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Wallace, Darwin, and the Practice of Author(s): Melinda B. Fagan Source: Journal of the History of , Vol. 40, No. 4 (Dec., 2007), pp. 601-635 Published by: Springer Stable URL: http://www.jstor.org/stable/29737514 . Accessed: 02/06/2013 13:59

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This content downloaded from 150.135.114.31 on Sun, 2 Jun 2013 13:59:18 PM All use subject to JSTOR Terms and Conditions Journal of the History of Biology (2007) 40:601-635 ? Springer 2007 DOI 10.1007/s 10739-007-9126-8

Wallace, Darwin, and the Practice of Natural History

MELINDA B. FAGAN Department of History and Indiana University Goodbody Hall 130, 1011 East Third St Bloomington, IN 47405 USA

E-mail: me fagan @ indiana, edu

Abstract. There is a pervasive contrast in the early natural history writings of the co-discoverers of natural selection, Alfred R?ssel Wallace and . In his writings from South America and (1848-1852, 1854-1862), Wallace consistently emphasized species and genera, and separated these descriptions from his rarer and briefer discussions of individual . In contrast, Darwin's writings during the Beagle voyage (1831-1836) emphasized individual organisms, and mingled descriptions of individuals and groups. The contrast is explained by the different practices of the two naturalists in the field. Wallace and Darwin went to the field with different educational experiences and social connections, constrained by different responsibilities and theoretical interests. These in turn resulted in different natural history practices; i.e., different habits and working routines in the field. Wallace's intense collecting activities aimed at a complete inventory of different species and their distributions at many localities. Darwin's less intense collecting practice focused on detailed of individual organisms. These different practices resulted in different material, textual and conceptual products. Placing natural history practices at the center of analysis reveals connections among these diverse products, and throws light on Wallace and Darwin's respective treatment of individuals and groups in natural history. In particular, this approach clarifies the relation between individuals and groups in Wallace's theory of natural selection, and provides an integrative starting point for further investigations of the broader social factors that shaped Victorian natural history practices and their scientific products.

Keywords: Alfred R?ssel Wallace, Charles Darwin, natural history, scientific practice, natural selection, specimen collecting, Beagle voyage, Malay Archipelago

Introduction

Then a host of new species burst upon me, revealing the richness of the country, and its intimate connection with New Guinea. Paridisea apoda, L., Pregia, L., Microglossus aterrimus, Wagl.,

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Brachyurus Macklotti, Temm., B. novae guineas, Schlegel, Tany siptera, sp., Eurystomus gularis, Vieill., Carpophaga, n.s., with several small flycatchers, thrushes, and shrikes, and that most magnificent of the swallow-tribe, Macropteryx mystaceus, Less., were what I now obtained... The following families are abundant in species and in individuals. They are everywhere common " (italics in original)1

- I shot a condor, it measured from tip to tip of wing 8 & 1/2 feet; - from beak to tail 4 feet. They are magnificent birds; when seated on a pinnacle over some steep precipice, sultan-like they view the plains beneath them. I believe these birds are never found excepting where there are perpendicular cliffs: further up the river, where the lava is 8 & 900 feet above the bed of the river, I found a regular breeding place; it was a fine sight to see between ten & twenty of these Condors start heavily from their resting spot & then wheel away in majestic circles.*"

These passages illustrate a pervasive contrast in the early writings of Alfred R?ssel Wallace and Charles Darwin. Both naturalists spent years in the field before independently developing their theories of natural selection.3 Wallace worked as a specimen collector in South America (1848-1852) and theMalay Archipelago (present-day Indonesia; 1854? 1862), while Darwin acquired his specimens during the Beagle voyage (1831-1836), primarily from South America. In his notes, essays and correspondence from the field, Wallace consistently emphasized species and genera, and separated these descriptions from his rarer and briefer discussions of individual organisms. The first passage above, from an 1857 article describing collecting in the Aru Islands, is typical: Wallace provides an enthusiastic litany of species, families and genera. It is easy to miss his distinction at the end of the passage, between families, species and individuals, in ternis of "abundance." Yet this too is

1 Wallace, 1857d, pp. 476, 479. 2 Keynes, 1988, p. 237. 3 Darwin and Wallace, 1858, Darwin, 1968 [1859]. McKinney (1972, pp. 97-155), Brackman (1980) and Brooks (1984, pp. 200-269) argue that Darwin drew on Wallace's 1858 essay for his own theory. Browne (1980), Beddall (1988), Kohn (1985, pp. 245 257), and Raby (2001, pp. 129-142) effectively criticize this 'conspiracy theory.' The current consensus is that the theories were conceived independently {cf. Browne 2003, pp. 14-45).

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characteristic of Wallace's writings from the field. At a given locality, families contain more or fewer species, and species contain more or fewer individual organisms. Wallace did not collapse or confuse these levels, but carefully distinguished between different sorts of abundance. In general, his natural history writing emphasized species, with clear distinctions between individual organisms and groups.4 By contrast, as illustrated by the second passage, Darwin's notes and correspondence during the Beagle voyage emphasized individual organisms and mingled descriptions of individuals and groups. The representative journal entry above, from April 1834, is laced with carefully observed details of individual birds: measurements of wing span, numbers at the breeding site. Though individual details are prominent, Darwin also discussed the "magnificent birds" as a group, shifting rapidly from the condor he shot, to condors in general, to particular views of condors. General observations are juxtaposed with anecdotes of particular events. These mingled modes of description make for a vivid and engaging style that is difficult to parse precisely. Emphasis on individual details and ambiguity as to level of description are characteristic of Darwin's writings from the field. The contrast in the two naturalists' writings from the field thus has two aspects. First, Wallace emphasized groups of organisms, while Darwin described many details of individual organisms. Second, Wal? lace clearly distinguished between groups and individuals, while Darwin was more ambiguous. Both aspects can be explained by differences in natural history practice. Wallace and Darwin's contrasting habits and working routines in the field were shaped in turn by their different circumstances and motivations. The two naturalists went to the field with different training and social connections, different finances and responsibilities, and different theoretical interests. These contrasts led Wallace and Darwin to practice natural history at different intensities, using different methods and standards. Wallace's intense collecting activities were aimed at obtaining a complete inventory of different species and their distributions at many localities. Darwin's less intense

4 I do not engage the metaphysical debate concerning biological groups and indi? viduals (see Hull, 1989; Ghiselin, 1997; Gould, 2002). I assume everyday notions of groups and individual organisms, the latter being simply and that live on the earth, and the former assemblages of these treated as entities in their own right. Populations, varieties, races, species, genera, and higher taxa are groups in this sense. My argument requires only that the distinction between individual organisms and groups is robustly drawn, whatever its metaphysical basis.

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but meticulous collecting practices aimed at detailed of new and interesting individual organisms. A number of historians have recently examined the staggering scope of Wallace's collecting activities, as well as his pioneering role in bio .5 Darwin's Beagle voyage has been extensively studied; re? cent studies have highlighted its importance for Darwin's scientific networking and subsequent career.6 This study goes beyond previous work by placing natural history practice at the center of analysis, offering an integrative account of the contrast between Wallace and Darwin that reveals links between their respective social contexts, nat? ural history practices, and theories of natural selection. Over time, the naturalist's routine practice yields material, literary and conceptual products: collections of specimens; writings pertaining to animals and plants; concepts and theories about these objects of study. In the case of Darwin and Wallace, the collections, writings and theories resulting from their respective practices show interrelated contrasts. Focusing on their practices in the field thus provides an integrated explanation of these different products, linking social and scientific factors without conflating them. More specifically, the pervasive contrast in Wallace and Darwin's writings from the field, their differential treatment of individuals and groups, emerges from social and theoretical constraints on their respective practices. The contrast in their writings in turn throws light on the conceptual frameworks within which their theories of natural selection developed. In this paper, I aim to establish the contrast inWallace and Darwin's writings from the field, and explain it in terms of their differing circumstances and motivations. This sets up an integrative framework for examining their theories of natural selec? tion, and for clarifying the relation between individuals and groups in Wallace's own theory.

