Charles Darwin Solves the “Riddle of the Flower”;Or,Why Don't Historians of Biology Know about the Birds and the Bees?

Knuth Paul , Handbuch der Blütenbiologie; unter Zugrundelegung von Hermann Müllers Werk: “Die Befruchtung der Blumen durch Insekten” (Leipzig, 1898–1905). The first two volumes were translated into English: Knuth Paul , Handbook of flower pollination, based upon Hermann Müller's work “The fertilisation of flowers by insects”, transl. by Ainsworth J. R. Davis (Oxford, 1906–9).

Darwin Charles , On the various contrivances by which British and foreign orchids are fertilised by insects, and on the good effect of intercrossing (London, 1862).

Müller Hermann , The fertilisation of flowers, transl. by Thompson D. W. , with preface by Charles Darwin (London, 1883), 599–630.

Knuth , Handbook (ref. 1), i, pp. vii, 212–380.

Editorial policy, in The correspondence of Charles Darwin, ed. by Burkhardt Frederick (Cambridge, 1985–2008; hereafter CCD), xii, p. xxxv. At the risk of anachronism I distinguish pollination from fertilisation in this essay.

[ Coulter John M. ], “Knuth's handbook”, Botanical gazette, xlii (1906), 494–5.

de Chadarevian Soraya , “Laboratory science versus country-house experiment: The controversy between Julius Sachs and Charles Darwin”, The British journal for the history of science, xxix (1996), 17–41, p. 18.

Smith Jonathan , Charles Darwin and Victorian visual culture (Cambridge, 2006), 137; Kohn David , Darwin's garden: An evolutionary adventure (New York, 2008), 9.

Kohn David is currently working to fill this gap with a major study.

Baker Herbert G. , “Charles Darwin and the perennial flax — A controversy and its implications”, Huntia, ii (1965), 141–61; Baker Herbert G. , “Anthecology: Old Testament, New Testament, Apocrypha”, New Zealand journal of botany, xvii (1979), 1979–40; Baker Herbert G. , “An outline of the history of anthecology, or pollination biology”, in Pollination biology, ed. by Real Leslie (Orlando, 1983), 7–28.

Pancaldi Giuliano , Darwin in Italy: Science across cultural frontiers, transl. by Morelli Ruey Brodine (Bloomington, 1991), 107–36.

Junker Thomas , Darwinismus und Botanik: Rezeption, Kritik und theoretische Alternativen im Deutschland des 19. Jahrhunderts (Stuttgart, 1989), 138–50, 261–87.

Allan Mea , Darwin and his flowers: The key to natural selection (London, 1977).

Ayers Peter , The aliveness of plants: The Darwins at the dawn of plant science (London, 2008).

Smith , Darwin and Victorian visual culture (ref. 8).

Other sources include: Müller , Fertilisation (ref. 3), 4–11; Knuth , Handbook (ref. 1), i, 8–11; Darwin Francis , “Darwin's work on the movements of plants”, in Darwin and modern science, ed. by Seward A. C. (Cambridge, 1909), 385–100; Goebel K. , “The biology of flowers”, ibid., 401–23; Irvine William , Apes, angels, and Victorians: The story of Darwin, Huxley, and evolution (Lanham, MD, 1955), 186–92; Dupree A. Hunter , Asa Gray: American botanist, friend of Darwin (Baltimore, 1988), 304–5, 355–60; Whitehouse H. L. K. , “Cross- and self-fertilization in plants”, in Darwin's biological work: Some aspects reconsidered, ed. by Bell P. R. (New York, 1959), 207–61; Ghiselin Michael T. , The triumph of the Darwinian method (Mineola, NY, 2003), 131–59; Desmond Adrian Moore James , Darwin (New York, 1991), 509–11; De Chadarevian , “Laboratory science versus country-house experiment” (ref. 7); Browne Janet , Charles Darwin: The power of place (New York, 2002), 165–78, 182–3, 192–5; Kohn David Murrell Gina Parker John Whitehorn Mark , “What Henslow taught Darwin”, Nature, cdxxxvi (2005), 2005–5; Kohn , Darwin's garden (ref. 8); Smocovitis Vassiliki Betty , “Darwin's botany in the Origin of species”, in The Cambridge companion to the “Origin of species”, ed. by Ruse Michael Richards Robert J. (Cambridge, 2009), 216–36; and Schneckenburger Stefan , “Darwin und die Blütenbiologie” in Darwin und die Botanik, ed. by Stöcklin Jürg Höxtermann Ekkehard (Rangsdorf, 2009), 76–101.

