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Darwin and the political economists: Divergence of character

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Conclusion

Several stages can be identified in Darwin's effort to formulate natural selection. The first stage corresponded, roughly speaking, to the period up to 1844. It was characterized by Darwin's attempt to base his model of geographic speciation on an individualistic dynamics, with species understood as reproductively isolated populations. Toward the end of this period, Darwin's ignorance of the laws of variations and heredity led him to adopt varieties and species as the units of variations. This had the extremely important effect of stimulating him to consider the process of speciation as involving populations. At the end of this period, Darwin also began to regard adaptation as being exclusively toward places in the economy of nature. Thus he faced the problem of integrating the process of natural selection with the process of speciation. Individual variants were the units that fueled the first process, whereas varieties produced new speices. There was no link between adaptation and speciation, except whatever could be supplied by a quasi-historical, developmental idea of optimizing the amount of life.

In the second stage, I contend, Darwin's reading of Milne-Edwards crystallized his previous insights into a coherent whole. Milne-Edwards' comments on the advantage of functional specialization could readily be understood in terms of the advantage accruing to the individual, relative to other members of its species, from occupying a different niche. Milne-Edwards' discussion of the division of labor suggested that organisms which moved into unoccupied niches would enjoy reduced competition, and hence a differential advantage in survival and reproduction; thus they would induce the species to do likewise. Rather than base his explanation on an analogy with the artificial economy, Darwin chose the principle of the optimalization of the amount of life per unit area as the overall explanatory principle. The difficulties connected with integrating different levels of description were therefore circumvented, insofar as the problem of diversity and speciation was concerned. Although natural selection considered individuals as the units of selection, and the units of variations were varieties and species, the dynamics of the process understood in terms of natural selection, competition, division of labor and niches could give a plausible account of how individual advantage could be transferred to the species, and how diversity resulted from this mechanism. The problem of the different levels of descriptions was confined to how the properties of variations in individuals (in particular, the frequency of variations and their transmission) were responsible for the assumed variability characteristic of varieties and species. This problem Darwin never solved.

A third stage occurred in 1858 with the amalgamation of the tree-of-life vizualization of the process of speciation. Speciation, geographic distribution, and systematics were all then embedded in a conceptual matrix with vast explanatory powers.

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References

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  3. He had read it in Paley's Natural Theology while an undergraduate at Cambridge and in 1833 in Humboldt's Political Essay on the Kingdom of New Spain (New York: Riley, 1811) in Buenos Aires, while the Beagle docked there; and in the summer of 1838, when Darwin was led to Malthus by way of Quetelet, he was also rereading Humboldt's Political Essay. The C transmutation notebook has the following entry on p. 268: “Humboldt: New Spain much about castes etc.”; and on the same page appears: “Find out from Statistical Society where M. Quetelet has published his laws about sexes relative to age of marriages.” All four transmutation notebooks are transcribed in Gavin de Beer, ed., “Darwin's Notebooks on Transmutation of Species, Bull. Brit. Mus. (Nat. Hist.) (Hist. Ser.), 2, (1960), De Beer, M. J. Rowlands, and B. M. Skramorski, ed., “Pages Excised by Darwin,” ibid., 3 (1967), 129–176 (excised pages). De Beer's “First,” “Second,” “Third,” and “Fourth” notebooks correspond to Darwin's B, C, D, and E notebooks. Throughout I will cite them by Darwin's letter and page number followed by a lower-case e in the case of excised pages: e.g., C, p. 123.

  4. See, e.g., Elie Halévy, The Growth of Philosophic Radicalism, (Boston: Beacon Press, 1955); Edwin Cannan, A History of the Theories of Production and Distribution in English Political Economy from 1776 to 1848, 3rd ed. (London: Staples Press, 1917).

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  11. Darwin's indebtedness to Scottish philosophy merits a thorough study. Scottish philosophy from Hume to Dugalt Stewart and Thomas Browne rejected the notion that one could explain anything by referring to ultimate principles or processes whose existence and operation could not be ascertained from the observation of particular instances. And Hume's notion of causality played a central role in the development of the “social” sciences and in ascertaining mechanisms of evolution. See Norman Kemp Smith, The Philosophy of David Hume (London: Macmillan, 1941); Thomas Browne's “Inquiry into the Relation of Cause and Effects,” in Geoffrey Keynes, ed., The Works of Sir Thomas Browne, 6 vols., (London: Faber and Gwyer, 1928); J. B. Morrell, “The University of Edinburgh in the Late Eighteenth Century,” Isis, 62 (1971), 158–171. For remarks on the British character of Darwinian evolution see E. Mayr, “The Nature of the Darwinian Revolution,” Science, 176 (1971), 981–989, reprinted in Mayr's Evolution and the Diversity of Life, pp. 277–296. The attempt to characterize Darwinian evolutionary theory as British has a long history. It has been observed repeatedly that James Cowles Prichard, William Lawrence, Joseph Adams, James Anderson, William Marshall, and Charles Wells, among others, were looking for a mechanism to explain the origin of organic diversity based solely on individual heredity and variation. See Herbert Hayes Odom, “Ground Work for Darwinism: Theories of Heredity and Variation in Great Britain, 1790–1820,” Ph.D. diss., Harvard University, 1972; P. J. Darlington, Darwin's Place in History (Oxford: Blackwell, 1959), pp. 19–24; Jacques Roger, Les sciences de la vie dans la pensée française du XVIII e siècle (Paris: A Colin, 1963); and J. B. Morrell, “Individualism and the Structure of British Science in 1830,” Hist. Stud. Phys. Sci., 3 (1971), 183–204.

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  12. Both the “Sketch of 1842” and the “Essay of 1844” appear in Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), and in Charles Darwin and A. R. Wallace, Evolution by Natural Selection (Cambridge: Cambridge University Press, 1958). My citations refer to the Darwin and Wallace book.

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  13. De Beer, ed., “Darwin's Notebooks on Transmutation of Species”; Gavin de Beer and M. J. Rowlands, ed., “Addenda and Corrigenda,” Bull. Brit. Mus. (Nat. Hist.) Hist Ser., 2 (1961); De Beer, Rowlands, and Skramorski, ed., “Pages Excised by Darwin,” pp. 129–176.

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  15. Ghiselin, The Triumph of the Darwinian Method.

  16. In the “Essay of 1844” Darwin does speak of perfect adaptation with reference to sexual selection: “The most rigorous males, implying perfect adaptation, must generally gain the victory in their several contexts. This kind of selection, however, is less rigorous than the other [i.e., natural]” (p. 121). For a sharply contrasting view, see Dov Ospovat, “Darwin after Malthus,” address at the History of Science meeting, Madison, Wisconsin, Oct. 31, 1978, and “Perfect Adaptation and Teleological Explanation: Approaches to the Problem of the History of Life in Mid-Nineteenth Century,” Stud. Hist. Biol., 2 (1978), 33–56.

  17. Darwin and Wallace, Evolution by Natural Selection, p. 41. The quotation occurs in the opening sentence of the “Sketch of 1842.”

  18. Darwin's interest in hermaphrodism is already evident in the notebooks. See, e.g., B, p. 96, C, p. 245, D, p. 174, and E, pp. 70–71. See the listings in “Concordance-Darwin Manuscripts at Cambridge University Library,” ed. P. H. Barrett (Michigan State University, 1977). I thank Professor Barrett for making a copy of the “Concordance” available to me.

  19. Box 49 of Darwin's papers at Cambridge University Library contains his notes on dichogamy in plants (the maturing of the anthers and stigmas of individual flower at different times) dated 1841. At that time Darwin viewed dichogamy not only as a device to reduce the self-fertilization of individual flowers but also as a means of favoring the crossing of different individuals.

  20. In his Autobiography, LLD, p. 74, Darwin wrote, “I had noticed in 1838 or 1839 the dimorphism of Linum flairim, and had at first thought that it was merely a case of unmeaning variability. But on examining the common species of Primula I found that the two forms were much too regular and constant to be thus viewed.”

