Wallace, Darwin, and the Theory of Natural Selection: a Study in the Development of Ideas and Attitudes Author(S): Barbara G

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Wallace, Darwin, and the Theory of Natural Selection: A Study in the Development of Ideas and Attitudes Author(s): Barbara G. Beddall Source: Journal of the History of Biology, Vol. 1, No. 2 (Autumn, 1968), pp. 261-323 Published by: Springer Stable URL: https://www.jstor.org/stable/4330498 Accessed: 24-07-2018 01:35 UTC

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A Study in the Development of Ideas and Attitudes

BARBARA G. BEDDALL

2502 Bronson Road, Fairfield, Connecticut

INTRODUCTION

On 1 July 1908 the Linnean Society of London commemorated the reading before the Society fifty years earlier of the Darwin- Wallace joint papers, "On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection." 1 On the first occasion only some thirty Fellows and guests had been present at a quiet, unheralded meeting; the authors themselves were absent. Now there was a large and distinguished gathering celebrating the historic event. Two of the original cast were present, the naturalist Alfred Russel Wallace (1823-1913) and the botanist Sir Joseph Dalton Hooker (1817-1911). The other two, the biologist Charles Robert Darwin (1809-1882) and the geologist Sir Charles Lyell (1797-1875) had been dead for many years. Hooker, now a venerable nonagenarian, spoke of his "half- century-old real or fancied memories" of that June in 1858 when his old friend Darwin received Wallace's paper on natural selection. He based his account on Sir Francis Darwin's Life and Letters of Charles Darwin, remarking with some uneasiness that, beyond the letters from Darwin to himself and to Lyell, no other documentary evidence existed of the events of those turbulent weeks before the reading of the papers. Despite a search, the letters to Darwin from Hooker and Lyell could not be found, "and, most surprising of all, Mr. Wallace's letter and its enclosure have disappeared." 2

1. Linnean Society of London, The Darwin-Wallace Celebration held on 1st July, 1908, by the Linnean Society of London (London; The Society, 1908). 2. Francis Darwin, ed., The Life and Letters of Charles Darwin, including an Autobiographical Chapter (London: Murray, 1887; reprinted

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Hooker was troubled by the meagerness of the evidence, but Marchant, Wallace's first biographer, was unconcerned, and most people have concurred in his opinion that the eight extant letters received from Darwin while Wallace was in the Malay Archipelago "explain themselves and reveal the inner story of the independent discovery of the theory of Natural Selection." 3 But do they? A second question pertains to the relationship between Wal- lace and Darwin. Their recollections are often taken to be accurate reflections of their earlier thoughts and actions, but details may have been altered and the emphasis changed. Both men did come to play the roles assigned to them by history, but the making of the myth, in which they both participated, has obscured some of the facts. This study will emphasize Wallace and the influences on him. It will attempt to disentangle the various lines of evidence, to trace Wallace's progress toward the discovery of the theory of natural selection, to throw some light on what happened both before and after June 1858, and to suggest some alterna- tives to commonly accepted theories about these events. It is based on a re-evaluation and reinterpretation of contemporary sources, both published and unpublished. Because the aim has been to concentrate on primary source material, exhaustive reference to every author who has written since on these subjects has not been attempted. In particular, three of Wallace's published papers are considered in detail: "On the Law which has regulated the Intro- duction of New Species," "Note on the Theory of Permanent and Geographical Varieties," and "On the Tendency of Varieties to depart indefinitely from the Original Type." A number of previously published letters bear on questions raised here. For ease of reference, information about these letters has been arranged in tabular form in the Appendix and the letters numbered consecutively. They will be referred to by numbers in brackets in the text, with further discussion when required in appropriate footnotes. Use has also been made of a manuscript notebook kept by Wallace during his travels in the Malay Archipelago, and I am grateful to Mr. Thomas O'Grady and the Council of the Linnean Society of London for permission to quote from it.

in 2 vols., New York, Basic Books, 1959); Linnean Society of London, The Darwin-Wallace Celebration, p. 16. See also notes 115 and 139. 3. James Marchant, Alfred Russel Wallace: Letters and Reminiscences (New York: Harper, 1916), p. 105.

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I. AN INQUIRING MIND

He who in place of reasoning, employs authority, assumes that those to whom he addresses himself are incapable of forming a judgment of their own. If they submit to this insult, may it not be presumed they acknowledge the justice of it?

Wallace, "Notebook, 1855-1859," from Jeremy Bentham's Book of Fallacies4

Not until 1841, when he was eighteen years of age, did Alfred Russel Wallace, the eminent naturalist, begin his solitary study of the natural world around him. As a frequently unemployed and always impecunious surveyor, he turned to the study of plants to fill his leisure time:

But what occupied me chiefly and became more and more the solace and delight of my lonely rambles among the moors and mountains, was my first introduction to the variety, the beauty, and the mystery of nature as manifested i the vegetable kingdom.5 Wallace's early years and education were quite undistin- guished. The eighth child of an increasingly impoverished family, he was born on 8 January 1823 in the remote village of Usk in Monmouthshire, Wales. When he was five, the family moved to Hertford, near London, and it was at the Hertford Grammar School that he received his "very ordinary education." This ended when he was almost fourteen, and after that he was more or less on his own. Despite his commonplace upbring- ing, however, he had received a priceless gift from his father: a love of books and reading-a key to the world for anyone who wants to use it. After a few months spent with his brother John in London in the spring of 1837, Wallace joined his oldest brother, Wil- liam, to learn land surveying. But these were lean years for William, just before the rush of activity brought on by the construction of railroads, and he often had difficulty in finding enough work for himself and his younger brother. During one lull in 1839, Wallace spent some months learning the watch- making trade. Fortunately, business changes brought this to an end before Wallace was formally apprenticed, and he returned to surveying with William.

4. Alfred Russel Wallace, "Notebook, 1855-1859," MS, Linnean So- ciety of London, p. 102. This and other quotations are reproduced with permission from the Wallace and other manuscript material in the Library of the Linnean Society of London. 5. Alfred Russel Wallace, My Life: A Record of Events and Opinions (New York; Dodd, Mead, 1905), I, 191.

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Two years later, in 1841, Wallace purchased his first book on natural history, a shilling pamphlet on botany published by the improbably named Society for Promoting the Diffusion of Useful Knowledge. This quickly became his constant com- panion on his rambles through the countryside. Eager now to learn more, he was attracted by an advertisement for a textbook by one of England's experts, The Elements of Botany by John Lindley. But this expensive purchase, which arrived in July 1842, was a disappointment because it described all the orders of plants without indicating the British species. Not deterred, Wallace, with the aid of Loudon's Encyclopaedia of Plants, set about annotating his copy of Lindley, thereby giving himself the rudiments of a botanical education.6 How Wallace first happened upon Darwin's Voyage of the Beagle is not known, but he tipped quotations from the first edition into his copy of Lindley. He was apparently struck by Darwin's comment that to receive the fullest enjoyment from the passing scene, "a traveller should be a botanist, for in all views plants form the chief embellishment." 7 Still another purchase made about the same time was Swainson's Treatise on the Geography and Classification of Animals, a remarkable assembly indeed for someone with no biological background whatever.8 William's business did not improve, and Wallace again left to look for something else. Early in 1844 he settled upon teaching at the Collegiate School in Leicester. Teaching proved enjoyable enough, but this period is important for other reasons. The town of Leicester boasted a good library, and Wallace was soon spending his free time there reading, among other things, Humboldt's Personal Narrative of Travels and Malthus' Essay on the Principle of Population.9

6. John Lindley, The Elements of Botany, Structural, Physiological, Systematical, and Medical: Being a Fourth Edition of The Outline of the First Principles of Botany (London: Taylor and Walton, 1841); Wallace's copy is in the library of the Linnean Society of London. John Claudius Loudon, Encyclopaedia of Plants (London: Longman, 1829-1840). 7. Charles Darwin, Journal and Remarks, 1832-1836 (vol. 3 of Robert Fitzroy's Narrative of the Surveying Voyages of His Majesty's Ships 'Ad- venture' and 'Beagle,' . . . London: H. Colburn, 1839), p. 604. Because the titles of the first and second editions differ, the short title used here for both will be Voyage of the Beagle. See also notes 39-41, 43 and Appendix, 69. 8. William Swainson, Treatise on the Geography and Classification of Animals (London: Longman, 1835). Wallace's copy is in the Library of the Linnean Society of London. 9. Alexander von Humboldt and Aim6 Bonpland, Personal Narrative of Travels to the Equinoctial Regions of the New Continent, tr. H. M. Wil- liams (London: Longman, 1814-1829); which edition Wallace read is

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Most important of all was the friendship of another young amateur naturalist, the entomologist Henry Walter Bates (1825- 1892). Bates, two years younger than Wallace, had similarly finished his formal education at the usual early age. But, though apprenticed to a hosiery manufacturer in Leicester, he had enrolled in the local Mechanics' Institute, one of many such schools set up for the further education of workingmen. Here he made the acquaintance of several naturalists and soon plunged enthusiastically into the study of entomology. By the time he met Wallace he had already published his first short paper.'0 For the first time Wallace had someone with whom to talk over his discoveries. And Bates soon introduced him to the wholly new world of insects. Wallace was overjoyed. In short order he equipped himself with collecting bottles and pins and a copy of Stephens' Manual of British Coleoptera and em- barked on this fascinating new study.'1 This happy period ended abruptly when Wallace's brother William died unexpectedly in February 1846.12 Wallace left the school to help settle his affairs, and in January 1847 his second brother, John, joined him in carrying on the business. Wallace was irked by the details of management, formerly handled by his oldest brother. And he was also cut off from his new-found friends in Leicester, as he wrote plaintively to Bates [1].13 The time was not wasted, however. Wallace's letters to Bates show the extraordinary progress in his reading [1-4I14: Dar- not known. Thomas Robert Malthus, An Essay on the Principle of Popula- tion, 6th ed. (London: Murray, 1826). Which edition Wallace read is not known; in 1908 he took his references from the 6th. According to De Beer ("Darwin's Notebook," part 4, 30n), this is the edition that Darwin read. 10. Henry Walter Bates, "Note on Coleopterous Insects Frequenting Damp Places," Zoologist, 1 (1843), 114-115. 11. James Francis Stephens, Manual of British Coleoptera (London: Longman, 1839). 12. There is some question about the date of William's death. In Wal- lace's autobiography (My Life, I, 239), he gives it as 1846, but he also said he spent only one year at Leicester and correspondingly longer at Neath. In an attempt to straighten out this discrepancy, Poulton persuaded Wallace's son that William must have died in 1845; see E. B. P[oulton], "Alfred Russel Wallace, 1823-1913," Proc. Roy. Soc. London, [95B] (1923-1924), viiin. 13. Numbers in brackets refer to the letters listed in the Appendix. Where additional discussion of the letters is required, it will be found in appropriate footnotes. 14. The dates of letters nos. 2 and 3 are uncertain, and there is no com- pelling evidence to settle the matter. McKinney, following Clodd, has placed them in 1845; see H. Lewis McKinney, "Alfred Russel Wallace and the Discovery of Natural Selection," J. Hist. Med. Allied Sci., 21 (1966),

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL win again, Lyell's Principles of Geology, Vestiges of the Natural History of Creation by the then anonymous author, Robert Chambers, Lawrence's Lectures on Man, and Prichard's Physical History of Man.15 Wallace was already caught up in the more philosophical questions of evolution, the origin and distribution of species, and the differences between species and varieties. What a giant step forward from the shilling pamphlet on botany purchased only six years beforel But an even greater step was in store.

Late in 1847, Wallace and Bates read an "unpretending volume," A Voyage up the River Amazon, by a young American amateur naturalist, W. H. Edwards.'6 Already dissatisfied with their prospects at home, they wondered if they could make such a trip. Inquiries at the British Museum about the feasi- bility of the scheme brought assurances that there was a ready market for anything they might collect in this little-known region. At once they determined to go. But even as early as this their interests were not limited to collecting. Wallace was already interested not only in the differences between species and varieties, but also in the origin of species, and just before they left he wrote to Bates that he had begun to "feel rather dissatisfied with a mere local collection; little is to be learnt by it. I should like to take some one family to study thoroughly, principally with a view to the theory of the origin of species. By that means I am strongly of opinion that some definite results might be arrived at [3, 4]." 17 The two young adventurers left England in April 1848, arriving a month later at Pari (now Belem) at the mouth of the

337 n25. See also note 12. Marchant, in his Wallace, pp. 73-74, puts them in 1847, and that date is used here. 15. Charles Lyell, Principles of Geology: or the Modern Changes of the Earth and Its Inhabitants (London: Murray, 1830-1833, and later editions). Robert Chambers, Vestiges of the Natural History of Creation (New York: Wiley and Putnam, 1845, from the 1st English ed., 1844). Darwin, among others, held a low opinion of the Vestiges; see Appendix, 14 and note 92. William Lawrence, Lectures on Physiology, Zoology, and the Natural History of Man (London: Printed for J. Callow, 1819, and later editions). James Cowles Prichard, Researches into the Physical History of Man (London: J. & A. Arch, 1813, and later editions). Darwin also read both Lawrence and Prichard (De Beer, "Darwin's Notebook," part 2, 107n). 16. William Henry Edwards, A Voyage up the River Amazon (London: Murray, 1847). 17. See Appendix; also note 97. McKinney ("Wallace and Natural Selection," p. 337) has also pointed out Wallace's early interest in species, but he overlooked the significance of the related question of species and varieties; see Sec. III and note 105.

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Amazon River. (It is worth noting that they had no benefactors, financial or otherwise, and that they had no resource but themselves. Darwin, on the other hand, was attached to an official government mission and was able to pay his personal expenses himself.) Wallace remained for four years, returning to Eng- land in October 1852; Bates stayed eleven, not arriving home until July 1859. For the first few months, while they were becoming familiar with their new surroundings, they traveled together. Then they parted company, each traveling independently from then on. Wallace journeyed up the Amazon and the Rio Negro to the place where the latter joins the Orinoco, a spot already made famous as the farthest point reached by Humboldt com- ing from the other direction. Bates chose to go to the Upper Amazon, spending considerable time at Ega, 1400 miles up- stream from the Atlantic Ocean. These intrepid explorers were entirely on their own in the strange and oftentimes forbidding tropics of Brazil, contending with an uncertain supply of food, hazardous travel, indolent or dishonest natives (spoiled, they thought, by being half-civilized), illness, and isolation. They financed their exploits by the sale in Europe of duplicate collections, mainly of insects. Unfortunately, the scientific and financial results of Wal- lace's travels were disappointingly meager, but with reason. Owing to a misunderstanding, the specimens collected during the last two years were still waiting unshipped when he returned to Barra (now Manaus) on his way back to England. And so, unhappily, these as well as the richest part of his private collections were on board with him when the ship caught fire at sea. Nearly everything was lost as passengers and crew rushed to leave the burning vessel, some 700 miles off Ber- muda. Wallace and his companions were forced to spend ten days in open boats before they were finally rescued by a passing freighter. As a consequence of this disaster, Wallace had to piece together the few papers and two books that he wrote on his return home from letters, a few rescued notes, and his memory. He pursued his growing interest in the geographical distribution of animals in papers on the monkeys and the butterflies of the Amazon Valley, remarking on limits and barriers to their distribution and on the importance of labelling specimens with the exact locality where they were found for the proper study of this distribution.18 Besides a small book on the palm 18. Alfred Russel Wallace, "On the Monkeys of the Amazon," Proc. Zool. Soc. London, 20 (1852), 107-110; and "On the Habits of the Butter-

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL trees of the region, he also wrote the story of his expedition, A Narrative of Travels on the Amazon and Rio Negro.19 It is a tale of high adventure but, due to the loss of so much of his material, it is less rewarding from a scientific point of view. Bates was more fortunate, although he took good care to send his last collections home on three ships instead of one. On his return, he wrote a series of papers on the insect fauna of the Amazon Valley, in one of which he developed his famous theory of protective coloration, still known as Batesian mim- icry.20 His account of his travels, The Naturalist on the River Amazons, was more scientific than Wallace's and also, to Wal- lace's chagrin, more successfUl.21

But Wallace's misfortune proved a boon after all. He had counted on the money from the sale of the lost collections; without it he faced a return to surveying or another trip. For- tunately for both himself and the world, he chose the latter course, deciding after much study that the Malayan region offered the opportunities he sought. And so, in the spring of 1854, he set off again, on the eight-year exploration of the tropics on the other side of the world "which constituted the central and controlling incident of my life." 22 Once again Wallace had to make do with things as he found them, accommodate himself to local customs, learn native lan- guages, find food and shelter where he could, and use any available means of travel. As a practical matter, the weather and traveling arrangements largely determined his itinerary. The first two years were spent around Singapore and in Borneo, the next five in the area from Celebes to northern New Guinea, and the last year in Timor, Java, and Sumatra. A particular aim this time was a more thorough investiga- tion of the problems of the geographical distribution of animals. Wallace had chosen well, for the lessons of an island world flies of the Amazon Valley," Trans. Entomol. Soc. London, N.S. 2 (1852- 1853), 253-264. 19. Alfred Russel Wallace, Palm Trees of the Amazon and Their Uses (London: J. Van Voorst, 1853); and A Narrative of Travels on the Ama- zon and Rio Negro (London: Reeve, 1853). 20. Henry Walter Bates, "Contributions to an Insect Fauna of the Ama- zon Valley. Lepidoptera: Heliconidae," Trans. Linn. Soc. London, 23 (1862), 495-515. 21. Henry Walter Bates, The Naturalist on the River Amazons (London: Murray, 1863). Many of Bates' more interesting comments on the origin of species and on geographical distribution were omitted from the 2nd abridged edition (1864). 22. Wallace, My Life, I, 336.

