linked to adaptive characteristics. His estimate that the 2392:

in the 1930s and 1940s that a model of heredity became completely integrated with a model of variation. This modern evolutionary synthesis had been dubbed Neo Darwinian Evolution because it encompasses

862:. There was growing support for such ideas among dissident anatomists and the general public, but during the first half of the 19th century the English scientific establishment was closely tied to the 3741:

was voted the most influential academic book ever written. It was hailed as "the supreme demonstration of why academic books matter" and "a book which has changed the way we think about everything".

3420:

claiming that species had an inherent tendency to change in a particular direction, and forms of neo-Lamarckism in which inheritance of acquired characteristics led to progress. The minority view of

2671:. Darwin discusses rudimentary organs, such as the wings of flightless birds and the rudiments of pelvis and leg bones found in some snakes. He remarks that some rudimentary organs, such as teeth in 2339:; in Chapter I Darwin admitted, "The laws governing inheritance are quite unknown." He accepted a version of the inheritance of acquired characteristics (which after Darwin's death came to be called 3062:, in which investigation could begin with the intuitive idea that species were fixed objects created by design. Early support for Darwin's ideas came from the findings of field naturalists studying 3444:, argued that action was needed to correct social and economic inequities to level the playing field before natural selection could improve humanity further. Some political commentaries, including 7096:(February to July 1838) pp. 196–197 "Man in his arrogance thinks himself a great work worthy the interposition of a deity, more humble & I believe truer to consider him created from animals." 1791:

felt it excused the lack of references, but wanted to keep "natural selection" which was "constantly used in all works on Breeding", and hoped "to retain it with Explanation, somewhat as thus",—

3734:

and the bicentenary of Darwin's birth were scheduled for 2009. They celebrated the ideas which "over the last 150 years have revolutionised our understanding of nature and our place within it".

2427:

tail of a giraffe, which serves as a fly-flapper, and, on the other hand, organs of such wonderful structure, as the eye, of which we hardly as yet fully understand the inimitable perfection?

2128:

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.

3222:, thought that humans shared a common ancestor with apes, but higher mental faculties could not have evolved through a purely material process. Darwin published his own explanation in the 2227:

of each species, that archetype being a fixed ideal in the mind of God. Darwin and Wallace made variation among individuals of the same species central to understanding the natural world.

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are dense, but other parts are almost lyrical, and the case studies and observations are presented in a narrative style unusual in serious scientific books, which broadened its audience.

2729:

many fields of natural history that were inexplicable under the alternative concept that species had been individually created. The structure of Darwin's argument showed the influence of

2568:, posed a problem. Darwin had no doubt that earlier seas had swarmed with living creatures, but stated that he had no satisfactory explanation for the lack of fossils. Fossil evidence of 1361:

were friendly and civilised, yet to Darwin their relatives on the island seemed "miserable, degraded savages", and he no longer saw an unbridgeable gap between humans and animals. As the

3555:, it had dropped a number of hints about human's animal ancestry and quickly became central to the debate, as mental and moral qualities were seen as spiritual aspects of the immaterial 1172:. Lamarck thought there was an inherent progressive tendency driving organisms continuously towards greater complexity, in parallel but separate lineages with no perceptible extinction. 1775:, and met with him to find if he would be willing to publish. On 28 March Darwin wrote to Lyell asking about progress, and offering to give Murray assurances "that my Book is not more 2479:

created instincts, but as small consequences of one general law, leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.

1585:) on ways that plant seeds and animals might disperse across oceans to colonise distant islands. By 1856, his theory was much more sophisticated, with a mass of supporting evidence. 1534:

Hooker was persuaded to take away a copy of the "Essay" in January 1847, and eventually sent a page of notes giving Darwin much-needed feedback. Reminded of his lack of expertise in

9761: 5710: 3476:

The book produced a wide range of religious responses at a time of changing ideas and increasing secularisation. The issues raised were complex and there was a large middle ground.

3120:. Despite this, Huxley strongly supported Darwin on evolution; though he called for experiments to show whether natural selection could form new species, and questioned if Darwin's 2857:. He added notes on sexual selection to his "big book on species", and in mid-1857 he added a section heading "Theory applied to Races of Man", but did not add text on this topic. 2510:

could not have evolved through natural selection because intermediate stages would not have been adaptive. Darwin proposed scenarios for the incremental evolution of each feature.