  21. The centrality of biogeography in the notebooks has been stressed by Limoges, La sélection naturelle, pp. 57–59, and by R. C. Stauffer, “Ecology in the long Manuscript Version of Darwin's Origin of Species and Linnaeus' Economy of Nature,” Proc. Amer. Phil. Soc., 104 (1960), 235–241, and “Haeckel, Darwin, and Ecology,” Quart. Rev. Biol., 32 (1937), 138–144. See also P. Vorzimmer, “Darwin's Ecology and Its Influence upon His Theory,” Isis, 56 (1965), 148–155; F. Egerton, “Humboldt, Darwin, and Population,” J. Hist. Biol., 3 (1970), 326–360; Egerton, Studies of Animal Population from Lamarck to Darwin,” J. Hist. Biol., 1 (1968), 255–259. For earlier but post-Origin, nontechnical presentations of biogeography, see Asa Gray, “Species as to Variation, Geographical Distribution, and Succession,” Amer. J. Sci. and Arts, May 1863, reprinted in Asa Gray, Darwiniana, ed. A. H. Dupree (Cambridge, Mass.: Harvard University Press, 1963). Biogeography also played a major role in Wallace's development of natural selection theory. In 1876, with Darwin's encouragement, Wallace wrote The Geographical Distribution of Animals (London: Macmillan, 1876), noting in the preface: “I am well aware that this first outline of a great subject is, in parts, very meagre and sketchy; and, though perhaps overburdened with some kinds of detail, yet leaves many points most inadequately treated. It is therefore with some hesitation that I venture to express the hope that I have made some approach to the standard of excellence I have aimed at; — which was, that my book should bear a similar relation to the eleventh and twelfth chapters of the Origin of Species, as Mr. Darwin's Animals and Plants Under Domestication does to the first chapter of that work. Should it be judged worthy of such a rank, my long, and often wearisome labours, will be well repaid.” See also A. R. Wallace and W. T. Thiselton-Dyer, “The Distribution of Life, Animal and Vegetable in Space and Time,” Humboldt Library of Popular Science Literature, January 1885; and Thiselton-Dyer's “Geographical Distribution of Plants” and Hans Gadow's “Geographical Distribution of Animals,” chap. 16 and 17 in A. C. Seward, ed., Darwin and Modern Science (Cambridge: Cambridge University Press, 1909). For other interesting articles on biogeography, see David Starr Jordan, “Isolation as a Factor in Organic Evolution,” in Fifty Years of Darwinism: Modern Aspects of Evolution (New York: Henry Holt, 1909); J. B. S. Haldane, “Natural Selection,” in P. R. Bell, ed., Darwin's Biological Work: Some Aspects Reconsidered (Cambridge: Cambridge University Press, 1959); and P. J. Darlington, Jr., “Darwin and Zoogeography,” Proc. Amer. Phil. Soc., 103 (1959), 307–319.

  22. The notebooks contain many references to A. von Humboldt and A. Bonpland's Essai sur la géographie des plantes (Paris, 1805) and to Humboldt's later essays “on the laws observed in the distribution of vegetable forms.” The latter essays (originally written in French) were translated into English and appeared in the Phil. Mag. J., 47 (1816), 446, and in the Edinburgh Phil. J., 6 (1822), 273. The earlier essay also appeared in 1817 as De Distributione Geographica Plantarum (Paris); there Humboldt showed that in plants the average number of species in genera is larger by a factor of two in France as compared with the number in Lapland. Humboldt believed that the number of species in each order is determined by a mathematical law and that this law remains constant throughout any geological epoch. Darwin appreciated Humboldt's efforts to use these quantitative relations to predict plant distributions in similar climatic regions. The E notebook contains many questions relating to quantitative biogeographical data.

  23. Schweber, “Origin of the Origin,” p. 264.

  24. In reviewing John Richardson's Fauna Boreali-America in the Edinburgh Review, 52 (1831), 328–360, James Wilson wrote an essay on the geographical distribution of animals that surveyed the field as of 1830, before the publication of Lyell's Principles of Geology. Wilson wrote: “A knowledge of the various phenomena presented by the different groups of animals and plants, in accordance with the latitude, the longitude, and the altitude of their position constitutes the science ... and forms one of the most interesting and important branches of natural history ... The geographical distribution of animals presents a wide field for speculation, although the modes by which that distribution has been effected will probably remain ever concealed from human knowledge. Their gradual extension by natural means, from a single center of creation, scarcely falls within the sphere of credibility; and thus the creation of various groups of species over different points of the earth's surface ... or the removal and dispersion, by supernatural agency, of the greater proportion of existing species from an original center seem to be the two points ... to be illustrated ... It is for the naturalist ... to collect an ample, accurate and extended series of facts ... and, by comparing and combining these determinant observations, to deduce the laws in accordance with which species and genera are now dispersed over the surface of the earth.” As a footnote to this last sentence, Wilson added: “One of the most important of those preliminary enquiries which are essential to a proper comprehension of zoological geography, consists of the investigation and ascertainment (at least approximately) of the limits which nature has assigned to the variation in the specific characters of animals, and the establishment of fixed and determinate principles, by reference to which it may be discovered whether certain distinctions were sufficient to constitute a specific difference, or were merely the result of climate, or some peculiar or accidental combination of circumstances ... One of the chief difficulties ... in tracing the distribution of widely-extended species, arises from the uncertainty under which naturalists labour, from the want of a positive and assured test to ascertain whether a certain character should be regarded as expressive of specific distinction, or ought rather to be ranked as within the legitimate range of individual variation ... Where we have acquired a knowledge of the habits and economy of a species, and of the individuals of that species, wheresoever found, and if these are uniformly the same under different and far-removed localities, then a distinction in plumage should be regarded as insufficient to constitute a specific difference between them; but when we find the individual from one country or continent characterized and distinguished by some peculiarity in their instinctive habits, or modes of life, as well as by a cognizable different of aspect, we are then authorized to infer that they are specifically distinct, and are entitled to rank them accordingly. We have entered into these apparently trifling details, because we are aware that some modern writers deny that any species is widely distributed.”

  25. In the Autobiography, Darwin indicates that “during my last year at Cambridge, I read with care and profound interest Humboldt's Personal Narrative” (LLD, p. 47). He read passages from it aloud to his friends on their excursions with Henslow. Darwin's personal copy of vols. I and II of A. Von Humboldt and A. Bonplan's Personal Narrative of Travels to the Equinoctial Regions of the New Continent during the Years 1799–1804, trans. Helen Williams, 7 vols. in 9 (London, 1814–1829), was given to him by Henslow and is inscribed “J. S. Henslow to his friend C. Darwin on his departure from England upon a voyage round the World 21 Sept. 1831.”

  26. Darwin noted in his Autobiography that he had met Robert Brown several times before boarding the Beagle, and he wrote Henslow that he had received advice from Brown about microscopes. Among Darwin's reprint collection is a copy of Brown's Observations on the Organs and Mode of Fecundation in Orchideae and Asclepiadeae (London: Richard Taylor, 1831), inscribed “Given to me by Mr. Brown on Friday, December 9th, 1831.” In view of Brown's travels as a naturalist and his phytogeographic interests it is not unlikely that Brown would have stressed the importance of biogeography in talking to Darwin. Darwin's letters to Henslow during the voyage of the Beagle indicate that the Dictionnaire classique d'histoire naturelle (Paris: Rey et Granier, 1822–1831) (in which Candolle had written the article on plant geography and Humboldt that on geognosy) and Humboldt's Voyages aux regions equinoxiales were aboard, and, moreover, that Darwin was reading them. See Nora Barlow, ed., Darwin and Henslow: The Growth of an Idea, Letters, 1831–1860 (Berkeley: University of California Press, 1867), in particular letter 1, p. 26, letter 19, p. 54. Candolle's essay on plant geography had originally appeared in 1820 as the Essai elémentaire de géographie botanique; in this form, it was well known and had proved very influential. It contained the profound statement: “All the theory of geographical botany rests on the particular idea one holds about the origin of living things and the permance of species.”

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  27. Charles Lyell, Principles of Geology, vol. II (London: John Murray, 1832).

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  28. Chap. 5 of vol. II of Charles Lyell, Principles of Geology, vol. II (London: John Murray, 1832) p. 66, begins with:. “Next to determining the question whether species have a real existence, the consideration of the laws which regulate their geographical distribution is a subject of primary importance to the geologist. It is only by studying these laws with attention, by observing the position which groups of species occupy at present, and inquiring how these may be varied in the course of time by migrations, by changes in physical geography, and other causes, that we can hope to learn whether the duration of species be limited, or in what manner the state of the animate world is affected by the endless vicissitudes of the inanimate.” Vol. II of Lyell's Principles is also important because it contains many references to Candolle, Brown, and Humboldt, which would have stimulated Darwin to read these works.

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  29. aMrs. Lyell, ed., Life, Letters, and Journals of Sir Charles Lyell, Bart., 2 vols. (London: John Murray, 1881), I, 246.

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  30. See the entries under these names in Barrett's “Concordance” to the notebooks.

  31. William Swainson, A Treatise on the Geography and Classification of Animals (London: Longmans, 1835); see B, pp. 67, 92, and 276.

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  32. William Swainson, A Treatise on the Geography and Classification of Animals (London: Longmans, 1835), pp. 1–2.

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  33. The tree-of-life and “coral”-of-life metaphor and diagrams are introduced early in the B notebook, pp. 21–24; see the discussion in Gruber and Barrett, Darwin on Man, (New York: E. P. Dutton, 1974); H. Gruber, “Darwin's Tree of Nature and Other Images of Wide Scope.” in On Aesthetics in Science, ed. J. Wechsler (Cambridge, Mass.: M.I.T. Press, 1978).

  34. In Lyell's Principles of Geology, which Darwin studied carefully in its original form in the first edition of 1830–1833 and thereafter in its various revisions, the Tertiary period received special attention. The fossil record of the mollusks of the Tertiary was the basis of the time reckoning Lyell had developed. See M. Rudwick, The Meaning of Fossils: Episodes in the History of Palaeontology (New York: American Elsevier, 1972); M. Rudwick, “The Strategy of Lyell's Principles of Geology,” Isis, 61 (1970), 4–33; L. G. Wilson, Charles Lyell, The Years to 1841: The Revolution in Geology (New Haven: Yale University Press, 1972); and particularly M. Rudwick, “Charles Lyell's Dream of a Statistical Palaeontology,” Palaeontology, 21 (1978), 225–244.

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  35. W. F. Cannon, “The Uniformitarian-Catastrophist Debate,” Isis, 51 (1960), 38–55, and “The Problem of Miracles in the 1830's,” Vict. Stud., 4 (1960), 5–32; see also Life of Lyell, I, 467.