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are even more vivid than those of a continental region; Darwin had also discovered this when he visited the islands of the Galapagos Archipelago. Although Wallace was again support- ing himself by the sale of duplicate collections, his theoretical interests were never far from his mind. The most spectacular result of this second trip was the independent discovery of the theory of natural selection, but this was only one of many important contributions to evolutionary theory. And happily enough, his discerning and well- written account of the region, The Malay Archipelago, became a worthy rival to Darwin's Voyage of the Beagle and Bates's Naturalist on the River Amazons.23 The three papers of special importance in tracing the development of Wallace's ideas on natural selection will now be considered in detail.

II. THE "LAW"

To discover how the extinct species have from time to time been replaced by new ones down to the very latest geological period, is the most difficult, and at the same time the most interesting problem in the natural history of the earth. The present inquiry, which seeks to elim- inate from known facts a law which has determined, to a certain degree, what species could and did appear at a given epoch, may, it is hoped, be considered as one step in the right direction towards a complete solution of it.

Wallace, "On the Law which has regulated the Introduction of New Species"

Wallace's "powerful essay," 24 "On the Law which has regulated the Introduction of New Species," was published in Sep- tember 1855, but to his disappointment it attracted little notice at the time.25 He had written it in the preceding February during a lull in his collecting activities, induced by the publication of Edward Forbes's theory of polarity, and he had hoped at the least for some comment from this brilliant young naturalist; unknown to Wallace, however, Forbes had died in November 1854. Except for Bates's, other response was mini- mal, if not disparaging. Wallace phrased his 'law" as follows: Every species has come into existence coincident both in space and time with a pre-existing closely allied species.26 Although the evolutionary

23. Alfred Russel Wallace, 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). 24. Thomas Henry Huxley, in Darwin, Life and Letters, 1, 539. 25. Alfred Russel Wallace, "On the Law which has regulated the Intro- duction of New Species," Ann. Mag. Nat. Hist. [2], 16 (1855), 184-196.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL implications of this statement are obvious, Wallace at this time had no suggestions on a mechanism of change. His speculations were rooted in his long-standing interest in the geographical distribution of animals and colored by his extensive practical experience in the tropics of two hemispheres. The study of zoogeography, in which Wallace was to become an acknowledged master, was still clouded in mystery when he first came upon it in Swainson's Treatise on the subject:

We may, indeed, build a theory upon every thing in nature: but the more we investigate, the stronger will be our con- viction in the following deduction: -That the primary causes which have led to different regions of the earth being peopled by different races of animals, and the laws by which their dispersion is regulated, must be for ever hid from human research [to which Wallace wrote "no" in the mar- gin]. This conclusion is strengthened by the inference which will be drawn from the facts we shall subsequently state; an inference so well expressed by a very intelligent writer, that we shall give it nearly in his own words. "It appears that various tribes of organised beings were originally placed by the Creator in certain regions, for which they are by their nature peculiarly adapted [Wallace's underlining]." 27

In a modification of the quinarian system of William Sharpe Macleay, Swainson divided the earth into five regions according to what he believed to be the five major races of mankind; animal groups were likewise divided into fives. The divisions were mathematical, the reasons not only unknown but un- knowable. But Wallace questioned Swainson from the first, noting that "there appears not to be the slightest reason for believing a priori that all groups of animals are divided into the same number of types of forms or divisions .. 28 Over the years Wallace's ideas matured. Traces of this can be found, as mentioned earlier, in his papers on the monkeys and the butterflies of the Amazon Valley, as well as in his Narrative, but he was undoubtedly handicapped by the loss of so much of his South American material. The appearance of Forbes's paper spurred Wallace to collect and organize his thoughts, for Forbes's theory of polarity as an explanation of organic changes through geologic time seemed as untenable

26. Ibid., p. 186. 27. Swainson, Treatise on Geography, p. 9. 28. Note written by Wallace in his copy of Swainson's Treatise on Geography, p. 223.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection to him as had Swainson's theories about the present geographical distribution of animals. A pervading influence on Wallace, one might almost say the pervading influence, was Charles Lyell's Principles of Geology. Wallace had read Lyell at least as early as 1846, and he had accepted wholeheartedly Lyell's application of the principle of uniformitarianism to geology. But Lyell's influence was more profound than that. From the notebook kept by Wallace during his Malayan travels, it seems that he had with him a copy of the fourth edition of Lyell's Principles, and the discussion here refers to this edition.29 In a large measure, Wallace's "Notebook" is a long, private argument with Lyell, refuting many of the latter's biological theories. Wallace was thus stimulated to broaden and deepen his own thinking. It should be noted in Lyell's defense, however, that he was neither a trained biologist nor a collector of living specimens (although he had collected insects as a youth), and that he lacked the extensive first-hand experience with living things acquired through long years by a Lamarck, a Darwin, or a Wallace. It has often been said that Lyell nearly stumbled onto evolutionary theory himself; it has even been suggested that he was an evolutionist in secret, at least at first.30 But in fact he was far removed at many critical points. He had missed altogether the crux of Wallace's paper-the relationship of species in both time and space-and so a theory of descent was not only not a logical deduction, it was irrelevant. On his own terms he believed he had applied the principle of uniformitarianism to organic changes by showing the gradual extinction and creation of species. But he believed firmly in the stability of species, scarcely a good foundation for a theory of evolution. Lyell was well acquainted, however, with Lamarck's theory of the transmutation of species, having first read it in 1827. Although he found the ideas provocative, he confessed in a letter to Mantell that "I read him rather as I hear an advocate on the wrong side, to know what can be made of the case in good hands."31 He included a lengthy summary in his Prin-

29. Lyell, Principles of Geology, 4th ed. (London: Murray, 1835). See also note 4. 30. Gertrude Himmelfarb, Darwin and the Darwinian Revolution (Garden City, N.Y.: Doubleday, 1959), pp. 180-189. An early proponent of this point of view was Thomas Huxley, in Darwin, Life and Letters, I, 543-548. 31. Katherine Lyell, ed., Life, Letters and Journals of Sir Charles Lyell (London: Murray, 1881), I, 168.

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ciples, at first in protest against the pretension that species were "capable . . . of being indefinitely modified in the course of a long series of generations," but later to give due honor to Lamarck and his evolutionary theories.32 Wallace and Dar- win both read Lyell's summary. Parenthetically, it might be observed that Lyell accepted to a limited degree what later generations have considered the hallmark of Lamarckism, the inheritance of acquired characteristics. In Lyell's case, this meant the possible inheritance of acquired habits that remained within the strict limits of predetermined variation. To a limited extent, Darwin was also to accept this Lamarckian tenet, but Wallace never did. Lyell's view of geographical distribution was more compre- hensive than Swainson's. He had broadened the scope to in- clude time and change, although there was no hint of evolu- don:

If the views which I have taken are just, there will be no difficulty in explaining why the habitations of so many species are now restrained within exceedingly narrow limits. Every local revolution, such as those contemplated in the preceding chapter, tends to circumscribe the range of some species, while it enlarges that of others; and if we axe led to infer that new species originate in one spot only, each must require time to diffuse itself over a wide area. It will follow, therefore, from the adoption of this hypothesis, that the recent origin of some species, and the high antiquity of others, are equally consistent with the general fact of their limited distribution, some being local, because they have not existed long enough to admit of their wide dissemination; others, because circumstances in the animate or inanimate world have occurred to restrict the range which they may once have obtained. As considerable modification in the relative levels of land and sea have taken place in certain regions since the existing species were in being, we can feel no surprise that the zoologist and botanist have hitherto found it difficult to refer the geographical distribution of species to any clear and determinate principles, since they have usually speculated on the phenomena, upon the assumption that the physical geography of the globe had undergone no material altera- tion since the introduction of the species now living. So long as this assumption was made, the facts relating to the

32. Lyell, Principles of Geology, lst ed. (1832), II,1; 10th ed. (1868), II, 246.

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geography of plants and animals appeared capricious in the extreme, and by many the subject was pronounced to be so full of mystery and anomalies, that the establishment of a satisfactory theory was hopeless.33

Although Lyell's biological theories were ambiguous and ill- defined, his contributions to geology had added new dimensions to the history of the earth. In particular, he had expanded on the uniformitarian theory of the Scottish geologist James Hut- ton (1726-1797) that attempted "to explain the former changes of the earth's crust by reference exclusively to natural agents," 34 and he had vigorously attacked the commonly accepted belief that life on the earth had begun in the year 4004 B.C., as calculated by James Usher, Archbishop of Armagh. Wallace's efforts (and Darwin's too) would have been crippled without Lyell. Evolution was advocated, however, in a book published not long afterward, Vestiges of the Natural History of Creation. But its anonymous author, Robert Chambers, "a private person with limited opportunities for study," was no biologist either, and his popular work was ridiculed by scientists. Nev- ertheless, Wallace thought his hypothesis ingenious, as he wrote to Bates [2, 3]. For somewhat different reasons, two editions of Darwin's Voyage of the Beagle should also be included here. Wallace read the first edition (1839) quite early, perhaps as early as 1842 according to his first letter to Bates and the note in his copy of Lindley; and from evidence in the "Notebook," he took a copy of the second (1845) along with him to the Malay Archipelago.

Wallace began his paper "On the Law" by noting the long- continued series of geologic changes. Then, applying the uniformitarian principle to organic changes, he proposed "a like gradation and natural sequence from one geological epoch to another." 35 (Lyell had suggested the slow and gradual extinction and creation of species, but with no hint of descent.) From a series of propositions relating to "organic geography and geology," Wallace then deduced his 'law," which supported a hypothesis that might explain the past and present distribution of life upon the earth that had occurred to him, he said, about ten years earlier. First of the four main questions illuminated by Wallace's

33. Ibid., 4th ed., III, 165-166. 34. Ibid., 12th ed. (1875), 1, 73. 35. Wallace, "On the Law," p. 184.

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'law" was that perennial problem, "the system of natural af- finities." 36 If the 'law" were true, species would be related to closely allied species which had preceded them. This relationship could rarely be expressed for long by a straight line. The divergence, uneven rates of change, and extinction of species, complicated by the fragmentary fossil record, could be better represented by a branching "as intricate as the twigs of a gnarled oak." 37 Artificial systems of classification based on circles or a fixed number of divisions were unnecessary con- trivances. Lamarck would have agreed. He believed that species were defined by the gaps between them that were produced by extinction and by the fragmentary record, and that two or more diverging species would merge going backward in time. To Lyell this would have been unthinkable, convinced as he was of the real and permanent existence of species in nature. Next, in answer to the "singular phenomena" of the distribution of animals and plants in space, Wallace offered some original suggestions.38 He clearly recognized the part played by geographical isolation in the origin of peculiar forms of life in long-isolated places, and also the divergence from a widespread "antitype" that results in two or more representa- tive forms in different regions of the world. He brought forward as examples the problems posed by the inhabitants of both ancient and recent island groups and mountain ranges. Of particular interest are Wallace's suggestions regarding the Galapagos Islands, where the "phenomena . . . have not hitherto received any, even a conjectural explanation." 39 Dar- win had mentioned in the first edition of his Voyage of the Beagle the many peculiarities of distribution that he had found in this small archipelago, six hundred miles off the coast of Ecuador, concluding only: "But there is not space in this work, to enter on this curious subject." 40 Although Darwin expanded on this in the second edition of the Voyage, his remarks were again inconclusive:

The only light which I can throw on this remarkable difference in the inhabitants of the different islands, is, that very strong currents of the sea running in a westerly and W.N.W. direction must separate, as far as transportal by

36. Ibid., pp. 186-188. 37. "Divergence" was an important part of Darwin's theory. See also notes 105, 111-113, 125-127, and Wallace, "On the Law," p. 187. 38. Ibid., pp. 188-190. 39. Ibid., p. 188. 40. Darwin, Voyage of the Beagle (1839), p. 475.

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sea is concemed, the southern islands from the northern ones; and between these northern islands a strong N.W. current was observed, which must effectually separate James and Albemarle Islands. As the archipelago is free to a most remarkable degree from gales of wind, neither the birds, insects, nor lighter seeds, would be blown from island to island. And lastly, the profound depth of the ocean between the islands, and their apparently recent (in a geological sense) volcanic origin, render it highly unlikely that they were ever united; and this, probably, is a far more important consideration than any other, with respect to the geographical distribution of their inhabitants. Reviewing the facts here given, one is astonished at the amount of creative force, if such an expression may be used, displayed on these small, barren, and rocky islands; and still more so, at its diverse and yet analogous action on points so near each other. I have said that the Galapagos Archipelago might be called a satellite attached to America, but it should rather be called a group of satellites, physically similar, organically distinct, yet intimately related to each other, and all related in a marked, though much lesser degree, to the great American continent.41

In other words, these islands were geologically recent, separated not only by deep ocean but also by currents, and without winds strong enough to have blown birds, insects, or seeds from one island to another. With no means of dispersal and a limited amount of time, it is no wonder that Darwin seemed puzzled. Actually, his private thinking was much in advance of this public statement, for he had already worked out his theory of natural selection as an explanation of the "creative force" at work in the islands, but Wallace had no way of knowing this.42 Wallace suggested, on the contrary, that these were islands of high antiquity that had been peopled by the agency of wind and water, as other islands were peopled, and that, the pre- existing species having died out, only variously modified prototypes now remained. The islands of the Malay Archipelago differed in being separated by shallow seas (this was written shortly before Wallace discovered in the summer of 1856 the division between the Australian and Oriental Regions since

41. Charles Darwin, Journal of Researches into the Natural History and Geology of the Countries visited during the Voyage of H.M.S. Beagle round the World, 2nd ed. (London: Murray, 1845), p. 398. 42. Charles Darwin and Alfred R. Wallace, Evolution by Natural Selec- tion (Cambridge: Cambridge University Press, 1958), pp. 116-121.

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Although we have as yet considered one class only of the causes (the organic) by which species may become extermin- ated, yet it cannot but appear evident that the continued action of these alone, throughout myriads of future ages, must work an entire change in the state of the organic creation, not merely on the continents and islands, where the power of man is chiefly exerted, but in the great oceans, where his controul [sic] is almost unknown. The mind is pre-

43. Wallace, "Notebook," p. 60. See also notes 81, 103, and 162, and Appendix, 69. 44. Wallace, "On the Law," pp. 190-195.