983:

This slowly effected process results in populations changing to adapt to their environments, and ultimately, these variations accumulate over time to form new species (inference).

2007:. In 1862, Bronn produced a second edition based on the third English edition and Darwin's suggested additions, but then died of a heart attack. Darwin corresponded closely with 1451:

Darwin now had the basic framework of his theory of natural selection, but he was fully occupied with his career as a geologist and held back from compiling it until his book on

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Journal of Researches into the Natural History and Geology of the Countries Visited During the Voyage of HMS Beagle Round the World, Under the Command of Captain Fitz Roy, R.N.

3187:

plan in a continuous "ordained becoming", with new species appearing by natural birth. Others that rejected natural selection, but supported "creation by birth", included the

3142:

over natural theology, praising Darwin for "extending the domination of Science over regions of thought into which she has, as yet, hardly penetrated" and coining the term "

2181:

environmental change is necessary to generate variation. The opening two sentences of On The Origin demonstrate this point and also show that Darwin did not see the role of

10498: 10482: 10081:, the 6th edition with additions and corrections (final text), the first American edition, and translations into Danish, Dutch, French, German, Polish, Russian and Spanish 7255:

one notion of beauty & negroes another" from Darwin, C. R. Notebook M : . CUL-DAR125.- Transcribed by Kees Rookmaaker, edited by Paul Barrett. (Darwin Online, p.

6521:, Quote: "… I gave, however, a tolerably clear sketch of this principle in the first edition of the 'Origin of Species,' and I there stated that it was applicable to man." 2289:. He did not suggest that every favourable variation must be selected, nor that the favoured animals were better or higher, but merely more adapted to their surroundings. 1806:

to review the chapters. At Lyell's suggestion, Elwin recommended that, rather than "put forth the theory without the evidence", the book should focus on observations upon

1399:. Darwin began speculating, in a series of notebooks, on the possibility that "one species does change into another" to explain these findings, and around July sketched a 2132:

Starting with the third edition, Darwin prefaced the introduction with a sketch of the historical development of evolutionary ideas. In that sketch he acknowledged that

2371:, offspring inheriting characteristics of a previous mate of the female parent, and accepted this process as increasing the variation available for natural selection. 10052: 1557:

Darwin's barnacle studies convinced him that variation arose constantly and not just in response to changed circumstances. In 1854, he completed the last part of his

5531: 2941:, which had been rejected by scientists, but had influenced a wide public readership into believing that nature and human society were governed by natural laws. The 2488:

have been by-products of adaptive changes to other features, and that often features seemed non-adaptive because their function was unknown, as shown by his book on

2136:

had, unknown to Wallace or himself, anticipated the concept of natural selection in an appendix to a book published in 1831; in the fourth edition he mentioned that

1722:, and focused on ecological pressures that kept different varieties adapted to local conditions. Some historians have suggested that Wallace was actually discussing 3440:

While some, like Spencer, used analogy from natural selection as an argument against government intervention in the economy to benefit the poor, others, including

3214:

as the "great hippopotamus test", Huxley showed that Owen was incorrect in asserting that ape brains lacked a structure present in human brains. Others, including

2586:

were unchanged since the earliest fossils. The pace of natural selection would depend on variability and change in the environment. This distanced his theory from

10042: 1481:

in January 1844, and by July had rounded out his "sketch" into a 230-page "Essay", to be expanded with his research results and published if he died prematurely.

11673: 10425: 2745:, where if you could argue that a proposed mechanism successfully explained various phenomena you could then use those arguments as evidence for that mechanism. 2376: 465: 5504: 10562: 2235:

In Chapter III, Darwin asks how varieties "which I have called incipient species" become distinct species, and in answer introduces the key concept he calls "

2207:

published in 1831, future editions of On the Origin of Species contained these first two sentences unchanged and continued to omit the correct explanation of

8367: 2160:. Darwin discusses contemporary opinions on the origins of different breeds under cultivation to argue that many have been produced from common ancestors by 2312:

features such as lion manes, deer antlers, peacock tails, bird songs, and the bright plumage of some male birds. He analysed sexual selection more fully in

10442: 8886: 3801: 2875: 2314: 6851:, Quote: "… this implies that man must be included with other organic beings in any general conclusion respecting his manner of appearance on this earth." 2540:

is a very rare occurrence, spread over vast periods of time; since few areas had been geologically explored, there could only be fragmentary knowledge of

8735:

The Autobiography of Charles Darwin 1809–1882. With the Original Omissions Restored. Edited and with Appendix and Notes by his Granddaughter Nora Barlow

2272:

interdependencies, and notes that competition is most severe between closely related forms "which fill nearly the same place in the economy of nature".

