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Login to your account Email Password Forgot password? Keep me logged in New User Institutional Login Change Password Old Password New Password Too Short Weak Medium Strong Very Strong Too Long YOUR PASSWORD MUST HAVE 8 CHARACTERS OR MORE AND CONTAIN 3 OF THE FOLLOWING: * a lower case character, * an upper case character, * a special character * or a digit Too Short Congrats! Your password has been changed Create a new account Email Returning user Can't sign in? Forgot your password? Enter your email address below and we will send you the reset instructions Email Cancel If the address matches an existing account you will receive an email with instructions to reset your password. Close Request Username Can't sign in? Forgot your username? Enter your email address below and we will send you your username Email Close If the address matches an existing account you will receive an email with instructions to retrieve your username Anywhere * Anywhere * This Journal Quick Search anywhere Enter words, phrases, DOI, keywords, authors, etc... SearchGo Quick Search in Journals Enter words, phrases, DOI, keywords, authors, etc... SearchGo Advanced Search Skip main navigationJournal menuClose Drawer MenuOpen Drawer Menu Home * All Journals * Biographical Memoirs * Biology Letters * Interface * Interface Focus * Notes and Records * Open Biology * Philosophical Transactions A * Philosophical Transactions B * Proceedings A * Proceedings B * Royal Society Open Science * * * * * * * * Sign in Institutional Access * 0 Cart * Search Skip main navigationJournal menuClose Drawer MenuOpen Drawer Menu Home * Home * Content * Latest articles * Latest issue * All content * Subject collections * Blog posts * Information for * Authors * Guest organizers * Reviewers * Readers * Institutions * About us * About the journal * Editorial board * Author benefits * Policies * Journal metrics * Open access * Sign up * Purchase * eTOC alerts * RSS feeds * Newsletters * Request a free trial * Submit You have access MoreSections * Figures * Related * References * Details View PDF View PDF Tools * Add to favorites * Download Citations * Track Citations Share Share on * Facebook * Twitter * Linked In * Reddit * Email Cite this article * Tanghe Koen B. 2019On The Origin of Species: The story of Darwin's titleNotes Rec.7383–100http://doi.org/10.1098/rsnr.2018.0015 SECTION * Abstract * A lacuna in the Darwin literature * Natural selection * The original title * The influence of John Murray on the modification of the original title * Four other modifications * Conclusion * Acknowledgements * Footnotes You have accessResearch articles ON THE ORIGIN OF SPECIES: THE STORY OF DARWIN'S TITLE Koen B. Tanghe Koen B. Tanghe Blandijnberg 2, Department of Philosophy and Moral Sciences, 9000 Gent, Belgium kbt.ugent@gmail.com Google Scholar Find this author on PubMed Search for more papers by this author Koen B. Tanghe Koen B. Tanghe Blandijnberg 2, Department of Philosophy and Moral Sciences, 9000 Gent, Belgium kbt.ugent@gmail.com Google Scholar Find this author on PubMed Search for more papers by this author Published:22 August 2018https://doi.org/10.1098/rsnr.2018.0015 ABSTRACT The genesis of Charles Darwin's book On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (1859) is well known, and the changes that it underwent in subsequent editions are well documented. However, less is known or has been published about the genesis of its original title and about the seven modifications that it subsequently underwent. That original title was much longer than the title of the unfinished big ‘Species Book’ that preceded and inspired The Origin: Natural Selection. Why did Darwin use an extended version of this elegant, short title for The Origin? And what was the rationale behind the later modifications? Contrary to what is often claimed or implied, the criticism of his publisher, John Murray, does not offer the only and certainly not the full answer to the latter question. Anyhow, you have a capital title, and some think this the most difficult part of a book. —Charles Darwin, Letter of 5 November 1860.1 A LACUNA IN THE DARWIN LITERATURE Since the emergence of a Darwin industry, in the wake of the 1959 commemoration of the centennial of the publication of On the Origin of Species, analyses and reviews of this flourishing field of study have been published on a regular basis.2 However, whereas there exist excellent online resources about Darwin's work, an encyclopaedic and systematic online overview of the huge literature that the Darwin industry has produced and still is producing on Darwin's life, background, work and influence is sorely lacking.3 Such an overview could not only greatly help scholars in their research of classic topics in the study of Darwin, but might also allow them to identify topics that have not yet received much systematic attention, such as the ‘Historical sketch’ that was added to the third English edition of The Origin (1861).4 As Johnson points out, ‘Somewhat surprisingly, little systematic attention has been paid to the Historical Sketch in the literature, as far as I can tell.’5 Even the most careful biographies ‘say little about the genesis and various transformations of and motivations for the [Historical] Sketch’.6 Much the same can be said about the title of The Origin: even the most careful biographies do not give a complete account of its convoluted genesis. Desmond and Moore merely point out that Darwin's publisher, John Murray, being a practical man, was ‘more concerned with the title’ (than with the orthodoxy of Darwin's theory).7 One page later, they add that Darwin's title ‘continued to evolve under Murray's selective pressure. It had slimmed down to On the Origin of Species and Varieties by Means of Natural Selection, when Darwin improved matters more by docking “and Varieties”’.8 Janet Browne's account is longer but, as will be explained below, inaccurate: At the last minute [Darwin] adjusted the title according to Murray's recommendation. Darwin's first suggestion was rather too complicated: ‘An Abstract of an Essay on the Origin of Species and Varieties Through Natural Selection.’ Common sense surely suggested to Murray that the words ‘abstract,’ ‘essay,’ and ‘varieties’ should go, and that ‘natural selection,’ a term with which Murray thought the public would not be familiar, ought to be explained. The agreed-upon title was, however, hardly less cumbersome—On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life.9 Both accounts overestimate the role of Murray in the long genesis of the title of Darwin's book and leave much undiscussed. I will start my own analysis with the title of the never-finished manuscript that Darwin wrote before he started writing The Origin: Natural Selection.10 NATURAL SELECTION Francis Darwin called the two sketches of his father's theory of evolution of the early 1840s (1842 and 1844) ‘essays’.11 However, that is not how Darwin himself referred to them. The first manuscript was a rough pencil sketch, but even the second, much longer version was still characterized as a ‘written sketch of species theory’.12 Likewise, in a letter to his wife, Emma, he referred to ‘my sketch of my species theory’.13 Interestingly, a letter from 9 April 1843 from Charles Lyell was, on the other hand, already annotated with the remark ‘Species Book’.14 As is well known, Lyell was later instrumental in the publication of that Species Book (i.e. The Origin). On 1 May 1856, he suggested that Darwin should ‘publish some small fragment of [his] data pigeons if you please & so out with the theory & let it take date—& be cited—& understood’.15 For he feared that Darwin might be forestalled.16 The ever-vacillating Darwin half-heartedly agreed―he had been sorting his species notes since September 1854―but hesitated about the format in which to publish his theory.17 He was ‘fixed against any periodical or Journal, as I positively will not expose myself to an Editor or Council allowing a publication for which they might be abused’.18 He initially rather thought of ‘a very thin & little volume, giving a sketch of my views & difficulties’, although it was ‘really dreadfully unphilosophical to give a resumé, without exact