Circumstances and Motivations

Both Wallace and Darwin began their lifelong careers in natural history as amateur specimen collectors, but under very different circumstances. Three contrasts are significant here: training and social connections; finances and responsibilities; and theoretical interests. Darwin's

5 Camerini, 1996, 1997; Raby, 2001; Shermer, 2002; Fichman, 2004; Slotten, 2004. 6 Desmond and Moore, 1991; Browne, 1995, 2003; Keynes, 2003; Stott, 2003; Herbert, 2005.

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circumstances were fortunate. Born into a wealthy and well-connected family, he was provided with opportunities to train in natural history and to make social contacts with leading zoologists, botanists, and geologists of the day. He began work as a naturalist in Edinburgh, collecting and microscopically examining marine invertebrates, guided by the anatomist Robert Grant.7 Finding medical training uncongenial, Darwin moved to Cambridge, where his cousin William Darwin Fox introduced him to beetle-collecting and to John Henslow, the new Professor of Botany.8 In his final year, Darwin became acquainted with several other influential professors, notably Adam Sedgwick (Professor of ), who took him on a geological tour of Wales in the summer of 1831.9 These Cambridge connections and expeditions set the stage for Darwin's subsequent career. When Robert FitzRoy, Captain of H.M.S. Beagle, cast about for a gentleman companion and natural history sa? vant to accompany him on a voyage of mapping and circumnavigation, Henslow and his Cambridge circle recommended Darwin. Throughout the voyage (December 1831 to October 1836), Darwin occupied a privileged position as Captain FitzRoy's friend and com? panion. Since his wealthy father paid all expenses, including his keep on board, Darwin had no official duties or financial constraints. He was also well-supplied with equipment for collecting and storage, including a new Bancks .10 The Beagle's library was extensive, and his brother Erasmus sent Darwin whatever books he lacked.11 A number of crew members contributed to his natural history collection, and FitzRoy allowed him to make several extended trips inland.12 Darwin was thus at liberty to pursue his own research interests, and well-equipped to do so. He was appreciative of his good fortune, remarking in his first letter home during the Beagle voyage: "I verily believe no person ever went out better provided for collecting & observing in the different branches - of Natural hist. In a multitude of counsellors I certainly found

7 CCD 1, p. 538, 543 note 14. 8 CCD l,pp. 56,432. 9 Herbert, 2005, pp. 39-47. 10 The expense was considerable: nearly ?1,200 in drafts requested en route plus an initial outlay of ?600. In contrast, Wallace cleared 3? per sold; Stevens insured his Amazon collection at ?200 (Wallace, 1906, pp. 303-309; Slotten, 2004, pp. 42^3, 88 89). 11 CCD l,pp. 553-566. 12 CCD 1, pp. 540-542; Keynes, 1988; Darwin, 1987 [1839]. Summary in Browne, 1995, pp. 164-165, 226-228.

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good."13 The system that supported him also restricted his movements; Darwin was effectively a paying passenger on a government-sponsored voyage, with no say as to the expedition's route or schedule.14 His work in natural history was constrained by the Beagle's orders, but by little else. Well-equipped, well-trained and well-connected, Darwin was pre? pared to study "all branches of Natural History that I can possibly manage."15 Wallace's humble origins contrast sharply with Darwin's privileged background. His parents were originally middle class, but a series of bad business decisions caused the family finances to decline over the 1820s and 1830s.16 In 1837, when his parents could no longer support him, 14 a year-old Alfred was taken out of school to make living.17 In 1841, working as a land-surveyor in Wales, a business slump gave Wallace leisure for his first forays into natural history.18 He began by identifying orders of wildflowers with a shilling book on botanical classification, published by the Society for the Diffusion of Useful Knowledge.19 A few years later, working as a teacher in Leicester, Wallace met amateur entomologist (and apprentice in knitwear manufacturing) , who introduced him to beetle-collecting. After Wallace returned to Wales, the two corresponded about specimens and recent books. In late 1847, inspired by reading Vestiges of the Natural History of Creation (1844) and W. H. Edwards' A Voyage up the Amazon (1847), Wallace and Bates decided to journey to the Amazon to study "the origin of species," paying their way as professional specimen collectors.20 After stopping in London to acquire an agent and some hasty training in collection and preservation techniques, Wallace and Bates sailed for Brazil, arriving in Para on May 26, 1848.21 They collected as a team for a few months, then separated. Wallace spent most of the next fourteen years collecting specimens in the Amazon and the Malay Archipelago.

13 CCD 1, p. 202. 14 Browne, 1995, pp. 227-228. 15 Entry of December 13, 1831 (Keynes 1988, 14). See Gruber (1985) for more on Darwin's early theorizing. 16 Wallace, 1906, pp. 55-79. See references in note 5 for recent accounts of Wallace's early life. 17 Wallace, 1906, p. 191. 18 Moore, 1997. 19 Wallace, 1906, p. 191. See McOuat, 1996, 2001. 20 Wallace, 1906, pp. 254-255, 264; Wallace, 1969 [1853] p. xi. 21 Wallace, 1969 [1853] p. 1. For details of Wallace and Bates' training and equipment, see Slotten, 2004, pp. 42-45.

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During those years abroad, Wallace was self-supporting, largely isolated from English society, and had only sporadic contact with other naturalists. His access to books, periodicals and conversation was lim? ited and intermittent. Samuel Stevens, Wallace's London agent and treasurer of the Entomological Society of London, was his principal source of information and conduit to the London natural history source community.22 Stevens was also Wallace's sole of money and equipment, contingent upon satisfactory sale of specimens to " and amateurs."2^ Wallace thus worked under constant financial con? straint, concentrating on specimens that fetched good prices and plan? ning his movements according to anticipated profits or shortfalls from his collections.24 The "constant employment and ever-varying interest of a collector's life" was balanced by "craving for intellectual and congenial society."25 Wallace may have felt intellectually isolated, but he was hardly alone in the field.26 A diffuse "colonial network" provided some society in a few centers (such as Ternate and Malacca), and facilitated the vital assistance of native tribes.27 Though colonial network in South America had strong English ties, its East Indies counterpart was a dislocated and socially fluid mixture of travelers, miners, missionaries, doctors, and businessmen from many countries. Due to clearing of land around towns, Wallace often found it necessary to live outside European set? tlements while he collected. His day-to-day work thus depended more immediately on his assistants and tribal hunters: Malays, Papuans, Dyaks and men from other local tribes. With their support, provided his specimens sold well, Wallace could raise enough money to stay in the field. Apart from economic constraints, he was a free agent, at liberty to plan his own itinerary. In addition to these different socio-economic circumstances, Wallace and Darwin came to the field with different theoretical aims. Wallace's collecting career was initially motivated by his theoretical interest in the origin of species, apparently sparked by reading Vestiges. In December 1846, Wallace wrote to Bates that the transmutation theory proposed in that anonymous work "serves both as an incitement to the collection of

22 Camerini, 1996, pp. 62-64. Wallace also corresponded with Bates and botanist George Spruce. 23 George, 1979, pp. 503-514. 24 E.g., Wallace, 1859b, p. 129. 25 Wallace, 1856b, p. 5117. 26 See Camerini, 1996, 1997. 27 The term is borrowed from Browne, 1992.

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facts, and an object to which they can be applied when collected."28 A letter of September 1847 went further: "I begin 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 may be arrived at."29 In his 1863 memoir, Bates recalls this as the motivation for their trip:

Wallace...proposed to me a joint expedition to the river Amazon, for the purpose of exploring the Natural History of its banks; the plan being to make for ourselves a collection of objects, dispose of the duplicates in London to pay expenses, and gather facts, as Mr. Wallace expressed it in one of his letters, "toward solving the problem of the origin of species," a subject on which we had conversed and corresponded much together.30

Wallace's collecting was thus driven by both his scientific interests and his financial constraints. These dual motivations literally resulted in duplicate collections. As planned, Wallace made two collections on both expeditions: one sold to cover his expenses, the other set aside for his personal scientific use.31 There was never any question of duplicates for Darwin. A paying passenger on the Beagle, he owned all his collections outright and had no need to sell them.32 With no economic incentive to collect, and no official responsibility to do so, Darwin was free to indulge his own interests in natural history. Among these, geology took precedence. Darwin's Beagle correspondence contains many remarks to this effect: "I am seeing the country & collecting in every branch of Nat. History..." "But Geology carries the day;" "Since leaving Valparaiso, during this cruize [sic], I have done little excepting in Geology," pursuing it "even to the neglect of marine ."33 Unlike Wallace, Darwin began to consider the problem of the origin of species, and consider transmuta? tion as a possible answer, only after his collecting days had ended. He later dated his own first "vague doubts" about the "stability of species" to the end of the Beagle voyage, and began to keep species notebooks in

28 Wallace, 1906, p. 254. 29 Wallace, 1906, pp. 254-256. 30 Bates, 1863, p. i. 31 Wallace, 1869, p. vii; Wallace, 1906, p. 266, 306. 32 Browne, 1995, pp. 185, 206-209, 219, 230-232. 33 CCD 1, pp. 230, 232, 432, 436; see Herbert, 2005, pp. 1-47, for more on Darwin's geological background.