Comment Bernard , The painted panorama, transl. by Glasheen Anne-Marie (New York, 2000), 62–65.

London as it is to-day: Where to go, and what to see, during the Great Exhibition (London, 1851), 281.

Müller Hermann , “Die Wechselbeziehungen zwischen den Blumen und den ihre Kreuzung vermittelnden Insekten”, in Handbuch der Botanik, ed. by Schenk A. (Breslau, 1879), i, 1–112, p. 2.

Ruricola , “Humble-bees”, Gardeners' chronicle, 24 July 1841, 485.

Darwin Charles , “Humble-bees”, Gardeners' chronicle, 21 August 1841, 550.

Darwin to Herbert , 26 June 1839, CCD (ref. 5), ii, 201–2.

Brown Robert , “On the organs and mode of fucundation in Orchideæ and Asclepiadeæ”, Transactions of the Linnean Society of London, xvi (1833), 685–745, pp. 717, 733.

[ Brewster David ], “Life and correspondence of Sir James Edward Smith”, Edinburgh review, lvii (1833), 39–69, pp. 68–69.

Sprengel Christian Konrad , “Discovery of the secret of nature in the structure and fertilization of flowers”, transl. by Haase Peter , in Floral biology: Studies on floral evolution in animal-pollinated plants, ed. by Lloyd David G. Barrett Spencer C. H. (New York, 1996), 3–43, p. 41.

Baker , “Anthecology” (ref. 10), 432; Vogel Stefan , “Christian Konrad Sprengel's theory of the flower: The cradle of flower ecology”, in Lloyd Barrett (eds), Floral biology (ref. 25), 44–60, pp. 45–46.

Darwin Charles , The autobiography of Charles Darwin, 1809–1882, ed. by Barlow Nora (New York, 1958), 127.

Darwin to Hooker , [11–12 July 1845], CCD (ref. 5), iii, 217.

Sprengel , “Discovery” (ref. 25), 3, 13, 43. see also Nordenskiöld Erik , The history of biology: A survey, transl. by Eyre L. B. (London, 1929), 256–7; Müller , Fertilisation (ref. 3), 1–3.

On Darwin's use of ‘final cause’, see Lennox James G. , “Teleology”, in Keywords in evolutionary biology, ed. by Keller Evelyn Fox Lloyd Elisabeth A. (Cambridge, MA, 1995), 324–33, pp. 328–9.

Darwin to Oliver , 13 October [1862], CCD (ref. 5), x, 457; Darwin , Autobiography (ref. 27), 126–9.

Darwin Charles , “On the two forms, or dimorphic condition, in the species of Primula, and on their remarkable sexual relations” [read 21 November 1861], Journal of the proceedings of the Linnean Society (botany), vi (1862), 77–96, pp. 77–78; idem, The different forms of flowers on plants of the same species (London, 1877), 14.

Kohn , “What Henslow taught Darwin” (ref. 16), 644–5.

Henslow John Stevens , “On the specific identity of the primrose, oxlip, cowslip, and polyanthus”, Magazine of natural history, iii (1830), 406–9.

Bellon Richard , “Joseph Hooker takes a ‘fixed post’: Transmutation and the ‘present unsatisfactory state of systematic botany’, 1844–1860”, Journal of the history of biology, xxxix (2006), 1–39, pp. 9–13; Endersby Jim , Imperial nature: Joseph Hooker and the practices of Victorian science (Chicago, 2008), 154–9, 270–2.

Darwin Charles , On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life (London, 1859), 96–101.

Daubeny Charles , On the final causes of the sexuality of plants, with particular reference to Mr. Darwin's work on the origin of species (Oxford, 1860), 26–27.

