Butterfly People

by William R. Leach

  9. Scudder, BEUSC, vol. 1, p. 180.

  10. William Doherty, “Notes on Assam Butterflies,” Journal of the Asia Society of Bengal (1889): 126; and Doherty to Scudder, November 20, 1886, SS-BMS.

  11. Scudder, BEUSC, vol. 2, p. 778.

  12. For Edwards’s culminating analysis, see William Henry Edwards, BNA, vol. 2 (Philadelphia: American Entomological Society, 1868–72).

  13. Sir John Lubbock, Fifty Years of Science: Being an Address Delivered at York to the British Association for the Advancement of Science in 1881 (London: Macmillan, 1882). Praise came from other quarters as well—for instance, from Raphael Meldola, another influential British evolutionary naturalist. In 1882 Meldola edited and translated August Weismann’s Studies in the Theory of Descent (with an introduction by Charles Darwin), carefully incorporating into it a thirty-five-page appendix devoted entirely to an explication of Edwards’s work on Papilio ajax (plates included), a great compliment. Joseph Lintner, too, in his 1878 address before the American Association for the Advancement of Science, singled out his friend for praise: “I commend to you the labors of William Henry Edwards in working out the histories of some of those butterflies which appear under different forms at different seasons of the year. The untiring zeal with which the work has been prosecuted and is being continued deserves the commendation which it has received from the most eminent European Entomologists.” See Lintner, CE (August 1878): 173–74; and Weismann, Studies in the Theory of Descent (London, 1882), 126–60. See also, on the extent of Edwards’s achievement, W. Conner Sorensen’s breakthrough analysis in his Brethren of the Net: American Entomology, 1840–1880 (Tuscaloosa: University of Alabama Press, 1995), 214–35.

  14. E. O. Essig, College Entomology (New York: Macmillan, 1942), 503.

  15. For a more extensive, fascinating analysis of the satyrids, see William Henry Edwards’s BNA, vol. 2, “Satyrus II, III.” See also Charles Remington, “Suture-Zones of Hybrid Interaction Between Recently Joined Biotas,” Evolutionary Biology 1 (1968): 321–428.

  16. Edwards never explained, however, why yellow females continued to multiply in the face of the supposed advantage of the black female. Nor has anyone else, according to Michael Gochfeld and Joanna Burger, who write that there is no satisfactory explanation as to “why the polymorphism exists. We suspect that the predation pressure may never be high enough to eliminate the yellow females entirely.” See their Butterflies of New Jersey (New Brunswick, NJ: Rutgers University Press, 1997), 125.

  17. Edwards to Scudder, January 23, 1888; September 15, 1887; and January 2, 1888, SS-BMS.

  18. Edwards, “On Pieris bryoniae Ochsenheimer, and Its Derivative Forms in Europe and America,” Papilio 1, no. 6 (1881): 98; and Augustus Grote, “The Origin of Ornamentation in the Lepidoptera,” CE (June 1888): 114–17, and “On Specific Names,” CE (March 1889): 52.

  19. Grote, “The Origin of Ornamentation in the Lepidoptera,” 114–17.

  20. Scudder, Butterflies, 234.

  21. Scudder, excursus 50, “Variations in Habit and Life According to Locality and Season of the Year,” BEUSC, vol. 2, pp. 1415–17.

  22. Doherty, “A List of Butterflies Taken in Kumaon,” Journal of the Asiatic Society of Bengal (1886): 103–6.

  23. William Henry Edwards, “Libythea,” in BNA, vol. 2. For lists of butterfly enemies, see Scudder, “Enemies,” BEUSC, vol. 2, pp. 1612–13.

  24. H. C. T. Godfray, Parasitoids (Princeton, NJ: Princeton University Press, 1994), 16.

  25. Harold F. Greeney, “Emergence of Parasitic Flies from Adult Actinote diceus (Nymphalidae: Acraeinae) in Ecuador,” Journal of the Lepidopterists’ Society 55, no. 2 (2001): 79–80.