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  36. Malcolm J. Kottler, “Charles Darwin's Biological Species Concept and Theory of Geographic Speciation: The Transmutation Notebooks,” Ann. Sci., 35 (1978), 275–297.

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  37. L.von Buch, Description physique des isles Canaries, trans. C. Boulanger (Paris: F. G. Levrault, 1926). Von Buch repaid the compliment; writing to Humboldt after he had read Darwin's Journal of Researches, von Buch said that his views on the origin of species on earth “are very stimulated by the excellent and solid description of the Galapagos by Darwin.” He did not, however, agree with darwin's theory of the formation of coral atolls. See H. Beck, Alexander von Humboldt (Wiesbaden: F. Sterner, 1959), II, 301 n. 120.

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  38. In chap. 10 of the Origin Darwin wrote: “Mr. Clift many years ago showed that the fossil mammals from the Australian caves were closely allied to the living marsupials of that continent ... I was so much impressed with these facts that I strongly insisted, in 1839 and 1845, on this ‘law of the succession’, on ‘this wonderful relation in the same continent between the dead and living’”; see also F. Darwin and A. C. Steward, ed., More Letters of Charles Darwin, 2 vols. (London: John Murray, 1903), letter 87, to Charles Lyell, pp. 132–134; this book is hereafter cited as More Letters.

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  39. E, p. 114 is dated March 12, 1839. At some later date Darwin inserted after the sentence “Has nature any process analogous...” the words “makes the difficulty apparent by cross-questioning”.

  40. E. Forbes, “Report on the Mollusca and Radiata of the Aegean Sea, and on Their Distribution Considered as Bearing on Geology,” Report of the Thirteenth meeting of the British Association for the Advancement of Science, 13 (1843), 130–207.

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  41. Forbes's paper showed that like the land, the seabed was subdivided into zones and stations. Although the potential for species formation by geographical isolation existed on the sea floor, how such speciation occurred within a zone was not obvious. Forbes's work probably also reinforced Darwin's view that adaptations were toward ecological niches. Today it is clear that Darwin was tackling extremely difficult problems, many of which are still unsolved. The answer to the question of whether species originate allopatrically or sympatrically is an ambiguous one. The vocabulary which has emerged in recent years is revealing. In addition to allopatric and sympatric speciation, one speaks of semigeographic, semisympatric, statipatric, parapatric, alloparapatric speciation! See, e.g., G. L. Brush, “Modes of animal Speciation,” Ann. Rev. Ecol. and Syst., 6 (1975), 339–364.

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  42. “Essay of 1844,” p. 119.

  43. This point has been stressed by Ernst Mayr in private conversations.

  44. H. Milne-Edwards, Introduction à la zoologie générale ou considerations sur les tendances de la nature dans la constitution du règne animal (Paris: Victor Masson, 1851). pp. 7–8. The idea that “nature is like an artist making a thousand pictures from a single plan” is Goethe's.

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  45. The M and N notebooks have been transcribed by Paul Barrett and published in Gruber and Barrett, Darwin on Man.

  46. The position of the adjective “physiological” is important. As one of the editors of More Letters, Francis Darwin commented that Darwin's use of “division of physiological labour” in his letter to Hooker in 1854 was “a slip of the pen for physiological division of labour.” But in fact Darwin was correctly quoting Milne-Edwards. It was in his own writing that Darwin put “physiological” before “division.” See More Letters, I, 76.

  47. Darwin, Natural Selection, p. 233.

  48. Charles Darwin, The Foundations of the “Origin of Species”: Two Essay Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), p. 115–116.

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  49. Darwin and Wallace, Evolution by Natural Selection, pp. 264–267.

  50. R. C. Stauffer, in his masterly edition of Natural Selection indicates that chap. 6, “On Natural Selection,” was worked on at two different periods. The first draft (written on gray foolscap paper) was completed on March 31, 1857, and contained only a brief mention of the principle of divergence. The later additions and versions (written on bluish-gray paper) were made between April 14 and June 12, 1858, and in them Darwin devotes over forty pages to divergence. The reference to Milne-Edwards occurs in the second draft of the principle of divergence. It has been hypothesized by John L. Brooks (The American Philosophical Society Yearbook, 1968, pp. 534–535) that Darwin's concept of divergence was formulated in response to his receipt of Wallace's MS in 1858. Brooks suggested that Darwin did not obtain the MS on June 18, 1858, as is usually assumed, but on May 18, so that it was in Darwin's hands several weeks before June 12, the date Darwin gives in his private diary for the completion of the “note on divergence” that he inserted in chap. 6 of the long manuscript of which the Origin of Species is an abstract. H. L. McKinney in Wallace and Natural Selection (New Haven: Yale University Press, 1972), pp. 138–146, has conclusively shown that Darwin did not receive the Wallace MS before June 3, 1858. But there can be no doubt that Darwin had the concept of divergence of character before May or June of 1858. The letter of September 1857 to Asa Gray would be sufficient proof of that.

  51. More Letters, II, p. 99.

  52. The letter is reprinted in Darwin and Wallace, Natural Selection, pp. 264–267. According to A. H. Dupree, Asa Gray (Cambridge, Mass: Harvard University Press, 1968), pp. 246–247, 458–459n23, Darwin's letter to Gray was written by an amanuensis but corrected by Darwin. “It varies in detail from the version published at the Linnean Society.”

  53. More Letters, II, 109.

  54. Charles Darwin, and Alfred Wallace, “On the Tendency of Species to Form Varieties” and “On the Perpetuation of Varieties and Species by Natural Means of Selection,” J. Linn. Soc., 3 (1859), 45 (read July 1, 1858); see especially the “Abstract of a letter from C. Darwin, Esq., to Professor Asa Gray of Boston, U.S., dated September 5, 1857.”

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  55. De Beer, Evolution by Natural Selection, p. 140. It was George Bentham's paper on the fixity of species that was displaced by the reading of the Darwin-Wallace material at the Linnean Society on July 1, 1858. Bentham seems to be one of the few people who was struck by the Darwin-Wallace findings, for he altered his paper in the light of their presentation. Bentham was a naturalist whose book Labiatarum Genera et Species (London, 1832–1836) Darwin had studied in 1837–1839. On March 6, 1838, Darwin noted on p. 104 of the E notebook: “Mr. Bentham says in Sandwich Isld. he believes there are many cases of genera peculiar to the group having species peculiar to the separate islands. In his work on the Labiatae some of the species are described — capital case — for Sandwich Isld are very similar to Galapagos — study Flora. What general forms — are the Labiatae nearest to American or Indian Groups?”

  56. LLD, II, 210–211.

  57. Charles Darwin, A Monograph on the Subclass Cirripedia with Figures of All the Species: The Lepadidae or Pedunculated Cirripedes (London: The Ray Society, 1851).

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  58. Darwin, Natural Selection, p. 227.

  59. Ibid., p. 228.

  60. Ibid., p. 233.

  61. Ibid., p. 234.

  62. Ibid., p. 235.

  63. Ibid., p. 235.

  64. Ibid., p. 235.

  65. Ibid., pp. 236–237.

  66. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), p. 111.

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  67. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), pp. 111–126.

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  68. Charles Darwin, The Variation of Animals and Plants under Domestication 2 vols. (London: John Murray, 1868); the 2nd ed., revised, appeared in 1875.

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  69. Charles Darwin, The Variation of Animals Plants under Domestication, (London: John Murray, 1868); 2nd ed., p. 18.

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  70. That the insight into divergence of character may have occurred twice has previously been suggested by Gruber, Darwin on Man, p. 117.

  71. The first diagram appears in B, p. 26. A particularly penetrating discussion of the tree-of-life diagrams appears in Darwin on Man, pp. 117–118, 140–149, 195–198, and in Gruber's essay “Darwin's Three of Nature.”

  72. The second diagram is in B, p. 26; B, p. 25: “The tree of life should perhaps be called the coral of life; base of branches dead; so that passages cannot be seen.”

  73. The diagram is in B, p. 35; B, p. 36: “Case must be that one generation then should have as many living as now. To do this & to have many species (as is) requires extinction.”

  74. B, pp. 21–22: “Organized beings represent a tree, irregularly branched; some branches far more branched, — Hence Genera.” See also B, pp. 39–40.

  75. B, pp. 37–38: “With respect to extinction we can easy see that variety of ostrich Petise may not be well adapted, & thus perish out, or on other hand like Orpheus being favourable, many might be produced.”