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July 1837. During the voyage, Darwin's primary theoretical interests were geological, as he enthusiastically applied 's theories in various localities. Wallace and Darwin thus came to the field with different educational experience and connections, constrained by different responsibilities and theoretical motivations. These biographically specific contrasts are embedded in the wider social and scientific context, which changed considerably in the twenty years separating their field expeditions. Geology, botany and zoology, the three traditional branches of natural history, began to fragment into distinct disciplines, as publications and societies proliferated in London and in provincial centers.35 The establishment of rules for zoological nomenclature (in which Darwin participated), and the rise in prominence of the British in the 1840s, focused Wallace's practice by allowing him to target gaps in a stable taxonomic framework.36 Yet the methods and equipment of natural history collecting underwent little change.37 Wallace and Dar? win's different natural history practices reflected their different loca? tions, framed by their personal circumstances and motivations, within this wider context.

Natural History Practice

Wallace and Darwin's natural history practices differed in intensity, in the principal activities performed, and in the standards they used. Wallace's practice was more intense than Darwin's in at least two re? spects: overall length of time in the field, and daily working routine. Most of Darwin's work in natural history took place in the first four years of the voyage (late 1831 to late 1835) while the Beagle mapped the South American coastline.38 Darwin collected specimens mainly along the coast of South America and adjacent islands, spending slightly less than half the voyage onboard. Though most of his trips ashore were of fairly short duration, Darwin made eight significant inland expeditions

34 Darwin and Seward, 1903, Volume 1, p. 367; de Beer, 1960; Barrett et al. 1987; Herbert, 1980. 35 Allen, 1976; Farber, 2000, pp. 33, 47. 36 Desmond and Moore, 1991;McOuat, 1996, 2001. 37 Larsen, 1996; Allen, 2001. 38 CCD I pp. 540-542; Keynes, 1988; Darwin, 1987 [1839]. Summary in Browne, 1995, pp. 164-165, 226-228.

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on horseback, and lived for a month or more at four different collecting sites (Botofogo Bay, Maldonado, Valparaiso, and Chilo?).39 Though Wallace also spent four years in South America (1848-1852), his route along the Rio Negro never crossed Darwin's earlier path.40 He stayed at each collecting site for several weeks, usually a month or more. Travel in the tropics was arduous. Permission to travel in certain areas had to be secured, local assistants and porters abandoned expeditions without warning, the weather was frequently uncooperative, illness and injury were constant threats. In his eight years in the Malay Archipelago (1854-1862), Wallace undertook more than "eighty movements [of house] averaging one a month," criss-crossing back and forth among the islands, and frequently revisiting collecting sites.41 He later calcu? lated that "as I travelled about fourteen thousand miles within the Archipelago, and made sixty or seventy separate journeys, each involving some preparation and loss of time, I do not think that more than six years were really occupied in collecting."42 As well as being of longer duration, Wallace's collecting practice was more intense than Darwin's on a daily basis. Wallace's schedule in the field was rigorous:

Get up at half-past five, bath, and coffee. Sit down to arrange and put away my of the day before, and set them in a safe place to dry. Charles [Wallace's sometime assistant] mends our insect nets, fills our pin-cushions, and gets ready for th? day. Breakfast at eight; out to the jungle at nine. We have to walk about a quarter mile up a steep hill to reach it, and arrive dripping with perspira? tion. Then we wander about in the delightful shade along paths made by the Chinese wood-cutters till two or three in the after? noon, generally returning with fifty or sixty beetles, some very rare or beautiful, and perhaps a few butterflies. Change clothes and sit down to kill and pin insects, Charles doing the flies, wasps, and bugs; I do not trust him yet with beetles. Dinner at four, then at work again until six: coffee. Then read or talk, or if insects very numerous, work again till eight or nine. Then to bed.4"'

39 Ibid. 40 Wallace, 1969 [1853]. 41 Marchant, 1916, p. 68; Wallace, 1869. 42 Wallace, 1869, pp. vii-viii. 43 Letter from Singapore (written May 28, 1854). Wallace, 1906, pp. 337-338. A longer but essentially similar description is included inWallace, 1855a, pp. 4805-4807. Reports and letters from his Amazon travels indicate that Wallace's schedule there was essentially the same (Shermer 2002, pp. 60-61; Slotten 2004, p. 60).

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The entire routine was organized around collecting: capture of speci? mens in the morning and early afternoon, and preparation afterwards. A typical workday was about 12 h long, "with hardly half an hour's intermission, from 6 am till 6 pm, four or five of the hottest hours being spent entirely out of doors."44 Once Wallace's daily hunt with gun and net was finished, the second half of his workday began. Dozens of specimens had to be prepared and arranged: insects killed and pinned; birds, reptiles, and skinned and dissected, fur or feathers as or treated with or arsenic soap.4~ All had to be identified known new species, with the help of a few annotated classification texts.46 Then came labeling, taking notes, and protecting drying specimens from the ravages of insects. All this "fill[ed] up the whole time of one person, with two or three native assistants".47 Yet Wallace always sought to do more, observing with dissatisfaction that "a travelling collector of limited means like myself does so much less than might be expected, or than he would himself wish to do".48 Darwin's typical workday, on land or at sea, was more relaxed. Often seasick, he limited natural history work on the Beagle to calm days. The schedule was set with naval rigidity: - We breakfast at eight oclock. The invariable maxim is...bolt off the minute one has done eating, &c. At sea, when the weather is calm, I work at marine animals, with which the whole ocean - - abounds. if there is any sea up. I am either sick or contrive to - read some voyage or Travels. At one we dine....[comments on food and drink.] ...At 5 we have tea.49

Though Darwin's "work at marine animals" was more substantial than he suggests here (see below), it was not typically an all-day affair. Nor did he regularly work "with hardly half an hour's intermission" on? shore, though he did collect intensely during the Beagle's short visits to St. Jago (Sao Tiago), Bahia Blanca, Tierra del Fuego, and the Gal?? pagos Islands. When the Beagle's itinerary allowed him to spend a few weeks or months on land at a stretch, Darwin pursued his "Usual quiet 44 Wallace, 1859a, pp. 111-113. 45 Wallace, 1906, pp. 328-330. See also Wallace (1855a); Slotten (2004). For details of naturalists' equipment in the 1840s to 1860s see Larsen (1996); for an overview F?rber (2000). 46 Wallace's principal sources were Boisduval's Histoire Naturelle des Insectes (1836) and Bonaparte's Conspectus Generum Avium (1850); Wallace, 1906, pp. 327-329. 47 MM, .18 (July-August 1856). 48 Ibid. 49 - CCD 1, p. 248 (Letter to Susan Darwin, 14 July 7 August 1832).