  26. Archibald Weeks, “Method of Oviposition of Tachina,” Entomologica Americana 3 (August 1887): 126.

  27. On this history, Godfray, Parasitoids, 6–7; Richard R. Askew, Parasitic Insects (London: Heinemann, 1971), v, 117–21, 140–45; Nick Mills, “Parasitoids,” in Encyclopedia of Insects, ed. Vincent H. Resh and Ring T. Carde (San Diego: Academic Press, 2003), 845–47; and David Grimaldi and Michael S. Engel, Evolution of Insects (Cambridge: Cambridge University Press, 2005), 427.

  28. On Linnaeus and the Ichneumonidae, see James Duncan, The Natural History of British Butterflies (Edinburgh, 1835), 89–91.

  29. Johann Meigen, William Kirby, Henry Spence, and James Duncan, among others, offered grisly portraits.

  30. See Charles Darwin to Asa Gray, May 22, 1860, in The Correspondence of Charles Darwin, ed. Frederick Burkhardt et al., vol. 8 (Cambridge: Cambridge University Press, 1993), 224. I would like to thank Paul Farber for sharing with me this Darwin citation.

  31. Alpheus Packard, Guide to the Study of Insects (New York, [1869] 1876), 196–97.

  32. R. L. Duffus, The Innocents at Cedro: A Memoir of Thorstein Veblen and Some Others (New York: Macmillan, 1944), 52, 62–63. In the 1880s, Herbert Smith, a dedicated and respected young naturalist from upstate New York, studied Ichneumonidae in South America, where their numbers, he believed, reached many thousands of species (all unnamed). Smith’s collection for the British Museum, assembled in this decade, was the largest of its kind ever donated there (see Smith to Ezra Cresson, December 6, 1892, Academy of Natural Science Archives, Philadelphia). On the Smith and Edwards collections in the Natural History Museum in London, see Claude Morley, A Revision of the Ichneumonidae (London, 1912), vol. 1, preface and pp. 9–13; and vol. 2, pp. 16–17, 56–58, and 107.

  33. Howard cites Edwards’s work in his essay in Scudder, BEUSC, vol. 3, pp. 1880–81, 1883–84.

  34. For Edwards’s parasite collection, his sharing with Howard and Riley, and his accounts to Lintner, see Edwards to Lintner, July 22, December 24, and December 29, 1882, JL.

  35. See Riley to Scudder, first page and date missing (but probably early 1870s and certainly before 1877, the year Riley left Missouri for Washington), SS-BMS.

  36. BEUSC, vol. 3, pp. 1901–2, 1897–1911; Howard’s contribution is on pp. 1869–96, and Williston’s on pp. 1912–24.

  37. William Field, the curator of entomology at the Smithsonian in the 1940s and a highly regarded authority, cited this work in his Manual of Butterflies and Skippers of Kansas as the only source of information on American parasites (Lawrence, KS: Bulletin of the Department of Entomology, no. 12, May 15, 1938), 50.

  38. “The Enemies of Butterflies,” BEUSC, vol. 2, p. 1613.

  39. “Aglais milberti: The American tortoiseshell,” BEUSC, vol. 1, pp. 428–29.

  40. BEUSC, vol. 1, pp. 454–55.

  41. Edwards called this butterfly Thecla laeta, when Thecla was a still a viable genus for most hairstreaks. Today, the Latin name is Erora laeta, designated by Scudder in 1872 and retained by modern catalogs. Scudder also chose “the spring beauty” as the vernacular name, which has not been kept. Reflecting the ongoing struggle over nomenclature, some contemporary authorities have called it the “turquoise beauty” (James Scott, The Butterflies of North America [Stanford, CA: Stanford University Press, 1986], 385), others the “early hairstreak” (Alexander D. Klots, A Field Guide to the Butterflies [Boston: Houghton Mifflin, 1951], 129, and Robert Pyle, National Audubon Society Field Guide to North American Butterflies [New York: Knopf, 1998], 467). Edwards would have opposed these names. For Scudder’s names, see BEUSC, vol. 2, p. 819. See also Edwards, “Notes on the Collection of Butterflies Made by Mr. H. K. Morrison in Arizona, 1882,” Papilio 3 (January 1883): 1.

  42. “Limenitis I,” BNA, vol. 2.