  76. The evolution of Darwin's understanding of extinction is vividly spelled out in the two editions of the Voyage of the Beagle. In the first edition, the Journal of Researches into the Geology and Natural History of the Various Countries Visited by H. M. S. “Beagle” under the Command of Captain Fitz Roy, R. N., from 1832 to 1836 (London: Colburn, 1839), the MS of which was completed in 1837, before the opening of the transmutation notebooks, Darwin concludes his brief discussion of extinction with: “All that at present can be said with certainty, is that, as with the individual, so with the species, the hour of life has run its course, and is spent.” However, Darwin had written earlier in the discussion that “variations of climate and food, or introduction of enemies, or the increased numbers of other species, [are] the cause of the succession of races.” In the 2nd ed., the Journal of Researches into the Natural History of the Countries Visited during the Voyage of H. M. S. “Beagle” round the World (London: J. Murray, 1845), Darwin wrote, “Certainly no fact in the long history of the world is so startling as the wide and repeated extermination of its inhabitants.” The discussion on extinction is much longer here, and Darwin explicitly states his Malthusian insights and vividly expresses his commitments to allopatric speciation. He reiterates his strong commitment to a naturalistic explanation of extinction and mockingly suggests that we should not be surprised to find extinction preceded by very low population densities. His discussion echoes Paley's in Natural Theology: “to a patient at the close of his disease ... Death ... is only the last of a long train of changes; in his progress through which, it is possible that he may experience no shocks or sudden transitions” (The Works of William Paley, D. D. [Philadelphia: J. Woodward, 1831], p. 479). Darwin also stresses that extinction can be initiated by a chance event. It was clear to Darwin that his theory need not account for chance events such as the transport of a seed by winds or birds being carried by a storm to an island. They formed the initial conditions for the phenomena to be explained by the theory, which was to account for the regularities that occurred when a system was placed under specified circumstances. This point had been explicitly discussed in Herschel's Preliminary Discourse on Natural Philosophy (London: Lardner's Cabinet, 1831).

  77. Lyell's Principles of Geology (London, J. Murray), vol. I (1830), vol. II (1832), and especially vol. III (1833). See also Rudwick, Meaning of Fossils, and Rudwick, “Charles Lyell's Dream of a Statistical Palaeontology,” pp. 225, 244.

  78. See my discussion in “Origin of the Origin,” pp. 299–302. A typical entry of that period is the one Darwin wrote on October 4, 1838, in E, p. 4: “It cannot be objected to my theory, that the amount of change within historical times has been small — because change in form is solely adaptation of whole of one race to some change of circumstances; now we know how slowly & insensibly such changes are in progress — we feel interest in discovering a change of level of a few feet during the last two thousand years in Italy, but what change would such a change produce in climate vegetation &c — it is the circumstance of small physical changes & oscillations, not affecting organic forms, that the whole value of the geological chronology depends that most sublime discovery of the genus of men.” See also the section “On Time,” chap. 7 of Charles Babbage's Ninth Bridgewater Treatise (London: John Murray, 1838), which Darwin read in October 1838.

  79. E, p. 125: “No one but a practised geologist can really comprehend how old the world is, as the measurements refer not to revolutions of the sun & our lives, but to period necessary to form heap of pebbles &c &c: the succession of organism tells nothing about length of time, only orders of succession.”

  80. D, p. 140: “when we multiply the effects of earthquakes, elevating forces in raising continents, sea on beaches we really measure the rapidity of changes of forms & instincts in the animal kingdom.”

  81. The ecological concept of place is refered to in the notebooks, e.g., E, p. 114 (March 12, 1839): “we must recollect the multitude of plants introduced into our gardens ... which are propagated with very little care — & which might spread themselves as well as our wild plants, we see how full nature, how finely each holds its place.” Earlier, in his “Ornithological Notes,” which were very probably written during 1836 on the last leg of the Beagle voyage, Darwin had noted: “I have specimens [of Galápagos mockingbirds] from four of the larger islands ... The specimens from Chatham & Albermarle Isld appear to be the same; but the other two are different. In each Isld. each kind is exclusively found: habits of all are indistinguishable. When I recollect the fact that the form of the body, shape of scales & general size, the Spaniards can at once pronounce, from which Island and Tortoise may have been brought. When I see these Islands in sight of each other, & [but deleted] possessed of but a scanty stock of animals, tenanted by these birds, but slightly differing in structure & filling the same place [my italics] in Nature, I must suspect they are only varieties ... If there is the slightest foundations to these remarks the Zoology of Archipelagoes, will be well worth examining; for such facts [would inserted] undermine the stability of Species.” N. Barlow, ed., “Darwin's Ornithological Notes,” Bull. Brit. Mus. (Nat. Hist.), Hist. Ser., 2, no. 7 (1963), 203–262. For the dating of this passage see S. Herbert, “The Place of Man in the Development of Darwin's Theory of Transmutation, Part I, to July 1837,” J. Hist. Biol., 7 (1974), 217–258; G. Grinnell, “The Rise and Fall of Darwin's First Theory of Transmutation,” J. Hist. Biol., 7 (1974), 259–273; R. Colp, Charles Darwin and the Galápagos,” N.Y. J. Med., 77 (1977), 262–267.

  82. E, p. 57: “Every structure is capable of innumerable variations, as long as each shall be perfectly adapted to circumstances of times & from persisting owing to their slow formation these variations tend to accumulate on any structure.” It is interesting to note that strictly speaking the time scale for variation has meaning only for a population.

  83. Darwin to Hooker, Sunday, 1844. “The conclusion, which I have come at is that those areas, in which species are most numerous, have oftenest been divided and isolated from other areas, united and again divided; a process implying antiquity and some changes in the external conditions. This will justly sound very hypothetical. I cannot give my reasons in detail; but the most general conclusion, which the geographical distribution of all organic beings, appears to me to indicate is that isolation is the chief concomitant, or cause of the appearance of new forms (I well know there are some starring exceptions). Secondly from seeing how often the plants and animals swarm in country, when introduced in it, and from seeing what a vast number of plants will live, for instance in England if kept free from weeds, and native plants, I have been led to consider that the spreading and number of the organic beings of any country depend less on its external features, than on the number of forms, which have been there originally created or produced. I much doubt whether you will find it possible to explain the number of forms by proportional differences of exposure; and I cannot doubt if half the species in any country were destroyed or had not been created yet that country would appear to us fully peopled. With respect to original creation or production of new forms, I have said that isolation appears the chief element. Hence, with respect to terrestrial productions a tract of country, which had oftenest within the later geological periods subsided and been converted into islands and reunited, I should expect to contain most forms. “But such speculations are amusing only to one self and in this case useless as they do not show any direct line of observation. “Your geographical-law-letters require being read and reread, and I have only read your last twice, and so will hazard no remarks on it. You seem however to have put the case of ‘typical forms’ in a clearer point of view, than I ever saw it and stripped the word of half, if not all its mystery: I have long suspected, that typical and abnormal forms consist only of those, of which a greater or less variety have been created or modified-with this excellenty!! expressed sentence, I will conclude.” This letter is in the Hooker-Darwin correspondence of Cambridge University Library. I thank Peter Gautrey and the Librarian of the Library for permission to quote it. Part of the letter is included in LLD, I, 388–389.

  84. J. F. W. Herschel, “On the Astronomical Causes Which May Influence Geological Phenomena,” Trans. Geol. Soc. London, 3 (1932), 293–299. See also W. F. Cannon, “John Herschel and the Idea of Science,” J. Hist. Ideas, 22 (1961), 215–239; Rudwick, Meaning of Fossils, p. 187; Dov Ospovat, “Lyell's Theory of Climate,” J. Hist. Biol., 10 (1977), 317–399.

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  85. E, p. 52: “I from looking at all facts as inducing toward law of transmutation, cannot see the deductions which are possible.”

  86. See, e.g., E, p. 51: “Thinking of effects of my theory, laws will probably be discovered of correlation of parts, from the laws of variation of one part affecting another.” See also E, pp. 53–54 and p. 59: “Hurrah — ‘intermediate cause.’” In connection with the vera causa principle, see M. J. S. Hodge, “The Structure and Strategy of Darwin's Long Argument,” Brit. J. Hist. Science, 10 (1977), 237–246.

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  87. Charles Lyell, Principles of Geology, (London: John Murray, 1832). II, 130.

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  88. “Essay of 1844,” p. 118.

  89. In other words, the questions why do organisms reproduce so as to give rise to a geometrical rate of increase, and what the time constant is, are meaningful and must be answered by the theory of natural selection.

  90. See Schweber, “The Young Darwin,” on the relation between how natural selection operated and how the Newtonian description worked.

  91. The word tendency does not occur in Darwin's original MS. De Beer inserted it.

  92. I thank J. Hodge for a discussion of this point.

  93. Quoted in Stauffer, “Ecology in the Long Manuscript Version of Darwin's Origin of Species,” p. 237.

  94. See ibid., p. 238, for the use of the world place.

  95. See Mayr's stimulating 1974 essay, “The Evolution of Living Systems,” reprinted in Evolution and the Diversity of Life, pp. 16–25.

  96. In C, p. 147e Darwin says that the quantity of life on a planet depends on “subdivisions of stations & diversity.”

  97. Neither Herschel in his Preliminary Discourse nor Whewell in his History of the Inductive Sciences deals with these variational formulations of mechanics. I have not established whether Darwin knew about such principles in optics.

  98. HenryLord Brougham, Dissertations on Subjects of Sciences Connected with Natural Theology, 2 vols. (London: C. Knight, 1839).

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  99. Justus Liebig, Organic Chemistry in Its Application to Agriculture and Physiology, ed. Lyon Playfair (London: Taylor and Walton, 1840). Darwin's copy is at Cambridge University Library and is signed “Charles Darwin 1841” inside the front cover.

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  100. I shall discuss later the attempts to quantify and mathematize political economy. Here I remark only that Whewell was involved in such a project while Darwin was at Cambridge. See W. Whewell, “Mathematical Exposition of Some Doctrines of Political Economy,” Trans. Camb. Phil. Soc., 3 (1830), 191–230, and 4 (1833), 155–198.