Daubeny , On the final causes (ref. 37), 24.

Rupke Nicolaas A. , Richard Owen: Victorian naturalist (New Haven, 1994), 270–2; and Cosans Christopher E. , Owen's ape and Darwin's bulldog: Beyond Darwinism and creationism (Bloomington, 2009).

Darwin to Daubeny , 16 July [1860], CCD (ref. 5), xiii, 428–9.

Darwin , Origin (ref. 36), 1–2.

Darwin to Huxley , 21 [January 1860], CCD (ref. 5), viii, 43.

Darwin to Hooker , [13 March 1846], [22 January 1860], [26 February or 4 March 1860], 12 March [1860], 18 April [1860], 12 July [1860], CCD (ref. 5), iii, 45, viii, 45–46, 111–12, 127–8, 162–4, 285–7.

Darwin to Hooker , [26 February or 4 March 1860] and 18 April [1860], Darwin to Lyell, 12 September [1860], CCD (ref. 5), viii, 111–12, 162–4, 354–6.

Darwin to Malden , 15–16 June [1861], to Hooker, 13 October [1861] and 27 October [1861], and to Lindley, 25 October [1861], CCD (ref. 5), ix, 175, 304, 324, 321.

Darwin to Oliver , 20 October [1860], CCD (ref. 5), viii, 440.

Wilberforce Samuel , “Darwin's origin of species”, Quarterly review, cviii (1860), 225–64, p. 231.

Harvey William Henry , An inquiry into the probable origin of the human animal, on the principle of Mr. Darwin's theory of natural selection, and in opposition to the Lamarckian notion of a monkey parentage (Dublin, 1860), 6, 10, 23–24; idem, “Darwin on the origin of species”, Gardeners' chronicle, 18 February 1860, 145–6.

[ Owen Richard ], “Darwin on the origin of species”, Edinburgh review, cxi (1860), 487–532, 580.

Darwin to Murray , 24 September [1861]; see also Darwin to Gray, 23[–4] July [1862], CCD (ref. 5), ix, 279, x, 330.

Darwin , “On the two forms” (ref. 32), 91–92.

Darwin to Hooker , 19 June [1861], CCD (ref. 5), ix, 182.

Darwin to Lyell , 20 October [1861] and Darwin to Hooker, CCD (ref. 5), ix, 309, 345.

Darwin to Murray , 21 September [1861] and Murray to Darwin, 23 September 1861, CCD (ref. 5), ix, 272–3, 276.

Bentham George , diary mss., Royal Botanic Gardens, Kew, Archives, xvii, entry for 3 April 1862; Linnean Society general minute books, Linnean Society of London, Library, vii, 1850–62, entry for 3 April 1862.

Lindley John , The vegetable kingdom; or, the structure, classification and uses of plants, illustrated upon the natural system (London, 1846), 178.

Darwin Charles , “On the three remarkable sexual forms of Catasetum tridentatum, an orchid in the possession of the Linnean Society”, Journal of the proceedings of the Linnean Society (botany), vi (1862), 151–7.

Darwin , “On the two forms” (ref. 32), 84, 94.

Darwin to Murray , 21 September [1861], CCD (ref. 5), ix, 273.

Darwin , Orchids (ref. 2), 1–2.

Darwin , Orchids (ref. 2), 288.

[George Campbell (eighth Duke of Argyll)], “The supernatural”, Edinburgh review, cxvi (1862), 378–97; [John R. Leifchild], Review of Orchids by Darwin Charles , The Athenaeum, no. 1804 (1862), 683–5. For a more sympathetic view of Argyll's engagement with Orchids, see Thierry Hoquet's contribution on Darwin's botany in a special issue of Comptes rendues biologies on “Non-Darwinian Darwin”, ed. by Gros F. Gayon J. Veuille M. (forthcoming, 2010).

Darwin , Orchids (ref. 2), 289–95.