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pared by the contemplation of such future revolutions to look for the signs of others, of an analogous nature, in the monuments of the past. Instead of being astonished at the proofs there manifested of endless mutations in the animate world, they will appear to one who has thought profoundly on the fluctuations now in progress, to afford evidence in favour of the uniformity of the system, unless, indeed, we are precluded from speaking of uniformity when we charac- terize a principle of endless variation.45

Lamarck, on the other hand, considered that only a few large land animals at most had become extinct, and then through the agency of man. Even earlier than Lyell he had disagreed with the catastrophists, who thought that life had been wiped out at various times by universal cataclysms. Lyell suggested a course of gradual extinction and creation of species. But La- marck had proposed that earlier species had gradually been changed into present-day species through transmutation; extinction thus played no important part in his system. Lyell, however, disagreed completely with Lamarck:

To pursue this train of reasoning farther is unnecessary; the geologist has only to reflect on what has been said of the habitations and stations of organic beings in general, and to consider them in relation to those effects which were contemplated in the second book, as resulting from the ig- neous and aqueous causes now in action, and he will immediately perceive that, amidst the vicissitudes of the earth's surface, species cannot be immortal, but must perish, one after the other, like the individuals which compose them. There is no possibility of escaping from this conclusion, without resorting to some hypothesis as violent as that of Lamarck, who imagined, as we have before seen, that species are each of them endowed with indefinite powers of modifying their organization, in conformity to the endless changes of circumstances to which they are exposed.46

Wallace steered a middle course. He agreed with Lyell that some species might become extinct, but he also agreed with Lamarck that modified prototypes might remain. If extinction were the rule, as Lyell suggested, then some sort of "creation" was necessary to fill the gaps. Here Lyell was quite vague, his hypothesis on the original introduction of species reading as follows: Each species may have had its origin in a single pair, or

45. Lyell, Principles of Geology, 4th ed., III, 140-141. 46. Ibid., 155-156.

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individual, where an individual was sufficient, and species may have been created in succession at such times and in such places as to enable them to multiply and endure for an appointed period, and occupy an appointed space on the globe.47

This creation of new species was not readily seen because it was an infrequent occurrence taking place over a long period of time at different places on the earth. Lamarck's solution to this problem was the mutability of species, though the agency he suggested-inheritance of characteristics acquired through the will of the animal-was scoffed at even in his own day. The author of the Vestiges had still another suggestion:

The idea, then, which I form of the progress of organic life upon the globe- and the hypothesis is applicable to all similar theatres of vital being-is, that the simplest and most primitive type, under a law to which that of like-production is subordinate, gave birth to the type next above it, that this again produced the next higher, and so on to the very highest, the stages of advance being in all cases very small-namely, from one species only to another; so that the phenomenon has always been of a simple and modest character.48

Chambers' proposal that one species gave birth directly to the next higher brought derision upon his book. Nevertheless, he should at least be credited with having emphasized a natural method. Wallace, however, had no theory on how extinct species were replaced. For the present, he confined himself to "what species could and did appear." He saw that this was neither a random process nor a straight line from simple to complex, as proposed by the proponents of the theory of progressive development. A theory of gradual change combined with his 'law" would, he thought, account for the observed facts and even for apparent retrogressions. Lyell, as a geologist familar with the anomalies of the fossil record, had protested against the theory of progressive development because he disputed the accuracy of the facts: It was before remarked, that the theory of progressive development arose from an attempt to ingraft the doctrine of the transmutationists upon one of the most popular gen- eralizations in geologv. But modem geological researches

47. Ibid., 99-100. 48. Chambers, Vestiges, 1st ed., p. 167.

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have almost destroyed every appearance of that gradation in the successive groups of animate beings, which was supposed to indicate the slow progress of the organic world from the more simple to the more compound structure.49

Although evolutionary trends do exist (and Lyell later accepted a general progression), the record is far from straight- forward, and its message is not easily read. The theory of evolution, with natural selection as an agent of change, was to make this record more comprehensible, for it postulates successful adaptation rather than any necessary progression. Wal- lace was still several years away from this answer, but he was heading in the right direction. It was Forbes's metaphysical theory of polarity that had prompted Wallace to set out his own thoughts.50 According to this theory, the distribution of organized beings (genera rather than species) in time manifested a quality known as polarity: an "arrangement in opposite directions with a development of intensity towards the extremes of each." 51 This relation was used to explain the larger number of generic forms in the earlier epochs of the Palaeozoic (Silurian and Devonian) and in the later epochs of the Neozoic (Cretaceous, Tertiary, and present), compared with the smaller number to be found in the intervening time. Wallace objected to this theory on several grounds. First of all, it invoked an obscure and hypothetical cause when "the facts may be readily accounted for on the principles already laid down." 52 In typical Lyellian style, he stressed the vast amounts of time involved and the action of natural causes, and he went on to propose that the rates of creation and extinction of species were related to unequal rates of geologic change, more species being created during quiet periods of the earth's long history and more becoming extinct during violent periods. Even more to the point was Wallace's criticism that Forbes's theory presupposed the completeness of the geological record (an error also made by those who believed in the theory of progressive development). Wallace knew it was fragmentary, but he would have been surprised to know that a century later fossils are still thought to represent less than one per cent of the species that have existed. Polarity's foundation was fragiele at best.

49. Lyell, Principles of Geology, 4th ed., III, 14-15. 50. Edward Forbes, "On the Manifestation of Polarity in the Distribu- tion of Organized Beings in Time," Proc. Roy. Inst. London, 57 (October 1854), 332-337. 51. Ibid., p. 336. 52. Wallace, "On the Law," p. 192.

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The fourth and final point of Wallace's paper was another subject that had attracted much attention, the puzzling problem of rudimentary organs.53 Lyell later admitted that he had missed their significance and as a result had omitted from his summary most of the examples given by Lamarck.54 This is not surprising, however, because Lamarck had interpreted them in his own light, believing them to be the result of "the permanent disuse of an organ, arising from a change of habits, [which caused] a gradual shrinkage and ultimately the disap- pearance and even extinction of that organ." 56 Wallace, like Chambers, thought that rudimentary organs showed relation- ships, but he misinterpreted them, confusing vestigial with nascent organs. He did, however, ask the right question: "If each species has been created independently, and without any necessary relations with pre-existing species, what do these rudiments, these apparent imperfections mean?" se Wallace ended his paper grandly: "Granted the law, and many of the most important facts in Nature could not have been otherwise, but are almost as necessary deductions from it, as are the elliptic orbits of the planets from the law of gravitation." 67 The response to it was hardly encouraging. More than two years passed before Bates's letter congratulat- ing him arrived from the Upper Amazon [9]. In the meantime, his agent, Samuel Stevens, wrote that he had heard several naturalists object to Wallace's "theorizing" when what was needed was more facts.

III. THE 'NOTE"

. . . why should a special act of creation be required to call into existence an organism differing only in degree from another which has been produced by existing laws? Wallace, "Note on the Theory of Permanent and Geographical Varieties"

In the fall of 1857 Wallace sent off to the Zoologist a short paper entitled "Note on the Theory of Permanent and Geo- graphical Varieties." 58 The first part of his hypothesis had been his 'law" on "what species could and did appear"; the

53. Ibid., pp. 195-196. 54. Lyell, Principles of Geology, 12th ed., II, 274. 55. Jean Baptiste Lamarck, Zoological Philosophy: an Exposition with Regard to the Natural History of Animals, tr. Hugh Elliot (London: Mac- milan, 1914; reprinted, New York: Hafner, 1963), p. 115. 56. Wallace, "On the Law," p. 195. 57. Ibid., p. 196. 58. Alfred Russel Wallace, "Note on the Theory of Permanent and Geographical Varieties," Zoologist, 16 (1858), 5887-5888.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection second part concerned the distinction between species and varieties, as he wrote to Bates in January 1858 [19]. This question still exists, of course, but the premises are entirely different. Then, species were real and permanent, "created" with certain relatively fixed characteristics; varieties, on the other hand, were produced by ordinary generation within strict limits of variation. Varieties were, if anything, an inconvenience, inter- fering with the rigid definition of species. Today, however, species have only a relative permanence; they also come about by ordinary generation, and varieties may (though they do not always) lead to the formation of new species. The definition of species has been a troublesome problem at least as far back as the time of Aristotle. Not until the time of the English naturalist John Ray (1628-1705) was the term limited to what we would today recognize as a breeding unit, including sex, color, and age variants. Ray concluded his discussion of species in his Historia Plantarum with the observa- tion that "animals that differ in species preserve their distinct species permanently; one species never springs from the seed of another nor vice versa." 59 Linnaeus (1707-1778) was strongly influenced by Ray. A clear notion of what was meant by the term "species" was a necessity in organizing his Systema Naturae, first published in 1735. At first he was convinced of the permanence of species, but later, after he had become acquainted with the vast number of new forms brought home by explorers from all over the world, he was not so sure. Nevertheless, the belief in the creation of permanent species, which was closely intertwined with religion, was the generally accepted opinion in the first half of the nineteenth century, in spite of the heresies of people like Lamarck and Chambers. This vexed question had attracted Wallace's attention from the start, as can be seen from his early letters to Bates [3, 4]. It became a matter of daily concern when, as a collector, he was confronted over and over again with the task of properly identifying specimens. Unhampered by any strong re- ligious convictions, he worried at the problem as a dog does a bone, as his "Notebook" attests. Lamarck had clearly grasped the gradual nature of change in species through time. Chambers, although he believed in evolution, thought that it proceeded by sudden leaps from species to species rather than by the accumulation of small changes. But Lyell stood on the opposite side of the fence,

59. Barbara G. Beddall, "Historical Notes on Avian Classification," Syst. Zool., 6 (1957), 134.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL firmly convinced of the fixity of species and of strict limita- tions to variation. Had either Chambers or Lyell been trained biologists or collectors, exposed day after day to the endless variations to be found in nature, they might have modified their views. But Wallace benefited from Lyell's position, using it as a springboard from which to develop his own. Although Lyell was unaware that he had been cast in the role of principal antagonist, his importance to Wallace cannot be over- rated. It seems likely that the "Notebook" was intended as the basis of a projected book about which Wallace wrote to Darwin in the fall of 1857 and to Bates early in 1858 [16, 19].60 The entries of particular interest here were written between June 1855 (not long after the formulation of his "law") and No- vember 1857, assuming that the dates scattered here and there are accurate indicators of the time and order of writing. They cover an assortment of topics relating to evolution-proofs of design, the theory of progressive development, transmutation and special creation, the "balance of species," geological changes and gaps in the fossil record, and the doctrine of the morphology of plants-and reflect Wallace's progress toward a solution of the problem of the origin of species.6' The argument from design was teleological, presuming that a contrivance existed in accordance with a preconceived plan. Adaptation between structure and function was recognized, but it was thought that a structure was provided simply because a function required it. Wallace wondered, however, how an animal could have necessities before it came into existence? And how could it "continue to exist unless its structure enabled it to obtain food?" 62 He thought that the arguments brought forward as proofs of design were absurd; not only were they insulting to the intelligence of a Supreme Being, but they also placed narrow limits on His power. Wallace returned several times to the inconsistencies in the geological record that made the theory of progressive development so troublesome for Lyell, observing that "the supposed contradictions all arise from considering it necessary that the highest forms of one group should appear before the lowest of the next succeeding, not considering that each group goes on

60. The information that Wallace's plan for his book is on the reverse side of the fragment about the jaguars was sent me by Sydney Smith. 61. Wallace, "Notebook," pp. 12-100. See also McKinney ("Wallace and Natural Selection," pp. 342-347), who has also pointed out that this is essentially a long argument with Lyell. 62. Wallace, "Notebook," p. 12.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection progressing after other groups have branched from it." M Lyell's static concept of species did not allow for such a dynamic interpretation. Were there connecting forms between groups, and if so, what were they? Wallace remarked that "as long as these most important characters [of groups] remain undiminished, no al- terations of external form or habits can be held to shew any signs of a transition." 04 He questioned the popularly held transitional status of the seal, asking, "is not the Cetaceous group rather a modification of mammalia to an aquatic life than a link connecting them with fishes?" f5 He noted that neither the bat nor the hummingbird is a transition form either, because each contains the characteristic features of mammal or bird in a high state of development. The bat's wing is even less like a bird's wing than are less modified forelimbs of other mammals. Therefore, the highest forms of one group cannot be a transition to the lowest forms of the next. Lyell had also had trouble in accounting for the appearance of new species. Having decided in favor of the stability of species, he was obliged to settle for their extinction and "creation," admitting to only a limited amount of variation. Wallace was groping his way forward, however, questioning Lyell's assumptions:

Lyell says that varieties of some species may differ more than other species do from each other without shaking our confidence in the reality of species-But why should we have that confidence? Is it not a nice prepossession or preju- dice like that in favour of the stability of the earth which he has so ably argued against? In fact, what positive evidence have we that species only vary within certain limits? . . . we have no proof how the varieties of dogs were produced. All varieties we know of are produced at birth, the offspring differing from the parent. This offspring propagates its kind. Who can declare that it shall not produce a variety, which process continued at intervals will account for all the facts?66

Not only did Lyell believe that the amount of variation was strictly limited, but he also believed that it occurred over a brief period, after which no more changes took place no matter how long a time passed. Wallace thought that "Mr. Lyell must be very perplexed to know this." 67 Lvell did concede.

63. Ibid., p. 38. 64. Ibid., pp. 77-78; comma added. 65. Ibid., p. 76. 66. Ibid., pp. 39-40; commas added. See also Lyell, Principles of Geology, 4th ed., II, 435. 67. Wallace, "Notebook," p. 42.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL that time might bring about any metamorphosis, but only if such change were evidenced by a wholly new sense or organ. Wallace objected that "this would be taking a leap with a vengeance. We should have to get out of one class or order into another passing through many thousand species-If this is supposed to prove a change from one species to another, it can never be proved." 88 Lyell opposed Lamarck's theory of the transmutation of species, but Wallace defended his doctrine of their indefinite modifiability:

Many of Lamarck's views are quite untenable & it is easy to controvert them, but not so the simple question of a species being produced in time from a closely allied distinct species, which, however, may of course continue to exist as long or longer than the offshoot. Changes which we bring about artificially in short periods may have a tendency to revert to the parent stock, though this in animals is not proved. This is considered a grand test of a variety. But when the change has been produced by Nature during a long series of generations, as gradual as the changes of Geology, it by no means follows that they may not be permanent, & thus true species produced.69

Wallace was to return later to the "grand test of a variety"; it became the opening gun in the essay he sent to Darwin from Ternate in February 1858. Lyell had made many contributions to the tangle of questions surrounding the geographical distribution of animals. But a central problem-the "creation" of species-muddled his conclusions, and Wallace put his finger on one of the inconsistencies to which this led:

Lyell occupies much space in shewing how the species which are common to different & distant countries, might have been carried from one to the other by a variety of acci- dents. But this has never been felt to be a difficulty. The matter of wonder has always been that in distant countries of similar climate so many should be different. This he gets over by special creation of the species each in one spot as they are wanted. This is no doubt a very easy way of getting over it, but just as philosophical as to say that fossils of existing species

68. Ibid., p. 43; comma added. See also Lyell, Principles of Geology, 4th ed., II, 414. 69. Wallace, "Notebook," pp. 44 45; some commas added. See also note 100.