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  101. Darwin's discussion of alpine plants in the “Sketch’ is on pp. 65–67. For an interesting discussion of biogeography written somewhat after the “Essay”, see Darwin's review of Natural History of the Mammalia, by G. R. Waterhouse, Ann. Mag. Nat. Hist., Including Zool., bot., Geol., 19 (1847), 53–56 which is reprinted in The Collected papers of Charles Darwin, ed. Paul Barrett, p. 214–217.

  102. In the “Sketch”, p. 74, Darwin further noted: “According to mere chance every species may generate another, but if any species A, in changing gets an advantage and that advantage (whatever it may be, intellect, etc., or some particular structure of constitution) is inherited, A will be the progenitor of several genera or even families in the hard struggle of nature. A will go on beating out other forms, it might come that A would people earth”.

  103. Quoted in Stauffer, “Darwin, Linnaeus, and Ecology”, from Charles Darwin, ed., The Zoology of The Voyage of H.M.S. “Beagle”, under the Command of Captain Fitz Roy, R. N., during the Years 1832 to 1836 (London: Smith, Elder).

  104. Quoted in Stauffer, “Darwin, Linnaeus and Ecology”, from C. Darwin, ed., The Structure and Distribution of Coral Reefs, Being the First Part of the Geology of the Voyage of the “Beagle” (London: Smith, Elder 1842), p. 63.

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  105. “Essay of 1844”, p. 142; my italics. In the “Fair Copy Annoted by C.D.”, which is in the Cambridge University Library, this entry occurs on pp. 154–155. The last line reads: “where they consequently hold a more strictly limited place against a far severer struggle”, with “against a far severer struggle” crossed out.

  106. Darwin, Journal of Researches, 2nd ed., 2 vols. (New York: Harpers, 1871), I, 225.

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  107. “Essay of 1844”, pp. 118–119.

  108. Darwin-Hooker correspondence, Cambridge University Library, box 150.

  109. See Kottler, “Darwin's Biological Species Concept and Theory of Geographic Speciation”. Note that Darwin had suggested an individualistic mechanism of inheritance several times before 1842 (e.g., blood theories in the early note-books) and that his gemmule theory was formulated in the 1840–1842 period. Mayr has noted that Darwin often used the term “variety” in two different senses: deviating individuals and deviating populations. It would be interesting to see whether one can correlate his usage with his beliefs in mechanisms of inheritance; that is, whether when he believed that he had an explanation of inheritance based on an individualistic mechanism, such as, gemmules, variety refers to deviating individuals, and whether when he had less faith in an individualistic mechanism, varieties refer to deviating populations.

  110. Darwin-Hooker corespondence at the Cambridge University Library, box 150. See also Browne, “Charles Darwin-Joseph Hooker Correspondence”.

  111. Humboldt, De Distributione Geographica Plantarum.

  112. L.von Buch, Description physique des isles Canaries, (Paris: F. G. Levrault, 1926). On p. 147, Von Buch quotes the following ratios of species to genera: continental North Africa, 1:4.2; Canaries, 1:1.46; St. Helena, 1:1.5. He also compares the numbers of orders on continents with those on islands.

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  113. Darwin-Hooker correspondence, Cambridge University Library, box 150.

  114. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), p. 112.

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  115. Darwin-Hooker correspondence, Cambridge University Library, box 150; undated letter, early 1844.

  116. Some of these letters are reprinted in LLD, pp. 382–389, and in More Letters, II, 402–422.

  117. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), pp. 53–54; see also pp. 326 ff.

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  118. Janet Browne has investigated the biostatistical materials in boxes 15 and 16 of the Darwin papers at Cambridge University Library. Included there are materials on “Common and Large Genera Presenting Most Varieties,” miscellaneous tables of species, etc., all important for understanding Darwin's views on variations, biogeography in the mid 1850s, and his final formulation of the principle of divergence. See Browne, “Darwin's Botanical Arithmetic and the Principle of Divergence, 1854–1858,” J. Hist. Biol., 13 (1980), 53–89; I thank Dr. Browne for allowing me to read this paper before publication. In this connection see also Fred Somkin, “The Contributions of Sir John Lubbock, F. R. S., to the Origin of Species: Some Annotations to Darwin,” Notes Rec. Roy. Soc., 17 (1962), 183–191.

  119. In broad outline the changes have been sketched in Limoges' La sélection naturelle and in Mayr's Evolution and the Diversity of Life, but it would be interesting to know the specific way in which Darwin's botanical research during the 1840s affected his views on speciation.

  120. See, e.g., Thomas H. Huxley's comments on pp. 347–348 of his Life and Letters and p. 315 of LLD for Hooker's thoughts.

  121. Ghiselin, The Triumph of the Darwinian Method, pp. 103–105, 111, 117, 128–129. See also De Beer, Evolution by Natural Selection, pp. 135–156; Sydney Smith, “The Darwin Collection at Cambridge with One Example of Its Use: Charles Darwin and Cirripedes,” Actes du XI e congrès International d'Histoire des Sciences, 15 (1964), 96–100; Thaddeus J. Trenn, “Charles Darwin, Fossil Cirripedes and Robert Fitch, Presenting Sixteen Hitherto Unpublished Darwin Letters of 1849 to 1851,” Proc. Amer. Phil. Soc., 118 (1974), 471–491; A. E. Gunther, “J. E. Gray, Charles Darwin, and Cirrepedes,” Notes Rec. Roy. Soc., 34 (1979), 53–63.

  122. For the place of barnacles in nineteenth-century taxonomy, see Mary P. Winsor, “Barnacle Larvae in the Nineteenth Century: A Case Study in Taxonomic Theory, J. Hist. Med. Allied Sci., 29 (1969), 294–309, and Winsor, Starfish, Jellyfish, and the Order of Life.

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  123. R. Owen, Lectures on the Comparative Anatomy and Physiology of the Inverterbrate Animals (London, 1843). See also Owen's report to the 1846 meeting of the BAAS and its enlarged version, On the Archetype and Homologies of the Verterbrate Skeleton (London, 1848); Merz, History of European Thought, II, 258–259; and Roy M. MacLeod, “Evolutionism and Richard Owen, 1830–1868: An Episode in Darwin's Century,” Isis, 56 (1965), 259–280.

  124. The association of cirripedes with the problem of the evolution of sex dates at least to the 1838 period: “The sexual system of the Cirripedes is the more remarkable from their alliance to Articulata which are truly bisexual” (E, p. 60). See also E, pp. 71, 80, and especially 155: “My theory only requires that organic beings propagated by gemmation do not undergo metamorphosis, but to arrive at their present structure they must have been propagated by sexual commerce... The relation of the inference from some plants & some mollusca being hermaphrodite is, that intercourse every time is of no consequence in that degree of development. It is singular there is not true hermaphrodite on beings which have fluid sperma.” Darwin believed that hermaphroditic species eventually evolve into bisexual ones.

  125. What W. S. MacLeay had said of Cuvier in his Horae Eatomological; or, Essays on the Annulose Animals (London: A. Bagster 1819–1821) — “For skill in anatomical dissection, for accuracy of observation, and reference of means to their respective ends, perhaps no man living can be placed in competition with M. Cuvier ... it has been too often and too justly remarked, that no person of such transcendent talents and ingenuity ever made so little use of his observations towards a natural arrangement as M. Cuvier” (p. 326) — may well have struck a resonant chord in Darwin. Darwin had carefully studied MacLeay in the 1837–1839 period, and from a letter to Henslow in 1832 it is evident that he knew of MacLeay's work while on the Beagle. See Darwin and Henslow, p. 62, particularly Barlow's note 2. For the relevance of MacLeay's work to Darwin's cirripedes research, see Smith, “Darwin and Cirripedes”; barnacles held a central position in MacLeay's classification system. How influential MacLeay was is indicated by James Dwight Dana, Crustacea: US Exploring Expedition under the Command of Charles Wilkes, USN, 2 vols. (Philadelphia, 1852–1853). Darwin read this book on Sept. 20, 1853, and must have come across on p. 54 Dana's comments: “There are neither straight lines nor circles in nature, but main branching lines, with subordinate branches, and almost endless reticulations or anastomoses, by curves of all kinds and all grades of divergence and convergence.” Dana was commenting on the numerical complexities of the circular classificatory system of MacLeay, Swainson, and others, and indicated that it “was a brilliant scheme when first brought forward ... but as the first glare has now passed, we can perceive that while it attempted to rid science of the straight and rigid bars of artificial systems, it only modified the mode of coercion by bending bars into circles.” See also Winsor, “Starfish, Jellyfish, and the Order of Life”; and P. L. Heilbroner, “Circular Logic: The Context, Structure, and Influence of William Sharp MacLeay's Quinarism,” thesis, Dept. of the History of Science, Harvard University, 1976.

  126. J. B. Lamarck, Philosophie zoologique ou expositions des considérations relatives à l'histoire naturelle des animaux (Paris, 1809).

  127. G. Cuvier, Le règne animal d'après son organisation, 4 vols. (Paris, 1817).

  128. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), p. 440.

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  129. Quoted in W. Coleman, Georges Cuvier, Zoologist: A Study in the History of Evolution Theory (Cambridge, Mass.: Harvard University Press, 1964), p. 143.

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  130. “Essay of 1844,” p. 114.