references, of an unpublished work. But Lyell seemed to think I might do this …’.19 He was ‘extremely glad’ that his good friend Joseph Hooker gave his blessing to such a separate ‘Preliminary Essay … for Lyell seemed rather to doubt on this head’.20 On 14 May 1856 he began working on this new ‘species sketch’, although he was initially not even sure that he would publish it.21 Five months later, Darwin wrote to his second cousin W. D. Fox that when he began writing this sketch or essay, he had found it such unsatisfactory work that he had desisted and instead was ‘now drawing up [his] work as perfect as [his] materials of 19 years collecting suffice’, adding that, to his sorrow, ‘it will run to quite a big Book’.22 We do not know when exactly he chose a title for that ‘big Book’, nor, with certainty, why he chose the phrase ‘natural selection’. In a letter to Charles Lyell from 10 November 1856, he still called it his ‘big Book’.23 Ten months later, he articulated its title in an abstract of his ideas, enclosed in a letter to Asa Gray: Natural Selection.24 The essays or sketches from 1842 and 1844 were both formally divided into two parts, which differed both in subject and in length.25 The first and shorter part (three chapters) consisted of an elaboration of the analogy between artificial and natural selection. In the second and by far the longer part (seven chapters, conclusion included), Darwin mustered the evidence for his doctrine of common descent by reinterpreting various biological disciplines (palaeontology, geographical distribution, classification, morphology and embryology) in evolutionary terms. This binary structure of his argumentation reflected, on the one hand, the importance he attached to this reinterpretation and, on the other hand, the relative unimportance of his theory of natural selection for his ‘one long argument’ (in favour of transmutation). As he declared in a letter to Asa Gray from 11 May 1863: ‘Personally, of course, I care much about Natural Selection; but that seems to me utterly unimportant compared to question of Creation or Modification’.26 In a letter from 20 September 1859, he even begged Charles Lyell to keep his mind open about his theory ‘till you receive (in perhaps a fortnight's time) my latter chapters which are the most important of all on the favourable side’.27 This unequal division of his almost fully developed theory of evolution was also a logical reflection of its gestation: Darwin had first become an evolutionist and had only later developed his theory of natural selection. It was, of course, also the all-important second part of his theory that would, in the nineteenth century, prove most influential: it was largely thanks to this part that Darwin converted a majority of his learned contemporaries to evolution. Natural selection, by contrast, was, in the nineteenth century and the first decades of the twentieth century, not generally accepted as an important evolutionary mechanism or process. The binary division of the Sketch and the Essay was implicitly preserved in the ‘big Book’ that he began writing in May 1856 and, later, in The Origin.28 It is somewhat puzzling, therefore, that the former manuscript was entitled Natural Selection. Why did Darwin call it after its least important part (i.e. the part about natural selection)? It seems that it had gained somewhat in importance, since Darwin's notes for Natural Selection suggest that the second part would only have been as long as the first part.29 Still, his later remarks indicate that his transmutational reinterpretation of mid-nineteenth-century static natural history or ‘biology’ remained the main part of his argument. The question therefore remains: why did he call his big ‘Species Book’ after the least important part of his argument? Part of the answer undoubtedly is that ‘he care[d] much about Natural Selection’. Also, since 1839, natural selection had been ‘the enduring core element’ of his evolutionary theorizing.30 Lyell may also have been a motivating factor behind Darwin's choice. After Darwin had explained to Lyell his theory of evolution and had shown him his pigeon breeds, during a visit in April 1856, Lyell noted in his scientific journal: ‘With Darwin: On the Formation of Species by Natural Selection’.31 He realized that what he called the natural selection theory explained the pattern, observed by Alfred Russel Wallace, that new species are most allied to those immediately preceding in time, and used ‘natural selection as a shorthand for Darwin's theory of evolution’.32 When the writing of Natural Selection was interrupted, in June 1858, by the arrival of a package from Ternate that contained Alfred Russel Wallace's rudimentary version of his own theory, Darwin panicked. Lyell and Hooker hastily arranged a presentation of the Darwin–Wallace theory at a meeting of the Linnean Society. The paper was published in the Journal of the Proceedings of the Linnean Society on 20 August 1858.33 By then, Darwin was already hard at work at an abstract of his big ‘Species Book’. Hooker had suggested a new article, of 30 pages or so, for the Linnean Society Journal. In a letter to T. C. Eyton from 4 August 1858, Darwin spoke of ‘a long abstract on my notions about Species & Varieties, to be read in parts before Linnean Socy’ that would be published ‘late in the autumn’.34 History repeated itself though: the journal article became a book, in the same way that, in 1856, his ‘preliminary Essay’ had soon turned into ‘quite a big Book’. On 12 October 1858, Darwin already expected that his abstract would ‘run into a small volume, which will have to be published separately’.35 Its full title was enclosed in a letter that he sent to Charles Lyell on 28 March 1859: ‘An Abstract of an Essay on the Origin of Species and Varieties Through Natural Selection’ (figure 1).36 Figure 1. The original title of On the Origin of Species. This proposed title page for what eventually became The Origin of Species was enclosed in a letter that Darwin sent to Charles Lyell on 28 March 1859. The text reads: ‘An abstract of an Essay on the Origin of Species and Varieties Through Natural Selection by Charles Darwin M. A Fellow of the Royal, Geological & Linn. Socy. London & & & & 1859’. (From John van Wyhe (ed.), The complete work of Charles Darwin online, 2002, http://darwin-online.org.uk.) * Download figure * Open in new tab * Download PowerPoint THE ORIGINAL TITLE One striking and paradoxical characteristic of this original title is that it was much longer than the title of the longer but unfinished manuscript that preceded The Origin. It included two new phrases: ‘abstract of an essay’ and ‘on the origin of species and varieties’. Darwin clearly wanted to present explicitly The Origin as an abstract.37 Indeed, for a surprisingly long time after the publication of the first edition of The Origin, he harboured the intention of publishing a longer version of his ‘abstract of an essay’. It would have hardly a page in common, & might bear a quite distinct title; & I shd like to produce that volume by volume (perhaps 3 thinnish octavos) as I have it ready.— Much of the M.S is roughly ready, but I daresay it would take me two years to prepare 1st vol.