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are remains of real animals while those which are not like any species now existing are special creations & not fossil animals at all.70

Wallace turned again to the peculiarities of distribution to be found on the Galapagos Islands. Lyell's theories were inadequate to the task of explaining them, as Wallace pointed out:

In a small group of islands not very distant from the mai land, like the Galapagos, we find animals & plants different from those of any other country but resembling those of the nearest land. If they are special creations, why should they resemble those of the nearest land? Does not that fact point to an origin from that land? Again in these islands we find species peculiar to each island, & not one of them containing all the species found in the others as would be the case had one been peopled with new creations & the others left to become peopled by winds, currents, etc., from it. Here we must suppose special creations in each island of peculiar species though the islands are all exactly similar in structure, soil, & climate, & some of them within sight of each other, a work of supererogation one would suppose, as they must inevitably in time become peopled from each other, & contrary to what takes place elsewhere-Ireland is peopled from England. It may be said it is a mystery which we cannot explain, but do we not thus make unnecessary systems and difficulties by supposing special creations contrary to the present course of nature? For we must conclude the course of nature in peopling islands in the ocean to be uni- form & that all islands distant from others should now be stocked with animals & plants equally peculiar. But we know this not to be the case. Volcanic islands recently produced & coral islands far in the ocean contai stragglers from the nearest land & no other, nothing peculiarl Now we can hardly suppose that islands would be left for ages to become stocked in this manner, & then the new & peculiar creations be introduced just when they were not wanted. According to Mr. Lyell's own arguments, they would hardly be able to hold their own against the previous occu- piers of the soil & there would have to be a special extermina- tion of them to make room for the new & peculiar species. We must therefore suppose that such islands as St. Helena & the Galapagos were stocked with their peculiar species

70. Ibid., pp. 45-46; see also Lyell, Principles of Geology, 4th ed., III, 22-97.

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immediately on their being raised from the ocean, & they would then have a chance of keeping out the new comers which might be blown accidentally on their shores. This supposition will certainly explain the present condition of those islands but it has the disadvantage of being contrary to the present order of nature, for none of the islands which we have any reason to believe have been formed, since a very late geological era, are inhabited by peculiar species. They generally have not one species peculiar to themselves. On the other hand, islands which are thus peculiarly inhabited, appear to be of a considerable antiquity [in a mar- ginal note, Wallace wrote, "this must be proved"]. A long succession of generations appears therefore to have been requisite, to produce those peculiar productions found nowhere else but allied to those of the nearest land. The change like every other change in nature was no doubt gradual, & the supposition that other species were successively produced closely allied to those previously existing, & that while this was going on, the original or some of the first formed species died out, exactly accords with the facts as we find them & the process of peopling new islands at the present day.7'

How far removed Lyell was from evolutionary theory is brought out at still another point. He thought that when climatic changes did cause the extinction of local species, any new inhabitants would be "perfectly dissimilar in their forms, habits, and organization." 72 In other words, there would be a wholesale extinction followed by a renewal with altogether different species. Wallace pointed out, however, that the new species would more likely be modified forms of those previously existing: "It would be an extraordinary thing if while the modifications of the surface took [place] by natural causes now in operation, & the extinction of species was the natural result of the same causes, yet the reproduction & introduction of new species required special acts of creation, or some process which does not present itself in the ordinary course of nature." (At this point in his "Notebook," Wallace inserted the followng note: "Introduce this and disprove all Lyell's arguments first at the commencement of my last chapter" -evidence that this was part of his proposed book.)73 When environmental conditions changed, Lyell thought that new species already adapted to these new conditions would come in from surrounding areas before those in residence had

71. Wallace, "Notebook," pp. 46-49; some commas added. 72. Lyell, Principles of Geology, 4th ed., III, 154. 73. Wallace, "Notebook," pp. 50-51.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection time to change (if they changed at all). Wallace agreed that "if the change took place rapidly, the exact results Lyell pre- dicts might follow"; "but," he went on, 'how the same results could follow from an excessively gradual change it is impossible to understand." 74 Lyell and Lamarck both agreed that plenty of time was available, but they differed on what would happen. Lyell believed that species could change, but only to a limited degree and within the space of a few generations. Not only were they incapable of greater change, but such change would also be precluded by the immigration of other species already adapted to the new environment. Lamarck, on the other hand, believed that species did change gradually through time in response to changes in the environment, but his agency, the wills of the animals themselves, was suspect. Whatever their deficiencies (perhaps because of them), the beliefs of both these men were valuable steppingstones along Wallace's way.

In the meantime, Wallace was reading as widely as circumstances permitted: Edward Blyth on the classification of varieties, Richard Owen on the varieties of color in man, and Leopold von Buch on the flora of the Canary Islands.75 Be- cause von Buch's perceptive comments also influenced Darwin, Wallace's notes on them are worth quoting:

On continents the individuals of one kind of plant disperse themselves very far, and by the difference of stations of nourishment & of soil produce varieties, which at such a distance not being crossed by other varieties & thus brought back to the primitive type, become at length permanent & distinct species. Then if by chance in other directions they meet with another variety equally changed in its march, the two have become very distinct species & are no longer sus- ceptible of intermixture . . . He then shews that plants on the exposed peak of Teneriffe where they can meet & cross do not form varieties or species, while others such as Pyre- thrum & Cineraria living in sheltered valleys & low grounds often have closely allied species confined to one valley or one island.7f

74. Ibid., p. 52; commas added. See also Lyell, Principles of Geology, 4th ed., III, 161-163. 75. Christian Leopold von Buch, Physicalische Beschreibung der Can- arischen Inseln (Berlin: K. Akademie der Wissenschaften, 1825). An earlier report on the flora of the Canary Islands appeared in the Abhand- lungen of the Society (1816-1817), 337-384. 76. Wallace, "Notebook," p. 90.

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Darwin in his own "Notebook" says, "Von Buch distinctly states that permanent varieties become species, pp. 147-150,-not being crossed with others." 77 Still another clue attracted Wallace's attention, and he asked:

What is the import of the doctrine of Morphology of plants? . . .For if stamens & petals & carpels have been in every case independently created as such, it is absurd to say they are modifications or developments of any thing else, & the absurdity is still greater if that of which they are said to be the development came into existence after them. In that case all the beautiful facts of morphology are a delusion & a snare, as much so as fossils would be were they really not the remains of living things but chance imitations of them. The natural inference of an unprejudiced person however would be that both are true records of the progress of the organic world. Nature seems to tell us that as organs are occasionally changed & modified now, in individual plants, we may learn how the actual changes have taken place in the species of plants. A key is offered us to a mystery we could otherwise never have laid open, why should we refuse to use it?78

Sometime during these musings, Wallace wrote out his "Note on the Theory of Permanent and Geographical Varieties," following up an interest expressed many years earlier to Bates, and it was published early in 1858. Although he had been pondering a wide range of subjects, he limited the "Note" to showing the logical inconsistency in the suggestion that geographical varieties had permanent characters. If varieties differ from species only in the minuteness of the permanent characters, then the difference between them is merely a quan- titative one and the dividing line becomes very difficult to dis- tinguish. The only qualitative difference that Wallace could discover was that of the pernanence vs. the impermanence of variations. But this was no better, for some groups so formed were called special creations and others not. "Strange that such widely different origins produce such identical results." 79 If varieties are known to be produced by ordinary generation, why should species only slightly different be produced by special creation? "If there is no other character [than one of mere

77. Gavin de Beer, ed., "Darwin's Notebooks on Transmutation of Species," Bull. Brit. Mus. (Nat. Hist.), Hist. Ser., 2 (1960), 61. Darwin's first notebook was written between July 1837 and February 1838. 78. Wallace, "Notebook," pp. 97-100. 79. Wallace, "Note," p. 5888. See also Appendix, 3.

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IV. AND DARWIN [Darwin] is now preparing for publication his great work on species and varieties, for which he has been collecting information twenty years. He may save me the trouble of writing the second part of my hypothesis by proving that there is no difference in nature between the origin of species and varieties, or he may give me trouble by arriving at another conclusion, but at all events his facts will be given me to work upon. Wallace to Bates, 4 January 1858, from Marchant, Alfred Russel Wallace Eighteen hundred and fifty-eight was the year in which the careers of Wallace and Darwin collided. Wallace had initiated the correspondence between them when he wrote his first letter to Darwin in October 1856 [8].81 Why he wrote is not known; perhaps, as suggested earlier, he hoped that Darwin would be interested in his speculations on the Galapagos Islands puzzle. Although it was Wallace's third letter to Darwin that brought with it Wallace's discovery of the theory of natural selection, even this first one may have been disquieting. The contents of Wallace's letter can be partially surmised from Darwin's answer written in the following May [11]: the paper in the Annals, domestic versus wild varieties (a crucial point in the development of Wallace's ideas), hybrid sterility, and the effects of climatic changes. As to the paper, Darwin agreed "to the truth of almost every word"; he had, in fact, already pondered the same problems-classification, extinction and creation, and rudimentary organs-as can be seen from his own notebooks and his essay written in 1844. There was less agreement, however, on other subjects. Darwin later wrote Lyell (June 1858) that he and Wallace differed only in "that I was led to my views from what artificial selection has done for domestic animals [231." But they both made use of domesticated animals, although for different ends: Darwin to show that variation existed and could be channeled, Wallace to show that the usual definition of species, based on domesticated animals, did not apply to wild animals. (And, as Wallace wrote many years later, "it has always been considered a weakness in Darwin's work that he based his theory, pri- marily, on the evidence of variation in domesticated animals and cultivated plants.")82 Nor did Wallace and Darwin ever agree on the thorny problem of the sterility of hybrids.

80. Ibid., p. 5888. 81. See also Appendix, 69 and notes 43 and 162. 82. Alfred Russel Wallace, Dar-winism: An Exposition of the Theory of Natural Selection, with Some of its Applications (New York: Humboldt, 1889), p. iv.

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There was also a difference of opinion on the effect of climatic changes. In 1844 Darwin had written "that probably such changes of external conditions would, from acting on the re- productive system, cause the organization of the beings most affected to become, as under domestication, plastic." 83 He had since modified his opinion, and in his letter to Wallace he agreed "on the little effect of 'climatal conditions."' Wallace, however, never did ascribe any such direct influence to climatic conditions in causing variations. Nevertheless, in spite of some disagreement it was plain, as Darwin said, that they had thought much alike. Darwin received Wallace's first letter late in April 1857. He was then deep in writing "his great work on species and varieties," having started almost exactly a year earlier at the urging of Lyell, with the strong support of the botanist Joseph Hooker [5, 6, 7]. As Lyell wrote not long afterward, "Part of the MS. of [Darwin's] projected work was read to Dr. Hooker as early as 1844, and some of the principal results were com- municated to me on several occasions. Dr. Hooker and I had repeatedly urged him to publish without delay." 84 It has recently been proposed, on evidence from Lyell's own notebooks, that it was actually Wallace's paper in the Annals that prompted Lyell to reconsider the subject of species and to prod Darwin to publication, and, furthermore, that it was at this time that Darwin explained his theory to Lyell.88 Darwin had protested to Lyell that he did not like writing for priority, but had admitted that he "certainly should be vexed if any one were to publish my doctrines before me [5]"; and to his cousin, W. D. Fox, he had confided that he wished he "could set less value on the bauble fame [10]." By 31 March 1857, he had completed six chapters of the projected work, and he may already have been at work on the seventh, com- paring species and varieties, when Wallace's first letter arrived in late April. The coincidence in their thinking may have put Darwin on his guard. On 1 May 1857, Darwin wrote in answer (to some speculations by Wallace?) that he had been working for nearly twenty years "on the question how and in what way do species and varieties differ from each other [11]." He did not elaborate on

83. Darwin and Wallace, Evolution, p. 119. Text differs slightly from that in joint papers; see note 133. 84. Charles Lyell, The Geological Evidences of the Antiquity of Man, with Remarks on Theories of the Origin of Species by Variation, 1st ed. (London: Murray, 1863), p. 408. 85. McKinney, "Wallace and Natural Selection," pp. 350-352.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection the point, however, and left it up in the air. (Wallace had not yet written his own "Note" on the subject, and it is not known whether Darwin ever did see it.) As was his usual practice, Darwin included several requests for information, asking Wallace, among other things, to let him know "if you should, after receiving this, stumble on any curious domestic breed" of poultry.86 Perhaps as a result of this inquiry, Wallace made a few notes in his "Notebook," some time before November 1857, on unusual breeds of ducks.87 This small point will be brought up later in connection with Wallace's recollections of the beginning of his correspondence with Darwin. More important at the moment is Darwin's protestation that "it is really impossible to explain my views (in the compass of a letter) on the causes and means of variation in a state of nature." Was it really impossible? Two years earlier, in April 1855, Darwin had begun to cor- respond with the noted American botanist Asa Gray, whom he had once met briefly at Kew, and Gray had been providing him with many valuable comments. On 20 July 1857, not long after receiving his first letter from Wallace, Darwin wrote again to Gray, saying, "I should like to tell you (and I do not think I have) how I view my work," condensing into a few sentences the gist of his theory [12].88 A recent biographer of Gray suggests that Darwin felt it necessary to let Gray in on his secret to ensure the continuance of this useful correspondence, but this does not seem to be a necessary assumption.89 More to the point, Darwin seemed rather to fear that Gray would despise him and his crotchets, and in his next letter (5 September) he confessed that he had been afraid that Gray might think him "worth no more notice or assistance" because of his unorthodox views [14].90 (Even in March 1860 Darwin considered Gray to be a convert to his views only "to some extent."))91 Along with this answer to Gray, Darwin sent a copy of an outline that he had made of his theory, "as you seem interested in the subject." This is the famous extract published the fol-

86. The paragraph containing this request is published in Marchant, Wallace, p. 108, but it was omitted from the Darwin, Life and Letters, I, 454. See also note 87. 87. Wallace, "Notebook," p. 91. See also Appendix, 69, Marchant, Wallace, p. 86, and notes 161 and 162. 88. See also note 146. 89. A. Hunter Dupree, Asa Gray, 1810-1888 (Cambridge: Harvard Uni- versity Press, 1959), p. 244. 90. See also note 146. 91. Darwin, Life and Letters, II, 87.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL lowing year as part of the Darwin-Wallace joint papers. Dar- win concluded his letter to Gray with the following request:

You will, perhaps, think it paltry in me, when I ask you not to mention my doctrine; the reason is, if any one, like the author of the 'Vestiges,' were to hear of them, he might easily work them in, and then I should have to quote from a work perhaps despised by naturalists, and this would greatly injure any chance of my views being received by those alone whose opinion I value [14]192

Why did Darwin send this statement to Gray, who responded that it was "grievously hypothetical [15, 17]" rather than to Wallace (who was hardly at this time, however, a scientific peer), who would have understood? Was it Wallace rather than Chambers of whom he was afraid? If so, a recent outline of his views, including the important addition on divergence, mailed to the eminent American botanist might protect his ideas. And, whatever his intentions may have been, this indeed was the result. Finally, on 29 November 1857, Darwin thanked Gray for his help, remarking that "every criticism from a good man is of value to me [17]." 93 But the subject was apparently not pursued, and this concludes the series of letters about Darwin's theory. Wallace was "much gratified" by Darwin's first letter, as he wrote Bates in January 1858, but he was no wiser than before (had he asked?) about Darwin's opinion for or against a "difference in nature between the origin of species and varieties [19]." 94 In any case, he had pretty wel made up his own mind already. Judging from Darwin's next answer, Wallace in reply once again brought up his paper in the Annals and also remarked on various problems relating to the geographical distribution of animals. But the only piece on Wallace's side of this early correspondence that is still in existence is a snippet from his answering letter of 27 September 1857, on the breeding habits of jaguars and his plan for his book [16].9r For the rest, Wal-

92. This paragraph has been variously interpreted; see Dupree, Asa Gray, p. 246 and Himmelfarb, Darwin, p. 207. 93. See note 146. 94. See also Loren Eiseley, Darwin's Century and the Men Who Discov- ered It (Garden City, N.Y.: Doubleday, 1958), p. 291, where he considers this correspondence "stimulating" to Wallace. But Wallace received no such positive statement as was given to Gray. 95. Unpublished fragment in the Cambridge University Library; see also note 60. For Darwin's methods of filing his letters, see notes 140-143.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection lace's opinions must be conjectured from Darwin's letters to him. By the time Darwin answered Wallace's second letter on 22 December 1857, he had almost completed the ninth chapter of his book, this one on hybridism. He assured Wallace that his paper had not gone unnoticed, Lyell and the zoologist Edward Blyth having called it to his attention; none of these men, however, had bothered to write Wallace about it. Darwin passed up his second and last opportunity to tell Wallace about his theory, saying that "though agreeing with you on your conclusions in that paper, I believe I go much further than you; but it is too long a subject to enter on my speculative no- tions [18]." It seems unlikely, however, that Wallace received this letter before the end of February 1858; most of the letters exchanged by the two men took anywhere from three to six months to reach the recipient. In the meantime, in his January 1858 letter to Bates, Wal- lace told him of his plans, explaining that "I have prepared the plan and written portions of an extensive work embracing the subject in all its bearings and endeavouring to prove what in the paper [in the Annals] I only indicated." He did not seem to be overly concemed about Darwin's conclusion on the origin of species and varieties, remarking that Darwin might save him the trouble of proving that there was no difference in their nature [19]. It has been claimed that Wallace, "duly warned off" by Dar- win's first letter, had nevertheless continued his own work on the subject.96 It should be objected that this was no private preserve of Darwin's. Many people were interested in it, as Wallace was well aware. Darwin gave Wallace no hint of a solution to the problem; why should he not continue with what had been a consuming interest for many years? Bates himself, on his return home, wrote that one of their purposes in going to the Amazon was to "gather facts, as Mr. Wallace expressed it in one of his letters, 'towards solving the problem of the origin of species,' a subject on which we had conversed and corres- ponded much together." 97 If Darwin had been working on the problem for twenty years, Wallace had been working on it for at least ten, the major difference being that Darwin had long had a theory against which he was collecting facts, while Wallace was still actively searching for one. By now Wallace had concluded that there was no qualitative difference in the origin of species and varieties and that they