  131. Ibid., p. 247.

  132. LLD, I, 397.

  133. Charles Darwin, A Monograph on the Sub-Class Cirripedia: The Balanidae (London: Ray Society, 1854), p. 155.

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  134. Charles Darwin, A Monograph on the Sub-Class Cirripedia: The Balanidae (London: Ray Society, 1854), p. XI.

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  135. Autobiography, LLD, p. 66.

  136. See the Preface in Darwin's Cirripedia: The Lepadidae, where the thanks those who had sent him specimens. Some of the letters requesting barnacle specimens are at the American Philosophical Society. See, e.g., letters 71 and 72 to Gray and no. 76 to Milne-Edwards, as quoted in P. T. Carroll, ed., An Annotated Calender of the Letters of Charles Darwin in the Library of the American Philosophical Society (Wilmington, Del.: Scholarly Resources, 1976).

  137. Covington became Darwin's attendant on the voyage of the Beagle, and his clerk and anamuensis after their return to England. Covington later settled in New South Wales, Australia, and sent Darwin one of the best collections of barnacles. “I have received a vast number of collections from different places, but never one so rich from one locality,” Darwin wrote Covington; see Gavin de Beer, ed., “Some Unpublished Letters of Charles Darwin,” Notes Rec. Roy. Soc. London, 13 (1958), 19.

  138. Darwin was familiar with Gould's book the Cicindelidae of Massachusetts (Boston, 1833) and his impressive conchological work “Results of an Examination of the Species of Shells of Mass. and Their Geographical Distribution,” Boston J. Nat. Hist., 3 (1840), 483–494. Darwin had studied vols. I to IV (“entire”) of the Boston Journal on May 30, 1847, and earlier (on May 31, 1845) had read vol. IV “p. 377 to end.” See P. J. Vorzimmer, “The Darwin Reading Notebooks,” J. Hist. Biol., 10 (1977), 107–153.

  139. This unpublished letter is in the Houghton Library of Harvard University, and is quoted here by permission of W. H. Bond, the librarian.

  140. This letter, dated Sept. 1, 1848, is no. 76 in the Annotated Calendar of the Letters of Charles Darwin.

  141. This letter is in the Houghton Library of Harvard University, and is quoted here by permission.

  142. Contrast Darwin's restrained presentation (admittedly for a “professional” audience) with Huxley's and Owen's presentation of their views to lay audiences. T. H. Huxley, in his famous lecture “On the Method of Studying Zoology,” delivered to the science classes at the South Kensighton Museum in the early 1850s, said, “Unity of plant everywhere lies hidden under the mask of diversity of structure—the complex is everywhere evolved of the simple.” Richard Owen, aswering questions in 1851 before the English Public School Commission, which was seeking adivce on curricular changes, said that “the fundamental principles of classification in natural history are as certain [as those of mathematics] ... There is neither fluctuation nor speculation [in natural history]. The principles of natural history are already as settled and fixed as can be needed for its use as a disciplinary science.” Both these presentations are reprinted in E. L. Youmans, ed., The Culture Demanded by Modern Life: A Series of Addresses and Arguments on the Claims of Scientific Education (New York: Appelton, 1897).

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  143. Ghiselin, The Triumph of the Darwinian Method, pp. 103–130.

  144. M. T. Ghiselin and Linda Jaffee, “Phylogenetic Classification in Darwin's Monograph on the Sub-Class Cirripedia,” Syst. Zool., 22 (1973), 132–140.

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  145. Barrett in his “Concordance” to the notebooks lists entries for Milne-Edwards. For an early entry see B, p. 112; for one after Darwin read Malthus, see E, p. 25.

  146. H. Milne-Edwards, “Considerations sur quelques principes relatifs à la classification naturelle des animaux,” Ann. Sci. Nat., 3rd ser. (1844), 1, 66–69. See also H. Milne-Edwards, “Rapport sur une serie de memoires de M. A. de Quatrefages relatifs à l'organisation des animaux sans vertebres,” Ann. Sci. Nat., 3rd. ser. (1844), 5–9.

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  147. In Vorzimmer, “The Darwin Reading Notebooks,” the following readings are indicated: “Jan. 30, 1847 — M. Edwards Geog. Distribution of Crustaceae, 3d Tom of Suite [?] de Buffon; Nov. 26, 1852 — M. Edwards Introduct. Zoolog. Gener. 1851.” Both monographs on the Cirripedia give ample further proof of Darwin's careful study of Milne-Edwards' published works.

  148. In a letter to Milne-Edwards dated Sept. 1, 1848, Darwin thanks him for his kindness at the Oxford BAAS meeting in June 1847. See letter 76 in Annotated Calendar of the Letters of Charles Darwin.

  149. See the Athenaeum, 1143 (22 Sept. 22, 1849), 966. This exchange is reprinted in P. H. Barrett, ed., The Collected Papers of Charles Darwin, 2 vols. (Chicago: University of Chicago Press, 1977), I, 250–251. The standing of Milne-Edwards in the scientific community is indicated by the following passage in the “Transactions of Section D, Natural History, Including Physiology,” of the BAAS report for 1849: “The comments made by the emminent naturalist H. Milne-Edwards, Member of the Academie des Sciences in Paris on this communication [M. Barrandi's paper on the metamorphosis of certain recently discovered trilobites] must have so much weight, that a deviation is made from the ordinary practice in giving this abstract of them in a note” — Milne-Edwards' comments are then given.

  150. Dictionnaire classique d'histoire naturelle; “Organisation” is in vol. XII (1827), pp. 332–344; “Nerfs” is in vol. XI (1827), pp. 529–534.

  151. Darwin's copy of the Dictionnaire classique is now at Down House. Mr. Titheradge, the curator of Down House, informs me that the entries “Organisation” and “Nerfs” bear no annotations.

  152. That Darwin was reading the Dictionnaire classique is clear from his letters to Henslow. On May 18, 1832, for instance, he wrote: “I am well off in books. The Dic. Class. is most useful” (Barlow, ed., Darwin and Henslow, p. 54; see also letter no. 22, from Henslow to Darwin). Darwin's “Diary of Observations on Zoology of the Places Visited during the Voyage of the H.M.S. Beagle” is kept in boxes 30, 31, and 32 of Darwin's Papers at the Cambridge University Library. The early observations are primarily on invertebrates.

  153. H. Milne-Edwards: Elemens de zoologie: leçons sur l'anatomie, la physiologie, la classification des moeurs des animaux (Paris: Crochard, 1834), Darwin did not own this book. It is not listed in Rutherford's Catalogue of the Library of Charles Darwin in the Botany School (Cambridge: Cambridge University Press, 1908). In a catalogue of the Victor Masson Books (Anciènne Maison Crochard) that Darwin owned, which now is at Cambridge University Library, Darwin checked off the Milne-Edwards book which he owned, but not Elemens de zoologie.

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  154. H. Milne-Edwards: Elemens de zoologie: leçons sur l'anatomie, la physiologie, la classification des moeurs des animaux (Paris: Crochard, 1834), I, 8.

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  155. H. Milne-Edwards, Histoire naturelle des crustacées comprenant l'anatomie, la physiologie, et la classification de ces animaux, 3 vols. (Paris: Librairie Encyclopedique de Roret, 1834–1840). Vol I appeared in 1834, vol. II in 1837, and vol. III in 1840. Darwin's annoted copies are at Cambridge University Library.

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  156. H. Milne-Edwards, Histoire naturelle des crustacées comprenant l'anatomie, la physiologie, et la classification de ces animaux, 3 vols. (Paris: Librairie Encyclopedique de Roret, 1834–1840), I, 6.

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  157. H. Milne-Edwards, Histoire naturelle des crustacées comprenant l'anatomie, la physiologie, et la classification de ces animaux, 3 vols. (Paris: Librairie Encyclopedique de Roret, 1834–1840), p. 555.

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  163. Limoges, La sélection naturelle and “Darwin, Milne-Edwards, et le principle de divergence.”

  164. On Erasmus Darwin, see C. Darwin's introduction in Ernst Krause, ed., The Life of Erasmus Darwin Together with an Essay on His Scientific Works (New York: D. Appelton, 1880); see also Gruber's “Essay on a Psychological Study of Scientific Creativity,” in Darwin on Man; R. E. Colp, To Be an Invalid: The Illness of Charles Darwin (Chicago: University of Chicago Press, 1977), chap. 1; Desmond King-Hale, Erasmus Darwin (London: Macmillan, 1963); and Hesketh Pearson, Doctor Darwin (London: Dent, 1930). For Robert W. Darwin, see Eliza Meteyard, A Group of Englishmen (1895 to 1915), Being Records of the Younger Wedgwoods and Their Friends (London: Longmans, Green, 1871); “where religious creeds and political opinions were concerned, Meteyard wrote, the inhabitants of Shrewsbury “were alike narrow and bigoted to excess” (p. 257).

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  165. Meteyard, A Group of Englishmen, p. 265.

  166. J. C. Wedgwood, A History of the Wedgwood Family (London: St. Catherine Press, 1909); E. Meteyard, The Life of Josiah Wedgwood, 2 vols. (London, 1865–1866). For a brief introduction to this period, see Gruber and Barrett, Darwin on Man, pp. 17–33; for another excellent study, see J. D. Y. Peel, Herbert Spencer: The Evolution of a Sociologist (New York: Basic Books, 1971), esp. chap. 2 on Erasmus Darwin.