—.38 On 25 December 1859, he wrote about this project to W. D. Fox: ‘I am going soon to begin my bigger book, which I shall publish as 3 separate volumes, with distinct titles, but with a general title in addition’.39 This tripartite scheme represented, as Hodge puts it, ‘a convenient division of Darwin's personal labour, rather than a natural articulation in his public argument’.40 That articulation was, as we saw, binary. The rationale for the term ‘abstract’ is, consequently, clear: the socially savvy Darwin wanted to emphasize that The Origin was incomplete and imperfect. The term ‘essay’, by contrast, is more problematic. It almost certainly refers to the unfinished Natural Selection manuscript, since The Origin was clearly inspired by it. This, as a matter of fact, explains why the all-important second part of his argument had, in The Origin, shrunk even further in relative size (versus Natural Selection): it now occupied only about one-third of the entire book (chapters IX–XIII). The reason is that, when the writing of Natural Selection was interrupted by the arrival of Wallace's package, Darwin had not yet begun writing the second part of his book, apart from a section on geographical distribution. His ‘Abstract of an Essay’ was, consequently, an ‘Abstract of an Unfinished Manuscript’. Put differently: ‘essay’ was a misnomer. The Origin could maybe be called an essay (i.e. a short and non-technical book); Natural Selection could not. Another question is why Darwin did not simply choose the title ‘Abstract of an Essay on Natural Selection’. Why did he replace the phrase ‘natural selection’ with the longer phrase ‘(an essay) on the origin of species and varieties through natural selection’? This is, without a doubt, the main and most intriguing change in the trajectory between Natural Selection and The Origin of Species (1872). It confronts us with two questions: why did Darwin decide to refer to the question of the origin of organic beings and why did he first refer to the origin of both species and varieties? One of the possible reasons why he chose a longer and more explanatory title is that the shorter and less scholarly The Origin targeted a broader public than Natural Selection. He may also have been inspired by the title of a previous publication that is structurally reminiscent of ‘On the Origin of Species and Varieties through Natural Selection’. As he pointed out, in a letter of 11 May 1856 to Hooker, it was not the first time that Lyell had urgently advised him to publish a preliminary sketch.41 On 31 May 1837, Darwin's coral theory was first presented at a meeting of the Geological Society of London under the title ‘On certain areas of elevation and subsidence in the Pacific and Indian Oceans, as deduced from the study of coral formations’ and, later that year, published in the Proceedings of the Geological Society of London.42 Lyell's influence may even have been more direct. As pointed out above, in his journal he used the phrase ‘On the Formation of Species by Natural Selection’ to refer to Darwin's theory. Lastly, the title of the aforementioned Darwin–Wallace joint paper may also have been an inspiration: ‘On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection’. This title can maybe also help explain why Darwin referred, in the original title of his abstract, to the origin of both species and varieties.43 His theory had, of course, always been intimately connected to the question of the taxonomic difference between species and varieties. The reason is simple: varieties could only be ‘incipient species’ if there was no clear distinction between varieties and species. The fact that such a distinction could not easily be made was, from the beginning, a very important element in Darwin's argument against the fixity of species, as can be seen from a letter that he wrote to Henry Denny on 7 November 1844: ‘I am deeply interested in everything connected with geographical distribution, & the differences between species and varieties’.44 He sometimes even referred to The Origin as his ‘book on species & varieties’.45 THE INFLUENCE OF JOHN MURRAY ON THE MODIFICATION OF THE ORIGINAL TITLE Darwin's original title underwent seven modifications (figure 2). One word was replaced (‘through’ became ‘by means of’) and four terms or phrases were deleted (‘an abstract of’, ‘an essay’, ‘and varieties’ and ‘on’). Lastly, Darwin added a subtitle. The first version read ‘the preservation of favoured races’, the second ‘the preservation of favoured races in the struggle for life’. Only three of these modifications were inspired by a (known) criticism from John Murray, although it must be added that these were the three most important modifications. Figure 2. The seven modifications of the original title of On the Origin of Species, with the earliest known date of the modification and the source. Additions are in bold, deletions are struck through, replacements are underlined. The words and phrases between brackets were added or deleted, or replaced other words, at some unknown point between the end of March 1859 and 24 November 1859 (the publication date of the book). There are, to the best of my knowledge, no known sources for the reason(s) why Darwin made modifications 4, 5, 6 and 7. * Download figure * Open in new tab * Download PowerPoint At the beginning of March 1859, Darwin had asked Lyell whether John Murray might be interested in publishing his abstract. Murray had not only published Lyell's books but had also issued the second edition of Darwin's successful Journal of Researches.46 Lyell thought this ‘an excellent idea’ and payed Murray ‘one of his most persuasive social calls’.47 In a subsequent letter to Lyell (28 March), Darwin asked his friend whether he had already spoken to Murray (he fancied that he had done so from an ‘expression in Lady Lyell's note’).48 He apparently soon received an affirmative answer, for in two letters—one to Lyell (30 March) and one to Murray (31 March)—he expressed his delight that Murray had agreed to publish his ‘work on the Origin of Species’, without even having read the manuscript.49 However, Murray was less enthusiastic about Darwin's title. The letter that Darwin sent to Lyell on 30 March shows that Murray did indeed object to the term ‘abstract’, as pointed out by Browne, but he did not object to the words ‘essay’ and ‘varieties’.50 Also, his objection to the phrase ‘natural selection’ was, in contrast with what Browne suggests, not constructive: he did not suggest that Darwin explained the term but simply objected to it. That may be the reason why Darwin, at a certain point in time, toyed with the idea of a completely different title, ‘On the Mutability of Species’, as suggested by a tentative title-page sketch (figure 3).51 Figure 3. A tentative sketch of the title page of On the Origin of Species, with an alternative title and additional notes. The text reads: ‘On The Mutability of Species “Whewell” by C. Darwin, M. A, F.R.S. John Murray. 1860!!’ It is not clear why Darwin emphasized the year 1860 (see, in this respect, note 51). (From John van Wyhe (ed.), The complete work of Charles Darwin online, 2002, http://darwin-online.org.uk.) * Download figure * Open in new tab * Download PowerPoint Darwin agreed to delete the term ‘abstract’ but he hoped to retain the phrase ‘natural selection’, ‘with Explanation, somewhat as thus,—Through Natural Selection or the preservation of favoured Races’.52 The reason why he thought that the phrase ‘natural selection’ was not problematic is that it was ‘constantly used in all works on Breeding, & I am surprised that it is not familiar to Murray’.53 Both statements make it clear that his full subtitle, ‘the Preservation of Favoured Races in the Struggle for Life’, was not conceived and intended as an alternative for the main title ‘On the Origin of Species by Means of Natural Selection’, as Richard Dawkins claims, but as an explanation of the, at the time, relatively novel phrase ‘natural selection’ (although it did have implications for the meaning of the phrase ‘origin of species’, as will become clear below).54 The explanation, through a subtitle, of the phrase ‘natural selection’ was also, without a doubt, the most important and most remarkable modification of the original title of Darwin's abstract. I will first concentrate upon the specific words of which it is composed and subsequently discuss the possible reason why Darwin chose this particular definition of natural selection. ‘Preservation’ was Darwin's standard term: he used it at least 20 times in the first edition of The Origin, whereas the term ‘survival’ was used only once (likewise, the term ‘preserved’ appears more than 10 times more often than the term ‘survived’).55 The term ‘favoured’ was a synonym of—and sometimes used in combination with—‘selected’. In his chapter on natural selection, for example, Darwin spoke of plants that ‘would be continually favoured or selected’.56 In his chapter on ‘difficulties on theory’, he wrote: On the absence or rarity of transitional varieties.