Darwin , “On the three remarkable sexual forms” (ref. 57), 154–5; Darwin , Orchids (ref. 2), 241–4. He made the same point in gentler language in the Origin (ref. 36), 453. The “distinguished naturalist” in question was the great Swiss botanist, Augustin-Pyramus de Candolle. See his Théorie élémentaire de la botanique, 2nd edn (Paris, 1819), 185–6.

Darwin , Orchids (ref. 2), 306–7.

Darwin , Orchids (ref. 2), 351–2.

Darwin , Orchids (ref. 2), 288.

Darwin , Orchids (ref. 2), 157–8.

Darwin to Gray , 22 January [1862], CCD (ref. 5), x, 40.

Darwin to Hooker , 14 December [1859], CCD (ref. 5), vii, 431.

Bellon Richard , “Inspiration in the harness of daily labor: Darwin, orchids and the triumph of evolution, 1858–1872”, paper delivered at the conference “The reception of Darwinism: Transcultural differences”, held at Boston University, 4 April 2009; and “Why naturalists were right to reject Darwin's theory (in 1858)”, paper delivered at the conference “Darwin and the boundaries of science”, held at New York University, 17 April 2009. An article elaborating this thesis is in preparation.

Darwin to Bates , 3 December [1861], CCD (ref. 5), ix, 363.

Browne , Darwin (ref. 16), 194.

Huxley Thomas , “Obituary”, Proceedings of the Royal Society of London, xliv (1888), pp. i–xxv, p. xxi.

Hooker to Darwin , 28 June 1862, CCD (ref. 5), x, 275.

[ Hooker Joseph ], Review of Orchids by Darwin Charles , Natural history review, ii (1862), 371–6, p. 371.

[ Hooker Joseph ], Review of Orchids by Darwin Charles , Gardeners' chronicle, 23 August, 13 September, 27 September 1862, 789–90, 863, 910, pp. 789–90.

Gray Asa , Review of Orchids by Darwin Charles , American journal of science and arts, xxxiv (1862), 138–44, p. 144; idem, “Fertilization of orchids through the agency of insects”, American journal of science and arts, xxxiv (1862), 1862–9, pp. 422, 429.

Gray to Darwin , 4 and 13 October 1862, CCD (ref. 5), x, 445.

Gray to Darwin , 2–3 July 1862, Darwin to Gray, 23[–4] July [1862], CCD (ref. 5), x, 292, 330.

Hooker , Review, Gardeners' chronicle (ref. 77), 789–90.

Hooker Joseph , “Presidential address”, in Report of the thirty-eighth meeting of the British Association for the Advancement of Science; held at Norwich in August, 1868 (London, 1869), pp. lviii–lxxv, p. lxviii.

Quoted in Ellegård Alvar , Darwin and the general reader: The reception of Darwin's theory of evolution in the British periodical press, 1859–1872 (Chicago, 1990), 72. see also Daubeny Charles , “The president's address”, Reports and transactions of the Devonshire Association for the Advancement of Science, Literature, and Art, iv (1865), 1865–29, p. 11; and Daubeny to Darwin , 5 July 1862, CCD (ref. 5), x, 301–2.

Balfour John Hutton , Outlines of botany, 2nd edn (Edinburgh, 1862), 356–9; [ Berkeley Miles Joseph , “Fertilization of orchids”, London review and weekly journal of politics, arts and sciences, iv (1862), 1862–4; idem, “Presidential address to biology section”, in Report of the thirty-eighth meeting of the British Association for the Advancement of Science; held at Norwich in August, 1868 (London, 1869), 83–87, pp. 86–87; Gosse Philip Henry , “Microscopic observations on some seeds of orchids”, Journal of horticulture and cottage gardener, iv (1863), 1863–8.

Bellon Richard , “‘The great question in agitation’: George Bentham and the origin of species”, Archives of natural history, xxx (2003), 282–97.

Bentham George , “Note on the stigmatic apparatus of Goodenovieæ” [read 19 December 1867], Journal of the proceedings of the Linnean Society (botany), x (1869), 203–6.

Bentham George , “On Inocarpus”, Journal of the proceedings of the Linnean Society (botany), vi (1862), 146–50, p. 147.