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  168. See, e.g., W. J. Bate, Coleridge (New York: Macmillan, 1968), pp. 85–88, 126–127. Apparently, Darwin took a special interest in the group of “radicals” who were close to the younger Josiah Wedgwood at the beginning of the century — Godwin, Southey, Wordsworth, Coleridge. Darwin was reading Godwin and Shelley in 1841 and 1842; and his reading notebook for that period records: “1841 — July 23, Godwin's Answer to Malthus (Shelley says is victorious and decisive). December 4, Godwin on Population. 1842 — September 26, Shelley's Letters and Essays.” His liking of the poetry of Wordsworth is well known. In 1848 Darwin commented in his reading notebook: “Cottle's Recollection of Coleridge & Southey (very good).”

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  169. Darwin, Autobiography, LLD, pp. 38–39: “I was also attached to and greatly revered my uncle Jos; he has silent and reserved ... but he sometimes talked openly with me. He was the very type of an upright man, with the clearest judgment. I do not believe that any power on earth could have made him swerve an inch from what he considered the right course.”

  170. Ibid., p. 38.

  171. Ibid.

  172. Vorzimmer, “The Darwin Reading Notebooks,” pp. 125, 134, 136. James Mackintosh, Dissertation on the Progress of Ethical Philosophy Chiefly during the Seventeenth and Eighteenth Century, 2nd ed. (Edinburgh: Black, 1836); R. J. Mackintosh, ed., Memoirs of the Life of Sir James Mackintosh, 2 vols. (London: E. Maxon, 1835); R. J. Mackintosh, ed., The Miscellaneous Works of Sir James Mackintosh, 3 vols. (London: Longman, Brown, 1846). James Mackintosh's History of England from Earliest Times to the Year 1588 (Lardner's Cabinet Encyclopedia 1830) may have been aboard the Beagle. Fitz Roy and Darwin quote from it in “The Moral State of Tahiti,” South African Christian Recorder, 2 (1836), 221–238. See Barrett, ed. Collected Papers of Charles Darwin, I, 19–38. See also E. Manier, The Young Darwin and His Cultural Circle (Dordrecht: D. Reidel, 1978).

  173. G. M. Young, Victorian England: Portrait of an Age (Oxford: Oxford University Press, 1953), p. 6.

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  174. Francis Gled Stanes Waugh, Members of the Athenaeum Club, 1824 to 1887, (privately printed).

  175. Life of Lyell, I, 356.

  176. Ibid., p. 322.

  177. Ibid., p. 264.

  178. Ibid., p. 44.

  179. Ibid., p. 356. In this connection see also M. Rudwick, “Poulett Scrope on the Volcanoes of Auvergne: Lyellian Time and Political Economy,” Brit. J. Hist. Sci., 7 (1974), 205–242.

  180. See Gavin de Beer, ed., “Darwin's Journal,” Bull. Brit. Mus. (Nat. Hist.), Hist. Ser., 2, no. 1 (1959), for a record of Darwin's attendance at the various BAAS meetings.

  181. See, e.g., the letter Darwin wrote from the 1849 Birmingham meeting of the BAAS, LLD, I, 346–347.

  182. John Herschel, A Manual of Scientific Inquiry (London: John Murray, 1849).

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  184. R. E. Colp, “Charles Darwin: Slavery and the American Civil War,” Harvard Lib. Bull., 26, no. 4 (1978), 471–489.

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  185. LLD, I, 309–310. See also Darwin's impassioned statement in the 2nd edition of his Journal of Researches, II, 302–304. Darwin's view of the American Civil War is recorded in Dupree, Asa Gray. By the end of the conflict, Darwin was the only British scientist to whom Gray could write about his antislavery views. See also Colp, “Charles Darwin: Slavery and the American Civil War,” pp. 471–489.

  186. Barlow, ed., Darwin and Henslow, pp. 156–157.

  187. Not until the early 1830s did Paley and his Utilitarianism fall into disrepute among the clerics at Cambridge, See, e.g., the sharp attack on Paley in Adam Sedgwick, A Discourse on the Studies of the University (Cambridge: Cambridge University Press 1833). J. S. Mill's rebuttal is reprinted is Dissertations and Discussions (Boston: W. V. Spencer, 1864), pp. 121–185.

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  188. Darwin, Autobiography, LLD, pp. 40–41. In 1859, writing to John Lubbock, Darwin noted: “I do not think I hardly ever admired a book more than Paley's Natural Theology. I could almost formally have said it by heart” (LLD, II, 219).

  189. The Works of William Paley, Moral and Political Philosophy, pp. 149–150.

  190. Ibid., p. 150.

  191. Ibid., p. 156.

  192. See for examples the N and the M notebooks, transcribed in Gruber and Barrett, Darwin on Man. See also E. Manier, The Young Darwin and His Cultural Circle (Dordrecht: D. Reidel, 1978).

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  194. The Works of William Paley, pp. 149 f.

  195. For Stewart's “Life of Adam Smith,” see Stewart's edition of Smith's Essays on Philosophic Subjects (London: T. Cadell and W. Davis, 1795); Smith's Theory of Moral Sentiments was published in 1759 (Edinburgh, A. Millar).

  196. Adam Smith, The Wealth of Nations (New York: Modern Library, 1937), chap. 2.

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  197. J. R. McCulloch, The Principles of Political Economy with a Sketch of the Rise and Progress of the Science, 2nd edition (London: Longman's, 1830). The reading notebook indicates that Darwin also read Mandeville's Fable of the Bees in May 1840 and abstracted it. For a recent assessment of McCulloch, see D. P. O'Brien, J. R. McCulloch: A Study in Classical Economics (London: George Allen & Unwin, 1970). See also Louis Dumont, From Mandeville to Marx (Chicago: University of Chicago Press, 1977).

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  198. J. R. McCulloch, The Principles of Political Economy with a Sketch of the Rise and Progress of the Science, 2nd edition (London: Longman's, 1830), p. 15.

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  199. J. R. McCulloch, The Principles of Political Economy with a Sketch of the Rise and Progress of the Science, 2nd edition (London: Longman's, 1830), p. 16.

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  201. A. Keith, Darwin Revalued (London: Watts, 1955), particularly chap. 18. Of course other Englishmen did likewise without having read McCulloch. The point is that McCulloch reflected a viewpoint that the elder and the younger Wedgwood had acted upon earlier in the century. See, e.g., the Correspondence of Josiah Wedgwood, 1781–1794 with an appendix containing some letters on canals and Bentley's pamphlet on island navigation. (Manchester: E. J. Morten, 1906). See also G. R. Porter, The Progress of the Nation, in Its Various Social and Economical Relations (London: Charles Knight, 1838), sections III and VI in particular, from which the following is taken: “There is not any circumstance connected with the internal condition of England which more strongly excites the admiration and the envy of foreigners than the degree of perfection to which we have brought our means of internal communication. The skill and labour that have been applied to this object are among the chief causes of that high degree of activity which characterizes and pervades the productive classes in every part of the country.”

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  202. Halévy, The Rise of Philosophical Radicalism.

  203. J. R. McCulloch, The Principles of Political Economy with a Sketch of the Rise and Progress of the Science, 2nd edition (London: Longman's, 1830), p. 149.

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  204. J. R. McCulloch, The Principles of Political Economy with a Sketch of the Rise and Progress of the Science, 2nd edition (London: Longman's, 1830), pp. 536–537.

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  205. Francis Palgrave, “Life and Works of Sismondi,” Quart. Rev., 72 (1843), 299–356. See also E. Halévy, Sismondi (Paris: Librairie Felix Alcan, 1935).

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  206. J. C. L.de Sismondi, Political Economy and the Philosophy of Government with a historical notice of his life and writings by M. Mignet (London: John Chapman, 1847).

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  207. Sismondi, preface to New Principles of Political Economy and the light Which They May Cast on the Crisis Which England Is at This Time Experiencing, reprinted in Sismondi, Political Economy.

  208. J. C. L.de Sismondi, Political Economy and the Philosophy of Government (London: John Chapman, 1847), p. 72.

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  209. In Darwin Revalued Keith says that Darwin purchased the farm of Beesby in Lincolnshire as an investment in 1845. His father advanced him £ 3,529 for the purchase. See also LLD, I, 311–312.

  210. C. Babbage, On the Economy of Machinery and Manufactures (London: J. Murray, 1832).

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  211. C. Babbage, On the Economy of Machinery and Manufactures, 4th ed. enlarged (London: J. Murray, 1846), pp. 175–176.

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  212. Liebig's Organic Chemistry was published in 1840.

  213. Darwin's copy is in the Cambridge University Library. Most of the annotations were made in 1841. Upon rereading the book in 1856, the only fact which warranted a note pinned in the back of the book relates to the fact that the blood of the different races of man have different odour.

  214. Darwin's growing interest in botany is attested by box 49 of Darwin's papers at Cambridge University Library. For a delightful overview of Darwin's botanical works, see M. Allen, Darwin and His Flowers: The Key to Natural Selection (New York: Taplinger, 1977).

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  215. Liebig, Organic Chemistry, preface.