—As natural selection acts solely by the preservation of profitable modifications, each new form will tend in a fully-stocked country to take the place of, and finally to exterminate, its own less improved parent or other less-favoured forms with which it comes into competition.57 The term ‘race’ was a synonym for ‘variety’, ‘breed’ or ‘form’. Surprisingly, it (or the plural ‘races’) was used much less frequently in the text of The Origin (in this particular meaning) than ‘variety’ (or ‘varieties’), ‘breed(s)’ or ‘form(s)’. Furthermore, ‘race’ had anthropological connotations, whereas Darwin avoided, in The Origin, the subject of humans, ‘as so surrounded with prejudices’.58 The question of human races was even ‘blowing up as an emotive issue in the 1850s’.59 So why did Darwin choose this particular, not unproblematic word for his subtitle instead of a more neutral and, in the text of The Origin, more common alternative such as ‘variety’?60 One possible explanation is that he wanted to make the link with the artificial selection of races of domesticated plants and animals (for a second possible explanation, see below). Indeed, in the text of The Origin, he often used the phrase ‘domestic race’. Also, the second part of his Essay of 1844 was headed ‘On the evidence favourable and opposed to the view that species are naturally formed races, descended from common stocks’ (my italics). Likewise, in his Sketch of 1842, he promised that a discussion of ‘whether the characters and relations of animated beings are such as favour the idea of wild species being races descended from a common stock’ would form ‘the second part of this sketch’.61 It is not known when exactly Darwin added the phrase ‘in the Struggle for Life’ to his subtitle. It was may be already inserted in the title that he sent to Murray, ‘(with some remarks on separate page)’, enclosed in a letter from 5 April 1859 (unfortunately, neither the title nor the remarks seem to have been preserved).62 It is quite clear, though, why he added it: it was through the struggle for life of, and within, geometrically increasing populations that the selective actions of breeders or ‘the preservation of their favoured races’ was mimicked by nature: In the preservation of favoured individuals and races, during the constantly-recurrent Struggle for Existence, we see the most powerful and ever-acting means of selection. The struggle for existence inevitably follows from the high geometrical ratio of increase which is common to all organic beings.63 Let us now proceed to the more important and intriguing question as to why Darwin chose, in his subtitle, this specific definition of natural selection: ‘the Preservation of Favoured Races in the Struggle for Life’. The natural process that he tried to convey through his analogy with artificial selection is quite complex and multifaceted and can be described in various ways. Bock points out, in this respect, that ‘Darwin used natural selection in two distinct meanings’.64 He emphasized either the causes of natural selection or its consequences. Paradoxically, he generally used the phrase ‘natural selection’ in its causative meaning, but he focused on the consequences of natural selection in his most precise definitions.65 These, however, are certainly not the only two meanings or ways in which the phrase is (implicitly) used in The Origin.66 For example, Darwin sometimes referred to the selection of individual organisms, elsewhere to the selection of a clearly distinguished collection of individuals (i.e. a breed, form, variety or race) and, in a few cases, to the selection ‘of favoured individuals and races’.67 He also realized that natural selection could cause the transmutation of an entire species (what we call anagenesis or vertical evolution) or the transmutation of a separate variety or race (cladogenesis or horizontal evolution) (see, in this respect, figure 4). The process of cladogenesis (or, as Darwin called it, divergence) starts out with the selection of individual organisms that are only slightly different from other members of a species but leads, after a number of generations, to the creation of a distinct, ‘favoured’ variety or race. Figure 4. Darwin's diagram of divergence of taxa. The only figure in On the Origin of Species, it shows the divergent evolution, the non-divergent evolution or the extinction of various species (A–L) of a large genus. Each horizontal line (I–XIV) represents a thousand or more generations. Species A and I produce, after thousands or millions of generations, through divergent selection, 14 new species (a14–m14 and n14–z14). The six new species descended from I, and the eight descended from A, form distinct genera or even distinct sub-families. All the other species, except F (non-divergent evolution), become extinct without leaving descendant species. (From Wikimedia Commons.) * Download figure * Open in new tab * Download PowerPoint This is, of course, also the specific kind of natural selection that formed the subject of Darwin's subtitle: it referred to the preservation or selection of a group of organisms that was already clearly distinguished (i.e. a race). Indeed, in 1860, Louis Agassiz argued that Darwin did not substantiate the specific assertion that his subtitle implied: The assertion of Darwin, which has crept into the title of his work, is, that favoured races are preserved, while all his facts go only to substantiate the assertion that favoured individuals have a better chance in the struggle for life than others.68 Darwin did not show that these favoured individuals of a specific race diverged ‘from their specific type; and neither [he] nor anybody else has furnished a single fact to show that they go on diverging’.69 The reason why Darwin chose this specific definition of natural selection is obvious: divergence of character was, as he put in The Origin, ‘of high importance on my theory, and explains, as I believe, several important facts’.70 The idea was that, when organisms compete for scarce resources, natural selection should favour the individuals that most differ from their competitors since those individuals could occupy a new ‘station’ in ‘the economy of nature’ and thus escape the severe competition in the old ‘station’. Consequently, individuals that compete should, over time, diverge or become more dissimilar and develop into ‘favoured races’. Darwin's often-ridiculed race of black bears that was transformed into a giant, insect-eating whale offers a good example of such a ‘favoured race’: In North America the black bear was seen by Hearne swimming for hours with widely open mouth, thus catching, like a whale, insects in the water. Even in so extreme a case as this, if the supply of insects were constant, and if better adapted competitors did not already exist in the country, I can see no difficulty in a race of bears being rendered, by natural selection, more and more aquatic in their structure and habits, with larger and larger mouths, till a creature was produced as monstrous as a whale.71 In a letter of 5 September 1857 to Asa Gray, he presented divergence as ‘one other principle’ (next to the principle of selection) and the ‘means by which nature makes her species’: One other principle, which may be called the principle of divergence plays, I believe, an important part in the origin of species. The same spot will support more life if occupied by very diverse forms: we see this in the many generic forms in a square yard of turf (I have counted 20 species belonging to 18 genera),—or in the plants and insects, on any little uniform islet, belonging almost to as many genera and families as to species. … This, I believe, to be the origin of the classification or arrangement of all organic beings at all times.72 In a letter to Hooker of 8 June 1858 (written while he was still working on his big ‘Species Book’), Darwin remarked: ‘I will try to leave out all allusion to genera coming in & out in this part, till when I discuss the “principle of Divergence”, which with “Natural Selection” is the keystone of my Book & I have very great confidence it is sound’.73 He also described divergence in great detail in The Origin.74 It is significant that it was discussed, in both The Origin and Natural Selection, in the chapter dedicated to natural selection (respectively chapter 4, titled ‘Natural selection’, and chapter 6, titled ‘On natural selection’): divergent selection was a crucial part of the principle of divergence (see also note 70). The