96. Himmelfarb, Darwin, p. 236. 97. Bates, The Naturalist, p. iii; see also Appendix, 4 and note 17.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL were formed gradually by natural means with a relationship in space and time to what had gone before, but without any necessary progression. Only a short distance separated him from his goal, a "'theory of the origin of species." In February 1858 the final step was taken. Wallace was ly- ing ill in Ternate in the Moluccas, mulling as usual over his problem, when at last he found the key to the puzzle: the theory of natural selection. Much has been made of the fact that the solution came to him while he was ill but, as he im- patiently remarked later, he "had no idea whatever of 'dying,' -as it was not a serious illness [69]." Indeed, he had frequently been sick, and sometimes much sicker. Although not iden- tified at the time, the disease has recently been referred to as malaria.98 As soon as he was able, Wallace wrote out his theory with the title, "On the Tendency of Varieties to depart indefinitely from the Original Type," and sent it to Darwin with the request that it be shown to Lyell.99 Had he chosen any other recipient, the results would have been different. Gray's response to Darwin's disclosure, for instance, had been that it was too hypothetical. But to Darwin, Wallace's paper was little short of a calamity. In his "Note," Wallace had been struggling with the supposed distinction between permanent varieties and permanently invariable species. Now he saw his way clear. Belief in variation within strict limits and reversion to the original type was based on domestic animals and then applied to wild animals. But domestic animals are artificial and unable to maintain themselves without the help of man. If allowed to go wild, they must either return to something similar to the original type or become extinct. Wild animals, on the other hand, must be adapted to their environment. Their every faculty is constantly exercised in keeping themselves alive. Any improve- ment in organization is quickly taken advantage of, and a new variety is thus superior to its predecessor. Being superior, it "could not return to the original form; for that form is an inferior one, and could never compete with it for existence." 100 Quite the opposite is true of domestic animals, which are inferior from the point of view of maintaining themselves in the wild. Thus varieties in nature tend to depart indefinitely from the original tvDe.

98. Julian S. Huxley and H. B. D. Kettlewell, Charles Darwin and his World (New York: Viking Press, 1965), p. 74. 99. Alfred Russel Wallace, "On the Tendency of Varieties to depart indefinitely from the Original Type," J. Linn. Soc. London (Zool.), 3 (1858), 53-62. 100. Ibid., p. 58. See also note 69.

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Several years earlier, Wallace had found puzzling Lyell's belief that the balance of species was preserved by plants, insects, mammals, and birds adapted to the purpose, and he had worried over this problem in his "Notebook":

This phrase is utterly without meaning. Some species are very rare, others very abundant-where is the balance? Some species exclude all others in particular tracts-where is the balance-When the locust devastates vast regions, & causes the death of animals & man, what is the meaning of saying the balance is preserved . . . To human apprehension there is no balance but a struggle in which one often exterminates the other-When animals and plants become extinct, where is the balance. If any state can be imagined proving a want of balance, then a balance may perhaps be admitted, but what state is that?10'

Now, two years later, Wallace had his answer. In the "struggle for existence," many individuals must perish annually. Even the least fecund species would soon overrun the earth if its numbers went unchecked. Those that survive are the ones best adapted to obtain food and to withstand their enemies and the seasonal changes in the weather; those that die are the young, the old, and the sick. In applying this not only to individuals but also to species, Wallace thought he had an answer to why some species are rare while others are abundant. Besides this, the animal population of a country cannot increase materially if conditions remain the same. In the "struggle for existence," therefore, those individuals and species best adapted to maintain themselves survive. It is well known that Wallace and Darwin both read Mal- thus' famous Essay on the Principle of Population, Wallace before leaving Leicester and Darwin after returning home from his voyage. Malthus' interest was in the moral perfectibility of man, and it was in this light that he discussed the checks to population growth-famine, war, disease, and vice-but Wal- lace and Darwin were both impressed by the implications of these checks. The exact extent of Malthus' influence is hard to determine and has been the subject of much debate, but at the least his forceful presentation was widely known. Darwin read Malthus in the fall of 1838. There is no indication of this in the first edition of his Voyage of the Beagle,

101. Wallace, "Notebook," pp. 49-50; some commas added. See also Lyell, Principles of Geology, 4th ed., III, 98-120, where he discusses the "checks and counter-checks which nature has appointed to preserve the balance of power amongst species."

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL however, because though published in 1839, the writing of it was largely finished by June 1837. Darwin had tried vainly to account for the extinction of so many peculiar South American forms that were related to the present inhabitants, concluding only that:

On such grounds it does not seem a necessary conclusion, that the extinction of species, more than their creation, should exclusively depend on the nature (altered by physical changes) of their country. 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.102

This comment was greatly enlarged for the second edition (1845) and clearly shows the influence of Malthus:

Nevertheless, if we consider the subject under another point of view, it will appear less perplexing. We do not steadily bear in mind, how profoundly ignorant we are of the conditions of existence of every animal; nor do we always remember, that some check is constantly preventing the too rapid increase of every organized being left in a state of nature. The supply of food, on an average, remains constant; yet the tendency in every animal to increase by propagation is geometrical; and its surprising effects have nowhere been more astonishingly shown, than in the case of the European animals run wild during the last few cen- turies in America. Every animal in a state of nature regu- larly breeds; yet in a species long established, any great increase in numbers is obviously impossible, and must be checked by some means. We are, nevertheless, seldom able with certainty to tell in any given species, at what period of life, or at what period of the year, or whether only at long intervals, the check falls; or, again, what is the precise nature of the check. Hence probably it is, that we feel so little surprise at one, of two species closely allied in habits, being rare and the other abundant in the same district; or, again, that one should be abundant in one district, and another, filling the same place in the economy of nature, should be abundant in a neighboring district, differing very little in its conditions. If asked how this is, one immediately replies that it is determined by some slight difference in climate, food, or the number of enemies: yet how rarely, if ever, we can point out the precise cause and manner of action of the check I We are, therefore, driven to the conclusion, that

102. Darwin, Voyage of the Beagle (1839), p. 212.

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causes generally quite inappreciable by us, determine whether a given species shall be abundant or scanty in numbers. . . .If then, as appears probable, species first become rare and then extinct-if the too rapid increase of every species, even the most favoured, is steadily checked, as we must admit, though how and when it is hard to say-and if we see, without the smallest surprise, though unable to assign the precise reason, one species abundant and another closely-allied species rare in the same district-why should we feel such great astonishment at the rarity being carried a step further to extinction?103 Wallace did not have Malthus with him, but he did have Darwin. Familiar as he already was with the Malthusian arguments, he must have noticed their inclusion in the second edition of the Voyage of the Beagle. These principles, first suggested in 1798, were now used by Wallace and Darwin in a rather different context. But Malthus alone was not enough. As Wallace justly pointed out many years later, "along with Mal- thus I had read, and been even more deeply impressed by, Sir Charles Lyell's immortal 'Principles of Geology.'" 104 The final and most important point of Wallace's paper was the application of the concepts he had developed to varieties. Even slight variations would have an effect, either favorable or unfavorable, and under changed physical conditions a better- adapted variety might survive its parent species. (This would be true only of wild varieties, however, for domesticated animals turned wild are rarely able to maintain themselves.) This process repeated would lead to "progression and continued divergence." 105 At last Wallace had a mechanism that explained the knotty problem of progression which had so baffled Lyell, and the equally puzzling problem of divergence; and it could also replace Lamarck's generally discredited theory that progressive changes were due to the wills of the animals themselves.

Darwin's approach was a little different. Because Wallace is the focus of this study, discussion here will be limited to the two selections from Darwin's writings that became part of the joint papers in 1858. The first was an extract from an essay written in 1844.106 Darwin had remarked there that De Can- 103. Ibid., (1845), pp. 174-176. 104. Linnean Society of London, The Darwin-Wallace Celebration, p. 118. 105. Wallace, "On the Tendency," p. 59. See also note 37. 106. Charles Darwin, "Extract from an Unpublished Work on Species, by C. Darwin, Esq., consisting of a portion of a Chapter entitled 'On the

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL dolle's war of nature "is the doctrine of Malthus applied in most cases with ten-fold force" 107 (he was to use a similar phrase in the Origin of Species).108 Under these circumstances, slight variations caused directly by changed physical conditions might lead to small improvements in organisms through the natural selection of those better adapted.109 Darwin also discussed sexual selection, another subject on which he and Wallace were to differ strongly. The brief abstract enclosed with the letter to Asa Gray sum- marized Darwin's conclusions at this later date, 1857.110 Se- lection of variations by man is recognized in the propagation of domestic animals. Physical conditions are known to have changed over a great length of time. These changed conditions have caused variations to occur in organisms in a state of nature, although Darwin was no longer certain that this was the sole cause. Finally, there is a natural power comparable to that of man which selects those that survive in the struggle for life, a power which Darwin called Natural Selection. In a country undergoing changes, these slight variations would be selected and gradually accumulated, leading to new varieties adapted to the new conditions. At the end, Darwin added a paragraph on his principle of divergence. The solution to this problem-"that the varying offspring of each species will try (only few will succeed) to seize on as many and as diverse places in the economy of nature as possible" "'-had not occured to him until 1852,112 'long after I had come to Down," 113 and so was not yet a part of a formal statement of his views. He knew from Wal- lace's paper in the Annals that he too was trying to solve this problem.

Variation of Organic Beings in a state of Nature; on the Natural Means of Selection; on the Comparison of Domestic Races and true Species,'" J. Linn. Soc. London (Zool.), 3 (1858), 46-50. See also note 133. 107. Ibid., p. 47. 108. Charles Darwin, On the Origin of Species by Means of Natural Selection, or the Preservation of favoured Races in the Struggle for Life (London: Murray, 1859; facsimile reprint, Cambridge: Harvard University Press, 1964), p. 63. 109. See also note 83. 110. Charles Darwin, "Abstract of a Letter from C. Darwin, Esq., to Prof. Asa Gray, Boston, U.S., dated Down, September 5th, 1857," J. Linn. Soc. London (Zool.), 3 (1858), 50-53. See also Appendix, 14 and notes 88-93. 111. Ibid., pp. 52-53. See also note 37. 112. Gavin de Beer, Charles Darwin: a Scientific Biography (Garden City, N.Y.: Doubleday, 1964), p. 140. 113. Darwin, Life and Letters, I, 69.

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V. THE JOINT PAPERS

I was not aware before that your father had been so distressed-or rather disturbed-by my sending him my essay from Ternate.

Wallace to Francis Darwin, 20 November 1887

"June 14th Pigeons (interrupted)." 114 Such is the cryptic note in Darwin's "Journal" indicating the receipt of Wallace's paper on 18 June 1858. By this time he had completed twelve chapters of his book and was at work on the thirteenth, and his distress at being thus forestalled can easily be imagined. The story of this most dramatic moment has often been re- counted. But, as Hooker observed, the details stem entirely from some of the letters written at the time by Darwin to himself and to Lyell [22-26, 29, 33-36];115 all other documentary evidence, the letters from Wallace, Lyell, and Hooker to Darwin, as well as the manuscript of Wallace's paper, has disappeared. The facts, consequently, are difficult to determine, and the circumstances have been variously interpreted. There is no way of ascertaining exactly why Wallace sent his paper to Darwin; certainly he could not have anticipated the result. With no hint from Darwin, he could not have re- alized that he had stumbled onto the very foundation of Dar- win's work. Darwin, on the other hand, must have had a fair notion of Wallace's progress from his published papers and perhaps a warning of this disaster from his letters, the first of which may indeed have precipitated the sketch sent to Gray. Wallace, who had been away from England for many years, was a self-educated collector from outside the regular establishment, and he had few personal contacts among the scientific elite. By dint of his own efforts, he had finally established a correspondence with one of its members, Charles Darwin. He may well have hoped for some useful criticism from Darwin and Lyell, but he could hardly have expected to be catapulted into the front ranks himself. From the evidence, it appears that Wallace sent his paper to Darwin with the request that it be forwarded to Lyell, "should he think it sufficiently novel and interesting [20]." 11I Darwin's own letter to Lyell, written on 18 June, said only

114. Gavin de Beer, ed., "Darwin's Journal," Bull. Brit. Mus. (Nat. Hist.), Hist. Ser., 2 (1959), 14. 14 June 1858 was the day on which Darwin began this chapter. 115. See also note 139. 116. Charles Lyell and Joseph D. Hooker, "[Letter communicating the Darwin-Wallace Papers to the Linnean Society]," 30 June 1858, J. Linn. Soc. London (Zool.), 3 (1858), 46.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL that Wallace "has to-day sent me the enclosed, and asked me to forward it to you [22]." Lyell's recollection later was that the paper had been brought to him by Hooker, who then suggested some sort of joint publication.117 In Leonard Huxley's biography of Hooker, it is stated that Darwin "had first confided Wallace's unexpected letter" to him, and that he first suggested joint publication and also the getting of Lyell's opinion.118 Whatever the precise details, Hooker, Darwin's "most intimate friend," accepted the larger share of the re- sponsibility for what happened. Darwin was now in a dilemma: "He does not say he wishes me to publish, but I shall, of course, at once write and offer to send to any journal. So all my originality, whatever it may amount to, will be smashed [22]." In fact, Darwin did start a letter to Wallace giving up his claims to priority, but he never finished it, for his old friends, Hooker and Lyell, suggested a compromise. If Darwin's and Wallace's roles had been reversed, as they could have been, Wallace would have had no one to help him resolve the difficulty. A week later, on 25 and 26 June, Darwin wrote Lyell again. A joint publication of some kind had apparently been proposed, although "Wallace says nothing about publication" and Dar- win was properly hesitant about the proprieties involved. He mentioned a copy of his sketch sent to Gray "about a year ago . . . (owing to correspondence on several points)." ("Cor- respondence" here probably means actual letter-writing rather than agreement, because the sketch "gives most imperfectly only the means of change," a subject new to Gray.) He also enclosed the letter from Wallace and requested that Lyell get Hooker's opinion; some of the confusion on this point has already been mentioned [20, 23, 24, 26]. By 29 June, Darwin had their answers. He was to send to Hooker both Wallace's paper (which Lyell must already have returned to him) and his own sketch sent to Asa Gray. Although they had apparently not asked for it, he also sent along his much more extensive sketch (230 pages), written in 1844, to show by notes in Hook- er's handwriting that he had read it [25, 26].119 Darwin's letters were usually written with an intensity of feeling lacking in the more pedestrian efforts of many of his contemporaries. At this moment they were shrill with anxiety and doubt. But Darwin was being sorely tried. He was troubled not only by Wallace's communication but also by severe illness

117. Lyell, Principles of Geology, 10th ed., II, 278. 118. Leonard Huxley, ed., Life and Letters of Sir Joseph Dalton Hooker (London: Murray, 1918), II, 465. 119. See also note 84.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection in his family. An infant son died of scarlet fever on 28 June, and a daughter was seriously ill with diphtheria. Finding a suitable forum for the papers at this time of year would ordinarily have presented still another problem. But both Hooker and Lyell, as Fellows of the Linnean Society of London, knew that one had unexpectedly become available. Robert Brown, a leading botanist, former President and then Council Member of the Society, had died on 10 June. Out of respect, the last meeting of the old session, held on 17 June, was ad- journed before the reading of the scheduled papers. But Brown had to be replaced on the Council within three months, and, as the new session would not start until November, it was decided to hold the extra meeting on 1 July. Without consulting anyone else, Hooker and Lyell transmitted their selections to the Secretary of the Society on 30 June, to be read by him the next day. As Hooker said in 1908: It cannot fail to be noticed that all these inter-communica- tions between Mr. Darwin, Sir Charles Lyell, and myself were conducted by correspondence, no two of us having met in the interval between June the 18th and July the 1st, when I met Lyell at the evening meeting of the Linnean Society; and no fourth individual had any cognisance of our proceedings.120

This was not an occasion of "mutual nobility," 121 nor was it "a monument to the natural generosity of both the great biologists," 122 as is so often claimed. It was clearly not mutual because Wallace's paper was read without his knowledge or consent, and he knew nothing about it until October. Nor does it seem to have been particularly noble. However just Darwin's claims to priority, he was a gainer, not a loser, from the decision. Wallace had no opportunity to be either noble or generous. Wallace, "a gentleman attached to the study of Natural His- tory," was not unknown to the Linnean Society. The first two volumes of the Society's Journal (Zoology) were largely taken up with descriptions (written by others) of the collections of insects he had sent home from Singapore, Malacca, and Sara- wak. The Society was later to publish some of Wallace's most

120. Linnean Society of London, The Darwin-Wallace Celebration, p. 15. This and the information of Brown (Ibid., pp. 14-15) were omitted in Marchant, Wallace, p. 98. 121. Eiseley, Darwin's Century, p. 292. 122. Julian S. Huxley, "Alfred Russel Wallace," Dictionary of National Biography, Supplement 1912-1921 (London: Oxford University Press, 1927), p. 547.