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occupations; one days collecting & the next arranging."50 He alternated days of collecting with days of describing and preserving his terrestrial specimens, rather than doing both each day. Collecting and processing sur? specimens at this relaxed pace gave Darwin time to appreciate his roundings: "Commonly I ride some few miles, put my horse & start by - some track into the impenetrable mass of vegetation. Whilst seated on a tree, & eating my luncheon in the sublime solitude of the forest, the no to at pleasure I experience is unspeakable."51 With need work Wallace's grueling pace, Darwin's natural history practice was less intensive, overall and on a daily basis. Wallace and Darwin's work in the field also consisted of different principal activities. They collected, arranged, and described different objects, using different methods. Wallace's typical quarry were insects (beetles or butterflies), which he hunted with nets. His field equipment consisted of a net, collecting box, forceps and bottles of arrack for or preservation.52 More rarely, he shot trapped birds, reptiles and mammals, and gathered land-snail shells. In South America, Wallace also collected fish and plants, and kept a small live menagerie.53 During his last 4 years in the Malay Archipelago, Wallace sought to acquire specimens of all the species of Paradiseae (Birds of Paradise). To this end, he made five different voyages, each of which took the better part of a year, to the Aru Islands or New Guinea/4 He thus collected in several branches of the solidifying discipline of zoology, primarily in and . Unlike Darwin, Wallace did not collect geological specimens. Nor, despite admiration for Darwin's Journal of Researches, did he emulate that work's extensive geological discus? sions.55 Botany was another matter. Wallace had a life-long interest in "botanizing." His reasons for focusing on zoology in the field were financial, as stated bluntly in a letter to his childhood friend George Silk: "I cannot afford to collect plants. I have to work for a living, and plants would not pay unless I collect nothing else, which I cannot do, being too much interested in zoology."56 Throughout his time in the

50 Keynes, 1988, pp. 66, 160. 51 CCD 1, p. 247 (Letter to Catherine, 5 July 1832, Rio de Janeiro). 52 Wallace, 1855a, p. 4805; MJb (March 1858). Wallace used forceps, two gauze panels joined by a hinge, to capture bees, wasps and moths. Further details in Larsen 1996, pp. 373-375. 53 Wallace, 1852, pp. 3641-3643; Wallace, 1969 [1853], 1906, pp. 303-309. 54 Wallace, 1862, 1869, p. 575. See also Camerini, 2002, pp. 103-141. 55 Marchant, 1916, p. 21. 56 Wallace, 1906, pp. 191, 411-412.

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field, Wallace tailored his collecting activities according to what would sell, concentrating on groups that fetched good prices: tropical birds, butterflies, and beetles. However, economic pressures did not com? pletely determine Wallace's collecting practice. His theoretical interest in characterizing the species in a given area, their relative abundance, and relations between the species composition at different localities, remained strong throughout his collecting career. In letters to the Entomological Society of London, for example, Wallace complains that amateur naturalists neglect "small and obscure...groups" in favor of "large and handsome" species, obscuring "a true idea of the Ento? mology of this country."57 Wallace practiced what he preached: at each collecting site, he sought as many different species as could be found, showy or drab, new or familiar. He often revisited sites, returning at different seasons to get a more complete sampling, and recorded precise details of his collecting locales. Darwin's principal collecting activities were quite different. For one thing, his geological interests often trumped his zoology and botany.58 This is evident from the relative frequency and length of geological (compared to zoological) discussion in Darwin's diary, notes and cor? respondence between 1831 and 1836, as well as from many of his own remarks.59 Accordingly, much of his time onshore was occupied with geological investigations rather than zoological and botanical collecting. Darwin's geologizing often involved covering large areas of terrain on horseback; such trips did not afford opportunities for extensive zoo? logical collecting. Bones and seeds might have withstood the shocks of hard riding, but most and specimens would have been dust in the saddlebags, even if there had been time to collect them. Since these activities did not mesh well, Darwin's focus on geology meant less zoological and botanical work. After geologizing, Darwin's main natural history interest during the Beagle voyage was marine zoology. His maritime collecting method was one of the first recorded uses of a plankton net: "a bag four feet deep, made of bunting, & attached to [a] semicircular bow this by lines is kept - upright, & dragged behind the vessel. this evening it brought up a 57 Wallace, 1856b, pp. 5113-5117; Wallace, 1858b, pp. 6120-6124; see also MJc (16 July 1858). 58 See Herbert, 2005, especially pp. 98-128. 59 CCD 1, pp. 232, 418, 432, 495. Darwin's geology notes were four times longer than those for zoology (Keynes, 2000, p. ix). Well over half of the latter concern marine invertebrates; descriptions of single specimens run to 5-6 pages. Vertebrates typically get only a few lines. See Herbert (2005, pp. 140-176) for Darwin's geological writings during the Beagle voyage.

This content downloaded from 150.135.114.31 on Sun, 2 Jun 2013 13:59:18 PM All use subject to JSTOR Terms and Conditions 614 MELINDA B. FAGAN mass of small animals, & tomorrow I look forward to a greater har? vest."60 Darwin examined his harvests of marine invertebrates under his microscope, making superbly detailed observations in his zoology notebook.61 Eager to describe new7 species, he also attempted to identify each specimen using the considerable resources of the Beagle's library. Onshore, where one could not simply wait for one's net to fill, Darwin still spent comparatively little time hunting for specimens as opposed to making detailed observations: "I find one hours collecting keeps me in full employment for the rest of the day."62 In contrast, Wallace's "extensive collections of birds & insects" demanded "constant personal attention", leaving him little time to observe individual specimens in detail.63 Darwin also relied on others to do his collecting to a much greater extent than Wallace. Though Wallace worked mostly alone in South America, he employed regular assistants in the Archipelago: Charles Allen, the son of an English carpenter, and Ali, a young Malay hired in 1855, who remained with Wallace for the remainder of his travels.64 Both young men helped Wallace shoot and prepare specimens, though Ali was apparently much more competent. Wallace also intermittently engaged one or two native hunters and guides for short periods in particular localities. He sometimes paid local hunters for specimens, notably Paradise birds, which were caught alive using specialized methods. But when his health permitted, Wallace worked alongside his assistants, adding their labors to his own. He did not use assistants to pursue different activities.65 Darwin did employ assistants to collect while he geologized, or microscopically examined his marine harvests. In July 1833, while living in Maldonado, he hired Syms Covington, a Beagle crew member, to assist him in ornithology. Regarding their arrangement, Darwin wrote frankly to Fox: "You ask me about Ornithology; my labours in it are - very simple. I have taught, my servant to shoot & skin birds, & I give

60 Keynes, 1988, p. 21. 61 For further detail, see Sloan (1985, pp. 87-103). 62 Keynes, 1988, p. 64. 63 Malay Diary I, 35 (1 Oct 1856). 64 Wallace, 1906, pp. 338-340, 382-383; Camerini, 1996, 1997. Other assistants, such as Baderoon, a young man of Macassar, worked for Wallace for shorter periods than Charles and Ali. 65 See, for example, MJa, 35 (1 Oct 1856).

This content downloaded from 150.135.114.31 on Sun, 2 Jun 2013 13:59:18 PM All use subject to JSTOR Terms and Conditions WALLACE. DARWIN, AND THE PRACTICE OF NATURAL HISTORY 615 him money. I collect reptiles, small quadrupeds, & fishes industriously. The invertebrate marine animals, are however my delight..."66 Darwin was also willing and able to pay for specimens, epitomizing his policy as "if gold or galloping will get them, they will be mine."67 Darwin's plankton net and social networks, as well as his wealth and position, enabled him to pursue his interests in geology and to make detailed microscopic observations of particular specimens. Wallace, in contrast, spent more of his time actively hunting for specimens in the jungles and fields, and less on detailed observations. He was keenly aware of this tradeoff, noting in his personal journal that

to make any thing like extensive collections of birds & insects, keeping brief notes of the most interesting facts connected with them will fill up the whole time of one person, with two or three native assistants. He absolutely cannot do much else, and is often even obliged to abridge his notes in order to secure the safe pres? ervation of his specimens.68

Finally, Wallace and Darwin applied different standards to their col? lecting practices. Darwin, with his preference for painstaking micro? scopic observation, aimed to thoroughly describe a few specimens of a species rather than collect a whole "series." A letter to Henslow makes this explicit: "Most assuredly I might collect a far greater number of specimens of Invertebrate animals if I took less time over each: But I have come to the conclusion, that 2 animals with their original colour & shape noted down, will be more valuable to Naturalists than 6 with only dates & place."69 Ever meticulous, Darwin took pains to record any features likely to degrade after preservation or drying, and dissected duplicates when possible. Representing a species with "2 animals," however meticulously de? scribed, was not Wallace's way. One reason for this was his concen? tration on insects and birds, groups far better-characterized in the 1850s than marine invertebrates in the 1830s. But Wallace's concern for series of specimens was not merely due to passive uptake of the natural history 66 CCD 1, p. 316 (Letter to Fox: 23 May 1833, Maldonado). Also CCD 1, pp. 312, 314, 321. 67 CCD 1, p. 398. See also Keynes, 1988, p. 160. 68 MM, 18 (July-August 1856). 69 CCD 1, p. 251; also Keynes, 1988, p. 207. At the time Darwin believed that most variation in animals and plants occurred between species and varieties, not within them; he thus may have seen no benefit to representing species with "series" (Darwin 1987 [1844]; Stott, 2003, pp. 146, 241-242). Moreover, as marine invertebrate was in an unsettled state, characterizing new animals was a sensible and productive strategy.