Harvey to Gray , 3 November 1860, in Memoirs of W. H. Harvey, M.D., F.R.S., ed. by Fischer Lydia Jane (London, 1869), 337–8.

Browne , Darwin (ref. 16), 104, 109–10, 175.

CCD (ref. 5), x, 672, 677.

Harvey to Darwin , 3 February 1863, 8 November [1864], 10 November 1864 and 11 November 1864, CCD (ref. 5), xi, 113, xii, 406–10.

Haughton Samuel , “Biogenesis”, Natural history review, vii (1860), 23–32.

For an analysis of the importance of friendship in the practice of Victorian natural history, see Endersby Jim , “Sympathetic science: Charles Darwin, Joseph Hooker, and the passions of Victorian naturalists”, Victorian studies, li (2009), 299–320.

Lightman Bernard , Victorian popularizers of science: Designing nature for new audiences (Chicago, 2007), 107, 156–8.

See the extensive bibliographies in Müller , Fertilisation (ref. 3), 599–603, and Knuth , Handbook (ref. 1), i, 212–372, for citations to published work by the above naturalists except Becker and Oliver. For them, see Becker Lydia , “On alteration in the structure of Lychnis diurnal, observed in connexion with the development of a parasitic fungus”, in Report of the thirty-ninth meeting of the British Association for the Advancement of Science; held at Exeter in August, 1869 (London, 1870), Transactions, 106; and [ Oliver Daniel ], “Dimorphic flowers”, Natural history review, iv (1864), 1864–8.

Reisz E. G. , “Henry Trimen”, in Dictionary of nineteenth-century British scientists, ed. by Lightman Bernard (Bristol, 2004), iv, 2033–4.

Darwin to Trimen , 31 January [1863], CCD (ref. 5), xi, 103.

Müller , Fertilisation (ref. 3), 19–21; Knuth , Handbook (ref. 1), i, 12–14; Darwin Charles , The effects of cross and self fertilisation in the vegetable kingdom (London, 1876), 6.

Knuth , Handbook (ref. 1), i, 12.

Delpino to Darwin , 5 September 1867, CCD (ref. 5), xv, 516–17; Pancaldi , Darwin in Italy (ref. 11), 107–9.

Crüger to Darwin , 21 January 1864, CCD (ref. 5), xii, 24. See also Crüger to Darwin, 23 February, 23 April and 8 August 1863, and Darwin to Crüger, 25 January and 25 May 1863, CCD (ref. 5), xi, 76, 163–5, 349–50, 446–7, 585–7.

Crüger to Darwin , 21 January 1864, CCD (ref. 5), xii, 24.

Nordenskiöld , The history of biology (ref. 29), 391.

Treviranus L. C. , “The botany of the Antarctic voyage etc. by Joh. Dalton Hooker”, Botanische Zeitung, xix (1861), 133–5, 142–4.

Treviranus L. C. , “Ueber Dichogamie nach C. C. Sprengel und Ch. Darwin”, Botanische Zeitung, xxi (1863), 1–7, 9–16; idem, “On the existence of two forms, and on their reciprocal sexual relations, in several species of the genus Linum by Charles Darwin”, Botanische Zeitung, xxi (1863), 1863–90.

Junker , Darwinismus und Botanik (ref. 12), 143–4.

Treviranus to Darwin , 12 February 1863, CCD (ref. 5), xi, 127.

Hildebrand to Darwin , 14 July 1862 and 16 July 1863, and Darwin to Hildebrand, 28 July [1863], CCD (ref. 5), x, 323, xi, 535–6, 567–8.

Hildebrand Friedrich , “Die Fruchtbildung der Orchideen, ein Beweis für die doppelte Wirkung des Pollen”, Botanische Zeitung, xxi (1863), 329–33, 337–45; Alefeld Friedrich , “Ueber Linum”, Botanische Zeitung, xxi (1863), 1863–2; von Mohl Hugo , “Einige Beobachtungen über dimorphe Blüthen”, Botanische Zeitung, xxi (1863), 1863–15, 321–8.

Knuth , Handbook (ref. 1), i, 10–11.