1128:

suggested that some similar species, such as horses and asses, or lions, tigers, and leopards, might be varieties descended from a common ancestor. The

3193: 2688:

with the sentence that "Light will be thrown on the origin of man and his history." Darwin ends with a passage that became well known and much quoted:

1523:

had significant influence on public opinion, and the intense debate helped to pave the way for the acceptance of the more scientifically sophisticated

1371:

showed that fossils of extinct species Darwin found in South America were allied to living species on the same continent. In March 1837, ornithologist

1108:

showed minor differences caused by local conditions. In God's benevolent design, carnivores caused mercifully swift death, but the suffering caused by

6317:

McBride, P. D., Gillman, L. N., & Wright, S. D. (2009). Current debates on the origin of species. Journal of Biological Education, 43(3), 104–107.

3399:

calculated that as it mixed traits, natural selection could not accumulate useful traits. Darwin tried to meet these objections in the fifth edition.

12902: 10450: 9305: 5725: 1236:

were influenced by Lamarck and Geoffroy, but most naturalists regarded their ideas of transmutation as a threat to divinely appointed social order.

9041:

Six Lectures to Working Men "On Our Knowledge of the Causes of the Phenomena of Organic Nature" (Republished in Volume II of his Collected Essays,

7972: 5558: 4653: 5702:

Het ontstaan der soorten van dieren en planten door middel van de natuurkeus of het bewaard blijven van bevoorregte rassen in de strijd des levens

2642:, a multilevel system of groups and sub-groups based on varying degrees of resemblance. After discussing classification issues, Darwin concludes: 12296: 10386: 10047: 8187: 3175:, an idealist who had shifted to the view in the 1850s that the history of life was the gradual unfolding of a divine plan. Owen's review of the 8600: 3326:, who viewed every species as a distinct fixed unit in the mind of the Creator, classifying as species what others considered merely varieties. 1997:

translators were expected to make their own arrangements with a local publisher. He welcomed the distinguished elderly naturalist and geologist

10466: 3210: 1605:

after five years, had enlarged these into a sketch in 1844, and "from that period to the present day I have steadily pursued the same object."

780: 13025: 10370: 4871: 1453: 11422: 3737:

In a survey conducted by a group of academic booksellers, publishers and librarians in advance of Academic Book Week in the United Kingdom,

3503:

defended evolutionary ideas by arguing that the introduction of new species should be considered a natural rather than a miraculous process.

12661: 10722: 4993: 2015:

published in 1862 added an introduction praising Darwin's ideas as an alternative to religious revelation and promoting ideas anticipating

1472:

Darwin continued to research and extensively revise his theory while focusing on his main work of publishing the scientific results of the

388: 10141: 8304:

is A02-2009-00000001-4. As a tribute to its influence, this work has been the first one to be registered by The International ISTC Agency.

1192:

strongly disputed such ideas, holding that unrelated, fixed species showed similarities that reflected a design for functional needs. His

12518: 10062: 9641: 9268: 8228: 3795: 2717: 1502: 1124:

in 1735 also viewed species as fixed according to the divine plan, but did recognize the hierarchical nature of different taxa. In 1766,

93: 2031:

was published in Swedish in 1871, Danish in 1872, Polish in 1873, Hungarian in 1873–1874, Spanish in 1877 and Serbian in 1878. By 1977,

1734:

Soon after the meeting, Darwin decided to write "an abstract of my whole work" in the form of one or more papers to be published by the

9327: 4834:

See letter to T. C. Eyton, 4 October (1858), in which CD first mentioned the possibility that his 'abstract' would form a small volume.