  43. Grote, “Moths and Moth-Catchers: Part I,” Popular Science Monthly (June 1885): 246–52.

  44. BEUSC, vol. 2, p. 1603.

  45. Excursus 51, “Southern Invaders,” BEUSC, vol. 2, pp. 1332–34.

  46. Excursus 46, “The Spread of a Butterfly in a New Region (with Map),” BEUSC, vol. 2, pp. 1175–90.

  47. For a full rendering of the history of the work on monarch migrations, see Lincoln Brower, “Understanding and Misunderstanding the Migration of the Monarch Butterfly,” Journal of the
Lepidopterists’ Society 49, no. 4 (1995): 304–85.

  48. Charles Valentine Riley, “A Swarm of Butterflies,” American Entomologist (September 1868): 28–29.

  49. For Scudder’s monarch life history, see BEUSC, vol. 1, pp. 720–48.

  50. Scudder to Charles Fernald, September 15, 1888, Charles Henry Fernald Papers (RG 40/11, C. H. Fernald), Special Collections and University Archives, University of Massachusetts, Amherst.

  51. BEUSC, vol. 2, p. 742.

  52. Grote, “Character of Protection and Defense in Insects,” CE (July 1888): 155.

  53. Grote, “On Insects Feigning Death,” CE (June 1888): 120, and “Characters of Protection and Defense in Insects,” CE, 155.

  54. BEUSC, vol. 2, pp. 1036–37.

  55. Doherty to his mother, April 18, 1883, JMH. Doherty wrote to Henry Elwes of his theory on “timidity as a source of protection” in 1889, which Elwes published in his article on Doherty’s butterflies for Transactions of the London Entomological Society (1891): 256–57.

  56. Grimaldi and Engel, Evolution of Insects, 603. For a recent account of all forms of defenses, see Graeme D. Ruxton, et al., Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals, and Mimicry (New York: Oxford University Press, 2004).

  57. Henry Bates, “Contributions to an Insect Fauna. Lepidoptera: Heliconidae,” Transactions of the London Entomological Society 23 (1862): 495–526. This discussion of Bates and mimicry owes a good deal to William C. Kimmler, “Mimicry: Views of Naturalists and Ecologists Before the Modern Synthesis,” in Dimensions of Darwinism, ed. Marjorie Grene (Cambridge: Cambridge University Press, 1986), 98–127. I would like to thank Mary C. Winsor of the University of Toronto for directing me to the source.

  58. On Bates, see Ruxton, et al., Avoiding Attack, 139–63. See also Thomas Belt, The Naturalist in Nicaragua (1874; repr., London, 1888); Major G. F. L. Marshall and Lionel de Nicéville, Butterflies of India (1881–82), vol. 1, p. 22; James J. Wood-Mason, “Description of Two New Species of Papilio from Northeastern India, with Preliminary Indication of an Apparently New and Remarkable Case of Mimicry Between the Two Distinct Groups Which They Represent,” Annals and Magazine of Natural History, 5th ser., 9 (1882); and Roland Trimen, “On Some Remarkable Mimetic Analogies Among African Butterflies,” Transactions of the Linnean Society 26 (1868): 497–522.

  59. Excursus 62, “Color Relations of Chrysalids and Their Surroundings,” BEUSC, vol. 2, pp. 1578–94. For the Wallace quote, see Andrew Berry, ed., Infinite Tropics: An Alfred Russel Wallace Anthology (London: Verso, 2002), 98–102, and David West, Fritz Müller: A Naturalist in Brazil (Blacksburg, VA: Pocahontas Press, 2003), 224–25. The Germans, too, had much to say, above all, Fritz Müller, an immigrant to Brazil living in Santa Catarina who, in the late 1870s, developed his own striking theory second only to Henry Bates’s theory in historical significance; Müller discovered that many unpalatable species mimicked the color and patterns of other unpalatable and often unrelated species, with a natural advantage accruing to both only if the number of mimics remained smaller than the models. On Müller, see Ruxton, et al., Avoiding Attack, 115–31, and West, Fritz Müller.