  216. Ibid., pp. 129–130.

  217. J. Liebig, Familiar Letters on Chemistry and Its Relation to Commerce, Physiology, and Agriculture (New York: Appelton, 1843). Darwin read this book on Nov. 20, 1844, and Oct. 14, 1851, according to his reading notebooks; see Vorzimmer, “The Darwin Reading Notebooks.” On Oct. 3, 1851, while in London visiting the Great Exhibition, Darwin wrote to his son William, “I am reading a Book on Chemistry called Familiar Letters on Chemistry and this makes me often think of you in the evenings.“ Liebig's Familiar Letters was very popular and went through several editions. The fourth edition had as its title Familiar Letters on Chemistry in Its Relation to Physiology, Dietetics, Agriculture, Commerce, and Political Economy (London: Walton and Maberly, 1859). It is very likely that Darwin read two different editions in 1843 and 1851.

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  218. J. Liebig, Familiar Letters on Chemistry and Its Relation to Commerce, Physiology, and Agriculture (New York: Appelton, 1843). p. 1.

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  219. J. Liebig, Familiar Letters on Chemistry and Its Relation to Commerce, Physiology, and Agriculture (New York: Appelton, 1843) p. 8.

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  220. Quoted in Asa Briggs, Victorian People: A Reassessment of Persons and Themes, 1851–67, rev. ed. (Chicago: University of Chicago Press, 1972), p. 16. The first essay in this book is “Crystal Palace and the Men of 1851.”

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  221. David Brewster's review of Charles Babbage, The Exposition of 1851 or, Views of the Industry, the Science, and the Government of England, London 1851, 2nd ed., appeared in the North Brit. Rev., 15 (1851) 529. C. Babbage's The Exposition of 1851 (London: J. Murray, 1851) was also reviewed in the June 14, 1851, issue of the Atheneaum and in many other Victorian periodicals.

  222. Prince Albert's speech was delivered at a banquet at the Mansion House on March 21, 1850. It is quoted in J. B. Bury, The Idea of Progress (London: Macmillan, 1932), p. 330. See also T. Martin, The Life of the Prince Consort, 5, vols. (London: Smith, Elder, 1975–1980), III, 247. For an account of Prince Albert's involvement in the Great Exhibition and its relation to his artistic interests, see R. Fulford, The Prince Consort (London: Macmillan, 1966), particularly pp. 203–225; and W. Ames, Prince Albert and Victorian Taste (New York: Viking Press, 1968).

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  223. Darwin's daughter recalled that “on July 30th [1851], my father and mother spent a week with Erasmus Darwin at his house in Park Street in order to see the Exhibition. George and I were also taken, but I at any rate did not make much of it, and remember staying at home to scrub the back stairs, as better fun ... My father enjoyed it immensely”; Emma Darwin, ed., A Century of Family Letters, 2 vols. (privately printed, 1904), II, 153. The public edition was published by J. Murray (London, 1915).

  224. See G. Holton, Thematic Origins of Scientific Thought: Kepler to Einstein (Cambridge, Mass.: Harvard University Press, 1973); and Holton, The Scientific Imagination: Case Studies (Cambridge: Cambridge University Press, 1978).

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  227. Charles Darwin, The Foundations of the “Origin of Species”: Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909) p. 27. The notion of a workshop may well have had its origin in Darwin's translation of Milne-Edwards' “Ateliers,” in the latter's discussion of the division of labor.

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  228. Of the many books on the subject, see, e.g., J. B. Bury, The Idea of Progress (London: Macmillan, 1932); F. E.Manuel, Shapes of Philosophical History (Stanford: Stanford University Press, 1968); and M. Mandelbaum, History, Man, and Reason (Baltimore: Johns Hopkins Press, 1971). I thank Ernst Mayr for calling Mandelbaum's book to my attention.

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  229. M. Foucault, Les mots et les choses (Paris: Gallimard, 1966).

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  230. See, e.g., the chapter on Condorcet in F. Manuel, The Prophets of Paris (Cambridge: Harvard University Press, 1962), and the introduction by F. Manuel to J. C. Herder, Reflections on the Philosophy of the History of Mankind (Chicago: University of Chicago Press, 1968).

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  231. “Statistics” had many different seses, ranging from the mere compilation of data to the handling of experimental data, but I shall not elaborate here on these differences. See Merz, History of European Thought, vol. II, the chapter “The Statistical View of Nature”.

  232. See Philip Abrams, The Origins of British Sociology, 1834–1914: An Essay with Selected Papers (Chicago: University of Chicago Press, 1968); M. J. Cullen, The Statistical Movement in Early Victorian Britain (Hassocks: Harvester Press, 1975); Victor L. Hilts, “Aliis Extererendum, or the Origins of the Statistical Society of London,” Isis, 69 (1978), 21–43.

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  233. The tradition, of course, goes back to Adam Smith and to the Physiocrats. One of the aspects in which Adam Smith differs from Senior, Jones, and McCulloch is in his view of the “objectivity” of political economy. The later political economists were at greater pains to separate political actions from political economy, and to clarify the relation of political economy to legislative political acts. Adam Smith also thought of each human being as at birth almost exactly the same in character and capacities as every other being: for him, it was the division of labor that alters characters. The later economists accepted greater variability and differences in people.

  234. For an account of the formation of the Statistical Section of the BAAS, see Hilts, “Aliis Extererendum”; Cullen, The Statistical Movement; and C. Babbage, Exposition of 1851, 2nd ed. (London: J. Murray, 1851).

  235. W. Whewell, Report of the Ninth Meeting of the British Association for the Advancement of Science, 3 (1833), xc-xci.

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  236. Rudwick, “Charles Lyell's Dream of a Statistical Palaeontology,” pp. 225–244.

  237. Janet Browne in an Imperial College Ph.D. dissertation (1978) has investigated phytogeography in the early nineteenth century.

  238. I have not compared the membership of the Political Economy Club and that of the Geological Society. See the centennial publication of the Political Economy Club: Political Economy Club Founded in London 1821, Minutes of Proceedings, 1899–1920: Roll of Members and Questions Discussed, 1821–1920, with Documents Bearing on the History of the Club, 6 vols. (London: Macmillan, 1900); M. J. Rudwick, “The Foundation of the Geological Society of London: Its Scheme for Co-operative Research and Its Struggle for Independence,” Brit. J. Hist. Sci., 1 (1963), 325–355; and J. B. Morrell, “London Institutions and Lyell's Career, 1820–1841,” Brit. J. Hist. Sci., 9 (1976), 132–146.

  239. Literary Remains Consisting of Lectures and Tracts on Political Economy of the Late Rev. Richard Jones, ed. William Whewell (London: John Murray, 1859), pp. 537–539; reprinted in 1964 by Augustus M. Kelley, New York.

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  240. J. Herschel, Preliminary Discourse on Natural Philosophy (London: Lardner's Cabinet, 1831).

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  241. Life Lyell, II, 37–39.

  242. See Frank W. Fetter, “Economic Controversy in the British Review, 1802–1850,” Economica, 0 (1965), 424–437; John Leonard Clive, Scotch Reviewers: The Edinburgh Review, 1802–1815 (London: Faber and Faber, 1957); W. Graham, English Literary Periodicals (New York: Thomas Nelson, 1930); and Michael Wolff, “Victorian Reviewers and Cultural Responsibility,” in P. Appleton, W. A. Madden, and M. Wolff, ed., Eighteen Fifty-Nine: Entering and Age of Crisis (Bloomington: Indiana University Press, 1959), pp. 269–289.

  243. Incidentally, G. H. Lewes and Herbert Spencer were also struck in 1852 by Milne-Edwards' concept of the physiological division of labor. In his Autobiography, Spencer writes: “Lewes... had brought with him a volume by Milne-Edwards, and in it for the first time I met with the expression—‘the physiological division of labour’. Though the conception was not new to me, as is shown towards the end of Social Statics, yet the mode of formulating it was; and the phrase thereafter played a part in my course of thought”; Herbert Spencer, An Autobiography, 2 vols. (New York: Appelton, 1904), I, 436–437. See also J. A. Thomson, Herbert Spencer (New York: E. P. Dutton, 1906), p. 30. In his Sea Side Studies (Edinburgh: Blackwood, 1890), p. 408, G. H. Lewes pointed out that the conception is to be found in Goethe, Zur Morphology (1807): “the French naturalist [Milne-Edwards] having the merit of application and abundant illustration of the law.” For an informative and insightful discussion of the background of Spencer's evolutionary theory, see J. D. Y. Peel, Herbert Spencer: The Evolution of a Sociologist (New York: Basic Books, 1971).

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  244. J. B. Say, in turn, was influenced by A. P. de Candolle; see Candolle's Mémoires et souvenirs de A. P. de Candolle, (Genèva, Paris: J. Cherbuliez, 1862). See also J. E. E. D. A. Acton, Historical Essays and Studies, ed. J. N. Figgis and R. V. Laurence (Freeport, N. Y.: Books for Library Presses, 1967).

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  246. Except for articles by Sismondi and Say, primarily in the Revue Encyclopedique, only the Revue Universelle in Geneva carried articles in French comparable to those found in the Victorian periodicals. Thomas Sowell, Say's Law: A Historical Analysis (Princeton: Princeton University Press, 1972), which chronicles the debate over Say's law in the English and French periodicals.

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Schweber, S.S. Darwin and the political economists: Divergence of character. J Hist Biol 13, 195–289 (1980). https://doi.org/10.1007/BF00125744

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