principle of divergence is also clearly illustrated in the only diagram included in his book (figure 4). It should therefore not surprise us that Darwin's subtitle defined ‘natural selection’ in terms of the selection of a clearly diverging group of organisms. This is maybe also another reason why he used the term ‘race’, for this was, in 1859, undoubtedly one of the best terms to refer to such a group of organisms. This, in turn, implies that the most common criticism of the title of The Origin—that Darwin failed to solve the problem ‘indicated by the title of his work’ (i.e. the origin of species) or that he remained largely silent about this problem—is not entirely correct.75 It is, as so often, a question of semantics. Through his subtitle, the phrase ‘origin of species’ acquired the very specific meaning of ‘origin of species through divergent selection’ and this process was, as just pointed out, central to The Origin. Consequently, Darwin did not fail to solve the problem indicated by the title of his work. However, he did indeed not offer a complete solution for the problem of the origin of new species through cladogenesis or horizontal evolution. Nor did he ‘prove that the principle of divergence plays a primary role in speciation’.76 However, it should immediately be added that modern research suggests that the importance of Darwinian character displacement should not be underestimated in a general theory on species diversification.77 FOUR OTHER MODIFICATIONS I can be brief about the four other modifications of the original title of The Origin. After comments from two friends about the position of the word ‘varieties’—that it ought to stand before ‘species’—and the absence of ‘genera & orders’ in the title, Darwin asked, in a letter of 10 September 1859, permission from Murray to delete the term ‘varieties’, because ‘The case of Species is the real important point; & the title, as now, is rather too long’.78 We do not know why and when he deleted the term ‘essay’. It may have been to further shorten the title but why did he then replace ‘through’ with the longer phrase ‘by means of’? Was he, once again, inspired by the Darwin–Wallace paper and, more particularly, by the last words of the title of this paper, ‘by natural means of selection’?79 Finally, in February 1872, Darwin dropped the last remnant of the introductory part of his original title: the preposition ‘On’. This last, subtle modification signals, or can be interpreted as signalling, the maturation or emancipation of a book that started out as a mere abstract of an ‘essay’. CONCLUSION On the Origin of Species is, without a doubt, one of the most famous and best-known book titles in history. Thanks to the long and complex history of Darwin's magnum opus and his tendency to vacillate, it is probably also a title with one of the longest gestation periods in the history of science book publishing. Thanks to the wealth of documents that Darwin left us, it is possible to reconstruct not only that gestation but also the reasoning behind it. Darwin certainly had good reasons to call his big ‘Species Book’ Natural Selection. Natural selection may not have been the most important component of his one ‘long argument’ but it was definitely its most brilliant component. It was also the common thread in his evolutionary theorizing. When he was forced to write swiftly an ‘Abstract of an Essay on Natural Selection’, instead of Natural Selection he made the historically crucial choice of inserting the phrase ‘the origin of species and varieties’ in his title. There are several non-exclusive factors that may have inspired this insertion: didactic reasons, the title of the article in which he elaborated his highly successful coral theory, the title of the Darwin–Wallace paper and/or Lyell's qualification, in 1856, of his theory as ‘On the Formation of Species by Natural Selection’. The main influence of John Murray was the addition of a subtitle to this title: ‘or the Preservation of Favoured Races in the Struggle for Life’. This referred to the kind of preservation or selection that was associated with the second keystone of Darwin's book: the principle of divergence. Other modifications were less crucial. Most enigmatic, maybe, was the replacement of ‘through’ with the longer phrase ‘by means of’. One might wonder whether this convoluted gestation process resulted in the best possible title for Darwin's book.80 Should he not, for example, have referred to the crucial second part of his book: his evolutionary reinterpretation of large parts of the contemporary knowledge about life and its history? However, that is not what this article was about. I have, through a detailed description of the trajectory between Natural Selection and The Origin and a tentative reconstruction of the reasoning behind this long and convoluted transmutation, merely tried to fill a small but not insignificant gap in the historiographical Darwin literature. ACKNOWLEDGEMENTS I would like to thank two anonymous reviewers for their helpful comments and recommendations. FOOTNOTES 1 Darwin to J. M. Rodwell, 5 November 1860, Darwin Correspondence Project, ‘Letter no. 2976', http://www.darwinproject.ac.uk/DCP-LETT-2976 (accessed 4 July 2018). 2 C. Darwin, On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life (John Murray, London, 1859). The study of Darwin, his work and his influence did not, of course, start in 1959. See, in this respect, J. C. Greene, ‘Reflections on the progress of Darwin studies’, Hist. Biol. 8(2), 243–273 (1975), at p. 243. He refers to the Isis Cumulative Bibliography: ‘for the years 1912–1919 the Bibliography lists 8 books and articles on Darwin and related subjects; 28 for the 1920's; 38 for the 1930's; 29 for the 1940's and 24 in the years 1950–1957’. However, these studies were not produced by a community of Darwin scholars but rather by isolated scholars and constituted a mere trickle in comparison to the torrent of the post-1959 Darwin industry. The 1959 centennial was not the only catalyst of the emergence of a Darwin industry. Greene refers in this respect to the crystallization of the modern evolutionary synthesis—which of course constituted a vindication of Darwin's ideas—and to the professionalization of the history and philosophy of science as an academic discipline. (Greene, op. cit. (this note), p. 248.) For analyses of the Darwin industry, see B. J. Loewenberg, ‘Darwin and Darwin studies, 1959–63’, Hist. Sci. 4(1), 15–54 (1965); M. Ruse, ‘The Darwin industry: a critical evaluation’, Hist. Sci. 12(1), 43–58 (1974); Greene, op. cit. (this note); T. Lenoir, ‘Essay review: the Darwin industry’, J. Hist. Biol. 20(1), 115–130 (1987); M. Ruse, ‘The Darwin industry: a guide’, Vic. Stud. 39(2), 217–235 (1996); M. C. Flannery, ‘The Darwin industry’, Am. Biol. Teach. 68(3), 163–166 (2006); D. Oldroyd, M. Ruse, P. Pearson and S. Herbert, ‘Review symposium: Darwin's geology: the end of the Darwin industry?’, Metascience 16(1), 25–50 (2007); J. van Wyhe, ‘Darwin online and the evolution of the Darwin industry’, Hist. Sci. 47(4), 459–473 (2009). 3 The main categorization of this literature could be non-hierarchical (i.e. based on a non-hierarchical list of subject matters) or hierarchical (i.e. based on the importance of the subject matter). In the latter case, Darwin biographies would undoubtedly figure among the most important subject matters. The timing of publication (i.e. possible delay) of The Origin and the differences between various editions are examples of subject matters of secondary importance. A third category, of least important topics, could encompass subjects such as Darwin's writing style and his much discussed, mysterious illness. 4 The ‘Historical sketch’ was prepared for the first authorized American edition of The Origin, published in May 1860, but it had already appeared in April 1860 as a preface to the first German edition. See C. N. Johnson, ‘The preface to Darwin's Origin of Species: the curious history of the “Historical sketch”’, J. Hist. Biol. 40(3), 529–556 (2007), at p. 530, n. 1. 5 Ibid., p. 531, n. 3. 6 Ibid. 7 A. Desmond and J. Moore, Darwin (Penguin, London, 1992), p. 474. 8 Ibid., p. 475. 9 J. Browne, Charles Darwin: the power of place (Alfred A. Knopf, New York, 2002), p. 81. 