  60. Edward Poulton, Colour of Animals (London, 1889), viii.

  61. Gerald H. Thayer, Concealing-Coloration in the Animal Kingdom: An Exposition of the Laws of Disguise Through Color and Pattern—Being a Summary of Abbott H. Thayer’s Disclosures, with an introductory essay by A. H. Thayer (New York, 1909/1918), 3–12; and BEUSC, vol. 2, chap. 26, “Butterflies and Moths,” pp. 212–40. On Thayer, see Sharon Kingsland, “Abbott Thayer and the Protective Coloration Debate,” Journal of the History of Biology 11, no. 2 (1978): 223–44; Hardy Blechman, Disruptive Pattern Material: An Encyclopedia of Camouflage (Tonowanda, NY: Firefly Books, 2004), 16–76; Nelson White, Abbott H. Thayer: Painter and Naturalist (Hartford: Connecticut Printers, 1951), 6–7; and Blum, Picturing Nature, 336–44. On Riley’s discoveries, see his “Imitative Butterflies,” American Entomologist 1, no. 10 (1869): 189–93.

  62. Grote, An Illustrated Essay on the Noctuidae of North America (London, 1882), 35. Grote quoted, word for word, from an earlier article by D. S. Kellicott, “An Example of Protective Mimicry,” which Grote had earlier published in the North American Entomologist 10 (1879): 1.

  63. Grote, “Notes on Species of Lepidoptera,” CE (March 1887): 52–54.

  64. Excursus 57, “Nests and Other Structures Made by Caterpillars,” BEUSC, vol. 2, pp. 1456–57; and excursus 74, “Odd Chrysalids,” vol. 2, p. 1750.

  65. Henry Smith to Samuel Henshaw, April 14, 1887, EML.

  66. Doherty, “A List of Butterflies Taken in Kumaon,” 106.

  67. Henry Elwes, “List of Diurnal Lepidoptera Taken by Mr. W. Doherty (of Cincinnati) in Burma,” Transactions of the London Entomological Society 21 (1891): 255.

  68. Excursus 8, “The Means Employed by Butterflies of the Genus Basilarchia for Perpetuation of the Species,” BEUSC, vol. 1, pp. 260–94.

  69. William Henry Edwards rejected the idea that the two purples were different species, arguing, instead, that they were the same species, with the red-spotted a form of the white admiral. Naturalists today agree with Edwards. See Paul Opler, Eastern Butterflies (New York: Houghton Mifflin, 1992), 189. Opler also rejects Scudder’s genus, Basilarchia, preferring instead the older European designation of Limenitis fabricius, which Edwards also embraced.

  70. Frederick Clarkson, “Probable Origin of the Word Butterfly,” CE 17 (March 1885): 45.

  71. Excursus 23, “Mimicry and Protective Resemblance; or, Butterflies in Disguise,” BEUSC, vol. 1, p. 719. This excursus was the longest Scudder wrote and the only one given a lengthy bibliography. See also his “Color Relations of Chrysalids and Their Surroundings,” vol. 2, pp. 1578–89, in which he praises the work of Edward Poulton.

  72. Excursus 26, “Hypermetamorphosis in Butterflies,” BEUSC, vol. 1, pp. 804–8.

  73. BNA, vol. 1, “Melitea I.”

  74. Edwards to Scudder, March 4, 1876, SS-BMS. The conflict over the E. phaeton webs is chronicled in the following Edwards letters to Scudder: February 3, February 14, February 17, February 25, and March 4, 1876, SS-BMS.

  75. He had discovered the adult form of the spring azure (as it was also called, although not by Edwards). See Tom Allen, The Butterflies of West Virginia and Their Caterpillars (Pittsburgh: University of Pittsburgh Press, 1997), 104–5. For a modern analysis of the relationship between ants and Lycaenidae, see Naomi Pierce et al., “The Ecology and Evolution of Ant Association in the Lycaenidae (Lepidoptera),” Annual Review of Entomology 47 (2002): 733–71.

  76. Edwards, “Notes on Lycaena pseudargiolous and Its Larval History,” CE (January 1878): 1–15.

  77. Edwards, Entomological Diary, June 10, 1878, WHE-SA.

  78. William Henry Edwards to Henry Edwards, November 22, 1877, HE. In his introduction to volume 1 of BEUSC, in a subsection titled, “The Larva or Caterpillar” (pp. 15–16), Scudder attributed the discovery of the relationship between the ants and lycaenid larvae to two Germans, Christian Pezold and Eugen Esper, in the mid-eighteenth century.