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Standards of his day. Rather, the combination of Wallace's theoretical and economic interests led him to collect whole series of specimens for particular species, from his first expeditions on the Rio Negro in the 1840s, to his hunt for Paradise birds in the Aru Islands over a decade later.70 The financial incentives to collect multiple specimens of partic? ularly salable species, such as showy Lepidoptera and Paradise birds, are obvious. But Wallace's theoretical interests, specifically his distinction of true species from "mere varieties" on the basis of constancy rather than degree of differentiation, also demanded multiple specimens of or each species.71 To determine what species he had, either for sale theoretical speculation, Wallace needed to examine multiple individuals. His field reports from 1856 onwards indicate that Wallace aimed to collect both male and female specimens of different stages of maturity.72 For some species, such as Paradise birds with ontogenetically variable plumage, a "good series" required over twenty specimens; while for many Cole?ptera he found three sufficient.73 Wallace and Darwin's natural history practices in the field thus differed in three significant ways. First, they worked at different inten? sities, both overall and on a daily basis. Second, their principal routine activities were different. Wallace spent 5 or 6 h each day hunting for specimens, and an equal amount of time processing and classifying his haul. Darwin spent much less time actively searching for specimens, allowing them to come to him via net or networking, and much more time minutely describing individual specimens, with special attention to marine invertebrates. Finally, they used different standards for collect? ing. Darwin aimed to meticulously describe one or two specimens per species and was most excited by novelty; while Wallace sought "a good series" of specimens to represent each species and a complete inventory of species at a given locality. These different practices were in turn shaped by the circumstances and motivations that brought Wallace and Darwin to the field: economic, social, and scientific.

Material Results

Their different practices, unsurprisingly, yielded different material results. Darwin's zoological and botanical collection from his Beagle

70 Wallace, 1850b, pp. 494-496. 71 Wallace, 1860, p. 107. 72 Wallace, 1850b, pp. 494-495; 1856c, 1857c, p. 415; 1861. 73 Wallace, 1869, p. 539.

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voyage was well-received by the natural history community of the late career. 1830s, and provided a solid basis for his subsequent illustrious His specimen lists from the Beagle show the following totals: 1509 specimens "in spirits of wine," and 3921 specimens "not in spirits."74 The total number of specimens listed is thus 5430. However, it is somewhat difficult to infer numbers of either species or individual organisms collected from these totals, as some specimen numbers refer to jars containing multiple species, multiple specimens of a single spe? cies, or parts of a single individual, separated after dissection.75 Such ambiguity is characteristic of Darwin's descriptions of animals and plants during the voyage, and contrasts with Wallace's crisp distinctions (see below). Ambiguities notwithstanding, the number and quality of his specimens was sufficient to launch Darwin as a respected natural his? torian, as his old mentors and new colleagues praised his work, and professional zoologists classified specimens of birds, insects, reptiles, fish, mammals and fossils.76 The extent of Wallace's material collection is staggering: over 125,000 specimens collected in the Malay Archipelago alone. His Amazon totals are much lower: roughly 10,000 specimens, mostly butterflies and beetles. This was not due to inexperience or lack of effort. Most of Wallace's South American collection, along with his journals, drawings and notes, was lost on the return voyage to England, when his ship caught fire and sank ten days into the voyage.77 Wallace's second expedition was more productive, yielding a stupendous total of 125,660 specimens, over 1000 of which represented entirely new species: 310 specimens, 100 reptiles, 8050 birds, 7500 shells, 13,100 Lepi doptera, 83,200 Cole?ptera, and 13,400 other insects.78 Wallace later

74 Keynes, 2000. Darwin's geological notebooks record 3913 specimens (Herbert, 2005, p. 99). 75 Desmond and Moore report Darwin's collection as consisting of 1529 species in wine spirits and 3907 dried specimens (Desmond and Moore, 1991, p. 129). But these as some specimen numbers include some redundant species, labeled "same (n)," and jars a of spirits are listed as containing multiple species. There is not one-to-one relationship between species and Darwin's specimen numbers. The Journal of Researches contains an advice on labeling and shipping specimens; however, these passages describe ideal, with the advantage of hindsight, rather than Darwin's actual practice during the voyage (Darwin, 1987 [1839], pp. 598-602). 76 CCD 1, pp. 512-514 (Letter to Henslow, 30-31 Oct 1836); Browne, 1995, pp. 348 on 354, 358-361; Keynes, 2000, pp. ix, xi. Most of Darwin's painstaking work marine a invertebrates remained unpublished; he took up barnacle systematics himself decade later (Love, 2002; Stott, 2003). 77 Wallace, 1852, pp. 3641-3643; 1906, pp. 269-270, 303-315. 78 Wallace, 1869, p. viii; Wallace 1906, pp. 362-363.

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estimated his private collection at approximately 3000 skins of about 1000 species, and approximately 20,000 beetles and butterflies of about 7000 species. Unlike Darwin, he appears to have organized his collection by numbering species, as well as keeping track of the number of specimens of each.79

Writings from the Field

Wallace and Darwin's natural history practices also yielded writings, though these were not (of course) produced in the same way as the collections described above. The relevant writings were produced con comitantly with those material collections, and concern the same ob? jects: living things, their habits and habitats. These writings result from natural history practice, in the sense that differences in the latter made a difference to the former. In what follows, I draw on Wallace's published early writings, including portions of his correspondence, as well as his personal journal from the Malay Archipelago (1856-1861), zoology notes (1855-58), and one theoretical notebook (1855-59).80 Comparable sources for Darwin include his Beagle diary, field notebooks, zoological and geological notes, catalogue of specimens, and his correspondence with family and friends. These various writings exhibit the contrast introduced above. Wallace consistently emphasized groups of organ? isms, while Darwin described many details of individual organisms. Also, Wallace clearly distinguished between groups and individuals, while Darwin was more ambiguous. Five lines of evidence show the pervasiveness of this contrast, and reveal the connections with the practices discussed above. First, and most coarsely, are the relative frequencies with which Wallace and Darwin mention groups vs. individual organisms in their writings from the field. Of 96 published papers and letter extracts written by Wallace between 1848 and 1862, twelve do not mention natural history (ten letter extracts, and two reports to the Royal Geo? graphic Society).81 The remaining 84 pieces can be subdivided as fol? lows: collecting reports (25), private correspondence (23), short articles

79 Wallace, 1850b, pp. 494-495; Zoology Notebook 1855-58. 80 See 'Manuscript Sources' below. Most of Wallace's Amazon writings were lost in the fire on the Helen. 81 Complete dataset available by request from author. See "The Alfred R?ssel Wal? lace Page" (Charles Smith) at http://www.wku.edu/~smithch/home.htm for a compre? hensive bibliography of Wallace's published writings.

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- (20), and longer articles regional, classificatory, or theoretical (16). All but one letter extract mention species or higher taxa, while only about half the pieces in each category mention individual organisms. When both are mentioned, groups (species, populations, higher taxa) are discussed at greater length and given greater prominence than individ? uals (organisms, specimens). Wallace's Malay diary exhibits the same pattern: of 78 separate discussions of natural history, over two-thirds focus on groups (species and higher taxa); only 4 are exclusively con? cerned with individual organisms. In contrast, about one-third of Darwin's discussions of natural history in his Beagle diary are exclu? sively concerned with individual organisms, while roughly three-fourths mention individual organisms, and about two-thirds mention species or higher taxa.82 The same pattern is observed in his Beagle correspon? dence. So, at this coarse level of analysis, Wallace's writing from the field emphasized groups over individual organisms, while Darwin's gave them roughly equal emphasis, with individuals predominating slightly. Second, in his correspondence Wallace consistently described his own activities in terms of working on species, while Darwin reported being "occupied by new & most interesting animals."83 For example, in a letter to Bates, Wallace described his work as connecting the affinities and geographical distribution "of a group, worked out species by spe? cies," and elsewhere asserted that " must be studied in detail, and it is the wonderful variety of the species of a group, their complicated relations and their endless modification of form, size, and colours, which constitute the pre-eminent charm of the entomologist's study."84 Darwin did not describe his activities in terms of species or other groups, but often wrote of "collecting & observing the numerous small animals in the sea" and his delight in observing "curious, & ...quite undescribed" creatures."85 Though it is difficult to determine whether such remarks refer to individual organisms or to groups, Darwin's focus was plainly not on species, but on new and interesting animals observed under his microscope. Third, Wallace's writings from the field are suffused with "species counts": tallies of the number of species in particular families or orders (usually insects or birds) collected at a given locality. These appear not only in Wallace's papers, zoology notes and reports to his agent, but also in correspondence with family and friends. For example, he

82 Details available by request from author. 83 CCD 1, p. 206. 84 Wallace, 1906, p. 358; Marchant, 1916, p. 56. 85 Keynes, 1988, p. 22; CCD 1, pp. 370; also CCD 1, pp. 278, 391.