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important papers, but he himself was not elected a Fellow until 1872. Darwin, on the other hand, was already a Fellow, and had just been elected to the Council in May. Some thirty people, perhaps more, out of a membership of over four hundred were present at the meeting. Although some of their names might not be recognized today, fully half of those listed in the Minutes as attending have rated notices in Britain's renowned Dictionary of National Biography, certainly a distinguished audience.123 The evening was a full one. The business of the meeting was transacted, and then came the reading of the papers, the joint papers by Darwin and Wallace followed by five of the six previously scheduled for 17 June. The joint papers were introduced by a letter from Lyell and Hooker explaining what they had done and why. The first selection was from Darwin's essay of 1844, an "Extract from an unpublished Work on Species, by C. Darwin, Esq., consisting of a portion of a Chap- ter entitled 'On the Variation of Organic Beings in a state of Nature; on the Natural Means of Selection; on the Comparison of Domestic Races and true Species."' Darwin appended to the published version a note that "this MS. work was never intended for publication, and therefore was not written with care." This was no hastily written summary, however, for Darwin had had it copied and bound, and he had also left instructions to his wife for its publication in the event of his premature death.124 Secondly came the "Abstract of a Letter from C. Darwin,

123. Listed in the minutes of the meeting, in the Society's Darwin- Wallace Celebration, pp. 81-86. The following can be found in the D.N.B.: Baird, William (1803-1872), zoologist; Ball, John (1818-1889), botanist; Baly, William (1814-1861), physician (visitor); Bell, Thomas (1792- 1880), dental surgeon and zoologist (President); Bennett, John Joseph (1801-1876), botanist (Sole Secretary), not listed as present, although he presumably read the papers; Bentham, George (1800-1884), botanist; Burchell, William John (1782?-1863), explorer and naturalist; Busk, George (1807-1886), physician and scientist (Under- (Zoological) Secre- tary), not listed as present, although Hooker later recalled that he was; Carpenter, William Benjamin (1813-1885), naturalist and physician; Currey, Frederick (1819-1881), mycologist; Fitton, William (1780-1861), physician and geologist; Henfrey, Arthur (1819-1859), botanist; Hooker, Joseph Dalton (1817-1911), botanist; Lyell, Charles (1797-1875), geologist; Salter, John William (1820-1869), geologist; Seeman, Berthold Carl (1825-1871), botanist and traveler; Ward, Nathaniel Bagshaw (1791-1868), botanist and physician. Others may have been present as the list ends with "etc., etc." 124. The date is incorrectly given as 1842 in the published papers. Darwin further confused the issue by using the date 1839 in a letter to Wallace; see Appendix, 43. For details of Darwin's plans, see Himmelfarb, Darwin, pp. 190-191.

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Esq., to Prof. Asa Gray, Boston, U.S., dated Down, September 5th, 1857," the outline of his theory of natural selection that Dar- win had sent to Gray. Francis Darwin was later of the opinion that the reason for the inclusion of this note was the discussion of the "principle of divergence," an important part of Darwin's theory not included in the 1844 essay.125 Hooker was aware that Darwin gave divergence equal prominence with natural selection as "the keystone of my book [21]," although he apparently did not understand the connection between them. In his own essay, "On the Flora of Australia," published at almost the same time as Darwin's Origin of Species, Hooker wrote that "the tendency of varieties, both in nature and under cultivation, when further varying, is rather to depart more and more widely from the original type than to revert to it." 126 Darwin objected that this was "without selection doubtful." 127 Third and last was Wallace's paper, "On the Tendency of Varieties to depart indefinitely from the Original Type." But no note was added to indicate that Wallace had not written for publication either. It was Hooker's recollection twenty-eight years later (whether accurately or no; certainly public reaction to the publication of the papers was almost nil) that the interest was intense, although there was no discussion. Thomas Bell, the President, though a personal friend of Darwin's, "was hostile to the end of his life." Neither of the Secretaries, George Busk and John J. Bennett, said anything, nor did the botanist George Bent- ham [67]. (Thomas Huxley, later to be "Darwin's bulldog," was not present, not being elected a Fellow until December 1858.) Bentham may have been silent, but his feelings were those of "severe pain and disappointment." His was the only one of the six previously scheduled papers that was not read. Many years later he recalled the events in a letter to Francis Darwin:

On the day that his [C. Darwin's] celebrated paper was read at the Linnean Society, July 1st, 1858, a long paper of mine had been set down for reading, in which, in commenting on the British Flora, I had collected a number of observa- tions and facts illustrating what I then believed to be a fixity in species, however difficult it might be to assign their limits, and showing a tendency of abnormal forms produced by cultivation or otherwise, to withdraw within those orig-

125. Darwin and Wallace, Evolution, p. 34. See also note 37. 126. Quoted in Francis Darwin, ed., More Letters of Charles Darwin: a Record of his Work in a Series of Hitherto Unpublished Letters (New York: Appleton, 1903), I, 134. 127. Ibid.

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inal limits when left to themselves. Most fortunately my paper had to give way to Mr. Darwin's and when once that was read, I felt bound to defer mine for reconsideration; I began to entertain doubts on the subject, and on the appearance of the 'Origin of Species,' I was forced, however re- luctantly, to give up my long-cherished convictions, the results of much labour and study, and I cancelled all that part of my paper which urged original fixity, and published only portions of the remainder in another form, chiefly in the 'Natural History Review.' I have since acknowledged on various occasions my full adoption of Mr. Darwin's views, and chiefly in my Presidential Address of 1863 [to the Lin- nean Society], and in my thirteenth and last address, issued in the form of a report to the British Association at its meeting at Belfast in 1874 [66].

Bentham had not given in easily; even in his Presidential Address in 1862 he was still struggling against the new doctrine:

I do not refer to those speculations on the origin of species, which have excited so much controversy; for the discussion of that question, when considered only with reference to the comparative plausibility of opposite hypothesis, is beyond the province of our Society. Attempts to bring it forward at our meetings were very judiciously checked by my predecessor [Bell] in this Chair, and I certainly should be sorry to see our time taken up by theoretical arguments not accompanied by the disclosure of new facts or observa- tions.128

Bell was a dental surgeon and zoologist, but "as a naturalist he was more at home in his study than in the field, and he made few original contributions of special value to zoology. As a writer, his chief merit is that of agreeable compilation." 129 In his own Presidential Address in 1859, he dismissed the joint papers altogether:

The year which has passed . . . has not been unproductive in contributions of interest and value, in those sciences to which we are professedly more particularly addicted, as well as in every other walk of scientific research. It has not, indeed, been marked by any of those striking discoveries which at once revolutionize, so to speak, the department of

128. Proc. Linn. Soc. London (1 November 1860-19 June 1862), p. Lxcxi. 129. G. T. Bettany, "Thomas Bell," D.N.B., II, 175.

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science on which they bear; it is only at remote intervals that we can reasonably expect any sudden and brilliant in- novation which shall produce a marked and permanent im- press on the character of any branch of knowledge, or confer a lasting and important service on mankind. A Bacon or a Newton, an Oersted or a Wheatstone, a Davy or a Daguerre, is an occasional phenomenon, whose existence and career seem to be especially appointed by Providence, for the purpose of effecting some great important change in the condition or pursuits of man.130

Darwin's immediate reaction to Wallace's paper was, hu- manly enough, great distress, followed by relief at the solution worked out by Hooker and Lyell. "But," he wrote to Hooker, "in truth it shames me that you should have lost time on a mere point of priority." (If it was not a point of priority, what, indeed, was the hurry?) Although he was not yet clear on exactly which of his papers had been read at the Linnean Society meeting, he was glad that Hooker planned to write Wallace about the affair, "as it would quite exonerate me [29]." Darwin was "more than satisfied" when he discovered what had been done: the strictly chronological (and alphabetical) arrangement of the papers meant that his preceded Wallace's (as does his name in the references to the published papers), when "I had thought that your letter and mine to Asa Gray were to be only an appendix to Wallace's paper [33]." 131 On 13 July Hooker and Darwin both sent letters to Wallace explaining the turn of events [31, 32]. Unhappily, these letters are missing, although Wallace carefully saved most of Dar- win's letters to him (and it is from them that we know of Wallace's early letters to Darwin). A few days later Darwin also thanked Lyell for his part, again expressing himself as "far more than satisfied [34]." He was pleased to have the public backing of men like Lyell and Hooker. It was only after some years of struggle, however, that Lyell became a "convert," while Hooker, although convinced of the action of natural selection, nevertheless vacillated on its importance. Even Darwin hedged as time went on. Of the four men, Wallace was to be the most steadfast, maintaining to the end his belief in "the overwhelm-

130. J. Linn. Soc. London (Zool.), 4 (1859), viii-ix. 131. The collective title for the joint papers and the accompanying letter is entered as follows: Charles Darwin and Alfred Russel 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. London (Zool.), 3 (1858), 45-62.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL ing importance of Natural Selection over all other agencies in the production of new species." 132 By 20 July Darwin had received the proof sheets, and he returned them to Hooker the next day, with "only a few corrections in the style [351." In fact, judging from the original texts as published in Evolution by Natural Selection and the Life and Letters of Charles Darwin, several hundred changes were made in both the 1844 sketch and the letter to Asa Gray, not only in the punctuation and wording but even in whole phrases.133 To have been scrupulously fair, it would seem that no changes should have been made at all. Darwin complained to Hooker that he had not been writing for publication [36], and a note to this effect was inserted in the Journal, as already mentioned. Wallace did not have these opportunities. It is not known who read the proof of his paper nor what became of his manuscript. In later reprintings he added phrases in footnotes that he would have inserted in the text, but he made no corrections then because of the historical importance of the document.134 The joint papers were published on 20 August 1858 in No. 9 of the third volume of the Linnean Society's Journal (Zoology). The sudden confrontation with Darwin threw Wallace into the limelight. But he had not stumbled upon the theory of natural selection by accident; he was to be neither a hanger-on nor a blind follower of Darwin's, and he was to make many valuable and original contributions of his own to evolutionary theory. His interests often paralleled those of Darwin, but his point of view frequently differed. Because of Darwin's illness and isolation at Down, their long and fruitful association is recorded in a correspondence that continued until Darwin's death.135 Wallace's first intimation of what had happened came when

132. Wallace, Darwinism, p. iv. 133. Compare the text of the joint papers with the 1844 excerpt in Darwin and Wallace, Evolution, pp. 116-121, and the letter to Asa Gray in Darwin, Life and Letters, I, 479-482. See also notes 83, 138, 150, and 158. 134. Wallace added two footnotes in the first reprinting in his Contri- butions to the Theory of Natural Selection (London: Macmillan, 1870); he omitted these and added a third in the second reprinting in Natural Selection and Tropical Nature (London: Macmillan, 1891), p. 27n, with the comment that "it must be remembered that the writer had no opportunity of correcting the proofs of this paper." 135. The complete extant correspondence is in Marchant, Wallace, pp. 107-262. The texts differ somewhat from those in Darwin, Life and Letters, besides the publication of sections omitted there. See also note 158.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection he received the letters from Hooker and Darwin in early Octo- ber 1858. At last he had aroused attention-attention that assured him, as he wrote his mother, "the acquaintance and assistance of these eminent men" on his return home. What- ever else Darwin's letter conveyed to Wallace, it was not the pain and anguish of those two final weeks in June. Wallace, unaware of the flurry he had aroused, told his mother that he was "highly gratified . . . I sent Mr. Darwin an essay on a subject on which he is now writing a great work. He showed it to Dr. Hooker and Sir C. Lyell, who thought so highly of it that they immediately read it before the Linnean Society [37]." And to an old boyhood friend, George Silk, Wallace crowed: "if you have any acquaintance who is a fellow of the Linnean Society, borrow the Journal of Proceedings for August last, and in the last article you will find some of my latest lucubra- tions, and also some complimentary remarks thereon by Sir Charles Lyell and Dr. Hooker, which (as I know neither of them) I am a little proud of [421." Almost thirty years passed before Wallace learned some of the details of Darwin's side of the story. Darwin, who received Wallace's answering letter in January 1859, was "extremely much pleased" with it; he had been "anxious to hear what your impression would be." He incorrectly referred to his own extracts as having been written in 1839, when in fact the first was written in 1844 and the second, the letter to Asa Gray, had been written in 1857, only a year and a half before [38, 43]. By November 1858 Wallace had received a copy of the Journal containing the papers and could read for himself Dar- win's "distinct and tangible idea." The tenor of Wallace's comments can be judged from Darwin's answer in the following April. Darwin agreed that Wallace was right in "that I came to the conclusion that selection was the principle of change from study of domesticated productions; and then, reading Malthus, I saw at once how to apply this principle. Geographi- cal distribution and geographical relations of extinct to recent inhabitants of South America first led me to the subject: especially the case of the Galapagos Islands [41, 45]." (But the letter in which this latter statement appeared was not published until 1903.) Again Darwin expressed his admiration for the manner in which Wallace had taken the publication of the papers. Actu- ally, it is hard to imagine what else Wallace could have done. Whatever reservations he might have had (and there is no indication that he had any), he was the forestaller, not the

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL forestalled, and he had at last achieved recognition from "two of the most eminent naturalists in England," a remarkable accomplishment for a self-educated collector. Darwin had set to work again almost immediately after the publication of the papers, making an "Abstract of Species book," and by November 1859 he could write Wallace that his publisher was sending him a copy of the Origin of Species, adding that "I do not think your share in the theory will be overlooked by the real judges, as Hooker, Lyell, Asa Gray, etc. [471." 136 Darwin had hoped that Wallace would feel that his Linnean paper was "fairly noticed" in the short Introduction and added that he would "allude" to the paper in the Annals in the body of the work [45]-this he did, but without giving either its title or the date (he always thought of this work as an abstract of the one he intended to write, and consequently he never gave proper references).1,37 But Darwin has never been accused of being overgenerous in his credits, particularly in the "Historical Sketch" later appended to the Origin of Species, and it may be that Wallace was too modest in his claims. Wallace wrote to congratulate Darwin on his book in Febru- ary 1860 [49], and Darwin sent the letter on to Lyell, remarking on "how admirably free [he was] from envy or jealousy. He must be a good fellow [53]."' This letter is among those missing, and so for Wallace's opinions it is necessary to turn to letters written to Bates in December 1860 and to his brother-in-law, Thomas Sims, in the following April. Perhaps better than anyone else, Wallace could appreciate the extraordinary amount of work involved, and he wrote Bates that he was thankful it had not been left for him to do. "Mr. Darwin has created a new science and a new philosophy, and I believe that never has such a complete illustration of a new branch of human knowledge been due to the labours and researches of a single man [55]." The letter to Sims is both a defense of Darwin and an explanation. Sims had apparently taken exception to Darwin's references to Wallace, and Wallace reproved him:

You quite misunderstand Mr. D.'s statement in the preface and his sentiments. I have, of course, been in correspondence with him since I first sent him my little essay. His conduct has been most liberal and disinterested. I think anyone who reads the Linnean Society papers and his book will

136. Darwin had sent copies of the joint papers to Wallace the preceding year; see Appendix, 40. 137. Darwin, Origin of Species, 1st ed., pp. 1-2, 355.