10 R. C. Stauffer, (ed.) Charles Darwin's Natural Selection: being the second part of his big Species Book written from 1856 to 1858 (Cambridge University Press, Cambridge, 1975). The first part of Natural Selection (two chapters) was cannibalized when Darwin wrote The Variation. C. Darwin, The Variation of Animals and Plants under Domestication, 2 vols (John Murray, London, 1868). Consequently, Stauffer's edition begins with chapter III, ‘Possibility of all organic beings crossing’. It comprises eight full chapters and one incomplete chapter on geographical distribution. Stauffer's division of Natural Selection into two parts is not to be confused with the bipartite structure of Darwin's argumentation (see note 25). A better title for the book that Stauffer edited might have been: The second part of Darwin's Natural Selection: chapters 3–11. 11 F. Darwin, (ed.) The foundations of The Origin of Species: two essays written in 1842 and 1844 (Cambridge University Press, Cambridge 1909). 12 In his personal journal, he wrote: ‘July 5th. Sent a written sketch of species theory (seven years after commencement[)] in about 230 pages to Mr. Fletcher to be copied’. Cambridge University Library, Darwin Online, DAR158.1–76, http://darwin-online.org.uk/content/frameset?pageseq=42&itemID=CUL-DAR158.1-76&viewtype=side (accessed 29 May 2018). 13 C. Darwin to E. Darwin, 5 July 1844, Darwin Correspondence Project, ‘Letter no. 761’, http://www.darwinproject.ac.uk/DCP-LETT-761 (accessed 12 March 2018). 14 C. Lyell to Darwin, 9 April 1843, Darwin Correspondence Project, ‘Letter no. 670’, http://www.darwinproject.ac.uk/DCP-LETT-670 (accessed 13 March 2018). 15 Lyell to Darwin, 1–2 May 1856, Darwin Correspondence Project, ‘Letter no. 1862’, http://www.darwinproject.ac.uk/DCP-LETT-1862 (accessed 16 March 2018). 16 It has long been believed that Lyell feared that Darwin would be scooped after reading Alfred Russel Wallace's essay on the introduction of new species: A. R. Wallace, ‘On the law which has regulated the introduction of new species’, Ann. Mag. Nat. Hist. 2nd Ser. 16, 184–196 (1855). However, this seems to be a myth. See, in this respect, J. van Wyhe, ‘The impact of A. R. Wallace's Sarawak Law paper reassessed’, Stud. Hist. Philos. Sci. C 60, 56–66 (2016). 17 D. Kohn, ‘Darwin's ambiguity: the secularization of biological meaning’, Br. J. Hist. Sci. 22, 215–239 (1989). 18 Darwin to J. D. Hooker, 9 May 1856, Darwin Correspondence Project, ‘Letter no. 1870’, http://www.darwinproject.ac.uk/DCP-LETT-1870 (accessed 16 March 2018). 19 Ibid. 20 Darwin to Hooker, 11 May 1856, Darwin Correspondence Project, ‘Letter no. 1874’, http://www.darwinproject.ac.uk/DCP-LETT-1874 (accessed 16 March 2018). 21 In his personal journal, he wrote: ‘May 14th Began by Lyells [sic] advice writing species sketch’. Cambridge University Library, Darwin Online, DAR158.1–76, http://darwin-online.org.uk/content/frameset?pageseq=66&itemID=CUL-DAR158.1-76&viewtype=text (accessed 29 May 2018). 22 Darwin to W. D. Fox, 3 October 1856, Darwin Correspondence Project, ‘Letter no. 1967’, http://www.darwinproject.ac.uk/DCP-LETT-1967 (accessed 16 March 2018). 23 Darwin to Lyell, 10 November 1856, Darwin Correspondence Project, ‘Letter no. 1984’, http://www.darwinproject.ac.uk/DCP-LETT-1984 (accessed 13 March 2018). 24 Darwin to A. Gray, 5 September 1857, Darwin Correspondence Project, ‘Letter no. 2136’, http://www.darwinproject.ac.uk/DCP-LETT-2136 (accessed 13 March 2018). 25 The Sketch (1842) seems to have had a tripartite plan (part 1 on variation under domestication, part 2 on variation under nature and part 3 on the reasons for and against believing that wild races ‘really have been produced, forming what are called species’), but, as editor, Francis Darwin ‘was being reasonable enough in dividing the Sketch not according to this tripartite plan … but according to the more fundamental bipartite structure of its argumentation’. M. J. S. Hodge, ‘Review: the structure and strategy of Darwin's “long argument”’, Br. J. Hist. Sci. 10(3), 237–246 (1977), at pp. 241–242. See also note 39. The ‘bigger work’ that Darwin intended to publish after the publication of On the Origin (1859) had a similar, tripartite structure. 26 Darwin to Gray, 11 May 1863, Darwin Correspondence Project, ‘Letter no. 4153’, http://www.darwinproject.ac.uk/DCP-LETT-4153 (accessed 18 December 2017). It was a reiteration of a letter that he had sent, a few days before, to the Athenæum, as a reaction to an anonymously published letter by Richard Owen: ‘Whether the naturalist believes in the views given by Lamarck, by Geoffroy St.-Hilaire, by the author of the “Vestiges”, by Mr. Wallace and myself, or in any other such view, signifies extremely little in comparison with the admission that species have descended from other species and have not been created immutable; for he who admits this as a great truth has a wide field opened to him for further inquiry. I believe, however, from what I see of the progress of opinion on the Continent, and in this country, that the theory of Natural Selection will ultimately be adopted, with, no doubt, many subordinate modifications and improvements’. Darwin to Athenæum, 5 May 1863, Darwin Correspondence Project, ‘Letter no. 4142’, http://www.darwinproject.ac.uk/DCP-LETT-4142 (accessed 19 December 2017). 27 Darwin to Lyell, 20 September 1859, Darwin Correspondence Project, ‘Letter no. 2492’, http://www.darwinproject.ac.uk/DCP-LETT-2492 (accessed 17 January 2018). 28 The Origin is indeed not explicitly divided into two parts, but chapters I–VIII correspond to Part I of the Sketch and the Essay and chapters IX–XIII (first edition) to Part II. It is often said that The Origin has a tripartite structure (see, e.g., R. Moore, ‘The persuasive Mr. Darwin’, Bioscience 47(2), 107–114 (1997)). Chapters VI–VIII, which deal with difficulties for Darwin's theory of natural selection (or even chapters VI–IX) are then considered as Part II. However, chapters VI–VIII belong to Part I of his argumentative structure, whereas chapter IX, on the imperfection of the geological record, opens Part II. It is correct, though, that Darwin, from the moment that he had been urged to publish something, had decided to give special attention to the difficulties in his theory: ‘If I publish anything it must be a very thin & little volume, giving a sketch of my views & difficulties; but it is really dreadfully unphilosophical to give a resumé, without exact references, of an unpublished work. But Lyell seemed to think I might do this, at the suggestion of friends, & on the ground which I might state that I had been at work for 18 years, & yet could not publish for several years, & especially as I could point out difficulties which seemed to me to require especial investigation’. Darwin to Hooker, 9 May 1856 (note 18). 29 D. Ospovat, The development of Darwin's theory: natural history, natural theology, and natural selection, 1838–1859 (Cambridge University Press, Cambridge, 1995), p. 88. 30 D. Partridge, ‘When did Darwin “clearly conceive” his theory of evolution?’ J. Nat. Hist. 52(1–2), 73–86 (2018), at p. 73. 31 L. G. Wilson, (ed.), Sir Charles Lyell's scientific journals on the species question (Yale University Press, New Haven, CT, 1970), p. 121. 32 Wallace, op. cit. (note 16); van Wyhe, op. cit. (note 16), p. 63. 33 C. Darwin, ‘On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. I. Extract from an unpublished work on species, II. Abstract of a letter from C. Darwin, Esq., to Prof. Asa Gray’, J. Proc. Linn. Soc. 3, 45–53 (1858); A. R. Wallace, ‘On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. III. On the tendency of varieties to depart indefinitely from the original type’, J. Proc. Linn. Soc. 3, 53–62 (1858). 34 Darwin to T. C. Eyton, 4 August 1858, Darwin Correspondence Project, ‘Letter no. 2319’, http://www.darwinproject.ac.uk/DCP-LETT-2319 (accessed 21 March 2018). 35 Darwin to Hooker, 12 October 1858, Darwin Correspondence Project, ‘Letter no. 2339’, http://www.darwinproject.ac.uk/DCP-LETT-2339 (accessed 20 March 2018). On 4 October 1858, he had already called it ‘an abstract of all my conclusions to be published as small book or read before Linn: Society’: Darwin to Eyton, 4 October 1858, Darwin Correspondence Project, ‘Letter no. 2333’, http://www.darwinproject.ac.uk/DCP-LETT-2333 (accessed 21 March 2018). 