  79. Doherty, “A List of Butterflies Taken in Kumaon,” 112–13.

  80. “Lycaena, II, III,” BNA.

  81. Excursus 21, “Companionship and Commensalism Among Caterpillars,” BEUSC, vol. 1, pp. 672–73; and excursus 35, vol. 2, pp. 962–64.

  82. Grote, “The Origin of Ornamentation in the Lepidoptera,” “The Friends and Associates of Caterpillars,” BEUSC, 114–17.

  83. See Karl Polyanyi, The Great Transformation (Boston: Beacon, 2001).

  84. Karen Halttunen, Confidence Men and Painted Women: A Study of Middle-Class Culture in America, 1830–1870 (New Haven, CT: Yale University Press, 1982), especially chapter 2, “Hypocrisy and Sincerity in a World of Strangers,” 33–55.

  85. Scudder, “Mimicry and Protective Resemblance; or Butterflies in Disguise,” BEUSC, vol. 1, pp. 713–19.

  86. As it turned out, however, Behr was wrong: the
butterfly survived into the 1940s. See Robert Pyle, “Bring Back the Xerces Blue,” Whole Earth (Spring 2001).

  87. Grote, “Collecting Noctuidae by Lake Erie,” Entomologist’s Record 6, no. 5 (1895): 100.

  88. Ibid., 98, and Grote’s introduction to Genesis I-II: An Essay on the Bible Narrative of Creation (New York, 1880), 17. I discovered the Rousseau quote in Jacques Roger, Buffon: A Life in Natural History, trans. Sarah Lucille Bonnefoi (Ithaca, NY: Cornell University Press, 1997), 301–2.

  89. Excursus 37, “Local Butterflies,” BEUSC, vol. 2, pp. 884–85; excursus 46, “The Spread of a Butterfly in a New Region,” BEUSC, vol. 2, p. 1175; and excursus 27, “The Best Localities for Collectors, BEUSC, vol. 2, pp. 817–19.

  90. Emory L. Kemp, The Great Kanawha Navigation (Pittsburgh: University of Pittsburgh Press, 2000), 24–62.

  91. See Scudder’s account of the life history Euphydryas phaeton in BEUSC, vol. 1, p. 705; and Edwards, Entomological Diary, May 22 and June 1, 1884, WHE-SA.

  92. Excursus 43, “Color Preferences of Butterflies: The Origins of Color in Butterflies,” BEUSC, vol. 2, pp. 1101–4.

  93. Excursus 15, “The Origins and Development of Ornamentation in Butterflies,” BEUSC, vol. 1, p. 517. Scudder’s preference for the beauty of butterflies compared to the aesthetic preferences of Charles Valentine Riley, also a gifted entomologist wide-ranging in his insect interests. To Scudder he wrote: “You know I see more beauty in the Heterocera and the other Orders than you do, so that my observations on the Rhopalocera are like stray shots—few and scattering.” See Riley to Scudder, 187?, first page of letter missing, written from the “office of the State Entomologist” of Missouri, St. Louis, SS-BMS.

  94. Excursus 70, “Sexual Diversity in the Form of the Scales,” BEUSC, vol. 2, pp. 1681–82.

  95. Ibid.; and Scudder, Butterflies, 235–38. In the mid-twentieth century, the great Swiss biologist Adolph Portmann would take a position very similar to Scudder’s, in defiance of the prevailing evolutionary wisdom. Portmann, too, accepted natural selection as causal but only for a special group of species that relied on protective coloration, warning signals, and deceptive resemblance, “a one-sided curio cabinet not to be compared to the wild profusion of form production in the garden of living creatures.” He also noted, “The scales of butterflies give the appearance of metal, of gold and blue textures, appearances whose sense surpasses any merely adaptive ability.” See Adolph Portmann, Essays in Philosophical Zoology (Lewiston, NY: Edwin Mellen, 1990), 27, and Animal Forms and Patterns: A Study of the Appearances of Animals (New York: Schocken, 1967), 122–23, 218–21. I would like to thank the noted lepidopterist Arthur Shapiro of the University of California, Davis, for introducing me to Portmann’s work.