Hildebrand Friedrich , Die Geschlecht-vereilung bei den Pflanzen und das Gesetz der Vermiedenen und unvortheilhaften stetigen Selbstbefruchtung (Leipzig, 1867), 3–4.

Junker , Darwinismus und Botanik (ref. 12), 145–7.

Müller Fritz , Für Darwin (Leipzig, 1864). see also West David , Fritz Müller: A naturalist in Brazil (Blacksburg, 2002), 116–41; Browne , Darwin (ref. 16), 259–60.

Müller F. to Darwin Charles , 12 August 1865, 31 August 1865, 10 October 1865 and 5 November 1865, CCD (ref. 5), xiii, 468–73.

Müller Fritz , “Notes on some of the climbing-plants near Desterro, in South Brazil” [read 7 December 1865], Journal of the proceedings of the Linnean Society (botany), ix (1866), 344–9.

West , Fritz Müller (ref. 113), 143–4.

Müller Fritz , “Ueber die Befruchtung der Martha (Posoqueria?) fragrans”, Botanische Zeitung, xxiv (1866), 129–33; Müller , Fertilisation (ref. 3), 305. See also Darwin to F. Müller, [before 10 December 1866], CCD (ref. 5), xiv, 415–16.

For a bibliography, see Knuth , Handbook (ref. 1), i, 316–18.

West , Fritz Müller (ref. 113), 143. see also Knuth , Handbook (ref. 1), i, 20–21.

Müller Hermann , Die Befruchtung der Blumen durch Insekten (Leipzig, 1873).

Darwin , Cross and self fertilisation (ref. 98), 6.

On Haeckel as the face of German Darwinism, see Richards Robert J. , The tragic sense of life: Ernst Haeckel and the struggle over evolutionary thought (Chicago, 2008); and Gliboff Sander Bronn H. G. , Ernst Haeckel and the origins of German Darwinism: A study in translation and transformation (Cambridge, MA, 2008).

Darwin to Huxley , 22 December [1866], CCD (ref. 5), xiv, 437.

Moore James , “Deconstructing Darwinism: The politics of evolution in the 1860s”, Journal of the history of biology, xxiv (1991), 353–408, pp. 384–6.

Müller , Fertilisation (ref. 3).

Kelly Alfred , The descent of Darwin: The popularization of Darwinism in Germany, 1860–1914 (Chapel Hill, 1981), 61–65.

Darwin , Cross and self fertilisation (ref. 98); Darwin , Different forms (ref. 32); Darwin Charles , The various contrivances by which orchids are fertilised by insects, 2nd edn (London, 1877), pp. vii–x.

Darwin Charles , The movements and habits of climbing plants (London, 1875); idem, Insectivorous plants (London, 1875); idem, The power of movement in plants (London, 1880).

Browne , Darwin (ref. 16), 494–5.

Darwin Charles , “Prefatory Notice” to The fertilisation of flowers by Müller Hermann (London, 1883), p. x.

Knuth , Handbook (ref. 1), i, pp. vi, 20–21.

Corsi Pietro , “Before Darwin: Transformist concepts in European natural history”, Journal of the history of biology, xxxviii (2005), 67–83.

Secord James A. , “Knowledge in transit”, Isis, xcv (2004), 654–72, p. 669.

Hooker to Harvey , 1860, in Life and letters of Joseph Dalton Hooker, ed. by Huxley Leonard (London, 1918), i, 517.

Desmond Adrian Moore James , Darwin's sacred cause: How a hatred of slavery shaped Darwin's views on human evolution (New York, 2009).

Hooker to Darwin , 6 October 1865, CCD (ref. 5), xi, 262–3.

Bennett to Darwin , 29 April 1864, CCD (ref. 5), xii, 159. Italics in the original.

Balfour John Hutton , “On dimorphic flowers of Cephaelis ipecacuanha, the ipecacuan plant”, Proceedings of the Royal Society of Edinburgh, vii (1872), 763–4; Gosse , “Microscopic observations” (ref. 84).

Balfour John Hutton , “On the fruiting of the ipecacuan plant (Cephaelis ipecacuanha, Rich.) in the Royal Botanic Garden”, Proceedings of the Royal Society of Edinburgh, vii (1872), 688–91, pp. 688, 691. For Balfour's debt to Darwin on dimorphism, see Balfour to Darwin, 14 January 1862, CCD (ref. 5), x, 17.

Gosse to Darwin , 5 April 1864, Darwin to Gosse, 7 April [1864], CCD (ref. 5), xii, 118, 126.

Gosse Edmund , Father and son: A study of two temperaments (London, 1907), 117.

Balfour , Outlines of botany (ref. 84), 358–9, 441.

Gray Asa , “Dimorphism in the genitalia of flowers”, American journal of science and arts, xxxiv (1862), 419–29, pp. 426–9. See also Asa Gray, Review of Orchids, American journal of science and arts, xxxiv (1862), 1862–44, pp. 139, 143.

Darwin to Gray , 8 June [1860], CCD (ref. 5), vii, 247–8.

Desmond Moore , Darwin's sacred cause (ref. 135), 327, 330–1.

Ruse Michael , Darwin and design (Cambridge, MA, 2003), 143.

See Knuth , Handbook (ref. 1), i, 222–3, 271–2.

Bennett Alfred William , “On the theory of natural selection looked at from a mathematical point of view”, in Report of the thirtieth meeting of the British Association for the Advancement of Science; held at Liverpool in September, 1870 (London, 1871), 130–1; idem, “Mimicry in plants”, Popular science review, xi (1872), 1872–10; Henslow George , The origin of plant structures by selfadaptation to the environment (London, 1895).

Bennett , “Mimicry” (ref. 148), 10.

Pancaldi , Darwin in Italy (ref. 11), 119–36; Dupree , Asa Gray (ref. 16), 227, 264–5.

Gray Asa , “Miscellanea”, American journal of science and arts, xviii (1878), 317.

Sabine to Phillips , 12 November 1863, in CCD (ref. 5), xi, 668–9. For accounts of the controversy over Darwin's nomination for the Copley Medal, see Bartholomew M. J. , “The award of the Copley Medal to Charles Darwin”, Notes and records of the Royal Society of London, xxx (1976), 1976–18; Burkhardt Frederick , “England and Scotland: The learned societies”, in The comparative reception of Darwinism, ed. by Glick Thomas F. (Chicago, 1988), 32–74, pp. 34–36; Barton Ruth , “‘An influential set of chaps’: The X-Club and Royal Society politics 1864–85”, The British journal for the history of science, xxiii (1990), 1990–81, pp. 61–62; “Darwin and the Copley Medal”, Appendix IV in CCD (ref. 5), xii, 509–27; Browne , Darwin (ref. 16), 244–7.

Hooker , “Presidential address” (ref. 82), pp. lxvi–lxviii.

“Michael Faraday” and “Charles Darwin”, in Portraits of men of eminence in literature, science, and art; with biographical memoirs (London, 1863–68), i, 147–51, v, 49–52.

Cleevley R. J. , “Alfred William Bennett”, in Dictionary of nineteenth-century British scientists, ed. by Lightman (ref. 96), i, 181–2.

Smith , Darwin and Victorian visual culture (ref. 8), 160–5; Lightman , Victorian popularizers (ref. 94), 279–89.

Knuth , Handbook (ref. 1), i, 213, 237, 297–8, 353.

Smith , Darwin and Victorian visual culture (ref. 8), 137–8.

Bateman James , The Orchidaceæ of Mexico and Guatemala (London, 1837–43), 6–7.

Bateman James , A monograph of Odontoglossum (London, 1874), p. iii.

Smith , Darwin and Victorian visual culture (ref. 8), 165–78.

Browne , Darwin (ref. 16), 477–8; Ayers , The aliveness of plants (ref. 14), 98–100.

De Chadarevian , “Laboratory science versus country-house experiment” (ref. 7). see also Ayers , The aliveness of plants (ref. 14), 97–114.

De Chadarevian , “Laboratory science versus country-house experiment” (ref. 7), 31.

Sachs Julius , Lectures on the physiology of plants, transl. by Ward H. Marshall (Oxford, 1887), 547, 674, 689–90, 698, 787–99.

Knuth , Handbook (ref. 1), i, p. v.

Baker , “An outline” (ref. 10), 14.

Knuth , Handbook (ref. 1), i, 22–23.

Cittadino Eugene , Nature as a laboratory: Darwinian plant ecology in the German Empire, 1880–1900 (Cambridge, 1990), especially pp. 1–8, 31, 37–50, 116–28.

Cittadino , Nature as a laboratory (ref. 169), 116.

Schimper A. F. W. , Plant-geography upon a physiological basis, transl. by Fisher William R. , rev. and ed. by Groom Percy Balfour Isaac Bayley (Oxford, 1903), 53, 120, 129, 138, 140, 153.

Cittadino , Nature as a laboratory (ref. 169), 146–57.

Warming Eugen , Oecology of plants: An introduction to the study of plant communities (Oxford, 1909), 83.

Cittadino , Nature as a laboratory (ref. 169), 149–51.

Clements Frederic E. Long Frances L. , Experimental pollination: An outline of the ecology of flowers and insects (Washington, DC, 1923), 3.

Kingsland Sharon E. , The evolution of American ecology, 1890–2000 (Baltimore, 2005), 141–5.

Bateson William , “The progress of genetic research”, in Report of the third International Conference 1906 on genetics, hybridisation (the cross-breeding of genera or species), the cross-breeding of varieties, and general plant-breeding, ed. by Wilks William (London, 1907), 90–97.

“List of invited guests”, Ibid., 22–28.

Richmond Marsha L. , “Women in the early history of genetics: William Bateson and the Newnham College Mendelians, 1900–1910”, Isis, xcii (2001), 55–90.

Bateson William Gregory R. P. , “On the inheritance of heterostylism in Primula”, Proceedings of the Royal Society of London, ser. B, lxxvi (1905), 581–6.

Barlow Nora , “Preliminary note on heterostylism in Oxalis and Lythrum”, Journal of genetics, iii (1913), 53–66.

Bateson , “The progress of genetic research” (ref. 177), 92.

[ Coulter ], “Knuth” (ref. 6).

Smocovitis , “Darwin's botany” (ref. 16), 235.

James Secord has categorized the move to study practice “as the single most important transformation in [the history of science] during the past twenty years”. See Secord , “Knowledge in transit” (ref. 133), 658.

Hooker , Review, Gardeners' chronicle (ref. 77), 790.

Kohler Robert , Lords of the fly: Drosophila genetics and the experimental life (Chicago, 1994); idem, Landscapes and labscapes: Exploring the lab-field border in biology (Chicago, 2002); idem, All creatures: Naturalists, collectors, and biodiversity, 1850–1950 (Princeton, 2006). As Kohler acknowledges (Lords of the fly, 3–6), his emphasis on practice builds upon the prior insights of other scholars, including Steven Shapin, Simon Schaffer, Owen Hannaway, William Cronon, Edward Thompson and Bruno Latour.

Kohler , Landscapes and labscapes (ref. 187), 98, 295.

Secord James A. , Victorian sensation: The extraordinary publication, reception, and secret authorship of Vestiges of the Natural History of Creation (Chicago, 2000), 520.

Secord , “Knowledge in transit” (ref. 133).

Endersby , Imperial nature (ref. 35), 6–8, 316, 326. For my argument about the importance of naturalhistory practice in the development and reception of Darwin's theory, see Bellon , “Joseph Hooker takes a ‘fixed post’” (ref. 35).

Darwin , Origin (ref. 36), 484–90.

Kohler , Landscapes and labscapes (ref. 187), 33.

Hull David L. , “Darwinism as a historical entity: A historiographic proposal”, in The Darwinian heritage, ed. by Kohn David (Princeton, 1985), 773–812, p. 776.

Bowler Peter J. , The non-Darwinian revolution: Reinterpreting a historical myth (Baltimore, 1988), 5.

Bowler has also referred to Darwin as a “figurehead” and a “steppingstone” to evolution. See Bowler , Non-Darwinian revolution (ref. 195), 67, 73.