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see it. I do back him up in his whole round of conclusions and look upon him as the Newton of Natural History.

Sims objected not only to the contents of the Origin of Spe- cies but even to its title. After explaining that Darwin had originally given him a different title (presumably in the letter of 13 July 1858), Wallace went on to give his own judgment of Darwin's accomplishment: "It is the vast chaos of facts, which are explicable and fall into beautiful order on the one theory, which are inexplicable and remain a chaos on the other, which I think must ultimately force Darwin's views on any and every reflecting mind." The letter is worth reading carefully because, in trying to convince Sims, Wallace showed himself a strong and articulate champion of these views [561. It is a pity that Wallace was not at home to take an active part in the controversy over the Origin of Species. He would have enjoyed the dispute which Darwin found so distasteful. But by the time he returned to England in 1862 the first heat of the battle was past. Darwin, at any rate, was pleased with Wallace's enthusiasm, and he thanked him for his "too high approbation of my book . . . most persons would in your position have felt bitter envy and jealousy [54]." 138

But the documentation of this famous episode leaves something to be desired. There are gaps in the record that not only are rarely noticed but also are being gradually obliterated in the frequent retelling of this episode. Most people agree with Marchant that Darwin's letters tell the whole story, but they form only part of the evidence. Missing are all the letters (except for one fragment) that Wallace sent to Darwin from the Malay Archipelago, Wallace's manuscript, the letters written to Darwin by Hooker and Lyell during those hectic weeks in June 1858, and the pertinent letters from Asa Gray.'39 The evidence on the disposition of the letters to Darwin is contradictory. Francis Darwin later recalled that his father "made a rule of keeping all letters that he received; this was a habit which he learnt from his father, and which he said had been of great use to him." 140 But, in describing his own work in compiling the Life and Letters of Charles Darwin, he gave a different account:

138. "Bitter envy" changed to "some envy" in Darwin, Life and Letters; see also note 150. 139. The missing letters are nos. 8, 13, 15, 20, 27, 28, 30, 38, 39, 41, 44, 48, 49, marked with an asterisk in the Appendix. Letters nos. 31 and 32 from Darwin and Hooker to Wallace are also missing. 140. Darwin, Life and Letters, I, 97.

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Of letters addressed to my father I have not made much use. It was his custom to file all letters received, and when his slender stock of files ("spits" as he called them) was ex- hausted, he would burn the letters of several years, in order that he might make use of the liberated "spits." This process, carried on for years, destroyed nearly all letters received before 1862. After that date he was persuaded to keep the more interesting letters, and these are preserved in an ac- cessible form.141

Many of the letters received before 1862 do exist, however, as can easily be checked in the published collections of Darwin's letters and in the list of unpublished material in the Cambridge University Library.142 Another version of what happened to the letters is given in a biography of Hooker:

In one of his letters Darwin makes special mention of pre- serving his friend's [i.e. Hooker's] letters. The answers to scientific questions are detached and placed among the memoranda of that subject; the other parts are put away among his general correspondence, so that it would only be a matter of half an hour to rearrange them in case of need. In spite of his care, however, a large number of the earlier letters from Hooker have disappeared wholly or in part.143 The one snippet from Wallace's letter of 27 September 1857 shows that Darwin must have followed this practice with his letters also, at least at first [16]. But Darwin was meticulous (indeed, it is from his own care in answering Wallace that the dates of Wallace's letters to him can be so easily determined). It seems surprising that all the material relating to the most dramatic (not to say traumatic) moment in his life should disappear. The dating of Darwin's own letters presents still another problem. As his son observed: He rarely dated his letters, so that but for the Diary [Jour- nal] it would have been all but impossible to unravel the history of his books. It has also enabled me to assign dates to many letters which would otherwise have been shom of half their value,144 [and]

141. Ibid., xviii-xix. 142. Cambridge University Library, Handlist of Darwin Papers at the University Library, Cambridge (Cambridge: Cambridge University Press, 1960). 143. L. Huxley, Life and Letters ... Hooker, I, 436. 144. Darwin, Life and Letters, I, xviii.

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Mr. Darwin, who was careful in other things, generally omitted the date in familiar correspondence, and it is often only by treating a letter as a detective studies a crime that we can make sure of its date. Fortunately, however, Sir Joseph Hooker and others of Darwin's correspondents were accustomed to add the date on which the letters were received.145

Some of the crucial letters on the theory of natural selection from Darwin to Asa Gray have recently been redated by Du- pree in a different connection, a redating that can be supported by an examination of the texts. These letters have appeared in different collections of Darwin's letters and have been dated 1856 [12], 1857 [14], and 1859 [17], respectively, but they seem aUl to have been written in 1857, after Darwin received Wallace's first letter. Put together and read in sequence, they seem to form a natural unit, lending support to the theory that the revelation to Gray was induced by Wallace.146

VI. EPILOGUE

I feel much satisfaction in having thus aided in bringing about the publication of this celebrated book, and with the ample recognition by Darwin himself of my independent discovery of "natural selection."

Wallace, Natural Selection and Tropical Nature

To a large extent the world has accepted at face value both Wallace's and Darwin's rather pious recollections of their close

145. Darwin, More Letters, I, x. 146. There are two ways to date the letters to Gray. One is by the sequence in the Darwin-Gray correspondence, and the other is from internal evidence. Both indicate that no. 12 was written in 1857. In particular, Darwin refers to a chapter on the continuous distribution of species, which "Hooker kindly read . . . over." According to Darwin's "Journal," he finished this section on 13 October 1856, having asked Hooker on 13 July 1856 if he would read it for him (Darwin, More Letters, I, 95). See also note 88. Although there is no question about the year (1857) of no. 14, Darwin did not date the copy he kept, and he thought he had sent it in October (Darwin, Life and Letters, I, 477n). The date is given as October in the letter of transmittal to the Linnean Society but as 5 September 1857 on the letter itself. See also note 90. Number 17 was probably put in 1859 because it concerns natural selection. But Darwin's book was published on 24 November, and Gray could not have received a copy, read it, and given Darwin his opinion in 5 days. Again, the sequence of letters shows that it belongs in 1857. It is followed by two others that have also been dated 1859 but which should be dated 1858, one on 21 February (Darwin, Life and Letters, I, 463-464) and another on 4 April (ibid., 510). The latter refers to a just finished chapter on instinct that, according to Darwin's "Journal," was finished on 9 March 1858. See also note 93.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL but not always unruffled relationship. The veil of Victorian propriety through which they came to view each other has, however, obscured some of their more human reactions to what must at times have been a trying entanglement. Public recognition by Darwin of Wallace's achievement can be divided into two parts, before and after publication of the Life and Letters of Charles Darwin in 1887, the first part consisting mainly of remarks in the "Historical Sketch" appended to the Origin of Species. This sketch (which has been called "the most unreliable account that ever will be written")'47 was added to the third edition in 1861 and expanded somewhat for the fourth in 1866.148 Wallace and the Linnean papers were given one sentence, while Wallace's paper in the Annals was not mentioned at all and Lyell was omitted altogether. Among the names added in 1866 were those of Patrick Matthew and William Wells, both contenders for the title of discoverer of the theory of natural selection. Matthew had written on natural selection in 1831, and he pressed his claim in the Gardners' Chronicle in 1860.149 Dar- win publicly acknowledged his anticipation of the theory, writing to Wallace that "he gives most clearly but very briefly ... our view of Natural Selection [51, 52, 54]."150 Wells' claim was made in 1865 by a "Mr. Rowley, of the United States," for "an account of a female . . . part of whose skin resembles that of a negro," a paper first read before the Royal Society in 1813 and published posthumously in 1818.'15 Wallace was surprised "that it should have struck no one that [his suggestion] was a great principle of universal application in Na- ture [60] I" Shortly before this, Wallace had modestly referred to "Mr. Darwin's celebrated theory of 'Natural Selection.' 152 Darwin had demurred, but Wallace had insisted that he would "always

147. C. D. Darlington, "The Origin of Darwinism," Sci. Amer., 200 (May 1959), 61. 148. Darwin, Origin of Species, 3rd ed., pp. v-xi; 4th ed., pp. xiii-xxi. 149. Patrick Matthew, Naval Timber and Arboriculture (Edinburgh: Longman, 1831); "Nature's Law of Selection," Gard. Chron. Agricul. Gaz., 20 (7 April 1860), pp. 312-313. 150. Darwin, Life and Letters, Jl, 95-96n. The remark on Patrick Matthew was omitted from Darwin, Life and Letters; see also note 138. 151. William Wells, Two Essays: one upon Single Vision with two Eyes; the other on Dew. A letter to . . . Lord Kenyon and an Account of a Female . . . part of whose Skin resembles that of a Negro (London: A. Constable, 1818). 152. Alfred Russel Wallace, "The Origin of Human Races and the Antiquity of Man deduced from Natural Selection," J. Anthropol. Soc. London, 2 (1864), clix.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection maintain it to be actually yours, and yours only [57, 58]." And yet in 1869 Wallace's reply to a request from the anthropolo- gist A. B. Meyer for his recollections of his part in the theory was brusque. After stating that he "was led to it by Malthus' views on population applied to animals," his terse account ended with the remark that his paper "was printed without my knowledge, and of course without any correction of proofs. I should, of course, like this fact to be stated [63]." Wallace objected to being classed with Matthew and Wells, "who made no further use of that principle, and failed to see its wide and immensely important applications," and he published a collection of his papers in 1870 to make the extent of his own contribution clear.153 But he did not then, nor did he ever, claim that he had worked out the theory in the detail that Darwin had. Darwin thanked Wallace in glowing terms for the kind words in his preface, having missed altogether the point about Matthew and Wells [65]. Lyell, in the meantime, thought Darwin had given short shrift not only to Wallace and himself, but also to Lamarck. He was astonished to find no mention of Wallace's paper in the Annals in Darwin's "Historical Sketch" [61], and he discussed this "next important effort to determine the manner in which new species may have originated" in some detail in the new edition of his famous Principles of Geology, published in 1867-1868.154 He had already credited Wallace with thinking out, "independently for himself, one of the most novel and important of Mr. Darwin's theories," in discussing the joint papers in his Antiquity of Man.155 And in his Principles of Geology, he now reprinted in its entirety his original summary of Lamarck, not to protest against his theory of the transmutation of species as before, but to "show how nearly the opinions taught by him at the commencement of this century resembled those now in vogue." 156 As for himself, Lyell wrote in a letter to Ernst Haeckel in November 1868 that he was obliged to him "for pointing out [in his History of Creation] how clearly I advocated a law of continuity even in the organic world, so far as possible without adopting Lamarck's theory of transmutation . . . I had certainly prepared the way in this country . . . for the reception of Darwin's gradual and insensible evolution of species [62]," and Huxley later agreed with him.157 In 1870, apparently in

153. Wallace, Contributions, p. iv. 154. Lyell, Principles of Geology, 10th ed., II, 276-281. 155. Lyell, Geological Evidences, 1st ed., pp. 408-409. 156. Lyell, Principles of Geology, 10th ed., II, 246n. 157. Darwin, Life and Letters, I, 543-544.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL reference to the preface to Wallace's compilation of his papers, Lyell again supported Wallace, writing him that "it is high time this modest assertion of your claims as an independent originator of Natural Selection should be published [64]."

Not until the publication of the Life and Letters of Charles Darwin in 1887, five years after Darwin's death, did Wallace receive full public recognition for his part in the theory of natural selection. The letters from Darwin to Hooker, Lyell, and Wallace himself were there for all to read (although neither the complete texts nor all of the early letters from Darwin to Wallace were included 158); and as further proof of Wal- lace's role in inducing Darwin to publish the Origin of Species, there was also Darwin's "Autobiography." For the first time, Wallace learned some of the details of those long-ago events. He was surprised to find that Darwin "had been so distressed- or rather disturbed" by his essay, and he wrote apologetically to Francis Darwin that he had always felt that he had received too much credit "for my mere sketch of a theory [68]." Darwin's attitude had also softened. Nearly twenty years after the event, in an autobiographical sketch written mainly for the eyes of his children, he could say that he "cared very little whether men attributed most originality to me or Wal- lace." 159 He had worried whether Wallace would consider the whole proceeding justifiable (about which both he and Hooker seemed to have some lingering doubts), not then knowing "how generous and noble was his disposition." 100 He mentioned once again that his own parts of the joint papers had not been intended for publication, while Wallace's was a model of clarity. He never recognized in any way that Wal- lace's was hastily written, that it had not been intended for publication either, or that Wallace had had no chance to proof- read it, but he did credit Wallace with giving him the impetus that produced the Origin of Species. Now that Wallace's position was secure, he began to em- broider his own recollections. At the time he wrote his essay, nearly thirty years before, he had had no idea either of its importance or of its impact on Darwin, and the earliest re-

158. Of the first eight extant letters from Darwin to Wallace, six are included in Darwin, Life and Letters (nos. 11, 18, 43, 46, 47, and 54), but only one (47) is complete. The seventh (45) is in Darwin, More Letters, and the eighth (50) is in Marchant, Wallace. Marchant includes the complete extant text of all of them, but his book was not published until 1916. The texts also differ slightly from those in Darwin, Life and Letters. See also notes 3, 87, 135, 138-143, 146, 150, and 171. 159. Darwin, Life and Letters, I, 71. 160. Ibid., 69.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms Wallace, Darwin and the Theory of Natural Selection quest for his memories had come from Meyer in 1869. But now the circumstances of that week so long ago assumed a new interest. Alfred Newton, ornithologist and zoologist, wrote inquiring for details to incorporate in his review of the Life and Letters of Charles Darwin, and Wallace obligingly responded. But the letter to Newton [69], written in 1887, contains a number of questionable statements. Wallace was now uncertain whether he had even read Darwin's Voyage of the Beagle at the time, when he had in fact read both editions and had the second one with him in the Malay Archipelago; scattered references to Darwin in his own early published works also show that he had read him with some care. He thought he had started the correspondence over some peculiar varieties of ducks. However, the letter itself shows that he was consulting the Life and Letters of Charles Darwin to refresh his memory, and the paragraph containing the request for "any curious breed" in Darwin's first letter to him was omitted there. Evidence from Wallace's "Notebook" also indicates that Darwin brought the subject up first [11].161 Notices in the Athenaeum that Darwin was interested in species and varieties seem improbable.162 Wallace again paid his respects to Mal- thus, further enshrining him in the annals of science. And finally, he referred to a "hot fit" of intermittent fever, although he later said the idea had come to him during a "cold fit." Wallace had returned only a few months before from a speaking tour of the United States. Interestingly enough, he had met Asa Gray during a month-long stay in Boston, and Gray had invited him to attend a meeting of the Cambridge Scientific Club. There, Gray showed his correspondence with Darwin before the publication of the Origin of Species,

161. See also notes 86 and 87. 162. No such notices located. Although it is possible that Wallace heard of Darwin's interest through his agent, Stevens (who had been a member with Darwin since 1837 of the Entomological Society), and he could have used them as an opening for his first letter in 1856 (8), this still would not account for the range of subjects he apparently discussed. Darwin answered: "By your letter and even still more by your paper in the Annals . . . ," indicating that the letter itself must have contained some related remarks (11). Sydney Smith has suggested (personal communication) that two letters from Darwin to W. B. Tegetmeier, dated 21 Nov. and 29 Nov. 1857, show that it was about the second date that Darwin received some poultry specimens collected by Wallace. Since Wallace received Darwin's letter (11) with this request in July (see 19), the shipment could have been in response to that request. This would leave unaffected the first letter Wallace wrote to Darwin in October 1856 (8), and the reason for it would then remain an open question.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms BARBARA G. BEDDALL and Wallace "related what led him to his theory of Natural Selection . . . The writings of Spencer, Vestiges of Creation, Lamarck?, but particularly of Malthus on population suggested his own view." 163 This also seems inaccurate: Wallace omitted Lyell and added Spencer, whose First Principles he read after his return from the Malay Archipelago, in September 1862.1'4 At any rate, perhaps heartened by the public recognition he had now received, Wallace expanded his series of lectures into a book, published in 1889, to which he gave the title, Darwinism: An Exposition of the Theory of Natural Selection. To him the terms were synonymous. Furthermore, he was still convinced of the prime importance of natural selection, although Darwin had staged a gradual retreat from this position. Next, in the introductory note to a chapter in a new compilation of his papers, Natural Selection and Tropical Nature, Wallace added further details of that now never-to-be-forgotten week, even to the temperature outside. Again he consulted the Life and Letters of Charles Darwin, and another probable error crept into the story. From the dates on which the first two Darwin letters were written, Wallace presumed that he had received them before sending his paper to Darwin, and he quoted from them both. It is unlikely, however, that the second letter could have reached him so quickly. But he was now satisfied, as can be seen from his concluding remark, "with the ample recognition by Darwin himself." 165 Wallace gave an even more detailed account in his own autobiography, written nearly fifty years after the event, adding, among other things, that he had asked Darwin to show his paper to Lyell "who had thought so highly of my former paper." This again presumes that Wallace had already received Darwin's second letter, which is doubtful. This frequently quoted-from account should be treated with caution.'66 In thanking Wallace for a copy of his Life, Hooker, now eighty-eight years old, remarked that "your citation of my letters and their contents are like dreams to me; but to tell you the truth, I am getting dull of memory as well as of hearing, and what is worse, in reading: what goes in at one eye goes out at the other"-perhaps an honest evaluation that could be applied to most octogenarians, including Wallace.'67

163. Cambridge Scientific Club, "Record Book," 17 November 1886, MS, Harvard University Archives; quoted with the perrmission of Harvard University. See also Dupree, Asa Gray, pp. 380, 473 n52. 164. Marchant, Wallace, p. 122. 165. Wallace, Natural Selection, pp. 20-21. 166. Wallace, My Life, I, 357-363. 167. Marchant, Wallace, p. 332.