4784: 3308:, which embodied a progressive, almost linear model of evolution. Darwin was cautious about such histories, and had already noted that 2469:

Chapter VII (of the first edition) addresses the evolution of instincts. His examples included two he had investigated experimentally:

2011:, who published an improved translation in 1867. Darwin's attempts to find a translator in France fell through, and the translation by 699: 9403:(June 1987), "Darwinian Gradualism and Its Limits: The development of Darwin's Views on the Rate and Pattern of Evolutionary Change", 9167: 8751: 8549: 7945: 5353: 5042: 1988:. Darwin was delighted by the popularity of the book, and asked Gray to keep any profits. Gray managed to negotiate a 5% royalty with 1621:

has suggested all these factors may have contributed, and notes Darwin's large output of books and busy family life during that time.

12932: 12674: 11987: 10597: 10570: 10304: 9407:, Humanities, Social Sciences and Law, vol. 20, no. 2, Springer Netherlands (published 6 November 2004), pp. 139–157, 7434: 7093: 5659: 5632: 4297: 2715:

had been the chief agent of change. He knew that his readers were already familiar with the concept of transmutation of species from

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opposed evolution in the late 19th and early 20th century, but other Catholic writers, starting with Mivart, pointed out that early

1771:

By mid-March 1859 Darwin's abstract had reached the stage where he was thinking of early publication; Lyell suggested the publisher

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Victorian Sensation: The Extraordinary Publication, Reception, and Secret Authorship of Vestiges of the Natural History of Creation

4233: 3240:

Haeckel showed a main trunk leading to mankind with minor branches to various animals, unlike Darwin's branching evolutionary tree.

2787:

as part of the natural processes he was investigating, and rejected divine intervention. In 1856, his "big book on species" titled

2039:

who added non-Darwinian ideas; he published the preliminaries and chapters 1–5 in 1902–1904, and his complete translation in 1920.

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had admitted, but the "explanation is manifest on the theory of the natural selection of successive slight modifications" showing

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On 5 April, Darwin sent Murray the first three chapters, and a proposal for the book's title. An early draft title page suggests

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The 6th is Darwin's final edition; there were minor modifications in the text of certain subsequent issues. See Freeman, R. B. "

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Darwin's early notebooks discussed how non-adaptive characteristics could be selected when animals or humans chose mates, with

2099:'s letter suggesting that the origin of species "would be found to be a natural in contradistinction to a miraculous process": 213: 7379: 7353: 7336: 7299: 6848: 5476: 5466: 5286: 4513: 4486: 4374: 1948:

clubbing together to buy the fifth edition at 15 shillings and wanted it made more widely available; the price was halved to 7

1419:

as well as established scientists. At the zoo he had his first sight of an ape, and was profoundly impressed by how human the

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aspects were felt to be relatively unimportant. The political economy of struggle was criticised as a British stereotype by

1711:

On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection

139:

On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection

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in 1860 where he disagreed with Darwin's 'argument'. There was much less controversy than had greeted the 1844 publication

2822:

describes it as his only discussion there of human origins, while noting that the book makes other references to humanity.

641: 9977: 9950: 9922: 9897: 9807: 9016: 8568: 7452: 5759: 5537: 5452: 4568: 3730:, an exhaustive external index published in 1981. Worldwide commemorations of the 150th anniversary of the publication of 2172:

breeding, noting that "he diversity of the breeds is something astonishing", yet all were descended from one species, the

907:

through natural selection became central to modern evolutionary theory, and it has now become the unifying concept of the

12396: 11772: 10692: 10629: 10506: 3188: 899:" from the 1880s to the 1930s, various other mechanisms of evolution were given more credit. With the development of the 9192: 4422: 1184:

when the environment acted on embryos, and that animal structures were determined by a constant plan as demonstrated by

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On the Origin of Species, by means of Natural Selection; or the Preservation of Favoured Races in the Struggle for Life

8928: 3883: 3750: 3708: 3679:. The theory explains the diversity of living organisms and their adaptation to the environment. It makes sense of the 3578: 3036: 2794:

28 March 1859, with his manuscript for the book well under way, Darwin wrote to Lyell offering the suggested publisher

2083:'s belief in a rational God who established a law-abiding cosmos. In the second edition, Darwin added an epigraph from 773: 502: 333: 20: 8663: