¹ Darwin, The Origin of Species (1876), ed. Barrett, Paul H. and Freeman, R. B. (London: William Pickering, 1886–1889), 16.182-4Google Scholar, 209, 366.

² Op. cit., Origin (1876), 16.2.

³ Cannon, H. Graham, Lamarck and Modern Genetics (Manchester: Manchester University Press, 1959), 35–36Google Scholar.

⁴ Op. cit., Origin (1876), 16.8. Similarly in The Descent of Man and Selection in Relation to Sex, ed. Barrett and Freeman (London: Pickering, 1889) 21.57Google Scholar, Darwin asks not whether but to what extent evolution is ruled by ‘the inherited effects of the increased use of certain parts’; cf. 21.65.

⁵ Op. cit., Origin (1876), 16.8–9. For convenience we include in square brackets the corresponding pages in The Origin of Species (New York: Random House, 1993) [29].

⁶ Op. cit., Origin (1876), 16.114 [175].

⁷ Op. cit., Origin (1876), 16.118–20 [181–83], where Darwin calls it ‘an obscure question’ how much of the ability of a plant species to acclimatize to a new setting ‘is due to mere habit, and how much to the natural selection of varieties having different innate constitutions.’

⁸ Op. cit., Origin (1876), 16.9 [29]. Darwin, of course, was as innocent of the role of hormones in gene expression as he was of the existence of genes and chromosomes. But the experience of fox breeders in Siberia has shown that selective breeding can yield floppy ears and even docile tempers in a captive population. Use and disuse, however, are not the cause.

⁹ Op. cit., Origin (1876), 16.116–7 [178–79].

¹⁰ Op. cit., Origin (1876), 16.114 [175–76].

¹¹ Op. cit., Descent, 21.37.

¹² Op. cit., Origin (1876), 16.220.

¹³ Op. cit., Origin (1876), 16.243.

¹⁴ Op. cit., Origin (1876), 16.182–4 [272–75].

¹⁵ For Darwin's argument from facial expression, see The Expression of the Emotions in Man and Animals (London: Murray, 1872)Google Scholar, supplementing the claims of The Descent of Man, published the previous year.

¹⁶ Kelvin had thrown all his authority as the master of the physics of heat into showing that neither the earth nor the sun could be old enough to give evolution the time needed for natural selection to do its work. It was not until fusion was understood to be the source of the sun's energy and radioactive decay within the earth's core was understood to be the source of our planet's residual heat that the age parameters of the solar system expanded broadly enough to allow the time span evolution needed. One visible effect of the impact of Kelvin's challenge on Darwin's work was Darwin's omission of a time scale from his graphic representation of the Tree of Life in Chapter 4 of the Origin. For Kelvin's challenge and its resolution, see Goodman, Creation and Evolution (London and New York: Routledge, 2010), 102–03Google Scholar.

¹⁷ Gould, Structure, 69.

¹⁸ Op. cit., Origin (1876), 16.10 [31].

¹⁹ Op. cit., Origin (1876), 16.32 [63].

²⁰ Op. cit., Origin (1876), 16.15 [39].

²¹ Op. cit., Origin (1876), 16.8 [28].

²² Op. cit., Origin (1876), 16.36 [68].

²³ Op. cit., Origin (1876), 16.112 [172]; in rounding out his argument, Darwin spoke of variability under conditions of domestication as ‘caused, or at least excited, by changed conditions of life, but often in so obscure a manner, that we are tempted to consider the variation as spontaneous’ Origin (1876), 16.426.

²⁴ Op. cit., Descent, 21.34–5.

²⁵ Op. cit., Descent, 21.36–7; cf. 21.92.

²⁶ Op. cit., Descent, 21.37–8.

²⁷ Op. cit., Descent, 22.237–9. Darwin included a full chapter (27) on what he called ‘the provisional hypothesis of pangenesis’ in his 1875 work Variations of Animals and Plants under Domestication: ‘I venture to advance the hypothesis of Pangenesis, which implies that every separate part of the whole organization reproduces itself. So that ovules, spermatozoa, and pollen grains, the fertilized egg or seed, as well as buds include and consist of a multitude of germs thrown off from each separate part or unit.’ ed. Barrett and Freeman, 20.303.

²⁸ Op. cit., Origin (1876) 16.6.

²⁹ Op. cit., Origin (1876) 16.5.

³⁰ Joravsky, David, ‘Inheritance of Acquired Characteristics,’ Dictionary of the History of Ideas, ed., Wiener, Philip (New York: Scribners, 1973), 2.617–18Google Scholar. Darwin cited the report in the second (1875) edition of The Variation of Animals and Plants under Domestication.

³¹ Op. cit., Descent, ed. Barrett and Freeman, 21.40.

³² Isaac La Peyrère introduced polygenism in an effort to reconcile biblical with other chronologies, and to explain the human presence in the New World, by positing human lines not descended from Adam. His bold Prae-Adamite hypothesis was very much the stuff of the early Enlightenment. See Popkin, Richard, Isaac La Peyrère - 1596–1676 (Leiden: Brill, 1987)Google Scholar. The idea was seized upon by the racial theorists of the Smithsonian movement in the run-up to the American Civil War. It survives among some twentieth century racialist anthropologists: Hooton, E. A., Up from the Ape (New York: Macmillan, 1954)Google Scholar and Coon, Carleton, The Living Races of Man (New York: Knopf, 1965)Google Scholar. See Goodman, M. J. and Goodman, L. E., ‘‘Particularly amongst the Sunburnt Nations…’ The Persistence of Stereotypes of Race in Bio-Science,’ International Journal of Group Tensions 19 (1989), 221–43, 365–84Google Scholar. Browman, David L. and Williams, Stephen in Anthropology at Harvard: A Biographical History, 1790–1940 (Cambridge, MA: Peabody Museum Press, 2013)Google Scholar mentions but does not stress the prominence of polygenism in the work of Agassiz, Hooton, and Coon.

³³ Op. cit. Descent, 21.182.

³⁴ Op. cit. Descent, 21.182; cf. 21.151.

³⁵ Op. cit. Descent, 21.183 [1.230].

³⁶ See Richards, Robert J., The Tragic Sense of Life: Ernst Haeckel and the Struggle over Evolutionary Thought (Chicago: University of Chicago Press, 2008)CrossRefGoogle Scholar.

³⁷ Op. cit. Descent, 21.187; cf. 21.181–2 [1.235; cf. 1.227–28].

³⁸ Op. cit. Descent, 21.4.

³⁹ Op. cit. Descent, 21.9; cf. 22.633.

⁴⁰ Op. cit. Descent, 21.9–10. He discusses the pros and cons at length at 21.173–87.

⁴¹ Op. cit. Descent, 21.133.

⁴² Op. cit. Descent, 21.21–2.

⁴³ Op. cit. Descent, 21.24.

⁴⁴ Op. cit. Descent, 21.26.

⁴⁵ Op. cit. Descent, 21.175–6.

⁴⁶ Op. cit. Descent, 21.24.

⁴⁷ Op. cit. Descent, 21.76–7; 21.166; 22.585–6. ‘He who rejects with scorn the belief that the shape of his own canines, and their occasional great development in other men, are due to our early progenitors having been provided with these formidable weapons will probably reveal by sneering the line of his descent. For though he no longer intends, nor has the power, to use these teeth as weapons, he will unconsciously retract his ‘snarling muscles’ (thus named by Sir C. Bell) so as to expose them ready for action, like a dog prepared to fight.’ Descent, 21.45 [1.127].

⁴⁸ Op. cit. Descent, 21.91.

⁴⁹ Op. cit. Descent, 21.134.

⁵⁰ Op. cit. Descent, 21.136.

⁵¹ Op. cit. Descent, 21.40.

⁵² Op. cit. Descent, 21.56.

⁵³ Op. cit. Descent, 21.165–6.

⁵⁴ Having ‘no doubt that the various races, when carefully compared and measured, differ much from each other as in the texture of the hair, the relative proportions of all parts of the body, the capacity of the lungs, the form and capacity of the skull, and even in the convolutions of the brain,’ Darwin argues: ‘The races differ also in constitution, in acclimatization and in liability to certain diseases. Their mental characteristics are likewise very distinct; chiefly as it would appear in their emotional, but partly in their intellectual faculties. Every one who has had the opportunity of comparison, must have been struck with the contrast between the taciturn, even morose, aborigines of S. America and the light-hearted, talkative negroes. There is a nearly similar contrast between the Malays and the Papuans, who live under the same physical conditions, and are separated from each other only by a narrow space of sea.’ Descent, 21.173–74.

⁵⁵ Op. cit. Descent, 21.2 and 21.138. Cf. op cit. Goodman and Goodman, ‘Especially Amongst the Sunburnt Nations…’

⁵⁶ Op. cit. Descent, 21.133, 138–9.

⁵⁷ Op. cit. Descent, 21.138–47.

⁵⁸ Op. cit. Descent, 21.137.

⁵⁹ Op. cit. Descent, 21.5. Despite the evidence of homology, embryology, and vestigial organs, Darwin had felt hampered by the relative lack of comparative evidence when discussing the evolution of a single species (Descent, 21.3). Prum, Richard, The Evolution of Beauty: How Darwin's Forgotten Theory of Mate Choice Shapes the Animal World – and Us (New York: Doubleday, 2017)Google Scholar documents the birth of Darwin's theory here. But sexual selection is hardly forgotten by biologists. See Richards, Eveleen, Darwin and the Making of Sexual Selection (Chicago: University of Chicago Press, 2017)CrossRefGoogle Scholar and the serious study by Cronin, Helena, The Ant and the Peacock Altruism and Sexual Selection from Darwin to Today (Cambridge: Cambridge University Press, 1991)Google Scholar.

⁶⁰ Op. cit. Descent, 21.5.

⁶¹ Op. cit. Descent, 21.97.

⁶² Op. cit. Descent, 21.179.

⁶³ Darwin, Notebook B, p. 24, quoted in Sulloway, Frank J., ‘Geographic Isolation in Darwin's Thinking: The Vicissitudes of a Crucial Idea,’ Studies in the History of Biology 3 (1979), 23Google ScholarPubMed. Cf. op. cit. Origin, 16.84, where speciation is ascribed to such behavioural processes in incipient species, ‘from haunting different stations, from breeding at slightly different seasons, or from the individuals of each variety pairing together.’

⁶⁴ Op. cit. Descent, 21.71–2.

⁶⁵ Op. cit. Descent, 22.596.

⁶⁶ Op. cit. Descent, 22.630; cf. 22.218.

⁶⁷ Op. cit. Descent, 22.596–600.

⁶⁸ Op. cit. Descent, 22.630.

⁶⁹ Op. cit. Descent, 22.630.

⁷⁰ Op. cit. Descent, Chapter 5.

⁷¹ Op. cit. Descent, 22.537.

⁷² Op. cit. Descent, 22.548.

⁷³ Op. cit. Descent, 22.620–1.

⁷⁴ Op. cit. Descent, 21.176–9.

⁷⁵ Op. cit. Descent, 21.178–9.

⁷⁶ Op. cit. Descent, 21.31.

⁷⁷ W. R. Greg, quoted in op. cit. Descent, 21.143. Darwin's prefatory comment to this citation: ‘the reckless, degraded, and often vicious members of society, tend to increase at a quicker rate than the provident and generally virtuous members.’ Darwin's Lamarckian penchant allows him to make vice an ethnic trait, just as the word squalor allows him to assimilate poverty to mental and moral weakness.

⁷⁸ Op. cit. Descent, 584–5.

⁷⁹ Op. cit. Descent, 22.587–8.

⁸⁰ Cf. Haller, John, Outcasts from Evolution (Chicago: University of Chicago Press, 1971)Google Scholar.

⁸¹ Op. cit. Descent, 21.71; cf. 21.120: ‘It is possible, or, as we shall hereafter see, even probable, that the habit of self-command may, like other habits, be inherited.’

⁸² Op. cit. Descent, 21.129–30.

⁸³ Op cit. Descent, 21.131.

⁸⁴ Op cit. Descent, 21.118.

⁸⁵ For the incoherence of the idea of heritable (im)morality, see Kant, Immanuel, Religion Within the Limits of Reason Alone, trans. Greene, T. M. and Hudson, H. H. (New York: Harper & Row, 1960), 16–21Google Scholar, cf. 22–49.

⁸⁶ Op cit. Descent, 21.135–7.

⁸⁷ Op cit. Descent, 21.138, 145–7.

⁸⁸ Op cit. Descent, 21.127–8.

⁸⁹ Op cit. Descent, 21.71.

⁹⁰ Op cit. Descent, 22.579–80.

⁹¹ Op cit. Descent, 21.74.

⁹² Op cit. Descent, 22.586–7.

⁹³ Op cit. Descent, 22.588.

⁹⁴ Op cit. Descent, 22.588.

⁹⁵ Op cit. Descent, 22.588.

⁹⁶ Cf. Hrdy, Sarah Blaffer's now classic The Woman that Never Evolved (Cambridge: Harvard University Press, 1999; first published, 1981)Google Scholar.

⁹⁷ Op cit. Descent, 22.580.

⁹⁸ Op cit. Descent, 21.187–8.

⁹⁹ Op cit. Descent, 21.188.

¹⁰⁰ Op cit. Descent, 21.162; cf. Darwin's letter to Lyell of October 11, 1859, Correspondence 7.345.

¹⁰¹ Op cit. Descent, 22.602–4.

¹⁰² Op cit. Descent, 22.605.

¹⁰³ I find it in the 10^th century Case of the Animals vs Man, for example, tr. L. E. Goodman and Richard McGregor (Oxford: Oxford University Press, 2009) 113.

¹⁰⁴ Desmond, Adrian and Moore, James, , in Darwin's Sacred Cause and the Quest for Human Origins (Chicago: University of Chicago Press, 2009)Google Scholar, see a quest for human unity as the humane ‘cause’ that motivated Darwin's pressing his evolutionary theory to include the human case. But this seems excessive and needlessly apologetic. The human case, as Darwin stressed, was the toughest nut to crack for his evolutionary theory, not only for the resistance roused in regard to human origins but for the special circumstances that seemed to mitigate natural selection. Indeed, it was the perceived insufficiency of natural selection in the human case that turned Darwin toward sexual selection to strengthen the argument and speed the course of evolution. I think it is clear that Darwin's writings are motivated neither by animus nor by benevolence but by his dedication to scientific inquiry and commitment to his powerful explanatory theory (so much so that one can understand the neglect of the Mendelian idea by early Darwinians and perhaps by Darwin himself in terms of his search for continuous rather than discrete vectors and degrees of heritable change). Darwin may worry about or cheer the rate and the constancy of evolutionary change, but he does not see the need to egg on what he takes to be the agents of such change. Nature, as he sees it, assures its outcome.

¹⁰⁵ Op. cit. Descent, 21.70.

¹⁰⁶ Op. cit. Descent, 21.199.

¹⁰⁷ Op. cit. Descent, 21.154–5, where seals and fish are similar in adaptations to their environment but sharply different in less critical respects. At 21.185 [1.233] Darwin applies his axiom to the human case: ‘it is improbable that the numerous and unimportant points of resemblance between the several races of man in bodily structure and mental faculties… should all have been independently acquired.’

¹⁰⁸ Op. cit. Descent (Princeton: Princeton University Press, 1981; facsimile of London: Murray, 1871) 1.248–49; cf. Descent, 21.204–5, where the text varies.

¹⁰⁹ Alfred Russel Wallace, for one, disputed the low appraisal of the moral character of tribal peoples that Darwin shared with Spencer. He found altruism as well as a concern for reputation among those ‘savages’ he had met, and they respected one another's rights, on the whole without formal laws or courts. See his The Malay Archipelago: The Land of the Orang-utan and the Bird of Paradise - A Narrative of Travel, with Studies of Man and Nature (London: Macmillan, 1869; revised, 1890; New York: Dover, 1962).

¹¹⁰ Schweitzer, Albert, On the Edge of the Primeval Forest (1922; reprinted, London: Black, 1956), 102Google Scholar.

¹¹¹ Radin, Paul, Primitive Man as Philosopher (New York: Appleton, 1927, 1955; reprinted, New York: Dover, 1957)Google Scholar.

¹¹² Darwin's readiness to typify the extreme and essentialize the exotic contrasts strikingly with his brilliant recognition of critical role of individual differences in animal morphology.

¹¹³ Mishnah Sanhedrin 4.5.

¹¹⁴ Op. cit. Descent, 21.169.

¹¹⁵ Op. cit. Descent, 21.129.

¹¹⁶ Op. cit. Descent, 21.103.

¹¹⁷ Op. cit. Descent, 21.129–30. O'Hear's remark cited above is to be found in his ‘Darwinian Tensions’, in Turning Images in Philosophy, Science and Religion, edited by Charles Taliaferro and Jil Evans (Oxford: Oxford University Press, 2011, 46–66) at 54.

¹¹⁸ Gardner, Howard, Frames of Mind: The Theory of Multiple Intelligences (New York: Basic Books, 1993; first edition, 1983)Google Scholar.

¹¹⁹ Op. cit. Descent, 21.69.

¹²⁰ Op. cit. Descent, 22.611.

¹²¹ Spencer, like Darwin, pictured ‘savages’ as impulsive, selfish (except when enticed by their childlike avidity for approval), and animal-like in their sexual appetites and parental instincts. Spencer read voluminously in the travel literature and early anthropology of his day. He also employed (and physically exhausted) three research assistants in scanning such literature for him, sharing with them the proceeds of the publications in which he used their work. Volume 1 of The Principles of Sociology has 2192 citations from 379 such sources. Yet he was spotty, selective, and often superficial in his use of sources. And most damaging was his subjectivity. For he read into the materials he encountered his own over-arching construction of the meanings of the data they presented, and he overlaid familiar stereotypes and presumptions as well – many of which were already well ensconced in what he read. So the anthropological research on which so much of his ‘descriptive sociology’ would depend became a kind of funhouse hall of mirrors, revealing little that was new but enlarging and distorting what he had begun with.

¹²² As Edward Westermarck (1862–1939), the Finnish sociologist and anthropologist, was the first to show, Darwin's ideas about primitive polygamy and promiscuity, and Engels’ related notions of primitive communism (of women), were unsupported by evidence from tribal societies. Such societies were typically monogamous, and no instance of norms of promiscuity could be found among them. Travelers’ tales, fantasies, misinterpretations of local norms and behaviors, and of the relations betwen the two account for most of what was once imagined on this score, and for the persistence of such notions even today.

¹²³ Op. cit. Descent, 22.619.

¹²⁴ Op. cit. Descent, 21.132.

¹²⁵ Op. cit. Descent, 22.611.

¹²⁶ Op. cit. Descent, 22.611–2. Cf. Spencer, Principles of Sociology (New York: Appleton, 1898; first published, 1876),1.622Google Scholar.

¹²⁷ Op. cit. Descent, 22.615–6.

¹²⁸ Op. cit. Descent, 22.617.

¹²⁹ Op. cit. Descent, 22.223–9.

¹³⁰ Op. cit. Descent, 21.174.

¹³¹ One can't help wondering what Darwin would have thought of our present realization that the Neanderthals were not wiped out but interbred with their Cro-Magnon contemporaries – and that their genes are traceable in many of us today.

¹³² Op. cit. Descent, 22.621–2.

¹³³ Op. cit. Descent, 21.140.

¹³⁴ Op. cit. Cannon, note 3, 27–28, citing Darwin's letter to T. H. Huxley, in Francis Darwin and A. C. Seward, eds., More Letters of Charles Darwin (1903) 1.125.

¹³⁵ Darwin himself pegs the maturation and improvement of the moral sense of humankind to the extension of moral regard to ‘the men of all races.’ But that view does not lead him to assign moral, mental, or existential equality to those men. The same generalization of benevolence, in his view, extends to ‘the imbecile, the maimed, and other useless members of society, and finally to the lower animals.’ Descent, (Princeton), 1.103.

¹³⁶ Thus the photo essays of Leni Riefenstahl on African peoples.

¹³⁷ Charles Darwin to W. Graham, July 3, 1881, Life and Letters, 1.316, cited in Himmelfarb, Gertrude, Darwin and the Darwinian Revolution (London: Chatto and Windus, 1959) 343Google Scholar. More fully: ‘Lastly, I could show fight [i.e., vigorously argue] on natural selection having done and doing more for the progress of civilisation than you seem inclined to admit. Remember what risk the nations of Europe ran, not so many centuries ago, of being overwhelmed by the Turks, and how ridiculous such an idea now is! The more civilized so-called Caucasian races have beaten the Turkish hollow in the struggle for existence. Looking to the world at no very distant date, what an endless number of the lower races will have been eliminated by the higher civilised races throughout the world.’ Darwin, Charles, The Life of Charles Darwin by Darwin, Francis (London: Senate, 1995, reprint of the 1902 John Murray edition), 64Google Scholar.

¹³⁸ Darwin, Descent (Princeton: Princeton University Press, 1981; facsimile of London: Murray, 1871) 1.239–40Google Scholar.*

¹³⁹ Humboldt, Alexander von, Cosmos: A Sketch of a Physical Description of the Universe, translated by Otté, E. C. (London: Bohn, 1849) 1.358Google Scholar.