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On 1 July 1908, the Linnean Society held a jubilee to com- memorate the reading of the joint papers fifty years before. It seems extraordinary that two of the protagonists, Wallace and Hooker, were still alive and able to participate. (Lyell had died in 1875 and Darwin in 1882.) Wallace, the first of the two to speak, repeated his story with some new elaborations, helping to perpetuate the (perhaps fictitious) report that his paper had come to Darwin 'like a thunderbolt from a cloud- less sky," but his account was modest and self-effacing.'68 He was, after all, a modest man. He had refused some of the honors offered him and had accepted others (such as an honor- ary degree from Oxford and a Fellowship in the Royal Society) only after strong urging. He had long since received the credit he felt due him as an independent originator of the theory of natural selection. In the published report of the proceedings, Wallace added some selections from the sixth edition of Malthus that he thought might have influenced him, remarking, however, that it was the over-all effect rather than particular details that he remembered. He concluded with a well-deserved tribute to Lyell who had, as he said, impressed him even more deeply than Malthus.'69 Then it was Hooker's turn. It is startling to discover that Hooker's first biographer, Leonard Huxley, referred to Hooker as the "sole survivor of those immediately concerned," dismiss- ing Wallace altogether (an oversight repeated by his latest biographer),170 and he mentioned just as casually that "one or two of the letters that then passed were missing." 171 But Hooker, in accepting the invitation to speak, was rather anxious not only about the expediency and propriety of telling the public what he had done, but also about the accuracy of his recollections. He appealed to Sir Francis Darwin and to Sir Leonard Lyell, Sir Charles's nephew, for help in finding additional documentary evidence. But none could be found, and he was forced to rely entirely on the partial story in the Life and Letters of Charles Darwin. He carefully noted this in his speech, apologiz- ing at the end for "the half-century-old real or fancied memories of a nonagenarian." 172 But Wallace had saved the early letters he received from

168. Linnean Society of London, The Darwin-Wallace Celebration, pp. 5-11. 169. Ibid., pp. 111-118. 170. Mea Allan, The Hookers of Kew, 1785-1911 (London: Michael Joseph, 1967), p. 248. 171. L. Huxley, Life and Letters . .. Hooker, II, 465. 172. Linnean Society of London, The Darwin-Wallace Celebration, pp. 11-16.

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Darwin (except for one), although the manuscript of his famous paper was never found. On the outside of an envelope in which he kept the letters, Wallace wrote:

The first 8 letters I received from Darwin-while in the Malay Archipelago. NB. The MSS. of my Paper sent to Darwin and printed in the Journal of the Linnean Society, was not returned to me, and seems to be lost. The proofs with the MSS. were perhaps sent to Sir Charles Lyell, or to the Secretary of the Linn. Soc. & may some day be found. It was written on thin foreign note paper.'73

However, "neither Wallace's part of this correspondence, nor the original MS. of his essay . . . has been discovered," as Marchant wrote after Wallace's death.'74 And so the story rests, with some questions still unanswered. Why did Wallace first write to Darwin? Why did Darwin send the outline of his theory of natural selection to Asa Gray? What became of the letters Darwin received from Wallace, Lyell, Hooker, and Gray? Where is Wallace's manuscript? The answers are in the missing material, and what really happened must remain speculation. The fact that much other material is also missing does not invalidate the point that evidence to support some commonly accepted explanations is inadequate or lacking and that other explanations are clearly in error.

ACKNOWLEDGMENTS I would like to express my appreciation to Dr. Ernst Mayr for critically reading the manuscript, and to Dr. Everett Men- delsohn for editorial help in preparing it for publication. I would also like to thank Miss Sandra Raphael, Librarian of the Linnean Society of London, for her kind assistance. Fi- nally, I would like to acknowledge my husband's encourage- ment and financial support, without which this would not have been done.

173. Marchant, Wallace, p. 106. 174. Ibid., p. 105.

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This content downloaded from 150.135.165.122 on Tue, 24 Jul 2018 01:35:11 UTC All use subject to https://about.jstor.org/terms APPENDIX * LETTERS

No.1 From - to Date2 Sources3 Comments

1. A. R. Wallace- 11 April 1846 W, 1, 255-256; Omission in M. H. W. Bates M, 21.

2. A. R. Wallace- 9 Nov. [1847] W, 1, 254; For dating, see H. W. Bates M, 73. note 14.

3. A. R. Wallace- 28 Dec. [18471 W, 1, 254-255; Omission in M; for H. W. Bates M, 73-74. dating, see note 14.

4. A. R. Wallace- [Early 1848] W, 1, 256-257; Omission in M. H. W. Bates M, 74-75.

5. C. Darwin- 3 May [1856] DLL, 1, 426-427; C. Lyell DLLE, 2, 67-68.

6. C. Darwin- 9 May [1856] DLL, 1, 427-428; 3. D. Hooker DLLE, 2, 68-69.

7. C. Darwin- 11 May [1856] DLL, 1,428-430; J. D. Hooker DLLE, 2, 69-71.

*8. A. R. Wallace- [10 Oct. 1856] Received late April C. Darwin 1857; see 11 and 69, and notes 43, 81, and 162.

9. H. W. Bates- 19 Nov. 1856 M, 52-53. Received July 1857; A. R. Wallace see 19.

10. C. Darwin- 22 Feb. 1857 DLL, 1, 452; W. D. Fox DLLE, 2, 94-95.

11. C. Darwin- 1 May 1857 DLL, 1, 452-454; Answer to 8, A. R. Wallace DLLE, 2, 95-96; received July 1857; M, 107-109. see 19. Omissions in DLL; see notes 86, 87, and 162.

12. C. Darwin- 20 July [1857] DLL, 1, 437-438; See also 14 and 17. A. Gray DLLE, 2, 78-80; For dating, see Du, 244-245,458 note 146. n22.

*13. A. Gray- [Aug. 1857] - See 14. C. Darwin

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No.1 From - to Date2 Sources8 Comments

14. C. Darwin- 5 Sept. [1857] JLZ, 3, 50-53; See also 12 and 17. A. Gray DLL, 1, 477-482; Many differences in DLLE, 2, 122-125; texts between JLZ Du, 246,458-459 and DLL. For dating, n23. see note 146; see also note 92. *15. A. Gray- [Autumn 1857] See 17. C. Darwin

16. A. R. Wallace- [27 Sept. 1857] CUL Unpublished frag- C. Darwin ment of answer to 11. Received Dec. 1857; see 18. See also notes 60, 95, 142, and 143.

17. C. Darwin- 29 Nov [1857] DML, 1, 126-127; Not published undl A. Gray Du, 247,459 n24. 1903. See also 12 and 14. For dating, see note 146.

18. C. Darwin- 22 Dec. 1857 DLL, 1,465-467; Answer to 16. A. R. Wallace DLLE, 2, 108-110; Omission in DLL. M, 109-111.

19. A. R. Wallace- 4 & 25 Jan. 1858 W, 1, 358-359; Answer to 9. H. W. Bates M, 53-55. Omissions in W, and texts differ slightly; here quoted from M. See also note 94.

*20. A. R. Wallace- [Feb. 1858] Received 18 June C. Darwin 1858; see 22. See also note 116.

21. C. Darwin- 8 June [1858] DML, 1, 109. Not published J. D. Hooker until 1903.

22. C. Darwin- 18 June [1858] DLL, 1, 473; C. Lyell DLLE, 2, 116-117.

23. C. Darwin- [25 June 1858] DLL, 1, 474-475; C. Lyell DLLE, 2, 117-118.

24. C. Darwin- 26 [June 1858] DLL, 1, 475; C. Lyell DLLE, 2, 118-119.

25. C. Darwin- [29 June 1858] DLL, 1, 476; J. D. Hooker DLLE, 2,119.

26. C. Darwin- [29 June 1858] DLL, 1, 476-477; J. D. Hooker DLLE, 2, 119-120.

*27. C. Lyell- [June 1858] See 25 and notes C. Darwin 3, 139-143, and 171.

*28. J. D. Hooker- [June 1858] See 25 and 26, and C. Darwin notes 3, 139-143, and 171.

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No.1 From - to Date2 Sources8 Comments

29. C. Darwin- S July [1858] DLL, 1, 482-484; J. D. Hooker DLLE, 2,124-127. *30. J. D. Hooker- [July 1858] - See 33. C. Darwin

*31. J. D. Hooker- [13? July 1858] Received early A. R. Wallace October 1858; see 33, 37, and 42. *32. C. Darwin- [13 July 1858] Answer to 20, A. R. Wallace received early October 1858; see 33,37, and 42.

33. C. Darwin- [13 July 1858] DLL, 1,484-485; J. D. Hooker DLLE, 2, 128-129.

34. C. Darwin- 18 July [1858] DLL, 1,485-486; C. Lyell DLLE, 2,129-130.

35. C. Darwin- 21 July [1858] DLL, 1, 486-487; J. D. Hooker DLLE, 2, 130-131.

36. C. Darwin- [5 Aug. 1858J DLL, 1,489-490; J. D. Hooker DLLE, 2,133.

37. A. R. Wallace- 6 Oct. 1858 W, 1,365; Omissions in W. his mother M, 57-58.

*38. A. R. Wallace- [Oct. 1858] Received [22] January C. Darwin 1859; see 43.

*39. A. R. Wallace- [Oct. 1858] Received [22] January J. D. Hooker 1859; see 43.

40. C. Darwin- 12 Oct. 1858 DLL, 1,494; J. D. Hooker DLLE, 2, 138.

*41. A. R. Wallace- [30 Nov. 1858] Received 6 April C. Darwin 1859; see 45.

42. A. R. Wallace- [Nov. 1858] W, 1,365-367. G. Silk

43. C. Darwin- 25 Jan. [1859] DLL, 1, 501-502; Answer to 38; A. R. Wallace DLLE, 2, 145-147; omissions in DLL. M, 111-112. *44. A. R. Wallace- ? Received 7 August C. Darwin 1859; see 46.

45. C. Darwin- 6 April 1859 DML, 1, 118-120; Answer to 41, but not A. R. Wallace M, 112-114. published until 1903.

46. C. Darwin- 9 Aug. 1859 DLL, 1, 516-517; Answer to 44; A. R. Wallace DLLE, 2, 161-162; omission in DLL. M, 114-115.

47. C. Darwin- 13 Nov. 1859 DLL, 2, 16-17; A. R. Wallace DLLE, 2, 220-221; M, 115-116.

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No.1 From - to Date2 Sources3 Comments

*48. A. R. Wallace- ? Received March? C. Darwin 1860; see 50. *49. A. R. Wallace- [16 Feb. 1860] Received 18 May C. Darwin 1860; see 54.

50. C. Darwin- 7 March 1860 M, 116. Answer to 48; not A. R. Wallace published until 1916.

51. C. Darwin- 10 April [1860] DLL, 2, 93-95; C. Lyell DLLE, 2, 300-301.

52. C. Darwin- [13 April 1860] DLL, 2, 95-96; J. D. Hooker DLLE, 2,301-303.

53. C. Darwin- 18 May [1860] DLL, 2, 101-102; C. Lyell DLLE, 2, 308-309.

54. C. Darwin- 18 May 1860 DLL, 2, 102-103; Answer to 49. A. R. Wallace DLLE, 2,309-310; Omissions and M, 117-118. changes in DLL; see notes 138, 150.

55. A. R. Wallace- 24 Dec. 1860 W, 1,373-375; Omissions in M. H. W. Bates M,59.

56. A. R. Wallace- 15 March 1861 M, 59-67. T. Sims

57. C. Darwin- 28 [May? 1864] DLL, 2,271-273; A. R. Wallace DLLE, 3,89-91; DML, 2,32-34; M, 127-128.

58. A. R. Wallace- 29 May [1864] DML, 2,34-37; C. Darwin M, 128-131.

59. C. Darwin- [Oct. 1865] DLL, 2, 225; J. D. Hooker DLLE, 3, 41.

60. A. R. Wallace- 19 Nov. 1866 M, 145-146. C. Darwin

61. C. Lyell- 4 April 1867 M, 279-281. A. R. Wallace

62. C. Lyell- 23 Nov. 1868 LLL, 2,435-437. E. Haeckel

63. A.R. Wallace- 22 Nov. 1869 Nat., 52 (1895), 415. Not published in its A. B. Meyer entirety until 1895.

64. C. Lyell- 15 Feb. [1870?] M, 288-289. A. R. Wallace

65. C. Darwin- 20 April [1870] DLL, 2, 301-302; A. R. Walace DLLE, 3,121; M, 206-207.

66. G. Bentham- 30 May 1882 DLL, 2, 87-88; F. Darwin DLLE, 2,293-294.

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No.1 From - to Date2 Sources8 Comments

67. J. D. Hooker- 22 Oct. 1886 HLL, 2, 300-302. F. Darwin

68. A. R. Wallace- 20 Nov. 1887 M, 295. F. Darwin

69. A. R. Wallace- 3 Dec. 1887 DAU, 200-201; See also notes 7, A. Newton DAUE, 189-190. 39-41,43,81,86, 87, 161, and 162.

1. Missing letters, known to have existed from other correspondence, are marked with an asterisk; see "Comments". 2. Uncertain dates of special interest here are discussed in the notes; see "Comments". 3. Sources are arranged chronologically. Most of them are fully cited in the notes, the rest here. They are abbreviated as follows: CUL Cambridge University Library. DAU Francis Darwin, ed., Charles Darwin: His Life Told in an Auto- biographical Chapter and in a Selected Series of His Published Letters (New York: D. Appleton, 1892). DAUE English edition of above (London: Murray, 1892). DLL F. Darwin, Life and Letters of Charles Darwin. DLLE English 3-volume edition of above (London: Murray, 1888). DML F. Darwin, More Letters of Charles Darwin. Du Dupree, Asa Gray. HLL L. Huxley, Life and Letters of Sir Joseph Dalton Hooker. JLZ Journal of the Linnean Society of London (Zoology). LLL K. Lyell, Life, Letters and Journals of Sir Charles Lyell. M Marchant, Alfred Russel Wallace. Nat. Nature. W Wallace, My Life.

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