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summed up six days' work in the K? Islands in terms of species "taken" in that time ("thirteen bird species, 194 insect species") and subdivided his Aru insect collection (1364 species) by family.86 When actual num? bers were not given, Wallace nearly always included a qualitative assessment of the species richness of his current site, often remarking on the relative abundance or scarcity of "fresh species," "new species," "handsome species," or "common species."87 Species counts are not prominent in Darwin's Beagle diary and correspondence.88 Instead, Darwin often recorded the numbers of individual animals killed or sighted: "we saw 40 or 50 ...small pretty Foxes"; "two monstrous whales were swimming within stone throw of the shore;" "this young man had shot 2 large bearded monkeys & had left another dead in the tree"; "the soldiers returned...[with] seven deer, 3 ostriches, and 40 of their eggs, many partridges & Armadilloes"; "several flocks of between 20 & 30 [Ostriches]"; and so on.89 Such remarks, tallies of individual animals seen or taken, appear over and over in Darwin's vignettes from the field. Fourth, Wallace and Darwin described themselves as "meeting" species and individuals, respectively. Wallace frequently referred to meeting species or higher taxa exhibiting characteristic behaviors or habits. For example, from Sarawak, Borneo (1854):

Imagine my delight in again meeting with many of my Singapore - friends, beautiful longicorns of the genera Astathes, Glenea and Clytus, the elegant Anthribidae, the pretty little Pericallus and Colliuris, and many other interesting insects. But my pleasure was increased as I daily got numbers of species, and many genera which I had not met with before.90

Wallace also attributed behaviors and habits to species rather than to individual organisms, remarking, for example: "For six weeks I have

86 Wallace, 1857d, p. 484. Similar counts occur frequently in Wallace's Malay diary and zoology notes. 87 Wallace, 1850b, pp. 494-496; 1906, p. 269. 88 The only example I know of is a letter from Maldonado, Brazil, where Darwin lived for several months: "We have got, almost every bird in this neighborhead [sic], about 80 in number and nearly 20 quadrupeds" (CCD 1, p. 321). 89 Keynes, 1988, pp. 187, 139, 71, 188, 237, 156. 90 Wallace, 1855a, pp. 4803-4807; also Wallace, 1854c, pp. 4394-4397; MJa (1 Jan 1857).

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almost daily seen Papilio Ulysses? or a new closely-allied species, but never a chance of him; he flies high and strong, only swooping down now and then, and off again to the treetops....I fear I shall never get him."91 An 1857 article on the Great Bird of Paradise begins: "Having enjoyed the rare privilege of a personal acquaintance with this remarkable bird in its native haunts, during my residence in the Aru Islands, I am enabled to give a more complete account of its habits..."92 Wallace then describes the mating displays, abundance, distribution and opulent plumage of this "most beautiful of all the beautiful winged forms which adorn the earth," all without mention of individual birds. Similarly, orang-utan traits and behaviors are attributed to the genus Mias rather than to individual orang-utans, though Wallace had extensive contact with the latter.93 Wallace's writings from Borneo (where he stayed for over a year, from late 1854 to early 1856) do include an exceptional case. In May 1855, Wallace adopted an infant orang after shooting and killing its mother. In several published letters (one anonymous) and in his 1869 memoir, Wallace described its behavior and facial expressions, attrib? uted emotions to it, and described his care for the "abnormal infant" in detail, repeatedly drawing parallels with human babies.94 Despite his efforts, the infant died after about three months, presumably of illness brought on by malnutrition. His writings concerning it are exceptions to Wallace's general tendency to emphasize groups rather than individuals in his writings from the field. However, even in this unusual case, Wallace's emphasis on species is evident. In his articles and letters to friends, colleagues, and family, Wallace persistently referred to the in? fant orang either as a "kind of baby," a "species of baby," "the mias" or "it." His popular article (in Chambers' Journal) is titled "A New Kind of Baby," and begins: "Not a newly born infant, but a really new baby, or, to speak as a naturalist, a new species of baby" (1856e, p. 325).95 Moreover, fondness for his "little pet" did not prevent Wal? lace from preserving its skin and skeleton as part of his orang-utan

91 Wallace, 1857b, p. 92. See also Wallace, 1854b, pp. 3884-3885; 1854c, pp. 4395 4397; 1857d, pp. 473-485; Malay Journal (e.g., entries from Oct 1856, Jan 1857, March 1857, Sept 1857). 92 Wallace, 1857c, p. 411. 93 Wallace, 1856c, d; 1869, pp. 51-53, 57-74. 94 Wallace, 1856a, e; 1869, pp. 53-57; 1906, pp. 343-345. 95 1856e, p. 325.

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collection, and one article concludes with an impersonal summary of "dimensions of young orang-utan."96 Thus Wallace's tendency to emphasize species appears even in this exceptional case. Darwin did not typically describe himself as "meeting" species, nor do his writings include lengthy discussions of the habits and behaviors of species as such. Instead, he vividly described meetings with individual animals and plants. General observations of natural history, such as those of Gal?pagos tortoises and lizards, are liberally sprinkled with individual encounters: "one I saw killed," "one large tortoise, which I watched," "one [egg] which Imeasured," "I have seen a large one which weighed twenty pounds," "I carried one to a deep pool...and threw it in several times" "I watched one for a long time...and pulled it by the tail," and many others.97 Thus, Wallace and Darwin tended to write about encounters with and habits of groups and individual organisms, respectively. Finally, Wallace and Darwin treated their respective 'de-emphasized entities' differently. Wallace wrote of individual specimens mainly to complain about having only one representative of a species, as in this typical letter to Stevens: "Here [Macassar] in two months I have got fifteen species...Of these six are represented by single specimens only, but of the rest I send you thirty fine specimens, and they will, I doubt not, contain something new."98 Since Wallace aimed to represent each species with a "good series" of specimens, a single specimen was not enough. He also occasionally described how particular individuals (most often orang-utans, unusual butterflies or Paradise birds) were or caught shot, thereby becoming his specimens.99 Wallace also wrote of individual animals when describing new or controversial observations about the habits of a given species or genus. These were based on his extensive observations of animal behavior in the wild, or on the many rough dissections he performed in the course of specimen preparation. For example, in several ornithological articles Wallace connected the fat, tough skin, and stomach contents of "a freshly killed bird," to the beautiful features of its species, which are the main emphasis.100 Though

96 Wallace 1869, p. 57; 1856a, p. 390. 97 Darwin, 1987 [1839], pp. 464-476, also pp. 126-127; Keynes, 1988, p. 272. Darwin's vivid and personal style has been linked to Romantic influences on his work (e.g., Herbert, 2005, pp. 131-135). 98 Wallace, 1857a, pp. 5652-5657. 99 E.g., Wallace, 1869, pp. 552-572, 572-575. 100 Wallace, 1861, pp. 288-290. See also Wallace, 1854a, 1854b, 1856c, 1856d, 1857b, 1860, 1861.