36 Darwin to Lyell, 28 March 1859, Darwin Correspondence Project, ‘Letter no. 2437’, http://www.darwinproject.ac.uk/DCP-LETT-2437 (accessed 24 March 2018). 37 As he declared in the introduction to The Origin: ‘No one can feel more sensible than I do of the necessity of hereafter publishing in detail all the facts, with references, on which my conclusions have been grounded; and I hope in a future work to do this’. Darwin, op. cit. (note 2), p. 2. 38 Darwin to J. Murray, 2 December 1859, Darwin Correspondence Project, ‘Letter no. 2566’, http://www.darwinproject.ac.uk/DCP-LETT-2566 (accessed 19 December 2017). 39 Darwin to Fox, 25 December 1859, Darwin Correspondence Project, ‘Letter no. 2604’, http://www.darwinproject.ac.uk/DCP-LETT-2604 (accessed 19 December 2017). See also Darwin to Murray, 22 December 1859, Darwin Correspondence Project, ‘Letter no. 2594’, http://www.darwinproject.ac.uk/DCP-LETT-2594 (accessed 19 December 2017). In The Variation, he called this book the first volume of that bigger work that he had announced in the introduction of The Origin and apologized for ‘the great delay in publishing this first work’, adding that it had ‘been caused by continued ill-health’ (Darwin, op. cit. (note 10), p. 2, n. 1). He also announced the two other, never-published volumes, one about ‘the Variation of organisms in a state of nature’ (p. 8) and one about ‘the several classes of facts’ (p. 9) that could be explained by the principle of natural selection. Stauffer points out, in this respect, that Darwin ‘did not abandon his long manuscript, nor write on the unused backs of the sheets for drafting other new publications as he so often did with other manuscripts’. Stauffer, op. cit. (note 10), p. 1. Indeed, he kept assembling notes for a big ‘Species Book’ until the 1870s (see Ospovat, op. cit. (note 29), p. 89). As Ospovat points out, the third volume (part 2 in his argument) would probably have been a very large one because his notes on geological succession, geographical distribution, morphology, and so forth were very extensive (see Darwin Manuscripts Project, DAR 205, https://tinyurl.com/y7ge5bux (accessed 29 May 2018)). Murray seems not to have been very enthusiastic about this project, which may help explain why it was never completed. See, in this respect, Browne, op. cit. (note 9), p. 97. 40 Hodge, op. cit. (note 25), p. 242. 41 Darwin to Hooker, 11 May 1856 (note 20). 42 C. R. Darwin, ‘On certain areas of elevation and subsidence in the Pacific and Indian oceans, as deduced from the study of coral formations’, Proc. Geol. Soc. Lond. 2, 552–554 (1837) (read 31 May 1837). It was later elaborated in Darwin's first monograph: C. R. Darwin, The Structure and Distribution of Coral Reefs. Being the First Part of the Geology of the Voyage of the Beagle, under the Command of Capt. Fitzroy, R.N. During the Years 1832 to 1836 (Smith Elder and Co., London, 1842). 43 In a letter that he sent on 18 May 1858 to Syms Covington, he declared that Natural Selection was his biggest work: ‘it treats on the origin of varieties of our domestic animals and plants, and on the origin of species in a state of nature’. However, this cannot have been the inspiration (and meaning) of the original title of his abstract, since domestic varieties are not produced through natural selection. Darwin to S. Covington, 18 May 1858, Darwin Correspondence Project, ‘Letter no. 2276’, http://www.darwinproject.ac.uk/DCP-LETT-2276 (accessed 13 March 2018). 44 Darwin to H. Denny, 7 November 1844, Darwin Correspondence Project, ‘Letter no. 787’, http://www.darwinproject.ac.uk/DCP-LETT-787 (accessed 21 March 2018). See also, in this respect, D. N. Stamos, Darwin and the nature of species (State University of New York Press, Albany, NY, 2007). 45 Darwin to F. Mackintosh Wedgwood, 18 [August 1856–January 1858], Darwin Correspondence Project, ‘Letter no. 1810’, http://www.darwinproject.ac.uk/DCP-LETT-1810 (accessed 21 March 2018). 46 C. Darwin, Journal of Researches into the Natural History and Geology of the Countries Visited During the Voyage of H.M.S. Beagle Round the World, under the Command of Capt. Fitz Roy, R.N., 2nd edn (John Murray, London, 1845). 47 Browne, op. cit. (note 9), p. 73. 48 Darwin to Lyell, op. cit. (note 36). 49 Darwin to Lyell, 30 March 1859, Darwin Correspondence Project, ‘Letter no. 2439’, http://www.darwinproject.ac.uk/DCP-LETT-2439 (accessed 8 December 2017). Darwin to Murray, 31 March 1859. Darwin Correspondence Project, ‘Letter no. 2441’, http://www.darwinproject.ac.uk/DCP-LETT-2441 (accessed 19 December 2017). 50 This certainly seems to be implied by what Darwin wrote to Lyell: he was ‘sorry about Murray objecting to term abstract as I look at it as only possible apology for not giving References & facts in full.—but I will defer to him & you’. Darwin to Lyell, 30 March 1859 (note 49). In note 2 to another letter by Darwin, addressed to Murray, the editor writes that Darwin had been advised ‘to drop the expression “Abstract of an essay” (letter to Charles Lyell, 30 March [1859])’. Darwin to Murray, 10 September 1859, Darwin Correspondence Project, ‘Letter no. 2488’, http://www.darwinproject.ac.uk/DCP-LETT-2488 (accessed 8 December 2017). That is not correct. In his letter to Lyell (30 March), Darwin only refers to the term ‘abstract’. 51 This interpretation of this draft title page is suggested by D. Ospovat, ‘God and natural selection: the Darwinian idea of design’, J. Hist. Biol. 13(2), 169–194 (1980), at p. 172, n. 10. The reference to Murray and to Whewell suggests that it is the correct interpretation. Although the verso of the half-title leaf of The Origin as published carries two quotations—one by Whewell and one by Bacon—whereas this draft title page only carries a quotation by Whewell. Also, this quotation was intended to be placed beneath the title. The emphasis on the year of publication as 1860 may even be incompatible with Ospovat's interpretation, since Darwin, to the best of my knowledge, never intended to publish The Origin in 1860 (the manuscript was finished in May 1859). Could it be that this was not an alternative title for The Origin but the ‘general title’ of the ‘bigger book’ that Darwin, in 1859, intended to ‘publish as 3 separate volumes, with distinct titles, but with a general title in addition’? 52 Darwin to Lyell, op. cit. (note 49). 53 Ibid. 54 R. Dawkins, Science in the soul: selected writings of a passionate rationalist (Bantam Press, London, 2017), p. 112. 55 The frequency with which (non-split) terms appear in The Origin can be determined through the search function of John van Wyhe's darwin-online.org.uk site. One should keep in mind, though, that terms are often used in different ways. 56 Darwin, op. cit. (note 2), p. 94. 57 Ibid., p. 172. 58 Darwin to A. R. Wallace, 22 December 1857, Darwin Correspondence Project, ‘Letter no. 2192’, http://www.darwinproject.ac.uk/DCP-LETT-2192 (accessed 27 March 2018). Darwin used the word ‘race’ mainly to refer to domestic or natural plants or animals. There are only three instances in The Origin where the term refers to ‘races of humans’. In one of these passages, Darwin foreshadowed his later book, The Descent of Man: ‘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 details my reasoning would appear frivolous’. Darwin, op. cit. (note 2), p. 199. C. Darwin, The Descent of Man, and Selection in Relation to Sex (John Murray, London, 1871). 59 Desmond and Moore, op. cit. (note 7), p. 442. See also A. Desmond and J. Moore, Darwin's sacred cause: race, slavery and the quest for human origins (Penguin Books, London, 2009). 60 With the benefit of hindsight, Darwin's choice was even less lucky or opportune. G. Himmelfarb, for example, points out that Darwin's subtitle ‘made a convenient motto for racists’ and that Darwin himself was ‘not averse to the idea that some races were more fit than others, and that this fitness was demonstrated in human history’. G. Himmelfarb, Darwin and the Darwinian revolution (Chatto & Windus, London, 1959), p. 416. M. Russell even claims that ‘favoured races’, translated, ‘meant his race, the Caucasians’. M. Russell, Beyond ramps: disability at the end of the social contract: a warning from an uppity crip (Common Courage Press, Monroe, ME, 1998), p. 19. Likewise, Dawkins writes that the word ‘race’ in the subtitle of The Origin ‘is sometimes misread in support of racialism’. Dawkins, op. cit. (note 54), p. 112. 