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the "beautiful winged forms" are highlighted, individual birds were the unlovely sources of Wallace's enthusiastic general observations. Simi? larly, his examination of "seventeen freshly killed Orangs" in Borneo served as the basis for lengthy descriptions of the appearance and habits as as of different orang species, well classification proposals.101 Individual organisms appear in Wallace's writing from the field in another way as well. Nearly all his communiqu?s mention the relative abundance of species of interest. For Wallace, species abundance and rarity were key operational terms: his own relative estimates, based on repeated sampling, of the number of individual members of a given species present at a collecting site. His practice of daily collecting at a single locality, usually for a month's time, yielded estimates of abun? dance that Wallace used to characterize sites and relate them to one another. For example, in Batchian "you may see hundreds of the common species [of Paradiseidce] to perhaps one of either of the rarer sorts;" while at Santarem "many common insects, such as Heliconia Melpomone and Agraulis Dido, [are] abundant, which we hardly ever saw at Para."102 Individual organisms thus appear obliquely in his writings from the field. This makes sense in the context of Wallace's practice. For 12 years, for 5 or 6 h a day, at nearly a hundred different sites, Wallace spent more time with birds and beetles than he did with other humans. For him, individual organisms were not irrelevant or absent, but simply too obvious to mention. They provided the medium for his work and suf? fused his daily routine. Certainly Wallace did not confuse individuals and groups, writing to Bates: "The individual abundance of beetles is not, however, so large as the number of species would indicate. I hardly collect on an average more than fifty beetles a day, in which number there will be from thirty to forty species."103 Nor did Wallace conflate higher taxa. His longer communiqu?s from the field include separate discussions of each taxonomic level, beginning with the most inclusive groups (birds, insects, shells), and proceeds to list and describe orders, genera, species, and (last of all) individual specimens.104 Wallace thus uses the concept of abundance to consistently distinguish between

101 Wallace, 1856c; 1856d. 102 Wallace 1859b, p. 113; Wallace, 1850a, pp. 156-157; also Wallace, 1849, pp. 74-75; Wallace, 1857d, p. 479, 484; Wallace, 1858b, p. 6124; Wallace, 1861, p. 285; Malay diary (entries of Jan 1857, Sept 1857) 103 Wallace, 1906, p. 352. 104 E.g., Wallace, 1857a, pp. 5652-5657; Wallace, 1906, p. 358; Marchant, 1916, pp. 53-55.

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species and individual organisms, species and genera, and so on. His frequent references to species abundance or rarity, coupled with his terse but significant remarks about individual organisms as the basis for his collection, indicate that individual organisms are not irrelevant or ab? sent in his work, but form the background for his writings. The relation between individual and species is complex, and traversed with much effort on Wallace's part. In contrast to Wallace's consistent (though often oblique) distinc? tions between species and individual specimens, Darwin's writings are strikingly ambiguous as to level of description. His discussions of nat? ural history shift rapidly between taxonomic levels, mingling general descriptions with engaging anecdotes about individual organisms, as in this diary entry:

In ascending the bare summit, I came close to two Guanaco & in - the course of my walk saw several more. These beautiful animals are truly alpine in their habits, & in their wildness well become the - surrounding landscape. I cannot imagine anything more graceful than their action: they start on a canter & when passing through - rough ground they dash at it like a thorough bred hunter. The noise they make is very peculiar & somewhat resembles the neig? hing of a colt.105

In addition, many of Darwin's descriptions of species are straightfor? ward generalizations from his descriptions of single specimens.106 Such ambiguity would come naturally to a meticulous observer of one or two specimens per species. The careful details noted in particular cases be? come the raw material for useful generalizations, which inform Dar? win's natural history practice and his writings as well. The most vivid examples are the familiar names Darwin gives to particularly significant species: a diamond beetle is "our old friend Crux Major," his most prized fossils are "Master Megatherium & Mastodon," and a small orange-colored barnacle, first called an "informed little monster," is later rechristened "Mr Arthrobalanus."107 However, Darwin did take an interest in species, particularly new ones. For example, in early 1834 Darwin acquired a specimen of a new species of ostrich in Patagonia, eventually named Rhea darwinii.l0H This

105 Keynes, 1988, pp. 126-127. Guanaco are the wild counterparts of llamas. 106 E.g., CCD 1, p. 236 (Keynes, 2000, pp. 14-16); CCD 1, p. 307 (Keynes, 2000, pp. 116-118). 107 CCD 1, pp. 232-233, 405; Keynes, 2000, pp. 305-307; CCD 3, pp. 305-308. 108 Keynes, 2000, p. 188; 2003, p. 186.

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smaller ostrich species overlapped in range with the larger, more com? mon Rhea americana, and had been described only by rumor before Darwin secured what became the type-specimen. Darwin was aware of the potential significance of his find, writing to Henslow "...what is of more general interest [than Darwin's observations of marine inverte? brates, described first] is the unquestionable (as it appears to me) exis? tence of another species of ostrich, besides the Struthio Rhea..."109 Yet in his diary and Zoology notes, Darwin discussed ostriches (of either species) mainly in terms of individuals sighted or killed, shifting rapidly between references to species and to individual organisms.110 This is in keeping with his tendency to sprinkle general descriptions with vivid anecdotes about individual organisms. One of these concerns Rhea darwinii itself: the future type specimen was shot by the Beagle's artist, Conrad Martens, and eaten for dinner. Fortunately, during the meal Darwin realized that the bird was a new species, and salvaged the head, neck, legs, wings, some large feathers, and most of the skin. These leftovers became the type specimen! To recapitulate: the pervasive contrast in writings from the field is supported by five lines of evidence. Wallace and Darwin's writings on natural history differ with respect to individual organisms and groups in frequency of mention, in self-descriptions of their work, in what they (literally) count and meet with in the field, and in how they draw dis? tinctions relevant to theorizing about natural history. The contrast is explained by Wallace and Darwin's different practices in the field. Wallace's emphasis on species and groups is understandable given his doubly motivated natural history practice: twelve years of collecting and arranging specimens, 12 h a day, aimed at obtaining a complete inventory of different species and their distributions at many localities. His clear distinctions between individual organisms and groups are also understandable in terms of his practice. Wallace's theoretical and financial ends demanded that species be represented by a "good series," which required multiple individuals, sometimes more than twenty. Wallace thus used populations of specimens to represent species, not one or two individuals, as Darwin did. Furthermore, the process of transforming an individual into a representative of its species was laborious. The individuals Wallace caught or shot provided raw material for his collection, w7hile species and other groups were in a sense the 'output' of his own work of cleaning, sorting, and classifying.

109 CCD 1, p. 370 (Letter to Henslow, March 1834, E. Falkland Islands). 110 Keynes, 1988, p. 156. See also Keynes, 2000, pp. 100-102, 188-190; 1988, pp. 104 105; CCD 1: 276; Darwin, 1987 [1839], pp. 105-110.

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The man who spent hours each day transforming individual organisms into representatives of groups would be the last person to conflate the two, or to minimize the distinction between them. Species and higher taxa were what Wallace worked towards, and receive more attention in his writings, but individual organisms are always, and necessarily, in the background. Darwin, lacking Wallace's motives, developed a different routine of natural history practice. His emphasis on individuals emerged from his routine practice, which involved relatively little time actively hunting for species, and more time carefully examining the minute features of specimens brought to him via net or network. He worked at a more sedate pace, lavished attention on each specimen he collected, and meticulously described their features. The distinctive features of his writing dovetail with these aspects of his practice. His habit of extrap? olating from one or two individuals to the characteristics of entire species is reflected in his tendency to mingle descriptions of individuals and groups. Detailed observations of each individual specimen, readily extrapolated to the species, result in descriptions rich in individual de? tail, in which the line between specimen and species tends to blur.

Conclusion: Wallace's theory

The contrast in Wallace and Darwin's writing, and the explanation in terms of their practice, is of interest in its own right. Moreover, this practice-based approach has broader significance as well. Historians of biology have recently called for integrative approaches to evolutionary theory that move beyond facile internalist-externalist dualisms.111 This account responds to such calls, connecting the wider social context which shaped Wallace and Darwin's different routines, with the results of their practice. This comparative approach can readily be extended to other naturalists (e.g., Robert Grant, Henry Walter Bates, Joseph Hooker, Richard Spruce, and Thomas Huxley) and to Wallace and Darwin's subsequent practices and the products of these (e.g., Darwin's years of Notebook theorizing, on plants, and barnacle re? search; Wallace's anthropological theories, investigations of spiritual? ism, and socialist activism). This approach also provides a starting point for further investigations of the broader social and scientific factors that shaped Darwin and Wallace's different practices (e.g., taxonomic the? ories and methods, imperial politics and colonialism).

111 See, e.g., Jones 2002, Hull 2005.

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The more narrowly focused contrast between Wallace and Darwin has theoretical significance as well. Differences in their natural history practice made for differences in their theories about the objects of that practice, which can be seen alongside the material and literary contrasts. In particular, the contrasting emphases and distinctions inWallace and Darwin's writings from the field indicate the conceptual frameworks of their theories of natural selection. A detailed comparison of the devel? opment and articulation of these theories is beyond the scope of this paper. In any case, Darwin's theory has been thoroughly examined in its own right, and the history of its development extends well beyond the Beagle voyage.112 However, Wallace's theory, fully developed during his years in the field, has received far less attention. The practice-based account can throw light on Wallace's theory of natural selection, and thereby improve our understanding of both theories, with their obvious similarities and subtle differences. As noted above, Wallace went to the field with the aim of testing the hypothesis of transmutation of species. There are parallels with Dar? win's work of the 1830s and 1840s, as well as a direct connection: Wallace was influenced by Malthus and Lyell, wrote out his ideas in species notebooks, and drew on Darwin's own Journal of Researches for geological and biogeographical information.113 His writings from the field include two theoretical essays: the 1855 "Law" paper (written in Borneo during the rainy season) and the Ternate paper of 1858 (written on Gilolo when Wallace was ill).114 The latter, entitled "On the ten? dency of varieties to depart indefinitely from the original type," was presented alongside Darwin's theory at the July 1, 1858 meeting of the Linnean Society, galvanizing Darwin to write (and quickly publish) the Origin of Species the following year. Though these events have been examined in detail by various authors, Wallace's theory itself has been left (to paraphrase Shermer) in the shadow of Darwin's.115 There is a longstanding (though not unanimous) historical and philosophical consensus that Darwin's theory focused on selection of

112 See Darwin and Wallace, 1959 [1858]; de Beer, 1960; Darwin, 1968 [1859]; Stauffer, 1975; Burkhardt and Smith, 1985; Barrett et al. 1987; Darwin, 1987 [1842-1844]. Ospovat (1981) provides a classic account of the development of Darwin's theory; see also Kohn (1985). 113 Fichman, 2004; Slotten, 2004. 114 Wallace, 1855b; Darwin and Wallace, 1959 [1858]. Wallace's classificatory essays, and a more thorough account of the development of his theory of natural selection, will be dealt with in future papers. For the timing and site of Wallace's 1858 essay, see McKinney (1972), Slotten (2004). 115 See references in note 3; also CCD 1, pp. xvii-xix, 107.

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individual organisms, Wallace's on differential survival of varieties.116 The difference is often attributed to confusion or oversight on Wallace's part. Bowler (1976) claims that Wallace "recognized the Malthusian concept of struggle at the individual level, but immediately translated this into what would happen when permanent varieties came into conflict." Gould (2002) judges him more harshly: "Wallace never comprehended the question of levels at all, as he searched for adapta? tion wherever he could find it, oblivious to any problems raised by the locus of its action." Slotten (following Kottler, 1985) circumspectly notes that "it is unclear if in 1858 [Wallace] considered competition among individuals to be as important as competition among subpop? ulations in a species."117 The received view of Wallace that emerges from these representative assessments is that he was either a group selectionist, misguided about the mechanism of selection, or both. One difficulty for this received view is that Wallace evidently did conceive of natural selection as acting on individuals. His first description of the mechanism is explicitly individualistic:

...so long as a country remains physically unchanged, the numbers of its animal population cannot materially increase. If one species does so, some others requiring the same kind of food must diminish in proportion. The numbers that die annually must be immense; and as the individual existence of each animal depends upon itself, - those that die must be the weakest the very young, the aged, and - the diseased, while those that prolong their existence can only be - the most perfect in health and vigour those who are best able to obtain food regularly, and avoid their numerous enemies. It is..."a struggle for existence," in which the weakest and least perfectly organized must always succumb (italics mine).118

Wallace then discusses the operation of natural selection in terms of relative abundance of different species of an "allied group," such that better-adapted varieties replace the parent species in a process of

116 Osborn, 1894, p. 245; Beddall, 1968; Bowler, 1976; Ruse, 1980; Gould, 1980; Kleiner, 1985; Gayon, 1998, pp. 19-59; Ruse, 1999, p. 233; Gould, 2002, pp. 126-137; Browne, 2003, p. 18. For 'neutral theories' see Kottler, 1985; Slotten, 2004, p. 159. For a dissenting view, see: Mayr, 1982, pp. 494-497. Bulmer (2005) rejects the group selec? tionist interpretation of Wallace, but criticizes him instead for "misunderstanding the population of competing species" (p. 133). I thank an anonymous reviewer for bringing Bulmer's article to my attention. 117 Bowler, 1976, p. 24; Gould, 2002, p.136; Slotten, 2004, p. 159. 118 Darwin and Wallace, 1959 [1858], pp. 56-57. All quotations in this section are from Wallace's 1858 Ternate essay, unless otherwise noted.

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"progression and continued divergence." The received view of Wallace as group selectionist is understandable, given that much of his essay is in terms of adaptation and replacement at the level of varieties. But, seen in the context of his practice and other writings from the field, Wallace's theory is neither confused nor misguided. Nor does it posit an addi? tional process occurring over and above selection on individual organisms. After describing selection on individual organisms (an unusual departure from his typical emphasis), Wallace shifts to species and varieties, the focus of most of his writing, which his routine practice led him to emphasize:

Now, let some alteration of physical conditions occur in the district - - a long period of drought [etc.]... any change in fact tending to render existence more difficult to the species in question, and tasking its utmost powers to avoid complete extermination; it is evident that, of all the individuals composing the species, those forming the least numerous and most feebly organized variety would suffer first, and, were the pressure severe, must soon become extinct. The same causes continuing in action, and parent species would next suffer, would gradually diminish in numbers, and with a recurrence of similar unfavourable conditions might also become extinct. The superior variety would alone remain, and on a return to favorable circumstances would rapidly increase in numbers and occupy the place of the extinct species and variety (p. 58).

As inWallace's other writings from the field, the individual and group levels are linked via the notion of abundance, the number of individuals composing a group. Wallace defines the best-adapted species as those that "obtain and preserve a superiority in population," while those that exhibit "some defect of power or organization ...must diminish in numbers, and, in extreme cases, become altogether extinct" (p. 57). Thus "continuance of the species and the keeping up of the average number of individuals" amount to the same thing (p. 55). The core of Wallace's theory, like Darwin's, is survival of the individuals that are best at obtaining food and avoiding predators. The two men thus rightly recognized their theories as essentially the same, despite their different emphases.119 The practice-based account provides a more nuanced and sympa? thetic interpretation of Wallace's theory, and clarifies its contrast with Darwin's individualistic focus. Though much changed in Darwin's practice between 1836 and 1858, his emphasis on individual organisms

119 E.g., CCD 7, 107; Wallace, 1906 (volume 2), pp. 16-22.

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and vivid examples remained a feature of his writing and theorizing.120 Wallace's routine practice of hunting, processing, and classifying shaped the theory that he conceived in the midst of these activities. In the material and literary results of his natural history practice, groups figure prominently, but individual organisms form the base or starting-point. The same pattern can be seen in his 1858 theory. Individual organisms are in the background of Wallace's theory, constituting the basis for the claims he makes about species and varieties. The practice-based account - thus places Wallace's theory of natural selection in its proper context - the field rather than leaving it in Darwin's shadow.

Acknowledgments

I thank Sander Gliboff, John Beatty, Jane Camerini, Steve Crowley, Michael Dickison, Elisabeth Lloyd, Rasmus Winther, the editor and two anonymous reviewers for insightful discussion and comments; the Linnean Society for permission to examine Wallace's unpublished MSS; and Jane Camerini for guidance on Wallace's published and unpublished writings. Earlier drafts of this paper were presented at the Vienna International Summer University workshop on "The Quest for Objectivity" (Vienna, Austria, July 26, 2004) and as the 2005 Hanson Prize Lecture at Indiana University (Bloomington, IN, January 20, 2006). Many thanks to participants on both occasions for helpful questions and comments. Any errors are mine.

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