61 Quoted in Hodge, op. cit., p. 242 (note 25), my italics. 62 Darwin to Murray, 5 April 1859, Darwin Correspondence Project, ‘Letter no. 2447’, http://www.darwinproject.ac.uk/DCP-LETT-2447 (accessed 19 December 2017). See also the letter that he sent to Murray on 2 April: Darwin Correspondence Project, ‘Letter no. 2445’, http://www.darwinproject.ac.uk/DCP-LETT-2445 (accessed 19 December 2017). 63 Darwin, op. cit. (note 2), p. 467. 64 J. W. Bock, ‘The Darwin–Wallace myth of 1858’, Proc. Zool. Soc. Lond. 62(1), 1–12 (2009), at p. 3. 65 For example, when he wrote that natural selection ‘is daily and hourly scrutinising, throughout the world, every variation, even the slightest; rejecting that which is bad, preserving and adding up all that is good’, he emphasized the causative or scrutinizing side of natural selection. Darwin, op. cit. (note 2), p. 84. In his most elaborate definition, however, he focused on the consequential or resultative dimension of natural selection: ‘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, and will generally be inherited by its offspring. The offspring, also, will thus have a better chance of surviving, for, of the many individuals of any species which are periodically born, but a small number can survive. 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. We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of Nature. But Natural Selection, as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man's feeble efforts, as the works of Nature are to those of Art’. Ibid., p. 61; see also Stauffer, op. cit. (note 10), p. 175. 66 Darwin had also already realized that selection can be endogenic (i.e. selection due to characteristics of the selected organism) or exogenic (i.e. selection of an organism due to the behaviour or characteristics of other organisms). That is how he explained the existence of castes of neuter insects with a profitable modification (i.e. profitable for the community to which they belonged). They evolved through the (exogenic) selection of their parents: ‘by the long-continued selection of the fertile parents which produced most neuters with the profitable modification, all the neuters ultimately came to have the desired character’. Darwin, op. cit. (note 2), p. 239. 67 Ibid., p. 467. 68 J. L. R. Agassiz, ‘Review of On the Origin of Species’, Am. J. Sci. Arts 30, 142–154 (1860), at p. 149, (my italics). 69 Ibid., p. 149. 70 Darwin, op. cit. (note 2), p. 111. The principle of divergence ‘grew out of work on classification’ (Ospovat, op. cit. (note 29), p. 171). The question that he initially posed ‘was not how one parent species gives rise to two descendant species, but rather “why the species of a large genus, will hereafter probably be a Family with several genera”’ (ibid.). It has been interpreted in various ways in the literature. D. Kohn argues that the principle of divergence was simply a special case or type of natural selection which he calls ‘divergent selection’. D. Kohn, ‘Darwin's keystone: the principle of divergence’, in The Cambridge companion to the Origin of Species (eds D. Kohn and M. Ruse), pp. 87–108 (Cambridge University Press, Cambridge, 2009). It certainly encompassed, or resulted in, that kind of selection. Put differently: Darwin's ‘principle of divergence’ encompassed the notion of divergent selection (i.e. the selection of the most divergent members of a population) and an explanation for divergent selection (i.e. organisms that differ from other members of the species can invade and occupy a new niche). For more literature on this principle, see J. Browne, ‘Darwin's botanical arithmetic and the principle of divergence, 1854–1858’, J. Hist. Biol. 13, 53–89 (1980); D. Kohn, ‘Darwin's principle of divergence as internal dialogue’, in The Darwinian heritage (ed. D. Kohn), pp. 245–258 (Princeton University Press, Princeton, 1985); E. Mayr, ‘Darwin's principle of divergence’, J. Hist. Biol. 25(3), 343–359 (1992); W. Tammone, ‘Competition, the division of labor, and Darwin's principle of divergence’, J. Hist. Biol. 28, 109–131 (1995); D. W. Pfennig and K. S. Pfennig, ‘Character displacement and the origins of diversity’, Am. Nat. 176(1), S26–S44 (2010); R. J. Richards, ‘Darwin's principles of divergence and natural selection: why Fodor was almost right’, Stud. Hist. Philos. Sci. C 43(1), 256–268 (2012). 71 Darwin, op. cit. (note 2), p. 184. 72 Darwin to op. cit. (note 24). 73 Darwin to Hooker, 8 June 1858, Darwin Correspondence Project, ‘Letter no. 2282’, http://www.darwinproject.ac.uk/DCP-LETT-2282 (accessed 19 December 2017). 74 Darwin, op. cit. (note 2), pp. 111–126. See also Stauffer, op. cit. (note 10), pp. 227–250. 75 ‘Darwin failed to solve the problem indicated by the title of his work. Although he demonstrated the modification of species in the time dimension, he never seriously attempted a rigorous analysis of the problem of the multiplication of species, the splitting of one species into two’. E. Mayr, Animal species and evolution (Belknap Press of Harvard University Press, Cambridge, MA, 1963), p. 12. In a similar vein, J. A. Coyne and H. A. Orr point out that ‘The Origin of Species, whose title and first paragraph imply that Darwin will have much to say about speciation … remains largely silent on the “mystery of mysteries”, and the little it does say about this mystery is seen by most modern evolutionists as muddled or wrong’. J. A. Coyne and H. A. Orr, Speciation (Sinauer, Sunderland, MA, 2004), p. 9. They believe that ‘for Darwin, the origin of species was identical to the origin of adaptations within species’ (ibid., p. 11). Coyne has even argued that The Origin of Species should have been called The Origin of Adaptations because Darwin ‘confused adaptation within lineages with the origin of new lineages’. J. A. Coyne, ‘Ernst Mayr and the Origin of Species’, Evolution 48(1), 19–30 (1994), at p. 19. 76 Mayr, op. cit. (note 70), p. 357. 77 Pfennig and Pfennig, op. cit. (note 70). 78 Darwin to Murray, op. cit. (note 50). 79 This was also the way in which he, in the Sketch, first formulated the idea of natural selection. 80 Another, historiographical question is whether or not Darwin later regretted having ended up with this specific title. He developed mixed feelings about the phrase ‘natural selection’ and sometimes used Herbert Spencer's phrase ‘the survival of the fittest’ but, as far as I know, never criticized the title of his magnum opus and he also never coined an alternative title for The Origin. The fact that the title of foreign editions of The Origin (e.g., Über die Entstehung der Arten im Thier- und Pflanzen-Reich durch natürliche Züchtung, oder, Erhaltung der vervollkommneten Rassen im Kampfe um's Daseyn, 1860) was, generally, very similar to the English title maybe also indicates that Darwin was not unhappy with his choice as foreign editions of books offer a good opportunity to launch a new, improved title. The Origin was translated in twenty-nine languages, a number that is higher than that of any other science book, except for the first books of Euclid (see http://darwin-online.org.uk/EditorialIntroductions/Freeman_OntheOriginofSpecies.html). © 2018 The Author(s) Published by the Royal Society. Previous Article Next Article VIEW FULL TEXT DOWNLOAD PDF RECOMMENDED ARTICLES 1. Essay Reviews - Charles Darwin: Imprimatur! Notes and Records, 1990 2. Introduction Edmund Brisco Ford et al., Proceedings B, 1956 3. 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Jones M, Rivett D, Pascual-García A and Bell T (2021) Relationships between community composition, productivity and invasion resistance in semi-natural bacterial microcosms, eLife, 10.7554/eLife.71811, 10 * None THIS ISSUE 20 March 2019 Volume 73Issue 1 * Article Information * DOI:https://doi.org/10.1098/rsnr.2018.0015 * Published by:Royal Society * Online ISSN:1743-0178 History: * Published online22/08/2018 * Published in print20/03/2019 License: © 2018 The Author(s) Published by the Royal Society. 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